Publications

Displaying 401 - 500 of 1006
  • Soutschek, A., Burke, C. J., Beharelle, A. R., Schreiber, R., Weber, S. C., Karipidis, I. I., Ten Velden, J., Weber, B., Haker, H., Kalenscher, T., & Tobler, P. N. (2017). The dopaminergic reward system underpins gender differences in social preferences. Nature Human Behaviour, 1, 819-827. doi:10.1038/s41562-017-0226-y.

    Abstract

    Women are known to have stronger prosocial preferences than men, but it remains an open question as to how these behavioural differences arise from differences in brain functioning. Here, we provide a neurobiological account for the hypothesized gender difference. In a pharmacological study and an independent neuroimaging study, we tested the hypothesis that the neural reward system encodes the value of sharing money with others more strongly in women than in men. In the pharmacological study, we reduced receptor type-specific actions of dopamine, a neurotransmitter related to reward processing, which resulted in more selfish decisions in women and more prosocial decisions in men. Converging findings from an independent neuroimaging study revealed gender-related activity in neural reward circuits during prosocial decisions. Thus, the neural reward system appears to be more sensitive to prosocial rewards in women than in men, providing a neurobiological account for why women often behave more prosocially than men.

    A large body of evidence suggests that women are often more prosocial (for example, generous, altruistic and inequality averse) than men, at least when other factors such as reputation and strategic considerations are excluded1,2,3. This dissociation could result from cultural expectations and gender stereotypes, because in Western societies women are more strongly expected to be prosocial4,5,6 and sensitive to variations in social context than men1. It remains an open question, however, whether and how on a neurobiological level the social preferences of women and men arise from differences in brain functioning. The assumption of gender differences in social preferences predicts that the neural reward system’s sensitivity to prosocial and selfish rewards should differ between women and men. Specifically, the hypothesis would be that the neural reward system is more sensitive to prosocial than selfish rewards in women and more sensitive to selfish than prosocial rewards in men. The goal of the current study was to test in two independent experiments for the hypothesized gender differences on both a pharmacological and a haemodynamic level. In particular, we examined the functions of the neurotransmitter dopamine using a dopamine receptor antagonist, and the role of the striatum (a brain region strongly innervated by dopamine neurons) during social decision-making in women and men using neuroimaging.

    The neurotransmitter dopamine is thought to play a key role in neural reward processing7,8. Recent evidence suggests that dopaminergic activity is sensitive not only to rewards for oneself but to rewards for others as well9. The assumption that dopamine is sensitive to both self- and other-related outcomes is consistent with the finding that the striatum shows activation for both selfish and shared rewards10,11,12,13,14,15. The dopaminergic response may represent a net signal encoding the difference between the value of preferred and unpreferred rewards8. Regarding the hypothesized gender differences in social preferences, this account makes the following predictions. If women prefer shared (prosocial) outcomes2, women’s dopaminergic signals to shared rewards will be stronger than to non-shared (selfish) rewards, so reducing dopaminergic activity should bias women to make more selfish decisions. In line with this hypothesis, a functional imaging study reported enhanced striatal activation in female participants during charitable donations11. In contrast, if men prefer selfish over prosocial rewards, dopaminergic activity should be enhanced to selfish compared to prosocial rewards. In line with this view, upregulating dopaminergic activity in a sample of exclusively male participants increased selfish behaviour in a bargaining game16. Thus, contrary to the hypothesized effect in women, reducing dopaminergic neurotransmission should render men more prosocial. Taken together, the current study tested the following three predictions: we expected the dopaminergic reward system (1) to be more sensitive to prosocial than selfish rewards in women and (2) to be more sensitive to selfish than prosocial rewards in men. As a consequence of these two predictions, we also predicted (3) dopaminoceptive regions such as the striatum to show stronger activation to prosocial relative to selfish rewards in women than in men.

    To test these predictions, we conducted a pharmacological study in which we reduced dopaminergic neurotransmission with amisulpride. Amisulpride is a dopamine antagonist that is highly specific for dopaminergic D2/D3 receptors17. After receiving amisulpride or placebo, participants performed an interpersonal decision task18,19,20, in which they made choices between a monetary reward only for themselves (selfish reward option) and sharing money with others (prosocial reward option). We expected that blocking dopaminergic neurotransmission with amisulpride, relative to placebo, would result in fewer prosocial choices in women and more prosocial choices in men. To investigate whether potential gender-related effects of dopamine are selective for social decision-making, we also tested the effects of amisulpride on time preferences in a non-social control task that was matched to the interpersonal decision task in terms of choice structure.

    In addition, because dopaminergic neurotransmission plays a crucial role in brain regions involved in value processing, such as the striatum21, a gender-related role of dopaminergic activity for social decision-making should also be reflected by dissociable activity patterns in the striatum. Therefore, to further test our hypothesis, we investigated the neural correlates of social decision-making in a functional imaging study. In line with our predictions for the pharmacological study, we expected to find stronger striatum activity during prosocial relative to selfish decisions in women, whereas men should show enhanced activity in the striatum for selfish relative to prosocial choices.

    Additional information

    Supplementary Information
  • Ye, Z., Stolk, A., Toni, I., & Hagoort, P. (2017). Oxytocin modulates semantic integration in speech comprehension. Journal of Cognitive Neuroscience, 29, 267-276. doi:10.1162/jocn_a_01044.

    Abstract

    Listeners interpret utterances by integrating information from multiple sources including word level semantics and world knowledge. When the semantics of an expression is inconsistent with his or her knowledge about the world, the listener may have to search through the conceptual space for alternative possible world scenarios that can make the expression more acceptable. Such cognitive exploration requires considerable computational resources and might depend on motivational factors. This study explores whether and how oxytocin, a neuropeptide known to influence socialmotivation by reducing social anxiety and enhancing affiliative tendencies, can modulate the integration of world knowledge and sentence meanings. The study used a betweenparticipant double-blind randomized placebo-controlled design. Semantic integration, indexed with magnetoencephalography through the N400m marker, was quantified while 45 healthymale participants listened to sentences that were either congruent or incongruent with facts of the world, after receiving intranasally delivered oxytocin or placebo. Compared with congruent sentences, world knowledge incongruent sentences elicited a stronger N400m signal from the left inferior frontal and anterior temporal regions and medial pFC (the N400m effect) in the placebo group. Oxytocin administration significantly attenuated the N400meffect at both sensor and cortical source levels throughout the experiment, in a state-like manner. Additional electrophysiological markers suggest that the absence of the N400m effect in the oxytocin group is unlikely due to the lack of early sensory or semantic processing or a general downregulation of attention. These findings suggest that oxytocin drives listeners to resolve challenges of semantic integration, possibly by promoting the cognitive exploration of alternative possible world scenarios.
  • Takashima, A., Bakker, I., Van Hell, J. G., Janzen, G., & McQueen, J. M. (2017). Interaction between episodic and semantic memory networks in the acquisition and consolidation of novel spoken words. Brain and Language, 167, 44-60. doi:10.1016/j.bandl.2016.05.009.

    Abstract

    When a novel word is learned, its memory representation is thought to undergo a process of consolidation and integration. In this study, we tested whether the neural representations of novel words change as a function of consolidation by observing brain activation patterns just after learning and again after a delay of one week. Words learned with meanings were remembered better than those learned without meanings. Both episodic (hippocampus-dependent) and semantic (dependent on distributed neocortical areas) memory systems were utilised during recognition of the novel words. The extent to which the two systems were involved changed as a function of time and the amount of associated information, with more involvement of both systems for the meaningful words than for the form-only words after the one-week delay. These results suggest that the reason the meaningful words were remembered better is that their retrieval can benefit more from these two complementary memory systems
  • Tan, Y., Martin, R. C., & Van Dyke, J. A. (2017). Semantic and syntactic interference in sentence comprehension: A comparison of working memory models. Frontiers in Psychology, 8: 198. doi:10.3389/fpsyg.2017.00198.

    Abstract

    This study investigated the nature of the underlying working memory system supporting sentence processing through examining individual differences in sensitivity to retrieval interference effects during sentence comprehension. Interference effects occur when readers incorrectly retrieve sentence constituents which are similar to those required during integrative processes. We examined interference arising from a partial match between distracting constituents and syntactic and semantic cues, and related these interference effects to performance on working memory, short-term memory (STM), vocabulary, and executive function tasks. For online sentence comprehension, as measured by self-paced reading, the magnitude of individuals' syntactic interference effects was predicted by general WM capacity and the relation remained significant when partialling out vocabulary, indicating that the effects were not due to verbal knowledge. For offline sentence comprehension, as measured by responses to comprehension questions, both general WM capacity and vocabulary knowledge interacted with semantic interference for comprehension accuracy, suggesting that both general WM capacity and the quality of semantic representations played a role in determining how well interference was resolved offline. For comprehension question reaction times, a measure of semantic STM capacity interacted with semantic but not syntactic interference. However, a measure of phonological capacity (digit span) and a general measure of resistance to response interference (Stroop effect) did not predict individuals' interference resolution abilities in either online or offline sentence comprehension. The results are discussed in relation to the multiple capacities account of working memory (e.g., Martin and Romani, 1994; Martin and He, 2004), and the cue-based retrieval parsing approach (e.g., Lewis et al., 2006; Van Dyke et al., 2014). While neither approach was fully supported, a possible means of reconciling the two approaches and directions for future research are proposed.
  • Tsuji, S., Fikkert, P., Minagawa, Y., Dupoux, E., Filippin, L., Versteegh, M., Hagoort, P., & Cristia, A. (2017). The more, the better? Behavioral and neural correlates of frequent and infrequent vowel exposure. Developmental Psychobiology, 59, 603-612. doi:10.1002/dev.21534.

    Abstract

    A central assumption in the perceptual attunement literature holds that exposure to a speech sound contrast leads to improvement in native speech sound processing. However, whether the amount of exposure matters for this process has not been put to a direct test. We elucidated indicators of frequency-dependent perceptual attunement by comparing 5–8-month-old Dutch infants’ discrimination of tokens containing a highly frequent [hɪt-he:t] and a highly infrequent [hʏt-hø:t] native vowel contrast as well as a non-native [hɛt-hæt] vowel contrast in a behavioral visual habituation paradigm (Experiment 1). Infants discriminated both native contrasts similarly well, but did not discriminate the non-native contrast. We sought further evidence for subtle differences in the processing of the two native contrasts using near-infrared spectroscopy and a within-participant design (Experiment 2). The neuroimaging data did not provide additional evidence that responses to native contrasts are modulated by frequency of exposure. These results suggest that even large differences in exposure to a native contrast may not directly translate to behavioral and neural indicators of perceptual attunement, raising the possibility that frequency of exposure does not influence improvements in discriminating native contrasts.

    Additional information

    dev21534-sup-0001-SuppInfo-S1.docx
  • Udden, J., Ingvar, M., Hagoort, P., & Petersson, K. M. (2017). Broca’s region: A causal role in implicit processing of grammars with crossed non-adjacent dependencies. Cognition, 164, 188-198. doi:10.1016/j.cognition.2017.03.010.

    Abstract

    Non-adjacent dependencies are challenging for the language learning machinery and are acquired later than adjacent dependencies. In this transcranial magnetic stimulation (TMS) study, we show that participants successfully discriminated between grammatical and non-grammatical sequences after having implicitly acquired an artificial language with crossed non-adjacent dependencies. Subsequent to transcranial magnetic stimulation of Broca’s region, discrimination was impaired compared to when a language-irrelevant control region (vertex) was stimulated. These results support the view that Broca’s region is engaged in structured sequence processing and extend previous functional neuroimaging results on artificial grammar learning (AGL) in two directions: first, the results establish that Broca’s region is a causal component in the processing of non-adjacent dependencies, and second, they show that implicit processing of non-adjacent dependencies engages Broca’s region. Since patients with lesions in Broca’s region do not always show grammatical processing difficulties, the result that Broca’s region is causally linked to processing of non-adjacent dependencies is a step towards clarification of the exact nature of syntactic deficits caused by lesions or perturbation to Broca’s region. Our findings are consistent with previous results and support a role for Broca’s region in general structured sequence processing, rather than a specific role for the processing of hierarchically organized sentence structure.
  • Udden, J., Snijders, T. M., Fisher, S. E., & Hagoort, P. (2017). A common variant of the CNTNAP2 gene is associated with structural variation in the left superior occipital gyrus. Brain and Language, 172, 16-21. doi:10.1016/j.bandl.2016.02.003.

    Abstract

    The CNTNAP2 gene encodes a cell-adhesion molecule that influences the properties of neural networks and the morphology and density of neurons and glial cells. Previous studies have shown association of CNTNAP2 variants with language-related phenotypes in health and disease. Here, we report associations of a common CNTNAP2 polymorphism (rs7794745) with variation in grey matter in a region in the dorsal visual stream. We tried to replicate an earlier study on 314 subjects by Tan and colleagues (2010), but now in a substantially larger group of more than 1700 subjects. Carriers of the T allele showed reduced grey matter volume in left superior occipital gyrus, while we did not replicate associations with grey matter volume in other regions identified by Tan et al (2010). Our work illustrates the importance of independent replication in neuroimaging genetic studies of language-related candidate genes.
  • Van der Ven, F., Takashima, A., Segers, A., & Verhoeven, L. (2017). Semantic priming in Dutch children: Word meaning integration and study modality effects. Language Learning, 67(3), 546-568. doi:10.1111/lang.12235.
  • Van Bergen, G., & Flecken, M. (2017). Putting things in new places: Linguistic experience modulates the predictive power of placement verb semantics. Journal of Memory and Language, 92, 26-42. doi:10.1016/j.jml.2016.05.003.

    Abstract

    A central question regarding predictive language processing concerns the extent to which linguistic experience modulates the process. We approached this question by investigating sentence processing in advanced second language (L2) users with different native language (L1) backgrounds. Using a visual world eye tracking paradigm, we investigated to what extent L1 and L2 participants showed anticipatory eye movements to objects while listening to Dutch placement event descriptions. L2 groups differed in the degree of similarity between Dutch and their L1 with respect to placement verb semantics: German, like Dutch, specifies object position in placement verbs (put.STAND vs. put.LIE), whereas English and French typically leave position underspecified (put). Results showed that German L2 listeners, like native Dutch listeners, anticipate objects that match the verbally encoded position immediately upon encountering the verb. French/English L2 participants, however, did not show any prediction effects, despite proper understanding of Dutch placement verbs. Our findings suggest that prior experience with a specific semantic contrast in one’s L1 facilitates prediction in L2, and hence adds to the evidence that linguistic experience modulates predictive sentence processing
  • Van Ekert, J., Wegman, J., Jansen, C., Takashima, A., & Janzen, G. (2017). The dynamics of memory consolidation of landmarks. Hippocampus, 27(4), 303-404. doi:10.1002/hipo.22698.

