Neurobiological Causal Models of Language Processing.

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.

Formal language hierarchy reflects different levels of cognitive complexity.

Formal language hierarchy describes levels of increasing syntactic complexity (adjacent dependencies, nonadjacent nested, nonadjacent crossed) of which the transcription into a hierarchy of cognitive complexity remains under debate. The cognitive foundations of formal language hierarchy have been contradicted by two types of evidence: First, adjacent dependencies are not easier to learn compared to nonadjacent; second, crossed nonadjacent dependencies may be easier than nested. However, studies providing these findings may have engaged confounds: Repetition monitoring strategies may have accounted for participants' high performance in nonadjacent dependencies, and linguistic experience may have accounted for the advantage of crossed dependencies. We conducted two artificial grammar learning experiments where we addressed these confounds by manipulating reliance on repetition monitoring and by testing participants inexperienced with crossed dependencies. Results showed relevant differences in learning adjacent versus nonadjacent dependencies and advantages of nested over crossed, suggesting that formal language hierarchy may indeed translate into a hierarchy of cognitive complexity. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

The Tripod neuron: a minimal structural reduction of the dendritic tree.

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. KEY POINTS: We present a neuron model, called the Tripod, with two segregated dendritic branches that are connected to an axosomatic compartment. Each branch implements inhibitory GABAergic and excitatory glutamatergic synaptic transmission, including voltage-gated NMDA receptors. Dendrites are modelled on relevant geometric and physiological parameters measured in human pyramidal cells. The neuron reproduces classical dendritic computations, such as coincidence detection and pathway selection via shunting inhibition, that are beyond the scope of point-neuron models. Under some conditions, dendritic NMDA spikes cause plateau potentials, and we show that they provide a form of short-term memory which is useful for sequence recognition. The dendritic structure of the Tripod neuron is sufficiently simple to be integrated into efficient network simulations and studied in a broad functional context.

Supramodal Sentence Processing in the Human Brain: fMRI Evidence for the Influence of Syntactic Complexity in More Than 200 Participants.

This study investigated two questions. One is: To what degree is sentence processing beyond single words independent of the input modality (speech vs. reading)? The second question is: Which parts of the network recruited by both modalities is sensitive to syntactic complexity? These questions were investigated by having more than 200 participants read or listen to well-formed sentences or series of unconnected words. A largely left-hemisphere frontotemporoparietal network was found to be supramodal in nature, i.e., independent of input modality. In addition, the left inferior frontal gyrus (LIFG) and the left posterior middle temporal gyrus (LpMTG) were most clearly associated with left-branching complexity. The left anterior temporal lobe showed the greatest sensitivity to sentences that differed in right-branching complexity. Moreover, activity in LIFG and LpMTG increased from sentence onset to end, in parallel with an increase of the left-branching complexity. While LIFG, bilateral anterior temporal lobe, posterior MTG, and left inferior parietal lobe all contribute to the supramodal unification processes, the results suggest that these regions differ in their respective contributions to syntactic complexity related processing. The consequences of these findings for neurobiological models of language processing are discussed.

Distinguishing Syntactic Operations in the Brain: Dependency and Phrase-Structure Parsing.

Finding the structure of a sentence-the way its words hold together to convey meaning-is a fundamental step in language comprehension. Several brain regions, including the left inferior frontal gyrus, the left posterior superior temporal gyrus, and the left anterior temporal pole, are supposed to support this operation. The exact role of these areas is nonetheless still debated. In this paper we investigate the hypothesis that different brain regions could be sensitive to different kinds of syntactic computations. We compare the fit of phrase-structure and dependency structure descriptors to activity in brain areas using fMRI. Our results show a division between areas with regard to the type of structure computed, with the left anterior temporal pole and left inferior frontal gyrus favouring dependency structures and left posterior superior temporal gyrus favouring phrase structures.

Neuronal spike-rate adaptation supports working memory in language processing.

