Publications

Abstract

In this study, we compared the performance of a long short-term memory (LSTM) neural network to the behavior of human participants on a language task that requires hierarchically structured knowledge. We show that humans interpret ambiguous noun phrases, such as second blue ball, in line with their hierarchical constituent structure. LSTMs, instead, only do
so after unambiguous training, and they do not systematically generalize to novel items. Overall, the results of our simulations indicate that a model can behave hierarchically without relying on hierarchical constituent structure.

Abstract

The endeavor to understand the human brain has seen more progress in the last few decades than in the previous two millennia. Still, our understanding of how the human brain relates to behavior in the real world and how this link is modulated by biological, social, and environmental factors is limited. To address this, we designed the Healthy Brain Study (HBS), an interdisciplinary, longitudinal, cohort study based on multidimensional, dynamic assessments in both the laboratory and the real world. Here, we describe the rationale and design of the currently ongoing HBS. The HBS is examining a population-based sample of 1,000 healthy participants (age 30-39) who are thoroughly studied across an entire year. Data are collected through cognitive, affective, behavioral, and physiological testing, neuroimaging, bio-sampling, questionnaires, ecological momentary assessment, and real-world assessments using wearable devices. These data will become an accessible resource for the scientific community enabling the next step in understanding the human brain and how it dynamically and individually operates in its bio-social context. An access procedure to the collected data and bio-samples is in place and published on https://www.healthybrainstudy.nl/en/data-and-methods.

https://www.trialregister.nl/trial/7955

Abstract

The ability to predict upcoming actions is a hallmark of cognition. It remains unclear, however, whether the predictive behaviour observed in controlled lab environments generalises to rich, everyday settings. In four virtual reality experiments, we tested whether a well-established marker of linguistic prediction (anticipatory eye movements) replicated when increasing the naturalness of the paradigm by means of immersing participants in naturalistic scenes (Experiment 1), increasing the number of distractor objects (Experiment 2), modifying the proportion of predictable noun-referents (Experiment 3), and manipulating the location of referents relative to the joint attentional space (Experiment 4). Robust anticipatory eye movements were observed for Experiments 1–3. The anticipatory effect disappeared, however, in Experiment 4. Our findings suggest that predictive processing occurs in everyday communication if the referents are situated in the joint attentional space. Methodologically, our study confirms that ecological validity and experimental control may go hand-in-hand in the study of human predictive behaviour.

Abstract

The present ERP study assessed whether grammatical aspect is used as a cue in online event comprehension, in particular when reading about events in which an object is visually changed. While perfective aspect cues holistic event representations, including an event's endpoint, progressive aspect highlights intermediate phases of an event. In a 2 × 3 design, participants read SVO sentences describing a change-of-state event (e.g., to chop an onion), with grammatical Aspect manipulated (perfective “chopped” vs progressive “was chopping”). Thereafter, they saw a Picture of an object either having undergone substantial state-change (SC; a chopped onion), no state-change (NSC; an onion in its original state) or an unrelated object (U; a cactus, acting as control condition). Their task was to decide whether the object in the Picture was mentioned in the sentence. We focused on N400 modulation, with ERPs time-locked to picture onset. U pictures elicited an N400 response as expected, suggesting detection of categorical mismatches in object type. For SC and NSC pictures, a whole-head follow-up analysis revealed a P300, implying people were engaged in detailed evaluation of pictures of matching objects. SC pictures received most positive responses overall. Crucially, there was an interaction of Aspect and Picture: SC pictures resulted in a higher amplitude P300 after sentences in the perfective compared to the progressive. Thus, while the perfective cued for a holistic event representation, including the resultant state of the affected object (i.e., the chopped onion) constraining object representations online, the progressive defocused event completion and object-state change. Grammatical aspect thus guided online event comprehension by cueing the visual representation(s) of an object's state.

Abstract

Brain connectivity plays a major role in the encoding, transfer, and
integration of sensory information. Interregional synchronization
of neural oscillations in the γ-frequency band has been suggested
as a key mechanism underlying perceptual integration. In a recent
study, we found evidence for this hypothesis showing that the
modulation of interhemispheric oscillatory synchrony by means of
bihemispheric high-density transcranial alternating current stimulation
(HD-TACS) affects binaural integration of dichotic acoustic features.
Here, we aimed to establish a direct link between oscillatory
synchrony, effective brain connectivity, and binaural integration.
We experimentally manipulated oscillatory synchrony (using bihemispheric
γ-TACS with different interhemispheric phase lags) and
assessed the effect on effective brain connectivity and binaural integration
(as measured with functional MRI and a dichotic listening
task, respectively). We found that TACS reduced intrahemispheric
connectivity within the auditory cortices and antiphase (interhemispheric
phase lag 180°) TACS modulated connectivity between the
two auditory cortices. Importantly, the changes in intra- and interhemispheric
connectivity induced by TACS were correlated with
changes in perceptual integration. Our results indicate that γ-band
synchronization between the two auditory cortices plays a functional
role in binaural integration, supporting the proposed role
of interregional oscillatory synchrony in perceptual integration.

