Publications

Abstract

When perceiving the world around us, we are constantly integrating pieces of information. The integrated experience consists of more than just the sum of its parts. For example, visual scenes are defined by a collection of objects as well as the spatial relations amongst them and sentence meaning is computed based on individual word semantic but also syntactic configuration. Having quantitative models of such integrated representations can help evaluate cognitive models of both language and scene perception. Here, we focus on language, and use a behavioral measure of perceived similarity as an approximation of integrated meaning representations. We collected similarity judgments of 200 subjects rating nouns or transitive sentences through an online multiple arrangement task. We find that perceived similarity between sentences is most strongly modulated by the semantic action category of the main verb. In addition, we show how non-negative matrix factorization of similarity judgment data can reveal multiple underlying dimensions reflecting both semantic as well as relational role information. Finally, we provide an example of how similarity judgments on sentence stimuli can serve as a point of comparison for artificial neural networks models (ANNs) by comparing our behavioral data against sentence similarity extracted from three state-of-the-art ANNs. Overall, our method combining the multiple arrangement task on sentence stimuli with matrix factorization can capture relational information emerging from integration of multiple words in a sentence even in the presence of strong focus on the verb.

Abstract

Recent artificial neural networks that process natural language achieve unprecedented performance in tasks requiring sentence-level understanding. As such, they could be interesting models of the integration of linguistic information in the human brain. We review works that compare these artificial language models with human brain activity and we assess the extent to which this approach has improved our understanding of the neural processes involved in natural language comprehension. Two main results emerge. First, the neural representation of word meaning aligns with the context-dependent, dense word vectors used by the artificial neural networks. Second, the processing hierarchy that emerges within artificial neural networks broadly matches the brain, but is surprisingly inconsistent across studies. We discuss current challenges in establishing artificial neural networks as process models of natural language comprehension. We suggest exploiting the highly structured representational geometry of artificial neural networks when mapping representations to brain data.

Abstract

Background: Despite a pervasive cortico-centric view in cognitive neuroscience, subcortical structures including the thalamus have been shown to be increasingly involved in higher cognitive functions. Previous structural and functional imaging studies demonstrated cortico-thalamo-cortical loops which may support various cognitive functions including language. However, large-scale functional connectivity of the thalamus during language tasks has not been examined before. Methods: The present study employed meta-analytic connectivity modeling to identify language-related coactivation patterns of the left and right thalami. The left and right thalami were used as regions of interest to search the BrainMap functional database for neuroimaging experiments with healthy participants reporting language-related activations in each region of interest. Activation likelihood estimation analyses were then carried out on the foci extracted from the identified studies to estimate functional convergence for each thalamus. A functional decoding analysis based on the same database was conducted to characterize thalamic contributions to different language functions. Results: The results revealed bilateral frontotemporal and bilateral subcortical (basal ganglia) coactivation patterns for both the left and right thalami, and also right cerebellar coactivations for the left thalamus, during language processing. In light of previous empirical studies and theoretical frameworks, the present connectivity and functional decoding findings suggest that cortico-subcortical-cerebellar-cortical loops modulate and fine-tune information transfer within the bilateral frontotemporal cortices during language processing, especially during production and semantic operations, but also other language (e.g., syntax, phonology) and cognitive operations (e.g., attention, cognitive control). Conclusion: The current findings show that the language-relevant network extends beyond the classical left perisylvian cortices and spans bilateral cortical, bilateral subcortical (bilateral thalamus, bilateral basal ganglia) and right cerebellar regions.

