Neural Dynamics of the Processing of Speech Features: Evidence for a Progression of Features from Acoustic to Sentential Processing.

When we listen to speech, our brain's neurophysiological responses "track" its acoustic features, but it is less well understood how these auditory responses are modulated by linguistic content. Here, we recorded magnetoencephalography (MEG) responses while subjects listened to four types of continuous-speech-like passages: speech-envelope modulated noise, English-like non-words, scrambled words, and narrative passage. Temporal response function (TRF) analysis provides strong neural evidence for the emergent features of speech processing in cortex, from acoustics to higher-level linguistics, as incremental steps in neural speech processing. Critically, we show a stepwise hierarchical progression of progressively higher order features over time, reflected in both bottom-up (early) and top-down (late) processing stages. Linguistically driven top-down mechanisms take the form of late N400-like responses, suggesting a central role of predictive coding mechanisms at multiple levels. As expected, the neural processing of lower-level acoustic feature responses is bilateral or right lateralized, with left lateralization emerging only for lexical-semantic features. Finally, our results identify potential neural markers of the computations underlying speech perception and comprehension.

Neural Representations of Non-native Speech Reflect Proficiency and Interference from Native Language Knowledge.

Learning to process speech in a foreign language involves learning new representations for mapping the auditory signal to linguistic structure. Behavioral experiments suggest that even listeners that are highly proficient in a non-native language experience interference from representations of their native language. However, much of the evidence for such interference comes from tasks that may inadvertently increase the salience of native language competitors. Here we tested for neural evidence of proficiency and native language interference in a naturalistic story listening task. We studied electroencephalography responses of 39 native speakers of Dutch (14 male) to an English short story, spoken by a native speaker of either American English or Dutch. We modeled brain responses with multivariate temporal response functions, using acoustic and language models. We found evidence for activation of Dutch language statistics when listening to English, but only when it was spoken with a Dutch accent. This suggests that a naturalistic, monolingual setting decreases the interference from native language representations, whereas an accent in the listener's own native language may increase native language interference, by increasing the salience of the native language and activating native language phonetic and lexical representations. Brain responses suggest that such interference stems from words from the native language competing with the foreign language in a single word recognition system, rather than being activated in a parallel lexicon. We further found that secondary acoustic representations of speech (after 200 ms latency) decreased with increasing proficiency. This may reflect improved acoustic-phonetic models in more proficient listeners.Significance Statement Behavioral experiments suggest that native language knowledge interferes with foreign language listening, but such effects may be sensitive to task manipulations, as tasks that increase metalinguistic awareness may also increase native language interference. This highlights the need for studying non-native speech processing using naturalistic tasks. We measured neural responses unobtrusively while participants listened for comprehension and characterized the influence of proficiency at multiple levels of representation. We found that salience of the native language, as manipulated through speaker accent, affected activation of native language representations: significant evidence for activation of native language (Dutch) categories was only obtained when the speaker had a Dutch accent, whereas no significant interference was found to a speaker with a native (American) accent.

Eelbrain, a Python toolkit for time-continuous analysis with temporal response functions.

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 using temporal 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.

Cortical networks for recognition of speech with simultaneous talkers.

The relative contributions of superior temporal vs. inferior frontal and parietal networks to recognition of speech in a background of competing speech remain unclear, although the contributions themselves are well established. Here, we use fMRI with spectrotemporal modulation transfer function (ST-MTF) modeling to examine the speech information represented in temporal vs. frontoparietal networks for two speech recognition tasks with and without a competing talker. Specifically, 31 listeners completed two versions of a three-alternative forced choice competing speech task: "Unison" and "Competing", in which a female (target) and a male (competing) talker uttered identical or different phrases, respectively. Spectrotemporal modulation filtering (i.e., acoustic distortion) was applied to the two-talker mixtures and ST-MTF models were generated to predict brain activation from differences in spectrotemporal-modulation distortion on each trial. Three cortical networks were identified based on differential patterns of ST-MTF predictions and the resultant ST-MTF weights across conditions (Unison, Competing): a bilateral superior temporal (S-T) network, a frontoparietal (F-P) network, and a network distributed across cortical midline regions and the angular gyrus (M-AG). The S-T network and the M-AG network responded primarily to spectrotemporal cues associated with speech intelligibility, regardless of condition, but the S-T network responded to a greater range of temporal modulations suggesting a more acoustically driven response. The F-P network responded to the absence of intelligibility-related cues in both conditions, but also to the absence (presence) of target-talker (competing-talker) vocal pitch in the Competing condition, suggesting a generalized response to signal degradation. Task performance was best predicted by activation in the S-T and F-P networks, but in opposite directions (S-T: more activation = better performance; F-P: vice versa). Moreover, S-T network predictions were entirely ST-MTF mediated while F-P network predictions were ST-MTF mediated only in the Unison condition, suggesting an influence from non-acoustic sources (e.g., informational masking) in the Competing condition. Activation in the M-AG network was weakly positively correlated with performance and this relation was entirely superseded by those in the S-T and F-P networks. Regarding contributions to speech recognition, we conclude: (a) superior temporal regions play a bottom-up, perceptual role that is not qualitatively dependent on the presence of competing speech; (b) frontoparietal regions play a top-down role that is modulated by competing speech and scales with listening effort; and (c) performance ultimately relies on dynamic interactions between these networks, with ancillary contributions from networks not involved in speech processing per se (e.g., the M-AG network).

