Publications

Displaying 901 - 954 of 954
  • De Vries, M., Ulte, C., Zwitserlood, P., Szymanski, B., & Knecht, S. (2010). Increasing dopamine levels in the brain improves feedback-based procedural learning in healthy participants: An artificial-grammar-learning experiment. Neuropsychologia, 48, 3193-3197. doi:10.1016/j.neuropsychologia.2010.06.024.

    Abstract

    Recently, an increasing number of studies have suggested a role for the basal ganglia and related dopamine inputs in procedural learning, specifically when learning occurs through trial-by-trial feedback (Shohamy, Myers, Kalanithi, & Gluck. (2008). Basal ganglia and dopamine contributions to probabilistic category learning. Neuroscience and Biobehavioral Reviews, 32, 219–236). A necessary relationship has however only been demonstrated in patient studies. In the present study, we show for the first time that increasing dopamine levels in the brain improves the gradual acquisition of complex information in healthy participants. We implemented two artificial-grammar-learning tasks, one with and one without performance feedback. Learning was improved after levodopa intake for the feedback-based learning task only, suggesting that dopamine plays a specific role in trial-by-trial feedback-based learning. This provides promising directions for future studies on dopaminergic modulation of cognitive functioning.
  • De Vries, B., Eising, E., Broos, L. A. M., Koelewijn, S. C., Todorov, B., Frants, R. R., Boer, J. M., Ferraro, M. D., Thoen, P. A. C., & Van Den Maagdenberg, A. (2014). RNA expression profiling in brains of familial hemiplegic migraine type 1 knock-in mice. Cephalalgia, 34(3), 174-182. doi:10.1177/0333102413502736.

    Abstract

    Background Various CACNA1A missense mutations cause familial hemiplegic migraine type 1 (FHM1), a rare monogenic subtype of migraine with aura. FHM1 mutation R192Q is associated with pure hemiplegic migraine, whereas the S218L mutation causes hemiplegic migraine, cerebellar ataxia, seizures, and mild head trauma-induced brain edema. Transgenic knock-in (KI) migraine mouse models were generated that carried either the FHM1 R192Q or the S218L mutation and were shown to exhibit increased CaV2.1 channel activity. Here we investigated their cerebellar and caudal cortical transcriptome. Methods Caudal cortical and cerebellar RNA expression profiles from mutant and wild-type mice were studied using microarrays. Respective brain regions were selected based on their relevance to migraine aura and ataxia. Relevant expression changes were further investigated at RNA and protein level by quantitative polymerase chain reaction (qPCR) and/or immunohistochemistry, respectively. Results Expression differences in the cerebellum were most pronounced in S218L mice. Particularly, tyrosine hydroxylase, a marker of delayed cerebellar maturation, appeared strongly upregulated in S218L cerebella. In contrast, only minimal expression differences were observed in the caudal cortex of either mutant mice strain. Conclusion Despite pronounced consequences of migraine gene mutations at the neurobiological level, changes in cortical RNA expression in FHM1 migraine mice compared to wild-type are modest. In contrast, pronounced RNA expression changes are seen in the cerebellum of S218L mice and may explain their cerebellar ataxia phenotype
  • Wang, L., Hagoort, P., & Yang, Y. (2009). Semantic illusion depends on information structure: ERP evidence. Brain Research, 1282, 50-56. doi:10.1016/j.brainres.2009.05.069.

    Abstract

    Next to propositional content, speakers distribute information in their utterances in such a way that listeners can make a distinction between new (focused) and given (non-focused) information. This is referred to as information structure. We measured event-related potentials (ERPs) to explore the role of information structure in semantic processing. Following different questions in wh-question-answer pairs (e.g. What kind of vegetable did Ming buy for cooking today? /Who bought the vegetables for cooking today?), the answer sentences (e.g., Ming bought eggplant/beef to cook today.) contained a critical word, which was either semantically appropriate (eggplant) or inappropriate (beef), and either focus or non-focus. The results showed a full N400 effect only when the critical words were in focus position. In non-focus position a strongly reduced N400 effect was observed, in line with the well-known semantic illusion effect. The results suggest that information structure facilitates semantic processing by devoting more resources to focused information.
  • Ward, M. E., McMahon, G., St Pourcain, B., Evans, D. M., Rietveld, C. A., Benjamin, D. J., Koellinger, P. D., Cesarini, D., Smith, G. D., Timpson, N. J., & Consortium}, {. S. G. A. (2014). Genetic variation associated with differential educational attainment in adults has anticipated associations with school performance in children. PLoS ONE, 9(7): e100248. doi:10.1371/journal.pone.0100248.

    Abstract

    Genome-wide association study results have yielded evidence for the association of common genetic variants with crude measures of completed educational attainment in adults. Whilst informative, these results do not inform as to the mechanism of these effects or their presence at earlier ages and where educational performance is more routinely and more precisely assessed. Single nucleotide polymorphisms exhibiting genome-wide significant associations with adult educational attainment were combined to derive an unweighted allele score in 5,979 and 6,145 young participants from the Avon Longitudinal Study of Parents and Children with key stage 3 national curriculum test results (SATS results) available at age 13 to 14 years in English and mathematics respectively. Standardised (z-scored) results for English and mathematics showed an expected relationship with sex, with girls exhibiting an advantage over boys in English (0.433 SD (95%CI 0.395, 0.470), p<10-10) with more similar results (though in the opposite direction) in mathematics (0.042 SD (95%CI 0.004, 0.080), p = 0.030). Each additional adult educational attainment increasing allele was associated with 0.041 SD (95%CI 0.020, 0.063), p = 1.79×10-04 and 0.028 SD (95%CI 0.007, 0.050), p = 0.01 increases in standardised SATS score for English and mathematics respectively. Educational attainment is a complex multifactorial behavioural trait which has not had heritable contributions to it fully characterised. We were able to apply the results from a large study of adult educational attainment to a study of child exam performance marking events in the process of learning rather than realised adult end product. Our results support evidence for common, small genetic contributions to educational attainment, but also emphasise the likely lifecourse nature of this genetic effect. Results here also, by an alternative route, suggest that existing methods for child examination are able to recognise early life variation likely to be related to ultimate educational attainment.
  • Warner, N., Smits, R., McQueen, J. M., & Cutler, A. (2005). Phonological and statistical effects on timing of speech perception: Insights from a database of Dutch diphone perception. Speech Communication, 46(1), 53-72. doi:10.1016/j.specom.2005.01.003.

