Publications

Displaying 1 - 100 of 112
  • Bakker-Marshall, I., Takashima, A., Schoffelen, J.-M., Van Hell, J. G., Janzen, G., & McQueen, J. M. (2018). Theta-band Oscillations in the Middle Temporal Gyrus Reflect Novel Word Consolidation. Journal of Cognitive Neuroscience, 30(5), 621-633. doi:10.1162/jocn_a_01240.

    Abstract

    Like many other types of memory formation, novel word learning benefits from an offline consolidation period after the initial encoding phase. A previous EEG study has shown that retrieval of novel words elicited more word-like-induced electrophysiological brain activity in the theta band after consolidation [Bakker, I., Takashima, A., van Hell, J. G., Janzen, G., & McQueen, J. M. Changes in theta and beta oscillations as signatures of novel word consolidation. Journal of Cognitive Neuroscience, 27, 1286–1297, 2015]. This suggests that theta-band oscillations play a role in lexicalization, but it has not been demonstrated that this effect is directly caused by the formation of lexical representations. This study used magnetoencephalography to localize the theta consolidation effect to the left posterior middle temporal gyrus (pMTG), a region known to be involved in lexical storage. Both untrained novel words and words learned immediately before test elicited lower theta power during retrieval than existing words in this region. After a 24-hr consolidation period, the difference between novel and existing words decreased significantly, most strongly in the left pMTG. The magnitude of the decrease after consolidation correlated with an increase in behavioral competition effects between novel words and existing words with similar spelling, reflecting functional integration into the mental lexicon. These results thus provide new evidence that consolidation aids the development of lexical representations mediated by the left pMTG. Theta synchronization may enable lexical access by facilitating the simultaneous activation of distributed semantic, phonological, and orthographic representations that are bound together in the pMTG.
  • Berkers, R. M. W. J., Ekman, M., van Dongen, E. V., Takashima, A., Barth, M., Paller, K. A., & Fernández, G. (2018). Cued reactivation during slow-wave sleep induces brain connectivity changes related to memory stabilization. Scientific Reports, 8: 16958. doi:10.1038/s41598-018-35287-6.

    Abstract

    Memory reprocessing following acquisition enhances memory consolidation. Specifically, neural activity during encoding is thought to be ‘replayed’ during subsequent slow-wave sleep. Such memory replay is thought to contribute to the functional reorganization of neural memory traces. In particular, memory replay may facilitate the exchange of information across brain regions by inducing a reconfiguration of connectivity across the brain. Memory reactivation can be induced by external cues through a procedure known as “targeted memory reactivation”. Here, we analysed data from a published study with auditory cues used to reactivate visual object-location memories during slow-wave sleep. We characterized effects of memory reactivation on brain network connectivity using graph-theory. We found that cue presentation during slow-wave sleep increased global network integration of occipital cortex, a visual region that was also active during retrieval of object locations. Although cueing did not have an overall beneficial effect on the retention of cued versus uncued associations, individual differences in overnight memory stabilization were related to enhanced network integration of occipital cortex. Furthermore, occipital cortex displayed enhanced connectivity with mnemonic regions, namely the hippocampus, parahippocampal gyrus, thalamus and medial prefrontal cortex during cue sound presentation. Together, these results suggest a neural mechanism where cue-induced replay during sleep increases integration of task-relevant perceptual regions with mnemonic regions. This cross-regional integration may be instrumental for the consolidation and long-term storage of enduring memories.

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    41598_2018_35287_MOESM1_ESM.doc
  • Dai, B., Chen, C., Long, Y., Zheng, L., Zhao, H., Bai, X., Liu, W., Zhang, Y., Liu, L., Guo, T., Ding, G., & Lu, C. (2018). Neural mechanisms for selectively tuning into the target speaker in a naturalistic noisy situation. Nature Communications, 9: 2405. doi:10.1038/s41467-018-04819-z.

    Abstract

    The neural mechanism for selectively tuning in to a target speaker while tuning out the others in a multi-speaker situation (i.e., the cocktail-party effect) remains elusive. Here we addressed this issue by measuring brain activity simultaneously from a listener and from multiple speakers while they were involved in naturalistic conversations. Results consistently show selectively enhanced interpersonal neural synchronization (INS) between the listener and the attended speaker at left temporal–parietal junction, compared with that between the listener and the unattended speaker across different multi-speaker situations. Moreover, INS increases significantly prior to the occurrence of verbal responses, and even when the listener’s brain activity precedes that of the speaker. The INS increase is independent of brain-to-speech synchronization in both the anatomical location and frequency range. These findings suggest that INS underlies the selective process in a multi-speaker situation through neural predictions at the content level but not the sensory level of speech.

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    Dai_etal_2018_sup.pdf
  • Degand, L., & Van Bergen, G. (2018). Discourse markers as turn-transition devices: Evidence from speech and instant messaging. Discourse Processes, 55, 47-71. doi:10.1080/0163853X.2016.1198136.

    Abstract

    In this article we investigate the relation between discourse markers and turn-transition strategies in face-to-face conversations and Instant Messaging (IM), that is, unplanned, real-time, text-based, computer-mediated communication. By means of a quantitative corpus study of utterances containing a discourse marker, we show that utterance-final discourse markers are used more often in IM than in face-to-face conversations. Moreover, utterance-final discourse markers are shown to occur more often at points of turn-transition compared with points of turn-maintenance in both types of conversation. From our results we conclude that the discourse markers in utterance-final position can function as a turn-transition mechanism, signaling that the turn is over and the floor is open to the hearer. We argue that this linguistic turn-taking strategy is essentially similar in face-to-face and IM communication. Our results add to the evidence that communication in IM is more like speech than like writing.
  • Duarte, R., Uhlmann, M., Van den Broek, D., Fitz, H., Petersson, K. M., & Morrison, A. (2018). Encoding symbolic sequences with spiking neural reservoirs. In Proceedings of the 2018 International Joint Conference on Neural Networks (IJCNN). doi:10.1109/IJCNN.2018.8489114.

    Abstract

    Biologically inspired spiking networks are an important tool to study the nature of computation and cognition in neural systems. In this work, we investigate the representational capacity of spiking networks engaged in an identity mapping task. We compare two schemes for encoding symbolic input, one in which input is injected as a direct current and one where input is delivered as a spatio-temporal spike pattern. We test the ability of networks to discriminate their input as a function of the number of distinct input symbols. We also compare performance using either membrane potentials or filtered spike trains as state variable. Furthermore, we investigate how the circuit behavior depends on the balance between excitation and inhibition, and the degree of synchrony and regularity in its internal dynamics. Finally, we compare different linear methods of decoding population activity onto desired target labels. Overall, our results suggest that even this simple mapping task is strongly influenced by design choices on input encoding, state-variables, circuit characteristics and decoding methods, and these factors can interact in complex ways. This work highlights the importance of constraining computational network models of behavior by available neurobiological evidence.
  • Eekhof, L. S., Eerland, A., & Willems, R. M. (2018). Readers’ insensitivity to tense revealed: No differences in mental simulation during reading of present and past tense stories. Collabra: Psychology, 4(1): 16. doi:10.1525/collabra.121.

    Abstract

    While the importance of mental simulation during literary reading has long been recognized, we know little about the factors that determine when, what, and how much readers mentally simulate. Here we investigate the influence of a specific text characteristic, namely verb tense (present vs. past), on mental simulation during literary reading. Verbs usually denote the actions and events that take place in narratives and hence it is hypothesized that verb tense will influence the amount of mental simulation elicited in readers. Although the present tense is traditionally considered to be more “vivid”, this study is one of the first to experimentally assess this claim. We recorded eye-movements while subjects read stories in the past or present tense and collected data regarding self-reported levels of mental simulation, transportation and appreciation. We found no influence of tense on any of the offline measures. The eye-tracking data showed a slightly more complex pattern. Although we did not find a main effect of sensorimotor simulation content on reading times, we were able to link the degree to which subjects slowed down when reading simulation eliciting content to offline measures of attention and transportation, but this effect did not interact with the tense of the story. Unexpectedly, we found a main effect of tense on reading times per word, with past tense stories eliciting longer first fixation durations and gaze durations. However, we were unable to link this effect to any of the offline measures. In sum, this study suggests that tense does not play a substantial role in the process of mental simulation elicited by literary stories.

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  • Eichert, N., Peeters, D., & Hagoort, P. (2018). Language-driven anticipatory eye movements in virtual reality. Behavior Research Methods, 50(3), 1102-1115. doi:10.3758/s13428-017-0929-z.

    Abstract

    Predictive language processing is often studied by measuring eye movements as participants look at objects on a computer screen while they listen to spoken sentences. The use of this variant of the visual world paradigm has shown that information encountered by a listener at a spoken verb can give rise to anticipatory eye movements to a target object, which is taken to indicate that people predict upcoming words. The ecological validity of such findings remains questionable, however, because these computer experiments used two-dimensional (2D) stimuli that are mere abstractions of real world objects. Here we present a visual world paradigm study in a three-dimensional (3D) immersive virtual reality environment. Despite significant changes in the stimulus material and the different mode of stimulus presentation, language-mediated anticipatory eye movements were observed. These findings thus indicate prediction of upcoming words in language comprehension in a more naturalistic setting where natural depth cues are preserved. Moreover, the results confirm the feasibility of using eye-tracking in rich and multimodal 3D virtual environments.

    Additional information

    13428_2017_929_MOESM1_ESM.docx
  • Ergin, R., Senghas, A., Jackendoff, R., & Gleitman, L. (2018). Structural cues for symmetry, asymmetry, and non-symmetry in Central Taurus Sign Language. In C. Cuskley, M. Flaherty, H. Little, L. McCrohon, A. Ravignani, & T. Verhoef (Eds.), Proceedings of the 12th International Conference on the Evolution of Language (EVOLANG XII) (pp. 104-106). Toruń, Poland: NCU Press. doi:10.12775/3991-1.025.
  • Ergin, R., Meir, I., Ilkbasaran, D., Padden, C., & Jackendoff, R. (2018). The Development of Argument Structure in Central Taurus Sign Language. Sign Language & Linguistics, 18(4), 612-639. doi:10.1353/sls.2018.0018.