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 provide 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 mechanism does not rely on persistent activity, excitatory feedback, or synaptic plasticity for storage. Instead, information is maintained in adaptive conductances that reduce firing rates and can be accessed directly without cued retrieval. Memory span is systematically related to both the time constant of adaptation and baseline levels of neuronal excitability. Interference 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 toward a computational neurobiology of language.

Implicit sequence learning is preserved in dyslexic children.

This study investigates the implicit sequence learning abilities of dyslexic children using an artificial grammar learning task with an extended exposure period. Twenty children with developmental dyslexia participated in the study and were matched with two control groups-one matched for age and other for reading skills. During 3 days, all participants performed an acquisition task, where they were exposed to colored geometrical forms sequences with an underlying grammatical structure. On the last day, after the acquisition task, participants were tested in a grammaticality classification task. Implicit sequence learning was present in dyslexic children, as well as in both control groups, and no differences between groups were observed. These results suggest that implicit learning deficits per se cannot explain the characteristic reading difficulties of the dyslexics.

Modality effects in implicit artificial grammar learning: An EEG study.

Recently, it has been proposed that sequence learning engages a combination of modality-specific operating networks and modality-independent computational principles. In the present study, we compared the behavioural and EEG outcomes of implicit artificial grammar learning in the visual vs. auditory modality. We controlled for the influence of surface characteristics of sequences (Associative Chunk Strength), thus focusing on the strictly structural aspects of sequence learning, and we adapted the paradigms to compensate for known frailties of the visual modality compared to audition (temporal presentation, fast presentation rate). The behavioural outcomes were similar across modalities. Favouring the idea of modality-specificity, ERPs in response to grammar violations differed in topography and latency (earlier and more anterior component in the visual modality), and ERPs in response to surface features emerged only in the auditory modality. In favour of modality-independence, we observed three common functional properties in the late ERPs of the two grammars: both were free of interactions between structural and surface influences, both were more extended in a grammaticality classification test than in a preference classification test, and both correlated positively and strongly with theta event-related-synchronization during baseline testing. Our findings support the idea of modality-specificity combined with modality-independence, and suggest that memory for visual vs. auditory sequences may largely contribute to cross-modal differences.

Broca's region: A causal role in implicit processing of grammars with crossed non-adjacent dependencies.

Non-adjacent dependencies are challenging for the language learning machinery and are acquired later than adjacent dependencies. In this transcranial magnetic stimulation (TMS) study, we show that participants successfully discriminated between grammatical and non-grammatical sequences after having implicitly acquired an artificial language with crossed non-adjacent dependencies. Subsequent to transcranial magnetic stimulation of Broca's region, discrimination was impaired compared to when a language-irrelevant control region (vertex) was stimulated. These results support the view that Broca's region is engaged in structured sequence processing and extend previous functional neuroimaging results on artificial grammar learning (AGL) in two directions: first, the results establish that Broca's region is a causal component in the processing of non-adjacent dependencies, and second, they show that implicit processing of non-adjacent dependencies engages Broca's region. Since patients with lesions in Broca's region do not always show grammatical processing difficulties, the result that Broca's region is causally linked to processing of non-adjacent dependencies is a step towards clarification of the exact nature of syntactic deficits caused by lesions or perturbation to Broca's region. Our findings are consistent with previous results and support a role for Broca's region in general structured sequence processing, rather than a specific role for the processing of hierarchically organized sentence structure.

Eye movements in implicit artificial grammar learning.

