Publications

Displaying 1701 - 1794 of 1794
  • Vromans, R. D., & Jongman, S. R. (2018). The interplay between selective and nonselective inhibition during single word production. PLoS One, 13(5): e0197313. doi:10.1371/journal.pone.0197313.

    Abstract

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

    Additional information

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

    Abstract

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

    Abstract

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

    Abstract

    Disrupted in schizophrenia 1 (DISC1) is a leading candidate susceptibility gene for schizophrenia, bipolar disorder, and recurrent major depression, which has been implicated in other psychiatric illnesses of neurodevelopmental origin, including autism. DISC1 was initially identified at the breakpoint of a balanced chromosomal translocation, t(1;11) (q42.1;14.3), in a family with a high incidence of psychiatric illness. Carriers of the translocation show a 50% reduction in DISC1 protein levels, suggesting altered DISC1 expression as a pathogenic mechanism in psychiatric illness. Altered DISC1 expression in the post-mortem brains of individuals with psychiatric illness and the frequent implication of non-coding regions of the gene by association analysis further support this assertion. Here, we provide the first characterisation of the DISC1 promoter region. Using dual luciferase assays, we demonstrate that a region -300bp to -177bp relative to the transcription start site (TSS) contributes positively to DISC1 promoter activity, whilst a region -982bp to -301bp relative to the TSS confers a repressive effect. We further demonstrate inhibition of DISC1 promoter activity and protein expression by FOXP2, a transcription factor implicated in speech and language function. This inhibition is diminished by two distinct FOXP2 point mutations, R553H and R328X, which were previously found in families affected by developmental verbal dyspraxia (DVD). Our work identifies an intriguing mechanistic link between neurodevelopmental disorders that have traditionally been viewed as diagnostically distinct but which do share varying degrees of phenotypic overlap.
  • Waller, D., Loomis, J. M., & Haun, D. B. M. (2004). Body-based senses enhance knowledge of directions in large-scale environments. Psychonomic Bulletin & Review, 11(1), 157-163.

    Abstract

    Previous research has shown that inertial cues resulting from passive transport through a large environment do not necessarily facilitate acquiring knowledge about its layout. Here we examine whether the additional body-based cues that result from active movement facilitate the acquisition of spatial knowledge. Three groups of participants learned locations along an 840-m route. One group walked the route during learning, allowing access to body-based cues (i.e., vestibular, proprioceptive, and efferent information). Another group learned by sitting in the laboratory, watching videos made from the first group. A third group watched a specially made video that minimized potentially confusing head-on-trunk rotations of the viewpoint. All groups were tested on their knowledge of directions in the environment as well as on its configural properties. Having access to body-based information reduced pointing error by a small but significant amount. Regardless of the sensory information available during learning, participants exhibited strikingly common biases.
  • Wang, L., Jensen, O., Van den Brink, D., Weder, N., Schoffelen, J.-M., Magyari, L., Hagoort, P., & Bastiaansen, M. C. M. (2012). Beta oscillations relate to the N400m during language comprehension. Human Brain Mapping, 33, 2898-2912. doi:10.1002/hbm.21410.

    Abstract

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

    Abstract

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

    Abstract

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

    Abstract

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

    Abstract

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

    Abstract

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

    Abstract

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

    Additional information

    1-s2.0-S0006322317321911-mmc1.pdf
  • 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., Jongman, A., Sereno, J., & Kemps, R. J. J. K. (2004). Incomplete neutralization and other sub-phonemic durational differences in production and perception: Evidence from Dutch. Journal of Phonetics, 32(2), 251-276. doi:10.1016/S0095-4470(03)00032-9.

    Abstract

    Words which are expected to contain the same surface string of segments may, under identical prosodic circumstances, sometimes be realized with slight differences in duration. Some researchers have attributed such effects to differences in the words’ underlying forms (incomplete neutralization), while others have suggested orthographic influence and extremely careful speech as the cause. In this paper, we demonstrate such sub-phonemic durational differences in Dutch, a language which some past research has found not to have such effects. Past literature has also shown that listeners can often make use of incomplete neutralization to distinguish apparent homophones. We extend perceptual investigations of this topic, and show that listeners can perceive even durational differences which are not consistently observed in production. We further show that a difference which is primarily orthographic rather than underlying can also create such durational differences. We conclude that a wide variety of factors, in addition to underlying form, can induce speakers to produce slight durational differences which listeners can also use in perception.
  • 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., 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.
  • Warren, C. M., Tona, K. D., Ouwekerk, L., Van Paridon, J., Poletiek, F. H., Bosch, J. A., & Nieuwenhuis, S. (2019). The neuromodulatory and hormonal effects of transcutaneous vagus nerve stimulation as evidenced by salivary alpha amylase, salivary cortisol, pupil diameter, and the P3 event-related potential. Brain Stimulation, 12(3), 635-642. doi:10.1016/j.brs.2018.12.224.