    Abstract

    One of the fundamental issues for a language is its capacity to express argument structure unambiguously. This study presents evidence for the emergence and the incremental development of these basic mechanisms in a newly developing language, Central Taurus Sign Language. Our analyses identify universal patterns in both the emergence and development of these mechanisms and in languagespecific trajectories.
  • Flecken, M., & Von Stutterheim, C. (2018). Sprache und Kognition: Sprachvergleichende und lernersprachliche Untersuchungen zur Ereigniskonzeptualisierung. In S. Schimke, & H. Hopp (Eds.), Sprachverarbeitung im Zweitspracherwerb (pp. 325-356). Berlin: De Gruyter. doi:10.1515/9783110456356-014.
  • Francisco, A. A., Takashima, A., McQueen, J. M., Van den Bunt, M., Jesse, A., & Groen, M. A. (2018). Adult dyslexic readers benefit less from visual input during audiovisual speech processing: fMRI evidence. Neuropsychologia, 117, 454-471. doi:10.1016/j.neuropsychologia.2018.07.009.

    Abstract

    The aim of the present fMRI study was to investigate whether typical and dyslexic adult readers differed in the neural correlates of audiovisual speech processing. We tested for Blood Oxygen-Level Dependent (BOLD) activity differences between these two groups in a 1-back task, as they processed written (word, illegal consonant strings) and spoken (auditory, visual and audiovisual) stimuli. When processing written stimuli, dyslexic readers showed reduced activity in the supramarginal gyrus, a region suggested to play an important role in phonological processing, but only when they processed strings of consonants, not when they read words. During the speech perception tasks, dyslexic readers were only slower than typical readers in their behavioral responses in the visual speech condition. Additionally, dyslexic readers presented reduced neural activation in the auditory, the visual, and the audiovisual speech conditions. The groups also differed in terms of superadditivity, with dyslexic readers showing decreased neural activation in the regions of interest. An additional analysis focusing on vision-related processing during the audiovisual condition showed diminished activation for the dyslexic readers in a fusiform gyrus cluster. Our results thus suggest that there are differences in audiovisual speech processing between dyslexic and normal readers. These differences might be explained by difficulties in processing the unisensory components of audiovisual speech, more specifically, dyslexic readers may benefit less from visual information during audiovisual speech processing than typical readers. Given that visual speech processing supports the development of phonological skills fundamental in reading, differences in processing of visual speech could contribute to differences in reading ability between typical and dyslexic readers.
  • Franken, M. K. (2018). Listening for speaking: Investigations of the relationship between speech perception and production. PhD Thesis, Radboud University, Nijmegen.

    Abstract

    Speaking and listening are complex tasks that we perform on a daily basis, almost without conscious effort. Interestingly, speaking almost never occurs without listening: whenever we speak, we at least hear our own speech. The research in this thesis is concerned with how the perception of our own speech influences our speaking behavior. We show that unconsciously, we actively monitor this auditory feedback of our own speech. This way, we can efficiently take action and adapt articulation when an error occurs and auditory feedback does not correspond to our expectation. Processing the auditory feedback of our speech does not, however, automatically affect speech production. It is subject to a number of constraints. For example, we do not just track auditory feedback, but also its consistency. If auditory feedback is more consistent over time, it has a stronger influence on speech production. In addition, we investigated how auditory feedback during speech is processed in the brain, using magnetoencephalography (MEG). The results suggest the involvement of a broad cortical network including both auditory and motor-related regions. This is consistent with the view that the auditory center of the brain is involved in comparing auditory feedback to our expectation of auditory feedback. If this comparison yields a mismatch, motor-related regions of the brain can be recruited to alter the ongoing articulations.

    Additional information

    full text via Radboud Repository
  • Franken, M. K., Eisner, F., Acheson, D. J., McQueen, J. M., Hagoort, P., & Schoffelen, J.-M. (2018). Self-monitoring in the cerebral cortex: Neural responses to pitch-perturbed auditory feedback during speech production. NeuroImage, 179, 326-336. doi:10.1016/j.neuroimage.2018.06.061.

    Abstract

    Speaking is a complex motor skill which requires near instantaneous integration of sensory and motor-related information. Current theory hypothesizes a complex interplay between motor and auditory processes during speech production, involving the online comparison of the speech output with an internally generated forward model. To examine the neural correlates of this intricate interplay between sensory and motor processes, the current study uses altered auditory feedback (AAF) in combination with magnetoencephalography (MEG). Participants vocalized the vowel/e/and heard auditory feedback that was temporarily pitch-shifted by only 25 cents, while neural activity was recorded with MEG. As a control condition, participants also heard the recordings of the same auditory feedback that they heard in the first half of the experiment, now without vocalizing. The participants were not aware of any perturbation of the auditory feedback. We found auditory cortical areas responded more strongly to the pitch shifts during vocalization. In addition, auditory feedback perturbation resulted in spectral power increases in the θ and lower β bands, predominantly in sensorimotor areas. These results are in line with current models of speech production, suggesting auditory cortical areas are involved in an active comparison between a forward model's prediction and the actual sensory input. Subsequently, these areas interact with motor areas to generate a motor response. Furthermore, the results suggest that θ and β power increases support auditory-motor interaction, motor error detection and/or sensory prediction processing.
  • Franken, M. K., Acheson, D. J., McQueen, J. M., Hagoort, P., & Eisner, F. (2018). Opposing and following responses in sensorimotor speech control: Why responses go both ways. Psychonomic Bulletin & Review, 25(4), 1458-1467. doi:10.3758/s13423-018-1494-x.

    Abstract

    When talking, speakers continuously monitor and use the auditory feedback of their own voice to control and inform speech production processes. When speakers are provided with auditory feedback that is perturbed in real time, most of them compensate for this by opposing the feedback perturbation. But some speakers follow the perturbation. In the current study, we investigated whether the state of the speech production system at perturbation onset may determine what type of response (opposing or following) is given. The results suggest that whether a perturbation-related response is opposing or following depends on ongoing fluctuations of the production system: It initially responds by doing the opposite of what it was doing. This effect and the non-trivial proportion of following responses suggest that current production models are inadequate: They need to account for why responses to unexpected sensory feedback depend on the production-system’s state at the time of perturbation.
  • De Groot, A. M. B., & Hagoort, P. (Eds.). (2018). Research methods in psycholinguistics and the neurobiology of language: A practical guide. Oxford: Wiley.
  • Hagoort, P. (2018). Prerequisites for an evolutionary stance on the neurobiology of language. Current Opinion in Behavioral Sciences, 21, 191-194. doi:10.1016/j.cobeha.2018.05.012.
  • Hahn, L. E., Benders, T., Snijders, T. M., & Fikkert, P. (2018). Infants' sensitivity to rhyme in songs. Infant Behavior and Development, 52, 130-139. doi:10.1016/j.infbeh.2018.07.002.

    Abstract

    Children’s songs often contain rhyming words at phrase endings. In this study, we investigated whether infants can already recognize this phonological pattern in songs. Earlier studies using lists of spoken words were equivocal on infants’ spontaneous processing of rhymes (Hayes, Slater, & Brown, 2000; Jusczyk, Goodman, & Baumann, 1999). Songs, however, constitute an ecologically valid rhyming stimulus, which could allow for spontaneous processing of this phonological pattern in infants. Novel children’s songs with rhyming and non-rhyming lyrics using pseudo-words were presented to 35 9-month-old Dutch infants using the Headturn Preference Procedure. Infants on average listened longer to the non-rhyming songs, with around half of the infants however exhibiting a preference for the rhyming songs. These results highlight that infants have the processing abilities to benefit from their natural rhyming input for the development of their phonological abilities.
  • Hasson, U., Egidi, G., Marelli, M., & Willems, R. M. (2018). Grounding the neurobiology of language in first principles: The necessity of non-language-centric explanations for language comprehension. Cognition, 180(1), 135-157. doi:10.1016/j.cognition.2018.06.018.

    Abstract

    Recent decades have ushered in tremendous progress in understanding the neural basis of language. Most of our current knowledge on language and the brain, however, is derived from lab-based experiments that are far removed from everyday language use, and that are inspired by questions originating in linguistic and psycholinguistic contexts. In this paper we argue that in order to make progress, the field needs to shift its focus to understanding the neurobiology of naturalistic language comprehension. We present here a new conceptual framework for understanding the neurobiological organization of language comprehension. This framework is non-language-centered in the computational/neurobiological constructs it identifies, and focuses strongly on context. Our core arguments address three general issues: (i) the difficulty in extending language-centric explanations to discourse; (ii) the necessity of taking context as a serious topic of study, modeling it formally and acknowledging the limitations on external validity when studying language comprehension outside context; and (iii) the tenuous status of the language network as an explanatory construct. We argue that adopting this framework means that neurobiological studies of language will be less focused on identifying correlations between brain activity patterns and mechanisms postulated by psycholinguistic theories. Instead, they will be less self-referential and increasingly more inclined towards integration of language with other cognitive systems, ultimately doing more justice to the neurobiological organization of language and how it supports language as it is used in everyday life.
  • Hervais-Adelman, A., Egorova, N., & Golestani, N. (2018). Beyond bilingualism: Multilingual experience correlates with caudate volume. Brain Structure and Function, 223(7), 3495-3502. doi:10.1007/s00429-018-1695-0.