    Abstract

    We report detailed analyses of a very large database on timing of speech perception collected by Smits et al. (Smits, R., Warner, N., McQueen, J.M., Cutler, A., 2003. Unfolding of phonetic information over time: A database of Dutch diphone perception. J. Acoust. Soc. Am. 113, 563–574). Eighteen listeners heard all possible diphones of Dutch, gated in portions of varying size and presented without background noise. The present report analyzes listeners’ responses across gates in terms of phonological features (voicing, place, and manner for consonants; height, backness, and length for vowels). The resulting patterns for feature perception differ from patterns reported when speech is presented in noise. The data are also analyzed for effects of stress and of phonological context (neighboring vowel vs. consonant); effects of these factors are observed to be surprisingly limited. Finally, statistical effects, such as overall phoneme frequency and transitional probabilities, along with response biases, are examined; these too exercise only limited effects on response patterns. The results suggest highly accurate speech perception on the basis of acoustic information alone.
  • Warner, N., Kim, J., Davis, C., & Cutler, A. (2005). Use of complex phonological patterns in speech processing: Evidence from Korean. Journal of Linguistics, 41(2), 353-387. doi:10.1017/S0022226705003294.

    Abstract

    Korean has a very complex phonology, with many interacting alternations. In a coronal-/i/ sequence, depending on the type of phonological boundary present, alternations such as palatalization, nasal insertion, nasal assimilation, coda neutralization, and intervocalic voicing can apply. This paper investigates how the phonological patterns of Korean affect processing of morphemes and words. Past research on languages such as English, German, Dutch, and Finnish has shown that listeners exploit syllable structure constraints in processing speech and segmenting it into words. The current study shows that in parsing speech, listeners also use much more complex patterns that relate the surface phonological string to various boundaries.
  • Warner, N., Fountain, A., & Tucker, B. V. (2009). Cues to perception of reduced flaps. Journal of the Acoustical Society of America, 125(5), 3317-3327. doi:10.1121/1.3097773.

    Abstract

    Natural, spontaneous speech (and even quite careful speech) often shows extreme reduction in many speech segments, even resulting in apparent deletion of consonants. Where the flap ([(sic)]) allophone of /t/ and /d/ is expected in American English, one frequently sees an approximant-like or even vocalic pattern, rather than a clear flap. Still, the /t/ or /d/ is usually perceived, suggesting the acoustic characteristics of a reduced flap are sufficient for perception of a consonant. This paper identifies several acoustic characteristics of reduced flaps based on previous acoustic research (size of intensity dip, consonant duration, and F4 valley) and presents phonetic identification data for continua that manipulate these acoustic characteristics of reduction. The results indicate that the most obvious types of acoustic variability seen in natural flaps do affect listeners' percept of a consonant, but not sufficiently to completely account for the percept. Listeners are affected by the acoustic characteristics of consonant reduction, but they are also very skilled at evaluating variability along the acoustic dimensions that realize reduction.

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  • Warner, N., Otake, T., & Arai, A. (2010). Intonational structure as a word-boundary cue in Tokyo Japanese. Language and Speech, 53, 107-131. doi:10.1177/0023830909351235.

    Abstract

    While listeners are recognizing words from the connected speech stream, they are also parsing information from the intonational contour. This contour may contain cues to word boundaries, particularly if a language has boundary tones that occur at a large proportion of word onsets. We investigate how useful the pitch rise at the beginning of an accentual phrase (APR) would be as a potential word-boundary cue for Japanese listeners. A corpus study shows that it should allow listeners to locate approximately 40–60% of word onsets, while causing less than 1% false positives. We then present a word-spotting study which shows that Japanese listeners can, indeed, use accentual phrase boundary cues during segmentation. This work shows that the prosodic patterns that have been found in the production of Japanese also impact listeners’ processing.
  • Warner, N., Luna, Q., Butler, L., & Van Volkinburg, H. (2009). Revitalization in a scattered language community: Problems and methods from the perspective of Mutsun language revitalization. International Journal of the Sociology of Language, 198, 135-148. doi:10.1515/IJSL.2009.031.

    Abstract

    This article addresses revitalization of a dormant language whose prospective speakers live in scattered geographical areas. In comparison to increasing the usage of an endangered language, revitalizing a dormant language (one with no living speakers) requires different methods to gain knowledge of the language. Language teaching for a dormant language with a scattered community presents different problems from other teaching situations. In this article, we discuss the types of tasks that must be accomplished for dormant-language revitalization, with particular focus on development of teaching materials. We also address the role of computer technologies, arguing that each use of technology should be evaluated for how effectively it increases fluency. We discuss methods for achieving semi-fluency for the first new speakers of a dormant language, and for spreading the language through the community.
  • Warner, N., McQueen, J. M., & Cutler, A. (2014). Tracking perception of the sounds of English. The Journal of the Acoustical Society of America, 135, 2295-3006. doi:10.1121/1.4870486.

    Abstract

    Twenty American English listeners identified gated fragments of all 2288 possible English within-word and cross-word diphones, providing a total of 538 560 phoneme categorizations. The results show orderly uptake of acoustic information in the signal and provide a view of where information about segments occurs in time. Information locus depends on each speech sound’s identity and phonological features. Affricates and diphthongs have highly localized information so that listeners’ perceptual accuracy rises during a confined time range. Stops and sonorants have more distributed and gradually appearing information. The identity and phonological features (e.g., vowel vs consonant) of the neighboring segment also influences when acoustic information about a segment is available. Stressed vowels are perceived significantly more accurately than unstressed vowels, but this effect is greater for lax vowels than for tense vowels or diphthongs. The dataset charts the availability of perceptual cues to segment identity across time for the full phoneme repertoire of English in all attested phonetic contexts.
  • Wassenaar, M., & Hagoort, P. (2005). Word-category violations in patients with Broca's aphasia: An ERP study. Brain and Language, 92, 117-137. doi:10.1016/j.bandl.2004.05.011.

