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Listening to speech in noise can require substantial mental effort, even among younger normal-hearing adults. The task-evoked pupil response (TEPR) has been shown to track the increased effort exerted to recognize words or sentences in increasing noise. However, few studies have examined the trajectory of listening effort across longer, more natural, stretches of speech, or the extent to which expectations about upcoming listening difficulty modulate the TEPR. Seventeen younger normal-hearing adults listened to 60-s-long audiobook passages, repeated three times in a row, at two different signal-to-noise ratios (SNRs) while pupil size was recorded. There was a significant interaction between SNR, repetition, and baseline pupil size on sustained listening effort. At lower baseline pupil sizes, potentially reflecting lower attention mobilization, TEPRs were more sustained in the harder SNR condition, particularly when attention mobilization remained low by the third presentation. At intermediate baseline pupil sizes, differences between conditions were largely absent, suggesting these listeners had optimally mobilized their attention for both SNRs. Lastly, at higher baseline pupil sizes, potentially reflecting over-mobilization of attention, the effect of SNR was initially reversed for the second and third presentations: participants initially appeared to disengage in the harder SNR condition, resulting in reduced TEPRs that recovered in the second half of the story. Together, these findings suggest that the unfolding of listening effort over time depends critically on the extent to which individuals have successfully mobilized their attention in anticipation of difficult listening conditions. 

Auditory cortical responses to speech obtained by magnetoencephalography (MEG) show robust speech tracking to the speaker's fundamental frequency in the high-gamma band (70–200 Hz), but little is currently known about whether such responses depend on the focus of selective attention. In this study 22 human subjects listened to concurrent, fixed-rate, speech from male and female speakers, and were asked to selectively attend to one speaker at a time, while their neural responses were recorded with MEG. The male speaker's pitch range coincided with the lower range of the high-gamma band, whereas the female speaker's higher pitch range had much less overlap, and only at the upper end of the high-gamma band. Neural responses were analyzed using the temporal response function (TRF) framework. As expected, the responses demonstrate robust speech tracking of the fundamental frequency in the high-gamma band, but only to the male's speech, with a peak latency of ~40 ms. Critically, the response magnitude depends on selective attention: the response to the male speech is significantly greater when male speech is attended than when it is not attended, under acoustically identical conditions. This is a clear demonstration that even very early cortical auditory responses are influenced by top-down, cognitive, neural processing mechanisms. 

Neural speech tracking has advanced our understanding of how our brains rapidly map an acoustic speech signal onto linguistic representations and ultimately meaning. It remains unclear, however, how speech intelligibility is related to the corresponding neural responses. Many studies addressing this question vary the level of intelligibility by manipulating the acoustic waveform, but this makes it difficult to cleanly disentangle the effects of intelligibility from underlying acoustical confounds. Here, using magnetoencephalography recordings, we study neural measures of speech intelligibility by manipulating intelligibility while keeping the acoustics strictly unchanged. Acoustically identical degraded speech stimuli (three-band noise-vocoded, ~20 s duration) are presented twice, but the second presentation is preceded by the original (nondegraded) version of the speech. This intermediate priming, which generates a “pop-out” percept, substantially improves the intelligibility of the second degraded speech passage. We investigate how intelligibility and acoustical structure affect acoustic and linguistic neural representations using multivariate temporal response functions (mTRFs). As expected, behavioral results confirm that perceived speech clarity is improved by priming. mTRFs analysis reveals that auditory (speech envelope and envelope onset) neural representations are not affected by priming but only by the acoustics of the stimuli (bottom–up driven). Critically, our findings suggest that segmentation of sounds into words emerges with better speech intelligibility, and most strongly at the later (~400 ms latency) word processing stage, in prefrontal cortex, in line with engagement of top–down mechanisms associated with priming. Taken together, our results show that word representations may provide some objective measures of speech comprehension.

 