    Abstract

    The multilingual brain implements mechanisms that serve to select the appropriate language as a function of the communicative environment. Engaging these mechanisms on a regular basis appears to have consequences for brain structure and function. Studies have implicated the caudate nuclei as important nodes in polyglot language control processes, and have also shown structural differences in the caudate nuclei in bilingual compared to monolingual populations. However, the majority of published work has focused on the categorical differences between monolingual and bilingual individuals, and little is known about whether these findings extend to multilingual individuals, who have even greater language control demands. In the present paper, we present an analysis of the volume and morphology of the caudate nuclei, putamen, pallidum and thalami in 75 multilingual individuals who speak three or more languages. Volumetric analyses revealed a significant relationship between multilingual experience and right caudate volume, as well as a marginally significant relationship with left caudate volume. Vertex-wise analyses revealed a significant enlargement of dorsal and anterior portions of the left caudate nucleus, known to have connectivity with executive brain regions, as a function of multilingual expertise. These results suggest that multilingual expertise might exercise a continuous impact on brain structure, and that as additional languages beyond a second are acquired, the additional demands for linguistic and cognitive control result in modifications to brain structures associated with language management processes.
  • Hervais-Adelman, A., Moser-Mercer, B., & Golestani, N. (2018). Commentary: Broca pars triangularis constitutes a “hub” of the language-control network during simultaneous language translation. Frontiers in Human Neuroscience, 12: 22. doi:10.3389/fnhum.2018.00022.

    Abstract

    A commentary on Broca Pars Triangularis Constitutes a “Hub” of the Language-Control Network during Simultaneous Language Translation by Elmer, S. (2016). Front. Hum. Neurosci. 10:491. doi: 10.3389/fnhum.2016.00491 Elmer (2016) conducted an fMRI investigation of “simultaneous language translation” in five participants. The article presents group and individual analyses of German-to-Italian and Italian-to-German translation, confined to a small set of anatomical regions previously reported to be involved in multilingual control. Here we take the opportunity to discuss concerns regarding certain aspects of the study.
  • Heyselaar, E., Mazaheri, A., Hagoort, P., & Segaert, K. (2018). Changes in alpha activity reveal that social opinion modulates attention allocation during face processing. NeuroImage, 174, 432-440. doi:10.1016/j.neuroimage.2018.03.034.

    Abstract

    Participants’ performance differs when conducting a task in the presence of a secondary individual, moreover the opinion the participant has of this individual also plays a role. Using EEG, we investigated how previous interactions with, and evaluations of, an avatar in virtual reality subsequently influenced attentional allocation to the face of that avatar. We focused on changes in the alpha activity as an index of attentional allocation. We found that the onset of an avatar’s face whom the participant had developed a rapport with induced greater alpha suppression. This suggests greater attentional resources are allocated to the interacted-with avatars. The evaluative ratings of the avatar induced a U-shaped change in alpha suppression, such that participants paid most attention when the avatar was rated as average. These results suggest that attentional allocation is an important element of how behaviour is altered in the presence of a secondary individual and is modulated by our opinion of that individual.

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  • Huettig, F., Lachmann, T., Reis, A., & Petersson, K. M. (2018). Distinguishing cause from effect - Many deficits associated with developmental dyslexia may be a consequence of reduced and suboptimal reading experience. Language, Cognition and Neuroscience, 33(3), 333-350. doi:10.1080/23273798.2017.1348528.

    Abstract

    The cause of developmental dyslexia is still unknown despite decades of intense research. Many causal explanations have been proposed, based on the range of impairments displayed by affected individuals. Here we draw attention to the fact that many of these impairments are also shown by illiterate individuals who have not received any or very little reading instruction. We suggest that this fact may not be coincidental and that the performance differences of both illiterates and individuals with dyslexia compared to literate controls are, to a substantial extent, secondary consequences of either reduced or suboptimal reading experience or a combination of both. The search for the primary causes of reading impairments will make progress if the consequences of quantitative and qualitative differences in reading experience are better taken into account and not mistaken for the causes of reading disorders. We close by providing four recommendations for future research.
  • Inacio, F., Faisca, L., Forkstam, C., Araujo, S., Bramao, I., Reis, A., & Petersson, K. M. (2018). Implicit sequence learning is preserved in dyslexic children. Annals of Dyslexia, 68(1), 1-14. doi:10.1007/s11881-018-0158-x.

    Abstract

    This study investigates the implicit sequence learning abilities of dyslexic children using an artificial grammar learning task with an extended exposure period. Twenty children with developmental dyslexia participated in the study and were matched with two control groups—one matched for age and other for reading skills. During 3 days, all participants performed an acquisition task, where they were exposed to colored geometrical forms sequences with an underlying grammatical structure. On the last day, after the acquisition task, participants were tested in a grammaticality classification task. Implicit sequence learning was present in dyslexic children, as well as in both control groups, and no differences between groups were observed. These results suggest that implicit learning deficits per se cannot explain the characteristic reading difficulties of the dyslexics.
  • Jacobs, A. M., & Willems, R. M. (2018). The fictive brain: Neurocognitive correlates of engagement in literature. Review of General Psychology, 22(2), 147-160. doi:10.1037/gpr0000106.

    Abstract

    Fiction is vital to our being. Many people enjoy engaging with fiction every day. Here we focus on literary reading as 1 instance of fiction consumption from a cognitive neuroscience perspective. The brain processes which play a role in the mental construction of fiction worlds and the related engagement with fictional characters, remain largely unknown. The authors discuss the neurocognitive poetics model (Jacobs, 2015a) of literary reading specifying the likely neuronal correlates of several key processes in literary reading, namely inference and situation model building, immersion, mental simulation and imagery, figurative language and style, and the issue of distinguishing fact from fiction. An overview of recent work on these key processes is followed by a discussion of methodological challenges in studying the brain bases of fiction processing
  • Kösem, A., Bosker, H. R., Takashima, A., Meyer, A. S., Jensen, O., & Hagoort, P. (2018). Neural entrainment determines the words we hear. Current Biology, 28, 2867-2875. doi:10.1016/j.cub.2018.07.023.

    Abstract

    Low-frequency neural entrainment to rhythmic input has been hypothesized as a canonical mechanism that shapes sensory perception in time. Neural entrainment is deemed particularly relevant for speech analysis, as it would contribute to the extraction of discrete linguistic elements from continuous acoustic signals. However, its causal influence in speech perception has been difficult to establish. Here, we provide evidence that oscillations build temporal predictions about the duration of speech tokens that affect perception. Using magnetoencephalography (MEG), we studied neural dynamics during listening to sentences that changed in speech rate. Weobserved neural entrainment to preceding speech rhythms persisting for several cycles after the change in rate. The sustained entrainment was associated with changes in the perceived duration of the last word’s vowel, resulting in the perception of words with different meanings. These findings support oscillatory models of speech processing, suggesting that neural oscillations actively shape speech perception.
  • Lam, N. H. L., Hulten, A., Hagoort, P., & Schoffelen, J.-M. (2018). Robust neuronal oscillatory entrainment to speech displays individual variation in lateralisation. Language, Cognition and Neuroscience, 33(8), 943-954. doi:10.1080/23273798.2018.1437456.

    Abstract

    Neural oscillations may be instrumental for the tracking and segmentation of continuous speech. Earlier work has suggested that delta, theta and gamma oscillations entrain to the speech rhythm. We used magnetoencephalography and a large sample of 102 participants to investigate oscillatory entrainment to speech, and observed robust entrainment of delta and theta activity, and weak group-level gamma entrainment. We show that the peak frequency and the hemispheric lateralisation of the entrainment are subject to considerable individual variability. The first finding may support the involvement of intrinsic oscillations in entrainment, and the second finding suggests that there is no systematic default right-hemispheric bias for processing acoustic signals on a slow time scale. Although low frequency entrainment to speech is a robust phenomenon, the characteristics of entrainment vary across individuals, and this variation is important for understanding the underlying neural mechanisms of entrainment, as well as its functional significance.
  • Lewis, A. G., Schriefers, H., Bastiaansen, M., & Schoffelen, J.-M. (2018). Assessing the utility of frequency tagging for tracking memory-based reactivation of word representations. Scientific Reports, 8: 7897. doi:10.1038/s41598-018-26091-3.

    Abstract

    Reinstatement of memory-related neural activity measured with high temporal precision potentially provides a useful index for real-time monitoring of the timing of activation of memory content during cognitive processing. The utility of such an index extends to any situation where one is interested in the (relative) timing of activation of different sources of information in memory, a paradigm case of which is tracking lexical activation during language processing. Essential for this approach is that memory reinstatement effects are robust, so that their absence (in the average) definitively indicates that no lexical activation is present. We used electroencephalography to test the robustness of a reported subsequent memory finding involving reinstatement of frequency-specific entrained oscillatory brain activity during subsequent recognition. Participants learned lists of words presented on a background flickering at either 6 or 15 Hz to entrain a steady-state brain response. Target words subsequently presented on a non-flickering background that were correctly identified as previously seen exhibited reinstatement effects at both entrainment frequencies. Reliability of these statistical inferences was however critically dependent on the approach used for multiple comparisons correction. We conclude that effects are not robust enough to be used as a reliable index of lexical activation during language processing.

    Additional information

    Lewis_etal_2018sup.docx
  • Lopopolo, A., Frank, S. L., Van den Bosch, A., Nijhof, A., & Willems, R. M. (2018). The Narrative Brain Dataset (NBD), an fMRI dataset for the study of natural language processing in the brain. In B. Devereux, E. Shutova, & C.-R. Huang (Eds.), Proceedings of LREC 2018 Workshop "Linguistic and Neuro-Cognitive Resources (LiNCR) (pp. 8-11). Paris: LREC.

    Abstract

    We present the Narrative Brain Dataset, an fMRI dataset that was collected during spoken presentation of short excerpts of three stories in Dutch. Together with the brain imaging data, the dataset contains the written versions of the stimulation texts. The texts are accompanied with stochastic (perplexity and entropy) and semantic computational linguistic measures. The richness and unconstrained nature of the data allows the study of language processing in the brain in a more naturalistic setting than is common for fMRI studies. We hope that by making NBD available we serve the double purpose of providing useful neural data to researchers interested in natural language processing in the brain and to further stimulate data sharing in the field of neuroscience of language.
  • Manahova, M. E., Mostert, P., Kok, P., Schoffelen, J.-M., & De Lange, F. P. (2018). Stimulus familiarity and expectation jointly modulate neural activity in the visual ventral stream. Journal of Cognitive Neuroscience, 30(9), 1366-1377. doi:10.1162/jocn_a_01281.

