Publications

Displaying 401 - 500 of 1209
  • Hagoort, P. (2008). Should psychology ignore the language of the brain? Current Directions in Psychological Science, 17(2), 96-101. doi:10.1111/j.1467-8721.2008.00556.x.

    Abstract

    Claims that neuroscientific data do not contribute to our understanding of psychological functions have been made recently. Here I argue that these criticisms are solely based on an analysis of functional magnetic resonance imaging (fMRI) studies. However, fMRI is only one of the methods in the toolkit of cognitive neuroscience. I provide examples from research on event-related brain potentials (ERPs) that have contributed to our understanding of the cognitive architecture of human language functions. In addition, I provide evidence of (possible) contributions from fMRI measurements to our understanding of the functional architecture of language processing. Finally, I argue that a neurobiology of human language that integrates information about the necessary genetic and neural infrastructures will allow us to answer certain questions that are not answerable if all we have is evidence from behavior.
  • Hagoort, P. (2008). The fractionation of spoken language understanding by measuring electrical and magnetic brain signals. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences, 363, 1055-1069. doi:10.1098/rstb.2007.2159.

    Abstract

    This paper focuses on what electrical and magnetic recordings of human brain activity reveal about spoken language understanding. Based on the high temporal resolution of these recordings, a fine-grained temporal profile of different aspects of spoken language comprehension can be obtained. Crucial aspects of speech comprehension are lexical access, selection and semantic integration. Results show that for words spoken in context, there is no ‘magic moment’ when lexical selection ends and semantic integration begins. Irrespective of whether words have early or late recognition points, semantic integration processing is initiated before words can be identified on the basis of the acoustic information alone. Moreover, for one particular event-related brain potential (ERP) component (the N400), equivalent impact of sentence- and discourse-semantic contexts is observed. This indicates that in comprehension, a spoken word is immediately evaluated relative to the widest interpretive domain available. In addition, this happens very quickly. Findings are discussed that show that often an unfolding word can be mapped onto discourse-level representations well before the end of the word. Overall, the time course of the ERP effects is compatible with the view that the different information types (lexical, syntactic, phonological, pragmatic) are processed in parallel and influence the interpretation process incrementally, that is as soon as the relevant pieces of information are available. This is referred to as the immediacy principle.
  • Hagoort, P. (1998). De electrofysiologie van taal: Wat hersenpotentialen vertellen over het menselijk taalvermogen. Neuropraxis, 2, 223-229.
  • Hagoort, P. (1998). De spreker als sprinter. Psychologie, 17, 48-49.
  • Li, X., Hagoort, P., & Yang, Y. (2008). Event-related potential evidence on the influence of accentuation in spoken discourse comprehension in Chinese. Journal of Cognitive Neuroscience, 20(5), 906-915. doi:10.1162/jocn.2008.20512.

    Abstract

    In an event-related potential experiment with Chinese discourses as material, we investigated how and when accentuation influences spoken discourse comprehension in relation to the different information states of the critical words. These words could either provide new or old information. It was shown that variation of accentuation influenced the amplitude of the N400, with a larger amplitude for accented than deaccented words. In addition, there was an interaction between accentuation and information state. The N400 amplitude difference between accented and deaccented new information was smaller than that between accented and deaccented old information. The results demonstrate that, during spoken discourse comprehension, listeners rapidly extract the semantic consequences of accentuation in relation to the previous discourse context. Moreover, our results show that the N400 amplitude can be larger for correct (new,accented words) than incorrect (new, deaccented words) information. This, we argue, proves that the N400 does not react to semantic anomaly per se, but rather to semantic integration load, which is higher for new information.
  • Hagoort, P., Ramsey, N. F., & Jensen, O. (2008). De gereedschapskist van de cognitieve neurowetenschap. In F. Wijnen, & F. Verstraten (Eds.), Het brein te kijk: Verkenning van de cognitieve neurowetenschap (pp. 41-75). Amsterdam: Harcourt Assessment.
  • Hagoort, P. (2004). Er is geen behoefte aan trompetten als gordijnen. In H. Procee, H. Meijer, P. Timmerman, & R. Tuinsma (Eds.), Bij die wereld wil ik horen! Zesendertig columns en drie essays over de vorming tot academicus (pp. 78-80). Amsterdam: Boom.
  • Hagoort, P., Hald, L. A., Bastiaansen, M. C. M., & Petersson, K. M. (2004). Integration of word meaning and world knowledge in language comprehension. Science, 304(5669), 438-441. doi:10.1126/science.1095455.

    Abstract

    Although the sentences that we hear or read have meaning, this does not necessarily mean that they are also true. Relatively little is known about the critical brain structures for, and the relative time course of, establishing the meaning and truth of linguistic expressions. We present electroencephalogram data that show the rapid parallel integration of both semantic and world
    knowledge during the interpretation of a sentence. Data from functional magnetic resonance imaging revealed that the left inferior prefrontal cortex is involved in the integration of both meaning and world knowledge. Finally, oscillatory brain responses indicate that the brain keeps a record of what makes a sentence hard to interpret.
  • Hagoort, P. (2004). Het zwarte gat tussen brein en bewustzijn. In N. Korteweg (Ed.), De oorsprong: Over het ontstaan van het leven en alles eromheen (pp. 107-124). Amsterdam: Boom.
  • Hagoort, P. (2008). Mijn omweg naar de filosofie. Algemeen Nederlands Tijdschrift voor Wijsbegeerte, 100(4), 303-310.
  • Hagoort, P. (2012). From ants to music and language [Preface]. In A. D. Patel, Music, language, and the brain [Chinese translation] (pp. 9-10). Shanghai: East China Normal University Press Ltd.
  • Hagoort, P., & Beckmann, C. F. (2019). Key issues and future directions: The neural architecture for language. In P. Hagoort (Ed.), Human language: From genes and brains to behavior (pp. 527-532). Cambridge, MA: MIT Press.
  • Hagoort, P. (1998). Hersenen en taal in onderzoek en praktijk. Neuropraxis, 6, 204-205.
  • Hagoort, P. (2012). Het muzikale brein. Speling: Tijdschrift voor bezinning. Muziek als bron van bezieling, 64(1), 44-48.
  • Hagoort, P. (2012). Het sprekende brein. MemoRad, 17(1), 27-30.

    Abstract

    Geen andere soort dan homo sapiens heeft in de loop van zijn evolutionaire geschiedenis een communicatiesysteem ontwikkeld waarin een eindig aantal symbolen samen met een reeks van regels voor het combineren daarvan een oneindig aantal uitdrukkingen mogelijk maakt. Dit natuurlijke taalsysteem stelt leden van onze soort in staat gedachten een uiterlijke vorm te geven en uit te wisselen met de sociale groep en, door de uitvinding van schriftsystemen, met de gehele samenleving. Spraak en taal zijn effectieve middelen voor het behoud van sociale cohesie in samenlevingen waarvan de groepsgrootte en de complexe sociale organisatie van dien aard is dat dit niet langer kan door middel van ‘vlooien’, de wijze waarop onze genetische buren, de primaten van de oude wereld, sociale cohesie bevorderen [1,2].
  • Hagoort, P. (2019). Introduction. In P. Hagoort (Ed.), Human language: From genes and brains to behavior (pp. 1-6). Cambridge, MA: MIT Press.
  • Hagoort, P. (1998). The shadows of lexical meaning in patients with semantic impairments. In B. Stemmer, & H. Whitaker (Eds.), Handbook of neurolinguistics (pp. 235-248). New York: Academic Press.
  • Hagoort, P. (2019). The meaning making mechanism(s) behind the eyes and between the ears. Philosophical Transactions of the Royal Society of London, Series B: Biological Sciences, 375: 20190301. doi:10.1098/rstb.2019.0301.

    Abstract

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

    Abstract

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

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  • Hagoort, P. (2008). Über Broca, Gehirn und Bindung. In Jahrbuch 2008: Tätigkeitsberichte der Institute. München: Generalverwaltung der Max-Planck-Gesellschaft. Retrieved from http://www.mpg.de/306524/forschungsSchwerpunkt1?c=166434.

    Abstract

    Beim Sprechen und beim Sprachverstehen findet man die Wortbedeutung im Gedächtnis auf und kombiniert sie zu größeren Einheiten (Unifikation). Solche Unifikations-Operationen laufen auf unterschiedlichen Ebenen der Sprachverarbeitung ab. In diesem Beitrag wird ein Rahmen vorgeschlagen, in dem psycholinguistische Modelle mit neurobiologischer Sprachbetrachtung in Verbindung gebracht werden. Diesem Vorschlag zufolge spielt der linke inferiore frontale Gyrus (LIFG) eine bedeutende Rolle bei der Unifi kation
  • Hallé, P., & Cristia, A. (2012). Global and detailed speech representations in early language acquisition. In S. Fuchs, M. Weirich, D. Pape, & P. Perrier (Eds.), Speech planning and dynamics (pp. 11-38). Frankfurt am Main: Peter Lang.

    Abstract

    We review data and hypotheses dealing with the mental representations for perceived and produced speech that infants build and use over the course of learning a language. In the early stages of speech perception and vocal production, before the emergence of a receptive or a productive lexicon, the dominant picture emerging from the literature suggests rather non-analytic representations based on units of the size of the syllable: Young children seem to parse speech into syllable-sized units in spite of their ability to detect sound equivalence based on shared phonetic features. Once a productive lexicon has emerged, word form representations are initially rather underspecified phonetically but gradually become more specified with lexical growth, up to the phoneme level. The situation is different for the receptive lexicon, in which phonetic specification for consonants and vowels seem to follow different developmental paths. Consonants in stressed syllables are somewhat well specified already at the first signs of a receptive lexicon, and become even better specified with lexical growth. Vowels seem to follow a different developmental path, with increasing flexibility throughout lexical development. Thus, children come to exhibit a consonant vowel asymmetry in lexical representations, which is clear in adult representations.
  • Hammarström, H. (2012). A full-scale test of the language farming dispersal hypothesis. In S. Wichmann, & A. P. Grant (Eds.), Quantitative approaches to linguistic diversity: Commemorating the centenary of the birth of Morris Swadesh (pp. 7-22). Amsterdam: Benjamins.

    Abstract

    Originally published in Diachronica 27:2 (2010) One attempt at explaining why some language families are large (while others are small) is the hypothesis that the families that are now large became large because their ancestral speakers had a technological advantage, most often agriculture. Variants of this idea are referred to as the Language Farming Dispersal Hypothesis. Previously, detailed language family studies have uncovered various supporting examples and counterexamples to this idea. In the present paper I weigh the evidence from ALL attested language families. For each family, I use the number of member languages as a measure of cardinal size, member language coordinates to measure geospatial size and ethnographic evidence to assess subsistence status. This data shows that, although agricultural families tend to be larger in cardinal size, their size is hardly due to the simple presence of farming. If farming were responsible for language family expansions, we would expect a greater east-west geospatial spread of large families than is actually observed. The data, however, is compatible with weaker versions of the farming dispersal hypothesis as well with models where large families acquire farming because of their size, rather than the other way around.
  • Hammarström, H. (2019). An inventory of Bantu languages. In M. Van de Velde, K. Bostoen, D. Nurse, & G. Philippson (Eds.), The Bantu languages (2nd). London: Routledge.

    Abstract

    This chapter aims to provide an updated list of all Bantu languages known at present and to provide individual pointers to further information on the inventory. The area division has some correlation with what are perceived genealogical relations between Bantu languages, but they are not defined as such and do not change whenever there is an update in our understanding of genealogical relations. Given the popularity of Guthrie codes in Bantu linguistics, our listing also features a complete mapping to Guthrie codes. The language inventory listed excludes sign languages used in the Bantu area, speech registers, pidgins, drummed/whistled languages and urban youth languages. Pointers to such languages in the Bantu area are included in the continent-wide overview in Hammarstrom. The most important alternative names, subvarieties and spelling variants are given for each language, though such lists are necessarily incomplete and reflect some degree of arbitrary selection.
  • Hammarström, H. (2012). [Review of Ferdinand von Mengden, Cardinal numerals: Old English from a cross-linguistic perspective]. Linguistic Typology, 16, 321-324. doi:10.1515/lity-2012-0010.
  • Hammarström, H., & van den Heuvel, W. (2012). Introduction to the LLM Special Issue 2012 on the History, contact and classification of Papuan languages. Language & Linguistics in Melanesia, 2012(Special Issue, Part 1), i-v.
  • Hammarström, H. (2012). Pronouns and the (Preliminary) Classification of Papuan languages. Language and linguistics in Melanesia, Special issue 2012 Part 2, 428-539. Retrieved from http://www.langlxmelanesia.com/hammarstrom428-539.pdf.

