Publications

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  • Willems, R. M., Toni, I., Hagoort, P., & Casasanto, D. (2009). Body-specific motor imagery of hand actions: Neural evidence from right- and left-handers. Frontiers in Human Neuroscience, 3: 39, pp. 39. doi:10.3389/neuro.09.039.2009.

    Abstract

    If motor imagery uses neural structures involved in action execution, then the neural correlates of imagining an action should differ between individuals who tend to execute the action differently. Here we report fMRI data showing that motor imagery is influenced by the way people habitually perform motor actions with their particular bodies; that is, motor imagery is ‘body-specific’ (Casasanto, 2009). During mental imagery for complex hand actions, activation of cortical areas involved in motor planning and execution was left-lateralized in right-handers but right-lateralized in left-handers. We conclude that motor imagery involves the generation of an action plan that is grounded in the participant’s motor habits, not just an abstract representation at the level of the action’s goal. People with different patterns of motor experience form correspondingly different neurocognitive representations of imagined actions.
  • Willems, R. M., & Hagoort, P. (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., Ozyurek, A., & Hagoort, P. (2009). Differential roles for left inferior frontal and superior temporal cortex in multimodal integration of action and language. Neuroimage, 47, 1992-2004. doi:10.1016/j.neuroimage.2009.05.066.

    Abstract

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

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

    Additional information

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

    Abstract

    It has been argued that we map observed actions onto our own motor system. Here we added to this issue by investigating whether hand preference influences the neural correlates of action observation of simple, essentially meaningless hand actions. Such an influence would argue for an intricate neural coupling between action production and action observation, which goes beyond effects of motor repertoire or explicit motor training, as has been suggested before. Indeed, parts of the human motor system exhibited a close coupling between action production and action observation. Ventral premotor and inferior and superior parietal cortices showed differential activation for left- and right-handers that was similar during action production as well as during action observation. This suggests that mapping observed actions onto the observer's own motor system is a core feature of action observation - at least for actions that do not have a clear goal or meaning. Basic differences in the way we act upon the world are not only reflected in neural correlates of action production, but can also influence the brain basis of action observation.
  • Willems, R. M., Van der Haegen, L., Fisher, S. E., & Francks, C. (2014). On the other hand: Including left-handers in cognitive neuroscience and neurogenetics. Nature Reviews Neuroscience, 15, 193-201. doi:10.1038/nrn3679.

    Abstract

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

    Abstract

    While most people prefer to use their right hand to brush their teeth, throw a ball, or hold a tennis racket, left-handers prefer to use their left hand. This is the case for around 10 per cent of all people. There was a time (not so long ago) when left-handers were stigmatized in Western (and other) communities: it was considered a bad sign if you were left-handed, and left-handed children were often forced to write with their right hand. This is nonsensical: there is nothing wrong with being left-handed, and trying to write with the non-preferred hand is frustrating for almost everybody. As a matter of fact, science can learn from left-handers, and in this paper, we discuss how this may be the case. We review why some people are left-handed and others are not, how left-handers' brains differ from right-handers’, and why scientists study left-handedness in the first place
  • Witteman, M. J., 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.
  • 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.
  • 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.
  • 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
  • Zhang, Y., Ding, R., Frassinelli, D., Tuomainen, J., Klavinskis-Whiting, S., & Vigliocco, G. (2023). The role of multimodal cues in second language comprehension. Scientific Reports, 13: 20824. doi:10.1038/s41598-023-47643-2.

    Abstract

    In face-to-face communication, multimodal cues such as prosody, gestures, and mouth movements can play a crucial role in language processing. While several studies have addressed how these cues contribute to native (L1) language processing, their impact on non-native (L2) comprehension is largely unknown. Comprehension of naturalistic language by L2 comprehenders may be supported by the presence of (at least some) multimodal cues, as these provide correlated and convergent information that may aid linguistic processing. However, it is also the case that multimodal cues may be less used by L2 comprehenders because linguistic processing is more demanding than for L1 comprehenders, leaving more limited resources for the processing of multimodal cues. In this study, we investigated how L2 comprehenders use multimodal cues in naturalistic stimuli (while participants watched videos of a speaker), as measured by electrophysiological responses (N400) to words, and whether there are differences between L1 and L2 comprehenders. We found that prosody, gestures, and informative mouth movements each reduced the N400 in L2, indexing easier comprehension. Nevertheless, L2 participants showed weaker effects for each cue compared to L1 comprehenders, with the exception of meaningful gestures and informative mouth movements. These results show that L2 comprehenders focus on specific multimodal cues – meaningful gestures that support meaningful interpretation and mouth movements that enhance the acoustic signal – while using multimodal cues to a lesser extent than L1 comprehenders overall.