    Abstract

    Prior knowledge about the visual world can change how a visual stimulus is processed. Two forms of prior knowledge are often distinguished: stimulus familiarity (i.e., whether a stimulus has been seen before) and stimulus expectation (i.e., whether a stimulus is expected to occur, based on the context). Neurophysiological studies in monkeys have shown suppression of spiking activity both for expected and for familiar items in object-selective inferotemporal cortex. It is an open question, however, if and how these types of knowledge interact in their modulatory effects on the sensory response. To address this issue and to examine whether previous findings generalize to noninvasively measured neural activity in humans, we separately manipulated stimulus familiarity and expectation while noninvasively recording human brain activity using magnetoencephalography. We observed independent suppression of neural activity by familiarity and expectation, specifically in the lateral occipital complex, the putative human homologue of monkey inferotemporal cortex. Familiarity also led to sharpened response dynamics, which was predominantly observed in early visual cortex. Together, these results show that distinct types of sensory knowledge jointly determine the amount of neural resources dedicated to object processing in the visual ventral stream.
  • Meyer, A. S., Alday, P. M., Decuyper, C., & Knudsen, B. (2018). Working together: Contributions of corpus analyses and experimental psycholinguistics to understanding conversation. Frontiers in Psychology, 9: 525. doi:10.3389/fpsyg.2018.00525.

    Abstract

    As conversation is the most important way of using language, linguists and psychologists should combine forces to investigate how interlocutors deal with the cognitive demands arising during conversation. Linguistic analyses of corpora of conversation are needed to understand the structure of conversations, and experimental work is indispensable for understanding the underlying cognitive processes. We argue that joint consideration of corpus and experimental data is most informative when the utterances elicited in a lab experiment match those extracted from a corpus in relevant ways. This requirement to compare like with like seems obvious but is not trivial to achieve. To illustrate this approach, we report two experiments where responses to polar (yes/no) questions were elicited in the lab and the response latencies were compared to gaps between polar questions and answers in a corpus of conversational speech. We found, as expected, that responses were given faster when they were easy to plan and planning could be initiated earlier than when they were harder to plan and planning was initiated later. Overall, in all but one condition, the latencies were longer than one would expect based on the analyses of corpus data. We discuss the implication of this partial match between the data sets and more generally how corpus and experimental data can best be combined in studies of conversation.

    Additional information

    Data_Sheet_1.pdf
  • Nieuwland, M. S., Politzer-Ahles, S., Heyselaar, E., Segaert, K., Darley, E., Kazanina, N., Von Grebmer Zu Wolfsthurn, S., Bartolozzi, F., Kogan, V., Ito, A., Mézière, D., Barr, D. J., Rousselet, G., Ferguson, H. J., Busch-Moreno, S., Fu, X., Tuomainen, J., Kulakova, E., Husband, E. M., Donaldson, D. I. and 3 moreNieuwland, M. S., Politzer-Ahles, S., Heyselaar, E., Segaert, K., Darley, E., Kazanina, N., Von Grebmer Zu Wolfsthurn, S., Bartolozzi, F., Kogan, V., Ito, A., Mézière, D., Barr, D. J., Rousselet, G., Ferguson, H. J., Busch-Moreno, S., Fu, X., Tuomainen, J., Kulakova, E., Husband, E. M., Donaldson, D. I., Kohút, Z., Rueschemeyer, S.-A., & Huettig, F. (2018). Large-scale replication study reveals a limit on probabilistic prediction in language comprehension. eLife, 7: e33468. doi:10.7554/eLife.33468.

    Abstract

    Do people routinely pre-activate the meaning and even the phonological form of upcoming words? The most acclaimed evidence for phonological prediction comes from a 2005 Nature Neuroscience publication by DeLong, Urbach and Kutas, who observed a graded modulation of electrical brain potentials (N400) to nouns and preceding articles by the probability that people use a word to continue the sentence fragment (‘cloze’). In our direct replication study spanning 9 laboratories (N=334), pre-registered replication-analyses and exploratory Bayes factor analyses successfully replicated the noun-results but, crucially, not the article-results. Pre-registered single-trial analyses also yielded a statistically significant effect for the nouns but not the articles. Exploratory Bayesian single-trial analyses showed that the article-effect may be non-zero but is likely far smaller than originally reported and too small to observe without very large sample sizes. Our results do not support the view that readers routinely pre-activate the phonological form of predictable words.

    Additional information

    Data sets
  • Niso, G., Gorgolewski, K. J., Bock, E., Brooks, T. L., Flandin, G., Gramfort, A., Henson, R. N., Jas, M., Litvak, V., Moreau, J. T., Oostenveld, R., Schoffelen, J.-M., Tadel, F., Wexler, J., & Baillet, S. (2018). MEG-BIDS, the brain imaging data structure extended to magnetoencephalography. Scientific Data, 5: 180110. doi:10.1038/sdata.2018.110.

    Abstract

    We present a significant extension of the Brain Imaging Data Structure (BIDS) to support the specific aspects of magnetoencephalography (MEG) data. MEG measures brain activity with millisecond temporal resolution and unique source imaging capabilities. So far, BIDS was a solution to organise magnetic resonance imaging (MRI) data. The nature and acquisition parameters of MRI and MEG data are strongly dissimilar. Although there is no standard data format for MEG, we propose MEG-BIDS as a principled solution to store, organise, process and share the multidimensional data volumes produced by the modality. The standard also includes well-defined metadata, to facilitate future data harmonisation and sharing efforts. This responds to unmet needs from the multimodal neuroimaging community and paves the way to further integration of other techniques in electrophysiology. MEGBIDS builds on MRI-BIDS, extending BIDS to a multimodal data structure. We feature several dataanalytics software that have adopted MEG-BIDS, and a diverse sample of open MEG-BIDS data resources available to everyone.
  • Palva, J. M., Wang, S. H., Palva, S., Zhigalov, A., Monto, S., Brookes, M. J., & Schoffelen, J.-M. (2018). Ghost interactions in MEG/EEG source space: A note of caution on inter-areal coupling measures. NeuroImage, 173, 632-643. doi:10.1016/j.neuroimage.2018.02.032.

    Abstract

    When combined with source modeling, magneto- (MEG) and electroencephalography (EEG) can be used to study long-range interactions among cortical processes non-invasively. Estimation of such inter-areal connectivity is nevertheless hindered by instantaneous field spread and volume conduction, which artificially introduce linear correlations and impair source separability in cortical current estimates. To overcome the inflating effects of linear source mixing inherent to standard interaction measures, alternative phase- and amplitude-correlation based connectivity measures, such as imaginary coherence and orthogonalized amplitude correlation have been proposed. Being by definition insensitive to zero-lag correlations, these techniques have become increasingly popular in the identification of correlations that cannot be attributed to field spread or volume conduction. We show here, however, that while these measures are immune to the direct effects of linear mixing, they may still reveal large numbers of spurious false positive connections through field spread in the vicinity of true interactions. This fundamental problem affects both region-of-interest-based analyses and all-to-all connectome mappings. Most importantly, beyond defining and illustrating the problem of spurious, or “ghost” interactions, we provide a rigorous quantification of this effect through extensive simulations. Additionally, we further show that signal mixing also significantly limits the separability of neuronal phase and amplitude correlations. We conclude that spurious correlations must be carefully considered in connectivity analyses in MEG/EEG source space even when using measures that are immune to zero-lag correlations.
  • Pascucci, D., Hervais-Adelman, A., & Plomp, G. (2018). Gating by induced A-Gamma asynchrony in selective attention. Human Brain Mapping, 39(10), 3854-3870. doi:10.1002/hbm.24216.

    Abstract

    Visual selective attention operates through top–down mechanisms of signal enhancement and suppression, mediated by a-band oscillations. The effects of such top–down signals on local processing in primary visual cortex (V1) remain poorly understood. In this work, we characterize the interplay between large-s cale interactions and local activity changes in V1 that orchestrat es selective attention, using Granger-causality and phase-amplitude coupling (PAC) analysis of EEG source signals. The task required participants to either attend to or ignore oriented gratings. Results from time-varying, directed connectivity analysis revealed frequency-specific effects of attentional selection: bottom–up g-band influences from visual areas increased rapidly in response to attended stimuli while distributed top–down a-band influences originated from parietal cortex in response to ignored stimuli. Importantly, the results revealed a critical interplay between top–down parietal signals and a–g PAC in visual areas. Parietal a-band influences disrupted the a–g coupling in visual cortex, which in turn reduced the amount of g-band outflow from visual area s. Our results are a first demon stration of how directed interactions affect cross-frequency coupling in downstream areas depending on task demands. These findings suggest that parietal cortex realizes selective attention by disrupting cross-frequency coupling at target regions, which prevents them from propagating task-irrelevant information.
  • Peeters, D. (2018). A standardized set of 3D-objects for virtual reality research and applications. Behavior Research Methods, 50(3), 1047-1054. doi:10.3758/s13428-017-0925-3.

    Abstract

    The use of immersive virtual reality as a research tool is rapidly increasing in numerous scientific disciplines. By combining ecological validity with strict experimental control, immersive virtual reality provides the potential to develop and test scientific theory in rich environments that closely resemble everyday settings. This article introduces the first standardized database of colored three-dimensional (3D) objects that can be used in virtual reality and augmented reality research and applications. The 147 objects have been normed for name agreement, image agreement, familiarity, visual complexity, and corresponding lexical characteristics of the modal object names. The availability of standardized 3D-objects for virtual reality research is important, as reaching valid theoretical conclusions critically hinges on the use of well controlled experimental stimuli. Sharing standardized 3D-objects across different virtual reality labs will allow for science to move forward more quickly.
  • Peeters, D., & Dijkstra, T. (2018). Sustained inhibition of the native language in bilingual language production: A virtual reality approach. Bilingualism: Language and Cognition, 21(5), 1035-1061. doi:10.1017/S1366728917000396.