    Abstract

    An event-related brain potential experiment was carried out to investigate on-line syntactic processing in patients with Broca’s aphasia. Subjects were visually presented with sentences that were either syntactically correct or contained violations of word-category. Three groups of subjects were tested: Broca patients (N=11), non-aphasic patients with a right hemisphere (RH) lesion (N=9), and healthy aged-matched controls (N=15). Both control groups appeared sensitive to the violations of word-category as shown by clear P600/SPS effects. The Broca patients displayed only a very reduced and delayed P600/SPS effect. The results are discussed in the context of a lexicalist parsing model. It is concluded that Broca patients are hindered to detect on-line violations of word-category, if word class information is incomplete or delayed available.
  • Weber, K., & Indefrey, P. (2009). Syntactic priming in German–English bilinguals during sentence comprehension. Neuroimage, 46, 1164-1172. doi:10.1016/j.neuroimage.2009.03.040.

    Abstract

    A longstanding question in bilingualism is whether syntactic information is shared between the two language processing systems. We used an fMRI repetition suppression paradigm to investigate syntactic priming in reading comprehension in German–English late-acquisition bilinguals. In comparison to conventional subtraction analyses in bilingual experiments, repetition suppression has the advantage of being able to detect neuronal populations that are sensitive to properties that are shared by consecutive stimuli. In this study, we manipulated the syntactic structure between prime and target sentences. A sentence with a passive sentence structure in English was preceded either by a passive or by an active sentence in English or German. We looked for repetition suppression effects in left inferior frontal, left precentral and left middle temporal regions of interest. These regions were defined by a contrast of all non-target sentences in German and English versus the baseline of sentence-format consonant strings. We found decreases in activity (repetition suppression effects) in these regions of interest following the repetition of syntactic structure from the first to the second language and within the second language.
    Moreover, a separate behavioural experiment using a word-by-word reading paradigm similar to the fMRI experiment showed faster reading times for primed compared to unprimed English target sentences regardless of whether they were preceded by an English or a German sentence of the same structure.
    We conclude that there is interaction between the language processing systems and that at least some syntactic information is shared between a bilingual's languages with similar syntactic structures.

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  • Weber, A., Di Betta, A. M., & McQueen, J. M. (2014). Treack or trit: Adaptation to genuine and arbitrary foreign accents by monolingual and bilingual listeners. Journal of phonetics, 46, 34-51. doi:10.1016/j.wocn.2014.05.002.

    Abstract

    Two cross-modal priming experiments examined two questions about word recognition in foreign-accented speech: Does accent adaptation occur only for genuine accents markers, and does adaptation depend on language experience? We compared recognition of words spoken with canonical, genuinely-accented and arbitrarily-accented vowels. In Experiment 1, an Italian speaker pronounced vowels in English prime words canonically, or by lengthening /ɪ/ as in a genuine Italian accent (*/tri:k/ for trick), or by arbitrarily shortening /i:/ (*/trɪt/ for treat). Lexical-decision times to subsequent visual target words showed different priming effects in three listener groups. Monolingual native English listeners recognized variants with lengthened but not shortened vowels. Bilingual nonnative Italian-English listeners, who could not reliably distinguish vowel length, recognized both variants. Bilingual nonnative Dutch-English listeners also recognized both variants. In Experiment 2, bilingual Dutch-English listeners recognized Dutch words with genuinely- and arbitrarily-accented vowels (spoken by a native Italian with lengthened and shortened vowels respectively), but recognized words with canonical vowels more easily than words with accented vowels. These results suggest that adaptation to genuine accent markers arises for monolingual and bilingual listeners alike and can occur in native and nonnative languages, but that bilinguals can adapt to arbitrary accent markers better than monolinguals.
  • Wegener, C. (2005). Major word classes in Savosavo. Grazer Linguistische Studien, 64, 29-52.
  • Wegman, J., Fonteijn, H. M., van Ekert, J., Tyborowska, A., Jansen, C., & Janzen, G. (2014). Gray and white matter correlates of navigational ability in humans. Human Brain Mapping, 35(6), 2561-2572. doi:10.1002/hbm.22349.

    Abstract

    Humans differ widely in their navigational abilities. Studies have shown that self-reports on navigational abilities are good predictors of performance on navigation tasks in real and virtual environments. The caudate nucleus and medial temporal lobe regions have been suggested to subserve different navigational strategies. The ability to use different strategies might underlie navigational ability differences. This study examines the anatomical correlates of self-reported navigational ability in both gray and white matter. Local gray matter volume was compared between a group (N = 134) of good and bad navigators using voxel-based morphometry (VBM), as well as regional volumes. To compare between good and bad navigators, we also measured white matter anatomy using diffusion tensor imaging (DTI) and looked at fractional anisotropy (FA) values. We observed a trend toward higher local GM volume in right anterior parahippocampal/rhinal cortex for good versus bad navigators. Good male navigators showed significantly higher local GM volume in right hippocampus than bad male navigators. Conversely, bad navigators showed increased FA values in the internal capsule, the white matter bundle closest to the caudate nucleus and a trend toward higher local GM volume in the caudate nucleus. Furthermore, caudate nucleus regional volume correlated negatively with navigational ability. These convergent findings across imaging modalities are in line with findings showing that the caudate nucleus and the medial temporal lobes are involved in different wayfinding strategies. Our study is the first to show a link between self-reported large-scale navigational abilities and different measures of brain anatomy.
  • Wells, J. B., Christiansen, M. H., Race, D. S., Acheson, D. J., & MacDonald, M. C. (2009). Experience and sentence processing: Statistical learning and relative clause comprehension. Cognitive Psychology, 58(2), 250-271. doi:10.1016/j.cogpsych.2008.08.002.

