Parallel and distributed encoding of speech across human auditory cortex.

Speech perception is thought to rely on a cortical feedforward serial transformation of acoustic into linguistic representations. Using intracranial recordings across the entire human auditory cortex, electrocortical stimulation, and surgical ablation, we show that cortical processing across areas is not consistent with a serial hierarchical organization. Instead, response latency and receptive field analyses demonstrate parallel and distinct information processing in the primary and nonprimary auditory cortices. This functional dissociation was also observed where stimulation of the primary auditory cortex evokes auditory hallucination but does not distort or interfere with speech perception. Opposite effects were observed during stimulation of nonprimary cortex in superior temporal gyrus. Ablation of the primary auditory cortex does not affect speech perception. These results establish a distributed functional organization of parallel information processing throughout the human auditory cortex and demonstrate an essential independent role for nonprimary auditory cortex in speech processing.

Speaker-induced Suppression in EEG during a Naturalistic Reading and Listening Task.

Speaking elicits a suppressed neural response when compared with listening to others' speech, a phenomenon known as speaker-induced suppression (SIS). Previous research has focused on investigating SIS at constrained levels of linguistic representation, such as the individual phoneme and word level. Here, we present scalp EEG data from a dual speech perception and production task where participants read sentences aloud then listened to playback of themselves reading those sentences. Playback was separated into immediate repetition of the previous trial and randomized repetition of a former trial to investigate if forward modeling of responses during passive listening suppresses the neural response. Concurrent EMG was recorded to control for movement artifact during speech production. In line with previous research, ERP analyses at the sentence level demonstrated suppression of early auditory components of the EEG for production compared with perception. To evaluate whether linguistic abstractions (in the form of phonological feature tuning) are suppressed during speech production alongside lower-level acoustic information, we fit linear encoding models that predicted scalp EEG based on phonological features, EMG activity, and task condition. We found that phonological features were encoded similarly between production and perception. However, this similarity was only observed when controlling for movement by using the EMG response as an additional regressor. Our results suggest that SIS operates at a sensory representational level and is dissociated from higher order cognitive and linguistic processing that takes place during speech perception and production. We also detail some important considerations when analyzing EEG during continuous speech production.

Generalizable EEG Encoding Models with Naturalistic Audiovisual Stimuli.

In natural conversations, listeners must attend to what others are saying while ignoring extraneous background sounds. Recent studies have used encoding models to predict electroencephalography (EEG) responses to speech in noise-free listening situations, sometimes referred to as "speech tracking." Researchers have analyzed how speech tracking changes with different types of background noise. It is unclear, however, whether neural responses from acoustically rich, naturalistic environments with and without background noise can be generalized to more controlled stimuli. If encoding models for acoustically rich, naturalistic stimuli are generalizable to other tasks, this could aid in data collection from populations of individuals who may not tolerate listening to more controlled and less engaging stimuli for long periods of time. We recorded noninvasive scalp EEG while 17 human participants (8 male/9 female) listened to speech without noise and audiovisual speech stimuli containing overlapping speakers and background sounds. We fit multivariate temporal receptive field encoding models to predict EEG responses to pitch, the acoustic envelope, phonological features, and visual cues in both stimulus conditions. Our results suggested that neural responses to naturalistic stimuli were generalizable to more controlled datasets. EEG responses to speech in isolation were predicted accurately using phonological features alone, while responses to speech in a rich acoustic background were more accurate when including both phonological and acoustic features. Our findings suggest that naturalistic audiovisual stimuli can be used to measure receptive fields that are comparable and generalizable to more controlled audio-only stimuli.SIGNIFICANCE STATEMENT Understanding spoken language in natural environments requires listeners to parse acoustic and linguistic information in the presence of other distracting stimuli. However, most studies of auditory processing rely on highly controlled stimuli with no background noise, or with background noise inserted at specific times. Here, we compare models where EEG data are predicted based on a combination of acoustic, phonetic, and visual features in highly disparate stimuli-sentences from a speech corpus and speech embedded within movie trailers. We show that modeling neural responses to highly noisy, audiovisual movies can uncover tuning for acoustic and phonetic information that generalizes to simpler stimuli typically used in sensory neuroscience experiments.

Advances in human intracranial electroencephalography research, guidelines and good practices.