Auditory Word Comprehension Is Less Incremental in Isolated Words.

Partial speech input is often understood to trigger rapid and automatic activation of successively higher-level representations of words, from sound to meaning. Here we show evidence from magnetoencephalography that this type of incremental processing is limited when words are heard in isolation as compared to continuous speech. This suggests a less unified and automatic word recognition process than is often assumed. We present evidence from isolated words that neural effects of phoneme probability, quantified by phoneme surprisal, are significantly stronger than (statistically null) effects of phoneme-by-phoneme lexical uncertainty, quantified by cohort entropy. In contrast, we find robust effects of both cohort entropy and phoneme surprisal during perception of connected speech, with a significant interaction between the contexts. This dissociation rules out models of word recognition in which phoneme surprisal and cohort entropy are common indicators of a uniform process, even though these closely related information-theoretic measures both arise from the probability distribution of wordforms consistent with the input. We propose that phoneme surprisal effects reflect automatic access of a lower level of representation of the auditory input (e.g., wordforms) while the occurrence of cohort entropy effects is task sensitive, driven by a competition process or a higher-level representation that is engaged late (or not at all) during the processing of single words.

Cortical Tracking of Continuous Speech Under Bimodal Divided Attention.

Speech processing often occurs amid competing inputs from other modalities, for example, listening to the radio while driving. We examined the extent to which dividing attention between auditory and visual modalities (bimodal divided attention) impacts neural processing of natural continuous speech from acoustic to linguistic levels of representation. We recorded electroencephalographic (EEG) responses when human participants performed a challenging primary visual task, imposing low or high cognitive load while listening to audiobook stories as a secondary task. The two dual-task conditions were contrasted with an auditory single-task condition in which participants attended to stories while ignoring visual stimuli. Behaviorally, the high load dual-task condition was associated with lower speech comprehension accuracy relative to the other two conditions. We fitted multivariate temporal response function encoding models to predict EEG responses from acoustic and linguistic speech features at different representation levels, including auditory spectrograms and information-theoretic models of sublexical-, word-form-, and sentence-level representations. Neural tracking of most acoustic and linguistic features remained unchanged with increasing dual-task load, despite unambiguous behavioral and neural evidence of the high load dual-task condition being more demanding. Compared to the auditory single-task condition, dual-task conditions selectively reduced neural tracking of only some acoustic and linguistic features, mainly at latencies >200 ms, while earlier latencies were surprisingly unaffected. These findings indicate that behavioral effects of bimodal divided attention on continuous speech processing occur not because of impaired early sensory representations but likely at later cognitive processing stages. Crossmodal attention-related mechanisms may not be uniform across different speech processing levels.

Cortical tracking of voice pitch in the presence of multiple speakers depends on selective attention.

Voice pitch carries linguistic and 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 electroencephalography and electrocorticography results. The response tracked both the presence of pitch and the relative value of the speaker's fundamental frequency. In the two-talker mixture, the 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.

Bilaterally Reduced Rolandic Beta Band Activity in Minor Stroke Patients.

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.

Examining the context benefit in older adults: A combined behavioral-electrophysiologic word identification study.

