Publications

Abstract

Human language offers a variety of ways to create meaning, one of which is referring to entities, objects, or events in the world. One such meaning maker is understanding to whom or to what a pronoun in a discourse refers to. To understand a pronoun, the brain must access matching entities or concepts that have been encoded in memory from previous linguistic context. Models of language processing propose that internally stored linguistic concepts, accessed via exogenous cues such as phonological input of a word, are represented as (a)synchronous activities across a population of neurons active at specific frequency bands. Converging evidence suggests that delta band activity (1–3 Hz) is involved in temporal and representational integration during sentence processing. Moreover, recent advances in the neurobiology of memory suggest that recollection engages neural dynamics similar to those which occurred during memory encoding. Integrating from these two research lines, we here tested the hypothesis that neural dynamic patterns, especially in delta frequency range, underlying referential meaning representation, would be reinstated during pronoun resolution. By leveraging neural decoding techniques (i.e., representational similarity analysis) on a magnetoencephalogram data set acquired during a naturalistic story-listening task, we provide evidence that delta-band activity underlies referential meaning representation. Our findings suggest that, during spoken language comprehension, endogenous linguistic representations such as referential concepts may be proactively retrieved and represented via activation of their underlying dynamic neural patterns.

Abstract

When we understand language, we recognize words and combine them into sentences. In this article, we explore the hypothesis that listeners use probabilistic information about words to build syntactic structure. Recent work has shown that lexical probability and syntactic structure both modulate the delta-band (<4 Hz) neural signal. Here, we investigated whether the neural encoding of syntactic structure changes as a function of the distributional properties of a word. To this end, we analyzed MEG data of 24 native speakers of Dutch who listened to three fairytales with a total duration of 49 min. Using temporal response functions and a cumulative model-comparison approach, we evaluated the contributions of syntactic and distributional features to the variance in the delta-band neural signal. This revealed that lexical surprisal values (a distributional feature), as well as bottom-up node counts (a syntactic feature) positively contributed to the model of the delta-band neural signal. Subsequently, we compared responses to the syntactic feature between words with high- and low-surprisal values. This revealed a delay in the response to the syntactic feature as a consequence of the surprisal value of the word: high-surprisal values were associated with a delayed response to the syntactic feature by 150–190 ms. The delay was not affected by word duration, and did not have a lexical origin. These findings suggest that the brain uses probabilistic information to infer syntactic structure, and highlight an importance for the role of time in this process.

Abstract

From a brain's-eye-view, when a stimulus occurs and what it is are interrelated aspects of interpreting the perceptual world. Yet in practice, the putative perceptual inferences about sensory content and timing are often dichotomized and not investigated as an integrated process. We here argue that neural temporal dynamics can influence what is perceived, and in turn, stimulus content can influence the time at which perception is achieved. This computational principle results from the highly interdependent relationship of what and when in the environment. Both brain processes and perceptual events display strong temporal variability that is not always modeled; we argue that understanding—and, minimally, modeling—this temporal variability is key for theories of how the brain generates unified and consistent neural representations and that we ignore temporal variability in our analysis practice at the peril of both data interpretation and theory-building. Here, we review what and when interactions in the brain, demonstrate via simulations how temporal variability can result in misguided interpretations and conclusions, and outline how to integrate and synthesize what and when in theories and models of brain computation.

Abstract

Neural oscillations reflect fluctuations in excitability, which biases the percept of ambiguous sensory input. Why this bias occurs is still not fully understood. We hypothesized that neural populations representing likely events are more sensitive, and thereby become active on earlier oscillatory phases, when the ensemble itself is less excitable. Perception of ambiguous input presented during less-excitable phases should therefore be biased toward frequent or predictable stimuli that have lower activation thresholds. Here, we show such a frequency bias in spoken word recognition using psychophysics, magnetoencephalography (MEG), and computational modelling. With MEG, we found a double dissociation, where the phase of oscillations in the superior temporal gyrus and medial temporal gyrus biased word-identification behavior based on phoneme and lexical frequencies, respectively. This finding was reproduced in a computational model. These results demonstrate that oscillations provide a temporal ordering of neural activity based on the sensitivity of separable neural populations.

