Publications

Displaying 1 - 78 of 78
  • 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.

    Additional information

    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.
  • 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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    Data Accessibility
  • 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., 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.
  • 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., 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.
  • 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.

    Additional information

    mmc1.docx
  • 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
  • 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.

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    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). 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.
  • 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.
  • Allen, S., Ozyurek, A., Kita, S., Brown, A., Furman, R., Ishizuka, T., & Fujii, M. (2007). Language-specific and universal influences in children's syntactic packaging of manner and path: A comparison of English, Japanese, and Turkish. Cognition, 102, 16-48. doi:10.1016/j.cognition.2005.12.006.

    Abstract

    Different languages map semantic elements of spatial relations onto different lexical and syntactic units. These crosslinguistic differences raise important questions for language development in terms of how this variation is learned by children. We investigated how Turkish-, English-, and Japanese-speaking children (mean age 3;8) package the semantic elements of Manner and Path onto syntactic units when both the Manner and the Path of the moving Figure occur simultaneously and are salient in the event depicted. Both universal and language-specific patterns were evident in our data. Children used the semantic-syntactic mappings preferred by adult speakers of their own languages, and even expressed subtle syntactic differences that encode different relations between Manner and Path in the same way as their adult counterparts (i.e., Manner causing vs. incidental to Path). However, not all types of semantics-syntax mappings were easy for children to learn (e.g., expressing Manner and Path elements in two verbal clauses). In such cases, Turkish- and Japanese-speaking children frequently used syntactic patterns that were not typical in the target language but were similar to patterns used by English-speaking children, suggesting some universal influence. Thus, both language-specific and universal tendencies guide the development of complex spatial expressions.
  • Bramão, I., Mendonça, A., Faísca, L., Ingvar, M., Petersson, K. M., & Reis, A. (2007). The impact of reading and writing skills on a visuo-motor integration task: A comparison between illiterate and literate subjects. Journal of the International Neuropsychological Society, 13(2), 359-364. doi:10.1017/S1355617707070440.

    Abstract

    Previous studies have shown a significant association between reading skills and the performance on visuo-motor tasks. In order to clarify whether reading and writing skills modulate non-linguistic domains, we investigated the performance of two literacy groups on a visuo-motor integration task with non-linguistic stimuli. Twenty-one illiterate participants and twenty matched literate controls were included in the experiment. Subjects were instructed to use the right or the left index finger to point to and touch a randomly presented target on the right or left side of a touch screen. The results showed that the literate subjects were significantly faster in detecting and touching targets on the left compared to the right side of the screen. In contrast, the presentation side did not affect the performance of the illiterate group. These results lend support to the idea that having acquired reading and writing skills, and thus a preferred left-to-right reading direction, influences visual scanning. (JINS, 2007, 13, 359–364
  • Furman, R., & Ozyurek, A. (2007). Development of interactional discourse markers: Insights from Turkish children's and adults' narratives. Journal of Pragmatics, 39(10), 1742-1757. doi:10.1016/j.pragma.2007.01.008.

    Abstract

    Discourse markers (DMs) are linguistic elements that index different relations and coherence between units of talk (Schiffrin, Deborah, 1987. Discourse Markers. Cambridge University Press, Cambridge). Most research on the development of these forms has focused on conversations rather than narratives and furthermore has not directly compared children's use of DMs to adult usage. This study examines the development of three DMs (şey ‘uuhh’, yani ‘I mean’, işte ‘y’know’) that mark interactional levels of discourse in oral Turkish narratives in 60 Turkish children (3-, 5- and 9-year-olds) and 20 Turkish-speaking adults. The results show that the frequency and functions of DMs change with age. Children learn şey, which mainly marks exchange level structures, earliest. However, yani and işte have multi-functions such as marking both information states and participation frameworks and are consequently learned later. Children also use DMs with different functions than adults. Overall, the results show that learning to use interactional DMs in narratives is complex and goes beyond age 9, especially for multi-functional DMs that index an interplay of discourse coherence at different levels.
  • Gisselgard, J., Uddén, J., Ingvar, M., & Petersson, K. M. (2007). Disruption of order information by irrelevant items: A serial recognition paradigm. Acta Psychologica, 124(3), 356-369. doi:10.1016/j.actpsy.2006.04.002.

