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

An important, but often ignored distinction in the analysis of EEG signals is that between evoked activity and induced activity. Whereas evoked activity reflects the summation of transient post-synaptic potentials triggered by an event, induced activity, which is mainly oscillatory in nature, is thought to reflect changes in parameters controlling dynamic interactions within and between brain structures. We hypothesize that induced activity may yield information about the dynamics of cell assembly formation, activation and subsequent uncoupling, which may play a prominent role in different types of memory operations. We then describe a number of analysis tools that can be used to study the reactivity of induced rhythmic activity, both in terms of amplitude changes and of phase variability.

We briefly discuss how alpha, gamma and theta rhythms are thought to be generated, paying special attention to the hypothesis that the theta rhythm reflects dynamic interactions between the hippocampal system and the neocortex. This hypothesis would imply that studying the reactivity of scalp-recorded theta may provide a window on the contribution of the hippocampus to memory functions.

We review studies investigating the reactivity of scalp-recorded theta in paradigms engaging episodic memory, spatial memory and working memory. In addition, we review studies that relate theta reactivity to processes at the interface of memory and language. Despite many unknowns, the experimental evidence largely supports the hypothesis that theta activity plays a functional role in cell assembly formation, a process which may constitute the neural basis of memory formation and retrieval. The available data provide only highly indirect support for the hypothesis that scalp-recorded theta yields information about hippocampal functioning. It is concluded that studying induced rhythmic activity holds promise as an additional important way to study brain function.

Abstract

In two studies subjects were required to read Dutch sentences that in some cases contained a syntactic violation, in other cases a semantic violation. All syntactic violations were word category violations. The design excluded differential contributions of expectancy to influence the syntactic violation effects. The syntactic violations elicited an Anterior Negativity between 300 and 500 ms. This negativity was bilateral and had a frontal distribution. Over posterior sites the same violations elicited a P600/SPS starting at about 600 ms. The semantic violations elicited an N400 effect. The topographic distribution of the AN was more frontal than the distribution of the classical N400 effect, indicating that the underlying generators of the AN and the N400 are, at least to a certain extent, non-overlapping. Experiment 2 partly replicated the design of Experiment 1, but with differences in rate of presentation and in the distribution of items over subjects, and without semantic violations. The word category violations resulted in the same effects as were observed in Experiment 1, showing that they were independent of some of the specific parameters of Experiment 1. The discussion presents a tentative account of the functional differences in the triggering conditions of the AN and the P600/SPS.

Abstract

To understand spoken language requires that the brain provides rapid access to different kinds of knowledge, including the sounds and meanings of words, and syntax. Syntax specifies constraints on combining words in a grammatically well formed manner. Agrammatic patients are deficient in their ability to use these constraints, due to a lesion in the perisylvian area of the languagedominant hemisphere. We report a study on real-time auditory sentence processing in agrammatic comprehenders, examining
their ability to accommodate damage to the language system. We recorded event-related brain potentials (ERPs) in agrammatic comprehenders, nonagrammatic aphasics, and age-matched controls. When listening to sentences with grammatical violations, the agrammatic aphasics did not show the same syntax-related ERP effect as the two other subject groups. Instead, the waveforms of the agrammatic aphasics were dominated by a meaning-related ERP effect, presumably reflecting their attempts to achieve understanding by the use of semantic constraints. These data demonstrate that although agrammatic aphasics are impaired in their ability to exploit syntactic information in real time, they can reduce the consequences of a syntactic deficit by exploiting a semantic route. They thus provide evidence for the compensation of a syntactic deficit by a stronger reliance on another route in mapping
sound onto meaning. This is a form of plasticity that we refer to as multiple-route plasticity.

Abstract

Syntax is one of the components in the architecture of language processing that allows the listener/reader to bind single-word information into a unified interpretation of multiword utterances. This paper discusses ERP effects that have been observed in relation to syntactic processing. The fact that these effects differ from the semantic N400 indicates that the brain honors the distinction between semantic and syntactic binding operations. Two models of syntactic processing attempt to account for syntax-related ERP effects. One type of model is serial, with a first phase that is purely syntactic in nature (syntax-first model). The other type of model is parallel and assumes that information immediately guides the interpretation process once it becomes available. This is referred to as the immediacy model. ERP evidence is presented in support of the latter model. Next, an explicit computational model is proposed to explain the ERP data. This Unification Model assumes that syntactic frames are stored in memory and retrieved on the basis of the spoken or written word form input. The syntactic frames associated with the individual lexical items are unified by a dynamic binding process into a structural representation that spans the whole utterance. On the basis of a meta-analysis of imaging studies on syntax, it is argued that the left posterior inferior frontal cortex is involved in binding syntactic frames together, whereas the left superior temporal cortex is involved in retrieval of the syntactic frames stored in memory. Lesion data that support the involvement of this left frontotemporal network in syntactic processing are discussed.