    Abstract

    A series of articles by Ross (1995, 2001, 2005) use pronoun sim- ilarities to gauge relatedness between various Papuan microgroups, arguing that the similarities could not be the result of chance or bor- rowing. I argue that a more appropriate manner of calculating chance gives a signicantly dierent result: when cross-comparing a pool of languages the prospects for chance matches of rst and second person pronouns are very good. Using pronoun form data from over 3000 lan- guages and over 300 language families inside and outside New Guinea, I show that there is, nevertheless, a tendency for Papuan pronouns to use certain consonants more often in 1P and 2P SG forms than in the rest of the world. This could reect an underlying family. An alter- native explanation is the established Papuan areal feature of having a small consonant inventory, which results in a higher functional load on the remaining consonants, which is, in turn, reected in the enhanced popularity of certain consonants in pronouns of those languages. A test of surface forms (i.e., non-reconstructed forms) favours the latter explanation.
  • Hammarström, H., & Nordhoff, S. (2012). The languages of Melanesia: Quantifying the level of coverage. In N. Evans, & M. Klamer (Eds.), Melanesian languages on the edge of Asia: Challenges for the 21st Century (pp. 13-33). Honolulu: University of Hawai'i Press. Retrieved from http://hdl.handle.net/10125/4559.
  • Han, J.-I., & Verdonschot, R. G. (2019). Spoken-word production in Korean: A non-word masked priming and phonological Stroop task investigation. Quarterly Journal of Experimental Psychology, 72(4), 901-912. doi:10.1177/1747021818770989.

    Abstract

    Speech production studies have shown that phonological unit initially used to fill the metrical frame during phonological encoding is language specific, that is, a phoneme for English and Dutch, an atonal syllable for Mandarin Chinese, and a mora for Japanese. However, only a few studies chronometrically investigated speech production in Korean, and they obtained mixed results. Korean is particularly interesting as there might be both phonemic and syllabic influences during phonological encoding. The purpose of this study is to further examine the initial phonological preparation unit in Korean, employing a masked priming task (Experiment 1) and a phonological Stroop task (Experiment 2). The results showed that significant onset (and onset-plus, that is, consonant-vowel [CV]) effects were found in both experiments, but there was no compelling evidence for a prominent role for the syllable. When the prime words were presented in three different forms related to the targets, namely, without any change, with re-syllabified codas, and with nasalised codas, there were no significant differences in facilitation among the three forms. Alternatively, it is also possible that participants may not have had sufficient time to process the primes up to the point that re-syllabification or nasalisation could have been carried out. In addition, the results of a Stroop task demonstrated that the onset phoneme effect was not driven by any orthographic influence. These findings suggest that the onset segment and not the syllable is the initial (or proximate) phonological unit used in the segment-to-frame encoding process during speech planning in Korean.

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    stimuli for experiment 1 and 2
  • Hanique, I., & Ernestus, M. (2012). The role of morphology in acoustic reduction. Lingue e linguaggio, 2012(2), 147-164. doi:10.1418/38783.

    Abstract

    This paper examines the role of morphological structure in the reduced pronunciation of morphologically complex words by discussing and re-analyzing data from the literature. Acoustic reduction refers to the phenomenon that, in spontaneous speech, phonemes may be shorter or absent. We review studies investigating effects of the repetition of a morpheme, of whether a segment plays a crucial role in the identification of its morpheme, and of a word's morphological decomposability. We conclude that these studies report either no effects of morphological structure or effects that are open to alternative interpretations. Our analysis also reveals the need for a uniform definition of morphological decomposability. Furthermore, we examine whether the reduction of segments in morphologically complex words correlates with these segments' contribution to the identification of the whole word, and discuss previous studies and new analyses supporting this hypothesis. We conclude that the data show no convincing evidence that morphological structure conditions reduction, which contrasts with the expectations of several models of speech production and of morphological processing (e.g., weaver++ and dual-route models). The data collected so far support psycholinguistic models which assume that all morphologically complex words are processed as complete units.
  • Hanulikova, A., & Dietrich, R. (2008). Die variable Coda in der slowakisch-deutschen Interimsprache. In M. Tarvas (Ed.), Tradition und Geschichte im literarischen und sprachwissenschaftlichen Kontext (pp. 119-130). Bern: Peter Lang.
  • Hanulikova, A., Dediu, D., Fang, Z., Basnakova, J., & Huettig, F. (2012). Individual differences in the acquisition of a complex L2 phonology: A training study. Language Learning, 62(Supplement S2), 79-109. doi:10.1111/j.1467-9922.2012.00707.x.

    Abstract

    Many learners of a foreign language (L2) struggle to correctly pronounce newly-learned speech sounds, yet many others achieve this with apparent ease. Here we explored how a training study of learning complex consonant clusters at the very onset of the L2 acquisition can inform us about L2 learning in general and individual differences in particular. To this end, adult Dutch native speakers were trained on Slovak words with complex consonant clusters (e.g., pstruh /pstrux/‘trout’, štvrť /ʃtvrc/ ‘quarter’) using auditory and orthographic input. In the same session following training, participants were tested on a battery of L2 perception and production tasks. The battery of L2 tests was repeated twice more with one week between each session. In the first session, an additional battery of control tests was used to test participants’ native language (L1) skills. Overall, in line with some previous research, participants showed only weak learning effects across the L2 perception tasks. However, there were considerable individual differences across all L2 tasks, which remained stable across sessions. Only two participants showed overall high L2 production performance that fell within 2 standard deviations of the mean ratings obtained for an L1 speaker. The mispronunciation detection task was the only perception task which significantly predicted production performance in the final session. We conclude by discussing several recommendations for future L2 learning studies.
  • Hanulikova, A., & Weber, A. (2012). Sink positive: Linguistic experience with th substitutions influences nonnative word recognition. Attention, Perception & Psychophysics, 74(3), 613-629. doi:10.3758/s13414-011-0259-7.

    Abstract

    We used eyetracking, perceptual discrimination, and production tasks to examine the influences of perceptual similarity and linguistic experience on word recognition in nonnative (L2) speech. Eye movements to printed words were tracked while German and Dutch learners of English heard words containing one of three pronunciation variants (/t/, /s/, or /f/) of the interdental fricative /θ/. Irrespective of whether the speaker was Dutch or German, looking preferences for target words with /θ/ matched the preferences for producing /s/ variants in German speakers and /t/ variants in Dutch speakers (as determined via the production task), while a control group of English participants showed no such preferences. The perceptually most similar and most confusable /f/ variant (as determined via the discrimination task) was never preferred as a match for /θ/. These results suggest that linguistic experience with L2 pronunciations facilitates recognition of variants in an L2, with effects of frequency outweighing effects of perceptual similarity.
  • Hanulikova, A., Van Alphen, P. M., Van Goch, M. M., & Weber, A. (2012). When one person’s mistake is another’s standard usage: The effect of foreign accent on syntactic processing. Journal of Cognitive Neuroscience, 24(4), 878-887. doi:10.1162/jocn_a_00103.

    Abstract

    How do native listeners process grammatical errors that are frequent in non-native speech? We investigated whether the neural correlates of syntactic processing are modulated by speaker identity. ERPs to gender agreement errors in sentences spoken by a native speaker were compared with the same errors spoken by a non-native speaker. In line with previous research, gender violations in native speech resulted in a P600 effect (larger P600 for violations in comparison with correct sentences), but when the same violations were produced by the non-native speaker with a foreign accent, no P600 effect was observed. Control sentences with semantic violations elicited comparable N400 effects for both the native and the non-native speaker, confirming no general integration problem in foreign-accented speech. The results demonstrate that the P600 is modulated by speaker identity, extending our knowledge about the role of speaker's characteristics on neural correlates of speech processing.
  • Harmon, Z., Idemaru, K., & Kapatsinski, V. (2019). Learning mechanisms in cue reweighting. Cognition, 189, 76-88. doi:10.1016/j.cognition.2019.03.011.

    Abstract

    Feedback has been shown to be effective in shifting attention across perceptual cues to a phonological contrast in speech perception (Francis, Baldwin & Nusbaum, 2000). However, the learning mechanisms behind this process remain obscure. We compare the predictions of supervised error-driven learning (Rescorla & Wagner, 1972) and reinforcement learning (Sutton & Barto, 1998) using computational simulations. Supervised learning predicts downweighting of an informative cue when the learner receives evidence that it is no longer informative. In contrast, reinforcement learning suggests that a reduction in cue weight requires positive evidence for the informativeness of an alternative cue. Experimental evidence supports the latter prediction, implicating reinforcement learning as the mechanism behind the effect of feedback on cue weighting in speech perception. Native English listeners were exposed to either bimodal or unimodal VOT distributions spanning the unaspirated/aspirated boundary (bear/pear). VOT is the primary cue to initial stop voicing in English. However, lexical feedback in training indicated that VOT was no longer predictive of voicing. Reduction in the weight of VOT was observed only when participants could use an alternative cue, F0, to predict voicing. Frequency distributions had no effect on learning. Overall, the results suggest that attention shifting in learning the phonetic cues to phonological categories is accomplished using simple reinforcement learning principles that also guide the choice of actions in other domains.
  • Harneit, A., Braun, U., Geiger, L. S., Zang, Z., Hakobjan, M., Van Donkelaar, M. M. J., Schweiger, J. I., Schwarz, K., Gan, G., Erk, S., Heinz, A., Romanczuk‐Seiferth, N., Witt, S., Rietschel, M., Walter, H., Franke, B., Meyer‐Lindenberg, A., & Tost, H. (2019). MAOA-VNTR genotype affects structural and functional connectivity in distributed brain networks. Human Brain Mapping, 40(18), 5202-5212. doi:10.1002/hbm.24766.