    Abstract

    Irrelevant speech effect (ISE) is defined as a decrement in visually presented digit-list short-term memory performance due to exposure to irrelevant auditory material. Perhaps the most successful theoretical explanation of the effect is the changing state hypothesis. This hypothesis explains the effect in terms of confusion between amodal serial order cues, and represents a view based on the interference caused by the processing of similar order information of the visual and auditory materials. An alternative view suggests that the interference occurs as a consequence of the similarity between the visual and auditory contents of the stimuli. An important argument for the former view is the observation that ISE is almost exclusively observed in tasks that require memory for serial order. However, most short-term memory tasks require that both item and order information be retained in memory. An ideal task to investigate the sensitivity of maintenance of serial order to irrelevant speech would be one that calls upon order information but not item information. One task that is particularly suited to address this issue is serial recognition. In a typical serial recognition task, a list of items is presented and then probed by the same list in which the order of two adjacent items has been transposed. Due to the re-presentation of the encoding string, serial recognition requires primarily the serial order to be maintained while the content of the presented items is deemphasized. In demonstrating a highly significant ISE of changing versus steady-state auditory items in a serial recognition task, the present finding lends support for and extends previous empirical findings suggesting that irrelevant speech has the potential to interfere with the coding of the order of the items to be memorized.
  • Hagoort, P., & Van Berkum, J. J. A. (2007). Beyond the sentence given. Philosophical Transactions of the Royal Society. Series B: Biological Sciences, 362, 801-811.

    Abstract

    A central and influential idea among researchers of language is that our language faculty is organized according to Fregean compositionality, which states that the meaning of an utterance is a function of the meaning of its parts and of the syntactic rules by which these parts are combined. Since the domain of syntactic rules is the sentence, the implication of this idea is that language interpretation takes place in a two-step fashion. First, the meaning of a sentence is computed. In a second step, the sentence meaning is integrated with information from prior discourse, world knowledge, information about the speaker and semantic information from extra-linguistic domains such as co-speech gestures or the visual world. Here, we present results from recordings of event-related brain potentials that are inconsistent with this classical two-step model of language interpretation. Our data support a one-step model in which knowledge about the context and the world, concomitant information from other modalities, and the speaker are brought to bear immediately, by the same fast-acting brain system that combines the meanings of individual words into a message-level representation. Underlying the one-step model is the immediacy assumption, according to which all available information will immediately be used to co-determine the interpretation of the speaker's message. Functional magnetic resonance imaging data that we collected indicate that Broca's area plays an important role in semantic unification. Language comprehension involves the rapid incorporation of information in a 'single unification space', coming from a broader range of cognitive domains than presupposed in the standard two-step model of interpretation.
  • Hald, L. A., Steenbeek-Planting, E. G., & Hagoort, P. (2007). The interaction of discourse context and world knowledge in online sentence comprehension: Evidence from the N400. Brain Research, 1146, 210-218. doi:10.1016/j.brainres.2007.02.054.

    Abstract

    In an ERP experiment we investigated how the recruitment and integration of world knowledge information relate to the integration of information within a current discourse context. Participants were presented with short discourse contexts which were followed by a sentence that contained a critical word that was correct or incorrect based on general world knowledge and the supporting discourse context, or was more or less acceptable based on the combination of general world knowledge and the specific local discourse context. Relative to the critical word in the correct world knowledge sentences following a neutral discourse, all other critical words elicited an N400 effect that began at about 300 ms after word onset. However, the magnitude of the N400 effect varied in a way that suggests an interaction between world knowledge and discourse context. The results indicate that both world knowledge and discourse context have an effect on sentence interpretation, but neither overrides the other.
  • Janzen, G., Wagensveld, B., & Van Turennout, M. (2007). Neural representation of navigational relevance is rapidly induced and long lasting. Cerebral Cortex, 17(4), 975-981. doi:10.1093/cercor/bhl008.

