Responses In Auditory Cortex Research Articles

Adaptation of high-gamma responses in human auditory association cortex.

This study investigates adaptation of high-frequency cortical responses [>60 Hz; high-gamma (HG)] to simple and complex sounds in human nonprimary auditory cortex. We used intracranial electrocorticographic recordings to measure event-related changes in HG power as a function of stimulus probability. Tone and speech stimuli were presented in a series of traditional oddball and control paradigms. We hypothesized that HG power attenuates with stimulus repetition over multiple concurrent time scales in auditory association cortex. Time-frequency analyses were performed to identify auditory-responsive sites. Single-trial analyses and quantitative modeling were then used to measure trial-to-trial changes in HG power for high (frequent), low (infrequent), and equal (control) stimulus probabilities. Results show strong reduction of HG responses to frequently repeated tones and speech, with no differences in responses to infrequent and equal-probability stimuli. Adaptation of the HG frequent response, and not stimulus-acoustic differences or deviance-detection enhancement effects, accounted for the differential responses observed for frequent and infrequent sounds. Adaptation of HG responses showed a rapid onset (less than two trials) with slower adaptation between consecutive, repeated trials (2-10 s) and across trials in a stimulus block (∼7 min). The auditory-evoked N100 response also showed repetition-related adaptation, consistent with previous human scalp and animal single-unit recordings. These findings indicate that HG responses are highly sensitive to the regularities of simple and complex auditory events and show adaptation on multiple concurrent time scales in human auditory association cortex.

Speech sound discrimination training improves auditory cortex responses in a rat model of autism.

Children with autism often have language impairments and degraded cortical responses to speech. Extensive behavioral interventions can improve language outcomes and cortical responses. Prenatal exposure to the antiepileptic drug valproic acid (VPA) increases the risk for autism and language impairment. Prenatal exposure to VPA also causes weaker and delayed auditory cortex responses in rats. In this study, we document speech sound discrimination ability in VPA exposed rats and document the effect of extensive speech training on auditory cortex responses. VPA exposed rats were significantly impaired at consonant, but not vowel, discrimination. Extensive speech training resulted in both stronger and faster anterior auditory field (AAF) responses compared to untrained VPA exposed rats, and restored responses to control levels. This neural response improvement generalized to non-trained sounds. The rodent VPA model of autism may be used to improve the understanding of speech processing in autism and contribute to improving language outcomes.

Neuroimaging paradigms for tonotopic mapping (I): The influence of sound stimulus type

Although a consensus is emerging in the literature regarding the tonotopic organisation of auditory cortex in humans, previous studies employed a vast array of different neuroimaging protocols. In the present functional magnetic resonance imaging (fMRI) study, we made a systematic comparison between stimulus protocols involving jittered tone sequences with either a narrowband, broadband, or sweep character in order to evaluate their suitability for the purpose of tonotopic mapping. Data-driven analysis techniques were used to identify cortical maps related to sound-evoked activation and tonotopic frequency tuning. Principal component analysis (PCA) was used to extract the dominant response patterns in each of the three protocols separately, and generalised canonical correlation analysis (CCA) to assess the commonalities between protocols. Generally speaking, all three types of stimuli evoked similarly distributed response patterns and resulted in qualitatively similar tonotopic maps. However, quantitatively, we found that broadband stimuli are most efficient at evoking responses in auditory cortex, whereas narrowband and sweep stimuli offer the best sensitivity to differences in frequency tuning. Based on these results, we make several recommendations regarding optimal stimulus protocols, and conclude that an experimental design based on narrowband stimuli provides the best sensitivity to frequency-dependent responses to determine tonotopic maps. We forward that the resulting protocol is suitable to act as a localiser of tonotopic cortical fields in individuals, or to make quantitative comparisons between maps in dedicated tonotopic mapping studies.