    Abstract

    Many explanations of the difficulties associated with interpreting object relative clauses appeal to the demands that object relatives make on working memory. MacDonald and Christiansen [MacDonald, M. C., & Christiansen, M. H. (2002). Reassessing working memory: Comment on Just and Carpenter (1992) and Waters and Caplan (1996). Psychological Review, 109, 35-54] pointed to variations in reading experience as a source of differences, arguing that the unique word order of object relatives makes their processing more difficult and more sensitive to the effects of previous experience than the processing of subject relatives. This hypothesis was tested in a large-scale study manipulating reading experiences of adults over several weeks. The group receiving relative clause experience increased reading speeds for object relatives more than for subject relatives, whereas a control experience group did not. The reading time data were compared to performance of a computational model given different amounts of experience. The results support claims for experience-based individual differences and an important role for statistical learning in sentence comprehension processes.
  • Whitmarsh, S., Barendregt, H., Schoffelen, J.-M., & Jensen, O. (2014). Metacognitive awareness of covert somatosensory attention corresponds to contralateral alpha power. NeuroImage, 85(2), 803-809. doi:10.1016/j.neuroimage.2013.07.031.

    Abstract

    Studies on metacognition have shown that participants can report on their performance on a wide range of perceptual, memory and behavioral tasks. We know little, however, about the ability to report on one's attentional focus. The degree and direction of somatosensory attention can, however, be readily discerned through suppression of alpha band frequencies in EEG/MEG produced by the somatosensory cortex. Such top-down attentional modulations of cortical excitability have been shown to result in better discrimination performance and decreased response times. In this study we asked whether the degree of attentional focus is also accessible for subjective report, and whether such evaluations correspond to the amount of somatosensory alpha activity. In response to auditory cues participants maintained somatosensory attention to either their left or right hand for intervals varying randomly between 5 and 32seconds, while their brain activity was recorded with MEG. Trials were terminated by a probe sound, to which they reported their level of attention on the cued hand right before probe-onset. Using a beamformer approach, we quantified the alpha activity in left and right somatosensory regions, one second before the probe. Alpha activity from contra- and ipsilateral somatosensory cortices for high versus low attention trials were compared. As predicted, the contralateral somatosensory alpha depression correlated with higher reported attentional focus. Finally, alpha activity two to three seconds before the probe-onset was correlated with attentional focus. We conclude that somatosensory attention is indeed accessible to metacognitive awareness.
  • Widlok, T., & Burenhult, N. (2014). Sehen, riechen, orientieren. Spektrum der Wissenschaft, June 2014, 76-81.
  • Willems, R. M., Toni, I., Hagoort, P., & Casasanto, D. (2009). Body-specific motor imagery of hand actions: Neural evidence from right- and left-handers. Frontiers in Human Neuroscience, 3: 39, pp. 39. doi:10.3389/neuro.09.039.2009.

    Abstract

    If motor imagery uses neural structures involved in action execution, then the neural correlates of imagining an action should differ between individuals who tend to execute the action differently. Here we report fMRI data showing that motor imagery is influenced by the way people habitually perform motor actions with their particular bodies; that is, motor imagery is ‘body-specific’ (Casasanto, 2009). During mental imagery for complex hand actions, activation of cortical areas involved in motor planning and execution was left-lateralized in right-handers but right-lateralized in left-handers. We conclude that motor imagery involves the generation of an action plan that is grounded in the participant’s motor habits, not just an abstract representation at the level of the action’s goal. People with different patterns of motor experience form correspondingly different neurocognitive representations of imagined actions.
  • Willems, R. M., Hagoort, P., & Casasanto, D. (2010). Body-specific representations of action verbs: Neural evidence from right- and left-handers. Psychological Science, 21, 67-74. doi:10.1177/0956797609354072.

    Abstract

    According to theories of embodied cognition, understanding a verb like throw involves unconsciously simulating the action of throwing, using areas of the brain that support motor planning. If understanding action words involves mentally simulating one’s own actions, then the neurocognitive representation of word meanings should differ for people with different kinds of bodies, who perform actions in systematically different ways. In a test of the body-specificity hypothesis, we used functional magnetic resonance imaging to compare premotor activity correlated with action verb understanding in right- and left-handers. Righthanders preferentially activated the left premotor cortex during lexical decisions on manual-action verbs (compared with nonmanual-action verbs), whereas left-handers preferentially activated right premotor areas. This finding helps refine theories of embodied semantics, suggesting that implicit mental simulation during language processing is body specific: Right- and lefthanders, who perform actions differently, use correspondingly different areas of the brain for representing action verb meanings.
  • Willems, R. M., & Hagoort, P. (2009). Broca's region: Battles are not won by ignoring half of the facts. Trends in Cognitive Sciences, 13(3), 101. doi:10.1016/j.tics.2008.12.001.
  • Willems, R. M., Peelen, M. V., & Hagoort, P. (2010). Cerebral lateralization of face-selective and body-selective visual areas depends on handedness. Cerebral Cortex, 20, 1719-1725. doi:10.1093/cercor/bhp234.

    Abstract

    The left-hemisphere dominance for language is a core example of the functional specialization of the cerebral hemispheres. The degree of left-hemisphere dominance for language depends on hand preference: Whereas the majority of right-handers show left-hemispheric language lateralization, this number is reduced in left-handers. Here, we assessed whether handedness analogously has an influence upon lateralization in the visual system. Using functional magnetic resonance imaging, we localized 4 more or less specialized extrastriate areas in left- and right-handers, namely fusiform face area (FFA), extrastriate body area (EBA), fusiform body area (FBA), and human motion area (human middle temporal [hMT]). We found that lateralization of FFA and EBA depends on handedness: These areas were right lateralized in right-handers but not in left-handers. A similar tendency was observed in FBA but not in hMT. We conclude that the relationship between handedness and hemispheric lateralization extends to functionally lateralized parts of visual cortex, indicating a general coupling between cerebral lateralization and handedness. Our findings indicate that hemispheric specialization is not fixed but can vary considerably across individuals even in areas engaged relatively early in the visual system.
  • Willems, R. M., De Boer, M., De Ruiter, J. P., Noordzij, M. L., Hagoort, P., & Toni, I. (2010). A dissociation between linguistic and communicative abilities in the human brain. Psychological Science, 21, 8-14. doi:10.1177/0956797609355563.