    Abstract

    Previous studies have linked the low expression variant of a variable number of tandem repeat polymorphism in the monoamine oxidase A gene (MAOA‐L) to the risk for impulsivity and aggression, brain developmental abnormalities, altered cortico‐limbic circuit function, and an exaggerated neural serotonergic tone. However, the neurobiological effects of this variant on human brain network architecture are incompletely understood. We studied healthy individuals and used multimodal neuroimaging (sample size range: 219–284 across modalities) and network‐based statistics (NBS) to probe the specificity of MAOA‐L‐related connectomic alterations to cortical‐limbic circuits and the emotion processing domain. We assessed the spatial distribution of affected links across several neuroimaging tasks and data modalities to identify potential alterations in network architecture. Our results revealed a distributed network of node links with a significantly increased connectivity in MAOA‐L carriers compared to the carriers of the high expression (H) variant. The hyperconnectivity phenotype primarily consisted of between‐lobe (“anisocoupled”) network links and showed a pronounced involvement of frontal‐temporal connections. Hyperconnectivity was observed across functional magnetic resonance imaging (fMRI) of implicit emotion processing (pFWE = .037), resting‐state fMRI (pFWE = .022), and diffusion tensor imaging (pFWE = .044) data, while no effects were seen in fMRI data of another cognitive domain, that is, spatial working memory (pFWE = .540). These observations are in line with prior research on the MAOA‐L variant and complement these existing data by novel insights into the specificity and spatial distribution of the neurogenetic effects. Our work highlights the value of multimodal network connectomic approaches for imaging genetics.
  • Hartz, S. M., Short, S. E., Saccone, N. L., Culverhouse, R., Chen, L., Schwantes-An, T.-H., Coon, H., Han, Y., Stephens, S. H., Sun, J., Chen, X., Ducci, F., Dueker, N., Franceschini, N., Frank, J., Geller, F., Gubjartsson, D., Hansel, N. N., Jiang, C., Keskitalo-Vuokko, K. and 132 moreHartz, S. M., Short, S. E., Saccone, N. L., Culverhouse, R., Chen, L., Schwantes-An, T.-H., Coon, H., Han, Y., Stephens, S. H., Sun, J., Chen, X., Ducci, F., Dueker, N., Franceschini, N., Frank, J., Geller, F., Gubjartsson, D., Hansel, N. N., Jiang, C., Keskitalo-Vuokko, K., Liu, Z., Lyytikainen, L.-P., Michel, M., Rawal, R., Rosenberger, A., Scheet, P., Shaffer, J. R., Teumer, A., Thompson, J. R., Vink, J. M., Vogelzangs, N., Wenzlaff, A. S., Wheeler, W., Xiao, X., Yang, B.-Z., Aggen, S. H., Balmforth, A. J., Baumeister, S. E., Beaty, T., Bennett, S., Bergen, A. W., Boyd, H. A., Broms, U., Campbell, H., Chatterjee, N., Chen, J., Cheng, Y.-C., Cichon, S., Couper, D., Cucca, F., Dick, D. M., Foroud, T., Furberg, H., Giegling, I., Gu, F., Hall, A. S., Hallfors, J., Han, S., Hartmann, A. M., Hayward, C., Heikkila, K., Hewitt, J. K., Hottenga, J. J., Jensen, M. K., Jousilahti, P., Kaakinen, M., Kittner, S. J., Konte, B., Korhonen, T., Landi, M.-T., Laatikainen, T., Leppert, M., Levy, S. M., Mathias, R. A., McNeil, D. W., Medland, S. E., Montgomery, G. W., Muley, T., Murray, T., Nauck, M., North, K., Pergadia, M., Polasek, O., Ramos, E. M., Ripatti, S., Risch, A., Ruczinski, I., Rudan, I., Salomaa, V., Schlessinger, D., Styrkarsdottir, U., Terracciano, A., Uda, M., Willemsen, G., Wu, X., Abecasis, G., Barnes, K., Bickeboller, H., Boerwinkle, E., Boomsma, D. I., Caporaso, N., Duan, J., Edenberg, H. J., Francks, C., Gejman, P. V., Gelernter, J., Grabe, H. J., Hops, H., Jarvelin, M.-R., Viikari, J., Kahonen, M., Kendler, K. S., Lehtimaki, T., Levinson, D. F., Marazita, M. L., Marchini, J., Melbye, M., Mitchell, B., Murray, J. C., Nothen, M. M., Penninx, B. W., Raitakari, O., Rietschel, M., Rujescu, D., Samani, N. J., Sanders, A. R., Schwartz, A. G., Shete, S., Shi, J., Spitz, M., Stefansson, K., Swan, G. E., Thorgeirsson, T., Volzke, H., Wei, Q., Wichmann, H.-E., Amos, C. I., Breslau, N., Cannon, D. S., Ehringer, M., Grucza, R., Hatsukami, D., Heath, A., Johnson, E. O., Kaprio, J., Madden, P., Martin, N. G., Stevens, V. L., Stitzel, J. A., Weiss, R. B., Kraft, P., & Bierut, L. J. (2012). Increased genetic vulnerability to smoking at CHRNA5 in early-onset smokers. Archives of General Psychiatry, 69, 854-860. doi:10.1001/archgenpsychiatry.2012.124.

    Abstract

    CONTEXT Recent studies have shown an association between cigarettes per day (CPD) and a nonsynonymous single-nucleotide polymorphism in CHRNA5, rs16969968. OBJECTIVE To determine whether the association between rs16969968 and smoking is modified by age at onset of regular smoking. DATA SOURCES Primary data. STUDY SELECTION Available genetic studies containing measures of CPD and the genotype of rs16969968 or its proxy. DATA EXTRACTION Uniform statistical analysis scripts were run locally. Starting with 94 050 ever-smokers from 43 studies, we extracted the heavy smokers (CPD >20) and light smokers (CPD ≤10) with age-at-onset information, reducing the sample size to 33 348. Each study was stratified into early-onset smokers (age at onset ≤16 years) and late-onset smokers (age at onset >16 years), and a logistic regression of heavy vs light smoking with the rs16969968 genotype was computed for each stratum. Meta-analysis was performed within each age-at-onset stratum. DATA SYNTHESIS Individuals with 1 risk allele at rs16969968 who were early-onset smokers were significantly more likely to be heavy smokers in adulthood (odds ratio [OR] = 1.45; 95% CI, 1.36-1.55; n = 13 843) than were carriers of the risk allele who were late-onset smokers (OR = 1.27; 95% CI, 1.21-1.33, n = 19 505) (P = .01). CONCLUSION These results highlight an increased genetic vulnerability to smoking in early-onset smokers.

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  • Haun, D. B. M., & Call, J. (2008). Imitation recognition in great apes. Current Biology, 18(7), 288-290. doi:10.1016/j.cub.2008.02.031.

    Abstract

    Human infants imitate not only to acquire skill, but also as a fundamental part of social interaction [1] , [2] and [3] . They recognise when they are being imitated by showing increased visual attention to imitators (implicit recognition) and by engaging in so-called testing behaviours (explicit recognition). Implicit recognition affords the ability to recognize structural and temporal contingencies between actions across agents, whereas explicit recognition additionally affords the ability to understand the directional impact of one's own actions on others' actions [1] , [2] and [3] . Imitation recognition is thought to foster understanding of social causality, intentionality in others and the formation of a concept of self as different from other [3] , [4] and [5] . Pigtailed macaques (Macaca nemestrina) implicitly recognize being imitated [6], but unlike chimpanzees [7], they show no sign of explicit imitation recognition. We investigated imitation recognition in 11 individuals from the four species of non-human great apes. We replicated results previously found with a chimpanzee [7] and, critically, have extended them to the other great ape species. Our results show a general prevalence of imitation recognition in all great apes and thereby demonstrate important differences between great apes and monkeys in their understanding of contingent social interactions.
  • Haun, D. B. M., Rekers, Y., & Tomasello, M. (2012). Majority-biased transmission in chimpanzees and human children, but not orangutans. Current Biology, 22, 727-731. doi:10.1016/j.cub.2012.03.006.

    Abstract

    Cultural transmission is a key component of human evolution. Two of humans' closest living relatives, chimpanzees and orangutans, have also been argued to transmit behavioral traditions across generations culturally [ [1], [2] and [3]], but how much the process might resemble the human process is still in large part unknown. One key phenomenon of human cultural transmission is majority-biased transmission: the increased likelihood for learners to end up not with the most frequent behavior but rather with the behavior demonstrated by most individuals. Here we show that chimpanzees and human children as young as 2 years of age, but not orangutans, are more likely to copy an action performed by three individuals, once each, than an action performed by one individual three times. The tendency to acquire the behaviors of the majority has been posited as key to the transmission of relatively safe, reliable, and productive behavioral strategies [ [4], [5], [6] and [7]] but has not previously been demonstrated in primates.
  • Haworth, S., Shapland, C. Y., Hayward, C., Prins, B. P., Felix, J. F., Medina-Gomez, C., Rivadeneira, F., Wang, C., Ahluwalia, T. S., Vrijheid, M., Guxens, M., Sunyer, J., Tachmazidou, I., Walter, K., Iotchkova, V., Jackson, A., Cleal, L., Huffmann, J., Min, J. L., Sass, L. and 15 moreHaworth, S., Shapland, C. Y., Hayward, C., Prins, B. P., Felix, J. F., Medina-Gomez, C., Rivadeneira, F., Wang, C., Ahluwalia, T. S., Vrijheid, M., Guxens, M., Sunyer, J., Tachmazidou, I., Walter, K., Iotchkova, V., Jackson, A., Cleal, L., Huffmann, J., Min, J. L., Sass, L., Timmers, P. R. H. J., UK10K consortium, Davey Smith, G., Fisher, S. E., Wilson, J. F., Cole, T. J., Fernandez-Orth, D., Bønnelykke, K., Bisgaard, H., Pennell, C. E., Jaddoe, V. W. V., Dedoussis, G., Timpson, N. J., Zeggini, E., Vitart, V., & St Pourcain, B. (2019). Low-frequency variation in TP53 has large effects on head circumference and intracranial volume. Nature Communications, 10: 357. doi:10.1038/s41467-018-07863-x.

    Abstract

    Cranial growth and development is a complex process which affects the closely related traits of head circumference (HC) and intracranial volume (ICV). The underlying genetic influences affecting these traits during the transition from childhood to adulthood are little understood, but might include both age-specific genetic influences and low-frequency genetic variation. To understand these influences, we model the developmental genetic architecture of HC, showing this is genetically stable and correlated with genetic determinants of ICV. Investigating up to 46,000 children and adults of European descent, we identify association with final HC and/or final ICV+HC at 9 novel common and low-frequency loci, illustrating that genetic variation from a wide allele frequency spectrum contributes to cranial growth. The largest effects are reported for low-frequency variants within TP53, with 0.5 cm wider heads in increaser-allele carriers versus non-carriers during mid-childhood, suggesting a previously unrecognized role of TP53 transcripts in human cranial development.

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  • Hayano, K. (2004). Kaiwa ni okeru ninshikiteki ken’i no koushou: Shuujoshi yo, ne, odoroki hyouji no bunpu to kinou [Negotiation of Epistemic Authority in Conversation: on the use of final particles yo, ne and surprise markers]. Studies in Pragmatics, 6, 17-28.
  • Hayano, K. (2008). Talk and body: Negotiating action framework and social relationship in conversation. Studies in English and American Literature, 43, 187-198.
  • Hervais-Adelman, A., Carlyon, R. P., Johnsrude, I. S., & Davis, M. H. (2012). Brain regions recruited for the effortful comprehension of noise-vocoded words. Language and Cognitive Processes, 27(7-8), 1145-1166. doi:10.1080/01690965.2012.662280.

    Abstract

    We used functional magnetic resonance imaging (fMRI) to investigate the neural basis of comprehension and perceptual learning of artificially degraded [noise vocoded (NV)] speech. Fifteen participants were scanned while listening to 6-channel vocoded words, which are difficult for naive listeners to comprehend, but can be readily learned with appropriate feedback presentations. During three test blocks, we compared responses to potentially intelligible NV words, incomprehensible distorted words and clear speech. Training sessions were interleaved with the test sessions and included paired presentation of clear then noise-vocoded words: a type of feedback that enhances perceptual learning. Listeners' comprehension of NV words improved significantly as a consequence of training. Listening to NV compared to clear speech activated left insula, and prefrontal and motor cortices. These areas, which are implicated in speech production, may play an active role in supporting the comprehension of degraded speech. Elevated activation in the precentral gyrus during paired clear-then-distorted presentations that enhance learning further suggests a role for articulatory representations of speech in perceptual learning of degraded speech.
  • Hervais-Adelman, A., Kumar, U., Mishra, R. K., Tripathi, V. N., Guleria, A., Singh, J. P., Eisner, F., & Huettig, F. (2019). Learning to read recycles visual cortical networks without destruction. Science Advances, 5(9): eaax0262. doi:10.1126/sciadv.aax0262.

    Abstract

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

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  • Hervais-Adelman, A., Davis, M. H., Johnsrude, I. S., & Carlyon, R. P. (2008). Perceptual learning of noise vocoded words: Effects of feedback and lexicality. Journal of Experimental Psychology: Human Perception and Performance, 34(2), 460-474. doi:10.1037/0096-1523.34.2.460.

    Abstract

    Speech comprehension is resistant to acoustic distortion in the input, reflecting listeners' ability to adjust perceptual processes to match the speech input. This adjustment is reflected in improved comprehension of distorted speech with experience. For noise vocoding, a manipulation that removes spectral detail from speech, listeners' word report showed a significantly greater improvement over trials for listeners that heard clear speech presentations before rather than after hearing distorted speech (clear-then-distorted compared with distorted-then-clear feedback, in Experiment 1). This perceptual learning generalized to untrained words suggesting a sublexical locus for learning and was equivalent for word and nonword training stimuli (Experiment 2). These findings point to the crucial involvement of phonological short-term memory and top-down processes in the perceptual learning of noise-vocoded speech. Similar processes may facilitate comprehension of speech in an unfamiliar accent or following cochlear implantation.
  • Heyselaar, E., & Segaert, K. (2019). Memory encoding of syntactic information involves domain-general attentional resources. Evidence from dual-task studies. Quarterly Journal of Experimental Psychology, 72(6), 1285-1296. doi:10.1177/1747021818801249.

