Responses In Human Auditory Cortex Research Articles

P-14 40 Hz Steady-state responses in human auditory cortex depend on GABAergic neuronal inhibition

P-14 40 Hz Steady-state responses in human auditory cortex depend on GABAergic neuronal inhibition

Speech frequency-following response in human auditory cortex is more than a simple tracking

The human auditory cortex is recently found to contribute to the frequency following response (FFR) and the cortical component has been shown to be more relevant to speech perception. However, it is not clear how cortical FFR may contribute to the processing of speech fundamental frequency (F0) and the dynamic pitch. Using intracranial EEG recordings, we observed a significant FFR at the fundamental frequency (F0) for both speech and speech-like harmonic complex stimuli in the human auditory cortex, even in the missing fundamental condition. Both the spectral amplitude and phase coherence of the cortical FFR showed a significant harmonic preference, and attenuated from the primary auditory cortex to the surrounding associative auditory cortex. The phase coherence of the speech FFR was found significantly higher than that of the harmonic complex stimuli, especially in the left hemisphere, showing a high timing fidelity of the cortical FFR in tracking dynamic F0 in speech. Spectrally, the frequency band of the cortical FFR was largely overlapped with the range of the human vocal pitch. Taken together, our study parsed the intrinsic properties of the cortical FFR and reveals a preference for speech-like sounds, supporting its potential role in processing speech intonation and lexical tones.

Transient and sustained processing of musical consonance in auditory cortex and the effect of musicality.

In recent years, electroencephalography and magnetoencephalography (MEG) have both been used to investigate the response in human auditory cortex to musical sounds that are perceived as consonant or dissonant. These studies have typically focused on the transient components of the physiological activity at sound onset, specifically, the N1 wave of the auditory evoked potential and the auditory evoked field, respectively. Unfortunately, the morphology of the N1 wave is confounded by the prominent neural response to energy onset at stimulus onset. It is also the case that the perception of pitch is not limited to sound onset; the perception lasts as long as the note producing it. This suggests that consonance studies should also consider the sustained activity that appears after the transient components die away. The current MEG study shows how energy-balanced sounds can focus the response waves on the consonance-dissonance distinction rather than energy changes and how source modeling techniques can be used to measure the sustained field associated with extended consonant and dissonant sounds. The study shows that musical dyads evoke distinct transient and sustained neuromagnetic responses in auditory cortex. The form of the response depends on both whether the dyads are consonant or dissonant and whether the listeners are musical or nonmusical. The results also show that auditory cortex requires more time for the early transient processing of dissonant dyads than it does for consonant dyads and that the continuous representation of temporal regularity in auditory cortex might be modulated by processes beyond auditory cortex.NEW & NOTEWORTHY We report a magnetoencephalography (MEG) study on transient and sustained cortical consonance processing. Stimuli were long-duration, energy-balanced, musical dyads that were either consonant or dissonant. Spatiotemporal source analysis revealed specific transient and sustained neuromagnetic activity in response to the dyads; in particular, the morphology of the responses was shaped by the dyad's consonance and the listener's musicality. Our results also suggest that the sustained representation of stimulus regularity might be modulated by processes beyond auditory cortex.

Neural phase locking predicts BOLD response in human auditory cortex

Natural environments elicit both phase-locked and non-phase-locked neural responses to the stimulus in the brain. The interpretation of the BOLD signal to date has been based on an association of the non-phase-locked power of high-frequency local field potentials (LFPs), or the related spiking activity in single neurons or groups of neurons. Previous studies have not examined the prediction of the BOLD signal by phase-locked responses. We examined the relationship between the BOLD response and LFPs in the same nine human subjects from multiple corresponding points in the auditory cortex, using amplitude modulated pure tone stimuli of a duration to allow an analysis of phase locking of the sustained time period without contamination from the onset response. The results demonstrate that both phase locking at the modulation frequency and its harmonics, and the oscillatory power in gamma/high-gamma bands are required to predict the BOLD response. Biophysical models of BOLD signal generation in auditory cortex therefore require revision and the incorporation of both phase locking to rhythmic sensory stimuli and power changes in the ensemble neural activity.