    Abstract

    Bilinguals often switch languages as a function of the language background of their addressee. The control mechanisms supporting bilinguals' ability to select the contextually appropriate language are heavily debated. Here we present four experiments in which unbalanced bilinguals named pictures in their first language Dutch and their second language English in mixed and blocked contexts. Immersive virtual reality technology was used to increase the ecological validity of the cued language-switching paradigm. Behaviorally, we consistently observed symmetrical switch costs, reversed language dominance, and asymmetrical mixing costs. These findings indicate that unbalanced bilinguals apply sustained inhibition to their dominant L1 in mixed language settings. Consequent enhanced processing costs for the L1 in a mixed versus a blocked context were reflected by a sustained positive component in event-related potentials. Methodologically, the use of virtual reality opens up a wide range of possibilities to study language and communication in bilingual and other communicative settings.
  • Piai, V., Rommers, J., & Knight, R. T. (2018). Lesion evidence for a critical role of left posterior but not frontal areas in alpha–beta power decreases during context-driven word production. European Journal of Neuroscience, 48(7), 2622-2629. doi:10.1111/ejn.13695.

    Abstract

    Different frequency bands in the electroencephalogram are postulated to support distinct language functions. Studies have suggested that alpha–beta power decreases may index word-retrieval processes. In context-driven word retrieval, participants hear lead-in sentences that either constrain the final word (‘He locked the door with the’) or not (‘She walked in here with the’). The last word is shown as a picture to be named. Previous studies have consistently found alpha–beta power decreases prior to picture onset for constrained relative to unconstrained sentences, localised to the left lateral-temporal and lateral-frontal lobes. However, the relative contribution of temporal versus frontal areas to alpha–beta power decreases is unknown. We recorded the electroencephalogram from patients with stroke lesions encompassing the left lateral-temporal and inferior-parietal regions or left-lateral frontal lobe and from matched controls. Individual participant analyses revealed a behavioural sentence context facilitation effect in all participants, except for in the two patients with extensive lesions to temporal and inferior parietal lobes. We replicated the alpha–beta power decreases prior to picture onset in all participants, except for in the two same patients with extensive posterior lesions. Thus, whereas posterior lesions eliminated the behavioural and oscillatory context effect, frontal lesions did not. Hierarchical clustering analyses of all patients’ lesion profiles, and behavioural and electrophysiological effects identified those two patients as having a unique combination of lesion distribution and context effects. These results indicate a critical role for the left lateral-temporal and inferior parietal lobes, but not frontal cortex, in generating the alpha–beta power decreases underlying context- driven word production.
  • Poletiek, F. H., Conway, C. M., Ellefson, M. R., Lai, J., Bocanegra, B. R., & Christiansen, M. H. (2018). Under what conditions can recursion be learned? Effects of starting small in artificial grammar learning of recursive structure. Cognitive Science, 42(8), 2855-2889. doi:10.1111/cogs.12685.

    Abstract

    It has been suggested that external and/or internal limitations paradoxically may lead to superior learning, that is, the concepts of starting small and less is more (Elman, 1993; Newport, 1990). In this paper, we explore the type of incremental ordering during training that might help learning, and what mechanism explains this facilitation. We report four artificial grammar learning experiments with human participants. In Experiments 1a and 1b we found a beneficial effect of starting small using two types of simple recursive grammars: right‐branching and center‐embedding, with recursive embedded clauses in fixed positions and fixed length. This effect was replicated in Experiment 2 (N = 100). In Experiment 3 and 4, we used a more complex center‐embedded grammar with recursive loops in variable positions, producing strings of variable length. When participants were presented an incremental ordering of training stimuli, as in natural language, they were better able to generalize their knowledge of simple units to more complex units when the training input “grew” according to structural complexity, compared to when it “grew” according to string length. Overall, the results suggest that starting small confers an advantage for learning complex center‐embedded structures when the input is organized according to structural complexity.
  • Popov, T., Jensen, O., & Schoffelen, J.-M. (2018). Dorsal and ventral cortices are coupled by cross-frequency interactions during working memory. NeuroImage, 178, 277-286. doi:10.1016/j.neuroimage.2018.05.054.

    Abstract

    Oscillatory activity in the alpha and gamma bands is considered key in shaping functional brain architecture. Power increases in the high-frequency gamma band are typically reported in parallel to decreases in the low-frequency alpha band. However, their functional significance and in particular their interactions are not well understood. The present study shows that, in the context of an N-backworking memory task, alpha power decreases in the dorsal visual stream are related to gamma power increases in early visual areas. Granger causality analysis revealed directed interregional interactions from dorsal to ventral stream areas, in accordance with task demands. Present results reveal a robust, behaviorally relevant, and architectonically decisive power-to-power relationship between alpha and gamma activity. This relationship suggests that anatomically distant power fluctuations in oscillatory activity can link cerebral network dynamics on trial-by-trial basis during cognitive operations such as working memory
  • Popov, T., Oostenveld, R., & Schoffelen, J.-M. (2018). FieldTrip made easy: An analysis protocol for group analysis of the auditory steady state brain response in time, frequency, and space. Frontiers in Neuroscience, 12: 711. doi:10.3389/fnins.2018.00711.

    Abstract

    The auditory steady state evoked response (ASSR) is a robust and frequently utilized phenomenon in psychophysiological research. It reflects the auditory cortical response to an amplitude-modulated constant carrier frequency signal. The present report provides a concrete example of a group analysis of the EEG data from 29 healthy human participants, recorded during an ASSR paradigm, using the FieldTrip toolbox. First, we demonstrate sensor-level analysis in the time domain, allowing for a description of the event-related potentials (ERPs), as well as their statistical evaluation. Second, frequency analysis is applied to describe the spectral characteristics of the ASSR, followed by group level statistical analysis in the frequency domain. Third, we show how timeand frequency-domain analysis approaches can be combined in order to describe the temporal and spectral development of the ASSR. Finally, we demonstrate source reconstruction techniques to characterize the primary neural generators of the ASSR. Throughout, we pay special attention to explaining the design of the analysis pipeline for single subjects and for the group level analysis. The pipeline presented here can be adjusted to accommodate other experimental paradigms and may serve as a template for similar analyses.
  • Rommers, J., & Federmeier, K. D. (2018). Electrophysiological methods. In A. M. B. De Groot, & P. Hagoort (Eds.), Research methods in psycholinguistics and the neurobiology of language: A practical guide (pp. 247-265). Hoboken: Wiley.
  • Rommers, J., & Federmeier, K. D. (2018). Lingering expectations: A pseudo-repetition effect for words previously expected but not presented. NeuroImage, 183, 263-272. doi:10.1016/j.neuroimage.2018.08.023.

    Abstract

    Prediction can help support rapid language processing. However, it is unclear whether prediction has downstream consequences, beyond processing in the moment. In particular, when a prediction is disconfirmed, does it linger, or is it suppressed? This study manipulated whether words were actually seen or were only expected, and probed their fate in memory by presenting the words (again) a few sentences later. If disconfirmed predictions linger, subsequent processing of the previously expected (but never presented) word should be similar to actual word repetition. At initial presentation, electrophysiological signatures of prediction disconfirmation demonstrated that participants had formed expectations. Further downstream, relative to unseen words, repeated words elicited a strong N400 decrease, an enhanced late positive complex (LPC), and late alpha band power decreases. Critically, like repeated words, words previously expected but not presented also attenuated the N400. This “pseudorepetition effect” suggests that disconfirmed predictions can linger at some stages of processing, and demonstrates that prediction has downstream consequences beyond rapid on-line processing
  • Rommers, J., & Federmeier, K. D. (2018). Predictability's aftermath: Downstream consequences of word predictability as revealed by repetition effects. Cortex, 101, 16-30. doi:10.1016/j.cortex.2017.12.018.

    Abstract

    Stimulus processing in language and beyond is shaped by context, with predictability having a particularly well-attested influence on the rapid processes that unfold during the presentation of a word. But does predictability also have downstream consequences for the quality of the constructed representations? On the one hand, the ease of processing predictablewordsmight free up time or cognitive resources, allowing for relatively thorough processing of the input. On the other hand, predictabilitymight allowthe systemto run in a top-down “verificationmode”, at the expense of thorough stimulus processing. This electroencephalogram (EEG) study manipulated word predictability, which reduced N400 amplitude and inter-trial phase clustering (ITPC), and then probed the fate of the (un)predictable words in memory by presenting them again. More thorough processing of predictable words should increase repetition effects, whereas less thorough processing should decrease them. Repetition was reflected in N400 decreases, late positive complex (LPC) enhancements, and late alpha/beta band power decreases. Critically, prior predictability tended to reduce the repetition effect on the N400, suggesting less priming, and eliminated the repetition effect on the LPC, suggesting a lack of episodic recollection. These findings converge on a top-down verification account, on which the brain processes more predictable input less thoroughly. More generally, the results demonstrate that predictability hasmultifaceted downstreamconsequences beyond processing in the moment
  • Seeliger, K., Fritsche, M., Güçlü, U., Schoenmakers, S., Schoffelen, J.-M., Bosch, S. E., & Van Gerven, M. A. J. (2018). Convolutional neural network-based encoding and decoding of visual object recognition in space and time. NeuroImage, 180, 253-266. doi:10.1016/j.neuroimage.2017.07.018.