    Abstract

    Background

    Transcutaneous vagus nerve stimulation (tVNS) is a new, non-invasive technique being investigated as an intervention for a variety of clinical disorders, including epilepsy and depression. It is thought to exert its therapeutic effect by increasing central norepinephrine (NE) activity, but the evidence supporting this notion is limited.
    Objective

    In order to test for an impact of tVNS on psychophysiological and hormonal indices of noradrenergic function, we applied tVNS in concert with assessment of salivary alpha amylase (SAA) and cortisol, pupil size, and electroencephalograph (EEG) recordings.
    Methods

    Across three experiments, we applied real and sham tVNS to 61 healthy participants while they performed a set of simple stimulus-discrimination tasks. Before and after the task, as well as during one break, participants provided saliva samples and had their pupil size recorded. EEG was recorded throughout the task. The target for tVNS was the cymba conchae, which is heavily innervated by the auricular branch of the vagus nerve. Sham stimulation was applied to the ear lobe.
    Results

    P3 amplitude was not affected by tVNS (Experiment 1A: N=24; Experiment 1B: N=20; Bayes factor supporting null model=4.53), nor was pupil size (Experiment 2: N=16; interaction of treatment and time: p=0.79). However, tVNS increased SAA (Experiments 1A and 2: N=25) and attenuated the decline of salivary cortisol compared to sham (Experiment 2: N=17), as indicated by significant interactions involving treatment and time (p=.023 and p=.040, respectively).
    Conclusion

    These findings suggest that tVNS modulates hormonal indices but not psychophysiological indices of noradrenergic function.
  • Wassenaar, M., & Hagoort, P. (2007). Thematic role assignment in patients with Broca's aphasia: Sentence-picture matching electrified. Neuropsychologia, 45(4), 716-740. doi:10.1016/j.neuropsychologia.2006.08.016.

    Abstract

    An event-related brain potential experiment was carried out to investigate on-line thematic role assignment during sentence–picture matching in patients with Broca's aphasia. Subjects were presented with a picture that was followed by an auditory sentence. The sentence either matched the picture or mismatched the visual information depicted. Sentences differed in complexity, and ranged from simple active semantically irreversible sentences to passive semantically reversible sentences. ERPs were recorded while subjects were engaged in sentence–picture matching. In addition, reaction time and accuracy were measured. Three groups of subjects were tested: Broca patients (N = 10), non-aphasic patients with a right hemisphere (RH) lesion (N = 8), and healthy aged-matched controls (N = 15). The results of this study showed that, in neurologically unimpaired individuals, thematic role assignment in the context of visual information was an immediate process. This in contrast to patients with Broca's aphasia who demonstrated no signs of on-line sensitivity to the picture–sentence mismatches. The syntactic contribution to the thematic role assignment process seemed to be diminished given the reduction and even absence of P600 effects. Nevertheless, Broca patients showed some off-line behavioral sensitivity to the sentence–picture mismatches. The long response latencies of Broca's aphasics make it likely that off-line response strategies were used.
  • Wassenaar, M., Brown, C. M., & Hagoort, P. (2004). ERP-effects of subject-verb agreement violations in patients with Broca's aphasia. Journal of Cognitive Neuroscience, 16(4), 553-576. doi:10.1162/089892904323057290.

    Abstract

    This article presents electrophysiological data on on-line syntactic processing during auditory sentence comprehension in patients with Broca's aphasia. Event-related brain potentials (ERPs) were recorded from the scalp while subjects listened to sentences that were either syntactically correct or contained violations of subject-verb agreement. Three groups of subjects were tested: Broca patients (n = 10), nonaphasic patients with a right-hemisphere (RH) lesion (n = 5), and healthy agedmatched controls (n = 12). The healthy, control subjects showed a P600/SPS effect as response to the agreement violations. The nonaphasic patients with an RH lesion showed essentially the same pattern. The overall group of Broca patients did not show this sensitivity. However, the sensitivity was modulated by the severity of the syntactic comprehension impairment. The largest deviation from the standard P600/SPS effect was found in the patients with the relatively more severe syntactic comprehension impairment. In addition, ERPs to tones in a classical tone oddball paradigm were also recorded. Similar to the normal control subjects and RH patients, the group of Broca patients showed a P300 effect in the tone oddball condition. This indicates that aphasia in itself does not lead to a general reduction in all cognitive ERP effects. It was concluded that deviations from the standard P600/SPS effect in the Broca patients reflected difficulties with on-line maintaining of number information across clausal boundaries for establishing subject-verb agreement.
  • Wassenaar, M., Hagoort, P., & Brown, C. M. (1997). Syntactic ERP effects in Broca's aphasics with agrammatic comprehension. Brain and Language, 60, 61-64. doi:10.1006/brln.1997.1911.
  • Watson, L. M., Wong, M. M. K., Vowles, J., Cowley, S. A., & Becker, E. B. E. (2018). A simplified method for generating purkinje cells from human-induced pluripotent stem cells. The Cerebellum, 17(4), 419-427. doi:10.1007/s12311-017-0913-2.

    Abstract

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

    Additional information

    12311_2017_913_MOESM1_ESM.docx
  • Weber, A., & Cutler, A. (2004). Lexical competition in non-native spoken-word recognition. Journal of Memory and Language, 50(1), 1-25. doi:10.1016/S0749-596X(03)00105-0.