Since the second-half of the twentieth century, intracranial electroencephalography (iEEG), including both electrocorticography (ECoG) and stereo-electroencephalography (sEEG), has provided an intimate view into the human brain. At the interface between fundamental research and the clinic, iEEG provides both high temporal resolution and high spatial specificity but comes with constraints, such as the individual's tailored sparsity of electrode sampling. Over the years, researchers in neuroscience developed their practices to make the most of the iEEG approach. Here we offer a critical review of iEEG research practices in a didactic framework for newcomers, as well addressing issues encountered by proficient researchers. The scope is threefold: (i) review common practices in iEEG research, (ii) suggest potential guidelines for working with iEEG data and answer frequently asked questions based on the most widespread practices, and (iii) based on current neurophysiological knowledge and methodologies, pave the way to good practice standards in iEEG research. The organization of this paper follows the steps of iEEG data processing. The first section contextualizes iEEG data collection. The second section focuses on localization of intracranial electrodes. The third section highlights the main pre-processing steps. The fourth section presents iEEG signal analysis methods. The fifth section discusses statistical approaches. The sixth section draws some unique perspectives on iEEG research. Finally, to ensure a consistent nomenclature throughout the manuscript and to align with other guidelines, e.g., Brain Imaging Data Structure (BIDS) and the OHBM Committee on Best Practices in Data Analysis and Sharing (COBIDAS), we provide a glossary to disambiguate terms related to iEEG research.

Accuracy of omni-planar and surface casting of epileptiform activity for intracranial seizure localization.

OBJECTIVE: Intracranial electroencephalography (ICEEG) recordings are performed for seizure localization in medically refractory epilepsy. Signal quantifications such as frequency power can be projected as heatmaps on personalized three-dimensional (3D) reconstructed cortical surfaces to distill these complex recordings into intuitive cinematic visualizations. However, simultaneously reconciling deep recording locations and reliably tracking evolving ictal patterns remain significant challenges. METHODS: We fused oblique magnetic resonance imaging (MRI) slices along depth probe trajectories with cortical surface reconstructions and projected dynamic heatmaps using a simple mathematical metric of epileptiform activity (line-length). This omni-planar and surface casting of epileptiform activity approach (OPSCEA) thus illustrated seizure onset and spread among both deep and superficial locations simultaneously with minimal need for signal processing supervision. We utilized the approach on 41 patients at our center implanted with grid, strip, and/or depth electrodes for localizing medically refractory seizures. Peri-ictal data were converted into OPSCEA videos with multiple 3D brain views illustrating all electrode locations. Five people of varying expertise in epilepsy (medical student through epilepsy attending level) attempted to localize the seizure-onset zones. RESULTS: We retrospectively compared this approach with the original ICEEG study reports for validation. Accuracy ranged from 73.2% to 97.6% for complete or overlapping onset lobe(s), respectively, and ~56.1% to 95.1% for the specific focus (or foci). Higher answer certainty for a given case predicted better accuracy, and scorers had similar accuracy across different training levels. SIGNIFICANCE: In an era of increasing stereo-EEG use, cinematic visualizations fusing omni-planar and surface functional projections appear to provide a useful adjunct for interpreting complex intracranial recordings and subsequent surgery planning.

Human Superior Temporal Gyrus Organization of Spectrotemporal Modulation Tuning Derived from Speech Stimuli.

The human superior temporal gyrus (STG) is critical for speech perception, yet the organization of spectrotemporal processing of speech within the STG is not well understood. Here, to characterize the spatial organization of spectrotemporal processing of speech across human STG, we use high-density cortical surface field potential recordings while participants listened to natural continuous speech. While synthetic broad-band stimuli did not yield sustained activation of the STG, spectrotemporal receptive fields could be reconstructed from vigorous responses to speech stimuli. We find that the human STG displays a robust anterior-posterior spatial distribution of spectrotemporal tuning in which the posterior STG is tuned for temporally fast varying speech sounds that have relatively constant energy across the frequency axis (low spectral modulation) while the anterior STG is tuned for temporally slow varying speech sounds that have a high degree of spectral variation across the frequency axis (high spectral modulation). This work illustrates organization of spectrotemporal processing in the human STG, and illuminates processing of ethologically relevant speech signals in a region of the brain specialized for speech perception. SIGNIFICANCE STATEMENT: Considerable evidence has implicated the human superior temporal gyrus (STG) in speech processing. However, the gross organization of spectrotemporal processing of speech within the STG is not well characterized. Here we use natural speech stimuli and advanced receptive field characterization methods to show that spectrotemporal features within speech are well organized along the posterior-to-anterior axis of the human STG. These findings demonstrate robust functional organization based on spectrotemporal modulation content, and illustrate that much of the encoded information in the STG represents the physical acoustic properties of speech stimuli.