Abstract

This study investigated the effects of combined semantic and syntactic violations in relation to the effects of single semantic and single syntactic violations on language-related event-related brain potential (ERP) effects (N400 and P600/ SPS). Syntactic violations consisted of a mismatch in grammatical gender or number features of the definite article and the noun in sentence-internal or sentence-final noun phrases (NPs). Semantic violations consisted of semantically implausible adjective–noun combinations in the same NPs. Combined syntactic and semantic violations were a summation of these two respective violation types. ERPs were recorded while subjects read the sentences with the different types of violations and the correct control sentences. ERP effects were computed relative to ERPs elicited by the sentence-internal or sentence-final nouns. The size of the N400 effect to the semantic violation was increased by an additional syntactic violation (the syntactic boost). In contrast, the size of the P600/ SPS to the syntactic violation was not affected by an additional semantic violation. This suggests that in the absence of syntactic ambiguity, the assignment of syntactic structure is independent of semantic context. However, semantic integration is influenced by syntactic processing. In the sentence-final position, additional global processing consequences were obtained as a result of earlier violations in the sentence. The resulting increase in the N400 amplitude to sentence-final words was independent of the nature of the violation. A speeded anomaly detection task revealed that it takes substantially longer to detect semantic than syntactic anomalies. These results are discussed in relation to the latency and processing characteristics of the N400 and P600/SPS effects. Overall, the results reveal an asymmetry in the interplay between syntax and semantics during on-line sentence comprehension.

Abstract

Mental imagery is a cognitive process crucial to human reasoning. Numerous studies have characterized specific
instances of this cognitive ability, as evoked by visual imagery (VI) or motor imagery (MI) tasks. However, it
remains unclear which neural resources are shared between VI and MI, and which are exclusively related to MI.
To address this issue, we have used fMRI to measure human brain activity during performance of VI and MI
tasks. Crucially, we have modulated the imagery process by manipulating the degree of mental rotation necessary
to solve the tasks. We focused our analysis on changes in neural signal as a function of the degree of mental
rotation in each task.

Abstract

Spoken language comprehension requires rapid integration of information from multiple linguistic sources. In the present study we addressed the temporal aspects of this integration process by focusing on the time course of the selection of the appropriate meaning of lexical ambiguities (“bank”) in sentence contexts. Successful selection of the contextually appropriate meaning of the ambiguous word is dependent upon the rapid binding of the contextual information in the sentence to the appropriate meaning of the ambiguity. We used the N400 to identify the time course of this binding process. The N400 was measured to target words that followed three types of context sentences. In the concordant context, the sentence biased the meaning of the sentence-final ambiguous word so that it was related to the target. In the discordant context, the sentence context biased the meaning so that it was not related to the target. In the unrelated control condition, the sentences ended in an unambiguous noun that was unrelated to the target. Half of the concordant sentences biased the dominant meaning, and the other half biased the subordinate meaning of the sentence-final ambiguous words. The ISI between onset of the target word and offset of the sentence-final word of the context sentence was 100 ms in one version of the experiment, and 1250 ms in the second version. We found that (i) the lexically dominant meaning is always partly activated, independent of context, (ii) initially both dominant and subordinate meaning are (partly) activated, which suggests that contextual and lexical factors both contribute to sentence interpretation without context completely overriding lexical information, and (iii) strong lexical influences remain present for a relatively long period of time.

Abstract

In two ERP experiments, we assessed the impact of discourse-level information on the processing of an unfolding spoken sentence. Subjects listened to sentences like Jane told her brother that he was exceptionally quick/slow, designed such that the alternative critical words were equally acceptable within the local sentence context. In Experiment 1, these sentences were embedded in a discourse that rendered one of the critical words anomalous (e.g. because Jane’s brother had in fact done something very quickly). Relative to the coherent alternative, these discourse-anomalous words elicited a standard N400 effect that started at 150–200 ms after acoustic word onset. Furthermore, when the same sentences were heard in isolation in Experiment 2, the N400 effect disappeared. The results demonstrate that our listeners related the unfolding spoken words to the wider discourse extremely rapidly, after having heard the first two or three phonemes only, and in many cases well before the end of the word. In addition, the identical nature of discourse- and sentence-dependent N400 effects suggests that from the perspective of the word-elicited comprehension process indexed by the N400, the interpretive context delineated by a single unfolding sentence and a larger discourse is functionally identical.

Abstract

In two experiments, we explored the use of event-related brain potentials to selectively track the processes that establish reference during spoken language comprehension. Subjects listened to stories in which a particular noun phrase like "the girl" either uniquely referred to a single referent mentioned in the earlier discourse, or ambiguously referred to two equally suitable referents. Referentially ambiguous nouns ("the girl" with two girls introduced in the discourse context) elicited a frontally dominant and sustained negative shift in brain potentials, emerging within 300–400 ms after acoustic noun onset. The early onset of this effect reveals that reference to a discourse entity can be established very rapidly. Its morphology and distribution suggest that at least some of the processing consequences of referential ambiguity may involve an increased demand on memory resources. Furthermore, because this referentially induced ERP effect is very different from that of well-known ERP effects associated with the semantic (N400) and syntactic (e.g., P600/SPS) aspects of language comprehension, it suggests that ERPs can be used to selectively keep track of three major processes involved in the comprehension of an unfolding piece of discourse.