    Abstract

    Representations learned by deep convolutional neural networks (CNNs) for object recognition are a widely investigated model of the processing hierarchy in the human visual system. Using functional magnetic resonance imaging, CNN representations of visual stimuli have previously been shown to correspond to processing stages in the ventral and dorsal streams of the visual system. Whether this correspondence between models and brain signals also holds for activity acquired at high temporal resolution has been explored less exhaustively. Here, we addressed this question by combining CNN-based encoding models with magnetoencephalography (MEG). Human participants passively viewed 1,000 images of objects while MEG signals were acquired. We modelled their high temporal resolution source-reconstructed cortical activity with CNNs, and observed a feed-forward sweep across the visual hierarchy between 75 and 200 ms after stimulus onset. This spatiotemporal cascade was captured by the network layer representations, where the increasingly abstract stimulus representation in the hierarchical network model was reflected in different parts of the visual cortex, following the visual ventral stream. We further validated the accuracy of our encoding model by decoding stimulus identity in a left-out validation set of viewed objects, achieving state-of-the-art decoding accuracy.
  • Segaert, K., Mazaheri, A., & Hagoort, P. (2018). Binding language: Structuring sentences through precisely timed oscillatory mechanisms. European Journal of Neuroscience, 48(7), 2651-2662. doi:10.1111/ejn.13816.

    Abstract

    Syntactic binding refers to combining words into larger structures. Using EEG, we investigated the neural processes involved in syntactic binding. Participants were auditorily presented two-word sentences (i.e. pronoun and pseudoverb such as ‘I grush’, ‘she grushes’, for which syntactic binding can take place) and wordlists (i.e. two pseudoverbs such as ‘pob grush’, ‘pob grushes’, for which no binding occurs). Comparing these two conditions, we targeted syntactic binding while minimizing contributions of semantic binding and of other cognitive processes such as working memory. We found a converging pattern of results using two distinct analysis approaches: one approach using frequency bands as defined in previous literature, and one data-driven approach in which we looked at the entire range of frequencies between 3-30 Hz without the constraints of pre-defined frequency bands. In the syntactic binding (relative to the wordlist) condition, a power increase was observed in the alpha and beta frequency range shortly preceding the presentation of the target word that requires binding, which was maximal over frontal-central electrodes. Our interpretation is that these signatures reflect that language comprehenders expect the need for binding to occur. Following the presentation of the target word in a syntactic binding context (relative to the wordlist condition), an increase in alpha power maximal over a left lateralized cluster of frontal-temporal electrodes was observed. We suggest that this alpha increase relates to syntactic binding taking place. Taken together, our findings suggest that increases in alpha and beta power are reflections of distinct the neural processes underlying syntactic binding.
  • Silva, S., Folia, V., Inácio, F., Castro, S. L., & Petersson, K. M. (2018). Modality effects in implicit artificial grammar learning: An EEG study. Brain Research, 1687, 50-59. doi:10.1016/j.brainres.2018.02.020.

    Abstract

    Recently, it has been proposed that sequence learning engages a combination of modality-specific operating networks and modality-independent computational principles. In the present study, we compared the behavioural and EEG outcomes of implicit artificial grammar learning in the visual vs. auditory modality. We controlled for the influence of surface characteristics of sequences (Associative Chunk Strength), thus focusing on the strictly structural aspects of sequence learning, and we adapted the paradigms to compensate for known frailties of the visual modality compared to audition (temporal presentation, fast presentation rate). The behavioural outcomes were similar across modalities. Favouring the idea of modality-specificity, ERPs in response to grammar violations differed in topography and latency (earlier and more anterior component in the visual modality), and ERPs in response to surface features emerged only in the auditory modality. In favour of modality-independence, we observed three common functional properties in the late ERPs of the two grammars: both were free of interactions between structural and surface influences, both were more extended in a grammaticality classification test than in a preference classification test, and both correlated positively and strongly with theta event-related-synchronization during baseline testing. Our findings support the idea of modality-specificity combined with modality-independence, and suggest that memory for visual vs. auditory sequences may largely contribute to cross-modal differences.
  • Sjerps, M. J., Zhang, C., & Peng, G. (2018). Lexical Tone is Perceived Relative to Locally Surrounding Context, Vowel Quality to Preceding Context. Journal of Experimental Psychology: Human Perception and Performance, 44(6), 914-924. doi:10.1037/xhp0000504.

    Abstract

    Important speech cues such as lexical tone and vowel quality are perceptually contrasted to the distribution of those same cues in surrounding contexts. However, it is unclear whether preceding and following contexts have similar influences, and to what extent those influences are modulated by the auditory history of previous trials. To investigate this, Cantonese participants labeled sounds from (a) a tone continuum (mid- to high-level), presented with a context that had raised or lowered F0 values and (b) a vowel quality continuum (/u/ to /o/), where the context had raised or lowered F1 values. Contexts with high or low F0/F1 were presented in separate blocks or intermixed in 1 block. Contexts were presented following (Experiment 1) or preceding the target continuum (Experiment 2). Contrastive effects were found for both tone and vowel quality (e.g., decreased F0 values in contexts lead to more high tone target judgments and vice versa). Importantly, however, lexical tone was only influenced by F0 in immediately preceding and following contexts. Vowel quality was only influenced by the F1 in preceding contexts, but this extended to contexts from preceding trials. Contextual influences on tone and vowel quality are qualitatively different, which has important implications for understanding the mechanism of context effects in speech perception.
  • Stolk, A., Griffin, S., Van der Meij, R., Dewar, C., Saez, I., Lin, J. J., Piantoni, G., Schoffelen, J.-M., Knight, R. T., & Oostenveld, R. (2018). Integrated analysis of anatomical and electrophysiological human intracranial data. Nature Protocols, 13, 1699-1723. doi:10.1038/s41596-018-0009-6.

    Abstract

    Human intracranial electroencephalography (iEEG) recordings provide data with much greater spatiotemporal precision than is possible from data obtained using scalp EEG, magnetoencephalography (MEG), or functional MRI. Until recently, the fusion of anatomical data (MRI and computed tomography (CT) images) with electrophysiological data and their subsequent analysis have required the use of technologically and conceptually challenging combinations of software. Here, we describe a comprehensive protocol that enables complex raw human iEEG data to be converted into more readily comprehensible illustrative representations. The protocol uses an open-source toolbox for electrophysiological data analysis (FieldTrip). This allows iEEG researchers to build on a continuously growing body of scriptable and reproducible analysis methods that, over the past decade, have been developed and used by a large research community. In this protocol, we describe how to analyze complex iEEG datasets by providing an intuitive and rapid approach that can handle both neuroanatomical information and large electrophysiological datasets. We provide a worked example using an example dataset. We also explain how to automate the protocol and adjust the settings to enable analysis of iEEG datasets with other characteristics. The protocol can be implemented by a graduate student or postdoctoral fellow with minimal MATLAB experience and takes approximately an hour to execute, excluding the automated cortical surface extraction.
  • Tan, Y., & Martin, R. C. (2018). Verbal short-term memory capacities and executive function in semantic and syntactic interference resolution during sentence comprehension: Evidence from aphasia. Neuropsychologia, 113, 111-125. doi:10.1016/j.neuropsychologia.2018.03.001.

    Abstract

    This study examined the role of verbal short-term memory (STM) and executive function (EF) underlying semantic and syntactic interference resolution during sentence comprehension for persons with aphasia (PWA) with varying degrees of STM and EF deficits. Semantic interference was manipulated by varying the semantic plausibility of the intervening NP as subject of the verb and syntactic interference was manipulated by varying whether the NP was another subject or an object. Nine PWA were assessed on sentence reading times and on comprehension question performance. PWA showed exaggerated semantic and syntactic interference effects relative to healthy age-matched control subjects. Importantly, correlational analyses showed that while answering comprehension questions, PWA’ semantic STM capacity related to their ability to resolve semantic but not syntactic interference. In contrast, PWA’ EF abilities related to their ability to resolve syntactic but not semantic interference. Phonological STM deficits were not related to the ability to resolve either type of interference. The results for semantic STM are consistent with prior findings indicating a role for semantic but not phonological STM in sentence comprehension, specifically with regard to maintaining semantic information prior to integration. The results for syntactic interference are consistent with the recent findings suggesting that EF is critical for syntactic processing.
  • Tromp, J., Peeters, D., Meyer, A. S., & Hagoort, P. (2018). The combined use of Virtual Reality and EEG to study language processing in naturalistic environments. Behavior Research Methods, 50(2), 862-869. doi:10.3758/s13428-017-0911-9.

    Abstract

    When we comprehend language, we often do this in rich settings in which we can use many cues to understand what someone is saying. However, it has traditionally been difficult to design experiments with rich three-dimensional contexts that resemble our everyday environments, while maintaining control over the linguistic and non-linguistic information that is available. Here we test the validity of combining electroencephalography (EEG) and Virtual Reality (VR) to overcome this problem. We recorded electrophysiological brain activity during language processing in a well-controlled three-dimensional virtual audiovisual environment. Participants were immersed in a virtual restaurant, while wearing EEG equipment. In the restaurant participants encountered virtual restaurant guests. Each guest was seated at a separate table with an object on it (e.g. a plate with salmon). The restaurant guest would then produce a sentence (e.g. “I just ordered this salmon.”). The noun in the spoken sentence could either match (“salmon”) or mismatch (“pasta”) with the object on the table, creating a situation in which the auditory information was either appropriate or inappropriate in the visual context. We observed a reliable N400 effect as a consequence of the mismatch. This finding validates the combined use of VR and EEG as a tool to study the neurophysiological mechanisms of everyday language comprehension in rich, ecologically valid settings.
  • Udden, J., & Männel, C. (2018). Artificial grammar learning and its neurobiology in relation to language processing and development. In S.-A. Rueschemeyer, & M. G. Gaskell (Eds.), The Oxford Handbook of Psycholinguistics (2nd ed., pp. 755-783). Oxford: Oxford University Press.