Abstract

The language faculty is physically realized in the neurobiological infrastructure of the human brain. Despite significant efforts, an integrated understanding of this system remains a formidable challenge. What is missing from most theoretical accounts is a specification of the neural mechanisms that implement language function. Computational models that have been put forward generally lack an explicit neurobiological foundation. We propose a neurobiologically informed causal modeling approach which offers a framework for how to bridge this gap. A neurobiological causal model is a mechanistic description of language processing that is grounded in, and constrained by, the characteristics of the neurobiological substrate. It intends to model the generators of language behavior at the level of implementational causality. We describe key features and neurobiological component parts from which causal models can be built and provide guidelines on how to implement them in model simulations. Then we outline how this approach can shed new light on the core computational machinery for language, the long-term storage of words in the mental lexicon and combinatorial processing in sentence comprehension. In contrast to cognitive theories of behavior, causal models are formulated in the “machine language” of neurobiology which is universal to human cognition. We argue that neurobiological causal modeling should be pursued in addition to existing approaches. Eventually, this approach will allow us to develop an explicit computational neurobiology of language.

Abstract

The neural correlates of sentence production have been mostly studied with constraining task paradigms that introduce artificial task effects. In this study, we aimed to gain a better understanding of syntactic processing in spontaneous production vs. naturalistic comprehension. We extracted word-by-word metrics of phrase-structure building with top-down and bottom-up parsers that make different hypotheses about the timing of structure building. In comprehension, structure building proceeded in an integratory fashion and led to an increase in activity in posterior temporal and inferior frontal areas. In production, structure building was anticipatory and predicted an increase in activity in the inferior frontal gyrus. Newly developed production-specific parsers highlighted the anticipatory and incremental nature of structure building in production, which was confirmed by a converging analysis of the pausing patterns in speech. Overall, the results showed that the unfolding of syntactic processing diverges between speaking and listening.

Abstract

The neural infrastructure for sentence production and comprehension has been found to be mostly shared. The same regions are engaged during speaking and listening, with some differences in how strongly they activate depending on modality. In this study, we investigated how modality affects the connectivity between regions previously found to be involved in syntactic processing across modalities. We determined how constituent size and modality affected the connectivity of the pars triangularis of the left inferior frontal gyrus (LIFG) and of the left posterior temporal lobe (LPTL) with the pars opercularis of the LIFG, the anterior temporal lobe (LATL) and the rest of the brain. We found that constituent size reliably increased the connectivity across these frontal and temporal ROIs. Connectivity between the two LIFG regions and the LPTL was enhanced as a function of constituent size in both modalities, and it was upregulated in production possibly because of linearization and motor planning in the frontal cortex. The connectivity of both ROIs with the LATL was lower and only enhanced for larger constituent sizes, suggesting a contributing role of the LATL in sentence processing in both modalities. These results thus show that the connectivity among fronto-temporal regions is upregulated for syntactic structure building in both sentence production and comprehension, providing further evidence for accounts of shared neural resources for sentence-level processing across modalities.

Abstract

The neural representations for compositional processing have so far been mostly studied during sentence comprehension. In an fMRI study of sentence production, we investigated the brain representations for compositional processing during speaking. We used a rapid serial visual presentation sentence recall paradigm to elicit sentence production from the conceptual memory of an event. With voxel-wise encoding models, we probed the specificity of the compositional structure built during the production of each sentence, comparing an unstructured model of word meaning without relational information with a model that encodes abstract thematic relations and a model encoding event-specific relational structure. Whole-brain analyses revealed that sentence meaning at different levels of specificity was encoded in a large left frontal-parietal-temporal network. A comparison with semantic structures composed during the comprehension of the same sentences showed similarly distributed brain activity patterns. An ROI analysis over left fronto-temporal language parcels showed that event-specific relational structure above word-specific information was encoded in the left inferior frontal gyrus. Overall, we found evidence for the encoding of sentence meaning during sentence production in a distributed brain network and for the encoding of event-specific semantic structures in the left inferior frontal gyrus.

Abstract

Language is inherently multimodal. In spoken languages, combined spoken and visual signals (e.g., co-speech gestures) are an integral part of linguistic structure and language representation. This requires an extension of the parallel architecture, which needs to include the visual signals concomitant to speech. We present the evidence for the multimodality of language. In addition, we propose that distributional semantics might provide a format for integrating speech and co-speech gestures in a common semantic representation.