Abstract

Linguistic labels exert a particularly strong top-down influence on perception. The potency of this influence has been ascribed to their ability to evoke category-diagnostic features of concepts. In doing this, they facilitate the formation of a perceptual template concordant with those features, effectively biasing perceptual activation towards the labelled category. In this study, we employ a cueing paradigm with moving, point-light stimuli across three experiments, in order to examine how the number of biological motion features (form and kinematics) encoded in lexical cues modulates the efficacy of lexical top-down influence on perception. We find that the magnitude of lexical influence on biological motion perception rises as a function of the number of biological motion-relevant features carried by both cue and target. When lexical cues encode multiple biological motion features, this influence is robust enough to mislead participants into reporting erroneous percepts, even when a masking level yielding high performance is used.

Abstract

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

Abstract

We sketch four applications of Marr's levels-of-analysis methodology to the relations between logic and experimental data in the cognitive neuroscience of language and reasoning. The first part of the paper illustrates the explanatory power of computational level theories based on logic. We show that a Bayesian treatment of the suppression task in reasoning with conditionals is ruled out by EEG data, supporting instead an analysis based on defeasible logic. Further, we describe how results from an EEG study on temporal prepositions can be reanalyzed using formal semantics, addressing a potential confound. The second part of the article demonstrates the predictive power of logical theories drawing on EEG data on processing progressive constructions and on behavioral data on conditional reasoning in people with autism. Logical theories can constrain processing hypotheses all the way down to neurophysiology, and conversely neuroscience data can guide the selection of alternative computational level models of cognition.

Abstract

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

Abstract

During sentence level language comprehension, semantic and syntactic unification are functionally distinct operations. Nevertheless, both recruit roughly the same brain areas (spatially overlapping networks in the left frontotemporal cortex) and happen at the same time (in the first few hundred milliseconds after word onset). We tested the hypothesis that semantic and syntactic unification are segregated by means of neuronal synchronization of the functionally relevant networks in different frequency ranges: gamma (40 Hz and up) for semantic unification and lower beta (10–20 Hz) for syntactic unification. EEG power changes were quantified as participants read either correct sentences, syntactically correct though meaningless sentences (syntactic prose), or sentences that did not contain any syntactic structure (random word lists). Other sentences contained either a semantic anomaly or a syntactic violation at a critical word in the sentence. Larger EEG gamma-band power was observed for semantically coherent than for semantically anomalous sentences. Similarly, beta-band power was larger for syntactically correct sentences than for incorrect ones. These results confirm the existence of a functional dissociation in EEG oscillatory dynamics during sentence level language comprehension that is compatible with the notion of a frequency-based segregation of syntactic and semantic unification.

Abstract

Previous studies suggest that linguistic material can modulate visual perception, but it is unclear at which level of processing these interactions occur. Here we aim to dissociate between two competing models of language–perception interactions: a feed-forward and a feedback model. We capitalized on the fact that the models make different predictions on the role of feedback. We presented unmasked (aware) or masked (unaware) words implying motion (e.g. “rise,” “fall”), directly preceding an upward or downward visual motion stimulus. Crucially, masking leaves intact feed-forward information processing from low- to high-level regions, whereas it abolishes subsequent feedback. Under this condition, participants remained faster and more accurate when the direction implied by the motion word was congruent with the direction of the visual motion stimulus. This suggests that language–perception interactions are driven by the feed-forward convergence of linguistic and perceptual information at higher-level conceptual and decision stages.

Abstract

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

Abstract

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

Abstract

This study aims to test a prediction of recent
theoretical frameworks in speech motor control: if speech production targets are specified in auditory
terms, people with better auditory acuity should have more precise speech targets.
To investigate this, we had participants perform speech perception and production tasks in a counterbalanced order. To assess speech perception acuity, we used an adaptive speech discrimination
task. To assess variability in speech production, participants performed a pseudo-word reading task; formant values were measured for each recording.
We predicted that speech production variability to correlate inversely with discrimination performance.
The results suggest that people do vary in their production and perceptual abilities, and that better discriminators have more distinctive vowel production targets, confirming our prediction. This
study highlights the importance of individual
differences in the study of speech motor control, and sheds light on speech production-perception interaction.