    Abstract

    Four eye-tracking experiments examined lexical competition in non-native spoken-word recognition. Dutch listeners hearing English fixated longer on distractor pictures with names containing vowels that Dutch listeners are likely to confuse with vowels in a target picture name (pencil, given target panda) than on less confusable distractors (beetle, given target bottle). English listeners showed no such viewing time difference. The confusability was asymmetric: given pencil as target, panda did not distract more than distinct competitors. Distractors with Dutch names phonologically related to English target names (deksel, ‘lid,’ given target desk) also received longer fixations than distractors with phonologically unrelated names. Again, English listeners showed no differential effect. With the materials translated into Dutch, Dutch listeners showed no activation of the English words (desk, given target deksel). The results motivate two conclusions: native phonemic categories capture second-language input even when stored representations maintain a second-language distinction; and lexical competition is greater for non-native than for native listeners.
  • Weber, A., & Scharenborg, O. (2012). Models of spoken-word recognition. Wiley Interdisciplinary Reviews: Cognitive Science, 3, 387-401. doi:10.1002/wcs.1178.

    Abstract

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

    Abstract

    We present two eye-tracking experiments that investigate lexical frequency and semantic context constraints in spoken-word recognition in German. In both experiments, the pivotal words were pairs of nouns overlapping at onset but varying in lexical frequency. In Experiment 1, German listeners showed an expected frequency bias towards high-frequency competitors (e.g., Blume, ‘flower’) when instructed to click on low-frequency targets (e.g., Bluse, ‘blouse’). In Experiment 2, semantically constraining context increased the availability of appropriate low-frequency target words prior to word onset, but did not influence the availability of semantically inappropriate high-frequency competitors at the same time. Immediately after target word onset, however, the activation of high-frequency competitors was reduced in semantically constraining sentences, but still exceeded that of unrelated distractor words significantly. The results suggest that (1) semantic context acts to downgrade activation of inappropriate competitors rather than to exclude them from competition, and (2) semantic context influences spoken-word recognition, over and above anticipation of upcoming referents.
  • Weber, K., Christiansen, M., Indefrey, P., & Hagoort, P. (2019). Primed from the start: Syntactic priming during the first days of language learning. Language Learning, 69(1), 198-221. doi:10.1111/lang.12327.

    Abstract

    New linguistic information must be integrated into our existing language system. Using a novel experimental task that incorporates a syntactic priming paradigm into artificial language learning, we investigated how new grammatical regularities and words are learned. This innovation allowed us to control the language input the learner received, while the syntactic priming paradigm provided insight into the nature of the underlying syntactic processing machinery. The results of the present study pointed to facilitatory syntactic processing effects within the first days of learning: Syntactic and lexical priming effects revealed participants’ sensitivity to both novel words and word orders. This suggested that novel syntactic structures and their meaning (form–function mapping) can be acquired rapidly through incidental learning. More generally, our study indicated similar mechanisms for learning and processing in both artificial and natural languages, with implications for the relationship between first and second language learning.
  • Weber, K., Micheli, C., Ruigendijk, E., & Rieger, J. (2019). Sentence processing is modulated by the current linguistic environment and a priori information: An fMRI study. Brain and Behavior, 9(7): e01308. doi:10.1002/brb3.1308.

    Abstract

    Introduction
    Words are not processed in isolation but in rich contexts that are used to modulate and facilitate language comprehension. Here, we investigate distinct neural networks underlying two types of contexts, the current linguistic environment and verb‐based syntactic preferences.

    Methods
    We had two main manipulations. The first was the current linguistic environment, where the relative frequencies of two syntactic structures (prepositional object [PO] and double‐object [DO]) would either follow everyday linguistic experience or not. The second concerned the preference toward one or the other structure depending on the verb; learned in everyday language use and stored in memory. German participants were reading PO and DO sentences in German while brain activity was measured with functional magnetic resonance imaging.

    Results
    First, the anterior cingulate cortex (ACC) showed a pattern of activation that integrated the current linguistic environment with everyday linguistic experience. When the input did not match everyday experience, the unexpected frequent structure showed higher activation in the ACC than the other conditions and more connectivity from the ACC to posterior parts of the language network. Second, verb‐based surprisal of seeing a structure given a verb (PO verb preference but DO structure presentation) resulted, within the language network (left inferior frontal and left middle/superior temporal gyrus) and the precuneus, in increased activation compared to a predictable verb‐structure pairing.

    Conclusion
    In conclusion, (1) beyond the canonical language network, brain areas engaged in prediction and error signaling, such as the ACC, might use the statistics of syntactic structures to modulate language processing, (2) the language network is directly engaged in processing verb preferences. These two networks show distinct influences on sentence processing.

    Additional information

    Supporting information
  • 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.
  • Weekes, B. S., Abutalebi, J., Mak, H.-K.-F., Borsa, V., Soares, S. M. P., Chiu, P. W., & Zhang, L. (2018). Effect of monolingualism and bilingualism in the anterior cingulate cortex: a proton magnetic resonance spectroscopy study in two centers. Letras de Hoje, 53(1), 5-12. doi:10.15448/1984-7726.2018.1.30954.