    Abstract

    We investigate the type of attention (domain-general or language-specific) used during
    syntactic processing. We focus on syntactic priming: In this task, participants listen to a
    sentence that describes a picture (prime sentence), followed by a picture the participants need
    to describe (target sentence). We measure the proportion of times participants use the
    syntactic structure they heard in the prime sentence to describe the current target sentence as a
    measure of syntactic processing. Participants simultaneously conducted a motion-object
    tracking (MOT) task, a task commonly used to tax domain-general attentional resources. We
    manipulated the number of objects the participant had to track; we thus measured
    participants’ ability to process syntax while their attention is not-, slightly-, or overly-taxed.
    Performance in the MOT task was significantly worse when conducted as a dual-task
    compared to as a single task. We observed an inverted U-shaped curve on priming magnitude
    when conducting the MOT task concurrently with prime sentences (i.e., memory encoding),
    but no effect when conducted with target sentences (i.e., memory retrieval). Our results
    illustrate how, during the encoding of syntactic information, domain-general attention
    differentially affects syntactic processing, whereas during the retrieval of syntactic
    information domain-general attention does not influence syntactic processing
  • Hoedemaker, R. S., & Meyer, A. S. (2019). Planning and coordination of utterances in a joint naming task. Journal of Experimental Psychology: Learning, Memory, and Cognition, 45(4), 732-752. doi:10.1037/xlm0000603.

    Abstract

    Dialogue requires speakers to coordinate. According to the model of dialogue as joint action, interlocutors achieve this coordination by corepresenting their own and each other’s task share in a functionally equivalent manner. In two experiments, we investigated this corepresentation account using an interactive joint naming task in which pairs of participants took turns naming sets of objects on a shared display. Speaker A named the first, or the first and third object, and Speaker B named the second object. In control conditions, Speaker A named one, two, or all three objects and Speaker B remained silent. We recorded the timing of the speakers’ utterances and Speaker A’s eye movements. Interturn pause durations indicated that the speakers effectively coordinated their utterances in time. Speaker A’s speech onset latencies depended on the number of objects they named, but were unaffected by Speaker B’s naming task. This suggests speakers were not fully incorporating their partner’s task into their own speech planning. Moreover, Speaker A’s eye movements indicated that they were much less likely to attend to objects their partner named than to objects they named themselves. When speakers did inspect their partner’s objects, viewing times were too short to suggest that speakers were retrieving these object names as if they were planning to name the objects themselves. These results indicate that speakers prioritized planning their own responses over attending to their interlocutor’s task and suggest that effective coordination can be achieved without full corepresentation of the partner’s task.
  • Holler, J., & Levinson, S. C. (2019). Multimodal language processing in human communication. Trends in Cognitive Sciences, 23(8), 639-652. doi:10.1016/j.tics.2019.05.006.

    Abstract

    Multiple layers of visual (and vocal) signals, plus their different onsets and offsets, represent a significant semantic and temporal binding problem during face-to-face conversation.
    Despite this complex unification process, multimodal messages appear to be processed faster than unimodal messages.

    Multimodal gestalt recognition and multilevel prediction are proposed to play a crucial role in facilitating multimodal language processing.

    The basis of the processing mechanisms involved in multimodal language comprehension is hypothesized to be domain general, coopted for communication, and refined with domain-specific characteristics.
    A new, situated framework for understanding human language processing is called for that takes into consideration the multilayered, multimodal nature of language and its production and comprehension in conversational interaction requiring fast processing.
  • Holler, J., & Beattie, G. (2004). The interaction of iconic gesture and speech. In A. Cammurri, & G. Volpe (Eds.), Lecture Notes in Computer Science, 5th International Gesture Workshop, Genova, Italy, 2003; Selected Revised Papers (pp. 63-69). Heidelberg: Springer Verlag.
  • Hoogman, M., Rijpkema, M., Janss, L., Brunner, H., Fernandez, G., Buitelaar, J., Franke, B., & Arias-Vásquez, A. (2012). Current self-reported symptoms of attention deficit/hyperactivity disorder are associated with total brain volume in healthy adults. PLoS One, 7(2), e31273. doi:10.1371/journal.pone.0031273.

    Abstract

    Background Reduced total brain volume is a consistent finding in children with Attention Deficit/Hyperactivity Disorder (ADHD). In order to get a better understanding of the neurobiology of ADHD, we take the first step in studying the dimensionality of current self-reported adult ADHD symptoms, by looking at its relation with total brain volume. Methodology/Principal Findings In a sample of 652 highly educated adults, the association between total brain volume, assessed with magnetic resonance imaging, and current number of self-reported ADHD symptoms was studied. The results showed an association between these self-reported ADHD symptoms and total brain volume. Post-hoc analysis revealed that the symptom domain of inattention had the strongest association with total brain volume. In addition, the threshold for impairment coincides with the threshold for brain volume reduction. Conclusions/Significance This finding improves our understanding of the biological substrates of self-reported ADHD symptoms, and suggests total brain volume as a target intermediate phenotype for future gene-finding in ADHD.
  • Horemans, I., & Schiller, N. O. (2004). Form-priming effects in nonword naming. Brain and Language, 90(1-3), 465-469. doi:10.1016/S0093-934X(03)00457-7.

    Abstract

    Form-priming effects from sublexical (syllabic or segmental) primes in masked priming can be accounted for in two ways. One is the sublexical pre-activation view according to which segments are pre-activated by the prime, and at the time the form-related target is to be produced, retrieval/assembly of those pre-activated segments is faster compared to an unrelated situation. However, it has also been argued that form-priming effects from sublexical primes might be due to lexical pre-activation. When the sublexical prime is presented, it activates all form-related words (i.e., cohorts) in the lexicon, necessarily including the form-related target, which—as a consequence—is produced faster than in the unrelated case. Note, however, that this lexical pre-activation account makes previous pre-lexical activation of segments necessary. This study reports a nonword naming experiment to investigate whether or not sublexical pre-activation is involved in masked form priming with sublexical primes. The results demonstrated a priming effect suggesting a nonlexical effect. However, this does not exclude an additional lexical component in form priming.
  • Hörpel, S. G., & Firzlaff, U. (2019). Processing of fast amplitude modulations in bat auditory cortex matches communication call-specific sound features. Journal of Neurophysiology, 121(4), 1501-1512. doi:10.1152/jn.00748.2018.
  • Howe, L., Lawson, D. J., Davies, N. M., St Pourcain, B., Lewis, S. J., Smith, G. D., & Hemani, G. (2019). Genetic evidence for assortative mating on alcohol consumption in the UK Biobank. Nature Communications, 10: 5039. doi:10.1038/s41467-019-12424-x.

    Abstract

    Alcohol use is correlated within spouse-pairs, but it is difficult to disentangle effects of alcohol consumption on mate-selection from social factors or the shared spousal environment. We hypothesised that genetic variants related to alcohol consumption may, via their effect on alcohol behaviour, influence mate selection. Here, we find strong evidence that an individual’s self-reported alcohol consumption and their genotype at rs1229984, a missense variant in ADH1B, are associated with their partner’s self-reported alcohol use. Applying Mendelian randomization, we estimate that a unit increase in an individual’s weekly alcohol consumption increases partner’s alcohol consumption by 0.26 units (95% C.I. 0.15, 0.38; P = 8.20 × 10−6). Furthermore, we find evidence of spousal genotypic concordance for rs1229984, suggesting that spousal concordance for alcohol consumption existed prior to cohabitation. Although the SNP is strongly associated with ancestry, our results suggest some concordance independent of population stratification. Our findings suggest that alcohol behaviour directly influences mate selection.
  • Howe, L. J., Richardson, T. G., Arathimos, R., Alvizi, L., Passos-Bueno, M. R., Stanier, P., Nohr, E., Ludwig, K. U., Mangold, E., Knapp, M., Stergiakouli, E., St Pourcain, B., Smith, G. D., Sandy, J., Relton, C. L., Lewis, S. J., Hemani, G., & Sharp, G. C. (2019). Evidence for DNA methylation mediating genetic liability to non-syndromic cleft lip/palate. Epigenomics, 11(2), 133-145. doi:10.2217/epi-2018-0091.

    Abstract

    Aim: To determine if nonsyndromic cleft lip with or without cleft palate (nsCL/P) genetic risk variants influence liability to nsCL/P through gene regulation pathways, such as those involving DNA methylation. Materials & methods: nsCL/P genetic summary data and methylation data from four studies were used in conjunction with Mendelian randomization and joint likelihood mapping to investigate potential mediation of nsCL/P genetic variants. Results & conclusion: Evidence was found at VAX1 (10q25.3), LOC146880 (17q23.3) and NTN1 (17p13.1), that liability to nsCL/P and variation in DNA methylation might be driven by the same genetic variant, suggesting that genetic variation at these loci may increase liability to nsCL/P by influencing DNA methylation. Follow-up analyses using different tissues and gene expression data provided further insight into possible biological mechanisms.

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  • Hoymann, G. (2004). [Review of the book Botswana: The future of the minority languages ed. by Herman M. Batibo and Birgit Smieja]. Journal of African Languages and Linguistics, 25(2), 171-173. doi:10.1515/jall.2004.25.2.171.
  • Hribar, A., Haun, D. B. M., & Call, J. (2012). Children’s reasoning about spatial relational similarity: The effect of alignment and relational complexity. Journal of Experimental Child Psychology, 111, 490-500. doi:10.1016/j.jecp.2011.11.004.

    Abstract

    We investigated 4- and 5-year-old children’s mapping strategies in a spatial task. Children were required to find a picture in an array of three identical cups after observing another picture being hidden in another array of three cups. The arrays were either aligned one behind the other in two rows or placed side by side forming one line. Moreover, children were rewarded for two different mapping strategies. Half of the children needed to choose a cup that held the same relative position as the rewarded cup in the other array; they needed to map left–left, middle–middle, and right–right cups together (aligned mapping), which required encoding and mapping of two relations (e.g., the cup left of the middle cup and left of the right cup). The other half needed to map together the cups that held the same relation to the table’s spatial features—the cups at the edges, the middle cups, and the cups in the middle of the table (landmark mapping)—which required encoding and mapping of one relation (e.g., the cup at the table’s edge). Results showed that children’s success was constellation dependent; performance was higher when the arrays were aligned one behind the other in two rows than when they were placed side by side. Furthermore, children showed a preference for landmark mapping over aligned mapping.
  • Hubbard, R. J., Rommers, J., Jacobs, C. L., & Federmeier, K. D. (2019). Downstream behavioral and electrophysiological consequences of word prediction on recognition memory. Frontiers in Human Neuroscience, 13: 291. doi:10.3389/fnhum.2019.00291.

    Abstract

    When people process language, they can use context to predict upcoming information,
    influencing processing and comprehension as seen in both behavioral and neural
    measures. Although numerous studies have shown immediate facilitative effects
    of confirmed predictions, the downstream consequences of prediction have been
    less explored. In the current study, we examined those consequences by probing
    participants’ recognition memory for words after they read sets of sentences.
    Participants read strongly and weakly constraining sentences with expected or
    unexpected endings (“I added my name to the list/basket”), and later were tested on
    their memory for the sentence endings while EEG was recorded. Critically, the memory
    test contained words that were predictable (“list”) but were never read (participants
    saw “basket”). Behaviorally, participants showed successful discrimination between old
    and new items, but false alarmed to the expected-item lures more often than to new
    items, showing that predicted words or concepts can linger, even when predictions
    are disconfirmed. Although false alarm rates did not differ by constraint, event-related
    potentials (ERPs) differed between false alarms to strongly and weakly predictable words.
    Additionally, previously unexpected (compared to previously expected) endings that
    appeared on the memory test elicited larger N1 and LPC amplitudes, suggesting greater
    attention and episodic recollection. In contrast, highly predictable sentence endings that
    had been read elicited reduced LPC amplitudes during the memory test. Thus, prediction
    can facilitate processing in the moment, but can also lead to false memory and reduced
    recollection for predictable information.
  • Hubers, F., Cucchiarini, C., Strik, H., & Dijkstra, T. (2019). Normative data of Dutch idiomatic expressions: Subjective judgments you can bank on. Frontiers in Psychology, 10: 1075. doi:10.3389/fpsyg.2019.01075.

