Publications

Abstract

According to the language marker hypothesis language has provided homo sapiens with a rich symbolic system that plays a central role in interpreting signals delivered by our sensory apparatus, in shaping action goals, and in creating a powerful tool for reasoning and inferencing. This view provides an important correction on embodied accounts of language that reduce language to action, perception, emotion and mental simulation. The presence of a language system has, however, also important consequences for perception, action, emotion, and memory. Language stamps signals from perception, action, and emotional systems with rich cognitive markers that transform the role of these signals in the overall cognitive architecture of the human mind. This view does not deny that language is implemented by means of universal principles of neural organization. However, language creates the possibility to generate rich internal models of the world that are shaped and made accessible by the characteristics of a language system. This makes us less dependent on direct action-perception couplings and might even sometimes go at the expense of the veridicality of perception. In cognitive (neuro)science the pendulum has swung from language as the key to understand the organization of the human mind to the perspective that it is a byproduct of perception and action. It is time that it partly swings back again.

Abstract

EEG and eye-tracking provide complementary information when investigating language comprehension. Evidence that speech processing may be facilitated by speech prediction comes from the observation that a listener's eye gaze moves towards a referent before it is mentioned if the remainder of the spoken sentence is predictable. However, changes to the trajectory of anticipatory fixations could result from a change in prediction or an attention shift. Conversely, N400 amplitudes and concurrent spectral power provide information about the ease of word processing the moment the word is perceived. In a proof-of-principle investigation, we combined EEG and eye-tracking to study linguistic prediction in naturalistic, virtual environments. We observed increased processing, reflected in theta band power, either during verb processing - when the verb was predictive of the noun - or during noun processing - when the verb was not predictive of the noun. Alpha power was higher in response to the predictive verb and unpredictable nouns. We replicated typical effects of noun congruence but not predictability on the N400 in response to the noun. Thus, the rich visual context that accompanied speech in virtual reality influenced language processing compared to previous reports, where the visual context may have facilitated processing of unpredictable nouns. Finally, anticipatory fixations were predictive of spectral power during noun processing and the length of time fixating the target could be predicted by spectral power at verb onset, conditional on the object having been fixated. Overall, we show that combining EEG and eye-tracking provides a promising new method to answer novel research questions about the prediction of upcoming linguistic input, for example, regarding the role of extralinguistic cues in prediction during language comprehension.

Abstract

During listening, brain activity tracks the rhythmic structures of speech signals. Here, we directly dissociated the contribution of neural envelope tracking in the processing of speech acoustic cues from that related to linguistic processing. We examined the neural changes associated with the comprehension of Noise-Vocoded (NV) speech using magnetoencephalography (MEG). Participants listened to NV sentences in a 3-phase training paradigm: (1) pre-training, where NV stimuli were barely comprehended, (2) training with exposure of the original clear version of speech stimulus, and (3) post-training, where the same stimuli gained intelligibility from the training phase. Using this paradigm, we tested if the neural responses of a speech signal was modulated by its intelligibility without any change in its acoustic structure. To test the influence of spectral degradation on neural envelope tracking independently of training, participants listened to two types of NV sentences (4-band and 2-band NV speech), but were only trained to understand 4-band NV speech. Significant changes in neural tracking were observed in the delta range in relation to the acoustic degradation of speech. However, we failed to find a direct effect of intelligibility on the neural tracking of speech envelope in both theta and delta ranges, in both auditory regions-of-interest and whole-brain sensor-space analyses. This suggests that acoustics greatly influence the neural tracking response to speech envelope, and that caution needs to be taken when choosing the control signals for speech-brain tracking analyses, considering that a slight change in acoustic parameters can have strong effects on the neural tracking response.