When listening to degraded speech, listeners can use high-level semantic information to support recognition. The literature contains conflicting findings regarding older listeners' ability to benefit from semantic cues in recognizing speech, relative to younger listeners. Electrophysiologic (EEG) measures of lexical access (N400) often show that semantic context does not facilitate lexical access in older listeners; in contrast, auditory behavioral studies indicate that semantic context improves speech recognition in older listeners as much as or more than in younger listeners. Many behavioral studies of aging and the context benefit have employed signal degradation or alteration, whereas this stimulus manipulation has been absent in the EEG literature, a possible reason for the inconsistencies between studies. Here we compared the context benefit as a function of age and signal type, using EEG combined with behavioral measures. Non-native accent, a common form of signal alteration which many older adults report as a challenge in daily speech recognition, was utilized for testing. The stimuli included English sentences produced by native speakers of English and Spanish, containing target words differing in cloze probability. Listeners performed a word identification task while 32-channel cortical responses were recorded. Results show that older adults' word identification performance was poorer in the low-predictability and non-native talker conditions than the younger adults', replicating earlier behavioral findings. However, older adults did not show reduction or delay in the average N400 response as compared to younger listeners, suggesting no age-related reduction in predictive processing capability. Potential sources for discrepancies in the prior literature are discussed.

Parallel processing in speech perception with local and global representations of linguistic context.

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.

Neural Markers of Speech Comprehension: Measuring EEG Tracking of Linguistic Speech Representations, Controlling the Speech Acoustics.

When listening to speech, our brain responses time lock to acoustic events in the stimulus. Recent studies have also reported that cortical responses track linguistic representations of speech. However, tracking of these representations is often described without controlling for acoustic properties. Therefore, the response to these linguistic representations might reflect unaccounted acoustic processing rather than language processing. Here, we evaluated the potential of several recently proposed linguistic representations as neural markers of speech comprehension. To do so, we investigated EEG responses to audiobook speech of 29 participants (22 females). We examined whether these representations contribute unique information over and beyond acoustic neural tracking and each other. Indeed, not all of these linguistic representations were significantly tracked after controlling for acoustic properties. However, phoneme surprisal, cohort entropy, word surprisal, and word frequency were all significantly tracked over and beyond acoustic properties. We also tested the generality of the associated responses by training on one story and testing on another. In general, the linguistic representations are tracked similarly across different stories spoken by different readers. These results suggests that these representations characterize the processing of the linguistic content of speech.SIGNIFICANCE STATEMENT For clinical applications, it would be desirable to develop a neural marker of speech comprehension derived from neural responses to continuous speech. Such a measure would allow for behavior-free evaluation of speech understanding; this would open doors toward better quantification of speech understanding in populations from whom obtaining behavioral measures may be difficult, such as young children or people with cognitive impairments, to allow better targeted interventions and better fitting of hearing devices.

Does signal reduction imply predictive coding in models of spoken word recognition?

Pervasive behavioral and neural evidence for predictive processing has led to claims that language processing depends upon predictive coding. Formally, predictive coding is a computational mechanism where only deviations from top-down expectations are passed between levels of representation. In many cognitive neuroscience studies, a reduction of signal for expected inputs is taken as being diagnostic of predictive coding. In the present work, we show that despite not explicitly implementing prediction, the TRACE model of speech perception exhibits this putative hallmark of predictive coding, with reductions in total lexical activation, total lexical feedback, and total phoneme activation when the input conforms to expectations. These findings may indicate that interactive activation is functionally equivalent or approximant to predictive coding or that caution is warranted in interpreting neural signal reduction as diagnostic of predictive coding.

Poststroke acute dysexecutive syndrome, a disorder resulting from minor stroke due to disruption of network dynamics.

Stroke patients with small central nervous system infarcts often demonstrate an acute dysexecutive syndrome characterized by difficulty with attention, concentration, and processing speed, independent of lesion size or location. We use magnetoencephalography (MEG) to show that disruption of network dynamics may be responsible. Nine patients with recent minor strokes and eight age-similar controls underwent cognitive screening using the Montreal cognitive assessment (MoCA) and MEG to evaluate differences in cerebral activation patterns. During MEG, subjects participated in a visual picture-word matching task. Task complexity was increased as testing progressed. Cluster-based permutation tests determined differences in activation patterns within the visual cortex, fusiform gyrus, and lateral temporal lobe. At visit 1, MoCA scores were significantly lower for patients than controls (median [interquartile range] = 26.0 [4] versus 29.5 [3], P = 0.005), and patient reaction times were increased. The amplitude of activation was significantly lower after infarct and demonstrated a pattern of temporal dispersion independent of stroke location. Differences were prominent in the fusiform gyrus and lateral temporal lobe. The pattern suggests that distributed network dysfunction may be responsible. Additionally, controls were able to modulate their cerebral activity based on task difficulty. In contrast, stroke patients exhibited the same low-amplitude response to all stimuli. Group differences remained, to a lesser degree, 6 mo later; while MoCA scores and reaction times improved for patients. This study suggests that function is a globally distributed property beyond area-specific functionality and illustrates the need for longer-term follow-up studies to determine whether abnormal activation patterns ultimately resolve or another mechanism underlies continued recovery.