    Abstract

    The processing of idiomatic expressions is a topical issue in empirical research. Various factors have been found to influence idiom processing, such as idiom familiarity and idiom transparency. Information on these variables is usually obtained through norming studies. Studies investigating the effect of various properties on idiom processing have led to ambiguous results. This may be due to the variability of operationalizations of the idiom properties across norming studies, which in turn may affect the reliability of the subjective judgements. However, not all studies that collected normative data on idiomatic expressions investigated their reliability, and studies that did address the reliability of subjective ratings used various measures and produced mixed results. In this study, we investigated the reliability of subjective judgements, the relation between subjective and objective idiom frequency, and the impact of these dimensions on the participants’ idiom knowledge by collecting normative data of five subjective idiom properties (Frequency of Exposure, Meaning Familiarity, Frequency of Usage, Transparency, and Imageability) from 390 native speakers and objective corpus frequency for 374 Dutch idiomatic expressions. For reliability, we compared measures calculated in previous studies, with the D-coefficient, a metric taken from Generalizability Theory. High reliability was found for all subjective dimensions. One reliability metric, Krippendorff’s alpha, generally produced lower values, while similar values were obtained for three other measures (Cronbach’s alpha, Intraclass Correlation Coefficient, and the D-coefficient). Advantages of the D-coefficient are that it can be applied to unbalanced research designs, and to estimate the minimum number of raters required to obtain reliable ratings. Slightly higher coefficients were observed for so-called experience-based dimensions (Frequency of Exposure, Meaning Familiarity, and Frequency of Usage) than for content-based dimensions (Transparency and Imageability). In addition, fewer raters were required to obtain reliable ratings for the experience-based dimensions. Subjective and objective frequency appeared to be poorly correlated, while all subjective idiom properties and objective frequency turned out to affect idiom knowledge. Meaning Familiarity, Subjective and Objective Frequency of Exposure, Frequency of Usage, and Transparency positively contributed to idiom knowledge, while a negative effect was found for Imageability. We discuss these relationships in more detail, and give methodological recommendations with respect to the procedures and the measure to calculate reliability.

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    supplementary material
  • Huettig, F., & Altmann, G. T. M. (2004). The online processing of ambiguous and unambiguous words in context: Evidence from head-mounted eye-tracking. In M. Carreiras, & C. Clifton (Eds.), The on-line study of sentence comprehension: Eyetracking, ERP and beyond (pp. 187-207). New York: Psychology Press.
  • Huettig, F., & Hartsuiker, R. J. (2008). When you name the pizza you look at the coin and the bread: Eye movements reveal semantic activation during word production. Memory & Cognition, 36(2), 341-360. doi:10.3758/MC.36.2.341.

    Abstract

    Two eyetracking experiments tested for activation of category coordinate and perceptually related concepts when speakers prepare the name of an object. Speakers saw four visual objects in a 2 × 2 array and identified and named a target picture on the basis of either category (e.g., "What is the name of the musical instrument?") or visual-form (e.g., "What is the name of the circular object?") instructions. There were more fixations on visual-form competitors and category coordinate competitors than on unrelated objects during name preparation, but the increased overt attention did not affect naming latencies. The data demonstrate that eye movements are a sensitive measure of the overlap between the conceptual (including visual-form) information that is accessed in preparation for word production and the conceptual knowledge associated with visual objects. Furthermore, these results suggest that semantic activation of competitor concepts does not necessarily affect lexical selection, contrary to the predictions of lexical-selection-by-competition accounts (e.g., Levelt, Roelofs, & Meyer, 1999).
  • Huettig, F., & Pickering, M. (2019). Literacy advantages beyond reading: Prediction of spoken language. Trends in Cognitive Sciences, 23(6), 464-475. doi:10.1016/j.tics.2019.03.008.

    Abstract

    Literacy has many obvious benefits—it exposes the reader to a wealth of new information and enhances syntactic knowledge. However, we argue that literacy has an additional, often overlooked, benefit: it enhances people’s ability to predict spoken language thereby aiding comprehension. Readers are under pressure to process information more quickly than listeners, and reading provides excellent conditions, in particular a stable environment, for training the predictive system. It also leads to increased awareness of words as linguistic units, and more fine-grained phonological and additional orthographic representations, which sharpen lexical representations and facilitate predicted representations to be retrieved. Thus, reading trains core processes and representations involved in language prediction that are common to both reading and listening.
  • Huettig, F., & Guerra, E. (2019). Effects of speech rate, preview time of visual context, and participant instructions reveal strong limits on prediction in language processing. Brain Research, 1706, 196-208. doi:10.1016/j.brainres.2018.11.013.

    Abstract

    There is a consensus among language researchers that people can predict upcoming language. But do people always predict when comprehending language? Notions that “brains … are essentially prediction machines” certainly suggest so. In three eye-tracking experiments we tested this view. Participants listened to simple Dutch sentences (‘Look at the displayed bicycle’) while viewing four objects (a target, e.g. a bicycle, and three unrelated distractors). We used the identical visual stimuli and the same spoken sentences but varied speech rates, preview time, and participant instructions. Target nouns were preceded by definite gender-marked determiners, which allowed participants to predict the target object because only the targets but not the distractors agreed in gender with the determiner. In Experiment 1, participants had four seconds preview and sentences were presented either in a slow or a normal speech rate. Participants predicted the targets as soon as they heard the determiner in both conditions. Experiment 2 was identical except that participants were given only a one second preview. Participants predicted the targets only in the slow speech condition. Experiment 3 was identical to Experiment 2 except that participants were explicitly told to predict. This led only to a small prediction effect in the normal speech condition. Thus, a normal speech rate only afforded prediction if participants had an extensive preview. Even the explicit instruction to predict the target resulted in only a small anticipation effect with a normal speech rate and a short preview. These findings are problematic for theoretical proposals that assume that prediction pervades cognition.
  • Huettig, F., Mishra, R. K., & Olivers, C. N. (2012). Mechanisms and representations of language-mediated visual attention. Frontiers in Psychology, 2, 394. doi:10.3389/fpsyg.2011.00394.

    Abstract

    The experimental investigation of language-mediated visual attention is a promising way to study the interaction of the cognitive systems involved in language, vision, attention, and memory. Here we highlight four challenges for a mechanistic account of this oculomotor behavior: the levels of representation at which language-derived and vision-derived representations are integrated; attentional mechanisms; types of memory; and the degree of individual and group differences. Central points in our discussion are (a) the possibility that local microcircuitries involving feedforward and feedback loops instantiate a common representational substrate of linguistic and non-linguistic information and attention; and (b) that an explicit working memory may be central to explaining interactions between language and visual attention. We conclude that a synthesis of further experimental evidence from a variety of fields of inquiry and the testing of distinct, non-student, participant populations will prove to be critical.
  • Huisman, J. L. A., Majid, A., & Van Hout, R. (2019). The geographical configuration of a language area influences linguistic diversity. PLoS One, 14(6): e0217363. doi:10.1371/journal.pone.0217363.

    Abstract

    Like the transfer of genetic variation through gene flow, language changes constantly as a result of its use in human interaction. Contact between speakers is most likely to happen when they are close in space, time, and social setting. Here, we investigated the role of geographical configuration in this process by studying linguistic diversity in Japan, which comprises a large connected mainland (less isolation, more potential contact) and smaller island clusters of the Ryukyuan archipelago (more isolation, less potential contact). We quantified linguistic diversity using dialectometric methods, and performed regression analyses to assess the extent to which distance in space and time predict contemporary linguistic diversity. We found that language diversity in general increases as geographic distance increases and as time passes—as with biodiversity. Moreover, we found that (I) for mainland languages, linguistic diversity is most strongly related to geographic distance—a so-called isolation-by-distance pattern, and that (II) for island languages, linguistic diversity reflects the time since varieties separated and diverged—an isolation-by-colonisation pattern. Together, these results confirm previous findings that (linguistic) diversity is shaped by distance, but also goes beyond this by demonstrating the critical role of geographic configuration.
  • Hulten, A., Schoffelen, J.-M., Udden, J., Lam, N. H. L., & Hagoort, P. (2019). How the brain makes sense beyond the processing of single words – An MEG study. NeuroImage, 186, 586-594. doi:10.1016/j.neuroimage.2018.11.035.

    Abstract

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

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    1-s2.0-S1053811918321165-mmc1.pdf
  • Hunley, K., Dunn, M., Lindström, E., Reesink, G., Terrill, A., Healy, M. E., Koki, G., Friedlaender, F. R., & Friedlaender, J. S. (2008). Genetic and linguistic coevolution in Northern Island Melanesia. PLoS Genetics, 4(10): e1000239. doi:10.1371/journal.pgen.1000239.

    Abstract

    Recent studies have detailed a remarkable degree of genetic and linguistic diversity in Northern Island Melanesia. Here we utilize that diversity to examine two models of genetic and linguistic coevolution. The first model predicts that genetic and linguistic correspondences formed following population splits and isolation at the time of early range expansions into the region. The second is analogous to the genetic model of isolation by distance, and it predicts that genetic and linguistic correspondences formed through continuing genetic and linguistic exchange between neighboring populations. We tested the predictions of the two models by comparing observed and simulated patterns of genetic variation, genetic and linguistic trees, and matrices of genetic, linguistic, and geographic distances. The data consist of 751 autosomal microsatellites and 108 structural linguistic features collected from 33 Northern Island Melanesian populations. The results of the tests indicate that linguistic and genetic exchange have erased any evidence of a splitting and isolation process that might have occurred early in the settlement history of the region. The correlation patterns are also inconsistent with the predictions of the isolation by distance coevolutionary process in the larger Northern Island Melanesian region, but there is strong evidence for the process in the rugged interior of the largest island in the region (New Britain). There we found some of the strongest recorded correlations between genetic, linguistic, and geographic distances. We also found that, throughout the region, linguistic features have generally been less likely to diffuse across population boundaries than genes. The results from our study, based on exceptionally fine-grained data, show that local genetic and linguistic exchange are likely to obscure evidence of the early history of a region, and that language barriers do not particularly hinder genetic exchange. In contrast, global patterns may emphasize more ancient demographic events, including population splits associated with the early colonization of major world regions.
  • Hustá, C., Dalmaijer, E., Belopolsky, A., & Mathôt, S. (2019). The pupillary light response reflects visual working memory content. Journal of Experimental Psychology: Human Perception and Performance, 45(11), 1522-1528. doi:10.1037/xhp0000689.

    Abstract

    Recent studies have shown that the pupillary light response (PLR) is modulated by higher cognitive functions, presumably through activity in visual sensory brain areas. Here we use the PLR to test the involvement of sensory areas in visual working memory (VWM). In two experiments, participants memorized either bright or dark stimuli. We found that pupils were smaller when a prestimulus cue indicated that a bright stimulus should be memorized; this reflects a covert shift of attention during encoding of items into VWM. Crucially, we obtained the same result with a poststimulus cue, which shows that internal shifts of attention within VWM affect pupil size as well. Strikingly, the effect of VWM content on pupil size was most pronounced immediately after the poststimulus cue, and then dissipated. This suggests that a shift of attention within VWM momentarily activates an "active" memory representation, but that this representation quickly transforms into a "hidden" state that does not rely on sensory areas.

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    Supplementary_xhp0000689.docx
  • Iacozza, S., Meyer, A. S., & Lev-Ari, S. (2019). How in-group bias influences source memory for words learned from in-group and out-group speakers. Frontiers in Human Neuroscience, 13: 308. doi:10.3389/fnhum.2019.00308.

    Abstract

    Individuals rapidly extract information about others’ social identity, including whether or not they belong to their in-group. Group membership status has been shown to affect how attentively people encode information conveyed by those others. These findings are highly relevant for the field of psycholinguistics where there exists an open debate on how words are represented in the mental lexicon and how abstract or context-specific these representations are. Here, we used a novel word learning paradigm to test our proposal that the group membership status of speakers also affects how speaker-specific representations of novel words are. Participants learned new words from speakers who either attended their own university (in-group speakers) or did not (out-group speakers) and performed a task to measure their individual in-group bias. Then, their source memory of the new words was tested in a recognition test to probe the speaker-specific content of the novel lexical representations and assess how it related to individual in-group biases. We found that speaker group membership and participants’ in-group bias affected participants’ decision biases. The stronger the in-group bias, the more cautious participants were in their decisions. This was particularly applied to in-group related decisions. These findings indicate that social biases can influence recognition threshold. Taking a broader scope, defining how information is represented is a topic of great overlap between the fields of memory and psycholinguistics. Nevertheless, researchers from these fields tend to stay within the theoretical and methodological borders of their own field, missing the chance to deepen their understanding of phenomena that are of common interest. Here we show how methodologies developed in the memory field can be implemented in language research to shed light on an important theoretical issue that relates to the composition of lexical representations.