Artificial grammar learning (AGL) has been probed with forced-choice behavioral tests (active tests). Recent attempts to probe the outcomes of learning (implicitly acquired knowledge) with eye-movement responses (passive tests) have shown null results. However, these latter studies have not tested for sensitivity effects, for example, increased eye movements on a printed violation. In this study, we tested for sensitivity effects in AGL tests with (Experiment 1) and without (Experiment 2) concurrent active tests (preference- and grammaticality classification) in an eye-tracking experiment. Eye movements discriminated between sequence types in passive tests and more so in active tests. The eye-movement profile did not differ between preference and grammaticality classification, and it resembled sensitivity effects commonly observed in natural syntax processing. Our findings show that the outcomes of implicit structured sequence learning can be characterized in eye tracking. More specifically, whole trial measures (dwell time, number of fixations) showed robust AGL effects, whereas first-pass measures (first-fixation duration) did not. Furthermore, our findings strengthen the link between artificial and natural syntax processing, and they shed light on the factors that determine performance differences in preference and grammaticality classification tests. (PsycINFO Database Record

The effects of ordinal load on incidental temporal learning.

How can we grasp the temporal structure of events? A few studies have indicated that representations of temporal structure are acquired when there is an intention to learn, but not when learning is incidental. Response-to-stimulus intervals, uncorrelated temporal structures, unpredictable ordinal information, and lack of metrical organization have been pointed out as key obstacles to incidental temporal learning, but the literature includes piecemeal demonstrations of learning under all these circumstances. We suggest that the unacknowledged effects of ordinal load may help reconcile these conflicting findings, ordinal load referring to the cost of identifying the sequence of events (e.g., tones, locations) where a temporal pattern is embedded. In a first experiment, we manipulated ordinal load into simple and complex levels. Participants learned ordinal-simple sequences, despite their uncorrelated temporal structure and lack of metrical organization. They did not learn ordinal-complex sequences, even though there were no response-to-stimulus intervals nor unpredictable ordinal information. In a second experiment, we probed learning of ordinal-complex sequences with strong metrical organization, and again there was no learning. We conclude that ordinal load is a key obstacle to incidental temporal learning. Further analyses showed that the effect of ordinal load is to mask the expression of temporal knowledge, rather than to prevent learning.

Disentangling stimulus plausibility and contextual congruency: Electro-physiological evidence for differential cognitive dynamics.

Expectancy mechanisms are routinely used by the cognitive system in stimulus processing and in anticipation of appropriate responses. Electrophysiology research has documented negative shifts of brain activity when expectancies are violated within a local stimulus context (e.g., reading an implausible word in a sentence) or more globally between consecutive stimuli (e.g., a narrative of images with an incongruent end). In this EEG study, we examine the interaction between expectancies operating at the level of stimulus plausibility and at more global level of contextual congruency to provide evidence for, or against, a disassociation of the underlying processing mechanisms. We asked participants to verify the congruency of pairs of cross-modal stimuli (a sentence and a scene), which varied in plausibility. ANOVAs on ERP amplitudes in selected windows of interest show that congruency violation has longer-lasting (from 100 to 500ms) and more widespread effects than plausibility violation (from 200 to 400ms). We also observed critical interactions between these factors, whereby incongruent and implausible pairs elicited stronger negative shifts than their congruent counterpart, both early on (100-200ms) and between 400-500ms. Our results suggest that the integration mechanisms are sensitive to both global and local effects of expectancy in a modality independent manner. Overall, we provide novel insights into the interdependence of expectancy during meaning integration of cross-modal stimuli in a verification task.

The P600 in Implicit Artificial Grammar Learning.

The suitability of the artificial grammar learning (AGL) paradigm to capture relevant aspects of the acquisition of linguistic structures has been empirically tested in a number of EEG studies. Some have shown a syntax-related P600 component, but it has not been ruled out that the AGL P600 effect is a response to surface features (e.g., subsequence familiarity) rather than the underlying syntax structure. Therefore, in this study, we controlled for the surface characteristics of the test sequences (associative chunk strength) and recorded the EEG before (baseline preference classification) and after (preference and grammaticality classification) exposure to a grammar. After exposure, a typical, centroparietal P600 effect was elicited by grammatical violations and not by unfamiliar subsequences, suggesting that the AGL P600 effect signals a response to structural irregularities. Moreover, preference and grammaticality classification showed a qualitatively similar ERP profile, strengthening the idea that the implicit structural mere-exposure paradigm in combination with preference classification is a suitable alternative to the traditional grammaticality classification test.

