Publications

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

Abstract

At least three cognitive brain components are necessary in order for us to be able to produce and comprehend language: a Memory repository for the lexicon, a Unification buffer where lexical information is combined into novel structures, and a Control apparatus presiding over executive function in language. Here we describe the brain networks that support Memory and Unification in semantics. A dynamic account of their interactions is presented, in which a balance between the two components is sought at each word-processing step. We use the theory to provide an explanation of the N400 effect.

Abstract

Many children with Specific Language Impairment (SLI) show impairments in discriminating auditorily presented stimuli. The present study investigates whether these discrimination problems are speech specific or of a general auditory nature. This was studied by using a linguistic and nonlinguistic contrast that were matched for acoustic complexity in an active behavioral task and a passive ERP paradigm, known to elicit the mismatch negativity (MMN). In addition, attention skills and a variety of language skills were measured. Participants were 25 five-year-old Dutch children with SLI having receptive as well as productive language problems and 25 control children with typical speechand language development. At the behavioral level, the SLI group was impaired in discriminating the linguistic contrast as compared to the control group, while both groups were unable to distinguish the non-linguistic contrast. Moreover, the SLI group tended to have impaired attention skills which correlated with performance on most of the language tests. At the neural level, the SLI group, in contrast to the control group, did not show an MMN in response to either the linguistic or nonlinguistic contrast. The MMN data are consistent with an account that relates the symptoms in children with SLI to non-speech processing difficulties.

Abstract

The first objective of this study was to compare the brain network engaged by preference classification and the standard grammaticality classification after implicit artificial syntax acquisition by re-analyzing previously reported event-related fMRI data. The results show that preference and grammaticality classification engage virtually identical brain networks, including Broca’s region, consistent with previous behavioral findings. Moreover, the results showed that the effects related to artificial syntax in Broca’s region were essentially the same when masked with variability related to natural syntax processing in the same participants. The second objective was to explore CNTNAP2-related effects in implicit artificial syntax learning by analyzing behavioral and event-related fMRI data from a subsample. The CNTNAP2 gene has been linked to specific language impairment and is controlled by the FOXP2 transcription factor. CNTNAP2 is expressed in language related brain networks in the developing human brain and the FOXP2–CNTNAP2 pathway provides a mechanistic link between clinically distinct syndromes involving disrupted language. Finally, we discuss the implication of taking natural language to be a neurobiological system in terms of bounded recursion and suggest that the left inferior frontal region is a generic on-line sequence processor that unifies information from various sources in an incremental and recursive manner.

Abstract

During face-to-face communication, one does not only hear speech but also see a speaker's communicative hand movements. It has been shown that such hand gestures play an important role in communication where the two modalities influence each other's interpretation. A gesture typically temporally overlaps with coexpressive speech, but the gesture is often initiated before (but not after) the coexpressive speech. The present ERP study investigated what degree of asynchrony in the speech and gesture onsets are optimal for semantic integration of the concurrent gesture and speech. Videos of a person gesturing were combined with speech segments that were either semantically congruent or incongruent with the gesture. Although gesture and speech always overlapped in time, gesture and speech were presented with three different degrees of asynchrony. In the SOA 0 condition, the gesture onset and the speech onset were simultaneous. In the SOA 160 and 360 conditions, speech was delayed by 160 and 360 msec, respectively. ERPs time locked to speech onset showed a significant difference between semantically congruent versus incongruent gesture–speech combinations on the N400 for the SOA 0 and 160 conditions. No significant difference was found for the SOA 360 condition. These results imply that speech and gesture are integrated most efficiently when the differences in onsets do not exceed a certain time span because of the fact that iconic gestures need speech to be disambiguated in a way relevant to the speech context.