    Abstract

    Navigating through space is fundamental to human nature and requires the ability to retrieve relevant information from the remote past. With the passage of time, some memories become generic, capturing only a sense of familiarity. Yet, others maintain precision, even when acquired decades ago. Understanding the dynamics of memory consolidation is a major challenge to neuroscientists. Using functional magnetic resonance imaging, we systematically examined the effects of time and spatial context on the neural representation of landmark recognition memory. An equal number of male and female subjects (males N = 10, total N = 20) watched a route through a large-scale virtual environment. Landmarks occurred at navigationally relevant and irrelevant locations along the route. Recognition memory for landmarks was tested directly following encoding, 24 h later and 30 days later. Surprisingly, changes over time in the neural representation of navigationally relevant landmarks differed between males and females. In males, relevant landmarks selectively engaged the parahippocampal gyrus (PHG) regardless of the age of the memory. In females, the response to relevant landmarks gradually diminished with time in the PHG but strengthened progressively in the inferior frontal gyrus (IFG). Based on what is known about the functioning of the PHG and IFG, the findings of this study suggest that males maintain access to the initially formed spatial representation of landmarks whereas females become strongly dependent on a verbal representation of landmarks with time. Our findings yield a clear objective for future studies
  • Vanlangendonck, F. (2017). Finding common ground: On the neural mechanisms of communicative language production. PhD Thesis, Radboud University, Nijmegen.
  • Varma, S., Takashima, A., Krewinkel, S., Van Kooten, M., Fu, L., Medendorp, W. P., Kessels, R. P. C., & Daselaar, S. M. (2017). Non-interfering effects of active post-encoding tasks on episodic memory consolidation in humans. Frontiers in Behavioral Neuroscience, 11: 54. doi:10.3389/fnbeh.2017.00054.

    Abstract

    So far, studies that investigated interference effects of post-learning processes on episodic memory consolidation in humans have used tasks involving only complex and meaningful information. Such tasks require reallocation of general or encoding-specific resources away from consolidation-relevant activities. The possibility that interference can be elicited using a task that heavily taxes our limited brain resources, but has low semantic and hippocampal related long-term memory processing demands, has never been tested. We address this question by investigating whether consolidation could persist in parallel with an active, encoding-irrelevant, minimally semantic task, regardless of its high resource demands for cognitive processing. We distinguish the impact of such a task on consolidation based on whether it engages resources that are: (1) general/executive, or (2) specific/overlapping with the encoding modality. Our experiments compared subsequent memory performance across two post-encoding consolidation periods: quiet wakeful rest and a cognitively demanding n-Back task. Across six different experiments (total N = 176), we carefully manipulated the design of the n-Back task to target general or specific resources engaged in the ongoing consolidation process. In contrast to previous studies that employed interference tasks involving conceptual stimuli and complex processing demands, we did not find any differences between n-Back and rest conditions on memory performance at delayed test, using both recall and recognition tests. Our results indicate that: (1) quiet, wakeful rest is not a necessary prerequisite for episodic memory consolidation; and (2) post-encoding cognitive engagement does not interfere with memory consolidation when task-performance has minimal semantic and hippocampally-based episodic memory processing demands. We discuss our findings with reference to resource and reactivation-led interference theories
  • Araújo, S., Faísca, L., Reis, A., Marques, J. F., & Petersson, K. M. (2016). Visual naming deficits in dyslexia: An ERP investigation of different processing domains. Neuropsychologia, 91, 61-76. doi:10.1016/j.neuropsychologia.2016.07.007.

    Abstract

    Naming speed deficits are well documented in developmental dyslexia, expressed by slower naming times and more errors in response to familiar items. Here we used event-related potentials (ERPs) to examine at what processing level the deficits in dyslexia emerge during a discrete-naming task. Dyslexic and skilled adult control readers performed a primed object-naming task, in which the relationship between the prime and the target was manipulated along perceptual, semantic and phonological dimensions. A 3×2 design that crossed Relationship Type (Visual, Phonemic Onset, and Semantic) with Relatedness (Related and Unrelated) was used. An attenuated N/P190 – indexing early visual processing – and N300 – which index late visual processing – was observed to pictures preceded by perceptually related (vs. unrelated) primes in the control but not in the dyslexic group. These findings suggest suboptimal processing in early stages of object processing in dyslexia, when integration and mapping of perceptual information to a more form-specific percept in memory take place. On the other hand, both groups showed an N400 effect associated with semantically related pictures (vs. unrelated), taken to reflect intact integration of semantic similarities in both dyslexic and control readers. We also found an electrophysiological effect of phonological priming in the N400 range – that is, an attenuated N400 to objects preceded by phonemic related primes vs. unrelated – while it showed a more widespread distributed and more pronounced over the right hemisphere in the dyslexics. Topographic differences between groups might have originated from a word form encoding process with different characteristics in dyslexics compared to control readers.
  • Asaridou, S. S., Takashima, A., Dediu, D., Hagoort, P., & McQueen, J. M. (2016). Repetition suppression in the left inferior frontal gyrus predicts tone learning performance. Cerebral Cortex, 26(6), 2728-2742. doi:10.1093/cercor/bhv126.

    Abstract

    Do individuals differ in how efficiently they process non-native sounds? To what extent do these differences relate to individual variability in sound-learning aptitude? We addressed these questions by assessing the sound-learning abilities of Dutch native speakers as they were trained on non-native tone contrasts. We used fMRI repetition suppression to the non-native tones to measure participants' neuronal processing efficiency before and after training. Although all participants improved in tone identification with training, there was large individual variability in learning performance. A repetition suppression effect to tone was found in the bilateral inferior frontal gyri (IFGs) before training. No whole-brain effect was found after training; a region-of-interest analysis, however, showed that, after training, repetition suppression to tone in the left IFG correlated positively with learning. That is, individuals who were better in learning the non-native tones showed larger repetition suppression in this area. Crucially, this was true even before training. These findings add to existing evidence that the left IFG plays an important role in sound learning and indicate that individual differences in learning aptitude stem from differences in the neuronal efficiency with which non-native sounds are processed.
  • Backus, A., Schoffelen, J.-M., Szebényi, S., Hanslmayr, S., & Doeller, C. (2016). Hippocampal-prefrontal theta oscillations support memory integration. Current Biology, 26, 450-457. doi:10.1016/j.cub.2015.12.048.

    Abstract

    Integration of separate memories forms the basis of inferential reasoning - an essential cognitive process that enables complex behavior. Considerable evidence suggests that both hippocampus and medial prefrontal cortex (mPFC) play a crucial role in memory integration. Although previous studies indicate that theta oscillations facilitate memory processes, the electrophysiological mechanisms underlying memory integration remain elusive. To bridge this gap, we recorded magnetoencephalography data while participants performed an inference task and employed novel source reconstruction techniques to estimate oscillatory signals from the hippocampus. We found that hippocampal theta power during encoding predicts subsequent memory integration. Moreover, we observed increased theta coherence between hippocampus and mPFC. Our results suggest that integrated memory representations arise through hippocampal theta oscillations, possibly reflecting dynamic switching between encoding and retrieval states, and facilitating communication with mPFC. These findings have important implications for our understanding of memory-based decision making and knowledge acquisition
  • Bastos, A. M., & Schoffelen, J.-M. (2016). A tutorial review of functional connectivity analysis methods and their interpretational pitfalls. Frontiers in Systems Neuroscience, 9: 175. doi:10.3389/fnsys.2015.00175.

    Abstract

    Oscillatory neuronal activity may provide a mechanism for dynamic network coordination. Rhythmic neuronal interactions can be quantified using multiple metrics, each with their own advantages and disadvantages. This tutorial will review and summarize current analysis methods used in the field of invasive and non-invasive electrophysiology to study the dynamic connections between neuronal populations. First, we review metrics for functional connectivity, including coherence, phase synchronization, phase-slope index, and Granger causality, with the specific aim to provide an intuition for how these metrics work, as well as their quantitative definition. Next, we highlight a number of interpretational caveats and common pitfalls that can arise when performing functional connectivity analysis, including the common reference problem, the signal to noise ratio problem, the volume conduction problem, the common input problem, and the sample size bias problem. These pitfalls will be illustrated by presenting a set of MATLAB-scripts, which can be executed by the reader to simulate each of these potential problems. We discuss how these issues can be addressed using current methods.
  • Bramão, I., Reis, A., Petersson, K. M., & Faísca, L. (2016). Knowing that strawberries are red and seeing red strawberries: The interaction between surface colour and colour knowledge information. Journal of Cognitive Psychology, 28(6), 641-657. doi:10.1080/20445911.2016.1182171.

    Abstract

    his study investigates the interaction between surface and colour knowledge information during object recognition. In two different experiments, participants were instructed to decide whether two presented stimuli belonged to the same object identity. On the non-matching trials, we manipulated the shape and colour knowledge information activated by the two stimuli by creating four different stimulus pairs: (1) similar in shape and colour (e.g. TOMATO–APPLE); (2) similar in shape and dissimilar in colour (e.g. TOMATO–COCONUT); (3) dissimilar in shape and similar in colour (e.g. TOMATO–CHILI PEPPER) and (4) dissimilar in both shape and colour (e.g. TOMATO–PEANUT). The object pictures were presented in typical and atypical colours and also in black-and-white. The interaction between surface and colour knowledge showed to be contingent upon shape information: while colour knowledge is more important for recognising structurally similar shaped objects, surface colour is more prominent for recognising structurally dissimilar shaped objects.
  • Broersma, M., Carter, D., & Acheson, D. J. (2016). Cognate costs in bilingual speech production: Evidence from language switching. Frontiers in Psychology, 7: 1461. doi:10.3389/fpsyg.2016.01461.

    Abstract

    This study investigates cross-language lexical competition in the bilingual mental lexicon. It provides evidence for the occurrence of inhibition as well as the commonly reported facilitation during the production of cognates (words with similar phonological form and meaning in two languages) in a mixed picture naming task by highly proficient Welsh-English bilinguals. Previous studies have typically found cognate facilitation. It has previously been proposed (with respect to non-cognates) that cross-language inhibition is limited to low-proficient bilinguals; therefore, we tested highly proficient, early bilinguals. In a mixed naming experiment (i.e., picture naming with language switching), 48 highly proficient, early Welsh-English bilinguals named pictures in Welsh and English, including cognate and non-cognate targets. Participants were English-dominant, Welsh-dominant, or had equal language dominance. The results showed evidence for cognate inhibition in two ways. First, both facilitation and inhibition were found on the cognate trials themselves, compared to non-cognate controls, modulated by the participants' language dominance. The English-dominant group showed cognate inhibition when naming in Welsh (and no difference between cognates and controls when naming in English), and the Welsh-dominant and equal dominance groups generally showed cognate facilitation. Second, cognate inhibition was found as a behavioral adaptation effect, with slower naming for non-cognate filler words in trials after cognates than after non-cognate controls. This effect was consistent across all language dominance groups and both target languages, suggesting that cognate production involved cognitive control even if this was not measurable in the cognate trials themselves. Finally, the results replicated patterns of symmetrical switch costs, as commonly reported for balanced bilinguals. We propose that cognate processing might be affected by two different processes, namely competition at the lexical-semantic level and facilitation at the word form level, and that facilitation at the word form level might (sometimes) outweigh any effects of inhibition at the lemma level. In sum, this study provides evidence that cognate naming can cause costs in addition to benefits. The finding of cognate inhibition, particularly for the highly proficient bilinguals tested, provides strong evidence for the occurrence of lexical competition across languages in the bilingual mental lexicon.
  • Chu, M., & Kita, S. (2016). Co-thought and Co-speech Gestures Are Generated by the Same Action Generation Process. Journal of Experimental Psychology: Learning, Memory, and Cognition, 42(2), 257-270. doi:10.1037/xlm0000168.

    Abstract

    People spontaneously gesture when they speak (co-speech gestures) and when they solve problems silently (co-thought gestures). In this study, we first explored the relationship between these 2 types of gestures and found that individuals who produced co-thought gestures more frequently also produced co-speech gestures more frequently (Experiments 1 and 2). This suggests that the 2 types of gestures are generated from the same process. We then investigated whether both types of gestures can be generated from the representational use of the action generation process that also generates purposeful actions that have a direct physical impact on the world, such as manipulating an object or locomotion (the action generation hypothesis). To this end, we examined the effect of object affordances on the production of both types of gestures (Experiments 3 and 4). We found that individuals produced co-thought and co-speech gestures more often when the stimulus objects afforded action (objects with a smooth surface) than when they did not (objects with a spiky surface). These results support the action generation hypothesis for representational gestures. However, our findings are incompatible with the hypothesis that co-speech representational gestures are solely generated from the speech production process (the speech production hypothesis).
  • Coulson, S., & Lai, V. T. (Eds.). (2016). The metaphorical brain [Research topic]. Lausanne: Frontiers Media. doi:10.3389/978-2-88919-772-9.

    Abstract

    This Frontiers Special Issue will synthesize current findings on the cognitive neuroscience of metaphor, provide a forum for voicing novel perspectives, and promote new insights into the metaphorical brain.
  • Dimitrova, D. V., Chu, M., Wang, L., Ozyurek, A., & Hagoort, P. (2016). Beat that word: How listeners integrate beat gesture and focus in multimodal speech discourse. Journal of Cognitive Neuroscience, 28(9), 1255-1269. doi:10.1162/jocn_a_00963.

    Abstract

    Communication is facilitated when listeners allocate their attention to important information (focus) in the message, a process called "information structure." Linguistic cues like the preceding context and pitch accent help listeners to identify focused information. In multimodal communication, relevant information can be emphasized by nonverbal cues like beat gestures, which represent rhythmic nonmeaningful hand movements. Recent studies have found that linguistic and nonverbal attention cues are integrated independently in single sentences. However, it is possible that these two cues interact when information is embedded in context, because context allows listeners to predict what information is important. In an ERP study, we tested this hypothesis and asked listeners to view videos capturing a dialogue. In the critical sentence, focused and nonfocused words were accompanied by beat gestures, grooming hand movements, or no gestures. ERP results showed that focused words are processed more attentively than nonfocused words as reflected in an N1 and P300 component. Hand movements also captured attention and elicited a P300 component. Importantly, beat gesture and focus interacted in a late time window of 600-900 msec relative to target word onset, giving rise to a late positivity when nonfocused words were accompanied by beat gestures. Our results show that listeners integrate beat gesture with the focus of the message and that integration costs arise when beat gesture falls on nonfocused information. This suggests that beat gestures fulfill a unique focusing function in multimodal discourse processing and that they have to be integrated with the information structure of the message.
  • Francken, J. C. (2016). Viewing the world through language-tinted glasses: Elucidating the neural mechanisms of language-perception interactions. PhD Thesis, Radboud University, Nijmegen.
  • Frank, S. L., & Fitz, H. (2016). Reservoir computing and the Sooner-is-Better bottleneck [Commentary on Christiansen & Slater]. Behavioral and Brain Sciences, 39: e73. doi:10.1017/S0140525X15000783.