    Abstract

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

    Abstract

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

    Abstract

    Beller, Bender, and Medin question the necessity of including social anthropology within the cognitive sciences. We argue that there is great scope for fruitful rapprochement while agreeing that there are obstacles (even if we might wish to debate some of those specifically identified by Beller and colleagues). We frame the general problem differently, however: not in terms of the problem of reconciling disciplines and research cultures, but rather in terms of the prospects for collaborative deployment of expertise (methodological and theoretical) in problem-driven research. For the purposes of illustration, our focus in this article is on the evolution of cooperation
  • 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. (2004). Ethnography in language Documentation. Language Archive Newsletter, 1(3), 4-6.
  • Widlok, T., & Burenhult, N. (2014). Sehen, riechen, orientieren. Spektrum der Wissenschaft, June 2014, 76-81.
  • Willems, R. M., Ozyurek, A., & Hagoort, P. (2007). When language meets action: The neural integration of gesture and speech. Cerebral Cortex, 17(10), 2322-2333. doi:10.1093/cercor/bhl141.

    Abstract

    Although generally studied in isolation, language and action often co-occur in everyday life. Here we investigated one particular form of simultaneous language and action, namely speech and gestures that speakers use in everyday communication. In a functional magnetic resonance imaging study, we identified the neural networks involved in the integration of semantic information from speech and gestures. Verbal and/or gestural content could be integrated easily or less easily with the content of the preceding part of speech. Premotor areas involved in action observation (Brodmann area [BA] 6) were found to be specifically modulated by action information "mismatching" to a language context. Importantly, an increase in integration load of both verbal and gestural information into prior speech context activated Broca's area and adjacent cortex (BA 45/47). A classical language area, Broca's area, is not only recruited for language-internal processing but also when action observation is integrated with speech. These findings provide direct evidence that action and language processing share a high-level neural integration system.
  • 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., 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., & Francken, J. C. (2012). Embodied cognition: Taking the next step. Frontiers in Psychology, 3, 582. doi:10.3389/fpsyg.2012.00582.

    Abstract

    Recent years have seen a large amount of empirical studies related to ‘embodied cognition’. While interesting and valuable, there is something dissatisfying with the current state of affairs in this research domain. Hypotheses tend to be underspecified, testing in general terms for embodied versus disembodied processing. The lack of specificity of current hypotheses can easily lead to an erosion of the embodiment concept, and result in a situation in which essentially any effect is taken as positive evidence. Such erosion is not helpful to the field and does not do justice to the importance of embodiment. Here we want to take stock, and formulate directions for how it can be studied in a more fruitful fashion. As an example we will describe few example studies that have investigated the role of sensori-motor systems in the coding of meaning (‘embodied semantics’). Instead of focusing on the dichotomy between embodied and disembodied theories, we suggest that the field move forward and ask how and when sensori-motor systems and behavior are involved in cognition.
  • Willems, R. M., & Hagoort, P. (2007). Neural evidence for the interplay between language, gesture, and action: A review. Brain and Language, 101(3), 278-289. doi:10.1016/j.bandl.2007.03.004.

    Abstract

    Co-speech gestures embody a form of manual action that is tightly coupled to the language system. As such, the co-occurrence of speech and co-speech gestures is an excellent example of the interplay between language and action. There are, however, other ways in which language and action can be thought of as closely related. In this paper we will give an overview of studies in cognitive neuroscience that examine the neural underpinnings of links between language and action. Topics include neurocognitive studies of motor representations of speech sounds, action-related language, sign language and co-speech gestures. It will be concluded that there is strong evidence on the interaction between speech and gestures in the brain. This interaction however shares general properties with other domains in which there is interplay between language and action.
  • 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. (2007). The neural construction of a Tinkertoy [‘Journal club’ review]. The Journal of Neuroscience, 27, 1509-1510. doi:10.1523/JNEUROSCI.0005-07.2007.
  • 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
  • Winsvold, B. S., Palta, P., Eising, E., Page, C. M., The International Headache Genetics Consortium, Van den Maagdenberg, A. M. J. M., Palotie, A., & Zwart, J.-A. (2018). Epigenetic DNA methylation changes associated with headache chronification: A retrospective case-control study. Cephalalgia, 38(2), 312-322. doi:10.1177/0333102417690111.

    Abstract

    Background

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

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

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

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

    Abstract

    Researchers have suggested that the vocabularies of languages are oriented towards the communicative needs of language users. Here, we provide evidence demonstrating that the higher frequency of visual words in a large variety of English corpora is reflected in greater lexical differentiation—a greater number of unique words—for the visual domain in the English lexicon. In comparison, sensory modalities that are less frequently talked about, particularly taste and smell, show less lexical differentiation. In addition, we show that even though sensory language can be expected to change across historical time and between contexts of use (e.g., spoken language versus fiction), the pattern of visual dominance is a stable property of the English language. Thus, we show that across the board, precisely those semantic domains that are more frequently talked about are also more lexically differentiated, for perceptual experiences. This correlation between type and token frequencies suggests that the sensory lexicon of English is geared towards communicative efficiency.
  • Wirthlin, M., Chang, E. F., Knörnschild, M., Krubitzer, L. A., Mello, C. V., Miller, C. T., Pfenning, A. R., Vernes, S. C., Tchernichovski, O., & Yartsev, M. M. (2019). A modular approach to vocal learning: Disentangling the diversity of a complex behavioral trait. Neuron, 104(1), 87-99. doi:10.1016/j.neuron.2019.09.036.