    Abstract

    Successful navigation is facilitated by the presence of landmarks. Previous functional magnetic resonance imaging (fMRI) evidence indicated that the human parahippocampal gyrus automatically distinguishes between landmarks placed at navigationally relevant (decision points) and irrelevant locations (nondecision points). This storage of navigational relevance can provide a neural mechanism underlying successful navigation. However, an efficient wayfinding mechanism requires that important spatial information is learned quickly and maintained over time. The present study investigates whether the representation of navigational relevance is modulated by time and practice. Participants learned 2 film sequences through virtual mazes containing objects at decision and at nondecision points. One maze was shown one time, and the other maze was shown 3 times. Twenty-four hours after study, event-related fMRI data were acquired during recognition of the objects. The results showed that activity in the parahippocampal gyrus was increased for objects previously placed at decision points as compared with objects placed at nondecision points. The decision point effect was not modulated by the number of exposures to the mazes and independent of explicit memory functions. These findings suggest a persistent representation of navigationally relevant information, which is stable after only one exposure to an environment. These rapidly induced and long-lasting changes in object representation provide a basis for successful wayfinding.
  • Janzen, G., & Weststeijn, C. G. (2007). Neural representation of object location and route direction: An event-related fMRI study. Brain Research, 1165, 116-125. doi:10.1016/j.brainres.2007.05.074.

    Abstract

    The human brain distinguishes between landmarks placed at navigationally relevant and irrelevant locations. However, to provide a successful wayfinding mechanism not only landmarks but also the routes between them need to be stored. We examined the neural representation of a memory for route direction and a memory for relevant landmarks. Healthy human adults viewed objects along a route through a virtual maze. Event-related functional magnetic resonance imaging (fMRI) data were acquired during a subsequent subliminal priming recognition task. Prime-objects either preceded or succeeded a target-object on a preciously learned route. Our results provide evidence that the parahippocampal gyri distinguish between relevant and irrelevant landmarks whereas the inferior parietal gyrus, the anterior cingulate gyrus as well as the right caudate nucleus are involved in the coding of route direction. These data show that separated memory systems store different spatial information. A memory for navigationally relevant object information and a memory for route direction exist.
  • Kita, S., Ozyurek, A., Allen, S., Brown, A., Furman, R., & Ishizuka, T. (2007). Relations between syntactic encoding and co-speech gestures: Implications for a model of speech and gesture production. Language and Cognitive Processes, 22(8), 1212-1236. doi:10.1080/01690960701461426.

    Abstract

    Gestures that accompany speech are known to be tightly coupled with speech production. However little is known about the cognitive processes that underlie this link. Previous cross-linguistic research has provided preliminary evidence for online interaction between the two systems based on the systematic co-variation found between how different languages syntactically package Manner and Path information of a motion event and how gestures represent Manner and Path. Here we elaborate on this finding by testing whether speakers within the same language gesturally express Manner and Path differently according to their online choice of syntactic packaging of Manner and Path, or whether gestural expression is pre-determined by a habitual conceptual schema congruent with the linguistic typology. Typologically congruent and incongruent syntactic structures for expressing Manner and Path (i.e., in a single clause or multiple clauses) were elicited from English speakers. We found that gestural expressions were determined by the online choice of syntactic packaging rather than by a habitual conceptual schema. It is therefore concluded that speech and gesture production processes interface online at the conceptual planning phase. Implications of the findings for models of speech and gesture production are discussed
  • Marklund, P., Fransson, P., Cabeza, R., Petersson, K. M., Ingvar, M., & Nyberg, L. (2007). Sustained and transient neural modulations in prefrontal cortex related to declarative long-term memory, working memory, and attention. Cortex, 43(1), 22-37. doi:10.1016/S0010-9452(08)70443-X.

    Abstract

    Common activations in prefrontal cortex (PFC) during episodic and semantic long-term memory (LTM) tasks have been hypothesized to reflect functional overlap in terms of working memory (WM) and cognitive control. To evaluate a WM account of LTM-general activations, the present study took into consideration that cognitive task performance depends on the dynamic operation of multiple component processes, some of which are stimulus-synchronous and transient in nature; and some that are engaged throughout a task in a sustained fashion. PFC and WM may be implicated in both of these temporally independent components. To elucidate these possibilities we employed mixed blocked/event-related functional magnetic resonance imaging (fMRI) procedures to assess the extent to which sustained or transient activation patterns overlapped across tasks indexing episodic and semantic LTM, attention (ATT), and WM. Within PFC, ventrolateral and medial areas exhibited sustained activity across all tasks, whereas more anterior regions including right frontopolar cortex were commonly engaged in sustained processing during the three memory tasks. These findings do not support a WM account of sustained frontal responses during LTM tasks, but instead suggest that the pattern that was common to all tasks reflects general attentional set/vigilance, and that the shared WM-LTM pattern mediates control processes related to upholding task set. Transient responses during the three memory tasks were assessed relative to ATT to isolate item-specific mnemonic processes and were found to be largely distinct from sustained effects. Task-specific effects were observed for each memory task. In addition, a common item response for all memory tasks involved left dorsolateral PFC (DLPFC). The latter response might be seen as reflecting WM processes during LTM retrieval. Thus, our findings suggest that a WM account of shared PFC recruitment in LTM tasks holds for common transient item-related responses rather than sustained state-related responses that are better seen as reflecting more general attentional/control processes.
  • Menenti, L., & Burani, C. (2007). What causes the effect of age of acquisition in lexical processing? Quarterly Journal of Experimental Psychology, 60(5), 652-660. doi:10.1080/17470210601100126.