    Abstract

    Prior language input is not lost but integrated with the current input. This principle is demonstrated by “reservoir computing”: Untrained recurrent neural networks project input sequences onto a random point in high-dimensional state space. Earlier inputs can be retrieved from this projection, albeit less reliably so as more input is received. The bottleneck is therefore not “Now-or-Never” but “Sooner-is-Better.
  • Gerwien, J., & Flecken, M. (2016). First things first? Top-down influences on event apprehension. In A. Papafragou, D. Grodner, D. Mirman, & J. Trueswell (Eds.), Proceedings of the 38th Annual Meeting of the Cognitive Science Society (CogSci 2016) (pp. 2633-2638). Austin, TX: Cognitive Science Society.

    Abstract

    Not much is known about event apprehension, the earliest stage of information processing in elicited language production studies, using pictorial stimuli. A reason for our lack of knowledge on this process is that apprehension happens very rapidly (<350 ms after stimulus onset, Griffin & Bock 2000), making it difficult to measure the process directly. To broaden our understanding of apprehension, we analyzed landing positions and onset latencies of first fixations on visual stimuli (pictures of real-world events) given short stimulus presentation times, presupposing that the first fixation directly results from information processing during apprehension
  • Gijssels, T., Staum Casasanto, L., Jasmin, K., Hagoort, P., & Casasanto, D. (2016). Speech accommodation without priming: The case of pitch. Discourse Processes, 53(4), 233-251. doi:10.1080/0163853X.2015.1023965.

    Abstract

    People often accommodate to each other's speech by aligning their linguistic production with their partner's. According to an influential theory, the Interactive Alignment Model (Pickering & Garrod, 2004), alignment is the result of priming. When people perceive an utterance, the corresponding linguistic representations are primed, and become easier to produce. Here we tested this theory by investigating whether pitch (F0) alignment shows two characteristic signatures of priming: dose dependence and persistence. In a virtual reality experiment, we manipulated the pitch of a virtual interlocutor's speech to find out (a.) whether participants accommodated to the agent's F0, (b.) whether the amount of accommodation increased with increasing exposure to the agent's speech, and (c.) whether changes to participants' F0 persisted beyond the conversation. Participants accommodated to the virtual interlocutor, but accommodation did not increase in strength over the conversation, and it disappeared immediately after the conversation ended. Results argue against a priming-based account of F0 accommodation, and indicate that an alternative mechanism is needed to explain alignment along continuous dimensions of language such as speech rate and pitch.
  • Hagoort, P. (2016). MUC (Memory, Unification, Control): A Model on the Neurobiology of Language Beyond Single Word Processing. In G. Hickok, & S. Small (Eds.), Neurobiology of language (pp. 339-347). Amsterdam: Elsever. doi:10.1016/B978-0-12-407794-2.00028-6.

    Abstract

    A neurobiological model of language is discussed that overcomes the shortcomings of the classical Wernicke-Lichtheim-Geschwind model. It is based on a subdivision of language processing into three components: Memory, Unification, and Control. The functional components as well as the neurobiological underpinnings of the model are discussed. In addition, the need for extension beyond the classical core regions for language is shown. Attentional networks as well as networks for inferential processing are crucial to realize language comprehension beyond single word processing and beyond decoding propositional content.
  • Hagoort, P. (2016). Zij zijn ons brein. In J. Brockman (Ed.), Machines die denken: Invloedrijke denkers over de komst van kunstmatige intelligentie (pp. 184-186). Amsterdam: Maven Publishing.
  • Hartung, F., Burke, M., Hagoort, P., & Willems, R. M. (2016). Taking perspective: Personal pronouns affect experiential aspects of literary reading. PLoS One, 11(5): e0154732. doi:10.1371/journal.pone.0154732.

    Abstract

    Personal pronouns have been shown to influence cognitive perspective taking during comprehension. Studies using single sentences found that 3rd person pronouns facilitate the construction of a mental model from an observer’s perspective, whereas 2nd person pronouns support an actor’s perspective. The direction of the effect for 1st person pronouns seems to depend on the situational context. In the present study, we investigated how personal pronouns influence discourse comprehension when people read fiction stories and if this has consequences for affective components like emotion during reading or appreciation of the story. We wanted to find out if personal pronouns affect immersion and arousal, as well as appreciation of fiction. In a natural reading paradigm, we measured electrodermal activity and story immersion, while participants read literary stories with 1st and 3rd person pronouns referring to the protagonist. In addition, participants rated and ranked the stories for appreciation. Our results show that stories with 1st person pronouns lead to higher immersion. Two factors—transportation into the story world and mental imagery during reading—in particular showed higher scores for 1st person as compared to 3rd person pronoun stories. In contrast, arousal as measured by electrodermal activity seemed tentatively higher for 3rd person pronoun stories. The two measures of appreciation were not affected by the pronoun manipulation. Our findings underscore the importance of perspective for language processing, and additionally show which aspects of the narrative experience are influenced by a change in perspective.
  • St. John-Saaltink, E. (2016). When the past influences the present: Modulations of the sensory response by prior knowledge and task set. PhD Thesis, Radboud University, Nijmegen.
  • Kösem, A., Basirat, A., Azizi, L., & van Wassenhove, V. (2016). High frequency neural activity predicts word parsing in ambiguous speech streams. Journal of Neurophysiology, 116(6), 2497-2512. doi:10.1152/jn.00074.2016.

    Abstract

    During speech listening, the brain parses a continuous acoustic stream of information into computational units (e.g. syllables or words) necessary for speech comprehension. Recent neuroscientific hypotheses propose that neural oscillations contribute to speech parsing, but whether they do so on the basis of acoustic cues (bottom-up acoustic parsing) or as a function of available linguistic representations (top-down linguistic parsing) is unknown. In this magnetoencephalography study, we contrasted acoustic and linguistic parsing using bistable speech sequences. While listening to the speech sequences, participants were asked to maintain one of the two possible speech percepts through volitional control. We predicted that the tracking of speech dynamics by neural oscillations would not only follow the acoustic properties but also shift in time according to the participant’s conscious speech percept. Our results show that the latency of high-frequency activity (specifically, beta and gamma bands) varied as a function of the perceptual report. In contrast, the phase of low-frequency oscillations was not strongly affected by top-down control. While changes in low-frequency neural oscillations were compatible with the encoding of pre-lexical segmentation cues, high-frequency activity specifically informed on an individual’s conscious speech percept.

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  • Kunert, R., Willems, R. M., & Hagoort, P. (2016). An independent psychometric evaluation of the PROMS measure of music perception skills. PLoS One, 11(7): e0159103. doi:10.1371/journal.pone.0159103.

    Abstract

    The Profile of Music Perception Skills (PROMS) is a recently developed measure of perceptual music skills which has been shown to have promising psychometric properties. In this paper we extend the evaluation of its brief version to three kinds of validity using an individual difference approach. The brief PROMS displays good discriminant validity with working memory, given that it does not correlate with backward digit span (r = .04). Moreover, it shows promising criterion validity (association with musical training (r = .45), musicianship status (r = .48), and self-rated musical talent (r = .51)). Finally, its convergent validity, i.e. relation to an unrelated measure of music perception skills, was assessed by correlating the brief PROMS to harmonic closure judgment accuracy. Two independent samples point to good convergent validity of the brief PROMS (r = .36; r = .40). The same association is still significant in one of the samples when including self-reported music skill in a partial correlation (rpartial = .30; rpartial = .17). Overall, the results show that the brief version of the PROMS displays a very good pattern of construct validity. Especially its tuning subtest stands out as a valuable part for music skill evaluations in Western samples. We conclude by briefly discussing the choice faced by music cognition researchers between different musical aptitude measures of which the brief PROMS is a well evaluated example.
  • Kunert, R., Willems, R. M., & Hagoort, P. (2016). Language influences music harmony perception: effects of shared syntactic integration resources beyond attention. Royal Society Open Science, 3(2): 150685. doi:10.1098/rsos.150685.

    Abstract

    Many studies have revealed shared music–language processing resources by finding an influence of music harmony manipulations on concurrent language processing. However, the nature of the shared resources has remained ambiguous. They have been argued to be syntax specific and thus due to shared syntactic integration resources. An alternative view regards them as related to general attention and, thus, not specific to syntax. The present experiments evaluated these accounts by investigating the influence of language on music. Participants were asked to provide closure judgements on harmonic sequences in order to assess the appropriateness of sequence endings. At the same time participants read syntactic garden-path sentences. Closure judgements revealed a change in harmonic processing as the result of reading a syntactically challenging word. We found no influence of an arithmetic control manipulation (experiment 1) or semantic garden-path sentences (experiment 2). Our results provide behavioural evidence for a specific influence of linguistic syntax processing on musical harmony judgements. A closer look reveals that the shared resources appear to be needed to hold a harmonic key online in some form of syntactic working memory or unification workspace related to the integration of chords and words. Overall, our results support the syntax specificity of shared music–language processing resources.
  • Kunert, R. (2016). Internal conceptual replications do not increase independent replication success. Psychonomic Bulletin & Review, 23(5), 1631-1638. doi:10.3758/s13423-016-1030-9.

    Abstract

    Recently, many psychological effects have been surprisingly difficult to reproduce. This article asks why, and investigates whether conceptually replicating an effect in the original publication is related to the success of independent, direct replications. Two prominent accounts of low reproducibility make different predictions in this respect. One account suggests that psychological phenomena are dependent on unknown contexts that are not reproduced in independent replication attempts. By this account, internal replications indicate that a finding is more robust and, thus, that it is easier to independently replicate it. An alternative account suggests that researchers employ questionable research practices (QRPs), which increase false positive rates. By this account, the success of internal replications may just be the result of QRPs and, thus, internal replications are not predictive of independent replication success. The data of a large reproducibility project support the QRP account: replicating an effect in the original publication is not related to independent replication success. Additional analyses reveal that internally replicated and internally unreplicated effects are not very different in terms of variables associated with replication success. Moreover, social psychological effects in particular appear to lack any benefit from internal replications. Overall, these results indicate that, in this dataset at least, the influence of QRPs is at the heart of failures to replicate psychological findings, especially in social psychology. Variable, unknown contexts appear to play only a relatively minor role. I recommend practical solutions for how QRPs can be avoided.

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  • Lai, V. T., & Huettig, F. (2016). When prediction is fulfilled: Insight from emotion processing. Neuropsychologia, 85, 110-117. doi:10.1016/j.neuropsychologia.2016.03.014.

    Abstract

    Research on prediction in language processing has focused predominantly on the function of predictive context and less on the potential contribution of the predicted word. The present study investigated how meaning that is not immediately prominent in the contents of predictions but is part of the predicted words influences sentence processing. We used emotional meaning to address this question. Participants read emotional and neutral words embedded in highly predictive and non-predictive sentential contexts, with the two sentential contexts rated similarly for their emotional ratings. Event Related Potential (ERP) effects of prediction and emotion both started at ~200 ms. Confirmed predictions elicited larger P200s than violated predictions when the target words were non-emotional (neutral), but such effect was absent when the target words were emotional. Likewise, emotional words elicited larger P200s than neutral words when the target words were non-predictive, but such effect were absent when the contexts were predictive. We conjecture that the prediction and emotion effects at ~200 ms may share similar neural process(es). We suggest that such process(es) could be affective, where confirmed predictions and word emotion give rise to ‘aha’ or reward feelings, and/or cognitive, where both prediction and word emotion quickly engage attention

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  • Lam, N. H. L., Schoffelen, J.-M., Udden, J., Hulten, A., & Hagoort, P. (2016). Neural activity during sentence processing as reflected in theta, alpha, beta and gamma oscillations. NeuroImage, 142(15), 43-54. doi:10.1016/j.neuroimage.2016.03.007.

    Abstract

    We used magnetoencephalography (MEG) to explore the spatio-temporal dynamics of neural oscillations associated with sentence processing, in 102 participants. We quantified changes in oscillatory power as the sentence unfolded, and in response to individual words in the sentence. For words early in a sentence compared to those late in the same sentence, we observed differences in left temporal and frontal areas, and bilateral frontal and right parietal regions for the theta, alpha, and beta frequency bands. The neural response to words in a sentence differed from the response to words in scrambled sentences in left-lateralized theta, alpha, beta, and gamma. The theta band effects suggest that a sentential context facilitates lexical retrieval, and that this facilitation is stronger for words late in the sentence. Effects in the alpha and beta band may reflect the unification of semantic and syntactic information, and are suggestive of easier unification late in a sentence. The gamma oscillations are indicative of predicting the upcoming word during sentence processing. In conclusion, changes in oscillatory neuronal activity capture aspects of sentence processing. Our results support earlier claims that language (sentence) processing recruits areas distributed across both hemispheres, and extends beyond the classical language regions
  • Lartseva, A. (2016). Reading emotions: How people with Autism Spectrum Disorders process emotional language. PhD Thesis, Radboud University Nijmegen, Nijmegen.
  • Leonard, M., Baud, M., Sjerps, M. J., & Chang, E. (2016). Perceptual restoration of masked speech in human cortex. Nature Communications, 7: 13619. doi:10.1038/ncomms13619.

    Abstract

    Humans are adept at understanding speech despite the fact that our natural listening environment is often filled with interference. An example of this capacity is phoneme restoration, in which part of a word is completely replaced by noise, yet listeners report hearing the whole word. The neurological basis for this unconscious fill-in phenomenon is unknown, despite being a fundamental characteristic of human hearing. Here, using direct cortical recordings in humans, we demonstrate that missing speech is restored at the acoustic-phonetic level in bilateral auditory cortex, in real-time. This restoration is preceded by specific neural activity patterns in a separate language area, left frontal cortex, which predicts the word that participants later report hearing. These results demonstrate that during speech perception, missing acoustic content is synthesized online from the integration of incoming sensory cues and the internal neural dynamics that bias word-level expectation and prediction.

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  • Lewis, A. G., Schoffelen, J.-M., Schriefers, H., & Bastiaansen, M. C. M. (2016). A Predictive Coding Perspective on Beta Oscillations during Sentence-Level Language Comprehension. Frontiers in Human Neuroscience, 10: 85. doi:10.3389/fnhum.2016.00085.

    Abstract

    Oscillatory neural dynamics have been steadily receiving more attention as a robust and temporally precise signature of network activity related to language processing. We have recently proposed that oscillatory dynamics in the beta and gamma frequency ranges measured during sentence-level comprehension might be best explained from a predictive coding perspective. Under our proposal we related beta oscillations to both the maintenance/change of the neural network configuration responsible for the construction and representation of sentence-level meaning, and to top–down predictions about upcoming linguistic input based on that sentence-level meaning. Here we zoom in on these particular aspects of our proposal, and discuss both old and new supporting evidence. Finally, we present some preliminary magnetoencephalography data from an experiment comparing Dutch subject- and object-relative clauses that was specifically designed to test our predictive coding framework. Initial results support the first of the two suggested roles for beta oscillations in sentence-level language comprehension.
  • Lewis, A. G., Lemhӧfer, K., Schoffelen, J.-M., & Schriefers, H. (2016). Gender agreement violations modulate beta oscillatory dynamics during sentence comprehension: A comparison of second language learners and native speakers. Neuropsychologia, 89(1), 254-272. doi:10.1016/j.neuropsychologia.2016.06.031.