Abstract

Models of speech production explain event-related suppression of the auditory cortical
response as reflecting a comparison between auditory predictions and feedback. The present MEG
study was designed to test two predictions from this framework: 1) whether the reduced auditory
response varies as a function of the mismatch between prediction and feedback; 2) whether individual
variation in this response is predictive of speech-motor adaptation.
Participants alternated between online imitation and listening tasks. In the imitation task, participants
began each trial producing the same vowel (/e/) and subsequently listened to and imitated auditorilypresented
vowels varying in acoustic distance from /e/.
Results replicated suppression, with a smaller M100 during speaking than listening. Although we did
not find unequivocal support for the first prediction, participants with less M100 suppression were
better at the imitation task. These results are consistent with the enhancement of M100 serving as an
error signal to drive subsequent speech-motor adaptation.

Abstract

The genetic determinants of cerebral asymmetries are unknown. Sex differences in asymmetry of the planum temporale, that overlaps Wernicke’s classical language area, have been inconsistently reported. Meta-analysis of previous studies has suggested that publication bias established this sex difference in the literature. Using probabilistic definitions of cortical regions we screened over the cerebral cortex for sexual dimorphisms of asymmetry in 2337 healthy subjects, and found the planum temporale to show the strongest sex-linked asymmetry of all regions, which was supported by two further datasets, and also by analysis with the Freesurfer package that performs automated parcellation of cerebral cortical regions. We performed a genome-wide association scan meta-analysis of planum temporale asymmetry in a pooled sample of 3095 subjects, followed by a candidate-driven approach which measured a significant enrichment of association in genes of the ´steroid hormone receptor activity´ and 'steroid metabolic process' pathways. Variants in the genes and pathways identified may affect the role of the planum temporale in language cognition.

Abstract

Recipients process information from speech and co-speech gestures, but it is currently unknown how this processing is influenced by the presence of other important social cues, especially gaze direction, a marker of communicative intent. Such cues may modulate neural activity in regions associated either with the processing of ostensive cues, such as eye gaze, or with the processing of semantic information, provided by speech and gesture.
Participants were scanned (fMRI) while taking part in triadic communication involving two recipients and a speaker. The speaker uttered sentences that
were and were not accompanied by complementary iconic gestures. Crucially, the speaker alternated her gaze direction, thus creating two recipient roles: addressed (direct gaze) vs unaddressed (averted gaze) recipient. The comprehension of Speech&Gesture relative to SpeechOnly utterances recruited middle occipital, middle temporal and inferior frontal gyri, bilaterally. The calcarine sulcus and posterior cingulate cortex were sensitive to differences between direct and averted gaze. Most importantly, Speech&Gesture utterances, but not SpeechOnly utterances, produced additional activity in the right middle temporal gyrus when participants were addressed. Marking communicative intent with gaze direction modulates the processing of speech–gesture utterances in cerebral areas typically associated with the semantic processing of multi-modal communicative acts.

Abstract

Instrumental music and language are both syntactic systems, employing complex, hierarchically-structured sequences built using implicit structural norms. This organization allows listeners to understand the role of individual words or tones in the context of an unfolding sentence or melody. Previous studies suggest that the brain mechanisms of syntactic processing may be partly shared between music and language. However, functional neuroimaging evidence for anatomical overlap of brain activity involved in linguistic and musical syntactic processing has been lacking. In the present study we used functional magnetic resonance imaging (fMRI) in conjunction with an interference paradigm based on sung sentences. We show that the processing demands of musical syntax (harmony) and language syntax interact in Broca’s area in the left inferior frontal gyrus (without leading to music and language main effects). A language main effect in Broca’s area only emerged in the complex music harmony condition, suggesting that (with our stimuli and tasks) a language effect only becomes visible under conditions of increased demands on shared neural resources. In contrast to previous studies, our design allows us to rule out that the observed neural interaction is due to: (1) general attention mechanisms, as a psychoacoustic auditory anomaly behaved unlike the harmonic manipulation, (2) error processing, as the language and the music stimuli contained no structural errors. The current results thus suggest that two different cognitive domains—music and language—might draw on the same high level syntactic integration resources in Broca’s area.

Abstract

This study investigated the brain regions for the comprehension of implied emotion in sentences. Participants read negative sentences without negative words, for example, “The boy fell asleep and never woke up again,” and their neutral counterparts “The boy stood up and grabbed his bag.” This kind of negative sentence allows us to examine implied emotion derived at the sentence level, without associative emotion coming from word retrieval. We found that implied emotion in sentences, relative to neutral sentences, led to activation in some emotion-related areas, including the medial prefrontal cortex, the amygdala, and the insula, as well as certain language-related areas, including the inferior frontal gyrus, which has been implicated in combinatorial processing. These results suggest that the emotional network involved in implied emotion is intricately related to the network for combinatorial processing in language, supporting the view that sentence meaning is more than simply concatenating the meanings of its lexical building blocks.