    Abstract

    Vocal learning is a behavioral trait in which the social and acoustic environment shapes the vocal repertoire of individuals. Over the past century, the study of vocal learning has progressed at the intersection of ecology, physiology, neuroscience, molecular biology, genomics, and evolution. Yet, despite the complexity of this trait, vocal learning is frequently described as a binary trait, with species being classified as either vocal learners or vocal non-learners. As a result, studies have largely focused on a handful of species for which strong evidence for vocal learning exists. Recent studies, however, suggest a continuum in vocal learning capacity across taxa. Here, we further suggest that vocal learning is a multi-component behavioral phenotype comprised of distinct yet interconnected modules. Discretizing the vocal learning phenotype into its constituent modules would facilitate integration of findings across a wider diversity of species, taking advantage of the ways in which each excels in a particular module, or in a specific combination of features. Such comparative studies can improve understanding of the mechanisms and evolutionary origins of vocal learning. We propose an initial set of vocal learning modules supported by behavioral and neurobiological data and highlight the need for diversifying the field in order to disentangle the complexity of the vocal learning phenotype.

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  • 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. (2004). Technology and Tools for Language Documentation. Language Archive Newsletter, 1(4), 3-4.
  • Wittenburg, P. (2004). Training Course in Lithuania. Language Archive Newsletter, 1(2), 6-6.
  • 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.
  • Wittenburg, P., Dirksmeyer, R., Brugman, H., & Klaas, G. (2004). Digital formats for images, audio and video. Language Archive Newsletter, 1(1), 3-6.
  • Wittenburg, P. (2004). International Expert Meeting on Access Management for Distributed Language Archives. Language Archive Newsletter, 1(3), 12-12.
  • Wittenburg, P. (2004). Final review of INTERA. Language Archive Newsletter, 1(4), 11-12.
  • Wittenburg, P. (2004). LinguaPax Forum on Language Diversity, Sustainability, and Peace. Language Archive Newsletter, 1(3), 13-13.
  • Wittenburg, P. (2004). LREC conference 2004. Language Archive Newsletter, 1(3), 12-13.
  • Wittenburg, P. (2004). News from the Archive of the Max Planck Institute for Psycholinguistics. Language Archive Newsletter, 1(4), 12-12.
  • 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.
  • Wolf, M. C., Muijselaar, M. M. L., Boonstra, A. M., & De Bree, E. H. (2019). The relationship between reading and listening comprehension: Shared and modality-specific components. Reading and Writing, 32(7), 1747-1767. doi:10.1007/s11145-018-9924-8.

    Abstract

    This study aimed to increase our understanding on the relationship between reading and listening comprehension. Both in comprehension theory and in educational practice, reading and listening comprehension are often seen as interchangeable, overlooking modality-specific aspects of them separately. Three questions were addressed. First, it was examined to what extent reading and listening comprehension comprise modality-specific, distinct skills or an overlapping, domain-general skill in terms of the amount of explained variance in one comprehension type by the opposite comprehension type. Second, general and modality-unique subskills of reading and listening comprehension were sought by assessing the contributions of the foundational skills word reading fluency, vocabulary, memory, attention, and inhibition to both comprehension types. Lastly, the practice of using either listening comprehension or vocabulary as a proxy of general comprehension was investigated. Reading and listening comprehension tasks with the same format were assessed in 85 second and third grade children. Analyses revealed that reading comprehension explained 34% of the variance in listening comprehension, and listening comprehension 40% of reading comprehension. Vocabulary and word reading fluency were found to be shared contributors to both reading and listening comprehension. None of the other cognitive skills contributed significantly to reading or listening comprehension. These results indicate that only part of the comprehension process is indeed domain-general and not influenced by the modality in which the information is provided. Especially vocabulary seems to play a large role in this domain-general part. The findings warrant a more prominent focus of modality-specific aspects of both reading and listening comprehension in research and education.
  • Womelsdorf, T., Schoffelen, J.-M., Oostenveld, R., Singer, W., Desimone, R., Engel, A. K., & Fries, P. (2007). Modulation of neuronal interactions through neuronal synchronization. Science, 316, 1609-1612. doi:10.1126/science.1139597.

    Abstract

    Brain processing depends on the interactions between neuronal groups. Those interactions are governed by the pattern of anatomical connections and by yet unknown mechanisms that modulate the effective strength of a given connection. We found that the mutual influence among neuronal groups depends on the phase relation between rhythmic activities within the groups. Phase relations supporting interactions between the groups preceded those interactions by a few milliseconds, consistent with a mechanistic role. These effects were specific in time, frequency, and space, and we therefore propose that the pattern of synchronization flexibly determines the pattern of neuronal interactions.
  • Wong, M. M. K., Hoekstra, S. D., Vowles, J., Watson, L. M., Fuller, G., Németh, A. H., Cowley, S. A., Ansorge, O., Talbot, K., & Becker, E. B. E. (2018). Neurodegeneration in SCA14 is associated with increased PKCγ kinase activity, mislocalization and aggregation. Acta Neuropathologica Communications, 6: 99. doi:10.1186/s40478-018-0600-7.