    Abstract

    For native speakers, many studies suggest a link between oscillatory neural activity in the beta frequency range and syntactic processing. For late second language (L2) learners on the other hand, the extent to which the neural architecture supporting syntactic processing is similar to or different from that of native speakers is still unclear. In a series of four experiments, we used electroencephalography to investigate the link between beta oscillatory activity and the processing of grammatical gender agreement in Dutch determiner-noun pairs, for Dutch native speakers, and for German L2 learners of Dutch. In Experiment 1 we show that for native speakers, grammatical gender agreement violations are yet another among many syntactic factors that modulate beta oscillatory activity during sentence comprehension. Beta power is higher for grammatically acceptable target words than for those that mismatch in grammatical gender with their preceding determiner. In Experiment 2 we observed no such beta modulations for L2 learners, irrespective of whether trials were sorted according to objective or subjective syntactic correctness. Experiment 3 ruled out that the absence of a beta effect for the L2 learners in Experiment 2 was due to repetition of the target nouns in objectively correct and incorrect determiner-noun pairs. Finally, Experiment 4 showed that when L2 learners are required to explicitly focus on grammatical information, they show modulations of beta oscillatory activity, comparable to those of native speakers, but only when trials are sorted according to participants’ idiosyncratic lexical representations of the grammatical gender of target nouns. Together, these findings suggest that beta power in L2 learners is sensitive to violations of grammatical gender agreement, but only when the importance of grammatical information is highlighted, and only when participants' subjective lexical representations are taken into account.
  • Lockwood, G. (2016). Academic clickbait: Articles with positively-framed titles, interesting phrasing, and no wordplay get more attention online. The Winnower, 3: e146723.36330. doi:10.15200/winn.146723.36330.

    Abstract

    This article is about whether the factors which drive online sharing of non-scholarly content also apply to academic journal titles. It uses Altmetric scores as a measure of online attention to articles from Frontiers in Psychology published in 2013 and 2014. Article titles with result-oriented positive framing and more interesting phrasing receive higher Altmetric scores, i.e., get more online attention. Article titles with wordplay and longer article titles receive lower Altmetric scores. This suggests that the same factors that affect how widely non-scholarly content is shared extend to academia, which has implications for how academics can make their work more likely to have more impact.
  • Lockwood, G., Hagoort, P., & Dingemanse, M. (2016). How iconicity helps people learn new words: neural correlates and individual differences in sound-symbolic bootstrapping. Collabra, 2(1): 7. doi:10.1525/collabra.42.

    Abstract

    Sound symbolism is increasingly understood as involving iconicity, or perceptual analogies and cross-modal correspondences between form and meaning, but the search for its functional and neural correlates is ongoing. Here we study how people learn sound-symbolic words, using behavioural, electrophysiological and individual difference measures. Dutch participants learned Japanese ideophones —lexical sound-symbolic words— with a translation of either the real meaning (in which form and meaning show cross-modal correspondences) or the opposite meaning (in which form and meaning show cross-modal clashes). Participants were significantly better at identifying the words they learned in the real condition, correctly remembering the real word pairing 86.7% of the time, but the opposite word pairing only 71.3% of the time. Analysing event-related potentials (ERPs) during the test round showed that ideophones in the real condition elicited a greater P3 component and late positive complex than ideophones in the opposite condition. In a subsequent forced choice task, participants were asked to guess the real translation from two alternatives. They did this with 73.0% accuracy, well above chance level even for words they had encountered in the opposite condition, showing that people are generally sensitive to the sound-symbolic cues in ideophones. Individual difference measures showed that the ERP effect in the test round of the learning task was greater for participants who were more sensitive to sound symbolism in the forced choice task. The main driver of the difference was a lower amplitude of the P3 component in response to ideophones in the opposite condition, suggesting that people who are more sensitive to sound symbolism may have more difficulty to suppress conflicting cross-modal information. The findings provide new evidence that cross-modal correspondences between sound and meaning facilitate word learning, while cross-modal clashes make word learning harder, especially for people who are more sensitive to sound symbolism.

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  • Lockwood, G., Dingemanse, M., & Hagoort, P. (2016). Sound-symbolism boosts novel word learning. Journal of Experimental Psychology: Learning, Memory, and Cognition, 42(8), 1274-1281. doi:10.1037/xlm0000235.

    Abstract

    The existence of sound-symbolism (or a non-arbitrary link between form and meaning) is well-attested. However, sound-symbolism has mostly been investigated with nonwords in forced choice tasks, neither of which are representative of natural language. This study uses ideophones, which are naturally occurring sound-symbolic words that depict sensory information, to investigate how sensitive Dutch speakers are to sound-symbolism in Japanese in a learning task. Participants were taught 2 sets of Japanese ideophones; 1 set with the ideophones’ real meanings in Dutch, the other set with their opposite meanings. In Experiment 1, participants learned the ideophones and their real meanings much better than the ideophones with their opposite meanings. Moreover, despite the learning rounds, participants were still able to guess the real meanings of the ideophones in a 2-alternative forced-choice test after they were informed of the manipulation. This shows that natural language sound-symbolism is robust beyond 2-alternative forced-choice paradigms and affects broader language processes such as word learning. In Experiment 2, participants learned regular Japanese adjectives with the same manipulation, and there was no difference between real and opposite conditions. This shows that natural language sound-symbolism is especially strong in ideophones, and that people learn words better when form and meaning match. The highlights of this study are as follows: (a) Dutch speakers learn real meanings of Japanese ideophones better than opposite meanings, (b) Dutch speakers accurately guess meanings of Japanese ideophones, (c) this sensitivity happens despite learning some opposite pairings, (d) no such learning effect exists for regular Japanese adjectives, and (e) this shows the importance of sound-symbolism in scaffolding language learning
  • Lockwood, G., Hagoort, P., & Dingemanse, M. (2016). Synthesized Size-Sound Sound Symbolism. In A. Papafragou, D. Grodner, D. Mirman, & J. Trueswell (Eds.), Proceedings of the 38th Annual Meeting of the Cognitive Science Society (CogSci 2016) (pp. 1823-1828). Austin, TX: Cognitive Science Society.

    Abstract

    Studies of sound symbolism have shown that people can associate sound and meaning in consistent ways when presented with maximally contrastive stimulus pairs of nonwords such as bouba/kiki (rounded/sharp) or mil/mal (small/big). Recent work has shown the effect extends to antonymic words from natural languages and has proposed a role for shared cross-modal correspondences in biasing form-to-meaning associations. An important open question is how the associations work, and particularly what the role is of sound-symbolic matches versus mismatches. We report on a learning task designed to distinguish between three existing theories by using a spectrum of sound-symbolically matching, mismatching, and neutral (neither matching nor mismatching) stimuli. Synthesized stimuli allow us to control for prosody, and the inclusion of a neutral condition allows a direct test of competing accounts. We find evidence for a sound-symbolic match boost, but not for a mismatch difficulty compared to the neutral condition.
  • Michalareas, G., Vezoli, J., Van Pelt, S., Schoffelen, J.-M., Kennedy, H., & Fries, P. (2016). Alpha-Beta and Gamma Rhythms Subserve Feedback and Feedforward Influences among Human Visual Cortical Areas. Neuron, 82(2), 384-397. doi:10.1016/j.neuron.2015.12.018.

    Abstract

    Primate visual cortex is hierarchically organized. Bottom-up and top-down influences are exerted through distinct frequency channels, as was recently revealed in macaques by correlating inter-areal influences with laminar anatomical projection patterns. Because this anatomical data cannot be obtained in human subjects, we selected seven homologous macaque and human visual areas, and we correlated the macaque laminar projection patterns to human inter-areal directed influences as measured with magnetoencephalography. We show that influences along feedforward projections predominate in the gamma band, whereas influences along feedback projections predominate in the alpha-beta band. Rhythmic inter-areal influences constrain a functional hierarchy of the seven homologous human visual areas that is in close agreement with the respective macaque anatomical hierarchy. Rhythmic influences allow an extension of the hierarchy to 26 human visual areas including uniquely human brain areas. Hierarchical levels of ventral- and dorsal-stream visual areas are differentially affected by inter-areal influences in the alpha-beta band.
  • De Nooijer, J. A., & Willems, R. M. (2016). What can we learn about cognition from studying handedness? Insights from cognitive neuroscience. In F. Loffing, N. Hagemann, B. Strauss, & C. MacMahon (Eds.), Laterality in sports: Theories and applications (pp. 135-153). Amsterdam: Elsevier.

    Abstract

    Can studying left- and right-handers inform us about cognition? In this chapter, we give an overview of research showing that studying left- and right-handers is informative for understanding the way the brain is organized (i.e., lateralized), as there appear to be differences between left- and right-handers in this respect, but also on the behavioral level handedness studies can provide new insights. According to theories of embodied cognition, our body can influence cognition. Given that left- and right-handers use their bodies differently, this might reflect their performance on an array of cognitive tasks. Indeed, handedness can have an influence on, for instance, what side of space we judge as more positive, the way we gesture, how we remember things, and how we learn new words. Laterality research can, therefore, provide valuable information as to how we act and why
  • Peeters, D. (2016). Processing consequences of onomatopoeic iconicity in spoken language comprehension. In A. Papafragou, D. Grodner, D. Mirman, & J. Trueswell (Eds.), Proceedings of the 38th Annual Meeting of the Cognitive Science Society (CogSci 2016) (pp. 1632-1647). Austin, TX: Cognitive Science Society.

    Abstract

    Iconicity is a fundamental feature of human language. However its processing consequences at the behavioral and neural level in spoken word comprehension are not well understood. The current paper presents the behavioral and electrophysiological outcome of an auditory lexical decision task in which native speakers of Dutch listened to onomatopoeic words and matched control words while their electroencephalogram was recorded. Behaviorally, onomatopoeic words were processed as quickly and accurately as words with an arbitrary mapping between form and meaning. Event-related potentials time-locked to word onset revealed a significant decrease in negative amplitude in the N2 and N400 components and a late positivity for onomatopoeic words in comparison to the control words. These findings advance our understanding of the temporal dynamics of iconic form-meaning mapping in spoken word comprehension and suggest interplay between the neural representations of real-world sounds and spoken words.
  • Peeters, D., & Ozyurek, A. (2016). This and that revisited: A social and multimodal approach to spatial demonstratives. Frontiers in Psychology, 7: 222. doi:10.3389/fpsyg.2016.00222.
  • Poletiek, F. H., Fitz, H., & Bocanegra, B. R. (2016). What baboons can (not) tell us about natural language grammars. Cognition, 151, 108-112. doi:10.1016/j.cognition.2015.04.016.

    Abstract

    Rey et al. (2012) present data from a study with baboons that they interpret in support of the idea that center-embedded structures in human language have their origin in low level memory mechanisms and associative learning. Critically, the authors claim that the baboons showed a behavioral preference that is consistent with center-embedded sequences over other types of sequences. We argue that the baboons’ response patterns suggest that two mechanisms are involved: first, they can be trained to associate a particular response with a particular stimulus, and, second, when faced with two conditioned stimuli in a row, they respond to the most recent one first, copying behavior they had been rewarded for during training. Although Rey et al. (2012) ‘experiment shows that the baboons’ behavior is driven by low level mechanisms, it is not clear how the animal behavior reported, bears on the phenomenon of Center Embedded structures in human syntax. Hence, (1) natural language syntax may indeed have been shaped by low level mechanisms, and (2) the baboons’ behavior is driven by low level stimulus response learning, as Rey et al. propose. But is the second evidence for the first? We will discuss in what ways this study can and cannot give evidential value for explaining the origin of Center Embedded recursion in human grammar. More generally, their study provokes an interesting reflection on the use of animal studies in order to understand features of the human linguistic system.
  • Schoot, L., Heyselaar, E., Hagoort, P., & Segaert, K. (2016). Does syntactic alignment effectively influence how speakers are perceived by their conversation partner. PLoS One, 11(4): e015352. doi:10.1371/journal.pone.0153521.

    Abstract

    The way we talk can influence how we are perceived by others. Whereas previous studies have started to explore the influence of social goals on syntactic alignment, in the current study, we additionally investigated whether syntactic alignment effectively influences conversation partners’ perception of the speaker. To this end, we developed a novel paradigm in which we can measure the effect of social goals on the strength of syntactic alignment for one participant (primed participant), while simultaneously obtaining usable social opinions about them from their conversation partner (the evaluator). In Study 1, participants’ desire to be rated favorably by their partner was manipulated by assigning pairs to a Control (i.e., primed participants did not know they were being evaluated) or Evaluation context (i.e., primed participants knew they were being evaluated). Surprisingly, results showed no significant difference in the strength with which primed participants aligned their syntactic choices with their partners’ choices. In a follow-up study, we used a Directed Evaluation context (i.e., primed participants knew they were being evaluated and were explicitly instructed to make a positive impression). However, again, there was no evidence supporting the hypothesis that participants’ desire to impress their partner influences syntactic alignment. With respect to the influence of syntactic alignment on perceived likeability by the evaluator, a negative relationship was reported in Study 1: the more primed participants aligned their syntactic choices with their partner, the more that partner decreased their likeability rating after the experiment. However, this effect was not replicated in the Directed Evaluation context of Study 2. In other words, our results do not support the conclusion that speakers’ desire to be liked affects how much they align their syntactic choices with their partner, nor is there convincing evidence that there is a reliable relationship between syntactic alignment and perceived likeability.

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  • Schoot, L., Hagoort, P., & Segaert, K. (2016). What can we learn from a two-brain approach to verbal interaction? Neuroscience and Biobehavioral Reviews, 68, 454-459. doi:10.1016/j.neubiorev.2016.06.009.

    Abstract

    Verbal interaction is one of the most frequent social interactions humans encounter on a daily basis. In the current paper, we zoom in on what the multi-brain approach has contributed, and can contribute in the future, to our understanding of the neural mechanisms supporting verbal interaction. Indeed, since verbal interaction can only exist between individuals, it seems intuitive to focus analyses on inter-individual neural markers, i.e. between-brain neural coupling. To date, however, there is a severe lack of theoretically-driven, testable hypotheses about what between-brain neural coupling actually reflects. In this paper, we develop a testable hypothesis in which between-pair variation in between-brain neural coupling is of key importance. Based on theoretical frameworks and empirical data, we argue that the level of between-brain neural coupling reflects speaker-listener alignment at different levels of linguistic and extra-linguistic representation. We discuss the possibility that between-brain neural coupling could inform us about the highest level of inter-speaker alignment: mutual understanding
  • Segaert, K., Wheeldon, L., & Hagoort, P. (2016). Unifying structural priming effects on syntactic choices and timing of sentence generation. Journal of Memory and Language, 91, 59-80. doi:10.1016/j.jml.2016.03.011.

