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Simon, J. Z.

Publications and source records attributed to Simon, J. Z..

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Transformation from auditory to linguistic representations across auditory cortex is rapid and attention dependent for continuous speech

During speech perception, a central task of the auditory cortex is to analyze complex acoustic patterns to allow detection of the words that encode a linguistic message. It is generally thought that this process includes at least one intermediate, phonetic, level of representations [1-6], localized bilaterally in the superior temporal lobe [7-10]. Phonetic representations reflect a transition from acoustic to linguistic information, classifying acoustic patterns into linguistically meaningful units, which can serve as input to mechanisms that access abstract word representations [11-13]. While recent research has identified neural signals arising from successful recognition of individual words in continuous speech [14-17], no explicit neurophysiological signal has been found demonstrating the transition from acoustic/phonetic to symbolic, lexical representations. Here we report a response reflecting the incremental integration of phonetic information for word identification, dominantly localized to the left temporal lobe. The short response latency, approximately 110 ms relative to phoneme onset, suggests that phonetic information is used for lexical processing as soon as it becomes available. Responses also tracked word boundaries, confirming previous reports of immediate lexical segmentation [18,19]. These new results were further investigated using a cocktail-party paradigm [20,21] in which participants listened to a mix of two talkers, attending to one and ignoring the other. Analysis indicates neural lexical processing of only the attended, but not the unattended, speech stream. Thus, while responses to acoustic features reflect attention through selective amplification of attended speech, responses consistent with a lexical processing model reveal categorically selective processing.

neuroscience

Increased speech representation in older adults originates from early response in higher order auditory cortex

1Previous research has found that, paradoxically, while older adults have more difficulty comprehending speech in challenging circumstances than younger adults, their brain responses track the acoustic signal more robustly. Here we investigate this puzzle by using magnetoencephalography (MEG) source localization to determine the anatomical origin of this difference. Our results indicate that this robust tracking in older adults does not arise merely from having the same responses as younger adults but with larger amplitudes; instead they recruit additional regions, inferior to core auditory cortex, as part of an early response peak at ~ 30 ms relative to the acoustic signal.

neuroscience

The neural representation of missing speech and the influence of prior knowledge on cortical fidelity and latency

In naturally noisy listening conditions, for example at a cocktail party, noise disruptions may completely mask significant parts of a sentence, and yet listeners may still perceive the missing speech as being present. Here we demonstrate that dynamic speech-related auditory cortical activity, as measured by magnetoencephalography (MEG), which can ordinarily be used to directly reconstruct to the physical speech stimulus, can also be used to \"reconstruct\" acoustically missing speech. The extent to which this occurs depends on the extent that listeners are familiar with the missing speech, which is consistent with this neural activity being a dynamic representation of perceived speech even if acoustically absence. Our findings are two-fold: first, we find that when the speech is entirely acoustically absent, the acoustically absent speech can still be reconstructed with performance up to 25% of that of acoustically present speech without noise; and second, that this same expertise facilitates faster processing of natural speech by approximately 5 ms. Both effects disappear when listeners have no or very little prior experience with a given sentence. Our results suggest adaptive mechanisms of consolidation of detailed representations about speech, and the enabling of strong expectations this entails, as identifiable factors assisting automatic speech restoration over ecologically relevant timescales.

neuroscience

Neural Coding of Noisy and Reverberant Speech in Human Auditory Cortex

Speech communication in daily listening environments is complicated by the phenomenon of reverberation, wherein any sound reaching the ear is a mixture of the direct component from the source and multiple reflections off surrounding objects and the environment. The brain plays a central role in comprehending speech accompanied by such distortion, which, frequently, is further complicated by the presence of additional noise sources in the vicinity. Here, using magnetoencephalography (MEG) recordings from human subjects, we investigate the neural representation of speech in noisy, reverberant listening conditions as measured by phase-locked MEG responses to the slow temporal modulations of speech. Using systems-theoretic linear methods of stimulus encoding, we observe that the cortex maintains both distorted and distortion-free (cleaned) representations of speech. Also, we show that, while neural encoding of speech remains robust to additive noise in absence of reverberation, it is detrimentally affected by noise when present along with reverberation. Further, using linear methods of stimulus reconstruction, we show that theta-band neural responses are a likely candidate for the distortion free representation of speech, whereas delta band responses are more likely to carry non-speech specific information regarding the listening environment.

neuroscience

Real-Time Tracking of Selective Auditory Attention from M/EEG: A Bayesian Filtering Approach