Processing of auditory feedback in perisylvian and insular cortex.

When we speak, we not only make movements with our mouth, lips, and tongue, but we also hear the sound of our own voice. Thus, speech production in the brain involves not only controlling the movements we make, but also auditory and sensory feedback. Auditory responses are typically suppressed during speech production compared to perception, but how this manifests across space and time is unclear. Here we recorded intracranial EEG in seventeen pediatric, adolescent, and adult patients with medication-resistant epilepsy who performed a reading/listening task to investigate how other auditory responses are modulated during speech production. We identified onset and sustained responses to speech in bilateral auditory cortex, with a selective suppression of onset responses during speech production. Onset responses provide a temporal landmark during speech perception that is redundant with forward prediction during speech production. Phonological feature tuning in these "onset suppression" electrodes remained stable between perception and production. Notably, the posterior insula responded at sentence onset for both perception and production, suggesting a role in multisensory integration during feedback control.

Acoustic structure of the five perceptual dimensions of timbre in orchestral instrument tones.

Attempts to relate the perceptual dimensions of timbre to quantitative acoustical dimensions have been tenuous, leading to claims that timbre is an emergent property, if measurable at all. Here, a three-pronged analysis shows that the timbre space of sustained instrument tones occupies 5 dimensions and that a specific combination of acoustic properties uniquely determines gestalt perception of timbre. Firstly, multidimensional scaling (MDS) of dissimilarity judgments generated a perceptual timbre space in which 5 dimensions were cross-validated and selected by traditional model comparisons. Secondly, subjects rated tones on semantic scales. A discriminant function analysis (DFA) accounting for variance of these semantic ratings across instruments and between subjects also yielded 5 significant dimensions with similar stimulus ordination. The dimensions of timbre space were then interpreted semantically by rotational and reflectional projection of the MDS solution into two DFA dimensions. Thirdly, to relate this final space to acoustical structure, the perceptual MDS coordinates of each sound were regressed with its joint spectrotemporal modulation power spectrum. Sound structures correlated significantly with distances in perceptual timbre space. Contrary to previous studies, most perceptual timbre dimensions are not the result of purely temporal or spectral features but instead depend on signature spectrotemporal patterns.

Human song: Separate neural pathways for melody and speech.

Does the brain perceive song as speech with melody? A new study using intracranial recordings and functional brain imaging in humans suggests that it does not. Instead, singing, instrumental music, and speech are represented by different neural populations.

Dataset size considerations for robust acoustic and phonetic speech encoding models in EEG.

In many experiments that investigate auditory and speech processing in the brain using electroencephalography (EEG), the experimental paradigm is often lengthy and tedious. Typically, the experimenter errs on the side of including more data, more trials, and therefore conducting a longer task to ensure that the data are robust and effects are measurable. Recent studies used naturalistic stimuli to investigate the brain's response to individual or a combination of multiple speech features using system identification techniques, such as multivariate temporal receptive field (mTRF) analyses. The neural data collected from such experiments must be divided into a training set and a test set to fit and validate the mTRF weights. While a good strategy is clearly to collect as much data as is feasible, it is unclear how much data are needed to achieve stable results. Furthermore, it is unclear whether the specific stimulus used for mTRF fitting and the choice of feature representation affects how much data would be required for robust and generalizable results. Here, we used previously collected EEG data from our lab using sentence stimuli and movie stimuli as well as EEG data from an open-source dataset using audiobook stimuli to better understand how much data needs to be collected for naturalistic speech experiments measuring acoustic and phonetic tuning. We found that the EEG receptive field structure tested here stabilizes after collecting a training dataset of approximately 200 s of TIMIT sentences, around 600 s of movie trailers training set data, and approximately 460 s of audiobook training set data. Thus, we provide suggestions on the minimum amount of data that would be necessary for fitting mTRFs from naturalistic listening data. Our findings are motivated by highly practical concerns when working with children, patient populations, or others who may not tolerate long study sessions. These findings will aid future researchers who wish to study naturalistic speech processing in healthy and clinical populations while minimizing participant fatigue and retaining signal quality.

A Platform for Cognitive Monitoring of Neurosurgical Patients During Hospitalization.