    Abstract

    We investigated whether structural priming of production latencies is sensitive to the same factors known to influence persistence of structural choices: structure preference, cumulativity and verb repetition. In two experiments, we found structural persistence only for passives (inverse preference effect) while priming effects on latencies were stronger for the actives (positive preference effect). We found structural persistence for passives to be influenced by immediate primes and long lasting cumulativity (all preceding primes) (Experiment 1), and to be boosted by verb repetition (Experiment 2). In latencies we found effects for actives were sensitive to long lasting cumulativity (Experiment 1). In Experiment 2, in latencies we found priming for actives overall, while for passives the priming effects emerged as the cumulative exposure increased but only when also aided by verb repetition. These findings are consistent with the Two-stage Competition model, an integrated model of structural priming effects for sentence choice and latency
  • Shitova, N., Roelofs, A., Schriefers, H., Bastiaansen, M., & Schoffelen, J.-M. (2016). Using Brain Potentials to Functionally Localise Stroop-Like Effects in Colour and Picture Naming: Perceptual Encoding versus Word Planning. PLoS One, 11(9): e0161052. doi:10.1371/journal.pone.0161052.

    Abstract

    The colour-word Stroop task and the picture-word interference task (PWI) have been used extensively to study the functional processes underlying spoken word production. One of the consistent behavioural effects in both tasks is the Stroop-like effect: The reaction time (RT) is longer on incongruent trials than on congruent trials. The effect in the Stroop task is usually linked to word planning, whereas the effect in the PWI task is associated with either word planning or perceptual encoding. To adjudicate between the word planning and perceptual encoding accounts of the effect in PWI, we conducted an EEG experiment consisting of three tasks: a standard colour-word Stroop task (three colours), a standard PWI task (39 pictures), and a Stroop-like version of the PWI task (three pictures). Participants overtly named the colours and pictures while their EEG was recorded. A Stroop-like effect in RTs was observed in all three tasks. ERPs at centro-parietal sensors started to deflect negatively for incongruent relative to congruent stimuli around 350 ms after stimulus onset for the Stroop, Stroop-like PWI, and the Standard PWI tasks: an N400 effect. No early differences were found in the PWI tasks. The onset of the Stroop-like effect at about 350 ms in all three tasks links the effect to word planning rather than perceptual encoding, which has been estimated in the literature to be finished around 200–250 ms after stimulus onset. We conclude that the Stroop-like effect arises during word planning in both Stroop and PWI.
  • Silva, S., Petersson, K. M., & Castro, S. (2016). Rhythm in the brain: Is music special? In D. Da Silva Marques, & J. Avila-Toscano (Eds.), Neuroscience to neuropsychology: The study of the human brain (pp. 29-54). Barranquilla, Colombia: Ediciones CUR.
  • Silva, S., Reis, A., Casaca, L., Petersson, K. M., & Faísca, L. (2016). When the eyes no longer lead: Familiarity and length effects eye-voice span. Frontiers in Psychology, 7: 1720. doi:10.3389/fpsyg.2016.01720.

    Abstract

    During oral reading, the eyes tend to be ahead of the voice (eye-voice span, EVS). It has been hypothesized that the extent to which this happens depends on the automaticity of reading processes, namely on the speed of print-to-sound conversion. We tested whether EVS is affected by another automaticity component – immunity from interference. To that end, we manipulated word familiarity (high-frequency, lowfrequency, and pseudowords, PW) and word length as proxies of immunity from interference, and we used linear mixed effects models to measure the effects of both variables on the time interval at which readers do parallel processing by gazing at word N C 1 while not having articulated word N yet (offset EVS). Parallel processing was enhanced by automaticity, as shown by familiarity length interactions on offset EVS, and it was impeded by lack of automaticity, as shown by the transformation of offset EVS into voice-eye span (voice ahead of the offset of the eyes) in PWs. The relation between parallel processing and automaticity was strengthened by the fact that offset EVS predicted reading velocity. Our findings contribute to understand how the offset EVS, an index that is obtained in oral reading, may tap into different components of automaticity that underlie reading ability, oral or silent. In addition, we compared the duration of the offset EVS with the average reference duration of stages in word production, and we saw that the offset EVS may accommodate for more than the articulatory programming stage of word N.
  • Silva, S., Faísca, L., Araújo, S., Casaca, L., Carvalho, L., Petersson, K. M., & Reis, A. (2016). Too little or too much? Parafoveal preview benefits and parafoveal load costs in dyslexic adults. Annals of Dyslexia, 66(2), 187-201. doi:10.1007/s11881-015-0113-z.

    Abstract

    Two different forms of parafoveal dysfunction have been hypothesized as core deficits of dyslexic individuals: reduced parafoveal preview benefits (“too little parafovea”) and increased costs of parafoveal load (“too much parafovea”). We tested both hypotheses in a single eye-tracking experiment using a modified serial rapid automatized naming (RAN) task. Comparisons between dyslexic and non-dyslexic adults showed reduced parafoveal preview benefits in dyslexics, without increased costs of parafoveal load. Reduced parafoveal preview benefits were observed in a naming task, but not in a silent letter-finding task, indicating that the parafoveal dysfunction may be consequent to the overload with extracting phonological information from orthographic input. Our results suggest that dyslexics’ parafoveal dysfunction is not based on strict visuo-attentional factors, but nevertheless they stress the importance of extra-phonological processing. Furthermore, evidence of reduced parafoveal preview benefits in dyslexia may help understand why serial RAN is an important reading predictor in adulthood
  • Takashima, A., Hulzink, I., Wagensveld, B., & Verhoeven, L. (2016). Emergence of representations through repeated training on pronouncing novel letter combinations leads to efficient reading. Neuropsychologia, 89, 14-30. doi:10.1016/j.neuropsychologia.2016.05.014.

    Abstract

    Printed text can be decoded by utilizing different processing routes depending on the familiarity of the script. A predominant use of word-level decoding strategies can be expected in the case of a familiar script, and an almost exclusive use of letter-level decoding strategies for unfamiliar scripts. Behavioural studies have revealed that frequently occurring words are read more efficiently, suggesting that these words are read in a more holistic way at the word-level, than infrequent and unfamiliar words. To test whether repeated exposure to specific letter combinations leads to holistic reading, we monitored both behavioural and neural responses during novel script decoding and examined changes related to repeated exposure. We trained a group of Dutch university students to decode pseudowords written in an unfamiliar script, i.e., Korean Hangul characters. We compared behavioural and neural responses to pronouncing trained versus untrained two-character pseudowords (equivalent to two-syllable pseudowords). We tested once shortly after the initial training and again after a four days' delay that included another training session. We found that trained pseudowords were pronounced faster and more accurately than novel combinations of radicals (equivalent to letters). Imaging data revealed that pronunciation of trained pseudowords engaged the posterior temporo-parietal region, and engagement of this network was predictive of reading efficiency a month later. The results imply that repeated exposure to specific combinations of graphemes can lead to emergence of holistic representations that result in efficient reading. Furthermore, inter-individual differences revealed that good learners retained efficiency more than bad learners one month later

    Additional information

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  • Takashima, A., Van de Ven, F., Kroes, M. C. W., & Fernández, G. (2016). Retrieved emotional context influences hippocampal involvement during recognition of neutral memories. NeuroImage, 143, 280-292. doi:10.1016/j.neuroimage.2016.08.069.

    Abstract

    It is well documented that emotionally arousing experiences are better remembered than mundane events. This is thought to occur through hippocampus-amygdala crosstalk during encoding, consolidation, and retrieval. Here we investigated whether emotional events (context) also cause a memory benefit for simultaneously encoded non-arousing contents and whether this effect persists after a delay via recruitment of a similar hippocampus-amygdala network. Participants studied neutral pictures (content) encoded together with either an arousing or a neutral sound (that served as context) in two study sessions three days apart. Memory was tested in a functional magnetic resonance scanner directly after the second study session. Pictures recognised with high confidence were more often thought to have been associated with an arousing than with a neutral context, irrespective of the veridical source memory. If the retrieved context was arousing, an area in the hippocampus adjacent to the amygdala exhibited heightened activation and this area increased functional connectivity with the parahippocampal gyrus, an area known to process pictures of scenes. These findings suggest that memories can be shaped by the retrieval act. Memory structures may be recruited to a higher degree when an arousing context is retrieved, and this may give rise to confident judgments of recognition for neutral pictures even after a delay
  • Thalmeier, D., Uhlmann, M., Kappen, H. J., & Memmeshiemer, R.-M. (2016). Learning Universal Computations with Spikes. PLoS Computational Biology, 12(6): e1004895. doi:10.1371/journal.pcbi.1004895.

    Abstract

    Providing the neurobiological basis of information processing in higher animals, spiking neural networks must be able to learn a variety of complicated computations, including the generation of appropriate, possibly delayed reactions to inputs and the self-sustained generation of complex activity patterns, e.g. for locomotion. Many such computations require previous building of intrinsic world models. Here we show how spiking neural networks may solve these different tasks. Firstly, we derive constraints under which classes of spiking neural networks lend themselves to substrates of powerful general purpose computing. The networks contain dendritic or synaptic nonlinearities and have a constrained connectivity. We then combine such networks with learning rules for outputs or recurrent connections. We show that this allows to learn even difficult benchmark tasks such as the self-sustained generation of desired low-dimensional chaotic dynamics or memory-dependent computations. Furthermore, we show how spiking networks can build models of external world systems and use the acquired knowledge to control them.
  • Tromp, J., Hagoort, P., & Meyer, A. S. (2016). Pupillometry reveals increased pupil size during indirect request comprehension. Quarterly Journal of Experimental Psychology, 69, 1093-1108. doi:10.1080/17470218.2015.1065282.

    Abstract

    Fluctuations in pupil size have been shown to reflect variations in processing demands during lexical and syntactic processing in language comprehension. An issue that has not received attention is whether pupil size also varies due to pragmatic manipulations. In two pupillometry experiments, we investigated whether pupil diameter was sensitive to increased processing demands as a result of comprehending an indirect request versus a direct statement. Adult participants were presented with 120 picture–sentence combinations that could be interpreted either as an indirect request (a picture of a window with the sentence “it's very hot here”) or as a statement (a picture of a window with the sentence “it's very nice here”). Based on the hypothesis that understanding indirect utterances requires additional inferences to be made on the part of the listener, we predicted a larger pupil diameter for indirect requests than statements. The results of both experiments are consistent with this expectation. We suggest that the increase in pupil size reflects additional processing demands for the comprehension of indirect requests as compared to statements. This research demonstrates the usefulness of pupillometry as a tool for experimental research in pragmatics
  • Van den Hoven, E., Hartung, F., Burke, M., & Willems, R. M. (2016). Individual differences in sensitivity to style during literary reading: Insights from eye-tracking. Collabra, 2(1): 25, pp. 1-16. doi:10.1525/collabra.39.

    Abstract

    Style is an important aspect of literature, and stylistic deviations are sometimes labeled foregrounded, since their manner of expression deviates from the stylistic default. Russian Formalists have claimed that foregrounding increases processing demands and therefore causes slower reading – an effect called retardation. We tested this claim experimentally by having participants read short literary stories while measuring their eye movements. Our results confirm that readers indeed read slower and make more regressions towards foregrounded passages as compared to passages that are not foregrounded. A closer look, however, reveals significant individual differences in sensitivity to foregrounding. Some readers in fact do not slow down at all when reading foregrounded passages. The slowing down effect for literariness was related to a slowing down effect for high perplexity (unexpected) words: those readers who slowed down more during literary passages also slowed down more during high perplexity words, even though no correlation between literariness and perplexity existed in the stories. We conclude that individual differences play a major role in processing of literary texts and argue for accounts of literary reading that focus on the interplay between reader and text.
  • Van den Broek, G., Takashima, A., Wiklund-Hörnqvist, C., Karlsson-Wirebring, C., Segers, E., Verhoeven, L., & Nyberg, L. (2016). Neurocognitive mechanisms of the “testing effect”: A review. Trends in Neuroscience and Education, 5(2), 52-66. doi:10.1016/j.tine.2016.05.001.

    Abstract

    Memory retrieval is an active process that can alter the content and accessibility of stored memories. Of potential relevance for educational practice are findings that memory retrieval fosters better retention than mere studying. This so-called testing effect has been demonstrated for different materials and populations, but there is limited consensus on the neurocognitive mechanisms involved. In this review, we relate cognitive accounts of the testing effect to findings from recent brain-imaging studies to identify neurocognitive factors that could explain the testing effect. Results indicate that testing facilitates later performance through several processes, including effects on semantic memory representations, the selective strengthening of relevant associations and inhibition of irrelevant associations, as well as potentiation of subsequent learning.
  • Van der Ven, F., Takashima, A., Segers, E., Fernández, G., & Verhoeven, L. (2016). Non-symbolic and symbolic notation in simple arithmetic differentially involve intraparietal sulcus and angular gyrus activity. Brain Research, 1643, 91-102.

    Abstract

    Addition problems can be solved by mentally manipulating quantities for which the bilateral intraparietal sulcus (IPS) is likely recruited, or by retrieving the answer directly from fact memory in which the left angular gyrus (AG) and perisylvian areas may play a role. Mental addition is usually studied with problems presented in the Arabic notation (4+2), and less so with number words (four+two) or dots (:: +·.). In the present study, we investigated how the notation of numbers influences processing during simple mental arithmetic. Twenty-five highly educated participants performed simple arithmetic while their brain activity was recorded with functional magnetic resonance imaging. To reveal the effect of number notation, arithmetic problems were presented in a non-symbolic (Dots) or symbolic (Arabic; Words) notation. Furthermore, we asked whether IPS processing during mental arithmetic is magnitude specific or of a more general, visuospatial nature. To this end, we included perception and manipulation of non-magnitude formats (Colors; unfamiliar Japanese Characters). Increased IPS activity was observed, suggesting magnitude calculations during addition of non-symbolic numbers. In contrast, there was greater activity in the AG and perisylvian areas for symbolic compared to non-symbolic addition, suggesting increased verbal fact retrieval. Furthermore, IPS activity was not specific to processing of numerical magnitude but also present for non-magnitude stimuli that required mental visuospatial processing (Color-mixing; Character-memory measured by a delayed match-to-sample task). Together, our data suggest that simple non-symbolic sums are calculated using visual imagery, whereas answers for simple symbolic sums are retrieved from verbal memory.
  • Vanlangendonck, F., Willems, R. M., Menenti, L., & Hagoort, P. (2016). An early influence of common ground during speech planning. Language, Cognition and Neuroscience, 31(6), 741-750. doi:10.1080/23273798.2016.1148747.