Abstract

How we combine minimal linguistic units into larger structures remains an unresolved topic in neuroscience. Language processing involves the abstract construction of ‘vertical’ and ‘horizontal’ information simultaneously (e.g., phrase structure, morphological agreement), but previous paradigms have been constrained in isolating only one type of composition and have utilized poor spatiotemporal resolution. Using intracranial recordings, we report multiple experiments designed to separate phrase structure from morphosyntactic agreement. Epilepsy patients (n = 10) were presented with auditory two-word phrases grouped into pseudoword-verb (‘trab run’) and pronoun-verb either with or without Person agreement (‘they run’ vs. ‘they runs’). Phrase composition and Person violations both resulted in significant increases in broadband high gamma activity approximately 300ms after verb onset in posterior middle temporal gyrus (pMTG) and posterior superior temporal sulcus (pSTS), followed by inferior frontal cortex (IFC) at 500ms. While sites sensitive to only morphosyntactic violations were distributed, those sensitive to both composition types were generally confined to pSTS/pMTG and IFC. These results indicate that posterior temporal cortex shows the earliest sensitivity for hierarchical linguistic structure across multiple dimensions, providing neural resources for distinct windows of composition. This region is comprised of sparsely interwoven heterogeneous constituents that afford cortical search spaces for dissociable syntactic relations.

Abstract

During communication in real-life settings, our brain often needs to integrate auditory and visual information, and at the same time actively focus on the relevant sources of information, while ignoring interference from irrelevant events. The interaction between integration and attention processes remains poorly understood. Here, we use rapid invisible frequency tagging (RIFT) and magnetoencephalography (MEG) to investigate how attention affects auditory and visual information processing and integration, during multimodal communication. We presented human participants (male and female) with videos of an actress uttering action verbs (auditory; tagged at 58 Hz) accompanied by two movie clips of hand gestures on both sides of fixation (attended stimulus tagged at 65 Hz; unattended stimulus tagged at 63 Hz). Integration difficulty was manipulated by a lower-order auditory factor (clear/degraded speech) and a higher-order visual semantic factor (matching/mismatching gesture). We observed an enhanced neural response to the attended visual information during degraded speech compared to clear speech. For the unattended information, the neural response to mismatching gestures was enhanced compared to matching gestures. Furthermore, signal power at the intermodulation frequencies of the frequency tags, indexing non-linear signal interactions, was enhanced in left frontotemporal and frontal regions. Focusing on LIFG (Left Inferior Frontal Gyrus), this enhancement was specific for the attended information, for those trials that benefitted from integration with a matching gesture. Together, our results suggest that attention modulates audiovisual processing and interaction, depending on the congruence and quality of the sensory input.

Abstract

Words are not processed in isolation; instead, they are commonly embedded in phrases and sentences. The sentential context influences the perception and processing of a word. However, how this is achieved by brain processes and whether predictive mechanisms underlie this process remain a debated topic. Here, we employed an experimental paradigm in which we orthogonalized sentence context constraints and predictive validity, which was defined as the ratio of congruent to incongruent sentence endings within the experiment. While recording electroencephalography, participants read sentences with three levels of sentential context constraints (high, medium, and low). Participants were also separated into two groups that differed in their ratio of valid congruent to incongruent target words that could be predicted from the sentential context. For both groups, we investigated modulations of alpha power before, and N400 amplitude modulations after target word onset. The results reveal that the N400 amplitude gradually decreased with higher context constraints and cloze probability. In contrast, alpha power was not significantly affected by context constraint. Neither the N400 nor alpha power were significantly affected by changes in predictive validity.