Intracranial recordings in epilepsy patients are increasingly utilized to gain insight into the electrophysiological mechanisms of human cognition. There are currently several practical limitations to conducting research with these patients, including patient and researcher availability and the cognitive abilities of patients, which limit the amount of task-related data that can be collected. Prior studies have synchronized clinical audio, video, and neural recordings to understand naturalistic behaviors, but these recordings are centered on the patient to understand their seizure semiology and thus do not capture and synchronize audiovisual stimuli experienced by patients. Here, we describe a platform for cognitive monitoring of neurosurgical patients during their hospitalization that benefits both patients and researchers. We provide the full specifications for this system and describe some example use cases in perception, memory, and sleep research. We provide results obtained from a patient passively watching TV as proof-of-principle for the naturalistic study of cognition. Our system opens up new avenues to collect more data per patient using real-world behaviors, affording new possibilities to conduct longitudinal studies of the electrophysiological basis of human cognition under naturalistic conditions.

The revolution will not be controlled: natural stimuli in speech neuroscience.

Humans have a unique ability to produce and consume rich, complex, and varied language in order to communicate ideas to one another. Still, outside of natural reading, the most common methods for studying how our brains process speech or understand language use only isolated words or simple sentences. Recent studies have upset this status quo by employing complex natural stimuli and measuring how the brain responds to language as it is used. In this article we argue that natural stimuli offer many advantages over simplified, controlled stimuli for studying how language is processed by the brain. Furthermore, the downsides of using natural language stimuli can be mitigated using modern statistical and computational techniques.

Regional gray matter variation in male-to-female transsexualism.

Gender identity-one's sense of being a man or a woman-is a fundamental perception experienced by all individuals that extends beyond biological sex. Yet, what contributes to our sense of gender remains uncertain. Since individuals who identify as transsexual report strong feelings of being the opposite sex and a belief that their sexual characteristics do not reflect their true gender, they constitute an invaluable model to understand the biological underpinnings of gender identity. We analyzed MRI data of 24 male-to-female (MTF) transsexuals not yet treated with cross-sex hormones in order to determine whether gray matter volumes in MTF transsexuals more closely resemble people who share their biological sex (30 control men), or people who share their gender identity (30 control women). Results revealed that regional gray matter variation in MTF transsexuals is more similar to the pattern found in men than in women. However, MTF transsexuals show a significantly larger volume of regional gray matter in the right putamen compared to men. These findings provide new evidence that transsexualism is associated with distinct cerebral pattern, which supports the assumption that brain anatomy plays a role in gender identity.

Volumetric and shape analysis of the thalamus in first-episode schizophrenia.

Thalamic abnormalities have been implicated in the pathogenesis of schizophrenia, although the majority of studies used chronic samples treated extensively with antipsychotics. Moreover, the clinical and neuropsychological correlates of these abnormalities remain largely unknown. Using high-resolution MR imaging and novel methods for shape analysis, we investigated thalamic subregions in 35 (25 M/10 F) first-episode schizophrenia patients compared with 33 (23 M/10 F) healthy volunteers. The right and left thalami were traced bilaterally on coronal brain slices and volumes were compared between groups. In addition, regional abnormalities were identified by comparing distances, measured from homologous thalamic surface points to the central core of each individual's surface model, between groups in 3D space. Patients had significantly less total thalamic volume compared with healthy volunteers. Statistical mapping demonstrated most pronounced shape abnormalities in the pulvinar; however, estimated false discovery rates in these regions were sizable. Smaller thalamus volume was significantly correlated with worse overall neuropsychological functioning and specific deficits were observed in the language, motor, and executive domains. There were no significant associations between thalamus volume and positive or negative symptoms. Our findings suggest that thalamic abnormalities are evident at the onset of a first episode of schizophrenia prior to extensive pharmacologic intervention and that these abnormalities have neuropsychological correlates.

Alterations in functional activation in euthymic bipolar disorder and schizophrenia during a working memory task.