Visual naming deficits in dyslexia: An ERP investigation of different processing domains.

Naming speed deficits are well documented in developmental dyslexia, expressed by slower naming times and more errors in response to familiar items. Here we used event-related potentials (ERPs) to examine at what processing level the deficits in dyslexia emerge during a discrete-naming task. Dyslexic and skilled adult control readers performed a primed object-naming task, in which the relationship between the prime and the target was manipulated along perceptual, semantic and phonological dimensions. A 3×2 design that crossed Relationship Type (Visual, Phonemic Onset, and Semantic) with Relatedness (Related and Unrelated) was used. An attenuated N/P190 - indexing early visual processing - and N300 - which index late visual processing - was observed to pictures preceded by perceptually related (vs. unrelated) primes in the control but not in the dyslexic group. These findings suggest suboptimal processing in early stages of object processing in dyslexia, when integration and mapping of perceptual information to a more form-specific percept in memory take place. On the other hand, both groups showed an N400 effect associated with semantically related pictures (vs. unrelated), taken to reflect intact integration of semantic similarities in both dyslexic and control readers. We also found an electrophysiological effect of phonological priming in the N400 range - that is, an attenuated N400 to objects preceded by phonemic related primes vs. unrelated - while it showed a more widespread distributed and more pronounced over the right hemisphere in the dyslexics. Topographic differences between groups might have originated from a word form encoding process with different characteristics in dyslexics compared to control readers.

fMRI Syntactic and Lexical Repetition Effects Reveal the Initial Stages of Learning a New Language.

UNLABELLED: When learning a new language, we build brain networks to process and represent the acquired words and syntax and integrate these with existing language representations. It is an open question whether the same or different neural mechanisms are involved in learning and processing a novel language compared with the native language(s). Here we investigated the neural repetition effects of repeating known and novel word orders while human subjects were in the early stages of learning a new language. Combining a miniature language with a syntactic priming paradigm, we examined the neural correlates of language learning on-line using functional magnetic resonance imaging. In left inferior frontal gyrus and posterior temporal cortex, the repetition of novel syntactic structures led to repetition enhancement, whereas repetition of known structures resulted in repetition suppression. Additional verb repetition led to an increase in the syntactic repetition enhancement effect in language-related brain regions. Similarly, the repetition of verbs led to repetition enhancement effects in areas related to lexical and semantic processing, an effect that continued to increase in a subset of these regions. Repetition enhancement might reflect a mechanism to build and strengthen a neural network to process novel syntactic structures and lexical items. By contrast, the observed repetition suppression points to overlapping neural mechanisms for native and new language constructions when these have sufficient structural similarities. SIGNIFICANCE STATEMENT: Acquiring a second language entails learning how to interpret novel words and relations between words, and to integrate them with existing language knowledge. To investigate the brain mechanisms involved in this particularly human skill, we combined an artificial language learning task with a syntactic repetition paradigm. We show that the repetition of novel syntactic structures, as well as words in contexts, leads to repetition enhancement, whereas repetition of known structures results in repetition suppression. We thus propose that repetition enhancement might reflect a brain mechanism to build and strengthen a neural network to process novel syntactic regularities and novel words. Importantly, the results also indicate an overlap in neural mechanisms for native and new language constructions with sufficient structural similarities.

Too little or too much? Parafoveal preview benefits and parafoveal load costs in dyslexic adults.

