Publications

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  • Willems, R. M., & Hagoort, P. (2009). Broca's region: Battles are not won by ignoring half of the facts. Trends in Cognitive Sciences, 13(3), 101. doi:10.1016/j.tics.2008.12.001.
  • Willems, R. M., Peelen, M. V., & Hagoort, P. (2010). Cerebral lateralization of face-selective and body-selective visual areas depends on handedness. Cerebral Cortex, 20, 1719-1725. doi:10.1093/cercor/bhp234.

    Abstract

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

    Abstract

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

    Abstract

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

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

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

    Abstract

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

    Abstract

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

    Abstract

    The human capacity to communicate has been hypothesized to be causally dependent upon language. Intuitively this seems plausible since most communication relies on language. Moreover, intention recognition abilities (as a necessary prerequisite for communication) and language development seem to co-develop. Here we review evidence from neuroimaging as well as from neuropsychology to evaluate the relationship between communicative and linguistic abilities. Our review indicates that communicative abilities are best considered as neurally distinct from language abilities. This conclusion is based upon evidence showing that humans rely on different cortical systems when designing a communicative message for someone else as compared to when performing core linguistic tasks, as well as upon observations of individuals with severe language loss after extensive lesions to the language system, who are still able to perform tasks involving intention understanding
  • Willems, R. M. (2009). Neural reflections of meaning in gesture, language, and action. PhD Thesis, Radboud University Nijmegen, Nijmegen.
  • 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.
  • 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.
  • 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.
  • 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. (2012). The language networks of the brain. PhD Thesis, Radboud University Nijmegen, Nijmegen.

    Abstract

    In recent decades, neuroimaging studies on the neural infrastructure of language are usually (or mostly) conducted with certain on-line language processing tasks. These functional neuroimaging studies helped to localize the language areas in the brain and to investigate the brain activity during explicit language processing. However, little is known about what is going on with the language areas when the brain is ‘at rest’, i.e., when there is no explicit language processing running. Taking advantage of the fcMRI and DTI techniques, this thesis is able to investigate the language function ‘off-line’ at the neuronal network level and the connectivity among language areas in the brain. Based on patient studies, the traditional, classical model on the perisylvian language network specifies a “Broca’ area – Arcuate Fasciculus – Werinicke’s area” loop (Ojemann 1991). With the help of modern neuroimaging techniques, researchers have been able to track language pathways that involve more brain structures than are in the classical model, and relate them to certain language functions. In such a background, a large part of this thesis made a contribution to the study of the topology of the language networks. It revealed that the language networks form a topographical functional connectivity pattern in the left hemisphere for the right-handers. This thesis also revealed the importance of structural hubs, such as Broca’s and Wernicke’s areas, which have more connectivity to other brain areas and play a central role in the language networks. Furthermore, this thesis revealed both functionally and structurally lateralized language networks in the brain. The consistency between what is found in this thesis and what has been known from previous functional studies seems to suggest, that the human brain is optimized and ‘ready’ for the language function even when there is currently no explicit language-processing running.
  • 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.
  • 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.
  • 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.

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    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.
  • 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.

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    pmem_a_1510966_sm9257.pdf
  • 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. (2009). Het Corpus NGT. Levende Talen Magazine, 6, 44-45.

    Abstract

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

    Abstract

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

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