Dysfunctions in prefrontal cortical networks are thought to underlie working memory (WM) impairments consistently observed in both subjects with bipolar disorder and schizophrenia. It remains unclear, however, whether patterns of WM-related hemodynamic responses are similar in bipolar and schizophrenia subjects compared to controls. We used fMRI to investigate differences in blood oxygen level dependent activation during a WM task in 21 patients with euthymic bipolar I, 20 patients with schizophrenia, and 38 healthy controls. Subjects were presented with four stimuli (abstract designs) followed by a fifth stimulus and required to recall whether the last stimulus was among the four presented previously. Task-related brain activity was compared within and across groups. All groups activated prefrontal cortex (PFC), primary and supplementary motor cortex, and visual cortex during the WM task. There were no significant differences in PFC activation between controls and euthymic bipolar subjects, but controls exhibited significantly increased activation (cluster-corrected P < 0.05) compared to patients with schizophrenia in prefrontal regions including dorsolateral prefrontal cortex (DLPFC). Although the bipolar group exhibited intermediate percent signal change in a functionally defined DLPFC region of interest with respect to the schizophrenia and control groups, effects remained significant only between patients with schizophrenia and controls. Schizophrenia and bipolar disorder may share some behavioral, diagnostic, and genetic features. Differences in the patterns of WM-related brain activity across groups, however, suggest some diagnostic specificity. Both patient groups showed some regional task-related hypoactivation compared to controls across the brain. Within DLPFC specifically, patients with schizophrenia exhibited more severe WM-related dysfunction than bipolar subjects.

DTNBP1 is associated with imaging phenotypes in schizophrenia.

Dystrobrevin binding protein 1 (DTNBP1) has been identified as putative schizophrenia susceptibility gene, but it remains unknown whether polymorphisms relate to altered cerebral structure. We examined relationships between a previously implicated DTNBP1 risk variant [P1578] and global and segmented brain tissue volumes and regional cortical thickness in schizophrenia (n = 62; 24 risk carriers) and healthy subjects (n = 42; 11 risk carriers), across ethnic groups and within Caucasians. Schizophrenia patients showed similar brain volumes, but significantly reduced brain-size adjusted gray matter and CSF volumes and cortical thinning in a widespread neocortical distribution compared to controls. DTNBP1 risk was found associated with reduced brain volume, but not with tissue sub-compartments. Cortical thickness, which was weakly associated with brain size, showed regional variations in association with genetic risk, although effects were dominated by highly significant genotype by diagnosis interactions over broad areas of cortex. Risk status was found associated with regional cortical thinning in patients, particularly in temporal networks, but with thickness increases in controls. DTNBP1 effects for brain volume and cortical thickness appear driven by different neurobiological processes. Smaller brain volumes observed in risk carriers may relate to previously reported DTNBP1/cognitive function relationships irrespective of diagnosis. Regional cortical thinning in patient, but not in control risk carriers, may suggest that DTNBP1 interacts with other schizophrenia-related risk factors to affect laminar thickness. Alternatively, DTNBP1 may influence neural processes for which individuals with thicker cortex are less vulnerable. Although DTNBP1 relates to cortical thinning in schizophrenia, morphological changes in the disorder are influenced by additional genetic and/or environmental factors.

Fiber tractography reveals disruption of temporal lobe white matter tracts in schizophrenia.

Diffusion tensor imaging (DTI) studies have demonstrated abnormal anisotropic diffusion in schizophrenia. However, examining data with low spatial resolution and/or a low number of gradient directions and limitations associated with analysis approaches sensitive to registration confounds may have contributed to mixed findings concerning the regional specificity and direction of results. This study examined three major white matter tracts connecting lateral and medial temporal lobe regions with neocortical association regions widely implicated in systems-level functional and structural disturbances in schizophrenia. Using DTIstudio, a previously validated regions of interest tractography method was applied to 30 direction diffusion weighted imaging data collected from demographically similar schizophrenia (n=23) and healthy control subjects (n=22). The diffusion tensor was computed at each voxel after intra-subject registration of diffusion-weighted images. Three-dimensional tract reconstruction was performed using the Fiber Assignment by Continuous Tracking (FACT) algorithm. Tractography results showed reduced fractional anisotropy (FA) of the arcuate fasciculi (AF) and inferior longitudinal fasciculi (ILF) in patients compared to controls. FA changes within the right ILF were negatively correlated with measures of thinking disorder. Reduced volume of the left AF was also observed in patients. These results, which avoid registration issues associated with voxel-based analyses of DTI data, support that fiber pathways connecting lateral and medial temporal lobe regions with neocortical regions are compromised in schizophrenia. Disruptions of connectivity within these pathways may potentially contribute to the disturbances of memory, language, and social cognitive processing that characterize the disorder.

Mean diffusivity: a biomarker for CSF-related disease and genetic liability effects in schizophrenia.