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  • Ibarretxe-Antuñano, I. (2012). Placement and removal events in Basque and Spanish. In A. Kopecka, & B. Narasimhan (Eds.), Events of putting and taking: A crosslinguistic perspective (pp. 123-144). Amsterdam: Benjamins.

    Abstract

    This paper examines how placement and removal events are lexicalised and conceptualised in Basque and Peninsular Spanish. After a brief description of the main linguistic devices employed for the coding of these types of events, the paper discusses how speakers of the two languages choose to talk about these events. Finally, the paper focuses on two aspects that seem to be crucial in the description of these events (1) the role of force dynamics: both languages distinguish between different degrees of force, causality, and intentionality, and (2) the influence of the verb-framed lexicalisation pattern. Data come from six Basque and ten Peninsular Spanish native speakers.
  • IJzerman, H., Gallucci, M., Pouw, W., Weiβgerber, S. C., Van Doesum, N. J., & Williams, K. D. (2012). Cold-blooded loneliness: Social exclusion leads to lower skin temperatures. Acta Psychologica, 140(3), 283-288. doi:10.1016/j.actpsy.2012.05.002.

    Abstract

    Being ostracized or excluded, even briefly and by strangers, is painful and threatens fundamental needs. Recent work by Zhong and Leonardelli (2008) found that excluded individuals perceive the room as cooler and that they desire warmer drinks. A perspective that many rely on in embodiment is the theoretical idea that people use metaphorical associations to understand social exclusion (see Landau, Meier, & Keefer, 2010). We suggest that people feel colder because they are colder. The results strongly support the idea that more complex metaphorical understandings of social relations are scaffolded onto literal changes in bodily temperature: Being excluded in an online ball tossing game leads to lower finger temperatures (Study 1), while the negative affect typically experienced after such social exclusion is alleviated after holding a cup of warm tea (Study 2). The authors discuss further implications for the interaction between body and social relations specifically, and for basic and cognitive systems in general.
  • Ikram, M. A., Fornage, M., Smith, A. V., Seshadri, S., Schmidt, R., Debette, S., Vrooman, H. A., Sigurdsson, S., Ropele, S., Taal, H. R., Mook-Kanamori, D. O., Coker, L. H., Longstreth, W. T., Niessen, W. J., DeStefano, A. L., Beiser, A., Zijdenbos, A. P., Struchalin, M., Jack, C. R., Rivadeneira, F. and 37 moreIkram, M. A., Fornage, M., Smith, A. V., Seshadri, S., Schmidt, R., Debette, S., Vrooman, H. A., Sigurdsson, S., Ropele, S., Taal, H. R., Mook-Kanamori, D. O., Coker, L. H., Longstreth, W. T., Niessen, W. J., DeStefano, A. L., Beiser, A., Zijdenbos, A. P., Struchalin, M., Jack, C. R., Rivadeneira, F., Uitterlinden, A. G., Knopman, D. S., Hartikainen, A.-L., Pennell, C. E., Thiering, E., Steegers, E. A. P., Hakonarson, H., Heinrich, J., Palmer, L. J., Jarvelin, M.-R., McCarthy, M. I., Grant, S. F. A., St Pourcain, B., Timpson, N. J., Smith, G. D., Sovio, U., Nalls, M. A., Au, R., Hofman, A., Gudnason, H., van der Lugt, A., Harris, T. B., Meeks, W. M., Vernooij, M. W., van Buchem, M. A., Catellier, D., Jaddoe, V. W. V., Gudnason, V., Windham, B. G., Wolf, P. A., van Duijn, C. M., Mosley, T. H., Schmidt, H., Launer, L. J., Breteler, M. M. B., DeCarli, C., Consortiumthe Cohorts for Heart and Aging Research in Genomic Epidemiology (CHARGE) Consortium, & Early Growth Genetics (EGG) Consortium (2012). Common variants at 6q22 and 17q21 are associated with intracranial volume. Nature Genetics, 44(5), 539-544. doi:10.1038/ng.2245.

    Abstract

    During aging, intracranial volume remains unchanged and represents maximally attained brain size, while various interacting biological phenomena lead to brain volume loss. Consequently, intracranial volume and brain volume in late life reflect different genetic influences. Our genome-wide association study (GWAS) in 8,175 community-dwelling elderly persons did not reveal any associations at genome-wide significance (P < 5 × 10(-8)) for brain volume. In contrast, intracranial volume was significantly associated with two loci: rs4273712 (P = 3.4 × 10(-11)), a known height-associated locus on chromosome 6q22, and rs9915547 (P = 1.5 × 10(-12)), localized to the inversion on chromosome 17q21. We replicated the associations of these loci with intracranial volume in a separate sample of 1,752 elderly persons (P = 1.1 × 10(-3) for 6q22 and 1.2 × 10(-3) for 17q21). Furthermore, we also found suggestive associations of the 17q21 locus with head circumference in 10,768 children (mean age of 14.5 months). Our data identify two loci associated with head size, with the inversion at 17q21 also likely to be involved in attaining maximal brain size.
  • Indefrey, P., & Cutler, A. (2004). Prelexical and lexical processing in listening. In M. Gazzaniga (Ed.), The cognitive neurosciences III. (pp. 759-774). Cambridge, MA: MIT Press.

    Abstract

    This paper presents a meta-analysis of hemodynamic studies on passive auditory language processing. We assess the overlap of hemodynamic activation areas and activation maxima reported in experiments involving the presentation of sentences, words, pseudowords, or sublexical or non-linguistic auditory stimuli. Areas that have been reliably replicated are identified. The results of the meta-analysis are compared to electrophysiological, magnetencephalic (MEG), and clinical findings. It is concluded that auditory language input is processed in a left posterior frontal and bilateral temporal cortical network. Within this network, no processing leve l is related to a single cortical area. The temporal lobes seem to differ with respect to their involvement in post-lexical processing, in that the left temporal lobe has greater involvement than the right, and also in the degree of anatomical specialization for phonological, lexical, and sentence -level processing, with greater overlap on the right contrasting with a higher degree of differentiation on the left.
  • Indefrey, P., & Levelt, W. J. M. (2004). The spatial and temporal signatures of word production components. Cognition, 92(1-2), 101-144. doi:10.1016/j.cognition.2002.06.001.

    Abstract

    This paper presents the results of a comprehensive meta-analysis of the relevant imaging literature on word production (82 experiments). In addition to the spatial overlap of activated regions, we also analyzed the available data on the time course of activations. The analysis specified regions and time windows of activation for the core processes of word production: lexical selection, phonological code retrieval, syllabification, and phonetic/articulatory preparation. A comparison of the word production results with studies on auditory word/non-word perception and reading showed that the time course of activations in word production is, on the whole, compatible with the temporal constraints that perception processes impose on the production processes they affect in picture/word interference paradigms.
  • Indefrey, P. (1998). De neurale architectuur van taal: Welke hersengebieden zijn betrokken bij het spreken. Neuropraxis, 2(6), 230-237.
  • Indefrey, P. (2004). Hirnaktivierungen bei syntaktischer Sprachverarbeitung: Eine Meta-Analyse. In H. Müller, & G. Rickheit (Eds.), Neurokognition der Sprache (pp. 31-50). Tübingen: Stauffenburg.
  • Indefrey, P., Hellwig, F. M., Herzog, H., Seitz, R. J., & Hagoort, P. (2004). Neural responses to the production and comprehension of syntax in identical utterances. Brain and Language, 89(2), 312-319. doi:10.1016/S0093-934X(03)00352-3.

    Abstract

    Following up on an earlier positron emission tomography (PET) experiment (Indefrey et al., 2001), we used a scene description paradigm to investigate whether a posterior inferior frontal region subserving syntactic encoding for speaking is also involved in syntactic parsing during listening. In the language production part of the experiment, subjects described visually presented scenes
    using either sentences, sequences of noun phrases, or sequences of syntactically unrelated words. In the language comprehension part of the experiment, subjects were auditorily presented with the same kinds of utterances and judged whether they matched the visual scenes. We were able to replicate the previous finding of a region in caudal Broca s area that is sensitive to the complexity of
    syntactic encoding in language production. In language comprehension, no hemodynamic activation differences due to syntactic complexity were found. Given that correct performance in the judgment task did not require syntactic processing of the auditory stimuli, the results suggest that the degree to which listeners recruit syntactic processing resources in language comprehension may be a function of the syntactic demands of the task or the stimulus material.
  • Indefrey, P. (2012). Hemodynamic studies of syntactic processing. In M. Faust (Ed.), Handbook of the neuropsychology of language. Volume 1: Language processing in the brain: Basic science (pp. 209-228). Malden, MA: Wiley-Blackwell.
  • Indefrey, P., Gruber, O., Brown, C. M., Hagoort, P., Posse, S., & Kleinschmidt, A. (1998). Lexicality and not syllable frequency determine lateralized premotor activation during the pronunciation of word-like stimuli: An fMRI study. NeuroImage, 7, S4.
  • Ioana, M., Ferwerda, B., Farjadian, S., Ioana, L., Ghaderi, A., Oosting, M., Joosten, L. A., Van der Meer, J. W., Romeo, G., Luiselli, D., Dediu, D., & Netea, M. G. (2012). High variability of TLR4 gene in different ethnic groups of Iran. Innate Immunity, 18, 492-502. doi:10.1177/1753425911423043.

    Abstract

    Infectious diseases exert a constant evolutionary pressure on the innate immunity genes. TLR4, an important member of the Toll-like receptors family, specifically recognizes conserved structures of various infectious pathogens. Two functional TLR4 polymorphisms, Asp299Gly and Thr399Ile, modulate innate host defense against infections, and their prevalence between various populations has been proposed to be influenced by local infectious pressures. If this assumption is true, strong local infectious pressures would lead to a homogeneous pattern of these ancient TLR4 polymorphisms in geographically close populations, while a weak selection or genetic drift may result in a diverse pattern. We evaluated TLR4 polymorphisms in 15 ethnic groups of Iran, to assess whether infections exerted selective pressures on different haplotypes containing these variants. The Iranian subpopulations displayed a heterogeneous pattern of TLR4 polymorphisms, comprising various percentages of Asp299Gly and Thr399Ile alone or in combination. The Iranian sample as a whole showed an intermediate mixed pattern when compared with commonly found patterns in Africa, Europe, Eastern Asia and Americas. These findings suggest a weak or absent selection pressure on TLR4 polymorphisms in the Middle-East, that does not support the assumption of an important role of these polymorphisms in the host defence against local pathogens.
  • Ioumpa, K., Graham, S. A., Clausner, T., Fisher, S. E., Van Lier, R., & Van Leeuwen, T. M. (2019). Enhanced self-reported affect and prosocial behaviour without differential physiological responses in mirror-sensory synaesthesia. Philosophical Transactions of the Royal Society of London, Series B: Biological Sciences, 374: 20190395. doi:10.1098/rstb.2019.0395.

    Abstract

    Mirror-sensory synaesthetes mirror the pain or touch that they observe in other people on their own bodies. This type of synaesthesia has been associated with enhanced empathy. We investigated whether the enhanced empathy of people with mirror-sensory synesthesia influences the experience of situations involving touch or pain and whether it affects their prosocial decision making. Mirror-sensory synaesthetes (N = 18, all female), verified with a touch-interference paradigm, were compared with a similar number of age-matched control individuals (all female). Participants viewed arousing images depicting pain or touch; we recorded subjective valence and arousal ratings, and physiological responses, hypothesizing more extreme reactions in synaesthetes. The subjective impact of positive and negative images was stronger in synaesthetes than in control participants; the stronger the reported synaesthesia, the more extreme the picture ratings. However, there was no evidence for differential physiological or hormonal responses to arousing pictures. Prosocial decision making was assessed with an economic game assessing altruism, in which participants had to divide money between themselves and a second player. Mirror-sensory synaesthetes donated more money than non-synaesthetes, showing enhanced prosocial behaviour, and also scored higher on the Interpersonal Reactivity Index as a measure of empathy. Our study demonstrates the subjective impact of mirror-sensory synaesthesia and its stimulating influence on prosocial behaviour.