Two different forms of parafoveal dysfunction have been hypothesized as core deficits of dyslexic individuals: reduced parafoveal preview benefits ("too little parafovea") and increased costs of parafoveal load ("too much parafovea"). We tested both hypotheses in a single eye-tracking experiment using a modified serial rapid automatized naming (RAN) task. Comparisons between dyslexic and non-dyslexic adults showed reduced parafoveal preview benefits in dyslexics, without increased costs of parafoveal load. Reduced parafoveal preview benefits were observed in a naming task, but not in a silent letter-finding task, indicating that the parafoveal dysfunction may be consequent to the overload with extracting phonological information from orthographic input. Our results suggest that dyslexics' parafoveal dysfunction is not based on strict visuo-attentional factors, but nevertheless they stress the importance of extra-phonological processing. Furthermore, evidence of reduced parafoveal preview benefits in dyslexia may help understand why serial RAN is an important reading predictor in adulthood.

Lexical and sublexical orthographic processing: an ERP study with skilled and dyslexic adult readers.

This ERP study investigated the cognitive nature of the P1-N1 components during orthographic processing. We used an implicit reading task with various types of stimuli involving different amounts of sublexical or lexical orthographic processing (words, pseudohomophones, pseudowords, nonwords, and symbols), and tested average and dyslexic readers. An orthographic regularity effect (pseudowords-nonwords contrast) was observed in the average but not in the dyslexic group. This suggests an early sensitivity to the dependencies among letters in word-forms that reflect orthographic structure, while the dyslexic brain apparently fails to be appropriately sensitive to these complex features. Moreover, in the adults the N1-response may already reflect lexical access: (i) the N1 was sensitive to the familiar vs. less familiar orthographic sequence contrast; (ii) and early effects of the phonological form (words-pseudohomophones contrast) were also found. Finally, the later N320 component was attenuated in the dyslexics, suggesting suboptimal processing in later stages of phonological analysis.

Musical phrase boundaries, wrap-up and the closure positive shift.

We investigated global integration (wrap-up) processes at the boundaries of musical phrases by comparing the effects of well and non-well formed phrases on event-related potentials time-locked to two boundary points: the onset and the offset of the boundary pause. The Closure Positive Shift, which is elicited at the boundary offset, was not modulated by the quality of phrase structure (well vs. non-well formed). In contrast, the boundary onset potentials showed different patterns for well and non-well formed phrases. Our results contribute to specify the functional meaning of the Closure Positive Shift in music, shed light on the large-scale structural integration of musical input, and raise new hypotheses concerning shared resources between music and language.

You know when: event-related potentials and theta/beta power indicate boundary prediction in music.

Neuroscientific and musicological approaches to music cognition indicate that listeners familiarized in the Western tonal tradition expect a musical phrase boundary at predictable time intervals. However, phrase boundary prediction processes in music remain untested. We analyzed event-related potentials (ERPs) and event-related induced power changes at the onset and offset of a boundary pause. We made comparisons with modified melodies, where the pause was omitted and filled by tones. The offset of the pause elicited a closure positive shift (CPS), indexing phrase boundary detection. The onset of the filling tones elicited significant increases in theta and beta powers. In addition, the P2 component was larger when the filling tones started than when they ended. The responses to boundary omission suggest that listeners expected to hear a boundary pause. Therefore, boundary prediction seems to coexist with boundary detection in music segmentation.

Phonological markers of information structure: an fMRI study.

In this fMRI study we investigate the neural correlates of information structure integration during sentence comprehension in Dutch. We looked into how prosodic cues (pitch accents) that signal the information status of constituents to the listener (new information) are combined with other types of information during the unification process. The difficulty of unifying the prosodic cues into overall sentence meaning was manipulated by constructing sentences in which the pitch accent did (focus-accent agreement), and sentences in which the pitch accent did not (focus-accent disagreement) match the expectations for focus constituents of the sentence. In case of a mismatch, the load on unification processes increases. Our results show two anatomically distinct effects of focus-accent disagreement, one located in the posterior left inferior frontal gyrus (LIFG, BA6/44), and one in the more anterior-ventral LIFG (BA 47/45). Our results confirm that information structure is taken into account during unification, and imply an important role for the LIFG in unification processes, in line with previous fMRI studies.