Even though human experience unfolds continuously in time, it is not strictly linear; instead, it entails cascading processes building hierarchical cognitive structures. For instance, during speech perception, humans transform a continuously varying acoustic signal into phonemes, words, and meaning, and these levels all have distinct but interdependent temporal structures. Time-lagged regression usingtemporal response functions (TRFs) has recently emerged as a promising tool for disentangling electrophysiological brain responses related to such complex models of perception. Here, we introduce the Eelbrain Python toolkit, which makes this kind of analysis easy and accessible. We demonstrate its use, using continuous speech as a sample paradigm, with a freely available EEG dataset of audiobook listening. A companion GitHub repository provides the complete source code for the analysis, from raw data to group-level statistics. More generally, we advocate a hypothesis-driven approach in which the experimenter specifies a hierarchy of time-continuous representations that are hypothesized to have contributed to brain responses, and uses those as predictor variables for the electrophysiological signal. This is analogous to a multiple regression problem, but with the addition of a time dimension. TRF analysis decomposes the brain signal into distinct responses associated with the different predictor variables by estimating a multivariate TRF (mTRF), quantifying the influence of each predictor on brain responses as a function of time(-lags). This allows asking two questions about the predictor variables: (1) Is there a significant neural representation corresponding to this predictor variable? And if so, (2) what are the temporal characteristics of the neural response associated with it? Thus, different predictor variables can be systematically combined and evaluated to jointly model neural processing at multiple hierarchical levels. We discuss applications of this approach, including the potential for linking algorithmic/representational theories at different cognitive levels to brain responses through computational models with appropriate linking hypotheses. 

Abstract Cortical ischaemic strokes result in cognitive deficits depending on the area of the affected brain. However, we have demonstrated that difficulties with attention and processing speed can occur even with small subcortical infarcts. Symptoms appear independent of lesion location, suggesting they arise from generalized disruption of cognitive networks. Longitudinal studies evaluating directional measures of functional connectivity in this population are lacking. We evaluated six patients with minor stroke exhibiting cognitive impairment 6–8 weeks post-infarct and four age-similar controls. Resting-state magnetoencephalography data were collected. Clinical and imaging evaluations of both groups were repeated 6- and 12 months later. Network Localized Granger Causality was used to determine differences in directional connectivity between groups and across visits, which were correlated with clinical performance. Directional connectivity patterns remained stable across visits for controls. After the stroke, inter-hemispheric connectivity between the frontoparietal cortex and the non-frontoparietal cortex significantly increased between visits 1 and 2, corresponding to uniform improvement in reaction times and cognitive scores. Initially, the majority of functional links originated from non-frontal areas contralateral to the lesion, connecting to ipsilesional brain regions. By visit 2, inter-hemispheric connections, directed from the ipsilesional to the contralesional cortex significantly increased. At visit 3, patients demonstrating continued favourable cognitive recovery showed less reliance on these inter-hemispheric connections. These changes were not observed in those without continued improvement. Our findings provide supporting evidence that the neural basis of early post-stroke cognitive dysfunction occurs at the network level, and continued recovery correlates with the evolution of inter-hemispheric connectivity. 

Primary auditory cortex is a critical stage in the human auditory pathway, a gateway between subcortical and higher-level cortical areas. Receiving the output of all subcortical processing, it sends its output on to higher-level cortex. Non-invasive physiological recordings of primary auditory cortex using electroencephalography (EEG) and magnetoencephalography (MEG), however, may not have sufficient specificity to separate responses generated in primary auditory cortex from those generated in underlying subcortical areas or neighboring cortical areas. This limitation is important for investigations of effects of top-down processing (e.g., selective-attention-based) on primary auditory cortex: higher-level areas are known to be strongly influenced by top-down processes, but subcortical areas are often assumed to perform strictly bottom-up processing. Fortunately, recent advances have made it easier to isolate the neural activity of primary auditory cortex from other areas. In this perspective, we focus on time-locked responses to stimulus features in the high gamma band (70–150 Hz) and with early cortical latency (∼40 ms), intermediate between subcortical and higher-level areas. We review recent findings from physiological studies employing either repeated simple sounds or continuous speech, obtaining either a frequency following response (FFR) or temporal response function (TRF). The potential roles of top-down processing are underscored, and comparisons with invasive intracranial EEG (iEEG) and animal model recordings are made. We argue that MEG studies employing continuous speech stimuli may offer particular benefits, in that only a few minutes of speech generates robust high gamma responses from bilateral primary auditory cortex, and without measurable interference from subcortical or higher-level areas. 

Voice pitch carries linguistic as well as non-linguistic information. Previous studies have described cortical tracking of voice pitch in clean speech, with responses reflecting both pitch strength and pitch value. However, pitch is also a powerful cue for auditory stream segregation, especially when competing streams have pitch differing in fundamental frequency, as is the case when multiple speakers talk simultaneously. We therefore investigated how cortical speech pitch tracking is affected in the presence of a second, task-irrelevant speaker. We analyzed human magnetoencephalography (MEG) responses to continuous narrative speech, presented either as a single talker in a quiet background, or as a two-talker mixture of a male and a female speaker. In clean speech, voice pitch was associated with a right-dominant response, peaking at a latency of around 100 ms, consistent with previous EEG and ECoG results. The response tracked both the presence of pitch as well as the relative value of the speaker’s fundamental frequency. In the two-talker mixture, pitch of the attended speaker was tracked bilaterally, regardless of whether or not there was simultaneously present pitch in the speech of the irrelevant speaker. Pitch tracking for the irrelevant speaker was reduced: only the right hemisphere still significantly tracked pitch of the unattended speaker, and only during intervals in which no pitch was present in the attended talker’s speech. Taken together, these results suggest that pitch-based segregation of multiple speakers, at least as measured by macroscopic cortical tracking, is not entirely automatic but strongly dependent on selective attention. 

