Alpha and Beta Oscillations Differentially Support Word Production in a Rule-Switching Task.

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.

A tradeoff between acoustic and linguistic feature encoding in spoken language comprehension.

When we comprehend language from speech, the phase of the neural response aligns with particular features of the speech input, resulting in a phenomenon referred to as neural tracking. In recent years, a large body of work has demonstrated the tracking of the acoustic envelope and abstract linguistic units at the phoneme and word levels, and beyond. However, the degree to which speech tracking is driven by acoustic edges of the signal, or by internally-generated linguistic units, or by the interplay of both, remains contentious. In this study, we used naturalistic story-listening to investigate (1) whether phoneme-level features are tracked over and above acoustic edges, (2) whether word entropy, which can reflect sentence- and discourse-level constraints, impacted the encoding of acoustic and phoneme-level features, and (3) whether the tracking of acoustic edges was enhanced or suppressed during comprehension of a first language (Dutch) compared to a statistically familiar but uncomprehended language (French). We first show that encoding models with phoneme-level linguistic features, in addition to acoustic features, uncovered an increased neural tracking response; this signal was further amplified in a comprehended language, putatively reflecting the transformation of acoustic features into internally generated phoneme-level representations. Phonemes were tracked more strongly in a comprehended language, suggesting that language comprehension functions as a neural filter over acoustic edges of the speech signal as it transforms sensory signals into abstract linguistic units. We then show that word entropy enhances neural tracking of both acoustic and phonemic features when sentence- and discourse-context are less constraining. When language was not comprehended, acoustic features, but not phonemic ones, were more strongly modulated, but in contrast, when a native language is comprehended, phoneme features are more strongly modulated. Taken together, our findings highlight the flexible modulation of acoustic, and phonemic features by sentence and discourse-level constraint in language comprehension, and document the neural transformation from speech perception to language comprehension, consistent with an account of language processing as a neural filter from sensory to abstract representations.

Delta-Band Neural Responses to Individual Words Are Modulated by Sentence Processing.

To understand language, we need to recognize words and combine them into phrases and sentences. During this process, responses to the words themselves are changed. In a step toward understanding how the brain builds sentence structure, the present study concerns the neural readout of this adaptation. We ask whether low-frequency neural readouts associated with words change as a function of being in a sentence. To this end, we analyzed an MEG dataset by Schoffelen et al. (2019) of 102 human participants (51 women) listening to sentences and word lists, the latter lacking any syntactic structure and combinatorial meaning. Using temporal response functions and a cumulative model-fitting approach, we disentangled delta- and theta-band responses to lexical information (word frequency), from responses to sensory and distributional variables. The results suggest that delta-band responses to words are affected by sentence context in time and space, over and above entropy and surprisal. In both conditions, the word frequency response spanned left temporal and posterior frontal areas; however, the response appeared later in word lists than in sentences. In addition, sentence context determined whether inferior frontal areas were responsive to lexical information. In the theta band, the amplitude was larger in the word list condition ∼100 milliseconds in right frontal areas. We conclude that low-frequency responses to words are changed by sentential context. The results of this study show how the neural representation of words is affected by structural context and as such provide insight into how the brain instantiates compositionality in language.SIGNIFICANCE STATEMENT Human language is unprecedented in its combinatorial capacity: we are capable of producing and understanding sentences we have never heard before. Although the mechanisms underlying this capacity have been described in formal linguistics and cognitive science, how they are implemented in the brain remains to a large extent unknown. A large body of earlier work from the cognitive neuroscientific literature implies a role for delta-band neural activity in the representation of linguistic structure and meaning. In this work, we combine these insights and techniques with findings from psycholinguistics to show that meaning is more than the sum of its parts; the delta-band MEG signal differentially reflects lexical information inside and outside sentence structures.

Naturalistic Spoken Language Comprehension Is Supported by Alpha and Beta Oscillations.

Brain oscillations are prevalent in all species and are involved in numerous perceptual operations. α oscillations are thought to facilitate processing through the inhibition of task-irrelevant networks, while ß oscillations are linked to the putative reactivation of content representations. Can the proposed functional role of α and ß oscillations be generalized from low-level operations to higher-level cognitive processes? Here we address this question focusing on naturalistic spoken language comprehension. Twenty-two (18 female) Dutch native speakers listened to stories in Dutch and French while MEG was recorded. We used dependency parsing to identify three dependency states at each word: the number of (1) newly opened dependencies, (2) dependencies that remained open, and (3) resolved dependencies. We then constructed forward models to predict α and ß power from the dependency features. Results showed that dependency features predict α and ß power in language-related regions beyond low-level linguistic features. Left temporal, fundamental language regions are involved in language comprehension in α, while frontal and parietal, higher-order language regions, and motor regions are involved in ß. Critically, α- and ß-band dynamics seem to subserve language comprehension tapping into syntactic structure building and semantic composition by providing low-level mechanistic operations for inhibition and reactivation processes. Because of the temporal similarity of the α-ß responses, their potential functional dissociation remains to be elucidated. Overall, this study sheds light on the role of α and ß oscillations during naturalistic spoken language comprehension, providing evidence for the generalizability of these dynamics from perceptual to complex linguistic processes.SIGNIFICANCE STATEMENT It remains unclear whether the proposed functional role of α and ß oscillations in perceptual and motor function is generalizable to higher-level cognitive processes, such as spoken language comprehension. We found that syntactic features predict α and ß power in language-related regions beyond low-level linguistic features when listening to naturalistic speech in a known language. We offer experimental findings that integrate a neuroscientific framework on the role of brain oscillations as "building blocks" with spoken language comprehension. This supports the view of a domain-general role of oscillations across the hierarchy of cognitive functions, from low-level sensory operations to abstract linguistic processes.