Abstract

Affective behaviors enable social robots to not only establish better connections with humans but also serve as a tool for the robots to express their internal states. It has been well established that emotions are important to signal understanding in Human-Robot Interaction (HRI). This work aims to harness the power of Large Language Models (LLM) and proposes an approach to control the affective behavior of robots. By interpreting emotion appraisal as an Emotion Recognition in Conversation (ERC) tasks, we used GPT-3.5 to predict the emotion of a robot’s turn in real-time, using the dialogue history of the ongoing conversation. The robot signaled the predicted emotion using facial expressions. The model was evaluated in a within-subjects user study (N = 47) where the model-driven emotion generation was compared against conditions where the robot did not display any emotions and where it displayed incongruent emotions. The participants interacted with the robot by playing a card sorting game that was specifically designed to evoke emotions. The results indicated that the emotions were reliably generated by the LLM and the participants were able to perceive the robot’s emotions. It was found that the robot expressing congruent model-driven facial emotion expressions were perceived to be significantly more human-like, emotionally appropriate, and elicit a more positive impression. Participants also scored significantly better in the card sorting game when the robot displayed congruent facial expressions. From a technical perspective, the study shows that LLMs can be used to control the affective behavior of robots reliably in real-time. Additionally, our results could be used in devising novel human-robot interactions, making robots more effective in roles where emotional interaction is important, such as therapy, companionship, or customer service.

Abstract

Neuron models with explicit dendritic dynamics have shed light on mechanisms for coincidence detection, pathway selection and temporal filtering. However, it is still unclear which morphological and physiological features are required to capture these phenomena. In this work, we introduce the Tripod neuron model and propose a minimal structural reduction of the dendritic tree that is able to reproduce these computations. The Tripod is a three-compartment model consisting of two segregated passive dendrites and a somatic compartment modelled as an adaptive, exponential integrate-and-fire neuron. It incorporates dendritic geometry, membrane physiology and receptor dynamics as measured in human pyramidal cells. We characterize the response of the Tripod to glutamatergic and GABAergic inputs and identify parameters that support supra-linear integration, coincidence-detection and pathway-specific gating through shunting inhibition. Following NMDA spikes, the Tripod neuron generates plateau potentials whose duration depends on the dendritic length and the strength of synaptic input. When fitted with distal compartments, the Tripod encodes previous activity into a dendritic depolarized state. This dendritic memory allows the neuron to perform temporal binding, and we show that it solves transition and sequence detection tasks on which a single-compartment model fails. Thus, the Tripod can account for dendritic computations previously explained only with more detailed neuron models or neural networks. Due to its simplicity, the Tripod neuron can be used efficiently in simulations of larger cortical circuits.

Abstract

The meaning of a sentence can be understood, whether presented in written or spoken form. Therefore it is highly probable that brain processes supporting language comprehension are at least partly independent of sensory modality. To identify where and when in the brain language processing is independent of sensory modality, we directly compared neuromagnetic brain signals of 200 human subjects (102 males) either reading or listening to sentences. We used multiset canonical correlation analysis to align individual subject data in a way that boosts those aspects of the signal that are common to all, allowing us to capture word-by-word signal variations, consistent across subjects and at a fine temporal scale. Quantifying this consistency in activation across both reading and listening tasks revealed a mostly left hemispheric cortical network. Areas showing consistent activity patterns include not only areas previously implicated in higher-level language processing, such as left prefrontal, superior & middle temporal areas and anterior temporal lobe, but also parts of the control-network as well as subcentral and more posterior temporal-parietal areas. Activity in this supramodal sentence processing network starts in temporal areas and rapidly spreads to the other regions involved. The findings do not only indicate the involvement of a large network of brain areas in supramodal language processing, but also indicate that the linguistic information contained in the unfolding sentences modulates brain activity in a word-specific manner across subjects.