    Abstract

    The artificial grammar learning (AGL) paradigm enables systematic investigation of the acquisition of linguistically relevant structures. It is a paradigm of interest for language processing research, interfacing with theoretical linguistics, and for comparative research on language acquisition and evolution. This chapter presents a key for understanding major variants of the paradigm. An unbiased summary of neuroimaging findings of AGL is presented, using meta-analytic methods, pointing to the crucial involvement of the bilateral frontal operculum and regions in the right lateral hemisphere. Against a background of robust posterior temporal cortex involvement in processing complex syntax, the evidence for involvement of the posterior temporal cortex in AGL is reviewed. Infant AGL studies testing for neural substrates are reviewed, covering the acquisition of adjacent and non-adjacent dependencies as well as algebraic rules. The language acquisition data suggest that comparisons of learnability of complex grammars performed with adults may now also be possible with children.
  • Van den Broek, G., Takashima, A., Segers, E., & Verhoeven, L. (2018). Contextual Richness and Word Learning: Context Enhances Comprehension but Retrieval Enhances Retention. Language Learning, 68(2), 546-585. doi:10.1111/lang.12285.

    Abstract

    Learning new vocabulary from context typically requires multiple encounters during which word meaning can be retrieved from memory or inferred from context. We compared the effect of memory retrieval and context inferences on short‐ and long‐term retention in three experiments. Participants studied novel words and then practiced the words either in an uninformative context that required the retrieval of word meaning from memory (“I need the funguo”) or in an informative context from which word meaning could be inferred (“I want to unlock the door: I need the funguo”). The informative context facilitated word comprehension during practice. However, later recall of word form and meaning and word recognition in a new context were better after successful retrieval practice and retrieval practice with feedback than after context‐inference practice. These findings suggest benefits of retrieval during contextualized vocabulary learning whereby the uninformative context enhanced word retention by triggering memory retrieval.
  • Van Bergen, G., & Bosker, H. R. (2018). Linguistic expectation management in online discourse processing: An investigation of Dutch inderdaad 'indeed' and eigenlijk 'actually'. Journal of Memory and Language, 103, 191-209. doi:10.1016/j.jml.2018.08.004.

    Abstract

    Interpersonal discourse particles (DPs), such as Dutch inderdaad (≈‘indeed’) and eigenlijk (≈‘actually’) are highly frequent in everyday conversational interaction. Despite extensive theoretical descriptions of their polyfunctionality, little is known about how they are used by language comprehenders. In two visual world eye-tracking experiments involving an online dialogue completion task, we asked to what extent inderdaad, confirming an inferred expectation, and eigenlijk, contrasting with an inferred expectation, influence real-time understanding of dialogues. Answers in the dialogues contained a DP or a control adverb, and a critical discourse referent was replaced by a beep; participants chose the most likely dialogue completion by clicking on one of four referents in a display. Results show that listeners make rapid and fine-grained situation-specific inferences about the use of DPs, modulating their expectations about how the dialogue will unfold. Findings further specify and constrain theories about the conversation-managing function and polyfunctionality of DPs.
  • Van Campen, A. D., Kunert, R., Van den Wildenberg, W. P. M., & Ridderinkhof, K. R. (2018). Repetitive transcranial magnetic stimulation over inferior frontal cortex impairs the suppression (but not expression) of action impulses during action conflict. Psychophysiology, 55(3): e13003. doi:10.1111/psyp.13003.

    Abstract

    In the recent literature, the effects of noninvasive neurostimulation on cognitive functioning appear to lack consistency and replicability. We propose that such effects may be concealed unless dedicated, sensitive, and process-specific dependent measures are used. The expression and subsequent suppression of response capture are often studied using conflict tasks. Response-time distribution analyses have been argued to provide specific measures of the susceptibility to make fast impulsive response errors, as well as the proficiency of the selective suppression of these impulses. These measures of response capture and response inhibition are particularly sensitive to experimental manipulations and clinical deficiencies that are typically obfuscated in commonly used overall performance analyses. Recent work using structural and functional imaging techniques links these behavioral outcome measures to the integrity of frontostriatal networks. These studies suggest that the presupplementary motor area (pre-SMA) is linked to the susceptibility to response capture whereas the right inferior frontal cortex (rIFC) is associated with the selective suppression of action impulses. Here, we used repetitive transcranial magnetic stimulation (rTMS) to test the causal involvement of these two cortical areas in response capture and inhibition in the Simon task. Disruption of rIFC function specifically impaired selective suppression of conflicting action tendencies, whereas the anticipated increase of fast impulsive errors after perturbing pre-SMA function was not confirmed. These results provide a proof of principle of the notion that the selection of appropriate dependent measures is perhaps crucial to establish the effects of neurostimulation on specific cognitive functions.
  • Vanlangendonck, F., Takashima, A., Willems, R. M., & Hagoort, P. (2018). Distinguishable memory retrieval networks for collaboratively and non-collaboratively learned information. Neuropsychologia, 111, 123-132. doi:10.1016/j.neuropsychologia.2017.12.008.

    Abstract

    Learning often occurs in communicative and collaborative settings, yet almost all research into the neural basis of memory relies on participants encoding and retrieving information on their own. We investigated whether learning linguistic labels in a collaborative context at least partly relies on cognitively and neurally distinct representations, as compared to learning in an individual context. Healthy human participants learned labels for sets of abstract shapes in three different tasks. They came up with labels with another person in a collaborative communication task (collaborative condition), by themselves (individual condition), or were given pre-determined unrelated labels to learn by themselves (arbitrary condition). Immediately after learning, participants retrieved and produced the labels aloud during a communicative task in the MRI scanner. The fMRI results show that the retrieval of collaboratively generated labels as compared to individually learned labels engages brain regions involved in understanding others (mentalizing or theory of mind) and autobiographical memory, including the medial prefrontal cortex, the right temporoparietal junction and the precuneus. This study is the first to show that collaboration during encoding affects the neural networks involved in retrieval.
  • Vanlangendonck, F., Willems, R. M., & Hagoort, P. (2018). Taking common ground into account: Specifying the role of the mentalizing network in communicative language production. PLoS One, 13(10): e0202943. doi:10.1371/journal.pone.0202943.
  • Varma, S., Daselaar, S. M., Kessels, R. P. C., & Takashima, A. (2018). Promotion and suppression of autobiographical thinking differentially affect episodic memory consolidation. PLoS One, 13(8): e0201780. doi:10.1371/journal.pone.0201780.

    Abstract

    During a post-encoding delay period, the ongoing consolidation of recently acquired memories can suffer interference if the delay period involves encoding of new memories, or sensory stimulation tasks. Interestingly, two recent independent studies suggest that (i) autobiographical thinking also interferes markedly with ongoing consolidation of recently learned wordlist material, while (ii) a 2-Back task might not interfere with ongoing consolidation, possibly due to the suppression of autobiographical thinking. In this study, we directly compare these conditions against a quiet wakeful rest baseline to test whether the promotion (via familiar sound-cues) or suppression (via a 2-Back task) of autobiographical thinking during the post-encoding delay period can affect consolidation of studied wordlists in a negative or a positive way, respectively. Our results successfully replicate previous studies and show a significant interference effect (as compared to the rest condition) when learning is followed by familiar sound-cues that promote autobiographical thinking, whereas no interference effect is observed when learning is followed by the 2-Back task. Results from a post-experimental experience-sampling questionnaire further show significant differences in the degree of autobiographical thinking reported during the three post-encoding periods: highest in the presence of sound-cues and lowest during the 2-Back task. In conclusion, our results suggest that varying levels of autobiographical thought during the post-encoding period may modulate episodic memory consolidation.
  • Wang, L., Hagoort, P., & Jensen, O. (2018). Gamma oscillatory activity related to language prediction. Journal of Cognitive Neuroscience, 30(8), 1075-1085. doi:10.1162/jocn_a_01275.

    Abstract

    Using magnetoencephalography, the current study examined gamma activity associated with language prediction. Participants read high- and low-constraining sentences in which the final word of the sentence was either expected or unexpected. Although no consistent gamma power difference induced by the sentence-final words was found between the expected and unexpected conditions, the correlation of gamma power during the prediction and activation intervals of the sentence-final words was larger when the presented words matched with the prediction compared with when the prediction was violated or when no prediction was available. This suggests that gamma magnitude relates to the match between predicted and perceived words. Moreover, the expected words induced activity with a slower gamma frequency compared with that induced by unexpected words. Overall, the current study establishes that prediction is related to gamma power correlations and a slowing of the gamma frequency.
  • Wang, L., Hagoort, P., & Jensen, O. (2018). Language prediction is reflected by coupling between frontal gamma and posterior alpha oscillations. Journal of Cognitive Neuroscience, 30(3), 432-447. doi:10.1162/jocn_a_01190.

    Abstract

    Readers and listeners actively predict upcoming words during language processing. These predictions might serve to support the unification of incoming words into sentence context and thus rely on interactions between areas in the language network. In the current magnetoencephalography study, participants read sentences that varied in contextual constraints so that the predictability of the sentence-final words was either high or low. Before the sentence-final words, we observed stronger alpha power suppression for the highly compared with low constraining sentences in the left inferior frontal cortex, left posterior temporal region, and visual word form area. Importantly, the temporal and visual word form area alpha power correlated negatively with left frontal gamma power for the highly constraining sentences. We suggest that the correlation between alpha power decrease in temporal language areas and left prefrontal gamma power reflects the initiation of an anticipatory unification process in the language network.
  • Willems, R. M., & Van Gerven, M. (2018). New fMRI methods for the study of language. In S.-A. Rueschemeyer, & M. G. Gaskell (Eds.), The Oxford Handbook of Psycholinguistics (2nd ed., pp. 975-991). Oxford: Oxford University Press.
  • Willems, R. M., & Cristia, A. (2018). Hemodynamic methods: fMRI and fNIRS. In A. M. B. De Groot, & P. Hagoort (Eds.), Research methods in psycholinguistics and the neurobiology of language: A practical guide (pp. 266-287). Hoboken: Wiley.
  • 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. (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.
  • 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
  • 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., 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., 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., 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., 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., 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., 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., van Dam, W., Conant, L. L., Binder, J. R., & Desai, R. H. (2015). Familiarity differentially affects right hemisphere contributions to processing metaphors and literals. Frontiers in Human Neuroscience, 9: 44. doi:10.3389/fnhum.2015.00044.