    Abstract

    Three hypotheses for effects of age of acquisition (AoA) in lexical processing are compared: the cumulative frequency hypothesis (frequency and AoA both influence the number of encounters with a word, which influences processing speed), the semantic hypothesis (early-acquired words are processed faster because they are more central in the semantic network), and the neural network model (early-acquired words are faster because they are acquired when a network has maximum plasticity). In a regression study of lexical decision (LD) and semantic categorization (SC) in Italian and Dutch, contrary to the cumulative frequency hypothesis, AoA coefficients were larger than frequency coefficients, and, contrary to the semantic hypothesis, the effect of AoA was not larger in SC than in LD. The neural network model was supported.
  • Nieuwland, M. S., Petersson, K. M., & Van Berkum, J. J. A. (2007). On sense and reference: Examining the functional neuroanatomy of referential processing. NeuroImage, 37(3), 993-1004. doi:10.1016/j.neuroimage.2007.05.048.

    Abstract

    In an event-related fMRI study, we examined the cortical networks involved in establishing reference during language comprehension. We compared BOLD responses to sentences containing referentially ambiguous pronouns (e.g., “Ronald told Frank that he…”), referentially failing pronouns (e.g., “Rose told Emily that he…”) or coherent pronouns. Referential ambiguity selectively recruited medial prefrontal regions, suggesting that readers engaged in problem-solving to select a unique referent from the discourse model. Referential failure elicited activation increases in brain regions associated with morpho-syntactic processing, and, for those readers who took failing pronouns to refer to unmentioned entities, additional regions associated with elaborative inferencing were observed. The networks activated by these two referential problems did not overlap with the network activated by a standard semantic anomaly. Instead, we observed a double dissociation, in that the systems activated by semantic anomaly are deactivated by referential ambiguity, and vice versa. This inverse coupling may reflect the dynamic recruitment of semantic and episodic processing to resolve semantically or referentially problematic situations. More generally, our findings suggest that neurocognitive accounts of language comprehension need to address not just how we parse a sentence and combine individual word meanings, but also how we determine who's who and what's what during language comprehension.
  • Nieuwland, M. S., Otten, M., & Van Berkum, J. J. A. (2007). Who are you talking about? Tracking discourse-level referential processing with event-related brain potentials. Journal of Cognitive Neuroscience, 19(2), 228-236. doi:10.1162/jocn.2007.19.2.228.

    Abstract

    In this event-related brain potentials (ERPs) study, we explored the possibility to selectively track referential ambiguity during spoken discourse comprehension. Earlier ERP research has shown that referentially ambiguous nouns (e.g., “the girl” in a two-girl context) elicit a frontal, sustained negative shift relative to unambiguous control words. In the current study, we examined whether this ERP effect reflects “deep” situation model ambiguity or “superficial” textbase ambiguity. We contrasted these different interpretations by investigating whether a discourse-level semantic manipulation that prevents referential ambiguity also averts the elicitation of a referentially induced ERP effect. We compared ERPs elicited by nouns that were referentially nonambiguous but were associated with two discourse entities (e.g., “the girl” with two girls introduced in the context, but one of which has died or left the scene), with referentially ambiguous and nonambiguous control words. Although temporally referentially ambiguous nouns elicited a frontal negative shift compared to control words, the “double bound” but referentially nonambiguous nouns did not. These results suggest that it is possible to selectively track referential ambiguity with ERPs at the level that is most relevant to discourse comprehension, the situation model.
  • Otten, M., Nieuwland, M. S., & Van Berkum, J. J. A. (2007). Great expectations: Specific lexical anticipation influences the processing of spoken language. BMC Neuroscience, 8: 89. doi:10.1186/1471-2202-8-89.