    Abstract

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

    Additional information

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

    Abstract

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

    Abstract

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

    Abstract

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

    Abstract

    L'A. étudie les corrélations entre structure syntaxique et fonction pragmatique dans les alternances de voix en akatek, une langue maya appartenant au sous-groupe Q'anjob'ala. Les alternances pragmatiques de voix sont les mécanismes par lesquels les langues encodent les différents degrés de topicalité des deux principaux participants d'un événement sémantiquement transitif, l'agent et le patient. A l'aide d'une analyse quantitative, l'A. évalue la topicalité de ces participants et identifie les structures syntaxiques permettant d'exprimer les quatre principales fonctions de voix en akatek : active-directe, inverse, passive et antipassive
  • Zeshan, U. (2004). Interrogative constructions in sign languages - Cross-linguistic perspectives. Language, 80(1), 7-39.

    Abstract

    This article reports on results from a broad crosslinguistic study based on data from thirty-five signed languages around the world. The study is the first of its kind, and the typological generalizations presented here cover the domain of interrogative structures as they appear across a wide range of geographically and genetically distinct signed languages. Manual and nonmanual ways of marking basic types of questions in signed languages are investigated. As a result, it becomes clear that the range of crosslinguistic variation is extensive for some subparameters, such as the structure of question-word paradigms, while other parameters, such as the use of nonmanual expressions in questions, show more similarities across signed languages. Finally, it is instructive to compare the findings from signed language typology to relevant data from spoken languages at a more abstract, crossmodality level.
  • Zeshan, U. (2004). Hand, head and face - negative constructions in sign languages. Linguistic Typology, 8(1), 1-58. doi:10.1515/lity.2004.003.

    Abstract

    This article presents a typology of negative constructions across a substantial number of sign languages from around the globe. After situating the topic within the wider context of linguistic typology, the main negation strategies found across sign languages are described. Nonmanual negation includes the use of head movements and facial expressions for negation and is of great importance in sign languages as well as particularly interesting from a typological point of view. As far as manual signs are concerned, independent negative particles represent the dominant strategy, but there are also instances of irregular negation in most sign languages. Irregular negatives may take the form of suppletion, cliticisation, affixing, or internal modification of a sign. The results of the study lead to interesting generalisations about similarities and differences between negatives in signed and spoken languages.
  • Zheng, X., Roelofs, A., Farquhar, J., & Lemhöfer, K. (2018). Monitoring of language selection errors in switching: Not all about conflict. PLoS One, 13(11): e0200397. doi:10.1371/journal.pone.0200397.

    Abstract

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

    Abstract

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

    Additional information

    plcp_a_1363401_sm2537.docx
  • Zheng, X., & Lemhöfer, K. (2019). The “semantic P600” in second language processing: When syntax conflicts with semantics. Neuropsychologia, 127, 131-147. doi:10.1016/j.neuropsychologia.2019.02.010.

    Abstract

    In sentences like “the mouse that chased the cat was hungry”, the syntactically correct interpretation (the mouse chases the cat) is contradicted by semantic and pragmatic knowledge. Previous research has shown that L1 speakers sometimes base sentence interpretation on this type of knowledge (so-called “shallow” or “good-enough” processing). We made use of both behavioural and ERP measurements to investigate whether L2 learners differ from native speakers in the extent to which they engage in “shallow” syntactic processing. German learners of Dutch as well as Dutch native speakers read sentences containing relative clauses (as in the example above) for which the plausible thematic roles were or were not reversed, and made plausibility judgments. The results show that behaviourally, L2 learners had more difficulties than native speakers to discriminate plausible from implausible sentences. In the ERPs, we replicated the previously reported finding of a “semantic P600” for semantic reversal anomalies in native speakers, probably reflecting the effort to resolve the syntax-semantics conflict. In L2 learners, though, this P600 was largely attenuated and surfaced only in those trials that were judged correctly for plausibility. These results generally point at a more prevalent, but not exclusive occurrence of shallow syntactic processing in L2 learners.
  • 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.
  • Zhu, Z., Hagoort, P., Zhang, J. X., Feng, G., Chen, H.-C., Bastiaansen, M. C. M., & Wang, S. (2012). The anterior left inferior frontal gyrus contributes to semantic unification. NeuroImage, 60, 2230-2237. doi:10.1016/j.neuroimage.2012.02.036.

    Abstract

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

    Additional information

    Zhu_2012_suppl.dot
  • Zhu, Z., Bastiaansen, M. C. M., Hakun, J. G., Petersson, K. M., Wang, S., & Hagoort, P. (2019). Semantic unification modulates N400 and BOLD signal change in the brain: A simultaneous EEG-fMRI study. Journal of Neurolinguistics, 52: 100855. doi:10.1016/j.jneuroling.2019.100855.