    Additional information

    supplementary materials
  • Wu, S., Zhao, J., de Villiers, J., Liu, X. L., Rolfhus, E., Sun, X. N., Li, X. Y., Pan, H., Wang, H. W., Zhu, Q., Dong, Y. Y., Zhang, Y. T., & Jiang, F. (2023). Prevalence, co-occurring difficulties, and risk factors of developmental language disorder: First evidence for Mandarin-speaking children in a population-based study. The Lancet Regional Health - Western Pacific, 34: 100713. doi:10.1016/j.lanwpc.2023.100713.

    Abstract

    Background: Developmental language disorder (DLD) is a condition that significantly affects children's achievement but has been understudied. We aim to estimate the prevalence of DLD in Shanghai, compare the co-occurrence of difficulties between children with DLD and those with typical development (TD), and investigate the early risk factors for DLD.

    Methods: We estimated DLD prevalence using data from a population-based survey with a cluster random sampling design in Shanghai, China. A subsample of children (aged 5-6 years) received an onsite evaluation, and each child was categorized as TD or DLD. The proportions of children with socio-emotional behavior (SEB) difficulties, low non-verbal IQ (NVIQ), and poor school readiness were calculated among children with TD and DLD. We used multiple imputation to address the missing values of risk factors. Univariate and multivariate regression models adjusted with sampling weights were used to estimate the correlation of each risk factor with DLD.

    Findings: Of 1082 children who were approached for the onsite evaluation, 974 (90.0%) completed the language ability assessments, of whom 74 met the criteria for DLD, resulting in a prevalence of 8.5% (95% CI 6.3-11.5) when adjusted with sampling weights. Compared with TD children, children with DLD had higher rates of concurrent difficulties, including SEB (total difficulties score at-risk: 156 [17.3%] of 900 TD vs. 28 [37.8%] of 74 DLD, p < 0.0001), low NVIQ (3 [0.3%] of 900 TD vs. 8 [10.8%] of 74 DLD, p < 0.0001), and poor school readiness (71 [7.9%] of 900 TD vs. 13 [17.6%] of 74 DLD, p = 0.0040). After accounting for all other risk factors, a higher risk of DLD was associated with a lack of parent-child interaction diversity (adjusted odds ratio [aOR] = 3.08, 95% CI = 1.29-7.37; p = 0.012) and lower kindergarten levels (compared to demonstration and first level: third level (aOR = 6.15, 95% CI = 1.92-19.63; p = 0.0020)).

    Interpretation: The prevalence of DLD and its co-occurrence with other difficulties suggest the need for further attention. Family and kindergarten factors were found to contribute to DLD, suggesting that multi-sector coordinated efforts are needed to better identify and serve DLD populations at home, in schools, and in clinical settings.

    Funding: The study was supported by Shanghai Municipal Education Commission (No. 2022you1-2, D1502), the Innovative Research Team of High-level Local Universities in Shanghai (No. SHSMU-ZDCX20211900), Shanghai Municipal Health Commission (No.GWV-10.1-XK07), and the National Key Research and Development Program of China (No. 2022YFC2705201).
  • Zioga, I., Weissbart, H., Lewis, A. G., Haegens, S., & Martin, A. E. (2023). Naturalistic spoken language comprehension is supported by alpha and beta oscillations. The Journal of Neuroscience, 43(20), 3718-3732. doi:10.1523/JNEUROSCI.1500-22.2023.