    Abstract

    Background Recently several studies have shown that people use contextual information to make predictions about the rest of the sentence or story as the text unfolds. Using event related potentials (ERPs) we tested whether these on-line predictions are based on a message-based representation of the discourse or on simple automatic activation by individual words. Subjects heard short stories that were highly constraining for one specific noun, or stories that were not specifically predictive but contained the same prime words as the predictive stories. To test whether listeners make specific predictions critical nouns were preceded by an adjective that was inflected according to, or in contrast with, the gender of the expected noun. Results When the message of the preceding discourse was predictive, adjectives with an unexpected gender-inflection evoked a negative deflection over right-frontal electrodes between 300 and 600 ms. This effect was not present in the prime control context, indicating that the prediction mismatch does not hinge on word-based priming but is based on the actual message of the discourse. Conclusions When listening to a constraining discourse people rapidly make very specific predictions about the remainder of the story, as the story unfolds. These predictions are not simply based on word-based automatic activation, but take into account the actual message of the discourse.
  • Otten, M., & Van Berkum, J. J. A. (2007). What makes a discourse constraining? Comparing the effects of discourse message and scenario fit on the discourse-dependent N400 effect. Brain Research, 1153, 166-177. doi:10.1016/j.brainres.2007.03.058.

    Abstract

    A discourse context provides a reader with a great deal of information that can provide constraints for further language processing, at several different levels. In this experiment we used event-related potentials (ERPs) to explore whether discourse-generated contextual constraints are based on the precise message of the discourse or, more `loosely', on the scenario suggested by one or more content words in the text. Participants read constraining stories whose precise message rendered a particular word highly predictable ("The manager thought that the board of directors should assemble to discuss the issue. He planned a...[meeting]") as well as non-constraining control stories that were only biasing in virtue of the scenario suggested by some of the words ("The manager thought that the board of directors need not assemble to discuss the issue. He planned a..."). Coherent words that were inconsistent with the message-level expectation raised in a constraining discourse (e.g., "session" instead of "meeting") elicited a classic centroparietal N400 effect. However, when the same words were only inconsistent with the scenario loosely suggested by earlier words in the text, they elicited a different negativity around 400 ms, with a more anterior, left-lateralized maximum. The fact that the discourse-dependent N400 effect cannot be reduced to scenario-mediated priming reveals that it reflects the rapid use of precise message-level constraints in comprehension. At the same time, the left-lateralized negativity in non-constraining stories suggests that, at least in the absence of strong message-level constraints, scenario-mediated priming does also rapidly affect comprehension.
  • Ozyurek, A., & Kelly, S. D. (2007). Gesture, language, and brain. Brain and Language, 101(3), 181-185. doi:10.1016/j.bandl.2007.03.006.
  • Ozyurek, A., Willems, R. M., Kita, S., & Hagoort, P. (2007). On-line integration of semantic information from speech and gesture: Insights from event-related brain potentials. Journal of Cognitive Neuroscience, 19(4), 605-616. doi:10.1162/jocn.2007.19.4.605.

    Abstract

    During language comprehension, listeners use the global semantic representation from previous sentence or discourse context to immediately integrate the meaning of each upcoming word into the unfolding message-level representation. Here we investigate whether communicative gestures that often spontaneously co-occur with speech are processed in a similar fashion and integrated to previous sentence context in the same way as lexical meaning. Event-related potentials were measured while subjects listened to spoken sentences with a critical verb (e.g., knock), which was accompanied by an iconic co-speech gesture (i.e., KNOCK). Verbal and/or gestural semantic content matched or mismatched the content of the preceding part of the sentence. Despite the difference in the modality and in the specificity of meaning conveyed by spoken words and gestures, the latency, amplitude, and topographical distribution of both word and gesture mismatches are found to be similar, indicating that the brain integrates both types of information simultaneously. This provides evidence for the claim that neural processing in language comprehension involves the simultaneous incorporation of information coming from a broader domain of cognition than only verbal semantics. The neural evidence for similar integration of information from speech and gesture emphasizes the tight interconnection between speech and co-speech gestures.
  • Petersson, K. M., Silva, C., Castro-Caldas, A., Ingvar, M., & Reis, A. (2007). Literacy: A cultural influence on functional left-right differences in the inferior parietal cortex. European Journal of Neuroscience, 26(3), 791-799. doi:10.1111/j.1460-9568.2007.05701.x.

