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.
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Cortical Localization of the Auditory Temporal Response Function from MEG via Non-convex Optimization
The magnetoencephalography (MEG) response to continuous auditory stimuli, such as speech, is commonly described using a linear filter, the auditory temporal response function (TRF). Though components of the sensor level TRFs have been well characterized, the underlying neural sources responsible for these components are not well understood. In this work, we provide a unified framework for determining the TRFs of neural sources directly from the MEG data, by integrating the TRF and distributed forward source models into one, and casting the joint estimation task as a Bayesian optimization problem. Though the resulting problem emerges as non-convex, we propose efficient solutions that leverage recent advances in evidence maximization. We demonstrate the effectiveness of the resulting algorithm in both simulated and experimentally recorded MEG data from humans.
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- Award ID(s):
- 1734892
- NSF-PAR ID:
- 10088393
- Date Published:
- Journal Name:
- 2018 52nd Asilomar Conference on Signals, Systems, and Computers
- Page Range / eLocation ID:
- 373 - 378
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
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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.more » « less
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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.more » « less
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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.more » « less
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Identifying the directed connectivity that underlie networked activity between different cortical areas is critical for understanding the neural mechanisms behind sensory processing. Granger causality (GC) is widely used for this purpose in functional magnetic resonance imaging analysis, but there the temporal resolution is low, making it difficult to capture the millisecond-scale interactions underlying sensory processing. Magne- toencephalography (MEG) has millisecond resolution, but only provides low-dimensional sensor-level linear mixtures of neural sources, which makes GC inference challenging. Conventional methods proceed in two stages: First, cortical sources are estimated from MEG using a source localization technique, followed by GC inference among the estimated sources. However, the spatiotemporal biases in estimating sources propagate into the subsequent GC analysis stage, may result in both false alarms and missing true GC links. Here, we introduce the Network Localized Granger Causality (NLGC) inference paradigm, which models the source dynamics as latent sparse multivariate autoregressive processes and estimates their parameters directly from the MEG measurements, integrated with source localization, and employs the resulting parameter estimates to produce a precise statistical characterization of the detected GC links. We offer several theoretical and algorithmic innovations within NLGC and further examine its utility via comprehensive simulations and application to MEG data from an auditory task involving tone processing from both younger and older participants. Our simulation studies reveal that NLGC is markedly robust with respect to model mismatch, network size, and low signal-to-noise ratio, whereas the conventional two-stage methods result in high false alarms and mis-detections. We also demonstrate the advantages of NLGC in revealing the cortical network- level characterization of neural activity during tone processing and resting state by delineating task- and age-related connectivity changes.more » « less
- Free Publicly Accessible Full Text
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Accepted Manuscript1.0
- Conference Paper:
- https://doi.org/10.1109/ACSSC.2018.8645204