    Abstract

    Although language is an effective vehicle for communication, it is unclear how linguistic and communicative abilities relate to each other. Some researchers have argued that communicative message generation involves perspective taking (mentalizing), and—crucially—that mentalizing depends on language. We employed a verbal communication paradigm to directly test whether the generation of a communicative action relies on mentalizing and whether the cerebral bases of communicative message generation are distinct from parts of cortex sensitive to linguistic variables. We found that dorsomedial prefrontal cortex, a brain area consistently associated with mentalizing, was sensitive to the communicative intent of utterances, irrespective of linguistic difficulty. In contrast, left inferior frontal cortex, an area known to be involved in language, was sensitive to the linguistic demands of utterances, but not to communicative intent. These findings show that communicative and linguistic abilities rely on cerebrally (and computationally) distinct mechanisms
  • Willems, R. M., Ozyurek, A., & Hagoort, P. (2009). Differential roles for left inferior frontal and superior temporal cortex in multimodal integration of action and language. Neuroimage, 47, 1992-2004. doi:10.1016/j.neuroimage.2009.05.066.

    Abstract

    Several studies indicate that both posterior superior temporal sulcus/middle temporal gyrus (pSTS/MTG) and left inferior frontal gyrus (LIFG) are involved in integrating information from different modalities. Here we investigated the respective roles of these two areas in integration of action and language information. We exploited the fact that the semantic relationship between language and different forms of action (i.e. co-speech gestures and pantomimes) is radically different. Speech and co-speech gestures are always produced together, and gestures are not unambiguously understood without speech. On the contrary, pantomimes are not necessarily produced together with speech and can be easily understood without speech. We presented speech together with these two types of communicative hand actions in matching or mismatching combinations to manipulate semantic integration load. Left and right pSTS/MTG were only involved in semantic integration of speech and pantomimes. Left IFG on the other hand was involved in integration of speech and co-speech gestures as well as of speech and pantomimes. Effective connectivity analyses showed that depending upon the semantic relationship between language and action, LIFG modulates activation levels in left pSTS.

    This suggests that integration in pSTS/MTG involves the matching of two input streams for which there is a relatively stable common object representation, whereas integration in LIFG is better characterized as the on-line construction of a new and unified representation of the input streams. In conclusion, pSTS/MTG and LIFG are differentially involved in multimodal integration, crucially depending upon the semantic relationship between the input streams.

    Additional information

    Supplementary table S1
  • Willems, R. M., & Hagoort, P. (2009). Hand preference influences neural correlates of action observation. Brain Research, 1269, 90-104. doi:10.1016/j.brainres.2009.02.057.

    Abstract

    It has been argued that we map observed actions onto our own motor system. Here we added to this issue by investigating whether hand preference influences the neural correlates of action observation of simple, essentially meaningless hand actions. Such an influence would argue for an intricate neural coupling between action production and action observation, which goes beyond effects of motor repertoire or explicit motor training, as has been suggested before. Indeed, parts of the human motor system exhibited a close coupling between action production and action observation. Ventral premotor and inferior and superior parietal cortices showed differential activation for left- and right-handers that was similar during action production as well as during action observation. This suggests that mapping observed actions onto the observer's own motor system is a core feature of action observation - at least for actions that do not have a clear goal or meaning. Basic differences in the way we act upon the world are not only reflected in neural correlates of action production, but can also influence the brain basis of action observation.
  • Willems, R. M., Toni, I., Hagoort, P., & Casasanto, D. (2010). Neural dissociations between action verb understanding and motor imagery. Journal of Cognitive Neuroscience, 22(10), 2387-2400. doi:10.1162/jocn.2009.21386.

    Abstract

    According to embodied theories of language, people understand a verb like throw, at least in part, by mentally simulating throwing. This implicit simulation is often assumed to be similar or identical to motor imagery. Here we used fMRI totest whether implicit simulations of actions during language understanding involve the same cortical motor regions as explicit motor imagery Healthy participants were presented with verbs related to hand actions (e.g., to throw) and nonmanual actions (e.g., to kneel). They either read these verbs (lexical decision task) or actively imagined performing the actions named by the verbs (imagery task). Primary motor cortex showd effector-specific activation during imagery, but not during lexical decision. Parts of premotor cortex distinguished manual from nonmanual actions during both lexical decision and imagery, but there was no overlap or correlation between regions activated during the two tasks. These dissociations suggest that implicit simulation and explicit imagery cued by action verbs may involve different types of motor representations and that the construct of “mental simulation” should be distinguished from “mental imagery” in embodied theories of language.
  • Willems, R. M., & Varley, R. (2010). Neural insights into the relation between language and communication. Frontiers in Human Neuroscience, 4, 203. doi:10.3389/fnhum.2010.00203.

    Abstract

    The human capacity to communicate has been hypothesized to be causally dependent upon language. Intuitively this seems plausible since most communication relies on language. Moreover, intention recognition abilities (as a necessary prerequisite for communication) and language development seem to co-develop. Here we review evidence from neuroimaging as well as from neuropsychology to evaluate the relationship between communicative and linguistic abilities. Our review indicates that communicative abilities are best considered as neurally distinct from language abilities. This conclusion is based upon evidence showing that humans rely on different cortical systems when designing a communicative message for someone else as compared to when performing core linguistic tasks, as well as upon observations of individuals with severe language loss after extensive lesions to the language system, who are still able to perform tasks involving intention understanding
  • Willems, R. M., Van der Haegen, L., Fisher, S. E., & Francks, C. (2014). On the other hand: Including left-handers in cognitive neuroscience and neurogenetics. Nature Reviews Neuroscience, 15, 193-201. doi:10.1038/nrn3679.

    Abstract

    Left-handers are often excluded from study cohorts in neuroscience and neurogenetics in order to reduce variance in the data. However, recent investigations have shown that the inclusion or targeted recruitment of left-handers can be informative in studies on a range of topics, such as cerebral lateralization and the genetic underpinning of asymmetrical brain development. Left-handed individuals represent a substantial portion of the human population and therefore left-handedness falls within the normal range of human diversity; thus, it is important to account for this variation in our understanding of brain functioning. We call for neuroscientists and neurogeneticists to recognize the potential of studying this often-discarded group of research subjects.
  • Willems, R. M., & Francks, C. (2014). Your left-handed brain. Frontiers for Young Minds, 2: 13. doi:10.3389/frym.2014.00013.

