Publications

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  • 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.
  • Yoshihara, M., Nakayama, M., Verdonschot, R. G., & Hino, Y. (2020). The influence of orthography on speech production: Evidence from masked priming in word-naming and picture-naming tasks. Journal of Experimental Psychology: Learning, Memory, and Cognition, 46(8), 1570-1589. doi:10.1037/xlm0000829.

    Abstract

    In a masked priming word-naming task, a facilitation due to the initial-segmental sound overlap for 2-character kanji prime-target pairs was affected by certain orthographic properties (Yoshihara, Nakayama, Verdonschot, & Hino, 2017). That is, the facilitation that was due to the initial mora overlap occurred only when the mora was the whole pronunciation of their initial kanji characters (i.e., match pairs; e.g., /ka-se.ki/-/ka-rjo.ku/). When the shared initial mora was only a part of the kanji characters' readings, however, there was no facilitation (i.e., mismatch pairs; e.g., /ha.tu-a.N/-/ha.ku-bu.tu/). In the present study, we used a masked priming picture-naming task to investigate whether the previous results were relevant only when the orthography of targets is visually presented. In Experiment 1. the main findings of our word-naming task were fully replicated in a picture-naming task. In Experiments 2 and 3. the absence of facilitation for the mismatch pairs were confirmed with a new set of stimuli. On the other hand, a significant facilitation was observed for the match pairs that shared the 2 initial morae (in Experiment 4), which was again consistent with the results of our word-naming study. These results suggest that the orthographic properties constrain the phonological expression of masked priming for kanji words across 2 tasks that are likely to differ in how phonology is retrieved. Specifically, we propose that orthography of a word is activated online and constrains the phonological encoding processes in these tasks.
  • 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., & Lemhöfer, K. (2020). Language selection contributes to intrusion errors in speaking: Evidence from picture naming. Bilingualism: Language and Cognition, 23, 788-800. doi:10.1017/S1366728919000683.

    Abstract

    Bilinguals usually select the right language to speak for the particular context they are in, but sometimes the nontarget language intrudes. Despite a large body of research into language selection and language control, it remains unclear where intrusion errors originate from. These errors may be due to incorrect selection of the nontarget language at the conceptual level, or be a consequence of erroneous word selection (despite correct language selection) at the lexical level. We examined the former possibility in two language switching experiments using a manipulation that supposedly affects language selection on the conceptual level, namely whether the conversational language context was associated with the target language (congruent) or with the alternative language (incongruent) on a trial. Both experiments showed that language intrusion errors occurred more often in incongruent than in congruent contexts, providing converging evidence that language selection during concept preparation is one driving force behind language intrusion.
  • Zheng, X., Roelofs, A., Erkan, H., & Lemhöfer, K. (2020). Dynamics of inhibitory control during bilingual speech production: An electrophysiological study. Neuropsychologia, 140: 107387. doi:10.1016/j.neuropsychologia.2020.107387.

    Abstract

    Bilingual speakers have to control their languages to avoid interference, which may be achieved by enhancing the target language and/or inhibiting the nontarget language. Previous research suggests that bilinguals use inhibition (e.g., Jackson et al., 2001), which should be reflected in the N2 component of the event-related potential (ERP) in the EEG. In the current study, we investigated the dynamics of inhibitory control by measuring the N2 during language switching and repetition in bilingual picture naming. Participants had to name pictures in Dutch or English depending on the cue. A run of same-language trials could be short (two or three trials) or long (five or six trials). We assessed whether RTs and N2 changed over the course of same-language runs, and at a switch between languages. Results showed that speakers named pictures more quickly late as compared to early in a run of same-language trials. Moreover, they made a language switch more quickly after a long run than after a short run. This run-length effect was only present in the first language (L1), not in the second language (L2). In ERPs, we observed a widely distributed switch effect in the N2, which was larger after a short run than after a long run. This effect was only present in the L2, not in the L1, although the difference was not significant between languages. In contrast, the N2 was not modulated during a same-language run. Our results suggest that the nontarget language is inhibited at a switch, but not during the repeated use of the target language.

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  • 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.

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    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.
  • 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.
  • 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., Rudner, M., & Montell Magnusson, A. (2020). Concurrent affective and linguistic prosody with the same emotional valence elicits a late positive ERP response. European Journal of Neuroscience, 51(11), 2236-2249. doi:10.1111/ejn.14658.

    Abstract

    Change in linguistic prosody generates a mismatch negativity response (MMN), indicating neural representation of linguistic prosody, while change in affective prosody generates a positive response (P3a), reflecting its motivational salience. However, the neural response to concurrent affective and linguistic prosody is unknown. The present paper investigates the integration of these two prosodic features in the brain by examining the neural response to separate and concurrent processing by electroencephalography (EEG). A spoken pair of Swedish words—[ˈfɑ́ːsɛn] phase and [ˈfɑ̀ːsɛn] damn—that differed in emotional semantics due to linguistic prosody was presented to 16 subjects in an angry and neutral affective prosody using a passive auditory oddball paradigm. Acoustically matched pseudowords—[ˈvɑ́ːsɛm] and [ˈvɑ̀ːsɛm]—were used as controls. Following the constructionist concept of emotions, accentuating the conceptualization of emotions based on language, it was hypothesized that concurrent affective and linguistic prosody with the same valence—angry [ˈfɑ̀ːsɛn] damn—would elicit a unique late EEG signature, reflecting the temporal integration of affective voice with emotional semantics of prosodic origin. In accordance, linguistic prosody elicited an MMN at 300–350 ms, and affective prosody evoked a P3a at 350–400 ms, irrespective of semantics. Beyond these responses, concurrent affective and linguistic prosody evoked a late positive component (LPC) at 820–870 ms in frontal areas, indicating the conceptualization of affective prosody based on linguistic prosody. This study provides evidence that the brain does not only distinguish between these two functions of prosody but also integrates them based on language and experience.
  • 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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  • 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.
  • Zuidema, W., French, R. M., Alhama, R. G., Ellis, K., O'Donnell, T. J. O., Sainburgh, T., & Gentner, T. Q. (2020). Five ways in which computational modeling can help advance cognitive science: Lessons from artificial grammar learning. Topics in Cognitive Science, 12(3), 925-941. doi:10.1111/tops.12474.

    Abstract

    There is a rich tradition of building computational models in cognitive science, but modeling, theoretical, and experimental research are not as tightly integrated as they could be. In this paper, we show that computational techniques—even simple ones that are straightforward to use—can greatly facilitate designing, implementing, and analyzing experiments, and generally help lift research to a new level. We focus on the domain of artificial grammar learning, and we give five concrete examples in this domain for (a) formalizing and clarifying theories, (b) generating stimuli, (c) visualization, (d) model selection, and (e) exploring the hypothesis space.
  • 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.

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