Abstract

There is evidence from both behavior and brain activity that the way information is structured, through the use of focus, can up-regulate processing of focused constituents, likely to give prominence to the relevant aspects of the input. This is hypothesized to be universal, regardless of the different ways in which languages encode focus. In order to test this universalist hypothesis, we need to go beyond the more familiar linguistic strategies for marking focus, such as by means of intonation or specific syntactic structures (e.g., it-clefts). Therefore, in this study, we examine Makhuwa-Enahara, a Bantu language spoken in northern Mozambique, which uniquely marks focus through verbal conjugation. The participants were presented with sentences that consisted of either a semantically anomalous constituent or a semantically nonanomalous constituent. Moreover, focus on this particular constituent could be either present or absent. We observed a consistent pattern: Focused information generated a more negative N400 response than the same information in nonfocus position. This demonstrates that regardless of how focus is marked, its consequence seems to result in an upregulation of processing of information that is in focus.

Abstract

In Swedish, words are associated with either of two pitch contours known as Accent 1 and Accent 2. Using a psychometric test, we investigated how listeners judge pitch accent violations while interpreting discourse. Forty native speakers of Central Swedish were presented with auditory dialogues, where test words were appropriately or inappropriately accented in a given context, and asked to judge the correctness of sentences containing the test words. Data indicated a statistically significant effect of wrong accent pattern on the correctness judgment. Both Accent 1 and Accent 2 violations interfered with the coherent interpretation of discourse and were judged as incorrect by the listeners. Moreover, there was a statistically significant difference in the perceived correctness between the accent patterns. Accent 2 violations led to a lower correctness score compared to Accent 1 violations, indicating that the listeners were more sensitive to pitch accent violations in Accent 2 words than in Accent 1 words. This result is in line with the notion that Accent 2 is marked and lexically represented in Central Swedish. Taken together, these findings indicate that listeners use both Accent 1 and Accent 2 to arrive at the correct interpretation of the linguistic input, while assigning varying degrees of relevance to them depending on their markedness.

Abstract

Although the sentences that we hear or read have meaning, this does not necessarily mean that they are also true. Relatively little is known about the critical brain structures for, and the relative time course of, establishing the meaning and truth of linguistic expressions. We present electroencephalogram data that show the rapid parallel integration of both semantic and world
knowledge during the interpretation of a sentence. Data from functional magnetic resonance imaging revealed that the left inferior prefrontal cortex is involved in the integration of both meaning and world knowledge. Finally, oscillatory brain responses indicate that the brain keeps a record of what makes a sentence hard to interpret.

Abstract

Following up on an earlier positron emission tomography (PET) experiment (Indefrey et al., 2001), we used a scene description paradigm to investigate whether a posterior inferior frontal region subserving syntactic encoding for speaking is also involved in syntactic parsing during listening. In the language production part of the experiment, subjects described visually presented scenes
using either sentences, sequences of noun phrases, or sequences of syntactically unrelated words. In the language comprehension part of the experiment, subjects were auditorily presented with the same kinds of utterances and judged whether they matched the visual scenes. We were able to replicate the previous finding of a region in caudal Broca s area that is sensitive to the complexity of
syntactic encoding in language production. In language comprehension, no hemodynamic activation differences due to syntactic complexity were found. Given that correct performance in the judgment task did not require syntactic processing of the auditory stimuli, the results suggest that the degree to which listeners recruit syntactic processing resources in language comprehension may be a function of the syntactic demands of the task or the stimulus material.

Abstract

This study addresses the supralexical inferential processes underlying wellformedness judgements and latencies for a specic sublexical unit that appears in Dutch compounds, the interfix. Production studies have shown that the selection of interfixes in novel Dutch compounds and the speed of
this selection is primarily determined by the distribution of interfixes in existing compounds that share the left constituent with the target compound, i.e. the ‘‘left constituent family’’. In this paper, we consider the question whether constituent families also affect wellformedness decisions of novel as well as existing Dutch compounds in comprehension. We visually presented compounds containing interfixes that were either in line with the bias of the left constituent family or not. In the case of existing compounds, we also presented variants with replaced interfixes. As in production, the bias of the left constituent family emerged as a crucial predictor for both acceptance rates and response latencies. This result supports the hypothesis that, as in production, constituent families are (co-)activated in comprehension. We argue that this co-activation is part of a supralexical inferential process, and we discuss how our data might be interpreted within sublexical and supralexical theories of morphological processing.