Abstract

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

Abstract

A fundamental property of language is that it can be used to refer to entities in the extra-linguistic physical context of a conversation in order to establish a joint focus of attention on a referent. Typological and psycholinguistic work across a wide range of languages has put forward at least two different theoretical views on demonstrative reference. Here we contrasted and tested these two accounts by investigating the electrophysiological brain activity underlying the construction of indexical meaning in comprehension. In two EEG experiments, participants watched pictures of a speaker who referred to one of two objects using speech and an index-finger pointing gesture. In contrast with separately collected native speakers’ linguistic intuitions, N400 effects showed a preference for a proximal demonstrative when speaker and addressee were in a face-to-face orientation and all possible referents were located in the shared space between them, irrespective of the physical proximity of the referent to the speaker. These findings reject egocentric proximity-based accounts of demonstrative reference, support a sociocentric approach to deixis, suggest that interlocutors construe a shared space during conversation, and imply that the psychological proximity of a referent may be more important than its physical proximity.

Abstract

Comprehension of pointing gestures is fundamental to human communication. However, the neural mechanisms
that subserve the integration of pointing gestures and speech in visual contexts in comprehension
are unclear. Here we present the results of an fMRI study in which participants watched images of an
actor pointing at an object while they listened to her referential speech. The use of a mismatch paradigm
revealed that the semantic unication of pointing gesture and speech in a triadic context recruits left
inferior frontal gyrus. Complementing previous ndings, this suggests that left inferior frontal gyrus
semantically integrates information across modalities and semiotic domains.

Abstract

Emotions are often expressed metaphorically, and both emotion and metaphor are ways through which abstract meaning can be grounded in language. Here we investigate specifically whether motion-related verbs when used metaphorically are differentially sensitive to a preceding emotional context, as compared to when they are used in a literal manner. Participants read stories that ended with ambiguous action/motion sentences (e.g., he got it), in which the action/motion could be interpreted metaphorically (he understood the idea) or literally (he caught the ball) depending on the preceding story. Orthogonal to the metaphorical manipulation, the stories were high or low in emotional content. The results showed that emotional context modulated the neural response in visual motion areas to the metaphorical interpretation of the sentences, but not to their literal interpretations. In addition, literal interpretations of the target sentences led to stronger activation in the visual motion areas as compared to metaphorical readings of the sentences. We interpret our results as suggesting that emotional context specifically modulates mental simulation during metaphor processing

Abstract

In this study, we explore the possibility to predict the semantic category of words from brain signals in a free word generation task. Participants produced single words from different semantic categories in a modified semantic fluency task. A Bayesian logistic regression classifier was trained to predict the semantic category of words from single-trial MEG data. Significant classification accuracies were achieved using sensor-level MEG time series at the time interval of conceptual preparation. Semantic category prediction was also possible using source-reconstructed time series, based on minimum norm estimates of cortical activity. Brain regions that contributed most to classification on the source level were identified. These were the left inferior frontal gyrus, left middle frontal gyrus, and left posterior middle temporal gyrus. Additionally, the temporal dynamics of brain activity underlying the semantic preparation during word generation was explored. These results provide important insights about central aspects of language production

Abstract

Diffusion tensor imaging (DTI) and a longitudinal language learning approach were applied to investigate the relationship between the achieved second language (L2) proficiency during L2 learning and the reorganization of structural connectivity between core language areas. Language proficiency tests and DTI scans were obtained from German students before and after they completed an intensive 6-week course of the Dutch language. In the initial learning stage, with increasing L2 proficiency, the hemispheric dominance of the BA6-temporal pathway (mainly along the arcuate fasciculus) shifted from the left to the right hemisphere. With further increased proficiency, however, lateralization dominance was again found in the left BA6-temporal pathway. This result is consistent with reports in the literature that imply a stronger involvement of the right hemisphere in L2-processing especially for less proficient L2-speakers. This is the first time that a L2-proficiency-dependent laterality shift in structural connectivity of language pathways during L2 acquisition has been observed to shift from left to right, and back to left hemisphere dominance with increasing L2-proficiency. We additionally find that changes in fractional anisotropy values after the course are related to the time elapsed between the two scans. The results suggest that structural connectivity in (at least part of) the perisylvian language network may be subject to fast dynamic changes following language learning