    Abstract

    In order to communicate successfully, speakers have to take into account which information they share with their addressee, i.e. common ground. In the current experiment we investigated how and when common ground affects speech planning by tracking speakers’ eye movements while they played a referential communication game. We found evidence that common ground exerts an early, but incomplete effect on speech planning. In addition, we did not find longer planning times when speakers had to take common ground into account, suggesting that taking common ground into account is not necessarily an effortful process. Common ground information thus appears to act as a partial constraint on language production that is integrated flexibly and efficiently in the speech planning process.
  • Weber, K., Christiansen, M., Petersson, K. M., Indefrey, P., & Hagoort, P. (2016). fMRI syntactic and lexical repetition effects reveal the initial stages of learning a new language. The Journal of Neuroscience, 36, 6872-6880. doi:10.1523/JNEUROSCI.3180-15.2016.

    Abstract

    When learning a new language, we build brain networks to process and represent the acquired words and syntax and integrate these with existing language representations. It is an open question whether the same or different neural mechanisms are involved in learning and processing a novel language compared to the native language(s). Here we investigated the neural repetition effects of repeating known and novel word orders while human subjects were in the early stages of learning a new language. Combining a miniature language with a syntactic priming paradigm, we examined the neural correlates of language learning online using functional magnetic resonance imaging (fMRI). In left inferior frontal gyrus (LIFG) and posterior temporal cortex the repetition of novel syntactic structures led to repetition enhancement, while repetition of known structures resulted in repetition suppression. Additional verb repetition led to an
    increase in the syntactic repetition enhancement effect in language-related brain regions. Similarly the repetition of verbs led to repetition enhancement effects in areas related to lexical and semantic processing, an effect that continued to increase in a subset of these regions. Repetition enhancement might reflect a mechanism to build and strengthen a neural network to process novel syntactic structures and lexical items. By contrast, the observed repetition suppression points to overlapping neural mechanisms for native and new language constructions when these have sufficient structural similarities.
  • Weber, K., Luther, L., Indefrey, P., & Hagoort, P. (2016). Overlap and differences in brain networks underlying the processing of complex sentence structures in second language users compared to native speakers. Brain Connectivity, 6(4), 345-355. doi:10.1089/brain.2015.0383.

    Abstract

    When we learn a second language later in life do we integrate it with the established neural networks in place for the first language or is at least a partially new network recruited? While there is evidence that simple grammatical structures in a second language share a system with the native language, the story becomes more multifaceted for complex sentence structures. In this study we investigated the underlying brain networks in native speakers compared to proficient second language users while processing complex sentences. As hypothesized, complex structures were processed by the same large-scale inferior frontal and middle temporal language networks of the brain in the second language, as seen in native speakers. These effects were seen both in activations as well as task-related connectivity patterns. Furthermore, the second language users showed increased task-related connectivity from inferior frontal to inferior parietal regions of the brain, regions related to attention and cognitive control, suggesting less automatic processing for these structures in a second language.
  • Weber, K., Lau, E., Stillerman, B., & Kuperberg, G. (2016). The Yin and the Yang of Prediction: An fMRI Study of Semantic Predictive Processing. PLoS One, 11(3): 0148637. doi:10.1371/journal.pone.0148637.

    Abstract

    Probabilistic prediction plays a crucial role in language comprehension. When predictions are fulfilled, the resulting facilitation allows for fast, efficient processing of ambiguous, rapidly-unfolding input; when predictions are not fulfilled, the resulting error signal allows us to adapt to broader statistical changes in this input. We used functional Magnetic Resonance Imaging to examine the neuroanatomical networks engaged in semantic predictive processing and adaptation. We used a relatedness proportion semantic priming paradigm, in which we manipulated the probability of predictions while holding local semantic context constant. Under conditions of higher (versus lower) predictive validity, we replicate previous observations of reduced activity to semantically predictable words in the left anterior superior/middle temporal cortex, reflecting facilitated processing of targets that are consistent with prior semantic predictions. In addition, under conditions of higher (versus lower) predictive validity we observed significant differences in the effects of semantic relatedness within the left inferior frontal gyrus and the posterior portion of the left superior/middle temporal gyrus. We suggest that together these two regions mediated the suppression of unfulfilled semantic predictions and lexico-semantic processing of unrelated targets that were inconsistent with these predictions. Moreover, under conditions of higher (versus lower) predictive validity, a functional connectivity analysis showed that the left inferior frontal and left posterior superior/middle temporal gyrus were more tightly interconnected with one another, as well as with the left anterior cingulate cortex. The left anterior cingulate cortex was, in turn, more tightly connected to superior lateral frontal cortices and subcortical regions—a network that mediates rapid learning and adaptation and that may have played a role in switching to a more predictive mode of processing in response to the statistical structure of the wider environmental context. Together, these findings highlight close links between the networks mediating semantic prediction, executive function and learning, giving new insights into how our brains are able to flexibly adapt to our environment.

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    Data availability
  • Willems, R. M., & Jacobs, A. M. (2016). Caring about Dostoyevsky: The untapped potential of studying literature. Trends in Cognitive Sciences, 20(4), 243-245. doi:10.1016/j.tics.2015.12.009.

    Abstract

    Should cognitive scientists and neuroscientists care about Dostoyevsky? Engaging with fiction is a natural and rich behavior, providing a unique window onto the mind and brain, particularly for mental simulation, emotion, empathy, and immersion. With advances in analysis techniques, it is time that cognitive scientists and neuroscientists embrace literature and fiction.
  • Willems, R. M., Frank, S. L., Nijhoff, A. D., Hagoort, P., & Van den Bosch, A. (2016). Prediction during natural language comprehension. Cerebral Cortex, 26(6), 2506-2516. doi:10.1093/cercor/bhv075.

    Abstract

    The notion of prediction is studied in cognitive neuroscience with increasing intensity. We investigated the neural basis of 2 distinct aspects of word prediction, derived from information theory, during story comprehension. We assessed the effect of entropy of next-word probability distributions as well as surprisal. A computational model determined entropy and surprisal for each word in 3 literary stories. Twenty-four healthy participants listened to the same 3 stories while their brain activation was measured using fMRI. Reversed speech fragments were presented as a control condition. Brain areas sensitive to entropy were left ventral premotor cortex, left middle frontal gyrus, right inferior frontal gyrus, left inferior parietal lobule, and left supplementary motor area. Areas sensitive to surprisal were left inferior temporal sulcus (“visual word form area”), bilateral superior temporal gyrus, right amygdala, bilateral anterior temporal poles, and right inferior frontal sulcus. We conclude that prediction during language comprehension can occur at several levels of processing, including at the level of word form. Our study exemplifies the power of combining computational linguistics with cognitive neuroscience, and additionally underlines the feasibility of studying continuous spoken language materials with fMRI.

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    Supplementary Material
  • Zimmermann, M., Verhagen, L., De Lange, F., & Toni, I. (2016). The extrastriate body area computes desired goal states during action planning. eNeuro, 3(2): ENEURO.0020-16.2016. doi:10.1523/ENEURO.0020-16.2016.

    Abstract

    How do object perception and action interact at a neural level? Here we test the hypothesis that perceptual
    features, processed by the ventral visuoperceptual stream, are used as priors by the dorsal visuomotor stream to
    specify goal-directed grasping actions. We present three main findings, which were obtained by combining
    time-resolved transcranial magnetic stimulation and kinematic tracking of grasp-and-rotate object manipulations,
    in a group of healthy human participants (N 22). First, the extrastriate body area (EBA), in the ventral stream,
    provides an initial structure to motor plans, based on current and desired states of a grasped object and of the
    grasping hand. Second, the contributions of EBA are earlier in time than those of a caudal intraparietal region
    known to specify the action plan. Third, the contributions of EBA are particularly important when desired and
    current object configurations differ, and multiple courses of actions are possible. These findings specify the
    temporal and functional characteristics for a mechanism that integrates perceptual processing with motor
    planning.
  • Araújo, S., Faísca, L., Bramão, I., Reis, A., & Petersson, K. M. (2015). Lexical and sublexical orthographic processing: An ERP study with skilled and dyslexic adult readers. Brain and Language, 141, 16-27. doi:10.1016/j.bandl.2014.11.007.

    Abstract

    This ERP study investigated the cognitive nature of the P1–N1 components during orthographic processing. We used an implicit reading task with various types of stimuli involving different amounts of sublexical or lexical orthographic processing (words, pseudohomophones, pseudowords, nonwords, and symbols), and tested average and dyslexic readers. An orthographic regularity effect (pseudowords– nonwords contrast) was observed in the average but not in the dyslexic group. This suggests an early sensitivity to the dependencies among letters in word-forms that reflect orthographic structure, while the dyslexic brain apparently fails to be appropriately sensitive to these complex features. Moreover, in the adults the N1-response may already reflect lexical access: (i) the N1 was sensitive to the familiar vs. less familiar orthographic sequence contrast; (ii) and early effects of the phonological form (words-pseudohomophones contrast) were also found. Finally, the later N320 component was attenuated in the dyslexics, suggesting suboptimal processing in later stages of phonological analysis.
  • Araújo, S., Reis, A., Petersson, K. M., & Faísca, L. (2015). Rapid automatized naming and reading performance: A meta-analysis. Journal of Educational Psychology, 107(3), 868-883. doi:10.1037/edu0000006.

    Abstract

    Evidence that rapid naming skill is associated with reading ability has become increasingly prevalent in recent years. However, there is considerable variation in the literature concerning the magnitude of this relationship. The objective of the present study was to provide a comprehensive analysis of the evidence on the relationship between rapid automatized naming (RAN) and reading performance. To this end, we conducted a meta-analysis of the correlational relationship between these 2 constructs to (a) determine the overall strength of the RAN–reading association and (b) identify variables that systematically moderate this relationship. A random-effects model analysis of data from 137 studies (857 effect sizes; 28,826 participants) indicated a moderate-to-strong relationship between RAN and reading performance (r = .43, I2 = 68.40). Further analyses revealed that RAN contributes to the 4 measures of reading (word reading, text reading, non-word reading, and reading comprehension), but higher coefficients emerged in favor of real word reading and text reading. RAN stimulus type and type of reading score were the factors with the greatest moderator effect on the magnitude of the RAN–reading relationship. The consistency of orthography and the subjects’ grade level were also found to impact this relationship, although the effect was contingent on reading outcome. It was less evident whether the subjects’ reading proficiency played a role in the relationship. Implications for future studies are discussed.
  • Asaridou, S. S., Hagoort, P., & McQueen, J. M. (2015). Effects of early bilingual experience with a tone and a non-tone language on speech-music. PLoS One, 10(12): e0144225. doi:10.1371/journal.pone.0144225.

    Abstract

    We investigated music and language processing in a group of early bilinguals who spoke a tone language and a non-tone language (Cantonese and Dutch). We assessed online speech-music processing interactions, that is, interactions that occur when speech and music are processed simultaneously in songs, with a speeded classification task. In this task, participants judged sung pseudowords either musically (based on the direction of the musical interval) or phonologically (based on the identity of the sung vowel). We also assessed longer-term effects of linguistic experience on musical ability, that is, the influence of extensive prior experience with language when processing music. These effects were assessed with a task in which participants had to learn to identify musical intervals and with four pitch-perception tasks. Our hypothesis was that due to their experience in two different languages using lexical versus intonational tone, the early Cantonese-Dutch bilinguals would outperform the Dutch control participants. In online processing, the Cantonese-Dutch bilinguals processed speech and music more holistically than controls. This effect seems to be driven by experience with a tone language, in which integration of segmental and pitch information is fundamental. Regarding longer-term effects of linguistic experience, we found no evidence for a bilingual advantage in either the music-interval learning task or the pitch-perception tasks. Together, these results suggest that being a Cantonese-Dutch bilingual does not have any measurable longer-term effects on pitch and music processing, but does have consequences for how speech and music are processed jointly.

    Additional information

    Data Availability
  • Asaridou, S. S. (2015). An ear for pitch: On the effects of experience and aptitude in processing pitch in language and music. PhD Thesis, Radboud University Nijmegen, Nijmegen.
  • Baggio, G., van Lambalgen, M., & Hagoort, P. (2015). Logic as Marr's computational level: Four case studies. Topics in Cognitive Science, 7, 287-298. doi:10.1111/tops.12125.

    Abstract

    We sketch four applications of Marr's levels-of-analysis methodology to the relations between logic and experimental data in the cognitive neuroscience of language and reasoning. The first part of the paper illustrates the explanatory power of computational level theories based on logic. We show that a Bayesian treatment of the suppression task in reasoning with conditionals is ruled out by EEG data, supporting instead an analysis based on defeasible logic. Further, we describe how results from an EEG study on temporal prepositions can be reanalyzed using formal semantics, addressing a potential confound. The second part of the article demonstrates the predictive power of logical theories drawing on EEG data on processing progressive constructions and on behavioral data on conditional reasoning in people with autism. Logical theories can constrain processing hypotheses all the way down to neurophysiology, and conversely neuroscience data can guide the selection of alternative computational level models of cognition.
  • Bakker, I., Takashima, A., van Hell, J. G., Janzen, G., & McQueen, J. M. (2015). Tracking lexical consolidation with ERPs: Lexical and semantic-priming effects on N400 and LPC responses to newly-learned words. Neuropsychologia, 79, 33-41. doi:10.1016/j.neuropsychologia.2015.10.020.
  • Bašnákova, J., Van Berkum, J. J. A., Weber, K., & Hagoort, P. (2015). A job interview in the MRI scanner: How does indirectness affect addressees and overhearers? Neuropsychologia, 76, 79-91. doi:10.1016/j.neuropsychologia.2015.03.030.

    Abstract

    In using language, people not only exchange information, but also navigate their social world – for example, they can express themselves indirectly to avoid losing face. In this functional magnetic resonance imaging study, we investigated the neural correlates of interpreting face-saving indirect replies, in a situation where participants only overheard the replies as part of a conversation between two other people, as well as in a situation where the participants were directly addressed themselves. We created a fictional job interview context where indirect replies serve as a natural communicative strategy to attenuate one’s shortcomings, and asked fMRI participants to either pose scripted questions and receive answers from three putative job candidates (addressee condition) or to listen to someone else interview the same candidates (overhearer condition). In both cases, the need to evaluate the candidate ensured that participants had an active interest in comprehending the replies. Relative to direct replies, face-saving indirect replies increased activation in medial prefrontal cortex, bilateral temporo-parietal junction (TPJ), bilateral inferior frontal gyrus and bilateral middle temporal gyrus, in active overhearers and active addressees alike, with similar effect size, and comparable to findings obtained in an earlier passive listening study (Bašnáková et al., 2013). In contrast, indirectness effects in bilateral anterior insula and pregenual ACC, two regions implicated in emotional salience and empathy, were reliably stronger in addressees than in active overhearers. Our findings indicate that understanding face-saving indirect language requires additional cognitive perspective-taking and other discourse-relevant cognitive processing, to a comparable extent in active overhearers and addressees. Furthermore, they indicate that face-saving indirect language draws upon affective systems more in addressees than in overhearers, presumably because the addressee is the one being managed by a face-saving reply. In all, face-saving indirectness provides a window on the cognitive as well as affect-related neural systems involved in human communication.
  • Bastiaansen, M. C. M., & Hagoort, P. (2015). Frequency-based segregation of syntactic and semantic unification during online sentence level language comprehension. Journal of Cognitive Neuroscience, 27(11), 2095-2107. doi:10.1162/jocn_a_00829.