    Abstract

    While most people prefer to use their right hand to brush their teeth, throw a ball, or hold a tennis racket, left-handers prefer to use their left hand. This is the case for around 10 per cent of all people. There was a time (not so long ago) when left-handers were stigmatized in Western (and other) communities: it was considered a bad sign if you were left-handed, and left-handed children were often forced to write with their right hand. This is nonsensical: there is nothing wrong with being left-handed, and trying to write with the non-preferred hand is frustrating for almost everybody. As a matter of fact, science can learn from left-handers, and in this paper, we discuss how this may be the case. We review why some people are left-handed and others are not, how left-handers' brains differ from right-handers’, and why scientists study left-handedness in the first place
  • Witteman, M. J., & Segers, E. (2010). The modality effect tested in children in a user-paced multimedia environment. Journal of Computer Assisted Learning, 26, 132-142. doi:10.1111/j.1365-2729.2009.00335.x.

    Abstract

    The modality learning effect, according to Mayer (2001), proposes that learning is enhanced when information is presented in both the visual and auditory domain (e.g., pictures and spoken information), compared to presenting information solely in the visual channel (e.g., pictures and written text). Most of the evidence for this effect comes from adults in a laboratory setting. Therefore, we tested the modality effect with 80 children in the highest grade of elementary school, in a naturalistic setting. In a between-subjects design children either saw representational pictures with speech or representational pictures with text. Retention and transfer knowledge was tested at three moments: immediately after the intervention, one day after, and after one week. The present study did not find any evidence for a modality effect in children when the lesson is learner-paced. Instead, we found a reversed modality effect directly after the intervention for retention. A reversed modality effect was also found for the transfer questions one day later. This effect was robust, even when controlling for individual differences.
  • Witteman, M. J., Weber, A., & McQueen, J. M. (2014). Tolerance for inconsistency in foreign-accented speech. Psychonomic Bulletin & Review, 21, 512-519. doi:10.3758/s13423-013-0519-8.

    Abstract

    Are listeners able to adapt to a foreign-accented speaker who has, as is often the case, an inconsistent accent? Two groups of native Dutch listeners participated in a cross-modal priming experiment, either in a consistent-accent condition (German-accented items only) or in an inconsistent-accent condition (German-accented and nativelike pronunciations intermixed). The experimental words were identical for both groups (words with vowel substitutions characteristic of German-accented speech); additional contextual words differed in accentedness (German-accented or nativelike words). All items were spoken by the same speaker: a German native who could produce the accented forms but could also pass for a Dutch native speaker. Listeners in the consistent-accent group were able to adapt quickly to the speaker (i.e., showed facilitatory priming for words with vocalic substitutions). Listeners in the inconsistent-accent condition showed adaptation to words with vocalic substitutions only in the second half of the experiment. These results indicate that adaptation to foreign-accented speech is rapid. Accent inconsistency slows listeners down initially, but a short period of additional exposure is enough for them to adapt to the speaker. Listeners can therefore tolerate inconsistency in foreign-accented speech.
  • Wittenburg, P., Skiba, R., & Trilsbeek, P. (2005). The language archive at the MPI: Contents, tools, and technologies. Language Archives Newsletter, 5, 7-9.
  • Wittenburg, P. (2010). Archiving and accessing language resources. Concurrency and Computation: Practice and Experience, 22(17), 2354-2368. doi:10.1002/cpe.1605.

    Abstract

    Languages are among the most complex systems that evolution has created. With an unforeseen speed many of these unique results of evolution are currently disappearing: every two weeks one of the 6500 still spoken languages is dying and many are subject to extreme changes due to globalization. Experts understood the need to document the languages and preserve the cultural and linguistic treasures embedded in them for future generations. Also linguistic theory will need to consider the variation of the linguistic systems encoded in languages to improve our understanding of how human minds process language material, thus accessibility to all types of resources is increasingly crucial. Deeper insights into human language processing and a higher degree of integration and interoperability between resources will also improve our language processing technology. The DOBES programme is focussing on the documentation and preservation of language material. The Max Planck Institute developed the Language Archiving Technology to help researchers when creating, archiving and accessing language resources. The recently started CLARIN research infrastructure has as main goals to achieve a broad visibility and an easy
    accessibility of language resources.
  • Wnuk, E., & Burenhult, N. (2014). Contact and isolation in hunter-gatherer language dynamics: Evidence from Maniq phonology (Aslian, Malay Peninsula). Studies in Language, 38(4), 956-981. doi:10.1075/sl.38.4.06wnu.
  • Wnuk, E., & Majid, A. (2014). Revisiting the limits of language: The odor lexicon of Maniq. Cognition, 131, 125-138. doi:10.1016/j.cognition.2013.12.008.

    Abstract

    It is widely believed that human languages cannot encode odors. While this is true for English,
    and other related languages, data from some non-Western languages challenge this
    view. Maniq, a language spoken by a small population of nomadic hunter–gatherers in
    southern Thailand, is such a language. It has a lexicon of over a dozen terms dedicated
    to smell. We examined the semantics of these smell terms in 3 experiments (exemplar
    listing, similarity judgment and off-line rating). The exemplar listing task confirmed that
    Maniq smell terms have complex meanings encoding smell qualities. Analyses of the
    similarity data revealed that the odor lexicon is coherently structured by two dimensions.
    The underlying dimensions are pleasantness and dangerousness, as verified by the off-line
    rating study. Ethnographic data illustrate that smell terms have detailed semantics tapping
    into broader cultural constructs. Contrary to the widespread view that languages cannot
    encode odors, the Maniq data show odor can be a coherent semantic domain, thus shedding
    new light on the limits of language.
  • Wohlgemuth, J., & Dirksmeyer, T. (Eds.). (2005). Bedrohte Vielfalt. Aspekte des Sprach(en)tods – Aspects of language death. Berlin: Weißensee.