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  • Irizarri van Suchtelen, P. (2012). Dative constructions in the Spanish of heritage speakers in the Netherlands. In Z. Wąsik, & P. P. Chruszczewski (Eds.), Languages in contact 2011 (pp. 103-118). Wrocław: Philological School of Higher Education in Wrocław Publishing.

    Abstract

    Spanish can use dative as well as non-dative strategies to encode Possessors, Human Sources, Interestees (datives of interest) and Experiencers. In Dutch this optionality is virtually absent, restricting dative encoding mainly to the Recipient of a ditransitive. The present study examines whether this may lead to instability of the non-prototypical dative constructions in the Spanish of Dutch-Spanish bilinguals. Elicited data of 12 Chilean heritage informants from the Netherlands were analyzed. Whereas the evidence on the stability of dative Experiencers was not conclusive, the results indicate that the use of prototypical datives, dative External Possessors, dative Human Sources and datives of interest is fairly stable in bilinguals, except for those with limited childhood exposure to Spanish. It is argued that the consistent preference for non-dative strategies of this group was primarily attributable to instability of the dative clitic, which affected all constructions, even the encoding of prototypical indirect objects
  • Isaac, A., Schlobach, S., Matthezing, H., & Zinn, C. (2008). Integrated access to cultural heritage resources through representation and alignment of controlled vocabularies. Library Review, 57(3), 187-199.
  • Ischebeck, A., Indefrey, P., Usui, N., Nose, I., Hellwig, F. M., & Taira, M. (2004). Reading in a regular orthography: An fMRI study investigating the role of visual familiarity. Journal of Cognitive Neuroscience, 16(5), 727-741. doi:10.1162/089892904970708.

    Abstract

    In order to separate the cognitive processes associated with phonological encoding and the use of a visual word form lexicon in reading, it is desirable to compare the processing of words presented in a visually familiar form with words in a visually unfamiliar form. Japanese Kana orthography offers this possibility. Two phonologically equivalent but visually dissimilar syllabaries allow the writing of, for example, foreign loanwords in two ways, only one of which is visually familiar. Familiarly written words, unfamiliarly written words, and pseudowords were presented in both Kana syllabaries (yielding six conditions in total) to participants during an fMRI measurement with a silent articulation task (Experiment 1) and a phonological lexical decision task (Experiment 2) using an event-related design. Consistent over two experimental tasks, the three different stimulus types (familiar, unfamiliar, and pseudoword) were found to activate selectively different brain regions previously associated with phonological encoding and word retrieval or meaning. Compatible with the predictions of the dual-route model for reading, pseudowords and visually unfamiliar words, which have to be read using phonological assembly, caused an increase in brain activity in left inferior frontal regions (BA 44/47), as compared to visually familiar words. Visually familiar and unfamiliar words were found to activate a range of areas associated with lexico-semantic processing more strongly than pseudowords, such as the left and right temporo-parietal region (BA 39/40), a region in the left middle/inferior temporal gyrus (BA 20/21), and the posterior cingulate (BA 31).
  • Ishibashi, M. (2012). The expression of ‘putting’ and ‘taking’ events in Japanese: The asymmetry of Source and Goal revisited. In A. Kopecka, & B. Narasimhan (Eds.), Events of putting and taking: A crosslinguistic perspective (pp. 253-272). Amsterdam: Benjamins.

    Abstract

    This study explores the expression of Source and Goal in describing placement and removal events in adult Japanese. Although placement and removal events a priori represent symmetry regarding the orientation of motion, their (c)overt expressions actually exhibit multiple asymmetries at various structural levels. The results show that the expression of the Source is less frequent than the expression of the Goal, but, if expressed, morphosyntactically more complex, suggesting that ‘taking’ events are more complex than ‘putting’ events in their construal. It is stressed that finer linguistic analysis is necessary before explaining linguistic asymmetries in terms of non-linguistic foundations of spatial language.
  • Iyer, S., Sam, F. S., DiPrimio, N., Preston, G., Verheijen, J., Murthy, K., Parton, Z., Tsang, H., Lao, J., Morava, E., & Perlstein, E. O. (2019). Repurposing the aldose reductase inhibitor and diabetic neuropathy drug epalrestat for the congenital disorder of glycosylation PMM2-CDG. Disease models & mechanisms, 12(11): UNSP dmm040584. doi:10.1242/dmm.040584.

    Abstract

    Phosphomannomutase 2 deficiency, or PMM2-CDG, is the most common congenital disorder of glycosylation and affects over 1000 patients globally. There are no approved drugs that treat the symptoms or root cause of PMM2-CDG. To identify clinically actionable compounds that boost human PMM2 enzyme function, we performed a multispecies drug repurposing screen using a novel worm model of PMM2-CDG, followed by PMM2 enzyme functional studies in PMM2-CDG patient fibroblasts. Drug repurposing candidates from this study, and drug repurposing candidates from a previously published study using yeast models of PMM2-CDG, were tested for their effect on human PMM2 enzyme activity in PMM2-CDG fibroblasts. Of the 20 repurposing candidates discovered in the worm-based phenotypic screen, 12 were plant-based polyphenols. Insights from structure-activity relationships revealed epalrestat, the only antidiabetic aldose reductase inhibitor approved for use in humans, as a first-in-class PMM2 enzyme activator. Epalrestat increased PMM2 enzymatic activity in four PMM2-CDG patient fibroblast lines with genotypes R141H/F119L, R141H/E139K, R141H/N216I and R141H/F183S. PMM2 enzyme activity gains ranged from 30% to 400% over baseline, depending on genotype. Pharmacological inhibition of aldose reductase by epalrestat may shunt glucose from the polyol pathway to glucose-1,6-bisphosphate, which is an endogenous stabilizer and coactivator of PMM2 homodimerization. Epalrestat is a safe, oral and brain penetrant drug that was approved 27 years ago in Japan to treat diabetic neuropathy in geriatric populations. We demonstrate that epalrestat is the first small molecule activator ofPMM2 enzyme activity with the potential to treat peripheral neuropathy and correct the underlying enzyme deficiency in a majority of pediatric and adult PMM2-CDG patients.

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  • Jaeger, E., Leedham, S., Lewis, A., Segditsas, S., Becker, M., Rodenas-Cuadrado, P., Davis, H., Kaur, K., Heinimann, K., Howarth, K., East, J., Taylor, J., Thomas, H., & Tomlinson, I. (2012). Hereditary mixed polyposis syndrome is caused by a 40-kb upstream duplication that leads to increased and ectopic expression of the BMP antagonist GREM1. Nature Genetics, 44, 699-703. doi:10.1038/ng.2263.

    Abstract

    Hereditary mixed polyposis syndrome (HMPS) is characterized by apparent autosomal dominant inheritance of multiple types of colorectal polyp, with colorectal carcinoma occurring in a high proportion of affected individuals. Here, we use genetic mapping, copy-number analysis, exclusion of mutations by high-throughput sequencing, gene expression analysis and functional assays to show that HMPS is caused by a duplication spanning the 3' end of the SCG5 gene and a region upstream of the GREM1 locus. This unusual mutation is associated with increased allele-specific GREM1 expression. Whereas GREM1 is expressed in intestinal subepithelial myofibroblasts in controls, GREM1 is predominantly expressed in the epithelium of the large bowel in individuals with HMPS. The HMPS duplication contains predicted enhancer elements; some of these interact with the GREM1 promoter and can drive gene expression in vitro. Increased GREM1 expression is predicted to cause reduced bone morphogenetic protein (BMP) pathway activity, a mechanism that also underlies tumorigenesis in juvenile polyposis of the large bowel.
  • Janse, E., & Klitsch, J. (2004). Auditieve perceptie bij gezonde sprekers en bij sprekers met verworven taalstoornissen. Afasiologie, 26(1), 2-6.
  • Janse, E. (2012). A non-auditory measure of interference predicts distraction by competing speech in older adults. Aging, Neuropsychology and Cognition, 19, 741-758. doi:10.1080/13825585.2011.652590.

    Abstract

    In this study, older adults monitored for pre-assigned target sounds in a target talker's speech in a quiet (no noise) condition and in a condition with competing-talker noise. The question was to which extent the impact of the competing-talker noise on performance could be predicted from individual hearing loss and from a cognitive measure of inhibitory abilities, i.e., a measure of Stroop interference. The results showed that the non-auditory measure of Stroop interference predicted the impact of distraction on performance, over and above the effect of hearing loss. This suggests that individual differences in inhibitory abilities among older adults relate to susceptibility to distracting speech.
  • Janse, I., Bok, J., Hamidjaja, R. A., Hodemaekers, H. M., & van Rotterdam, B. J. (2012). Development and comparison of two assay formats for parallel detection of four biothreat pathogens by using suspension microarrays. PLoS One, 7(2), e31958. doi:10.1371/journal.pone.0031958.

    Abstract

    Microarrays provide a powerful analytical tool for the simultaneous detection of multiple pathogens. We developed diagnostic suspension microarrays for sensitive and specific detection of the biothreat pathogens Bacillus anthracis, Yersinia pestis, Francisella tularensis and Coxiella burnetii. Two assay chemistries for amplification and labeling were developed, one method using direct hybridization and the other using target-specific primer extension, combined with hybridization to universal arrays. Asymmetric PCR products for both assay chemistries were produced by using a multiplex asymmetric PCR amplifying 16 DNA signatures (16-plex). The performances of both assay chemistries were compared and their advantages and disadvantages are discussed. The developed microarrays detected multiple signature sequences and an internal control which made it possible to confidently identify the targeted pathogens and assess their virulence potential. The microarrays were highly specific and detected various strains of the targeted pathogens. Detection limits for the different pathogen signatures were similar or slightly higher compared to real-time PCR. Probit analysis showed that even a few genomic copies could be detected with 95% confidence. The microarrays detected DNA from different pathogens mixed in different ratios and from spiked or naturally contaminated samples. The assays that were developed have a potential for application in surveillance and diagnostics. © 2012 Janse et al.
  • Janse, E. (2008). Spoken-word processing in aphasia: Effects of item overlap and item repetition. Brain and Language, 105, 185-198. doi:10.1016/j.bandl.2007.10.002.

    Abstract

    Two studies were carried out to investigate the effects of presentation of primes showing partial (word-initial) or full overlap on processing of spoken target words. The first study investigated whether time compression would interfere with lexical processing so as to elicit aphasic-like performance in non-brain-damaged subjects. The second study was designed to compare effects of item overlap and item repetition in aphasic patients of different diagnostic types. Time compression did not interfere with lexical deactivation for the non-brain-damaged subjects. Furthermore, all aphasic patients showed immediate inhibition of co-activated candidates. These combined results show that deactivation is a fast process. Repetition effects, however, seem to arise only at the longer term in aphasic patients. Importantly, poor performance on diagnostic verbal STM tasks was shown to be related to lexical decision performance in both overlap and repetition conditions, which suggests a common underlying deficit.
  • Janse, E., & Adank, P. (2012). Predicting foreign-accent adaptation in older adults. Quarterly Journal of Experimental Psychology, 65, 1563-1585. doi:10.1080/17470218.2012.658822.

    Abstract

    We investigated comprehension of and adaptation to speech in an unfamiliar accent in older adults. Participants performed a speeded sentence verification task for accented sentences: one group upon auditory-only presentation, and the other group upon audiovisual presentation. Our questions were whether audiovisual presentation would facilitate adaptation to the novel accent, and which cognitive and linguistic measures would predict adaptation. Participants were therefore tested on a range of background tests: hearing acuity, auditory verbal short-term memory, working memory, attention-switching control, selective attention, and vocabulary knowledge. Both auditory-only and audiovisual groups showed improved accuracy and decreasing response times over the course of the experiment, effectively showing accent adaptation. Even though the total amount of improvement was similar for the auditory-only and audiovisual groups, initial rate of adaptation was faster in the audiovisual group. Hearing sensitivity and short-term and working memory measures were associated with efficient processing of the novel accent. Analysis of the relationship between accent comprehension and the background tests revealed furthermore that selective attention and vocabulary size predicted the amount of adaptation over the course of the experiment. These results suggest that vocabulary knowledge and attentional abilities facilitate the attention-shifting strategies proposed to be required for perceptual learning.
  • Janse, E. (2004). Word perception in fast speech: Artificially time-compressed vs. naturally produced fast speech. Speech Communication, 42, 155-173. doi:10.1016/j.specom.2003.07.001.