Functional organization of human auditory cortex: Investigation of response latencies through direct recordings

The model for functional organization of human auditory cortex is in part based on findings in non-human primates, where the auditory cortex is hierarchically delineated into core, belt and parabelt fields. This model envisions that core cortex directly projects to belt, but not to parabelt, whereas belt regions are a major source of direct input for auditory parabelt. In humans, the posteromedial portion of Heschl's gyrus (HG) represents core auditory cortex, whereas the anterolateral portion of HG and the posterolateral superior temporal gyrus (PLST) are generally interpreted as belt and parabelt, respectively. In this scheme, response latencies can be hypothesized to progress in serial fashion from posteromedial to anterolateral HG to PLST. We examined this hypothesis by comparing response latencies to multiple stimuli, measured across these regions using simultaneous intracranial recordings in neurosurgical patients. Stimuli were 100 Hz click trains and the speech syllable /da/. Response latencies were determined by examining event-related band power in the high gamma frequency range. The earliest responses in auditory cortex occurred in posteromedial HG. Responses elicited from sites in anterolateral HG were neither earlier in latency from sites on PLST, nor more robust. Anterolateral HG and PLST exhibited some preference for speech syllable stimuli compared to click trains. These findings are not supportive of a strict serial model envisioning principal flow of information along HG to PLST. In contrast, data suggest that a portion of PLST may represent a relatively early stage in the auditory cortical hierarchy.

Perceptual Gap Detection Is Mediated by Gap Termination Responses in Auditory Cortex

Understanding speech in the presence of background noise often becomes increasingly difficult with age. These age-related speech processing deficits reflect impairments in temporal acuity. Gap detection is a model for temporal acuity in speech processing in which a gap inserted in white noise acts as a cue that attenuates subsequent startle responses. Lesion studies have shown that auditory cortex is necessary for the detection of brief gaps, and auditory cortical neurons respond to the end of the gap with a characteristic burst of spikes called the gap termination response (GTR). However, it remains unknown whether and how the GTR plays a causal role in gap detection. Wetested this by optogenetically suppressing the activity of somatostatin- or parvalbumin-expressing inhibitory interneurons, or CaMKII-expressing excitatory neurons, in auditory cortex of behaving mice during specific epochs of a gap detection protocol. Suppressing interneuron activity during the postgap interval enhanced gap detection. Suppressing excitatory cells during this interval attenuated gap detection. Suppressing activity preceding the gap had the opposite behavioral effects, whereas prolonged suppression across both intervals had no effect on gap detection. In addition to confirming cortical involvement, we demonstrate here for the first time a causal relationship between postgap neural activity and perceptual gap detection. Furthermore, our results suggest that gap detection involves an ongoing comparison of pre- and postgap spiking activity. Finally, we propose a simple yet biologically plausible neural circuit that reproduces each of these neural and behavioral results.

Регистрация и анализ точных частотных ЭЭГ/МЭГ откликов аудиторной коры головного мозга человека в ответ на монауральную стимуляцию звуком с фиксированными частотными составляющими

Регистрация и анализ точных частотных ЭЭГ/МЭГ откликов аудиторной коры головного мозга человека в ответ на монауральную стимуляцию звуком с фиксированными частотными составляющими

Stimulating the lip motor cortex with transcranial magnetic stimulation.

Transcranial magnetic stimulation (TMS) has proven to be a useful tool in investigating the role of the articulatory motor cortex in speech perception. Researchers have used single-pulse and repetitive TMS to stimulate the lip representation in the motor cortex. The excitability of the lip motor representation can be investigated by applying single TMS pulses over this cortical area and recording TMS-induced motor evoked potentials (MEPs) via electrodes attached to the lip muscles (electromyography; EMG). Larger MEPs reflect increased cortical excitability. Studies have shown that excitability increases during listening to speech as well as during viewing speech-related movements. TMS can be used also to disrupt the lip motor representation. A 15-min train of low-frequency sub-threshold repetitive stimulation has been shown to suppress motor excitability for a further 15-20 min. This TMS-induced disruption of the motor lip representation impairs subsequent performance in demanding speech perception tasks and modulates auditory-cortex responses to speech sounds. These findings are consistent with the suggestion that the motor cortex contributes to speech perception. This article describes how to localize the lip representation in the motor cortex and how to define the appropriate stimulation intensity for carrying out both single-pulse and repetitive TMS experiments.

