Publications

Abstract

Studies of perception have long shown that the brain adds information to its sensory analysis of the physical environment. A touchstone example for humans is language use: to comprehend a physical signal like speech, the brain must add linguistic knowledge, including syntax. Yet, syntactic rules and representations are widely assumed to be atemporal (i.e., abstract and not bound by time), so they must be translated into time-varying signals for speech comprehension and production. Here, we test 3 different models of the temporal spell-out of syntactic structure against brain activity of people listening to Dutch stories: an integratory bottom-up parser, a predictive top-down parser, and a mildly predictive left-corner parser. These models build exactly the same structure but differ in when syntactic information is added by the brain—this difference is captured in the (temporal distribution of the) complexity metric “incremental node count.” Using temporal response function models with both acoustic and information-theoretic control predictors, node counts were regressed against source-reconstructed delta-band activity acquired with magnetoencephalography. Neural dynamics in left frontal and temporal regions most strongly reflect node counts derived by the top-down method, which postulates syntax early in time, suggesting that predictive structure building is an important component of Dutch sentence comprehension. The absence of strong effects of the left-corner model further suggests that its mildly predictive strategy does not represent Dutch language comprehension well, in contrast to what has been found for English. Understanding when the brain projects its knowledge of syntax onto speech, and whether this is done in language-specific ways, will inform and constrain the development of mechanistic models of syntactic structure building in the brain.

Abstract

Face-to-face communication is not only about ‘what’ is said but also ‘how’ it is said, both in speech and bodily signals. Beat gestures are rhythmic hand movements that typically accompany prosodic prominence in con-versation. Yet, it is still unclear how beat gestures influence language comprehension. On the one hand, beat gestures may share the same functional role of focus markers as prosodic prominence. Accordingly, they would drive attention towards the concurrent speech and highlight its content. On the other hand, beat gestures may trigger inferences of high speaker confidence, generate the expectation that the sentence content is correct and thereby elicit the commitment to the truth of the statement. This study directly disentangled the two hypotheses by evaluating additive and interactive effects of prosodic prominence and beat gestures on language comprehension. Participants watched videos of a speaker uttering sentences and judged whether each sentence was true or false. Sentences sometimes contained a world knowledge violation that may go unnoticed (‘semantic illusion’). Combining beat gestures with prosodic prominence led to a higher degree of semantic illusion, making more world knowledge violations go unnoticed during language comprehension. These results challenge current theories proposing that beat gestures are visual focus markers. To the contrary, they suggest that beat gestures automatically trigger inferences of high speaker confidence and thereby elicit the commitment to the truth of the statement, in line with Grice’s cooperative principle in conversation. More broadly, our findings also highlight the influence of metacognition on language comprehension in face-to-face ommunication.

Abstract

The increased interest in developing next-gen social robots has raised questions about the factors affecting the perception of robot emotions. This study investigates the impact of robot appearances (human-like, mechanical) and face regions (full-face, eye-region) on human perception of robot emotions. A between-subjects user study (N = 305) was conducted where participants were asked to identify the emotions being displayed in videos of robot faces, as well as a human baseline. Our findings reveal three important insights for effective social robot face design in Human-Robot Interaction (HRI): Firstly, robots equipped with a back-projected, fully animated face – regardless of whether they are more human-like or more mechanical-looking – demonstrate a capacity for emotional expression comparable to that of humans. Secondly, the recognition accuracy of emotional expressions in both humans and robots declines when only the eye region is visible. Lastly, within the constraint of only the eye region being visible, robots with more human-like features significantly enhance emotion recognition.

Abstract

To date, research has reliably shown that language can engage and modify perceptual processes in a top-down manner. However, our understanding of the cognitive and neural mechanisms underlying such top-down influences is still under debate. In this review, we provide an overview of findings from literature investigating the organization of semantic networks in the brain (spontaneous engagement of the visual system while processing linguistic information), and linguistic cueing studies (looking at the immediate effects of language on the perception of a visual target), in an effort to isolate such mechanisms. Additionally, we connect the findings from linguistic cueing studies to those reported in (nonlinguistic) literature on priors in perception, in order to find commonalities in neural processes allowing for top-down influences on perception. In doing so, we discuss the effects of language on perception in the context of broader, general cognitive and neural principles. Finally, we propose a way forward in the study of linguistic influences on perception.

Abstract

Prosody underpins various linguistic domains ranging from semantics and syntax to discourse. For instance, prosodic information in the form of lexical stress modifies meanings and, as such, syntactic contexts of words as in Turkish kaz-má "pickaxe" (noun) versus káz-ma "do not dig" (imperative). Likewise, prosody indicates the focused constituent of an utterance as the noun phrase filling the wh-spot in a dialogue like What did you eat? I ate----. In the present study, we investigated the relevance of such prosodic variations for discourse comprehension in Turkish. We aimed at answering how lexical stress and prosodic focus mismatches on critical noun phrases-resulting in grammatical anomalies involving both semantics and syntax and discourse-level anomalies, respectively-affect the perceived correctness of an answer to a question in a given context. To that end, 80 native speakers of Turkish, 40 participating in a psychometric experiment and 40 participating in an EEG experiment, were asked to judge the acceptability of prosodic mismatches that occur either separately or concurrently. Psychometric results indicated that lexical stress mismatch led to a lower correctness score than prosodic focus mismatch, and combined mismatch received the lowest score. Consistent with the psychometric data, EEG results revealed an N400 effect to combined mismatch, and this effect was followed by a P600 response to lexical stress mismatch. Conjointly, these results suggest that every source of prosodic information is immediately available and codetermines the interpretation of an utterance; however, semantically and syntactically relevant lexical stress information is assigned more significance by the language comprehension system compared with prosodic focus information.

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

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.