    Abstract

    Semantic unification during sentence comprehension has been associated with amplitude change of the N400 in event-related potential (ERP) studies, and activation in the left inferior frontal gyrus (IFG) in functional magnetic resonance imaging (fMRI) studies. However, the specificity of this activation to semantic unification remains unknown. To more closely examine the brain processes involved in semantic unification, we employed simultaneous EEG-fMRI to time-lock the semantic unification related N400 change, and integrated trial-by-trial variation in both N400 and BOLD change beyond the condition-level BOLD change difference measured in traditional fMRI analyses. Participants read sentences in which semantic unification load was parametrically manipulated by varying cloze probability. Separately, ERP and fMRI results replicated previous findings, in that semantic unification load parametrically modulated the amplitude of N400 and cortical activation. Integrated EEG-fMRI analyses revealed a different pattern in which functional activity in the left IFG and bilateral supramarginal gyrus (SMG) was associated with N400 amplitude, with the left IFG activation and bilateral SMG activation being selective to the condition-level and trial-level of semantic unification load, respectively. By employing the EEG-fMRI integrated analyses, this study among the first sheds light on how to integrate trial-level variation in language comprehension.
  • Ziegler, A., DeStefano, A. L., König, I. R., Bardel, C., Brinza, D., Bull, S., Cai, Z., Glaser, B., Jiang, W., Lee, K. E., Li, C. X., Li, J., Li, X., Majoram, P., Meng, Y., Nicodemus, K. K., Platt, A., Schwarz, D. F., Shi, W., Shugart, Y. Y. and 7 moreZiegler, A., DeStefano, A. L., König, I. R., Bardel, C., Brinza, D., Bull, S., Cai, Z., Glaser, B., Jiang, W., Lee, K. E., Li, C. X., Li, J., Li, X., Majoram, P., Meng, Y., Nicodemus, K. K., Platt, A., Schwarz, D. F., Shi, W., Shugart, Y. Y., Stassen, H. H., Sun, Y. V., Won, S., Wang, W., Wahba, G., Zagaar, U. A., & Zhao, Z. (2007). Data mining, neural nets, trees–problems 2 and 3 of Genetic Analysis Workshop 15. Genetic Epidemiology, 31(Suppl 1), S51-S60. doi:10.1002/gepi.20280.

    Abstract

    Genome-wide association studies using thousands to hundreds of thousands of single nucleotide polymorphism (SNP) markers and region-wide association studies using a dense panel of SNPs are already in use to identify disease susceptibility genes and to predict disease risk in individuals. Because these tasks become increasingly important, three different data sets were provided for the Genetic Analysis Workshop 15, thus allowing examination of various novel and existing data mining methods for both classification and identification of disease susceptibility genes, gene by gene or gene by environment interaction. The approach most often applied in this presentation group was random forests because of its simplicity, elegance, and robustness. It was used for prediction and for screening for interesting SNPs in a first step. The logistic tree with unbiased selection approach appeared to be an interesting alternative to efficiently select interesting SNPs. Machine learning, specifically ensemble methods, might be useful as pre-screening tools for large-scale association studies because they can be less prone to overfitting, can be less computer processor time intensive, can easily include pair-wise and higher-order interactions compared with standard statistical approaches and can also have a high capability for classification. However, improved implementations that are able to deal with hundreds of thousands of SNPs at a time are required.
  • Zoefel, B., Ten Oever, S., & Sack, A. T. (2018). The involvement of endogenous neural oscillations in the processing of rhythmic input: More than a regular repetition of evoked neural responses. Frontiers in Neuroscience, 12: 95. doi:10.3389/fnins.2018.00095.

    Abstract

    It is undisputed that presenting a rhythmic stimulus leads to a measurable brain response that follows the rhythmic structure of this stimulus. What is still debated, however, is the question whether this brain response exclusively reflects a regular repetition of evoked responses, or whether it also includes entrained oscillatory activity. Here we systematically present evidence in favor of an involvement of entrained neural oscillations in the processing of rhythmic input while critically pointing out which questions still need to be addressed before this evidence could be considered conclusive. In this context, we also explicitly discuss the potential functional role of such entrained oscillations, suggesting that these stimulus-aligned oscillations reflect, and serve as, predictive processes, an idea often only implicitly assumed in the literature.
  • Zora, H., Riad, T., & Ylinen, S. (2019). Prosodically controlled derivations in the mental lexicon. Journal of Neurolinguistics, 52: 100856. doi:10.1016/j.jneuroling.2019.100856.