Humans are able to identify and track a target speaker amid a cacophony of acoustic interference, which is often referred to as the cocktail party phenomenon. Results from several decades of studying this phenomenon have culminated in recent years in various promising attempts to decode the attentional state of a listener in a competing-speaker environment from non-invasive neuroimaging recordings such as magnetoencephalography (MEG) and electroencephalography (EEG). To this end, most existing approaches compute correlation-based measures by either regressing the features of each speech stream to the M/EEG channels (the decoding approach) or vice versa (the encoding approach). These procedures operate in an offline fashion, i.e., require the entire duration of the experiment and multiple trials to provide robust results. Therefore, they cannot be used in emerging applications such as smart hearing aid devices, where a single trial must be used in real-time to decode the attentional state. In this paper, we close this gap by developing an algorithmic pipeline for real-time decoding of the attentional state. Our proposed framework consists of three main modules: 1) Real-time and robust estimation of encoding or decoding coefficients, achieved by sparse adaptive filtering, 2) Extracting reliable markers of the attentional state, and thereby generalizing the widely-used correlation-based measures thereof, and 3) Devising a near real-time state-space estimator that translates the noisy and variable attention markers to robust and reliable estimates of the attentional state with minimal delay. Our proposed algorithms integrate various techniques including forgetting factor-based adaptive filtering, {ell}1-regularization, forward-backward splitting algorithms, fixed-lag smoothing, and Expectation Maximization. We validate the performance of our proposed framework using comprehensive simulations as well as application to experimentally acquired M/EEG data. Our results reveal that the proposed real-time algorithms perform nearly as accurate as the existing state-of-the-art offline techniques, while providing a significant degree of adaptivity, statistical robustness, and computational savings.

neuroscience

Representation of speech in noise in the aging midbrain and cortex: aging may dominate over hearing-loss

ObjectiveTo understand the effect of peripheral hearing loss on the representation of speech in noise in the aging midbrain and cortex.\n\nMethodsSubjects comprised 17 normal-hearing younger adults, 15 normal-hearing older adults and 14 hearing-impaired older adults. The midbrain response, measured with Frequency-Following Responses (FFRs), and the cortical response, measured with magnetoencephalography (MEG) responses, were recorded from subjects listening to speech in quiet and noise at varying signal to noise ratios (SNRs).\n\nResultsBoth groups of older listeners showed both weaker midbrain response amplitudes and overrepresentation of cortical responses compared to younger listeners. However, significant differences between the older groups were found in both midbrain-cortex relationships and in cortical processing durations, suggesting that hearing loss may alter reciprocal connections between lower and higher levels of the auditory pathway.\n\nConclusionsThe paucity of differences in midbrain or cortical responses between the two older groups suggest that age-related temporal processing deficits may contribute to older adults communication difficulties beyond what might be predicted from peripheral hearing loss alone.\n\nSignificanceClinical devices, such as hearing aids, should not ignore age-related temporal processing deficits in the design of algorithms to maximize user benefit.\n\nHIGHLIGHTSO_LIMild sensorineural hearing loss does not appear to significantly exacerbate already appreciable age-related deficits in midbrain speech-in-noise encoding.\nC_LIO_LIMild sensorineural hearing loss also does not appear to significantly exacerbate already appreciable age-related deficits in most measures of cortical speech-in-noise encoding.\nC_LIO_LICentral processing deficits caused by peripheral hearing loss in older adults are seen only in more subtle measures, including altered relationships between midbrain and cortex.\nC_LI

neuroscience

Neural source dynamics of brain responses to continuous stimuli: speech processing from acoustics to comprehension

Human experience often involves continuous sensory information that unfolds over time. This is true in particular for speech comprehension, where continuous acoustic signals are processed over seconds or even minutes. We show that brain responses to such continuous stimuli can be investigated in detail, for magnetoencephalography (MEG) data by combining linear kernel estimation with minimum norm source localization. Previous research has shown that the requirement to average data over many trials can be overcome by modeling the brain response as a linear convolution of the stimulus and a kernel, or response function, and estimating a kernel that predicts the response from the stimulus. However, such analysis has been typically restricted to sensor space. Here we demonstrate that this analysis can also be performed in neural source space. We first computed distributed minimum norm current source estimates for continuous MEG recordings, and then computed response functions for the current estimate at each source element, using the boosting algorithm with cross-validation. Permutation tests can then assess the significance of individual predictor variables as well as features of the corresponding spatio-temporal response functions. We demonstrate the viability of this technique by computing spatio-temporal response functions for speech stimuli, using predictor variables reflecting acoustic, lexical and semantic processing. Results indicate that processes related to comprehension of continuous speech can be differentiated anatomically as well as temporally: acoustic information engaged auditory cortex at short latencies, followed by responses over the central sulcus and inferior frontal gyrus, possibly related to somatosensory/motor cortex involvement in speech perception; lexical frequency was associated with a left-lateralized response in auditory cortex and subsequent bilateral frontal activity; and semantic composition was associated with bilateral temporal and frontal brain activity. We conclude that this technique can be used to study the neural processing of continuous stimuli in time and anatomical space with the millisecond temporal resolution of MEG. This suggests new avenues for analyzing neural processing of naturalistic stimuli, without the necessity of averaging over artificially short or truncated stimuli.