Abstract

Bodily mimicry often makes the mimickee have more positive feelings about the mimicker. Yet, little is known about the causes of mimicry’s social effects. When people mimic each other’s bodily movements face to face, they can either adopt a mirrorwise perspective (moving in the same absolute direction) or an anatomical perspective (moving in the same direction relative to their own bodies). Mirrorwise mimicry maximizes visuo-spatial similarity between the mimicker and mimickee, whereas anatomical mimicry maximizes the similarity in the states of their motor systems. To compare the social consequences of visuo-spatial and motoric similarity, we asked participants to converse with an embodied virtual agent (VIRTUO), who mimicked their head movements either mirrorwise, anatomically, or not at all. Compared to participants who were not mimicked, those who were mimicked mirrorwise tended to rate VIRTUO more positively, but those who were mimicked anatomically rated him more negatively. During face-to-face conversation, mirrorwise and anatomical mimicry have opposite social consequences. Results suggest that visuo-spatial similarity between mimicker and mimickee, not similarity in motor system activity, gives rise to the positive social effects of bodily mimicry.

Abstract

Language processing involves the ability to store and integrate pieces of
information in working memory over short periods of time. According to
the dominant view, information is maintained through sustained, elevated
neural activity. Other work has argued that short-term synaptic facilitation
can serve as a substrate of memory. Here, we propose an account where
memory is supported by intrinsic plasticity that downregulates neuronal
firing rates. Single neuron responses are dependent on experience and we
show through simulations that these adaptive changes in excitability pro-
vide memory on timescales ranging from milliseconds to seconds. On this
account, spiking activity writes information into coupled dynamic variables
that control adaptation and move at slower timescales than the membrane
potential. From these variables, information is continuously read back into
the active membrane state for processing. This neuronal memory mech-
anism does not rely on persistent activity, excitatory feedback, or synap-
tic plasticity for storage. Instead, information is maintained in adaptive
conductances that reduce firing rates and can be accessed directly with-
out cued retrieval. Memory span is systematically related to both the time
constant of adaptation and baseline levels of neuronal excitability. Inter-
ference effects within memory arise when adaptation is long-lasting. We
demonstrate that this mechanism is sensitive to context and serial order
which makes it suitable for temporal integration in sequence processing
within the language domain. We also show that it enables the binding of
linguistic features over time within dynamic memory registers. This work
provides a step towards a computational neurobiology of language.

Abstract

Speakers and listeners usually interact in larger discourses than single words or even single sentences. The goal of the present study was to identify the neural bases reflecting how the mental representation of the situation denoted in a multi-sentence discourse (situation model) is constructed and shared between speakers and listeners. An fMRI study using a variant of the ambiguous text paradigm was designed. Speakers (n=15) produced ambiguous texts in the scanner and listeners (n=27) subsequently listened to these texts in different states of ambiguity: preceded by a highly informative, intermediately informative or no title at all. Conventional BOLD activation analyses in listeners, as well as inter-subject correlation analyses between the speakers’ and the listeners’ hemodynamic time courses were performed. Critically, only the processing of disambiguated, coherent discourse with an intelligible situation model representation involved (shared) activation in bilateral lateral parietal and medial prefrontal regions. This shared spatiotemporal pattern of brain activation between the speaker and the listener suggests that the process of memory retrieval in medial prefrontal regions and the binding of retrieved information in the lateral parietal cortex constitutes a core mechanism underlying the communication of complex conceptual representations.

Abstract

Visual context facilitates perception, but how this is neurally implemented remains unclear. One example of contextual facilitation is found in reading, where letters are more easily identified when embedded in a word. Bottom-up models explain this word advantage as a post-perceptual decision bias, while top-down models propose that word contexts enhance perception itself. Here, we arbitrate between these accounts by presenting words and nonwords and probing the representational fidelity of individual letters using functional magnetic resonance imaging. In line with top-down models, we find that word contexts enhance letter representations in early visual cortex. Moreover, we observe increased coupling between letter information in visual cortex and brain activity in key areas of the reading network, suggesting these areas may be the source of the enhancement. Our results provide evidence for top-down representational enhancement in word recognition, demonstrating that word contexts can modulate perceptual processing already at the earliest visual regions.