Continuous speech processing.

Speech processing in the human brain is grounded in non-specific auditory processing in the general mammalian brain, but relies on human-specific adaptations for processing speech and language. For this reason, many recent neurophysiological investigations of speech processing have turned to the human brain, with an emphasis on continuous speech. Substantial progress has been made using the phenomenon of "neural speech tracking", in which neurophysiological responses time-lock to the rhythm of auditory (and other) features in continuous speech. One broad category of investigations concerns the extent to which speech tracking measures are related to speech intelligibility, which has clinical applications in addition to its scientific importance. Recent investigations have also focused on disentangling different neural processes that contribute to speech tracking. The two lines of research are closely related, since processing stages throughout auditory cortex contribute to speech comprehension, in addition to subcortical processing and higher order and attentional processes.

Neural speech restoration at the cocktail party: Auditory cortex recovers masked speech of both attended and ignored speakers.

Humans are remarkably skilled at listening to one speaker out of an acoustic mixture of several speech sources. Two speakers are easily segregated, even without binaural cues, but the neural mechanisms underlying this ability are not well understood. One possibility is that early cortical processing performs a spectrotemporal decomposition of the acoustic mixture, allowing the attended speech to be reconstructed via optimally weighted recombinations that discount spectrotemporal regions where sources heavily overlap. Using human magnetoencephalography (MEG) responses to a 2-talker mixture, we show evidence for an alternative possibility, in which early, active segregation occurs even for strongly spectrotemporally overlapping regions. Early (approximately 70-millisecond) responses to nonoverlapping spectrotemporal features are seen for both talkers. When competing talkers' spectrotemporal features mask each other, the individual representations persist, but they occur with an approximately 20-millisecond delay. This suggests that the auditory cortex recovers acoustic features that are masked in the mixture, even if they occurred in the ignored speech. The existence of such noise-robust cortical representations, of features present in attended as well as ignored speech, suggests an active cortical stream segregation process, which could explain a range of behavioral effects of ignored background speech.

High gamma cortical processing of continuous speech in younger and older listeners.

Neural processing along the ascending auditory pathway is often associated with a progressive reduction in characteristic processing rates. For instance, the well-known frequency-following response (FFR) of the auditory midbrain, as measured with electroencephalography (EEG), is dominated by frequencies from ∼100 Hz to several hundred Hz, phase-locking to the acoustic stimulus at those frequencies. In contrast, cortical responses, whether measured by EEG or magnetoencephalography (MEG), are typically characterized by frequencies of a few Hz to a few tens of Hz, time-locking to acoustic envelope features. In this study we investigated a crossover case, cortically generated responses time-locked to continuous speech features at FFR-like rates. Using MEG, we analyzed responses in the high gamma range of 70-200 Hz to continuous speech using neural source-localized reverse correlation and the corresponding temporal response functions (TRFs). Continuous speech stimuli were presented to 40 subjects (17 younger, 23 older adults) with clinically normal hearing and their MEG responses were analyzed in the 70-200 Hz band. Consistent with the relative insensitivity of MEG to many subcortical structures, the spatiotemporal profile of these response components indicated a cortical origin with ∼40 ms peak latency and a right hemisphere bias. TRF analysis was performed using two separate aspects of the speech stimuli: a) the 70-200 Hz carrier of the speech, and b) the 70-200 Hz temporal modulations in the spectral envelope of the speech stimulus. The response was dominantly driven by the envelope modulation, with a much weaker contribution from the carrier. Age-related differences were also analyzed to investigate a reversal previously seen along the ascending auditory pathway, whereby older listeners show weaker midbrain FFR responses than younger listeners, but, paradoxically, have stronger cortical low frequency responses. In contrast to both these earlier results, this study did not find clear age-related differences in high gamma cortical responses to continuous speech. Cortical responses at FFR-like frequencies shared some properties with midbrain responses at the same frequencies and with cortical responses at much lower frequencies.