Lateralized enhancement of auditory cortex activity and increased sensitivity to self-generated sounds.

Performing actions with auditory consequences modulates the response in auditory cortex to otherwise identical stimuli passively heard. Such modulation has been suggested to occur through a corollary discharge sent from the motor cortex during voluntary actions. However, the relationship between the effector used to generate the sound, type of modulation and changes in perceptual sensitivity are unclear. Here we use functional magnetic resonance imaging on healthy subjects and demonstrate bilateral enhancement in the auditory cortex to self-generated versus externally generated sounds. Furthermore, we find that this enhancement is stronger when the sound-producing hand is contralateral to the auditory cortex. At the behavioural level, binaural hearing thresholds are lower for self-generated sounds and monaural thresholds are lower for sounds triggered by the hand ipsilateral to the stimulated ear. Together with functional connectivity analysis, our results suggest that a corollary discharge sent from active motor cortex enhances activity in the auditory cortex and increases perceptual sensitivity in a lateralized manner.

The Relationship Between Acceptable Noise Level and Electrophysiologic Auditory Brainstem and Cortical Signal to Noise Ratios.

The following objectives of the study were formulated: i) to investigate differences in measured signal to noise ratios while recording speech-evoked auditory brainstem response (cABR) and cortical late latency response (LLR) in low and high acceptable noise level (ANL) groups; and ii) to compare peak to peak amplitude of cABR (V-A) and LLR (N1-P2) in low and high ANL groups. A total of 23 normal hearing participants was included in the study. One shot replicative and partly exploratory research design was utilized to study the effect of signal to noise ratio in a recorded waveform on afferent mechanism, assessed by cABR and LLR on participants having values of ANL of ≤7 (low ANL group) and ≥13 (high ANL group). There were no differences in signal to noise ratio in the recorded waveforms of cABR and LLR between low and high ANL groups at both brainstem and cortical levels. However, the peak to peak amplitude of V-A of cABR and N1-P2 of LLR were both statistically larger in the high ANL group compared to their counterpart. The signal to noise ratio in recorded waveforms did not differentiated cABR (V-A) or LLR (N1-P2) in low and high ANL groups. However, Larger peak to peak amplitudes in the high ANL group suggests differences higher processing centers in the upper brainstem to the auditory cortex. The findings of the study may be useful in determining the patient acceptability of noise.

Sensory and cognitive neurophysiology in rats, Part 1: Controlled tactile stimulation and micro-ECoG recordings in freely moving animals.

We have developed a setup for rats that allows for controlled sensory input to an animal engaged in a task while recording both electrophysiological signals and behavioral output. We record electrophysiological signals using a novel high-density micro-electrocorticography (micro-ECoG) grid that covers almost the whole somatosensory system. We dealt with the well-known difficulty that the rat uses its whisker system in an active (motor-controlled) way to explore its environment by designing a head-mounted device that stimulates the rat's snout in a way unaffected by whisker movements. We replicate the spatial specificity of early evoked responses in somatosensory and auditory cortex. In a companion paper (Cognitive Neurophysiology in Rats, Part 2: Validation and Demonstration) we validate our setup and show for the first time that the ECoG can be used to record evoked responses in a signal that reflects neural output (spiking activity). Compared with high-density wire recordings, micro-ECoG offers a much more stable signal without readjustments, and a much better scalability. Compared with head-fixed preparations, our head-mounted stimulator allows to stay closer to the rat's natural way of collecting sensory information. For perceptual and cognitive research, our setup provides a unique combination of possibilities that cannot be achieved in other setups for rodents.

Mechanisms of noise robust representation of speech in primary auditory cortex.