Abstract

Recent neuroimaging evidence suggests that the frequency of entrained oscillations in auditory cortices influences the perceived duration of speech segments, impacting word perception (Kösem et al. 2018). We further tested the causal influence of neural entrainment frequency during speech processing, by manipulating entrainment with continuous transcranial alternating
current stimulation (tACS) at distinct oscillatory frequencies (3 Hz and 5.5 Hz) above the auditory cortices. Dutch participants listened to speech and were asked to report their percept of a target Dutch word, which contained a vowel with an ambiguous duration. Target words
were presented either in isolation (first experiment) or at the end of spoken sentences (second experiment). We predicted that the tACS frequency would influence neural entrainment and
therewith how speech is perceptually sampled, leading to a perceptual over- or underestimation of the vowel’s duration. Whereas results from Experiment 1 did not confirm this prediction, results from experiment 2 suggested a small effect of tACS frequency on target word
perception: Faster tACS lead to more long-vowel word percepts, in line with the previous neuroimaging findings. Importantly, the difference in word perception induced by the different tACS frequencies was significantly larger in experiment 1 vs. experiment 2, suggesting that the
impact of tACS is dependent on the sensory context. tACS may have a stronger effect on spoken word perception when the words are presented in continuous speech as compared to when they are isolated, potentially because prior (stimulus-induced) entrainment of brain oscillations
might be a prerequisite for tACS to be effective.

Abstract

Perceiving speech requires the integration of different speech cues, that is, formants. When the speech signal is split so that different cues are presented to the right and left ear (dichotic listening), comprehension requires the integration of binaural information. Based on prior electrophysiological evidence, we hypothesized that the integration of dichotically presented speech cues is enabled by interhemispheric phase synchronization between primary and secondary auditory cortex in the gamma frequency band. We tested this hypothesis by applying transcranial alternating current stimulation (TACS) bilaterally above the superior temporal lobe to induce or disrupt interhemispheric gamma-phase coupling. In contrast to initial predictions, we found that gamma TACS applied in-phase above the two hemispheres (interhemispheric lag 0°) perturbs interhemispheric integration of speech cues, possibly because the applied stimulation perturbs an inherent phase lag between the left and right auditory cortex. We also observed this disruptive effect when applying antiphasic delta TACS (interhemispheric lag 180°). We conclude that interhemispheric phase coupling plays a functional role in interhemispheric speech integration. The direction of this effect may depend on the stimulation frequency.

Abstract

This neuroimaging study investigated the neural infrastructure of sentence-level language production. We compared brain activation patterns, as measured with BOLD-fMRI, during production of sentences that differed in verb argument structures (intransitives, transitives, ditransitives) and the lexical status of the verb (known verbs or pseudoverbs). The experiment consisted of 30 mini-blocks of six sentences each. Each mini-block started with an example for the type of sentence to be produced in that block. On each trial in the mini-blocks, participants were first given the (pseudo-)verb followed by three geometric shapes to serve as verb arguments in the sentences. Production of sentences with known verbs yielded greater activation compared to sentences with pseudoverbs in the core language network of the left inferior frontal gyrus, the left posterior middle temporalgyrus, and a more posterior middle temporal region extending into the angular gyrus, analogous to effects observed in language comprehension. Increasing the number of verb arguments led to greater activation in an overlapping left posterior middle temporal gyrus/angular gyrus area, particularly for known verbs, as well as in the bilateral precuneus. Thus, producing sentences with more complex structures using existing verbs leads to increased activation in the language network, suggesting some reliance on memory retrieval of stored lexical–syntactic information during sentence production. This study thus provides evidence from sentence-level language production in line with functional models of the language network that have so far been mainly based on single-word production, comprehension, and language processing in aphasia.