Mean diffusivity (MD), the rotationally invariant magnitude of water diffusion that is greater in cerebrospinal fluid (CSF) and smaller in organized brain tissue, has been suggested to reflect schizophrenia-associated cortical atrophy. Regional changes, associations with CSF, and the effects of genetic predisposition towards schizophrenia, however, remain uncertain. Six-direction diffusion tensor imaging DTI and high-resolution structural images were obtained from 26 schizophrenia patients, 36 unaffected first-degree patient relatives, 20 control subjects and 32 control relatives (N=114). Registration procedures aligned diffusion tensor imaging (DTI) data across imaging modalities. MD was averaged within lobar regions and the cingulate and superior temporal gyri. CSF volume and MD were highly correlated. Significant bilateral temporal, and superior temporal MD increases were observed in schizophrenia compared with unrelated control probands. First-degree relatives of schizophrenia probands showed larger MD measures compared with controls within bilateral superior temporal regions with CSF volume correction. Superior temporal lobe brain tissue deficits and proximal CSF enlargements are widely documented in schizophrenia. Larger MD indices in patients and their relatives may thus reflect similar pathophysiological mechanisms. However, persistence of regional MD effects after controlling for CSF volume, suggests that MD is a sensitive biological marker of disease and genetic liability, characterizing at least partially distinct aspects of brain structural integrity.

Spontaneous Neural Activity in the Superior Temporal Gyrus Recapitulates Tuning for Speech Features.

Background: Numerous studies have demonstrated that individuals exhibit structured neural activity in many brain regions during rest that is also observed during different tasks, however it is still not clear whether and how resting state activity patterns may relate to underlying tuning for specific stimuli. In the posterior superior temporal gyrus (STG), distinct neural activity patterns are observed during the perception of specific linguistic speech features. We hypothesized that spontaneous resting-state neural dynamics of the STG would be structured to reflect its role in speech perception, exhibiting an organization along speech features as seen during speech perception. Methods: Human cortical local field potentials were recorded from the superior temporal gyrus (STG) in 8 patients undergoing surgical treatment of epilepsy. Signals were recorded during speech perception and rest. Patterns of neural activity (high gamma power: 70-150 Hz) during rest, extracted with spatiotemporal principal component analysis, were compared to spatiotemporal neural responses to speech features during perception. Hierarchical clustering was applied to look for patterns in rest that corresponded to speech feature tuning. Results: Significant correlations were found between neural responses to speech features (sentence onsets, consonants, and vowels) and the spontaneous neural activity in the STG. Across subjects, these correlations clustered into five groups, demonstrating tuning for speech features-most robustly for acoustic onsets. These correlations were not seen in other brain areas, or during motor and spectrally-rotated speech control tasks. Conclusions: In this study, we present evidence that the RS structure of STG activity robustly recapitulates its stimulus-evoked response to acoustic onsets. Further, secondary patterns in RS activity appear to correlate with stimulus-evoked responses to speech features. The role of these spontaneous spatiotemporal activity patterns remains to be elucidated.

A Spatial Map of Onset and Sustained Responses to Speech in the Human Superior Temporal Gyrus.

To derive meaning from speech, we must extract multiple dimensions of concurrent information from incoming speech signals. That is, equally important to processing phonetic features is the detection of acoustic cues that give structure and context to the information we hear. How the brain organizes this information is unknown. Using data-driven computational methods on high-density intracranial recordings from 27 human participants, we reveal the functional distinction of neural responses to speech in the posterior superior temporal gyrus according to either onset or sustained response profiles. Though similar response types have been observed throughout the auditory system, we found novel evidence for a major spatial parcellation in which a distinct caudal zone detects acoustic onsets and a rostral-surround zone shows sustained, relatively delayed responses to ongoing speech stimuli. While posterior onset and anterior sustained responses are used substantially during natural speech perception, they are not limited to speech stimuli and are seen even for reversed or spectrally rotated speech. Single-electrode encoding of phonetic features in each zone depended upon whether the sound occurred at sentence onset, suggesting joint encoding of phonetic features and their temporal context. Onset responses in the caudal zone could accurately decode sentence and phrase onset boundaries, providing a potentially important internal mechanism for detecting temporal landmarks in speech and other natural sounds. These findings suggest that onset and sustained responses not only define the basic spatial organization of high-order auditory cortex but also have direct implications for how speech information is parsed in the cortex. VIDEO ABSTRACT.