    Abstract

    Swedish morphemes are classified as prosodically specified or prosodically unspecified, depending on lexical or phonological stress, respectively. Here, we investigate the allomorphy of the suffix -(i)sk, which indicates the distinction between lexical and phonological stress; if attached to a lexically stressed morpheme, it takes a non-syllabic form (-sk), whereas if attached to a phonologically stressed morpheme, an epenthetic vowel is inserted (-isk). Using mismatch negativity (MMN), we explored the neural processing of this allomorphy across lexically stressed and phonologically stressed morphemes. In an oddball paradigm, participants were occasionally presented with congruent and incongruent derivations, created by the suffix -(i)sk, within the repetitive presentation of their monomorphemic stems. The results indicated that the congruent derivation of the lexically stressed stem elicited a larger MMN than the incongruent sequences of the same stem and the derivational suffix, whereas after the phonologically stressed stem a non-significant tendency towards an opposite pattern was observed. We argue that the significant MMN response to the congruent derivation in the lexical stress condition is in line with lexical MMN, indicating a holistic processing of the sequence of lexically stressed stem and derivational suffix. The enhanced MMN response to the incongruent derivation in the phonological stress condition, on the other hand, is suggested to reflect combinatorial processing of the sequence of phonologically stressed stem and derivational suffix. These findings bring a new aspect to the dual-system approach to neural processing of morphologically complex words, namely the specification of word stress.
  • Zormpa, E., Meyer, A. S., & Brehm, L. (2019). Slow naming of pictures facilitates memory for their names. Psychonomic Bulletin & Review, 26(5), 1675-1682. doi:10.3758/s13423-019-01620-x.

    Abstract

    Speakers remember their own utterances better than those of their interlocutors, suggesting that language production is beneficial to memory. This may be partly explained by a generation effect: The act of generating a word is known to lead to a memory advantage (Slamecka & Graf, 1978). In earlier work, we showed a generation effect for recognition of images (Zormpa, Brehm, Hoedemaker, & Meyer, 2019). Here, we tested whether the recognition of their names would also benefit from name generation. Testing whether picture naming improves memory for words was our primary aim, as it serves to clarify whether the representations affected by generation are visual or conceptual/lexical. A secondary aim was to assess the influence of processing time on memory. Fifty-one participants named pictures in three conditions: after hearing the picture name (identity condition), backward speech, or an unrelated word. A day later, recognition memory was tested in a yes/no task. Memory in the backward speech and unrelated conditions, which required generation, was superior to memory in the identity condition, which did not require generation. The time taken by participants for naming was a good predictor of memory, such that words that took longer to be retrieved were remembered better. Importantly, that was the case only when generation was required: In the no-generation (identity) condition, processing time was not related to recognition memory performance. This work has shown that generation affects conceptual/lexical representations, making an important contribution to the understanding of the relationship between memory and language.
  • Zormpa, E., Brehm, L., Hoedemaker, R. S., & Meyer, A. S. (2019). The production effect and the generation effect improve memory in picture naming. Memory, 27(3), 340-352. doi:10.1080/09658211.2018.1510966.

    Abstract

    The production effect (better memory for words read aloud than words read silently) and the picture superiority effect (better memory for pictures than words) both improve item memory in a picture naming task (Fawcett, J. M., Quinlan, C. K., & Taylor, T. L. (2012). Interplay of the production and picture superiority effects: A signal detection analysis. Memory (Hove, England), 20(7), 655–666. doi:10.1080/09658211.2012.693510). Because picture naming requires coming up with an appropriate label, the generation effect (better memory for generated than read words) may contribute to the latter effect. In two forced-choice memory experiments, we tested the role of generation in a picture naming task on later recognition memory. In Experiment 1, participants named pictures silently or aloud with the correct name or an unreadable label superimposed. We observed a generation effect, a production effect, and an interaction between the two. In Experiment 2, unreliable labels were included to ensure full picture processing in all conditions. In this experiment, we observed a production and a generation effect but no interaction, implying the effects are dissociable. This research demonstrates the separable roles of generation and production in picture naming and their impact on memory. As such, it informs the link between memory and language production and has implications for memory asymmetries between language production and comprehension.

    Additional information

    pmem_a_1510966_sm9257.pdf
  • 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.
  • Zwaan, R. A., Van der Stoep, N., Guadalupe, T., & Bouwmeester, S. (2012). Language comprehension in the balance: The robustness of the action-compatibility effect (ACE). PLoS One, 7(2), e31204. doi:10.1371/journal.pone.0031204.

    Abstract

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

    Abstract

    This paper explores the expression of multiple entities in Turkish Sign Language (Türk İşaret Dili; TİD), a less well-studied sign language. It aims to provide a comprehensive description of the ways and frequencies in which entity plurality in this language is expressed, both within and outside the noun phrase. We used a corpus that includes both elicited and spontaneous data from native signers. The results reveal that most of the expressions of multiple entities in TİD are iconic, spatial strategies (i.e. localization and spatial plural predicate inflection) none of which, we argue, should be considered as genuine plural marking devices with the main aim of expressing plurality. Instead, the observed devices for localization and predicate inflection allow for a plural interpretation when multiple locations in space are used. Our data do not provide evidence that TİD employs (productive) morphological plural marking (i.e. reduplication) on nouns, in contrast to some other sign languages and many spoken languages. We relate our findings to expression of multiple entities in other signed languages and in spoken languages and discuss these findings in terms of modality effects on expression of multiple entities in human language.
  • Zwitserlood, I. (2010). Laat je vingers spreken: NGT en vingerspelling. Levende Talen Magazine, 2010(2), 46-47.
  • Zwitserlood, I. (2010). Het Corpus NGT en de dagelijkse lespraktijk (2). Levende Talen Magazine, 2010(3), 47-48.
  • 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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