Objective: The Temporal Response Function (TRF) is a linear model of neural activity time-locked to continuous stimuli, including continuous speech. TRFs based on speech envelopes typically have distinct components that have provided remarkable insights into the cortical processing of speech. However, current methods may lead to less than reliable estimates of single-subject TRF components. Here, we compare two established methods, in TRF component estimation, and also propose novel algorithms that utilize prior knowledge of these components, bypassing the full TRF estimation. Methods: We compared two established algorithms, ridge and boosting, and two novel algorithms based on Subspace Pursuit (SP) and Expectation Maximization (EM), which directly estimate TRF components given plausible assumptions regarding component characteristics. Single-channel, multi-channel, and source-localized TRFs were fit on simulations and real magnetoencephalographic data. Performance metrics included model fit and component estimation accuracy. Results: Boosting and ridge have comparable performance in component estimation. The novel algorithms outperformed the others in simulations, but not on real data, possibly due to the plausible assumptions not actually being met. Ridge had slightly better model fits on real data compared to boosting, but also more spurious TRF activity. Conclusion: Results indicate that both smooth (ridge) and sparse (boosting) algorithms perform comparably at TRF component estimation. The SP and EM algorithms may be accurate, but rely on assumptions of component characteristics. Significance: This systematic comparison establishes the suitability of widely used and novel algorithms for estimating robust TRF components, which is essential for improved subject-specific investigations into the cortical processing of speech. 

Stroke patients with hemiparesis display decreased beta band (13–25 Hz) rolandic activity, correlating to impaired motor function. However, clinically, patients without significant weakness, with small lesions far from sensorimotor cortex, exhibit bilateral decreased motor dexterity and slowed reaction times. We investigate whether these minor stroke patients also display abnormal beta band activity. Magnetoencephalographic (MEG) data were collected from nine minor stroke patients (NIHSS < 4) without significant hemiparesis, at ~1 and ~6 months postinfarct, and eight age-similar controls. Rolandic relative beta power during matching tasks and resting state, and Beta Event Related (De)Synchronization (ERD/ERS) during button press responses were analyzed. Regardless of lesion location, patients had significantly reduced relative beta power and ERS compared to controls. Abnormalities persisted over visits, and were present in both ipsi- and contra-lesional hemispheres, consistent with bilateral impairments in motor dexterity and speed. Minor stroke patients without severe weakness display reduced rolandic beta band activity in both hemispheres, which may be linked to bilaterally impaired dexterity and processing speed, implicating global connectivity dysfunction affecting sensorimotor cortex independent of lesion location. Findings not only illustrate global network disruption after minor stroke, but suggest rolandic beta band activity may be a potential biomarker and treatment target, even for minor stroke patients with small lesions far from sensorimotor areas. 

Speech processing is highly incremental. It is widely accepted that human listeners continuously use the linguistic context to anticipate upcoming concepts, words, and phonemes. However, previous evidence supports two seemingly contradictory models of how a predictive context is integrated with the bottom-up sensory input: Classic psycholinguistic paradigms suggest a two-stage process, in which acoustic input initially leads to local, context-independent representations, which are then quickly integrated with contextual constraints. This contrasts with the view that the brain constructs a single coherent, unified interpretation of the input, which fully integrates available information across representational hierarchies, and thus uses contextual constraints to modulate even the earliest sensory representations. To distinguish these hypotheses, we tested magnetoencephalography responses to continuous narrative speech for signatures of local and unified predictive models. Results provide evidence that listeners employ both types of models in parallel. Two local context models uniquely predict some part of early neural responses, one based on sublexical phoneme sequences, and one based on the phonemes in the current word alone; at the same time, even early responses to phonemes also reflect a unified model that incorporates sentence-level constraints to predict upcoming phonemes. Neural source localization places the anatomical origins of the different predictive models in nonidentical parts of the superior temporal lobes bilaterally, with the right hemisphere showing a relative preference for more local models. These results suggest that speech processing recruits both local and unified predictive models in parallel, reconciling previous disparate findings. Parallel models might make the perceptual system more robust, facilitate processing of unexpected inputs, and serve a function in language acquisition.