    Abstract

    About 5,000 languages are spoken in the world today. More than half of them have less than 10,000 speakers, a quarter of them even fewer than 1,000. The majority of these “small” languages will not live to see the end of this century; some estimates predict that no more than a dozen languages will still be spoken by the turn of the next millennium. This collection of papers approaches the subject of language extinction through five major topics: general aspects of language death, case studies, endangered subsystems, language protection and revitalization, language ecology. In 24 articles, the authors address the causes, manifestations, and consequences of language endangerment and extinction as well as the linguistic and social changes associated with it, drawing examples from a large number of languages.
  • Won, S.-O., Hu, I., Kim, M.-Y., Bae, J.-M., Kim, Y.-M., & Byun, K.-S. (2009). Theory and practice of Sign Language interpretation. Pyeongtaek: Korea National College of Rehabilitation & Welfare.
  • Xiang, H.-D., Fonteijn, H. M., Norris, D. G., & Hagoort, P. (2010). Topographical functional connectivity pattern in the perisylvian language networks. Cerebral Cortex, 20, 549-560. doi:10.1093/cercor/bhp119.

    Abstract

    We performed a resting-state functional connectivity study to investigate directly the functional correlations within the perisylvian language networks by seeding from 3 subregions of Broca's complex (pars opercularis, pars triangularis, and pars orbitalis) and their right hemisphere homologues. A clear topographical functional connectivity pattern in the left middle frontal, parietal, and temporal areas was revealed for the 3 left seeds. This is the first demonstration that a functional connectivity topology can be observed in the perisylvian language networks. The results support the assumption of the functional division for phonology, syntax, and semantics of Broca's complex as proposed by the memory, unification, and control (MUC) model and indicated a topographical functional organization in the perisylvian language networks, which suggests a possible division of labor for phonological, syntactic, and semantic function in the left frontal, parietal, and temporal areas.
  • Yang, Y., Dai, B., Howell, P., Wang, X., Li, K., & Lu, C. (2014). White and Grey Matter Changes in the Language Network during Healthy Aging. PLoS One, 9(9): e108077. doi: 10.1371/journal.pone.0108077.

    Abstract

    Neural structures change with age but there is no consensus on the exact processes involved. This study tested the hypothesis that white and grey matter in the language network changes during aging according to a “last in, first out” process. The fractional anisotropy (FA) of white matter and cortical thickness of grey matter were measured in 36 participants whose ages ranged from 55 to 79 years. Within the language network, the dorsal pathway connecting the mid-to-posterior superior temporal cortex (STC) and the inferior frontal cortex (IFC) was affected more by aging in both FA and thickness than the other dorsal pathway connecting the STC with the premotor cortex and the ventral pathway connecting the mid-to-anterior STC with the ventral IFC. These results were independently validated in a second group of 20 participants whose ages ranged from 50 to 73 years. The pathway that is most affected during aging matures later than the other two pathways (which are present at birth). The results are interpreted as showing that the neural structures which mature later are affected more than those that mature earlier, supporting the “last in, first out” theory.
  • Zeshan, U., & Panda, S. (2005). Professional course in Indian sign language. Mumbai: Ali Yavar Jung National Institute for the Hearing Handicapped.
  • Zeshan, U., Vasishta, M. N., & Sethna, M. (2005). Implementation of Indian Sign Language in educational settings. Asia Pacific Disability Rehabilitation Journal, 16(1), 16-40.

    Abstract

    This article reports on several sub-projects of research and development related to the use of Indian Sign Language in educational settings. In many countries around the world, sign languages are now recognised as the legitimate, full-fledged languages of the deaf communities that use them. In India, the development of sign language resources and their application in educational contexts, is still in its initial stages. The work reported on here, is the first principled and comprehensive effort of establishing educational programmes in Indian Sign Language at a national level. Programmes are of several types: a) Indian Sign Language instruction for hearing people; b) sign language teacher training programmes for deaf people; and c) educational materials for use in schools for the Deaf. The conceptual approach used in the programmes for deaf students is known as bilingual education, which emphasises the acquisition of a first language, Indian Sign Language, alongside the acquisition of spoken languages, primarily in their written form.
  • Zhang, J., Bao, S., Furumai, R., Kucera, K. S., Ali, A., Dean, N. M., & Wang, X.-F. (2005). Protein phosphatase 5 is required for ATR-mediated checkpoint activation. Molecular and Cellular Biology, 25, 9910-9919. doi:10.1128/​MCB.25.22.9910-9919.2005.

    Abstract

    In response to DNA damage or replication stress, the protein kinase ATR is activated and subsequently transduces genotoxic signals to cell cycle control and DNA repair machinery through phosphorylation of a number of downstream substrates. Very little is known about the molecular mechanism by which ATR is activated in response to genotoxic insults. In this report, we demonstrate that protein phosphatase 5 (PP5) is required for the ATR-mediated checkpoint activation. PP5 forms a complex with ATR in a genotoxic stress-inducible manner. Interference with the expression or the activity of PP5 leads to impairment of the ATR-mediated phosphorylation of hRad17 and Chk1 after UV or hydroxyurea treatment. Similar results are obtained in ATM-deficient cells, suggesting that the observed defect in checkpoint signaling is the consequence of impaired functional interaction between ATR and PP5. In cells exposed to UV irradiation, PP5 is required to elicit an appropriate S-phase checkpoint response. In addition, loss of PP5 leads to premature mitosis after hydroxyurea treatment. Interestingly, reduced PP5 activity exerts differential effects on the formation of intranuclear foci by ATR and replication protein A, implicating a functional role for PP5 in a specific stage of the checkpoint signaling pathway. Taken together, our results suggest that PP5 plays a critical role in the ATR-mediated checkpoint activation.
  • Zhernakova, A., Elbers, C. C., Ferwerda, B., Romanos, J., Trynka, G., Dubois, P. C., De Kovel, C. G. F., Franke, L., Oosting, M., Barisani, D., Bardella, M. T., Joosten, L. A. B., Saavalainen, P., van Heel, D. A., Catassi, C., Netea, M. G., Wijmenga, C., & Finnish Celiac Dis Study, G. (2010). Evolutionary and Functional Analysis of Celiac Risk Loci Reveals SH2B3 as a Protective Factor against Bacterial Infection. American Journal of Human Genetics, 86(6), 970-977. doi:10.1016/j.ajhg.2010.05.004.