Abstract

Whether the brain’s speech-production system is also involved in speech comprehension is a topic of much debate. Research has focused on whether motor areas are involved in listening, but overlap between speaking and listening might occur not only at primary sensory and motor levels, but also at linguistic levels (where semantic, lexical, and syntactic processes occur). Using functional MRI adaptation during speech comprehension and production, we found that the brain areas involved in semantic, lexical, and syntactic processing are mostly the same for speaking and for listening. Effects of primary processing load (indicative of sensory and motor processes) overlapped in auditory cortex and left inferior frontal cortex, but not in motor cortex, where processing load affected activity only in speaking. These results indicate that the linguistic parts of the language system are used for both speaking and listening, but that the motor system does not seem to provide a crucial contribution to listening.

Abstract

Defeasible inferences are inferences that can be revised in the light of new information. Although defeasible inferences are pervasive in everyday communication, little is known about how and when they are processed by the brain. This study examined the electrophysiological signature of defeasible reasoning using a modified version of the suppression task. Participants were presented with conditional inferences (of the type “if p, then q; p, therefore q”) that were preceded by a congruent or a disabling context. The disabling context contained a possible exception or precondition that prevented people from drawing the conclusion. Acceptability of the conclusion was indeed lower in the disabling condition compared to the congruent condition. Further, we found a large sustained negativity at the conclusion of the disabling condition relative to the congruent condition, which started around 250 msec and was persistent throughout the entire epoch. Possible accounts for the observed effect are discussed.

Abstract

Work on animals indicates that BOLD is preferentially sensitive to local field potentials, and that it correlates most strongly with gamma band neuronal synchronization. Here we investigate how the BOLD signal in humans performing a cognitive task is related to neuronal synchronization across different frequency bands. We simultaneously recorded EEG and BOLD while subjects engaged in a visual attention task known to induce sustained changes in neuronal synchronization across a wide range of frequencies. Trial-by-trial BOLD luctuations correlated positively with trial-by-trial fluctuations in high-EEG gamma power (60–80 Hz) and negatively with alpha and beta power. Gamma power on the one hand, and alpha and beta power on the other hand, independently contributed to explaining BOLD variance. These results indicate that the BOLD-gamma coupling observed in animals can be extrapolated to humans performing a task and that neuronal dynamics underlying high- and low-frequency synchronization contribute independently to the BOLD signal.

Abstract

Speakers tend to repeat syntactic structures across sentences, a phenomenon called syntactic priming. Although it has been suggested that repeating syntactic structures should result in speeded responses, previous research has focused on effects in response tendencies. We investigated syntactic priming effects simultaneously in response tendencies and response latencies for active and passive transitive sentences in a picture description task. In Experiment 1, there were priming effects in response tendencies for passives and in response latencies for actives. However, when participants' pre-existing preference for actives was altered in Experiment 2, syntactic priming occurred for both actives and passives in response tendencies as well as in response latencies. This is the first investigation of the effects of structure frequency on both response tendencies and latencies in syntactic priming. We discuss the implications of these data for current theories of syntactic processing.

Abstract

In individuals with ASD, difficulties with language comprehension are most evident when higher-level semantic-pragmatic language processing is required, for instance when context has to be used to interpret the meaning of an utterance. Until now, it is unclear at what level of processing and for what type of context these difficulties in language comprehension occur. Therefore, in the current fMRI study, we investigated the neural correlates of the integration of contextual information during auditory language comprehension in 24 adults with ASD and 24 matched control participants. Different levels of context processing were manipulated by using spoken sentences that were correct or contained either a semantic or world knowledge anomaly. Our findings demonstrated significant differences between the groups in inferior frontal cortex that were only present for sentences with a world knowledge anomaly. Relative to the ASD group, the control group showed significantly increased activation in left inferior frontal gyrus (LIFG) for sentences with a world knowledge anomaly compared to correct sentences. This effect possibly indicates reduced integrative capacities of the ASD group. Furthermore, world knowledge anomalies elicited significantly stronger activation in right inferior frontal gyrus (RIFG) in the control group compared to the ASD group. This additional RIFG activation probably reflects revision of the situation model after new, conflicting information. The lack of recruitment of RIFG is possibly related to difficulties with exception handling in the ASD group.