    Abstract

    During sentence level language comprehension, semantic and syntactic unification are functionally distinct operations. Nevertheless, both recruit roughly the same brain areas (spatially overlapping networks in the left frontotemporal cortex) and happen at the same time (in the first few hundred milliseconds after word onset). We tested the hypothesis that semantic and syntactic unification are segregated by means of neuronal synchronization of the functionally relevant networks in different frequency ranges: gamma (40 Hz and up) for semantic unification and lower beta (10–20 Hz) for syntactic unification. EEG power changes were quantified as participants read either correct sentences, syntactically correct though meaningless sentences (syntactic prose), or sentences that did not contain any syntactic structure (random word lists). Other sentences contained either a semantic anomaly or a syntactic violation at a critical word in the sentence. Larger EEG gamma-band power was observed for semantically coherent than for semantically anomalous sentences. Similarly, beta-band power was larger for syntactically correct sentences than for incorrect ones. These results confirm the existence of a functional dissociation in EEG oscillatory dynamics during sentence level language comprehension that is compatible with the notion of a frequency-based segregation of syntactic and semantic unification.
  • Bastos, A. M., Vezoli, J., Bosman, C. A., Schoffelen, J.-M., Oostenveld, R., Dowdall, J. R., De Weerd, P., Kennedy, H., & Fries, P. (2015). Visual areas exert feedforward and feedback influences through distinct frequency channels. Neuron, 85(2), 390-401. doi:10.1016/j.neuron.2014.12.018.

    Abstract

    Visual cortical areas subserve cognitive functions by interacting in both feedforward and feedback directions. While feedforward influences convey sensory signals, feedback influences modulate feedforward signaling according to the current behavioral context. We investigated whether these interareal influences are subserved differentially by rhythmic synchronization. We correlated frequency-specific directed influences among 28 pairs of visual areas with anatomical metrics of the feedforward or feedback character of the respective interareal projections. This revealed that in the primate visual system, feedforward influences are carried by theta-band ( approximately 4 Hz) and gamma-band ( approximately 60-80 Hz) synchronization, and feedback influences by beta-band ( approximately 14-18 Hz) synchronization. The functional directed influences constrain a functional hierarchy similar to the anatomical hierarchy, but exhibiting task-dependent dynamic changes in particular with regard to the hierarchical positions of frontal areas. Our results demonstrate that feedforward and feedback signaling use distinct frequency channels, suggesting that they subserve differential communication requirements.
  • Chang, F., Bauman, M., Pappert, S., & Fitz, H. (2015). Do lemmas speak German?: A verb position effect in German structural priming. Cognitive Science, 39(5), 1113-1130. doi:10.1111/cogs.12184.

    Abstract

    Lexicalized theories of syntax often assume that verb-structure regularities are mediated by lemmas, which abstract over variation in verb tense and aspect. German syntax seems to challenge this assumption, because verb position depends on tense and aspect. To examine how German speakers link these elements, a structural priming study was performed which varied syntactic structure, verb position (encoded by tense and aspect), and verb overlap. Abstract structural priming was found, both within and across verb position, but priming was larger when the verb position was the same between prime and target. Priming was boosted by verb overlap, but there was no interaction with verb position. The results can be explained by a lemma model where tense and aspect are linked to structural choices in German. Since the architecture of this lemma model is not consistent with results from English, a connectionist model was developed which could explain the cross-linguistic variation in the production system. Together, these findings support the view that language learning plays an important role in determining the nature of structural priming in different languages
  • Cronin, K. A., Acheson, D. J., Hernández, P., & Sánchez, A. (2015). Hierarchy is Detrimental for Human Cooperation. Scientific Reports, 5: 18634. doi:10.1038/srep18634.

    Abstract

    Studies of animal behavior consistently demonstrate that the social environment impacts cooperation, yet the effect of social dynamics has been largely excluded from studies of human cooperation. Here, we introduce a novel approach inspired by nonhuman primate research to address how social hierarchies impact human cooperation. Participants competed to earn hierarchy positions and then could cooperate with another individual in the hierarchy by investing in a common effort. Cooperation was achieved if the combined investments exceeded a threshold, and the higher ranked individual distributed the spoils unless control was contested by the partner. Compared to a condition lacking hierarchy, cooperation declined in the presence of a hierarchy due to a decrease in investment by lower ranked individuals. Furthermore, hierarchy was detrimental to cooperation regardless of whether it was earned or arbitrary. These findings mirror results from nonhuman primates and demonstrate that hierarchies are detrimental to cooperation. However, these results deviate from nonhuman primate findings by demonstrating that human behavior is responsive to changing hierarchical structures and suggests partnership dynamics that may improve cooperation. This work introduces a controlled way to investigate the social influences on human behavior, and demonstrates the evolutionary continuity of human behavior with other primate species.
  • Flecken, M., Carroll, M., Weimar, K., & Von Stutterheim, C. (2015). Driving along the road or heading for the village? Conceptual differences underlying motion event encoding in French, German, and French-German L2 users. Modern Language Journal, 99(S1), 100-122. doi:10.1111/j.1540-4781.2015.12181.x.

    Abstract

    The typological contrast between verb- and satellite-framed languages (Talmy, 1985) has set the basis for many empirical studies on L2 acquisition. The current analysis goes beyond this typology by looking in detail at the conceptualization of the path of motion in a motion event. We take as a starting point the cognitive salience of specific elements of motion events that are relevant when conceptualizing space. When expressing direction in French, specific spatial relations involving the entity in motion (its alignment and its distance toward a [potential] endpoint) are relevant, given a variety of path verbs in the lexicon expressing this information (e.g., se diriger vers, avancer to direct oneself toward,' to advance'). This is not the case in German (manner verbs in the lexicon mainly). In German, spatial information is packaged in adjuncts and particles and the path of motion is typically structured via features of the ground (entlanglaufen/fahren to walk/drive along') or endpoints (to walk/drive to/toward'). We investigate those fundamental differences in spatial conceptualization in French and German, as reflected in pre-articulatory patterns of attention allocation (measured with eye tracking) to moving entities and endpoints in motion scenes in an event description task. Our focus is on spatial conceptualization in an L2 (French L2 users of German), analyzing the extent to which these L2 users display target-like patterns or traces of L1 conceptualization transfer. Findings show that, in line with directional concepts expressed in verbs, L1 French speakers allocate more attention to entities in motion and endpoints, before utterance onset, than L1 German speakers do. The L2 German speakers pattern with L1 German speakers in the use of manner verbs, but they have not fully acquired the spatial concepts and means that structure the path of motion in the L2. This is reflected in pre-articulatory attention allocation patterns, according to which the L2 speakers pattern with native speakers of their L1 (French). The findings show a continued deep entrenchment of L1-based processing patterns and spatial frames of reference when speakers prepare for speech in an L2
  • Flecken, M., Walbert, K., & Dijkstra, T. (2015). ‘Right now, Sophie ∗swims in the pool?!’: Brain potentials of grammatical aspect processing. Frontiers in Psychology, 6: 1764. doi:10.3389/fpsyg.2015.01764.

    Abstract

    We investigated whether brain potentials of grammatical aspect processing resemble semantic or morpho-syntactic processing, or whether they instead are characterized by an entirely distinct pattern in the same individuals. We studied aspect from the perspective of agreement between the temporal information in the context (temporal adverbials, e.g., Right now) and a morpho-syntactic marker of grammatical aspect (e.g., progressive is swimming). Participants read questions providing a temporal context that was progressive (What is Sophie doing in the pool right now?) or habitual (What does Sophie do in the pool every Monday?). Following a lead-in sentence context such as Right now, Sophie…, we measured event-related brain potentials (ERPs) time-locked to verb phrases in four different conditions, e.g., (a) is swimming (control); (b) ∗is cooking (semantic violation); (c) ∗are swimming (morpho-syntactic violation); or (d)?swims (aspect mismatch); …in the pool.” The collected ERPs show typical N400 and P600 effects for semantics and morpho-syntax, while aspect processing elicited an Early Negativity (250–350 ms). The aspect-related Negativity was short-lived and had a central scalp distribution with an anterior onset. This differentiates it not only from the semantic N400 effect, but also from the typical LAN (Left Anterior Negativity), that is frequently reported for various types of agreement processing. Moreover, aspect processing did not show a clear P600 modulation. We argue that the specific context for each item in this experiment provided a trigger for agreement checking with temporal information encoded on the verb, i.e., morphological aspect marking. The aspect-related Negativity obtained for aspect agreement mismatches reflects a violated expectation concerning verbal inflection (in the example above, the expected verb phrase was Sophie is X-ing rather than Sophie X-s in condition d). The absence of an additional P600 for aspect processing suggests that the mismatch did not require additional reintegration or processing costs. This is consistent with participants’ post hoc grammaticality judgements of the same sentences, which overall show a high acceptability of aspect mismatch sentences.

    Additional information

    data sheet 1.docx
  • Flecken, M., Athanasopoulos, P., Kuipers, J. R., & Thierry, G. (2015). On the road to somewhere: Brain potentials reflect language effects on motion event perception. Cognition, 141, 41-51. doi:10.1016/j.cognition.2015.04.006.

    Abstract

    Recent studies have identified neural correlates of language effects on perception in static domains of experience such as colour and objects. The generalization of such effects to dynamic domains like motion events remains elusive. Here, we focus on grammatical differences between languages relevant for the description of motion events and their impact on visual scene perception. Two groups of native speakers of German or English were presented with animated videos featuring a dot travelling along a trajectory towards a geometrical shape (endpoint). English is a language with grammatical aspect in which attention is drawn to trajectory and endpoint of motion events equally. German, in contrast, is a non-aspect language which highlights endpoints. We tested the comparative perceptual saliency of trajectory and endpoint of motion events by presenting motion event animations (primes) followed by a picture symbolising the event (target): In 75% of trials, the animation was followed by a mismatching picture (both trajectory and endpoint were different); in 10% of trials, only the trajectory depicted in the picture matched the prime; in 10% of trials, only the endpoint matched the prime; and in 5% of trials both trajectory and endpoint were matching, which was the condition requiring a response from the participant. In Experiment 1 we recorded event-related brain potentials elicited by the picture in native speakers of German and native speakers of English. German participants exhibited a larger P3 wave in the endpoint match than the trajectory match condition, whereas English speakers showed no P3 amplitude difference between conditions. In Experiment 2 participants performed a behavioural motion matching task using the same stimuli as those used in Experiment 1. German and English participants did not differ in response times showing that motion event verbalisation cannot readily account for the difference in P3 amplitude found in the first experiment. We argue that, even in a non-verbal context, the grammatical properties of the native language and associated sentence-level patterns of event encoding influence motion event perception, such that attention is automatically drawn towards aspects highlighted by the grammar.
  • Francken, J. C., Meijs, E. L., Ridderinkhof, O. M., Hagoort, P., de Lange, F. P., & van Gaal, S. (2015). Manipulating word awareness dissociates feed-forward from feedback models of language-perception interactions. Neuroscience of consciousness, 1. doi:10.1093/nc/niv003.

    Abstract

    Previous studies suggest that linguistic material can modulate visual perception, but it is unclear at which level of processing these interactions occur. Here we aim to dissociate between two competing models of language–perception interactions: a feed-forward and a feedback model. We capitalized on the fact that the models make different predictions on the role of feedback. We presented unmasked (aware) or masked (unaware) words implying motion (e.g. “rise,” “fall”), directly preceding an upward or downward visual motion stimulus. Crucially, masking leaves intact feed-forward information processing from low- to high-level regions, whereas it abolishes subsequent feedback. Under this condition, participants remained faster and more accurate when the direction implied by the motion word was congruent with the direction of the visual motion stimulus. This suggests that language–perception interactions are driven by the feed-forward convergence of linguistic and perceptual information at higher-level conceptual and decision stages.
  • Francken, J. C., Meijs, E. L., Hagoort, P., van Gaal, S., & de Lange, F. P. (2015). Exploring the automaticity of language-perception interactions: Effects of attention and awareness. Scientific Reports, 5: 17725. doi:10.1038/srep17725.

    Abstract

    Previous studies have shown that language can modulate visual perception, by biasing and/
    or enhancing perceptual performance. However, it is still debated where in the brain visual and
    linguistic information are integrated, and whether the effects of language on perception are
    automatic and persist even in the absence of awareness of the linguistic material. Here, we aimed
    to explore the automaticity of language-perception interactions and the neural loci of these
    interactions in an fMRI study. Participants engaged in a visual motion discrimination task (upward
    or downward moving dots). Before each trial, a word prime was briefly presented that implied
    upward or downward motion (e.g., “rise”, “fall”). These word primes strongly influenced behavior:
    congruent motion words sped up reaction times and improved performance relative to incongruent
    motion words. Neural congruency effects were only observed in the left middle temporal gyrus,
    showing higher activity for congruent compared to incongruent conditions. This suggests that higherlevel
    conceptual areas rather than sensory areas are the locus of language-perception interactions.
    When motion words were rendered unaware by means of masking, they still affected visual motion
    perception, suggesting that language-perception interactions may rely on automatic feed-forward
    integration of perceptual and semantic material in language areas of the brain.
  • Francken, J. C., Kok, P., Hagoort, P., & De Lange, F. P. (2015). The behavioral and neural effects of language on motion perception. Journal of Cognitive Neuroscience, 27(1), 175-184. doi:10.1162/jocn_a_00682.

    Abstract

    Perception does not function as an isolated module but is tightly linked with other cognitive functions. Several studies have demonstrated an influence of language on motion perception, but it remains debated at which level of processing this modulation takes place. Some studies argue for an interaction in perceptual areas, but it is also possible that the interaction is mediated by "language areas" that integrate linguistic and visual information. Here, we investigated whether language-perception interactions were specific to the language-dominant left hemisphere by comparing the effects of language on visual material presented in the right (RVF) and left visual fields (LVF). Furthermore, we determined the neural locus of the interaction using fMRI. Participants performed a visual motion detection task. On each trial, the visual motion stimulus was presented in either the LVF or in the RVF, preceded by a centrally presented word (e.g., "rise"). The word could be congruent, incongruent, or neutral with regard to the direction of the visual motion stimulus that was presented subsequently. Participants were faster and more accurate when the direction implied by the motion word was congruent with the direction of the visual motion stimulus. Interestingly, the speed benefit was present only for motion stimuli that were presented in the RVF. We observed a neural counterpart of the behavioral facilitation effects in the left middle temporal gyrus, an area involved in semantic processing of verbal material. Together, our results suggest that semantic information about motion retrieved in language regions may automatically modulate perceptual decisions about motion.
  • Franken, M. K., McQueen, J. M., Hagoort, P., & Acheson, D. J. (2015). Assessing the link between speech perception and production through individual differences. In Proceedings of the 18th International Congress of Phonetic Sciences. Glasgow: the University of Glasgow.