    Abstract

    The current understanding of hemispheric interaction is limited. Functional hemispheric specialization is likely to depend on both genetic and environmental factors. In the present study we investigated the importance of one factor, literacy, for the functional lateralization in the inferior parietal cortex in two independent samples of literate and illiterate subjects. The results show that the illiterate group are consistently more right-lateralized than their literate controls. In contrast, the two groups showed a similar degree of left-right differences in early speech-related regions of the superior temporal cortex. These results provide evidence suggesting that a cultural factor, literacy, influences the functional hemispheric balance in reading and verbal working memory-related regions. In a third sample, we investigated grey and white matter with voxel-based morphometry. The results showed differences between literacy groups in white matter intensities related to the mid-body region of the corpus callosum and the inferior parietal and parietotemporal regions (literate > illiterate). There were no corresponding differences in the grey matter. This suggests that the influence of literacy on brain structure related to reading and verbal working memory is affecting large-scale brain connectivity more than grey matter per se.
  • Qin, S., Piekema, C., Petersson, K. M., Han, B., Luo, J., & Fernández, G. (2007). Probing the transformation of discontinuous associations into episodic memory: An event-related fMRI study. NeuroImage, 38(1), 212-222. doi:10.1016/j.neuroimage.2007.07.020.

    Abstract

    Using event-related functional magnetic resonance imaging, we identified brain regions involved in storing associations of events discontinuous in time into long-term memory. Participants were scanned while memorizing item-triplets including simultaneous and discontinuous associations. Subsequent memory tests showed that participants remembered both types of associations equally well. First, by constructing the contrast between the subsequent memory effects for discontinuous associations and simultaneous associations, we identified the left posterior parahippocampal region, dorsolateral prefrontal cortex, the basal ganglia, posterior midline structures, and the middle temporal gyrus as being specifically involved in transforming discontinuous associations into episodic memory. Second, we replicated that the prefrontal cortex and the medial temporal lobe (MTL) especially the hippocampus are involved in associative memory formation in general. Our findings provide evidence for distinct neural operation(s) that supports the binding and storing discontinuous associations in memory. We suggest that top-down signals from the prefrontal cortex and MTL may trigger reactivation of internal representation in posterior midline structures of the first event, thus allowing it to be associated with the second event. The dorsolateral prefrontal cortex together with basal ganglia may support this encoding operation by executive and binding processes within working memory, and the posterior parahippocampal region may play a role in binding and memory formation.
  • Reis, A., Faísca, L., Mendonça, S., Ingvar, M., & Petersson, K. M. (2007). Semantic interference on a phonological task in illiterate subjects. Scandinavian Journal of Psychology, 48(1), 69-74. doi:10.1111/j.1467-9450.2006.00544.x.

    Abstract

    Previous research suggests that learning an alphabetic written language influences aspects of the auditory-verbal language system. In this study, we examined whether literacy influences the notion of words as phonological units independent of lexical semantics in literate and illiterate subjects. Subjects had to decide which item in a word- or pseudoword pair was phonologically longest. By manipulating the relationship between referent size and phonological length in three word conditions (congruent, neutral, and incongruent) we could examine to what extent subjects focused on form rather than meaning of the stimulus material. Moreover, the pseudoword condition allowed us to examine global phonological awareness independent of lexical semantics. The results showed that literate performed significantly better than illiterate subjects in the neutral and incongruent word conditions as well as in the pseudoword condition. The illiterate group performed least well in the incongruent condition and significantly better in the pseudoword condition compared to the neutral and incongruent word conditions and suggest that performance on phonological word length comparisons is dependent on literacy. In addition, the results show that the illiterate participants are able to perceive and process phonological length, albeit less well than the literate subjects, when no semantic interference is present. In conclusion, the present results confirm and extend the finding that illiterate subjects are biased towards semantic-conceptual-pragmatic types of cognitive processing.
  • Snijders, T. M., Kooijman, V., Cutler, A., & Hagoort, P. (2007). Neurophysiological evidence of delayed segmentation in a foreign language. Brain Research, 1178, 106-113. doi:10.1016/j.brainres.2007.07.080.