Abstract

Synesthesia provides an elegant model to investigate neural mechanisms underlying individual differences in subjective experience in humans. In grapheme–color synesthesia, written letters induce color sensations, accompanied by activation of color area V4. Competing hypotheses suggest that enhanced V4 activity during synesthesia is either induced by direct bottom-up cross-activation from grapheme processing areas within the fusiform gyrus, or indirectly via higher-order parietal areas. Synesthetes differ in the way synesthetic color is perceived: “projector” synesthetes experience color externally colocalized with a presented grapheme, whereas “associators” report an internally evoked association. Using dynamic causal modeling for fMRI, we show that V4 cross-activation during synesthesia was induced via a bottom-up pathway (within fusiform gyrus) in projector synesthetes, but via a top-down pathway (via parietal lobe) in associators. These findings show how altered coupling within the same network of active regions leads to differences in subjective experience. Our findings reconcile the two most influential cross-activation accounts of synesthesia.

Abstract

To highlight relevant information in dialogues, both wh-question context and pitch accent in answers can be used, such that focused information gains more attention and is processed more elaborately. To evaluate the relative influence of context and pitch accent on the depth of semantic processing, we measured Event-Related Potentials (ERPs) to auditorily presented wh-question-answer pairs. A semantically incongruent word in the answer occurred either in focus or non-focus position as determined by the context, and this word was either accented or unaccented. Semantic incongruency elicited different N400 effects in different conditions. The largest N400 effect was found when the question-marked focus was accented, while the other three conditions elicited smaller N400 effects. The results suggest that context and accentuation interact. Thus accented focused words were processed more deeply compared to conditions where focus and accentuation mismatched, or when the new information had no marking. In addition, there seems to be sex differences in the depth of semantic processing. For males, a significant N400 effect was observed only when the question-marked focus was accented, reduced N400 effects were found in the other dialogues. In contrast, females produced similar N400 effects in all the conditions. These results suggest that regardless of external cues, females tend to engage in more elaborate semantic processing compared to males.

Abstract

We investigated how visual and linguistic information interact in the perception of emotion. We borrowed a phenomenon from film theory which states that presentation of an as such neutral visual scene intensifies the percept of fear or suspense induced by a different channel of information, such as language. Our main aim was to investigate how neutral visual scenes can enhance responses to fearful language content in parts of the brain involved in the perception of emotion. Healthy participants’ brain activity was measured (using functional magnetic resonance imaging) while they read fearful and less fearful sentences presented with or without a neutral visual scene. The main idea is that the visual scenes intensify the fearful content of the language by subtly implying and concretizing what is described in the sentence. Activation levels in the right anterior temporal pole were selectively increased when a neutral visual scene was paired with a fearful sentence, compared to reading the sentence alone, as well as to reading of non-fearful sentences presented with the same neutral scene. We conclude that the right anterior temporal pole serves a binding function of emotional information across domains such as visual and linguistic information.

Abstract

A debated issue in the relationship between language and thought is how our linguistic abilities are involved in understanding the intentions of others (‘mentalizing’). The results of both theoretical and empirical work have been used to argue that linguistic, and more specifically, grammatical, abilities are crucial in representing the mental states of others. Here we contribute to this debate by investigating how damage to the language system influences the generation and understanding of intentional communicative behaviors. Four patients with pervasive language difficulties (severe global or agrammatic aphasia) engaged in an experimentally controlled non-verbal communication paradigm, which required signaling and understanding a communicative message. Despite their profound language problems they were able to engage in recipient design as well as intention recognition, showing similar indicators of mentalizing as have been observed in the neurologically healthy population. Our results show that aspects of the ability to communicate remain present even when core capacities of the language system are dysfunctional