Humans and animals can reliably perceive behaviorally relevant sounds in noisy and reverberant environments, yet the neural mechanisms behind this phenomenon are largely unknown. To understand how neural circuits represent degraded auditory stimuli with additive and reverberant distortions, we compared single-neuron responses in ferret primary auditory cortex to speech and vocalizations in four conditions: clean, additive white and pink (1/f) noise, and reverberation. Despite substantial distortion, responses of neurons to the vocalization signal remained stable, maintaining the same statistical distribution in all conditions. Stimulus spectrograms reconstructed from population responses to the distorted stimuli resembled more the original clean than the distorted signals. To explore mechanisms contributing to this robustness, we simulated neural responses using several spectrotemporal receptive field models that incorporated either a static nonlinearity or subtractive synaptic depression and multiplicative gain normalization. The static model failed to suppress the distortions. A dynamic model incorporating feed-forward synaptic depression could account for the reduction of additive noise, but only the combined model with feedback gain normalization was able to predict the effects across both additive and reverberant conditions. Thus, both mechanisms can contribute to the abilities of humans and animals to extract relevant sounds in diverse noisy environments.

How phonetically selective is the human auditory cortex?

Responses in the human auditory cortex to natural speech reveal a dual character. Often they are categorically selective to phonetic elements, serving as a gateway to abstract linguistic representations. But at other times they reflect a distributed generalized spectrotemporal analysis of the acoustic features, as seen in early mammalian auditory cortices.

Degraded speech sound processing in a rat model of fragile X syndrome

Fragile X syndrome is the most common inherited form of intellectual disability and the leading genetic cause of autism. Impaired phonological processing in fragile X syndrome interferes with the development of language skills. Although auditory cortex responses are known to be abnormal in fragile X syndrome, it is not clear how these differences impact speech sound processing. This study provides the first evidence that the cortical representation of speech sounds is impaired in Fmr1 knockout rats, despite normal speech discrimination behavior. Evoked potentials and spiking activity in response to speech sounds, noise burst trains, and tones were significantly degraded in primary auditory cortex, anterior auditory field and the ventral auditory field. Neurometric analysis of speech evoked activity using a pattern classifier confirmed that activity in these fields contains significantly less information about speech sound identity in Fmr1 knockout rats compared to control rats. Responses were normal in the posterior auditory field, which is associated with sound localization. The greatest impairment was observed in the ventral auditory field, which is related to emotional regulation. Dysfunction in the ventral auditory field may contribute to poor emotional regulation in fragile X syndrome and may help explain the observation that later auditory evoked responses are more disturbed in fragile X syndrome compared to earlier responses. Rodent models of fragile X syndrome are likely to prove useful for understanding the biological basis of fragile X syndrome and for testing candidate therapies.

Degraded auditory processing in a rat model of autism limits the speech representation in non‐primary auditory cortex

Although individuals with autism are known to have significant communication problems, the cellular mechanisms responsible for impaired communication are poorly understood. Valproic acid (VPA) is an anticonvulsant that is a known risk factor for autism in prenatally exposed children. Prenatal VPA exposure in rats causes numerous neural and behavioral abnormalities that mimic autism. We predicted that VPA exposure may lead to auditory processing impairments which may contribute to the deficits in communication observed in individuals with autism. In this study, we document auditory cortex responses in rats prenatally exposed to VPA. We recorded local field potentials and multiunit responses to speech sounds in primary auditory cortex, anterior auditory field, ventral auditory field. and posterior auditory field in VPA exposed and control rats. Prenatal VPA exposure severely degrades the precise spatiotemporal patterns evoked by speech sounds in secondary, but not primary auditory cortex. This result parallels findings in humans and suggests that secondary auditory fields may be more sensitive to environmental disturbances and may provide insight into possible mechanisms related to auditory deficits in individuals with autism.