    Abstract

    Previous studies have shown that segmentation skills are language-specific, making it difficult to segment continuous speech in an unfamiliar language into its component words. Here we present the first study capturing the delay in segmentation and recognition in the foreign listener using ERPs. We compared the ability of Dutch adults and of English adults without knowledge of Dutch (‘foreign listeners’) to segment familiarized words from continuous Dutch speech. We used the known effect of repetition on the event-related potential (ERP) as an index of recognition of words in continuous speech. Our results show that word repetitions in isolation are recognized with equivalent facility by native and foreign listeners, but word repetitions in continuous speech are not. First, words familiarized in isolation are recognized faster by native than by foreign listeners when they are repeated in continuous speech. Second, when words that have previously been heard only in a continuous-speech context re-occur in continuous speech, the repetition is detected by native listeners, but is not detected by foreign listeners. A preceding speech context facilitates word recognition for native listeners, but delays or even inhibits word recognition for foreign listeners. We propose that the apparent difference in segmentation rate between native and foreign listeners is grounded in the difference in language-specific skills available to the listeners.
  • Takashima, A., Nieuwenhuis, I. L. C., Rijpkema, M., Petersson, K. M., Jensen, O., & Fernández, G. (2007). Memory trace stabilization leads to large-scale changes in the retrieval network: A functional MRI study on associative memory. Learning & Memory, 14, 472-479. doi:10.1101/lm.605607.

    Abstract

    Spaced learning with time to consolidate leads to more stabile memory traces. However, little is known about the neural correlates of trace stabilization, especially in humans. The present fMRI study contrasted retrieval activity of two well-learned sets of face-location associations, one learned in a massed style and tested on the day of learning (i.e., labile condition) and another learned in a spaced scheme over the course of one week (i.e., stabilized condition). Both sets of associations were retrieved equally well, but the retrieval of stabilized association was faster and accompanied by large-scale changes in the network supporting retrieval. Cued recall of stabilized as compared with labile associations was accompanied by increased activity in the precuneus, the ventromedial prefrontal cortex, the bilateral temporal pole, and left temporo–parietal junction. Conversely, memory representational areas such as the fusiform gyrus for faces and the posterior parietal cortex for locations did not change their activity with stabilization. The changes in activation in the precuneus, which also showed increased connectivity with the fusiform area, are likely to be related to the spatial nature of our task. The activation increase in the ventromedial prefrontal cortex, on the other hand, might reflect a general function in stabilized memory retrieval. This area might succeed the hippocampus in linking distributed neocortical representations.
  • Tendolkar, I., Arnold, J., Petersson, K. M., Weis, S., Brockhaus-Dumke, A., Van Eijndhoven, P., Buitelaar, J., & Fernández, G. (2007). Probing the neural correlates of associative memory formation: A parametrically analyzed event-related functional MRI study. Brain Research, 1142, 159-168. doi:10.1016/j.brainres.2007.01.040.

    Abstract

    The medial temporal lobe (MTL) is crucial for declarative memory formation, but the function of its subcomponents in associative memory formation remains controversial. Most functional imaging studies on this topic are based on a stepwise approach comparing a condition with and one without associative encoding. Extending this approach we applied additionally a parametric analysis by varying the amount of associative memory formation. We found a hippocampal subsequent memory effect of almost similar magnitude regardless of the amount of associations formed. By contrast, subsequent memory effects in rhinal and parahippocampal cortices were parametrically and positively modulated by the amount of associations formed. Our results indicate that the parahippocampal region supports associative memory formation as tested here and the hippocampus adds a general mnemonic operation. This pattern of results might suggest a new interpretation. Instead of having either a fixed division of labor between the hippocampus (associative memory formation) and the rhinal cortex (non-associative memory formation) or a functionally unitary MTL system, in which all substructures are contributing to memory formation in a similar way, we propose that the location where associations are formed within the MTL depends on the kind of associations bound: If visual single-dimension associations, as used here, can already be integrated within the parahippocampal region, the hippocampus might add a general purpose mnemonic operation only. In contrast, if associations have to be formed across widely distributed neocortical representations, the hippocampus may provide a binding operation in order to establish a coherent memory.
  • Van Berkum, J. J. A., Koornneef, A. W., Otten, M., & Nieuwland, M. S. (2007). Establishing reference in language comprehension: An electrophysiological perspective. Brain Research, 1146, 158-171. doi:10.1016/j.brainres.2006.06.091.