Abstract

Humans excel at extracting structurally-determined meaning from speech despite inherent physical variability. This study explores the brain’s ability to predict and understand spoken language robustly. It investigates the relationship between structural and statistical language knowledge in brain dynamics, focusing on phase and amplitude modulation. Using syntactic features from constituent hierarchies and surface statistics from a transformer model as predictors of forward encoding models, we reconstructed cross-frequency neural dynamics from MEG data during audiobook listening. Our findings challenge a strict separation of linguistic structure and statistics in the brain, with both aiding neural signal reconstruction. Syntactic features have a more temporally spread impact, and both word entropy and the number of closing syntactic constituents are linked to the phase-amplitude coupling of neural dynamics, implying a role in temporal prediction and cortical oscillation alignment during speech processing. Our results indicate that structured and statistical information jointly shape neural dynamics during spoken language comprehension and suggest an integration process via a cross-frequency coupling mechanism

Abstract

Electrophysiological brain activity has been shown to synchronize with the quasi-regular repetition of grammatical phrases in connected speech—so-called phrase-rate neural tracking. Current debate centers around whether this phenomenon is best explained in terms of the syntactic properties of phrases or in terms of syntax-external information, such as the sequential repetition of parts of speech. As these two factors were confounded in previous studies, much of the literature is compatible with both accounts. Here, we used electroencephalography (EEG) to determine if and when the brain is sensitive to both types of information. Twenty native speakers of Mandarin Chinese listened to isochronously presented streams of monosyllabic words, which contained either grammatical two-word phrases (e.g., catch fish, sell house) or non-grammatical word combinations (e.g., full lend, bread far). Within the grammatical conditions, we varied two structural factors: the position of the head of each phrase and the type of attachment. Within the non-grammatical conditions, we varied the consistency with which parts of speech were repeated. Tracking was quantified through evoked power and inter-trial phase coherence, both derived from the frequency-domain representation of EEG responses. As expected, neural tracking at the phrase rate was stronger in grammatical sequences than in non-grammatical sequences without syntactic structure. Moreover, it was modulated by both attachment type and head position, revealing the structure-sensitivity of phrase-rate tracking. We additionally found that the brain tracks the repetition of parts of speech in non-grammatical sequences. These data provide an integrative perspective on the current debate about neural tracking effects, revealing that the brain utilizes regularities computed over multiple levels of linguistic representation in guiding rhythmic computation.

Abstract

Research into the role of brain oscillations in basic perceptual and cognitive functions has suggested that the alpha rhythm reflects functional inhibition while the beta rhythm reflects neural ensemble (re)activation. However, little is known regarding the generalization of these proposed fundamental operations to linguistic processes, such as speech comprehension and production. Here, we recorded magnetoencephalography in participants performing a novel rule-switching paradigm. Specifically, Dutch native speakers had to produce an alternative exemplar from the same category or a feature of a given target word embedded in spoken sentences (e.g., for the word “tuna”, an exemplar from the same category—“seafood”—would be “shrimp”, and a feature would be “pink”). A cue indicated the task rule—exemplar or feature—either before (pre-cue) or after (retro-cue) listening to the sentence. Alpha power during the working memory delay was lower for retro-cue compared with that for pre-cue in the left hemispheric language-related regions. Critically, alpha power negatively correlated with reaction times, suggestive of alpha facilitating task performance by regulating inhibition in regions linked to lexical retrieval. Furthermore, we observed a different spatiotemporal pattern of beta activity for exemplars versus features in the right temporoparietal regions, in line with the proposed role of beta in recruiting neural networks for the encoding of distinct categories. Overall, our study provides evidence for the generalizability of the role of alpha and beta oscillations from perceptual to more “complex, linguistic processes” and offers a novel task to investigate links between rule-switching, working memory, and word production.

Abstract

Computation in neuronal assemblies is putatively reflected in the excitatory and inhibitory cycles of activation distributed throughout the brain. In speech and language processing, coordination of these cycles resulting in phase synchronization has been argued to reflect the integration of information on different timescales (e.g. segmenting acoustics signals to phonemic and syllabic representations; (Giraud and Poeppel 2012 Nat. Neurosci.15, 511 (doi:10.1038/nn.3063)). A natural extension of this claim is that phase synchronization functions similarly to support the inference of more abstract higher-level linguistic structures (Martin 2016 Front. Psychol.7, 120; Martin and Doumas 2017 PLoS Biol. 15, e2000663 (doi:10.1371/journal.pbio.2000663); Martin and Doumas. 2019 Curr. Opin. Behav. Sci.29, 77–83 (doi:10.1016/j.cobeha.2019.04.008)). Hale et al. (Hale et al. 2018 Finding syntax in human encephalography with beam search. arXiv 1806.04127 (http://arxiv.org/abs/1806.04127)) showed that syntactically driven parsing decisions predict electroencephalography (EEG) responses in the time domain; here we ask whether phase synchronization in the form of either inter-trial phrase coherence or cross-frequency coupling (CFC) between high-frequency (i.e. gamma) bursts and lower-frequency carrier signals (i.e. delta, theta), changes as the linguistic structures of compositional meaning (viz., bracket completions, as denoted by the onset of words that complete phrases) accrue. We use a naturalistic story-listening EEG dataset from Hale et al. to assess the relationship between linguistic structure and phase alignment. We observe increased phase synchronization as a function of phrase counts in the delta, theta, and gamma bands, especially for function words. A more complex pattern emerged for CFC as phrase count changed, possibly related to the lack of a one-to-one mapping between ‘size’ of linguistic structure and frequency band—an assumption that is tacit in recent frameworks. These results emphasize the important role that phase synchronization, desynchronization, and thus, inhibition, play in the construction of compositional meaning by distributed neural networks in the brain.