    Abstract

    Brain oscillations are prevalent in all species and are involved in numerous perceptual operations. α oscillations are thought to facilitate processing through the inhibition of task-irrelevant networks, while β oscillations are linked to the putative reactivation of content representations. Can the proposed functional role of α and β oscillations be generalized from low-level operations to higher-level cognitive processes? Here we address this question focusing on naturalistic spoken language comprehension. Twenty-two (18 female) Dutch native speakers listened to stories in Dutch and French while MEG was recorded. We used dependency parsing to identify three dependency states at each word: the number of (1) newly opened dependencies, (2) dependencies that remained open, and (3) resolved dependencies. We then constructed forward models to predict α and β power from the dependency features. Results showed that dependency features predict α and β power in language-related regions beyond low-level linguistic features. Left temporal, fundamental language regions are involved in language comprehension in α, while frontal and parietal, higher-order language regions, and motor regions are involved in β. Critically, α- and β-band dynamics seem to subserve language comprehension tapping into syntactic structure building and semantic composition by providing low-level mechanistic operations for inhibition and reactivation processes. Because of the temporal similarity of the α-β responses, their potential functional dissociation remains to be elucidated. Overall, this study sheds light on the role of α and β oscillations during naturalistic spoken language comprehension, providing evidence for the generalizability of these dynamics from perceptual to complex linguistic processes.
  • Zora, H., Wester, J. M., & Csépe, V. (2023). Predictions about prosody facilitate lexical access: Evidence from P50/N100 and MMN components. International Journal of Psychophysiology, 194: 112262. doi:10.1016/j.ijpsycho.2023.112262.

    Abstract

    Research into the neural foundation of perception asserts a model where top-down predictions modulate the bottom-up processing of sensory input. Despite becoming increasingly influential in cognitive neuroscience, the precise account of this predictive coding framework remains debated. In this study, we aim to contribute to this debate by investigating how predictions about prosody facilitate speech perception, and to shed light especially on lexical access influenced by simultaneous predictions in different domains, inter alia, prosodic and semantic. Using a passive auditory oddball paradigm, we examined neural responses to prosodic changes, leading to a semantic change as in Dutch nouns canon [ˈkaːnɔn] ‘cannon’ vs kanon [kaːˈnɔn] ‘canon’, and used acoustically identical pseudowords as controls. Results from twenty-eight native speakers of Dutch (age range 18–32 years) indicated an enhanced P50/N100 complex to prosodic change in pseudowords as well as an MMN response to both words and pseudowords. The enhanced P50/N100 response to pseudowords is claimed to indicate that all relevant auditory information is still processed by the brain, whereas the reduced response to words might reflect the suppression of information that has already been encoded. The MMN response to pseudowords and words, on the other hand, is best justified by the unification of previously established prosodic representations with sensory and semantic input respectively. This pattern of results is in line with the predictive coding framework acting on multiple levels and is of crucial importance to indicate that predictions about linguistic prosodic information are utilized by the brain as early as 50 ms.
  • Zormpa, E., Meyer, A. S., & Brehm, L. (2023). In conversation, answers are remembered better than the questions themselves. Journal of Experimental Psychology: Learning, Memory, and Cognition, 49(12), 1971-1988. doi:10.1037/xlm0001292.

    Abstract

    Language is used in communicative contexts to identify and successfully transmit new information that should be later remembered. In three studies, we used question–answer pairs, a naturalistic device for focusing information, to examine how properties of conversations inform later item memory. In Experiment 1, participants viewed three pictures while listening to a recorded question–answer exchange between two people about the locations of two of the displayed pictures. In a memory recognition test conducted online a day later, participants recognized the names of pictures that served as answers more accurately than the names of pictures that appeared as questions. This suggests that this type of focus indeed boosts memory. In Experiment 2, participants listened to the same items embedded in declarative sentences. There was a reduced memory benefit for the second item, confirming the role of linguistic focus on later memory beyond a simple serial-position effect. In Experiment 3, two participants asked and answered the same questions about objects in a dialogue. Here, answers continued to receive a memory benefit, and this focus effect was accentuated by language production such that information-seekers remembered the answers to their questions better than information-givers remembered the questions they had been asked. Combined, these studies show how people’s memory for conversation is modulated by the referential status of the items mentioned and by the speaker’s roles of the conversation participants.
  • De Zubicaray, G. I., Hartsuiker, R. J., & Acheson, D. J. (2014). Mind what you say—general and specific mechanisms for monitoring in speech production. Frontiers in Human Neuroscience, 8: 514. doi:10.3389%2Ffnhum.2014.00514.

    Abstract

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

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

    Abstract

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

    Abstract

    The Corpus NGT
  • Zwitserlood, I. (2009). Het Corpus NGT en de dagelijkse lespraktijk (1). Levende Talen Magazine, 8, 40-41.

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