    Abstract

    Celiac disease (CD) is an intolerance to dietary proteins of wheat, barley, and rye. CD may have substantial morbidity, yet it is quite common with a prevalence of 1%-2% in Western populations. It is not clear why the CD phenotype is so prevalent despite its negative effects on human health, especially because appropriate treatment in the form of a gluten-free diet has only been available since the 1950s, when dietary gluten was discovered to be the triggering factor. The high prevalence of CD might suggest that genes underlying this disease may have been favored by the process of natural selection. We assessed signatures of selection for ten confirmed CD-associated loci in several genome-wide data sets, comprising 8154 controls from four European populations and 195 individuals from a North African population, by studying haplotype lengths via the integrated haplotype score (iHS) method. Consistent signs of positive selection for CD-associated derived alleles were observed in three loci: IL12A, IL18RAP, and SH2B3. For the SH2B3 risk allele, we also show a difference in allele frequency distribution (F(st)) between HapMap phase II populations. Functional investigation of the effect of the SH2B3 genotype in response to lipopolysaccharide and muramyl dipeptide revealed that carriers of the SH2B3 rs3184504*A risk allele showed stronger activation of the NOD2 recognition pathway. This suggests that SH2B3 plays a role in protection against bacteria infection, and it provides a possible explanation for the selective sweep on SH2B3, which occurred sometime between 1200 and 1700 years ago.
  • De Zubicaray, G. I., Hartsuiker, R. J., & Acheson, D. J. (2014). Mind what you say—general and specific mechanisms for monitoring in speech production. Frontiers in Human Neuroscience, 8: 514. doi:10.3389%2Ffnhum.2014.00514.

    Abstract

    For most people, speech production is relatively effortless and error-free. Yet it has long been recognized that we need some type of control over what we are currently saying and what we plan to say. Precisely how we monitor our internal and external speech has been a topic of research interest for several decades. The predominant approach in psycholinguistics has assumed monitoring of both is accomplished via systems responsible for comprehending others' speech.

    This special topic aimed to broaden the field, firstly by examining proposals that speech production might also engage more general systems, such as those involved in action monitoring. A second aim was to examine proposals for a production-specific, internal monitor. Both aims require that we also specify the nature of the representations subject to monitoring.
  • Zumer, J. M., Scheeringa, R., Schoffelen, J.-M., Norris, D. G., & Jensen, O. (2014). Occipital alpha activity during stimulus processing gates the information flow to object-selective cortex. PLoS Biology, 12(10): e1001965. doi:10.1371/journal.pbio.1001965.

    Abstract

    Given the limited processing capabilities of the sensory system, it is essential that attended information is gated to downstream areas, whereas unattended information is blocked. While it has been proposed that alpha band (8–13 Hz) activity serves to route information to downstream regions by inhibiting neuronal processing in task-irrelevant regions, this hypothesis remains untested. Here we investigate how neuronal oscillations detected by electroencephalography in visual areas during working memory encoding serve to gate information reflected in the simultaneously recorded blood-oxygenation-level-dependent (BOLD) signals recorded by functional magnetic resonance imaging in downstream ventral regions. We used a paradigm in which 16 participants were presented with faces and landscapes in the right and left hemifields; one hemifield was attended and the other unattended. We observed that decreased alpha power contralateral to the attended object predicted the BOLD signal representing the attended object in ventral object-selective regions. Furthermore, increased alpha power ipsilateral to the attended object predicted a decrease in the BOLD signal representing the unattended object. We also found that the BOLD signal in the dorsal attention network inversely correlated with visual alpha power. This is the first demonstration, to our knowledge, that oscillations in the alpha band are implicated in the gating of information from the visual cortex to the ventral stream, as reflected in the representationally specific BOLD signal. This link of sensory alpha to downstream activity provides a neurophysiological substrate for the mechanism of selective attention during stimulus processing, which not only boosts the attended information but also suppresses distraction. Although previous studies have shown a relation between the BOLD signal from the dorsal attention network and the alpha band at rest, we demonstrate such a relation during a visuospatial task, indicating that the dorsal attention network exercises top-down control of visual alpha activity.
  • Zwitserlood, I., van den Bogaerde, B., & Terpstra, A. (2010). De Nederlandse Gebarentaal en het ERK. Levende Talen Magazine, 2010(5), 50-51.
  • Zwitserlood, I. (2010). De Nederlandse Gebarentaal, het Corpus NGT en het ERK. Levende Talen Magazine, 2010(8), 44-45.
  • Zwitserlood, I. (2010). Laat je vingers spreken: NGT en vingerspelling. Levende Talen Magazine, 2010(2), 46-47.
  • Zwitserlood, I. (2009). Het Corpus NGT. Levende Talen Magazine, 6, 44-45.

    Abstract

    The Corpus NGT
  • Zwitserlood, I. (2010). Het Corpus NGT en de dagelijkse lespraktijk (2). Levende Talen Magazine, 2010(3), 47-48.
  • Zwitserlood, I. (2009). Het Corpus NGT en de dagelijkse lespraktijk (1). Levende Talen Magazine, 8, 40-41.
  • Zwitserlood, I. (2010). Sign language lexicography in the early 21st century and a recently published dictionary of Sign Language of the Netherlands. International Journal of Lexicography, 23, 443-476. doi:10.1093/ijl/ecq031.

    Abstract

    Sign language lexicography has thus far been a relatively obscure area in the world of lexicography. Therefore, this article will contain background information on signed languages and the communities in which they are used, on the lexicography of sign languages, the situation in the Netherlands as well as a review of a sign language dictionary that has recently been published in the Netherlands.
  • Zwitserlood, I., & Crasborn, O. (2010). Wat kunnen we leren uit een Corpus Nederlandse Gebarentaal? WAP Nieuwsbrief, 28(2), 16-18.
  • Zwitserlood, I. (2010). Verlos ons van de glos. Levende Talen Magazine, 2010(7), 40-41.

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