    Abstract

    The role of the two hemispheres in processing metaphoric language is controversial. While some studies have reported a special role of the right hemisphere (RH) in processing metaphors, others indicate no difference in laterality relative to literal language. Some studies have found a role of the RH for novel/unfamiliar metaphors, but not conventional/familiar metaphors. It is not clear, however, whether the role of the RH is specific to metaphor novelty, or whether it reflects processing, reinterpretation or reanalysis of novel/unfamiliar language in general. Here we used functional magnetic resonance imaging (fMRI) to examine the effects of familiarity in both metaphoric and non-metaphoric sentences. A left lateralized network containing the middle and inferior frontal gyri, posterior temporal regions in the left hemisphere (LH), and inferior frontal regions in the RH, was engaged across both metaphoric and non-metaphoric sentences; engagement of this network decreased as familiarity decreased. No region was engaged selectively for greater metaphoric unfamiliarity. An analysis of laterality, however, showed that the contribution of the RH relative to that of LH does increase in a metaphorspecific manner as familiarity decreases. These results show that RH regions, taken by themselves, including commonly reported regions such as the right inferior frontal gyrus (IFG), are responsive to increased cognitive demands of processing unfamiliar stimuli, rather than being metaphor-selective. The division of labor between the two hemispheres, however, does shift towards the right for metaphoric processing. The shift results not because the RH contributes more to metaphoric processing. Rather, relative to its contribution for processing literals, the LH contributes less.
  • 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.”
  • Lewis, A. G., & Bastiaansen, M. C. M. (2015). A predictive coding framework for rapid neural dynamics during sentence-level language comprehension. Cortex, 68, 155-168. doi:10.1016/j.cortex.2015.02.014.

    Abstract

    There is a growing literature investigating the relationship between oscillatory neural dynamics measured using EEG and/or MEG, and sentence-level language comprehension. Recent proposals have suggested a strong link between predictive coding accounts of the hierarchical flow of information in the brain, and oscillatory neural dynamics in the beta and gamma frequency ranges. We propose that findings relating beta and gamma oscillations to sentence-level language comprehension might be unified under such a predictive coding account. Our suggestion is that oscillatory activity in the beta frequency range may reflect both the active maintenance of the current network configuration responsible for representing the sentence-level meaning under construction, and the top-down propagation of predictions to hierarchically lower processing levels based on that representation. In addition, we suggest that oscillatory activity in the low and middle gamma range reflect the matching of top-down predictions with bottom-up linguistic input, while evoked high gamma might reflect the propagation of bottom-up prediction errors to higher levels of the processing hierarchy. We also discuss some of the implications of this predictive coding framework, and we outline ideas for how these might be tested experimentally
  • Lewis, A. G., Wang, L., & Bastiaansen, M. C. M. (2015). Fast oscillatory dynamics during language comprehension: Unification versus maintenance and prediction? Brain and Language, 148, 51-63. doi:10.1016/j.bandl.2015.01.003.

    Abstract

    The role of neuronal oscillations during language comprehension is not yet well understood. In this paper we review and reinterpret the functional roles of beta- and gamma-band oscillatory activity during language comprehension at the sentence and discourse level. We discuss the evidence in favor of a role for beta and gamma in unification (the unification hypothesis), and in light of mounting evidence that cannot be accounted for under this hypothesis, we explore an alternative proposal linking beta and gamma oscillations to maintenance and prediction (respectively) during language comprehension. Our maintenance/prediction hypothesis is able to account for most of the findings that are currently available relating beta and gamma oscillations to language comprehension, and is in good agreement with other proposals about the roles of beta and gamma in domain-general cognitive processing. In conclusion we discuss proposals for further testing and comparing the prediction and unification hypotheses.
  • Lockwood, G., & Dingemanse, M. (2015). Iconicity in the lab: A review of behavioural, developmental, and neuroimaging research into sound-symbolism. Frontiers in Psychology, 6: 1246. doi:10.3389/fpsyg.2015.01246.

    Abstract

    This review covers experimental approaches to sound-symbolism—from infants to adults, and from Sapir’s foundational studies to twenty-first century product naming. It synthesizes recent behavioral, developmental, and neuroimaging work into a systematic overview of the cross-modal correspondences that underpin iconic links between form and meaning. It also identifies open questions and opportunities, showing how the future course of experimental iconicity research can benefit from an integrated interdisciplinary perspective. Combining insights from psychology and neuroscience with evidence from natural languages provides us with opportunities for the experimental investigation of the role of sound-symbolism in language learning, language processing, and communication. The review finishes by describing how hypothesis-testing and model-building will help contribute to a cumulative science of sound-symbolism in human language.
  • Lockwood, G., & Tuomainen, J. (2015). Ideophones in Japanese modulate the P2 and late positive complex responses. Frontiers in Psychology, 6: 933. doi:10.3389/fpsyg.2015.00933.

    Abstract

    Sound-symbolism, or the direct link between sound and meaning, is typologically and behaviorally attested across languages. However, neuroimaging research has mostly focused on artificial non-words or individual segments, which do not represent sound-symbolism in natural language. We used EEG to compare Japanese ideophones, which are phonologically distinctive sound-symbolic lexical words, and arbitrary adverbs during a sentence reading task. Ideophones elicit a larger visual P2 response and a sustained late positive complex in comparison to arbitrary adverbs. These results and previous literature suggest that the larger P2 may indicate the integration of sound and sensory information by association in response to the distinctive phonology of ideophones. The late positive complex may reflect the facilitated lexical retrieval of ideophones in comparison to arbitrary words. This account provides new evidence that ideophones exhibit similar cross-modal correspondences to those which have been proposed for non-words and individual sounds, and that these effects are detectable in natural language.
  • Moreno, I., De Vega, M., León, I., Bastiaansen, M. C. M., Lewis, A. G., & Magyari, L. (2015). Brain dynamics in the comprehension of action-related language. A time-frequency analysis of mu rhythms. Neuroimage, 109, 50-62. doi:10.1016/j.neuroimage.2015.01.018.

    Abstract

    EEG mu rhythms (8-13Hz) recorded at fronto-central electrodes are generally considered as markers of motor cortical activity in humans, because they are modulated when participants perform an action, when they observe another’s action or even when they imagine performing an action. In this study, we analyzed the time-frequency (TF) modulation of mu rhythms while participants read action language (“You will cut the strawberry cake”), abstract language (“You will doubt the patient´s argument”), and perceptive language (“You will notice the bright day”). The results indicated that mu suppression at fronto-central sites is associated with action language rather than with abstract or perceptive language. Also, the largest difference between conditions occurred quite late in the sentence, while reading the first noun, (contrast Action vs. Abstract), or the second noun following the action verb (contrast Action vs. Perceptive). This suggests that motor activation is associated with the integration of words across the sentence beyond the lexical processing of the action verb. Source reconstruction localized mu suppression associated with action sentences in premotor cortex (BA 6). The present study suggests (1) that the understanding of action language activates motor networks in the human brain, and (2) that this activation occurs online based on semantic integration across multiple words in the sentence.
  • Nijhoff, A. D., & Willems, R. M. (2015). Simulating fiction: Individual differences in literature comprehension revealed with fMRI. PLoS One, 10(2): e0116492. doi:10.1371/journal.pone.0116492.

    Abstract

    When we read literary fiction, we are transported to fictional places, and we feel and think along with the characters. Despite the importance of narrative in adult life and during development, the neurocognitive mechanisms underlying fiction comprehension are unclear. We used functional magnetic resonance imaging (fMRI) to investigate how individuals differently employ neural networks important for understanding others’ beliefs and intentions (mentalizing), and for sensori-motor simulation while listening to excerpts from literary novels. Localizer tasks were used to localize both the cortical motor network and the mentalizing network in participants after they listened to excerpts from literary novels. Results show that participants who had high activation in anterior medial prefrontal cortex (aMPFC; part of the mentalizing network) when listening to mentalizing content of literary fiction, had lower motor cortex activity when they listened to action-related content of the story, and vice versa. This qualifies how people differ in their engagement with fiction: some people are mostly drawn into a story by mentalizing about the thoughts and beliefs of others, whereas others engage in literature by simulating more concrete events such as actions. This study provides on-line neural evidence for the existence of qualitatively different styles of moving into literary worlds, and adds to a growing body of literature showing the potential to study narrative comprehension with neuroimaging methods.
  • Peeters, D. (2015). A social and neurobiological approach to pointing in speech and gesture. PhD Thesis, Radboud University, Nijmegen.
  • Peeters, D., Chu, M., Holler, J., Hagoort, P., & Ozyurek, A. (2015). Electrophysiological and kinematic correlates of communicative intent in the planning and production of pointing gestures and speech. Journal of Cognitive Neuroscience, 27(12), 2352-2368. doi:10.1162/jocn_a_00865.

    Abstract

    In everyday human communication, we often express our communicative intentions by manually pointing out referents in the material world around us to an addressee, often in tight synchronization with referential speech. This study investigated whether and how the kinematic form of index finger pointing gestures is shaped by the gesturer's communicative intentions and how this is modulated by the presence of concurrently produced speech. Furthermore, we explored the neural mechanisms underpinning the planning of communicative pointing gestures and speech. Two experiments were carried out in which participants pointed at referents for an addressee while the informativeness of their gestures and speech was varied. Kinematic and electrophysiological data were recorded online. It was found that participants prolonged the duration of the stroke and poststroke hold phase of their gesture to be more communicative, in particular when the gesture was carrying the main informational burden in their multimodal utterance. Frontal and P300 effects in the ERPs suggested the importance of intentional and modality-independent attentional mechanisms during the planning phase of informative pointing gestures. These findings contribute to a better understanding of the complex interplay between action, attention, intention, and language in the production of pointing gestures, a communicative act core to human interaction.

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