Neuro-current response functions: A unified approach to MEG source analysis under the continuous stimuli paradigm.

Characterizing the neural dynamics underlying sensory processing is one of the central areas of investigation in systems and cognitive neuroscience. Neuroimaging techniques such as magnetoencephalography (MEG) and Electroencephalography (EEG) have provided significant insights into the neural processing of continuous stimuli, such as speech, thanks to their high temporal resolution. Existing work in the context of auditory processing suggests that certain features of speech, such as the acoustic envelope, can be used as reliable linear predictors of the neural response manifested in M/EEG. The corresponding linear filters are referred to as temporal response functions (TRFs). While the functional roles of specific components of the TRF are well-studied and linked to behavioral attributes such as attention, the cortical origins of the underlying neural processes are not as well understood. In this work, we address this issue by estimating a linear filter representation of cortical sources directly from neuroimaging data in the context of continuous speech processing. To this end, we introduce Neuro-Current Response Functions (NCRFs), a set of linear filters, spatially distributed throughout the cortex, that predict the cortical currents giving rise to the observed ongoing MEG (or EEG) data in response to continuous speech. NCRF estimation is cast within a Bayesian framework, which allows unification of the TRF and source estimation problems, and also facilitates the incorporation of prior information on the structural properties of the NCRFs. To generalize this analysis to M/EEG recordings which lack individual structural magnetic resonance (MR) scans, NCRFs are extended to free-orientation dipoles and a novel regularizing scheme is put forward to lessen reliance on fine-tuned coordinate co-registration. We present a fast estimation algorithm, which we refer to as the Champ-Lasso algorithm, by leveraging recent advances in optimization, and demonstrate its utility through application to simulated and experimentally recorded MEG data under auditory experiments. Our simulation studies reveal significant improvements over existing methods that typically operate in a two-stage fashion, in terms of spatial resolution, response function reconstruction, and recovering dipole orientations. The analysis of experimentally-recorded MEG data without MR scans corroborates existing findings, but also delineates the distinct cortical distribution of the underlying neural processes at high spatiotemporal resolution. In summary, we provide a principled modeling and estimation paradigm for MEG source analysis tailored to extracting the cortical origin of electrophysiological responses to continuous stimuli.

The temporal dynamics of structure and content in sentence comprehension: Evidence from fMRI-constrained MEG.

Humans have a striking capacity to combine words into sentences that express new meanings. Previous research has identified key brain regions involved in this capacity, but little is known about the time course of activity in these regions, as hemodynamic methods such as fMRI provide little insight into temporal dynamics of neural activation. We performed an MEG experiment to elucidate the temporal dynamics of structure and content processing within four brain regions implicated by fMRI data from the same experiment: the temporo-parietal junction (TPJ), the posterior temporal lobe (PTL), the anterior temporal lobe (ATL), and the anterior inferior frontal gyrus (IFG). The TPJ showed increased activity for both structure and content near the end of the sentence, consistent with a role in incremental interpretation of event semantics. The PTL, a region not often associated with core aspects of syntax, showed a strong early effect of structure, consistent with predictive parsing models, and both structural and semantic context effects on function words. These results provide converging evidence that the PTL plays an important role in lexicalized syntactic processing. The ATL and IFG, regions traditionally associated with syntax, showed minimal effects of sentence structure. The ATL, PTL and IFG all showed effects of semantic content: increased activation for real words relative to nonwords. Our fMRI-guided MEG investigation therefore helps identify syntactic and semantic aspects of sentence comprehension in the brain in both spatial and temporal dimensions.

Rapid Transformation from Auditory to Linguistic Representations of 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 [1]. It is generally thought that this process includes at least one intermediate, phonetic, level of representations [2-6], localized bilaterally in the superior temporal lobe [7-9]. 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 [10, 11]. While recent research has identified neural signals arising from successful recognition of individual words in continuous speech [12-15], no explicit neurophysiological signal has been found demonstrating the transition from acoustic and/or 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 114 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 [16, 17]. These new results were further investigated using a cocktail-party paradigm [18, 19] 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.

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.