    Abstract

    This study aims to test a prediction of recent
    theoretical frameworks in speech motor control: if speech production targets are specified in auditory
    terms, people with better auditory acuity should have more precise speech targets.
    To investigate this, we had participants perform speech perception and production tasks in a counterbalanced order. To assess speech perception acuity, we used an adaptive speech discrimination
    task. To assess variability in speech production, participants performed a pseudo-word reading task; formant values were measured for each recording.
    We predicted that speech production variability to correlate inversely with discrimination performance.
    The results suggest that people do vary in their production and perceptual abilities, and that better discriminators have more distinctive vowel production targets, confirming our prediction. This
    study highlights the importance of individual
    differences in the study of speech motor control, and sheds light on speech production-perception interaction.
  • Franken, M. K., Hagoort, P., & Acheson, D. J. (2015). Modulations of the auditory M100 in an Imitation Task. Brain and Language, 142, 18-23. doi:10.1016/j.bandl.2015.01.001.

    Abstract

    Models of speech production explain event-related suppression of the auditory cortical
    response as reflecting a comparison between auditory predictions and feedback. The present MEG
    study was designed to test two predictions from this framework: 1) whether the reduced auditory
    response varies as a function of the mismatch between prediction and feedback; 2) whether individual
    variation in this response is predictive of speech-motor adaptation.
    Participants alternated between online imitation and listening tasks. In the imitation task, participants
    began each trial producing the same vowel (/e/) and subsequently listened to and imitated auditorilypresented
    vowels varying in acoustic distance from /e/.
    Results replicated suppression, with a smaller M100 during speaking than listening. Although we did
    not find unequivocal support for the first prediction, participants with less M100 suppression were
    better at the imitation task. These results are consistent with the enhancement of M100 serving as an
    error signal to drive subsequent speech-motor adaptation.
  • Guadalupe, T., Zwiers, M. P., Wittfeld, K., Teumer, A., Vasquez, A. A., Hoogman, M., Hagoort, P., Fernandez, G., Buitelaar, J., van Bokhoven, H., Hegenscheid, K., Völzke, H., Franke, B., Fisher, S. E., Grabe, H. J., & Francks, C. (2015). Asymmetry within and around the human planum temporale is sexually dimorphic and influenced by genes involved in steroid hormone receptor activity. Cortex, 62, 41-55. doi:10.1016/j.cortex.2014.07.015.

    Abstract

    The genetic determinants of cerebral asymmetries are unknown. Sex differences in asymmetry of the planum temporale, that overlaps Wernicke’s classical language area, have been inconsistently reported. Meta-analysis of previous studies has suggested that publication bias established this sex difference in the literature. Using probabilistic definitions of cortical regions we screened over the cerebral cortex for sexual dimorphisms of asymmetry in 2337 healthy subjects, and found the planum temporale to show the strongest sex-linked asymmetry of all regions, which was supported by two further datasets, and also by analysis with the Freesurfer package that performs automated parcellation of cerebral cortical regions. We performed a genome-wide association scan meta-analysis of planum temporale asymmetry in a pooled sample of 3095 subjects, followed by a candidate-driven approach which measured a significant enrichment of association in genes of the ´steroid hormone receptor activity´ and 'steroid metabolic process' pathways. Variants in the genes and pathways identified may affect the role of the planum temporale in language cognition.
  • Hagoort, P. (2015). Het talige brein. In A. Aleman, & H. E. Hulshoff Pol (Eds.), Beeldvorming van het brein: Imaging voor psychiaters en psychologen (pp. 169-176). Utrecht: De Tijdstroom.
  • Hagoort, P. (2015). Spiegelneuronen. In J. Brockmann (Ed.), Wetenschappelijk onkruid: 179 hardnekkige ideeën die vooruitgang blokkeren (pp. 455-457). Amsterdam: Maven Publishing.
  • Holler, J., Kokal, I., Toni, I., Hagoort, P., Kelly, S. D., & Ozyurek, A. (2015). Eye’m talking to you: Speakers’ gaze direction modulates co-speech gesture processing in the right MTG. Social Cognitive & Affective Neuroscience, 10, 255-261. doi:10.1093/scan/nsu047.

    Abstract

    Recipients process information from speech and co-speech gestures, but it is currently unknown how this processing is influenced by the presence of other important social cues, especially gaze direction, a marker of communicative intent. Such cues may modulate neural activity in regions associated either with the processing of ostensive cues, such as eye gaze, or with the processing of semantic information, provided by speech and gesture.
    Participants were scanned (fMRI) while taking part in triadic communication involving two recipients and a speaker. The speaker uttered sentences that
    were and were not accompanied by complementary iconic gestures. Crucially, the speaker alternated her gaze direction, thus creating two recipient roles: addressed (direct gaze) vs unaddressed (averted gaze) recipient. The comprehension of Speech&Gesture relative to SpeechOnly utterances recruited middle occipital, middle temporal and inferior frontal gyri, bilaterally. The calcarine sulcus and posterior cingulate cortex were sensitive to differences between direct and averted gaze. Most importantly, Speech&Gesture utterances, but not SpeechOnly utterances, produced additional activity in the right middle temporal gyrus when participants were addressed. Marking communicative intent with gaze direction modulates the processing of speech–gesture utterances in cerebral areas typically associated with the semantic processing of multi-modal communicative acts.
  • Horschig, J. M., Smolders, R., Bonnefond, M., Schoffelen, J.-M., Van den Munckhof, P., Schuurman, P. R., Cools, R., Denys, D., & Jensen, O. (2015). Directed communication between nucleus accumbens and neocortex in humans is differentially supported by synchronization in the theta and alpha band. PLoS One, 10(9): e0138685. doi:10.1371/journal.pone.0138685.

    Abstract

    Here, we report evidence for oscillatory bi-directional interactions between the nucleus accumbens and the neocortex in humans. Six patients performed a demanding covert visual attention task while we simultaneously recorded brain activity from deep-brain electrodes implanted in the nucleus accumbens and the surface electroencephalogram (EEG). Both theta and alpha oscillations were strongly coherent with the frontal and parietal EEG during the task. Theta-band coherence increased during processing of the visual stimuli. Granger causality analysis revealed that the nucleus accumbens was communicating with the neocortex primarily in the theta-band, while the cortex was communicating the nucleus accumbens in the alpha-band. These data are consistent with a model, in which theta- and alpha-band oscillations serve dissociable roles: Prior to stimulus processing, the cortex might suppress ongoing processing in the nucleus accumbens by modulating alpha-band activity. Subsequently, upon stimulus presentation, theta oscillations might facilitate the active exchange of stimulus information from the nucleus accumbens to the cortex.
  • Jiang, J., Chen, C., Dai, B., Shi, G., Liu, L., & Lu, C. (2015). Leader emergence through interpersonal neural synchronization. Proceedings of the National Academy of Sciences of the United States of America, 112(14), 4274-4279. doi:10.1073/pnas.1422930112.

    Abstract

    The neural mechanism of leader emergence is not well understood. This study investigated (i) whether interpersonal neural synchronization (INS) plays an important role in leader emergence, and (ii) whether INS and leader emergence are associated with the frequency or the quality of communications. Eleven three-member groups were asked to perform a leaderless group discussion (LGD) task, and their brain activities were recorded via functional near infrared spectroscopy (fNIRS)-based hyperscanning. Video recordings of the discussions were coded for leadership and communication. Results showed that the INS for the leader–follower (LF) pairs was higher than that for the follower–follower (FF) pairs in the left temporo-parietal junction (TPJ), an area important for social mentalizing. Although communication frequency was higher for the LF pairs than for the FF pairs, the frequency of leader-initiated and follower-initiated communication did not differ significantly. Moreover, INS for the LF pairs was significantly higher during leader-initiated communication than during follower-initiated communications. In addition, INS for the LF pairs during leader-initiated communication was significantly correlated with the leaders’ communication skills and competence, but not their communication frequency. Finally, leadership could be successfully predicted based on INS as well as communication frequency early during the LGD (before half a minute into the task). In sum, this study found that leader emergence was characterized by high-level neural synchronization between the leader and followers and that the quality, rather than the frequency, of communications was associated with synchronization. These results suggest that leaders emerge because they are able to say the right things at the right time.
  • Kunert, R., & Slevc, L. R. (2015). A commentary on: “Neural overlap in processing music and speech”. Frontiers in Human Neuroscience, 9: 330. doi:10.3389/fnhum.2015.00330.
  • Kunert, R., Willems, R. M., Casasanto, D., Patel, A. D., & Hagoort, P. (2015). Music and language syntax interact in Broca’s Area: An fMRI study. PLoS One, 10(11): e0141069. doi:10.1371/journal.pone.0141069.

    Abstract

    Instrumental music and language are both syntactic systems, employing complex, hierarchically-structured sequences built using implicit structural norms. This organization allows listeners to understand the role of individual words or tones in the context of an unfolding sentence or melody. Previous studies suggest that the brain mechanisms of syntactic processing may be partly shared between music and language. However, functional neuroimaging evidence for anatomical overlap of brain activity involved in linguistic and musical syntactic processing has been lacking. In the present study we used functional magnetic resonance imaging (fMRI) in conjunction with an interference paradigm based on sung sentences. We show that the processing demands of musical syntax (harmony) and language syntax interact in Broca’s area in the left inferior frontal gyrus (without leading to music and language main effects). A language main effect in Broca’s area only emerged in the complex music harmony condition, suggesting that (with our stimuli and tasks) a language effect only becomes visible under conditions of increased demands on shared neural resources. In contrast to previous studies, our design allows us to rule out that the observed neural interaction is due to: (1) general attention mechanisms, as a psychoacoustic auditory anomaly behaved unlike the harmonic manipulation, (2) error processing, as the language and the music stimuli contained no structural errors. The current results thus suggest that two different cognitive domains—music and language—might draw on the same high level syntactic integration resources in Broca’s area.
  • Lai, V. T., & Curran, T. (2015). Erratum to “ERP evidence for conceptual mappings and comparison processes during the comprehension of conventional and novel metaphors” [Brain Lang. 127 (3) (2013) 484–496]. Brain and Language, 149, 148-150. doi:10.1016/j.bandl.2014.11.001.
  • Lai, V. T., Willems, R. M., & Hagoort, P. (2015). Feel between the Lines: Implied emotion from combinatorial semantics. Journal of Cognitive Neuroscience, 27(8), 1528-1541. doi:10.1162/jocn_a_00798.

    Abstract

    This study investigated the brain regions for the comprehension of implied emotion in sentences. Participants read negative sentences without negative words, for example, “The boy fell asleep and never woke up again,” and their neutral counterparts “The boy stood up and grabbed his bag.” This kind of negative sentence allows us to examine implied emotion derived at the sentence level, without associative emotion coming from word retrieval. We found that implied emotion in sentences, relative to neutral sentences, led to activation in some emotion-related areas, including the medial prefrontal cortex, the amygdala, and the insula, as well as certain language-related areas, including the inferior frontal gyrus, which has been implicated in combinatorial processing. These results suggest that the emotional network involved in implied emotion is intricately related to the network for combinatorial processing in language, supporting the view that sentence meaning is more than simply concatenating the meanings of its lexical building blocks.
  • Lai, V. T., & Narasimhan, B. (2015). Verb representation and thinking-for-speaking effects in Spanish-English bilinguals. In R. G. De Almeida, & C. Manouilidou (Eds.), Cognitive science perspectives on verb representation and processing (pp. 235-256). Cham: Springer.

    Abstract

    Speakers of English habitually encode motion events using manner-of-motion verbs (e.g., spin, roll, slide) whereas Spanish speakers rely on path-of-motion verbs (e.g., enter, exit, approach). Here, we ask whether the language-specific verb representations used in encoding motion events induce different modes of “thinking-for-speaking” in Spanish–English bilinguals. That is, assuming that the verb encodes the most salient information in the clause, do bilinguals find the path of motion to be more salient than manner of motion if they had previously described the motion event using Spanish versus English? In our study, Spanish–English bilinguals described a set of target motion events in either English or Spanish and then participated in a nonlinguistic similarity judgment task in which they viewed the target motion events individually (e.g., a ball rolling into a cave) followed by two variants a “same-path” variant such as a ball sliding into a cave or a “same-manner” variant such as a ball rolling away from a cave). Participants had to select one of the two variants that they judged to be more similar to the target event: The event that shared the same path of motion as the target versus the one that shared the same manner of motion. Our findings show that bilingual speakers were more likely to classify two motion events as being similar if they shared the same path of motion and if they had previously described the target motion events in Spanish versus in English. Our study provides further evidence for the “thinking-for-speaking” hypothesis by demonstrating that bilingual speakers can flexibly shift between language-specific construals of the same event “on-the-fly.”
  • Lartseva, A., Dijkstra, T., & Buitelaar, J. (2015). Emotional language processing in Autism Spectrum Disorders: A systematic review. Frontiers in Human Neuroscience, 8: 991. doi:10.3389/fnhum.2014.00991.

    Abstract

    In his first description of Autism Spectrum Disorders (ASD), Kanner emphasized emotional impairments by characterizing children with ASD as indifferent to other people, self-absorbed, emotionally cold, distanced, and retracted. Thereafter, emotional impairments became regarded as part of the social impairments of ASD, and research mostly focused on understanding how individuals with ASD recognize visual expressions of emotions from faces and body postures. However, it still remains unclear how emotions are processed outside of the visual domain. This systematic review aims to fill this gap by focusing on impairments of emotional language processing in ASD.
    We systematically searched PubMed for papers published between 1990 and 2013 using standardized search terms. Studies show that people with ASD are able to correctly classify emotional language stimuli as emotionally positive or negative. However, processing of emotional language stimuli in ASD is associated with atypical patterns of attention and memory performance, as well as abnormal physiological and neural activity. Particularly, younger children with ASD have difficulties in acquiring and developing emotional concepts, and avoid using these in discourse. These emotional language impairments were not consistently associated with age, IQ, or level of development of language skills.
    We discuss how emotional language impairments fit with existing cognitive theories of ASD, such as central coherence, executive dysfunction, and weak Theory of Mind. We conclude that emotional impairments in ASD may be broader than just a mere consequence of social impairments, and should receive more attention in future research

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