Abstract

Human thought and language have extraordinary expressive power because meaningful parts can be assembled into more complex semantic structures. This partly underlies our ability to compose meanings into endlessly novel configurations, and sets us apart from other species and current computing devices. Crucially, human behaviour, including language use and linguistic data, indicates that composing parts into complex structures does not threaten the existence of constituent parts as independent units in the system: parts and wholes exist simultaneously yet independently from one another in the mind and brain. This independence is evident in human behaviour, but it seems at odds with what is known about the brain's exquisite sensitivity to statistical patterns: everyday language use is productive and expressive precisely because it can go beyond statistical regularities. Formal theories in philosophy and linguistics explain this fact by assuming that language and thought are compositional: systems of representations that separate a variable (or role) from its values (fillers), such that the meaning of a complex expression is a function of the values assigned to the variables. The debate on whether and how compositional systems could be implemented in minds, brains and machines remains vigorous. However, it has not yet resulted in mechanistic models of semantic composition: how, then, are the constituents of thoughts and sentences put and held together? We review and discuss current efforts at understanding this problem, and we chart possible routes for future research.

Abstract

Neither neurobiological nor process models of meaning composition specify the operator through which constituent parts are bound together into compositional structures. In this paper, we argue that a neurophysiological computation system cannot achieve the compositionality exhibited in human thought and language if it were to rely on a multiplicative operator to perform binding, as the tensor product (TP)-based systems that have been widely adopted in cognitive science, neuroscience and artificial intelligence do. We show via simulation and two behavioural experiments that TPs violate variable-value independence, but human behaviour does not. Specifically, TPs fail to capture that in the statements fuzzy cactus and fuzzy penguin, both cactus and penguin are predicated by fuzzy(x) and belong to the set of fuzzy things, rendering these arguments similar to each other. Consistent with that thesis, people judged arguments that shared the same role to be similar, even when those arguments themselves (e.g., cacti and penguins) were judged to be dissimilar when in isolation. By contrast, the similarity of the TPs representing fuzzy(cactus) and fuzzy(penguin) was determined by the similarity of the arguments, which in this case approaches zero. Based on these results, we argue that neural systems that use TPs for binding cannot approximate how the human mind and brain represent compositional information during processing. We describe a contrasting binding mechanism that any physiological or artificial neural system could use to maintain independence between a role and its argument, a prerequisite for compositionality and, thus, for instantiating the expressive power of human thought and language in a neural system.

Abstract

Humans learn to represent complex structures (e.g. natural language, music, mathematics) from experience with their environments. Often such structures are latent, hidden, or not encoded in statistics about sensory representations alone. Accounts of human cognition have long emphasized the importance of structured representations, yet the majority of contemporary neural networks do not learn structure from experience. Here, we describe one way that structured, functionally symbolic representations can be instantiated in an artificial neural network. Then, we describe how such latent structures (viz. predicates) can be learned from experience with unstructured data. Our approach exploits two principles from psychology and neuroscience: comparison of representations, and the naturally occurring dynamic properties of distributed computing across neuronal assemblies (viz. neural oscillations). We discuss how the ability to learn predicates from experience, to represent information compositionally, and to extrapolate knowledge to unseen data is core to understanding and modeling the most complex human behaviors (e.g. relational reasoning, analogy, language processing, game play).

Abstract

Language comprehension requires recovering meaning from linguistic form, even when the mapping between the two is indirect. A canonical example is ellipsis, the omission of information that is subsequently understood without being overtly pronounced. Comprehension of ellipsis requires retrieval of an antecedent from memory, without prior prediction, a property which enables the study of retrieval in situ ( Martin and McElree, 2008 and Martin and McElree, 2009). Sluicing, or inflectional-phrase ellipsis, in the presence of a conjunction, presents a test case where a competing antecedent position is syntactically licensed, in contrast with most cases of nonadjacent dependency, including verb–phrase ellipsis. We present speed–accuracy tradeoff and eye-movement data inconsistent with the hypothesis that retrieval is accomplished via a syntactically guided search, a particular variant of search not examined in past research. The observed timecourse profiles are consistent with the hypothesis that antecedents are retrieved via a cue-dependent direct-access mechanism susceptible to general memory variables.