neuroscience

Functional significance of spectrotemporal response functions obtained using magnetoencephalography

The spectrotemporal response function (STRF) model of neural encoding quantitatively associates dynamic auditory neural (output) responses to a spectrogram-like representation of a dynamic (input) stimulus. STRFs were experimentally obtained via whole-head human cortical responses to dynamic auditory stimuli using magnetoencephalography (MEG). The stimuli employed consisted of unpredictable pure tones presented at a range of rates. The predictive power of the estimated STRFs was found to be comparable to those obtained from the cortical single and multiunit activity literature. The STRFs were also qualitatively consistent with those obtained from electrophysiological studies in animal models; in particular their local-field-potential-generated spectral distributions and multiunit-activity-generated temporal distributions. Comparison of these MEG STRFs with others obtained using natural speech and music stimuli reveal a general structure consistent with common baseline auditory processing, including evidence for a transition in low-level neural representations of natural speech by 100 ms, when an appropriately chosen stimulus representation was used. It is also demonstrated that MEG-based STRFs contain information similar to that obtained using classic auditory evoked potential based approaches, but with extended applications to long-duration, non-repeated stimuli.\n\nAuthor summaryThe spectrotemporal response function (STRF) model of linking dynamic acoustic stimuli to dynamic neural responses is applied to whole-head non-invasive magnetoencephalography (MEG) recordings of the human auditory cortex. MEG STRFs were consistent predictors of neural activity, quantitatively and qualitatively, by comparison to those obtained from animal models using local field potential or multiunit activity as neural responses. Comparison of STRFs using stimuli as diverse as tone clouds, natural speech, and music revealed a common structure consistent with shared baseline auditory processing, when an appropriately chosen stimulus representation was used.

neuroscience

Dynamic cortical representations of perceptual filling-in for missing acoustic rhythm

In the phenomenon of perceptual filling-in, missing sensory information can be reconstructed via interpolation from adjacent contextual cues by what is necessarily an endogenous, not yet well understood, neural process. In this investigation, sound stimuli were chosen to allow observation of fixed cortical oscillations driven by contextual (but missing) sensory input, thus entirely reflecting endogenous neural activity. The stimulus employed was a 5 Hz frequency-modulated tone, with brief masker probes (noise bursts) occasionally added. For half the probes, the rhythmic frequency modulation was moreover removed. Listeners reported whether the tone masked by each probe was perceived as being rhythmic or not. Time-frequency analysis of neural responses obtained by magnetoencephalography (MEG) shows that for maskers without the underlying acoustic rhythm, trials where rhythm was nonetheless perceived show higher evoked sustained rhythmic power than trials for which no rhythm was reported. The results support a model in which perceptual filling-in is aided by differential co-modulations of cortical activity at rates directly relevant to human speech communication. We propose that the presence of rhythmically-modulated neural dynamics predicts the subjective experience of a rhythmically modulated sound in real time, even when the perceptual experience is not supported by corresponding sensory data.

neuroscience

Cortical Representations of Speech in a Multi-talker Auditory Scene

The ability to parse a complex auditory scene into perceptual objects is facilitated by a hierarchical auditory system. Successive stages in the hierarchy transform an auditory scene of multiple overlapping sources, from peripheral tonotopically-based representations in the auditory nerve, into perceptually distinct auditory-objects based representation in auditory cortex. Here, using magnetoencephalography (MEG) recordings from human subjects, both men and women, we investigate how a complex acoustic scene consisting of multiple speech sources is represented in distinct hierarchical stages of auditory cortex. Using systems-theoretic methods of stimulus reconstruction, we show that the primary-like areas in auditory cortex contain dominantly spectro-temporal based representations of the entire auditory scene. Here, both attended and ignored speech streams are represented with almost equal fidelity, and a global representation of the full auditory scene with all its streams is a better candidate neural representation than that of individual streams being represented separately. In contrast, we also show that higher order auditory cortical areas represent the attended stream separately, and with significantly higher fidelity, than unattended streams. Furthermore, the unattended background streams are more faithfully represented as a single unsegregated background object rather than as separated objects. Taken together, these findings demonstrate the progression of the representations and processing of a complex acoustic scene up through the hierarchy of human auditory cortex.\n\nSignificance StatementUsing magnetoencephalography (MEG) recordings from human listeners in a simulated cocktail party environment, we investigate how a complex acoustic scene consisting of multiple speech sources is represented in separate hierarchical stages of auditory cortex. We show that the primary-like areas in auditory cortex use a dominantly spectro-temporal based representation of the entire auditory scene, with both attended and ignored speech streams represented with almost equal fidelity. In contrast, we show that higher order auditory cortical areas represent an attended speech stream separately from, and with significantly higher fidelity than, unattended speech streams. Furthermore, the unattended background streams are represented as a single undivided background object rather than as distinct background objects.

neuroscience