Abstract

Catecholamine (CA) function has been widely implicated in cognitive functions that are tied to the prefrontal cortex and striatal areas. The present study investigated the effects of methylphenidate, which is a CA agonist, on the electroencephalogram (EEG) response related to semantic processing using a double-blind, placebo-controlled, randomized, crossover, within-subject design. Forty-eight healthy participants read semantically congruent or incongruent sentences after receiving 20-mg methylphenidate or a placebo while their brain activity was monitored with EEG. To probe whether the catecholaminergic modulation is task-dependent, in one condition participants had to focus on comprehending the sentences, while in the other condition, they only had to attend to the font size of the sentence. The results demonstrate that methylphenidate has a task-dependent effect on semantic processing. Compared to placebo, when semantic processing was task-irrelevant, methylphenidate enhanced the detection of semantic incongruence as indexed by a larger N400 amplitude in the incongruent sentences; when semantic processing was task-relevant, methylphenidate induced a larger N400 amplitude in the semantically congruent condition, which was followed by a larger late positive complex effect. These results suggest that CA-related neurotransmitters influence language processing, possibly through the projections between the prefrontal cortex and the striatum, which contain many CA receptors.

Abstract

Changes in oscillatory brain dynamics can be studied by means of induced band power (IBP) analyses, which quantify event-related changes in amplitude of frequency-specific EEG rhythms. Such analyses capture EEG phenomena that are not part of traditional event-related potential measures. The present study investigated whether IBP changes in the δ, θ, and α frequency ranges are sensitive to syntactic violations in sentences. Subjects read sentences that either were correct or contained a syntactic violation. The violations were either grammatical gender agreement violations, where a prenominal adjective was not appropriately inflected for the head noun's gender, or number agreement violations, in which a plural quantifier was combined with a singular head noun. IBP changes of the concurrently measured EEG were computed in five frequency bands of 2-Hz width, individually adjusted on the basis of subjects' α peak, ranging approximately from 2 to 12 Hz. Words constituting a syntactic violation elicited larger increases in θ power than the same words in a correct sentence context, in an interval of 300–500 ms after word onset. Of all the frequency bands studied, this was true for the θ frequency band only. The scalp topography of this effect was different for different violations: following number violations a left-hemispheric dominance was found, whereas gender violations elicited a right-hemisphere dominance of the θ power increase. Possible interpretations of this effect are considered in closing.

Abstract

By analyzing event-related changes in induced band power in narrow frequency bands of the human electroencephalograph, the present paper explores a possible functional role of the alpha and theta rhythms during the processing of words and of sentences. The results show a phasic power increase in the theta frequency range, together with a phasic power decrease in the alpha frequency range, following the presentation of words in a sentence. These effects may be related to word processing, either lexical or in relation to sentence context. Most importantly, there is a slow and highly frequency-specific increase in theta power as a sentence unfolds, possibly related to the formation of an episodic memory trace, or to incremental verbal working memory load.

Abstract

The control of language use has in its simplest form perhaps been most intensively studied using the color–word Stroop task. The authors review chronometric and neuroimaging evidence on Stroop task performance to evaluate two prominent, implemented models of control in naming and reading: GRAIN and WEAVER++. Computer simulations are reported, which reveal that WEAVER++ offers a more satisfactory account of the data than GRAIN. In particular, we report WEAVER++ simulations of the BOLD response in anterior cingulate cortex during Stroop performance. Aspects of single-word production and perception in the Stroop task are discussed in relation to the wider problem of the control of language use.

Abstract

This paper presents electrophysiological evidence of an impairment in the on-line processing of word class information in patients with Broca’s aphasia with agrammatic comprehension. Event-related brain potentials (ERPs) were recorded from the scalp while Broca patients and non-aphasic control subjects read open- and closed-class words that appeared one at a time on a PC screen. Separate waveforms were computed for open- and closed-class words. The non-aphasic control subjects showed a modulation of an early left anterior negativity in the 210–325 ms as a function of vocabulary class (VC), and a late left anterior negative shift to closed-class words in the 400–700 ms epoch. An N400 effect was present in both control subjects and Broca patients. We have taken the early electrophysiological differences to reflect the first availability of word-category information from the mental lexicon. The late differences can be related to post-lexical processing. In contrast to the control subjects, the Broca patients showed no early VC effect and no late anterior shift to closed-class words. The results support the view that an incomplete and/or delayed availability of word-class information might be an important factor in Broca’s agrammatic comprehension.