Abstract

Chronic fatigue syndrome (CFS) is characterized by a debilitating fatigue of unknown aetiology. Patients who suffer from CFS report a variety of physical complaints as well as neuropsychological complaints. Therefore, it is conceivable that the CNS plays a role in the pathophysiology of CFS. The purpose of this study was to investigate neural correlates of CFS, and specifically whether there exists a linkage between disturbances in the motor system and CFS. We measured behavioural performance and cerebral activity using rapid event-related functional MRI in 16 CFS patients and 16 matched healthy controls while they were engaged in a motor imagery task and a control visual imagery task. CFS patients were considerably slower on performance of both tasks, but the increase in reaction time with increasing task load was similar between the groups. Both groups used largely overlapping neural resources. However, during the motor imagery task, CFS patients evoked stronger responses in visually related structures. Furthermore, there was a marked between-groups difference during erroneous performance. In both groups, dorsal anterior cingulate cortex was specifically activated during error trials. Conversely, ventral anterior cingulate cortex was active when healthy controls made an error, but remained inactive when CFS patients made an error. Our results support the notion that CFS may be associated with dysfunctional motor planning. Furthermore, the between-groups differences observed during erroneous performance point to motivational disturbances as a crucial component of CFS.

Abstract

An event-related brain potential experiment was carried out to investigate the influence of semantic and syntactic context constraints on lexical selection and integration in spoken-word comprehension. Subjects were presented with constraining spoken sentences that contained a critical word that was either (a) congruent, (b) semantically and syntactically incongruent, but beginning with the same initial phonemes as the congruent critical word, or (c) semantically and syntactically incongruent, beginning with phonemes that differed from the congruent critical word. Relative to the congruent condition, an N200 effect reflecting difficulty in the lexical selection process was obtained in the semantically and syntactically incongruent condition where word onset differed from that of the congruent critical word. Both incongruent conditions elicited a large N400 followed by a left anterior negativity (LAN) time-locked to the moment of word category violation and a P600 effect. These results would best fit within a cascaded model of spoken-word processing, proclaiming an optimal use of contextual information during spokenword identification by allowing for semantic and syntactic processing to take place in parallel after bottom-up activation of a set of candidates, and lexical integration to proceed with a limited number of candidates that still match the acoustic input.

Abstract

This article presents electrophysiological data on on-line syntactic processing during auditory sentence comprehension in patients with Broca's aphasia. Event-related brain potentials (ERPs) were recorded from the scalp while subjects listened to sentences that were either syntactically correct or contained violations of subject-verb agreement. Three groups of subjects were tested: Broca patients (n = 10), nonaphasic patients with a right-hemisphere (RH) lesion (n = 5), and healthy agedmatched controls (n = 12). The healthy, control subjects showed a P600/SPS effect as response to the agreement violations. The nonaphasic patients with an RH lesion showed essentially the same pattern. The overall group of Broca patients did not show this sensitivity. However, the sensitivity was modulated by the severity of the syntactic comprehension impairment. The largest deviation from the standard P600/SPS effect was found in the patients with the relatively more severe syntactic comprehension impairment. In addition, ERPs to tones in a classical tone oddball paradigm were also recorded. Similar to the normal control subjects and RH patients, the group of Broca patients showed a P300 effect in the tone oddball condition. This indicates that aphasia in itself does not lead to a general reduction in all cognitive ERP effects. It was concluded that deviations from the standard P600/SPS effect in the Broca patients reflected difficulties with on-line maintaining of number information across clausal boundaries for establishing subject-verb agreement.