    Abstract

    The electrophysiology of language comprehension has long been dominated by research on syntactic and semantic integration. However, to understand expressions like "he did it" or "the little girl", combining word meanings in accordance with semantic and syntactic constraints is not enough--readers and listeners also need to work out what or who is being referred to. We review our event-related brain potential research on the processes involved in establishing reference, and present a new experiment in which we examine when and how the implicit causality associated with specific interpersonal verbs affects the interpretation of a referentially ambiguous pronoun. The evidence suggests that upon encountering a singular noun or pronoun, readers and listeners immediately inspect their situation model for a suitable discourse entity, such that they can discriminate between having too many, too few, or exactly the right number of referents within at most half a second. Furthermore, our implicit causality findings indicate that a fragment like "David praised Linda because..." can immediately foreground a particular referent, to the extent that a subsequent "he" is at least initially construed as a syntactic error. In all, our brain potential findings suggest that referential processing is highly incremental, and not necessarily contingent upon the syntax. In addition, they demonstrate that we can use ERPs to relatively selectively keep track of how readers and listeners establish reference.
  • Wassenaar, M., & Hagoort, P. (2007). Thematic role assignment in patients with Broca's aphasia: Sentence-picture matching electrified. Neuropsychologia, 45(4), 716-740. doi:10.1016/j.neuropsychologia.2006.08.016.

    Abstract

    An event-related brain potential experiment was carried out to investigate on-line thematic role assignment during sentence–picture matching in patients with Broca's aphasia. Subjects were presented with a picture that was followed by an auditory sentence. The sentence either matched the picture or mismatched the visual information depicted. Sentences differed in complexity, and ranged from simple active semantically irreversible sentences to passive semantically reversible sentences. ERPs were recorded while subjects were engaged in sentence–picture matching. In addition, reaction time and accuracy were measured. Three groups of subjects were tested: Broca patients (N = 10), non-aphasic patients with a right hemisphere (RH) lesion (N = 8), and healthy aged-matched controls (N = 15). The results of this study showed that, in neurologically unimpaired individuals, thematic role assignment in the context of visual information was an immediate process. This in contrast to patients with Broca's aphasia who demonstrated no signs of on-line sensitivity to the picture–sentence mismatches. The syntactic contribution to the thematic role assignment process seemed to be diminished given the reduction and even absence of P600 effects. Nevertheless, Broca patients showed some off-line behavioral sensitivity to the sentence–picture mismatches. The long response latencies of Broca's aphasics make it likely that off-line response strategies were used.
  • Willems, R. M., & Hagoort, P. (2007). Neural evidence for the interplay between language, gesture, and action: A review. Brain and Language, 101(3), 278-289. doi:10.1016/j.bandl.2007.03.004.

    Abstract

    Co-speech gestures embody a form of manual action that is tightly coupled to the language system. As such, the co-occurrence of speech and co-speech gestures is an excellent example of the interplay between language and action. There are, however, other ways in which language and action can be thought of as closely related. In this paper we will give an overview of studies in cognitive neuroscience that examine the neural underpinnings of links between language and action. Topics include neurocognitive studies of motor representations of speech sounds, action-related language, sign language and co-speech gestures. It will be concluded that there is strong evidence on the interaction between speech and gestures in the brain. This interaction however shares general properties with other domains in which there is interplay between language and action.
  • Willems, R. M., Ozyurek, A., & Hagoort, P. (2007). When language meets action: The neural integration of gesture and speech. Cerebral Cortex, 17(10), 2322-2333. doi:10.1093/cercor/bhl141.

    Abstract

    Although generally studied in isolation, language and action often co-occur in everyday life. Here we investigated one particular form of simultaneous language and action, namely speech and gestures that speakers use in everyday communication. In a functional magnetic resonance imaging study, we identified the neural networks involved in the integration of semantic information from speech and gestures. Verbal and/or gestural content could be integrated easily or less easily with the content of the preceding part of speech. Premotor areas involved in action observation (Brodmann area [BA] 6) were found to be specifically modulated by action information "mismatching" to a language context. Importantly, an increase in integration load of both verbal and gestural information into prior speech context activated Broca's area and adjacent cortex (BA 45/47). A classical language area, Broca's area, is not only recruited for language-internal processing but also when action observation is integrated with speech. These findings provide direct evidence that action and language processing share a high-level neural integration system.

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