Neural discrimination of degraded speech

Cochlear implants provide a degraded input to the auditory system. Despite this, cochlear implant users are able to discriminate speech sounds in quiet with a degree of accuracy comparable to that of normal hearing listeners. The neural bases of this phenomenon is not well understood. A set of vowel-consonant-vowel phoneme sequences, each produced by multiple talkers, were parametrically degraded using a noise vocoder. Neural responses were recorded in the guinea pig midbrain and cortex, and auditory nerve responses were generated using a computational model. The discriminability of these responses was quantified using a nearest neighbor classifier. When envelope modulations were limited to 16 Hz, classifier performance was qualitatively similar to that of human listeners for all brain regions. However, in the auditory nerve and the midbrain, the preservation of high rate envelope cues enabled the near perfect discrimination of speech tokens even for heavily spectrally degraded speech. High rate envelope cues do not appear to increase discriminability of auditory cortex responses. High rate envelope cues, represented up to the midbrain, are useful for discriminating speech tokens. However, qualitatively more consistent with perception, high rate envelope cues do not contribute to the discriminability of cortical neural responses.

Task-Dependent Decoding of Speaker and Vowel Identity from Auditory Cortical Response Patterns

Selective attention to relevant sound properties is essential for everyday listening situations. It enables the formation of different perceptual representations of the same acoustic input and is at the basis of flexible and goal-dependent behavior. Here, we investigated the role of the human auditory cortex in forming behavior-dependent representations of sounds. We used single-trial fMRI and analyzed cortical responses collected while subjects listened to the same speech sounds (vowels /a/, /i/, and /u/) spoken by different speakers (boy, girl, male) and performed a delayed-match-to-sample task on either speech sound or speaker identity. Univariate analyses showed a task-specific activation increase in the right superior temporal gyrus/sulcus (STG/STS) during speaker categorization and in the right posterior temporal cortex during vowel categorization. Beyond regional differences in activation levels, multivariate classification of single trial responses demonstrated that the success with which single speakers and vowels can be decoded from auditory cortical activation patterns depends on task demands and subject's behavioral performance. Speaker/vowel classification relied on distinct but overlapping regions across the (right) mid-anterior STG/STS (speakers) and bilateral mid-posterior STG/STS (vowels), as well as the superior temporal plane including Heschl's gyrus/sulcus. The task dependency of speaker/vowel classification demonstrates that the informative fMRI response patterns reflect the top-down enhancement of behaviorally relevant sound representations. Furthermore, our findings suggest that successful selection, processing, and retention of task-relevant sound properties relies on the joint encoding of information across early and higher-order regions of the auditory cortex.

Deconvolution of magnetic acoustic change complex (mACC)

ObjectiveThe aim of this study was to design a novel experimental approach to investigate the morphological characteristics of auditory cortical responses elicited by rapidly changing synthesized speech sounds. MethodsSix sound-evoked magnetoencephalographic (MEG) responses were measured to a synthesized train of speech sounds using the vowels /e/ and /u/ in 17 normal hearing young adults. Responses were measured to: (i) the onset of the speech train, (ii) an F0 increment; (iii) an F0 decrement; (iv) an F2 decrement; (v) an F2 increment; and (vi) the offset of the speech train using short (jittered around 135ms) and long (1500ms) stimulus onset asynchronies (SOAs). The least squares (LS) deconvolution technique was used to disentangle the overlapping MEG responses in the short SOA condition only. ResultsComparison between the morphology of the recovered cortical responses in the short and long SOAs conditions showed high similarity, suggesting that the LS deconvolution technique was successful in disentangling the MEG waveforms. Waveform latencies and amplitudes were different for the two SOAs conditions and were influenced by the spectro-temporal properties of the sound sequence. The magnetic acoustic change complex (mACC) for the short SOA condition showed significantly lower amplitudes and shorter latencies compared to the long SOA condition. The F0 transition showed a larger reduction in amplitude from long to short SOA compared to the F2 transition. Lateralization of the cortical responses were observed under some stimulus conditions and appeared to be associated with the spectro-temporal properties of the acoustic stimulus. ConclusionsThe LS deconvolution technique provides a new tool to study the properties of the auditory cortical response to rapidly changing sound stimuli. The presence of the cortical auditory evoked responses for rapid transition of synthesized speech stimuli suggests that the temporal code is preserved at the level of the auditory cortex. Further, the reduced amplitudes and shorter latencies might reflect intrinsic properties of the cortical neurons to rapidly presented sounds. SignificanceThis is the first demonstration of the separation of overlapping cortical responses to rapidly changing speech sounds and offers a potential new biomarker of discrimination of rapid transition of sound.