The neural response at the fundamental frequency of speech is modulated by word-level acoustic and linguistic information.

Spoken language comprehension requires rapid and continuous integration of information, from lower-level acoustic to higher-level linguistic features. Much of this processing occurs in the cerebral cortex. Its neural activity exhibits, for instance, correlates of predictive processing, emerging at delays of a few 100 ms. However, the auditory pathways are also characterized by extensive feedback loops from higher-level cortical areas to lower-level ones as well as to subcortical structures. Early neural activity can therefore be influenced by higher-level cognitive processes, but it remains unclear whether such feedback contributes to linguistic processing. Here, we investigated early speech-evoked neural activity that emerges at the fundamental frequency. We analyzed EEG recordings obtained when subjects listened to a story read by a single speaker. We identified a response tracking the speaker's fundamental frequency that occurred at a delay of 11 ms, while another response elicited by the high-frequency modulation of the envelope of higher harmonics exhibited a larger magnitude and longer latency of about 18 ms with an additional significant component at around 40 ms. Notably, while the earlier components of the response likely originate from the subcortical structures, the latter presumably involves contributions from cortical regions. Subsequently, we determined the magnitude of these early neural responses for each individual word in the story. We then quantified the context-independent frequency of each word and used a language model to compute context-dependent word surprisal and precision. The word surprisal represented how predictable a word is, given the previous context, and the word precision reflected the confidence about predicting the next word from the past context. We found that the word-level neural responses at the fundamental frequency were predominantly influenced by the acoustic features: the average fundamental frequency and its variability. Amongst the linguistic features, only context-independent word frequency showed a weak but significant modulation of the neural response to the high-frequency envelope modulation. Our results show that the early neural response at the fundamental frequency is already influenced by acoustic as well as linguistic information, suggesting top-down modulation of this neural response.

Cortical Tracking of Surprisal during Continuous Speech Comprehension.

Speech comprehension requires rapid online processing of a continuous acoustic signal to extract structure and meaning. Previous studies on sentence comprehension have found neural correlates of the predictability of a word given its context, as well as of the precision of such a prediction. However, they have focused on single sentences and on particular words in those sentences. Moreover, they compared neural responses to words with low and high predictability, as well as with low and high precision. However, in speech comprehension, a listener hears many successive words whose predictability and precision vary over a large range. Here, we show that cortical activity in different frequency bands tracks word surprisal in continuous natural speech and that this tracking is modulated by precision. We obtain these results through quantifying surprisal and precision from naturalistic speech using a deep neural network and through relating these speech features to EEG responses of human volunteers acquired during auditory story comprehension. We find significant cortical tracking of surprisal at low frequencies, including the delta band as well as in the higher frequency beta and gamma bands, and observe that the tracking is modulated by the precision. Our results pave the way to further investigate the neurobiology of natural speech comprehension.

Modulation of Speech-in-Noise Comprehension Through Transcranial Current Stimulation With the Phase-Shifted Speech Envelope.

Neural activity tracks the envelope of a speech signal at latencies from 50 ms to 300 ms. Modulating this neural tracking through transcranial alternating current stimulation influences speech comprehension. Two important variables that can affect this modulation are the latency and the phase of the stimulation with respect to the sound. While previous studies have found an influence of both variables on speech comprehension, the interaction between both has not yet been measured. We presented 17 subjects with speech in noise coupled with simultaneous transcranial alternating current stimulation. The currents were based on the envelope of the target speech but shifted by different phases, as well as by two temporal delays of 100 ms and 250 ms. We also employed various control stimulations, and assessed the signal-to-noise ratio at which the subject understood half of the speech. We found that, at both latencies, speech comprehension is modulated by the phase of the current stimulation. However, the form of the modulation differed between the two latencies. Phase and latency of neurostimulation have accordingly distinct influences on speech comprehension. The different effects at the latencies of 100 ms and 250 ms hint at distinct neural processes for speech processing.