    Abstract

    Natural fast speech differs from normal-rate speech with respect to its temporal pattern. Previous results showed that word intelligibility of heavily artificially time-compressed speech could not be improved by making its temporal pattern more similar to that of natural fast speech. This might have been due to the extrapolation of timing rules for natural fast speech to rates that are much faster than can be attained by human speakers. The present study investigates whether, at a speech rate that human speakers can attain, artificially time-compressed speech is easier to process if its timing pattern is similar to that of naturally produced fast speech. Our first experiment suggests, however, that word processing speed was slowed down, relative to linear compression. In a second experiment, word processing of artificially time-compressed speech was compared with processing of naturally produced fast speech. Even when naturally produced fast speech is perfectly intelligible, its less careful articulation, combined with the changed timing pattern, slows down processing, relative to linearly time-compressed speech. Furthermore, listeners preferred artificially time-compressed speech over naturally produced fast speech. These results suggest that linearly time-compressed speech has both a temporal and a segmental advantage over natural fast speech.
  • Jansma, B. M., & Schiller, N. O. (2004). Monitoring syllable boundaries during speech production. Brain and Language, 90(1-3), 311-317. doi:10.1016/S0093-934X(03)00443-7.

    Abstract

    This study investigated the encoding of syllable boundary information during speech production in Dutch. Based on Levelt's model of phonological encoding, we hypothesized segments and syllable boundaries to be encoded in an incremental way. In a selfmonitoring experiment, decisions about the syllable affiliation (first or second syllable) of a pre-specified consonant, which was the third phoneme in a word, were required (e.g., ka.No canoe vs. kaN.sel pulpit ; capital letters indicate pivotal consonants, dots mark syllable boundaries). First syllable responses were faster than second syllable responses, indicating the incremental nature of segmental encoding and syllabification during speech production planning. The results of the experiment are discussed in the context of Levelt 's model of phonological encoding.
  • Janssen, D. P., Roelofs, A., & Levelt, W. J. M. (2004). Stem complexity and inflectional encoding in language production. Journal of Psycholinguistic Research, 33(5), 365-381. doi:10.1023/B:JOPR.0000039546.60121.a8.

    Abstract

    Three experiments are reported that examined whether stem complexity plays a role in inflecting polymorphemic words in language production. Experiment 1 showed that preparation effects for words with polymorphemic stems are larger when they are produced among words with constant inflectional structures compared to words with variable inflectional structures and simple stems. This replicates earlier findings for words with monomorphemic stems (Janssen et al., 2002). Experiments 2 and 3 showed that when inflectional structure is held constant, the preparation effects are equally large with simple and compound stems, and with compound and complex adjectival stems. These results indicate that inflectional encoding is blind to the complexity of the stem, which suggests that specific inflectional rather than generic morphological frames guide the generation of inflected forms in speaking words.
  • Janssen, C., Segers, E., McQueen, J. M., & Verhoeven, L. (2019). Comparing effects of instruction on word meaning and word form on early literacy abilities in kindergarten. Early Education and Development, 30(3), 375-399. doi:10.1080/10409289.2018.1547563.

    Abstract

    Research Findings: The present study compared effects of explicit instruction on and practice with the phonological form of words (form-focused instruction) versus explicit instruction on and practice with the meaning of words (meaning-focused instruction). Instruction was given via interactive storybook reading in the kindergarten classroom of children learning Dutch. We asked whether the 2 types of instruction had different effects on vocabulary development and 2 precursors of reading ability—phonological awareness and letter knowledge—and we examined effects on these measures of the ability to learn new words with minimal acoustic-phonetic differences. Learners showed similar receptive target-word vocabulary gain after both types of instruction, but learners who received form-focused vocabulary instruction showed more gain in semantic knowledge of target vocabulary, phonological awareness, and letter knowledge than learners who received meaning-focused vocabulary instruction. Level of ability to learn pairs of words with minimal acoustic-phonetic differences predicted gain in semantic knowledge of target vocabulary and in letter knowledge in the form-focused instruction group only. Practice or Policy: A focus on the form of words during instruction appears to have benefits for young children learning vocabulary.
  • Janssen, R., Moisik, S. R., & Dediu, D. (2019). The effects of larynx height on vowel production are mitigated by the active control of articulators. Journal of Phonetics, 74, 1-17. doi:10.1016/j.wocn.2019.02.002.

    Abstract

    The influence of larynx position on vowel articulation is an important topic in understanding speech production, the present-day distribution of linguistic diversity and the evolution of speech and language in our lineage. We introduce here a realistic computer model of the vocal tract, constructed from actual human MRI data, which can learn, using machine learning techniques, to control the articulators in such a way as to produce speech sounds matching as closely as possible to a given set of target vowels. We systematically control the vertical position of the larynx and we quantify the differences between the target and produced vowels for each such position across multiple replications. We report that, indeed, larynx height does affect the accuracy of reproducing the target vowels and the distinctness of the produced vowel system, that there is a “sweet spot” of larynx positions that are optimal for vowel production, but that nevertheless, even extreme larynx positions do not result in a collapsed or heavily distorted vowel space that would make speech unintelligible. Together with other lines of evidence, our results support the view that the vowel space of human languages is influenced by our larynx position, but that other positions of the larynx may also be fully compatible with speech.

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  • Janzen, G., & Van Turennout, M. (2004). Selective neural representation of objects relevant for navigation. Nature Neuroscience, 7(6), 673-677. doi:10.1038/nn1257.

    Abstract

    As people find their way through their environment, objects at navigationally relevant locations can serve as crucial landmarks. The parahippocampal gyrus has previously been shown to be involved in object and scene recognition. In the present study, we investigated the neural representation of navigationally relevant locations. Healthy human adults viewed a route through a virtual museum with objects placed at intersections (decision points) or at simple turns (non-decision points). Event-related functional magnetic resonance imaging (fMRI) data were acquired during subsequent recognition of the objects in isolation. Neural activity in the parahippocampal gyrus reflected the navigational relevance of an object's location in the museum. Parahippocampal responses were selectively increased for objects that occurred at decision points, independent of attentional demands. This increase occurred for forgotten as well as remembered objects, showing implicit retrieval of navigational information. The automatic storage of relevant object location in the parahippocampal gyrus provides a part of the neural mechanism underlying successful navigation.
  • Janzen, G., Jansen, C., & Van Turennout, M. (2008). Memory consolidation of landmarks in good navigators. Hippocampus, 18, 40-47.

    Abstract

    Landmarks play an important role in successful navigation. To successfully find your way around an environment, navigationally relevant information needs to be stored and become available at later moments in time. Evidence from functional magnetic resonance imaging (fMRI) studies shows that the human parahippocampal gyrus encodes the navigational relevance of landmarks. In the present event-related fMRI experiment, we investigated memory consolidation of navigationally relevant landmarks in the medial temporal lobe after route learning. Sixteen right-handed volunteers viewed two film sequences through a virtual museum with objects placed at locations relevant (decision points) or irrelevant (nondecision points) for navigation. To investigate consolidation effects, one film sequence was seen in the evening before scanning, the other one was seen the following morning, directly before scanning. Event-related fMRI data were acquired during an object recognition task. Participants decided whether they had seen the objects in the previously shown films. After scanning, participants answered standardized questions about their navigational skills, and were divided into groups of good and bad navigators, based on their scores. An effect of memory consolidation was obtained in the hippocampus: Objects that were seen the evening before scanning (remote objects) elicited more activity than objects seen directly before scanning (recent objects). This increase in activity in bilateral hippocampus for remote objects was observed in good navigators only. In addition, a spatial-specific effect of memory consolidation for navigationally relevant objects was observed in the parahippocampal gyrus. Remote decision point objects induced increased activity as compared with recent decision point objects, again in good navigators only. The results provide initial evidence for a connection between memory consolidation and navigational ability that can provide a basis for successful navigation.
  • Janzen, G., Haun, D. B. M., & Levinson, S. C. (2012). Tracking down abstract linguistic meaning: Neural correlates of spatial frame of reference ambiguities in language. PLoS One, 7(2), e30657. doi:10.1371/journal.pone.0030657.

    Abstract

    This functional magnetic resonance imaging (fMRI) study investigates a crucial parameter in spatial description, namely variants in the frame of reference chosen. Two frames of reference are available in European languages for the description of small-scale assemblages, namely the intrinsic (or object-oriented) frame and the relative (or egocentric) frame. We showed participants a sentence such as “the ball is in front of the man”, ambiguous between the two frames, and then a picture of a scene with a ball and a man – participants had to respond by indicating whether the picture did or did not match the sentence. There were two blocks, in which we induced each frame of reference by feedback. Thus for the crucial test items, participants saw exactly the same sentence and the same picture but now from one perspective, now the other. Using this method, we were able to precisely pinpoint the pattern of neural activation associated with each linguistic interpretation of the ambiguity, while holding the perceptual stimuli constant. Increased brain activity in bilateral parahippocampal gyrus was associated with the intrinsic frame of reference whereas increased activity in the right superior frontal gyrus and in the parietal lobe was observed for the relative frame of reference. The study is among the few to show a distinctive pattern of neural activation for an abstract yet specific semantic parameter in language. It shows with special clarity the nature of the neural substrate supporting each frame of spatial reference
  • Jasmin, K., & Casasanto, D. (2012). The QWERTY Effect: How typing shapes the meanings of words. Psychonomic Bulletin & Review, 19, 499-504. doi:10.3758/s13423-012-0229-7.

    Abstract

    The QWERTY keyboard mediates communication for millions of language users. Here, we investigated whether differences in the way words are typed correspond to differences in their meanings. Some words are spelled with more letters on the right side of the keyboard and others with more letters on the left. In three experiments, we tested whether asymmetries in the way people interact with keys on the right and left of the keyboard influence their evaluations of the emotional valence of the words. We found the predicted relationship between emotional valence and QWERTY key position across three languages (English, Spanish, and Dutch). Words with more right-side letters were rated as more positive in valence, on average, than words with more left-side letters: the QWERTY effect. This effect was strongest in new words coined after QWERTY was invented and was also found in pseudowords. Although these data are correlational, the discovery of a similar pattern across languages, which was strongest in neologisms, suggests that the QWERTY keyboard is shaping the meanings of words as people filter language through their fingers. Widespread typing introduces a new mechanism by which semanntic changes in language can arise.
  • Jepma, M., Verdonschot, R. G., Van Steenbergen, H., Rombouts, S. A. R. B., & Nieuwenhuis, S. (2012). Neural mechanisms underlying the induction and relief of perceptual curiosity. Frontiers in Behavioral Neuroscience, 6: 5. doi:10.3389/fnbeh.2012.00005.

    Abstract

    Curiosity is one of the most basic biological drives in both animals and humans, and has been identified as a key motive for learning and discovery. Despite the importance of curiosity and related behaviors, the topic has been largely neglected in human neuroscience; hence little is known about the neurobiological mechanisms underlying curiosity. We used functional magnetic resonance imaging (fMRI) to investigate what happens in our brain during the induction and subsequent relief of perceptual curiosity. Our core findings were that (1) the induction of perceptual curiosity, through the presentation of ambiguous visual input, activated the anterior insula and anterior cingulate cortex (ACC), brain regions sensitive to conflict and arousal; (2) the relief of perceptual curiosity, through visual disambiguation, activated regions of the striatum that have been related to reward processing; and (3) the relief of perceptual curiosity was associated with hippocampal activation and enhanced incidental memory. These findings provide the first demonstration of the neural basis of human perceptual curiosity. Our results provide neurobiological support for a classic psychological theory of curiosity, which holds that curiosity is an aversive condition of increased arousal whose termination is rewarding and facilitates memory.

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