Attention Fine-Tunes Auditory–Motor Processing of Speech Sounds

The earliest stages of cortical processing of speech sounds take place in the auditory cortex. Transcranial magnetic stimulation (TMS) studies have provided evidence that the human articulatory motor cortex contributes also to speech processing. For example, stimulation of the motor lip representation influences specifically discrimination of lip-articulated speech sounds. However, the timing of the neural mechanisms underlying these articulator-specific motor contributions to speech processing is unknown. Furthermore, it is unclear whether they depend on attention. Here, we used magnetoencephalography and TMS to investigate the effect of attention on specificity and timing of interactions between the auditory and motor cortex during processing of speech sounds. We found that TMS-induced disruption of the motor lip representation modulated specifically the early auditory-cortex responses to lip-articulated speech sounds when they were attended. These articulator-specific modulations were left-lateralized and remarkably early, occurring 60-100 ms after sound onset. When speech sounds were ignored, the effect of this motor disruption on auditory-cortex responses was nonspecific and bilateral, and it started later, 170 ms after sound onset. The findings indicate that articulatory motor cortex can contribute to auditory processing of speech sounds even in the absence of behavioral tasks and when the sounds are not in the focus of attention. Importantly, the findings also show that attention can selectively facilitate the interaction of the auditory cortex with specific articulator representations during speech processing.

Substructure of Functional Network for Auditory Stream Segregation in Auditory Cortex

SUMMARYPerceptual integration and segregation of alternating tone sequences differing in frequency (ABA‐ABA‐...) depend on the frequency differences (ΔFs) between the A and B tones and the intertone intervals (ITIs) between successive tones. In the auditory cortex, tonotopic separation, forward suppression, and multisecond habituation have been considered as possible neural mechanisms of this perceptual phenomenon. These mechanisms, however, cannot completely account for van Noorden's perceptual boundary and the temporally continuous perception of auditory streaming. Here we examined the temporal changes of the functional network properties in the auditory cortex in response to tone sequences with different ΔFs and ITIs. Specifically, we recorded LFPs using microelectrode arrays from anesthetized rats and constructed the functional network based on phase synchrony in gamma‐band oscillation. Consequently, the networks consisted of subnetworks highly correlated with the place code of frequency, that is, a tonotopic map, and the subnetwork selective to B tones lasted for a prolonged period at large ΔF. Such characteristic substructures of the functional network are a possible candidate for the neural basis of auditory stream segregation.

The Effects of Salicylate on Auditory Evoked Potential Amplitwde from the Auditory Cortex and Auditory Brainstem

Tinnitus has often been studied using salicylate in animal models as they are capable of inducing temporary hearing loss and tinnitus. Studies have recently observed enhancement of auditory evoked responses of the auditory cortex (AC) post salicylate treatment which is also shown to be related to tinnitus like behavior in rats. The aim of this study was to observe if enhancements of the AC post salicylate treatment are also present at structures in the brainstem. Four male Sprague Dawley rats with AC implanted electrodes were tested for both AC and auditory brainstem response (ABR) recordings pre and post 250 mg/kg intraperitoneal injections of salicylate. The responses were recorded as the peak to trough amplitudes of P1-N1 (AC), ABR wave V, and ABR wave II. AC responses resulted in statistically significant enhancement of amplitude at 2 hours post salicylate with 90 dB stimuli tone bursts of 4, 8, 12, and 20kHz. Wave V of ABR responses at 90 dB resulted in a statistically significant reduction of amplitude 2 hours post salicylate and a mean decrease of amplitude of 31% for 16kHz. Wave II amplitudes at 2 hours post treatment were significantly reduced for 4, 12, and 20kHz stimuli at 90 dB SPL. Our results suggest that the enhancement changes of the AC related to salicylate induced tinnitus are generated superior to the level of the inferior colliculus and may originate in the AC.