Abstract

This study tests the recent claim that Broca’s aphasics are impaired in automatic lexical access, including the retrieval of word meaning. Subjects are required to perform a lexical decision on visually presented prime target pairs. Half of the word targets are preceded by a related word, half by an unrelated word. Primes and targets are presented with a long stimulus-onset-asynchrony (SOA) of 1400 msec and with a short SOA of 300 msec. Normal priming effects are observed in Broca’s aphasics for both SOAs. This result is discussed in the context of the claim that Broca’s aphasics suffer from an impairment in the automatic access of lexical–semantic information. It is argued that none of the current priming studies provides evidence supporting this claim, since with short SOAs priming effects have been reliably obtained in Broca’s aphasics. The results are more compatible with the claim that in many Broca’s aphasics the functional locus of their comprehension deficit is at the level of postlexical integration processes.

Abstract

Stimulus-related changes in cerebral blood oxygenation were measured using high-resolution functional magnetic resonance imaging sequentially covering visual occipital areas in contiguous sections. During dynamic imaging, healthy subjects silently viewed pseudowords, single false fonts, or length-matched strings of the same false fonts. The paradigm consisted of a sixfold alternation of an activation and a control task. With pseudowords as activation vs single false fonts as control, responses were seen mainly in medial occipital cortex. These responses disappeared when pseudowords were alternated with false font strings as the control and reappeared when false font strings instead of pseudowords served as activation and were alternated with single false fonts. The string-length contrast alone, therefore, is sufficient to account for the activation pattern observed in medial visual cortex when word-like stimuli are contrasted with single characters.

Abstract

In this study the N400 component of the event-related potential was used to investigate spoken sentence understanding in Broca's and Wernicke's aphasics. The aim of the study was to determine whether spoken sentence comprehension problems in these patients might result from a deficit in the on-line integration of lexical information. Subjects listened to sentences spoken at a normal rate. In half of these sentences, the meaning of the final word of the sentence matched the semantic specifications of the preceding sentence context. In the other half of the sentences, the sentence-final word was anomalous with respect to the preceding sentence context. The N400 was measured to the sentence-final words in both conditions. The results for the aphasic patients (n = 14) were analyzed according to the severity of their comprehension deficit and compared to a group of 12 neurologically unimpaired age-matched controls, as well as a group of 6 nonaphasic patients with a lesion in the right hemisphere. The nonaphasic brain damaged patients and the aphasic patients with a light comprehension deficit (high comprehenders, n = 7) showed an N400 effect that was comparable to that of the neurologically unimpaired subjects. In the aphasic patients with a moderate to severe comprehension deficit (low comprehenders, n = 7), a reduction and delay of the N400 effect was obtained. In addition, the P300 component was measured in a classical oddball paradigm, in which subjects were asked to count infrequent low tones in a random series of high and low tones. No correlation was found between the occurrence of N400 and P300 effects, indicating that changes in the N400 results were related to the patients' language deficit. Overall, the pattern of results was compatible with the idea that aphasic patients with moderate to severe comprehension problems are impaired in the integration of lexical information into a higher order representation of the preceding sentence context.

Abstract

The temporal properties of semantic and phonological processes in speech production were investigated in a new experimental paradigm using movement-related brain potentials. The main experimental task was picture naming. In addition, a 2-choice reaction go/no-go procedure was included, involving a semantic and a phonological categorization of the picture name. Lateralized readiness potentials (LRPs) were derived to test whether semantic and phonological information activated motor processes at separate moments in time. An LRP was only observed on no-go trials when the semantic (not the phonological) decision determined the response hand. Varying the position of the critical phoneme in the picture name did not affect the onset of the LRP but rather influenced when the LRP began to differ on go and no-go trials and allowed the duration of phonological encoding of a word to be estimated. These results provide electrophysiological evidence for early semantic activation and later phonological encoding.