Amplitude modulation rate dependent topographic organization of the auditory steady-state response in human auditory cortex

Periodic modulations of an acoustic feature, such as amplitude over a certain frequency range, leads to phase locking of neural responses to the envelope of the modulation. Using electrophysiological methods this neural activity pattern, also called the auditory steady-state response (aSSR), is visible following frequency transformation of the evoked response as a clear spectral peak at the modulation frequency. Despite several studies employing the aSSR that show, for example, strongest responses for ∼40 Hz and an overall right-hemispheric dominance, it has not been investigated so far to what extent within auditory cortex different modulation frequencies elicit aSSRs at a homogenous source or whether the localization of the aSSR is topographically organized in a systematic manner. The latter would be suggested by previous neuroimaging works in monkeys and humans showing a periodotopic organization within and across distinct auditory fields. However, the sluggishness of the signal from these neuroimaging works prohibit inferences with regards to the fine-temporal features of the neural response. In the present study, we employed amplitude-modulated (AM) sounds over a range between 4 and 85 Hz to elicit aSSRs while recording brain activity via magnetoencephalography (MEG). Using beamforming and a fine spatially resolved grid restricted to auditory cortical processing regions, our study revealed a topographic representation of the aSSR that depends on AM rate, in particular in the medial-lateral (bilateral) and posterior-anterior (right auditory cortex) direction. In summary, our findings confirm previous studies that showing different AM rates to elicit maximal response in distinct neural populations. They extend these findings however by also showing that these respective neural ensembles in auditory cortex actually phase lock their activity over a wide modulation frequency range.

Robustness to real-world background noise increases between primary and non-primary human auditory cortex

In everyday listening, the sounds from sources we seek to understand are frequently embedded in background noise, which often profoundly alters auditory nerve spiking. To recognize sources of interest the brain must be somewhat robust to the effects of these background noises. To study the neural basis of listening in real-world background noise, we measured fMRI responses in human auditory cortex to a diverse set of thirty natural sounds, presented in quiet as well as embedded in thirty different everyday background noise textures (e.g., a bustling coffee shop, crickets chirping, etc.). We quantified the noise-robustness of neural responses by correlating each voxel’s response to the natural sounds in quiet with its response to those same sounds superimposed on background noise. Responses in core regions (commonly identified with primary auditory cortex) were substantially affected by background noise. However, noise-robustness increased with distance from primary auditory cortex: nearby non-primary areas were slightly more robust, while more distal areas were hardly affected by the background noises. Our results provide a neural correlate of the noise robustness of real-world listening, and offer evidence of a hierarchical organization in human auditory cortex.

Sparse Spectro-Temporal Receptive Fields Based on Multi-Unit and High-Gamma Responses in Human Auditory Cortex.

Spectro-Temporal Receptive Fields (STRFs) were estimated from both multi-unit sorted clusters and high-gamma power responses in human auditory cortex. Intracranial electrophysiological recordings were used to measure responses to a random chord sequence of Gammatone stimuli. Traditional methods for estimating STRFs from single-unit recordings, such as spike-triggered-averages, tend to be noisy and are less robust to other response signals such as local field potentials. We present an extension to recently advanced methods for estimating STRFs from generalized linear models (GLM). A new variant of regression using regularization that penalizes non-zero coefficients is described, which results in a sparse solution. The frequency-time structure of the STRF tends toward grouping in different areas of frequency-time and we demonstrate that group sparsity-inducing penalties applied to GLM estimates of STRFs reduces the background noise while preserving the complex internal structure. The contribution of local spiking activity to the high-gamma power signal was factored out of the STRF using the GLM method, and this contribution was significant in 85 percent of the cases. Although the GLM methods have been used to estimate STRFs in animals, this study examines the detailed structure directly from auditory cortex in the awake human brain. We used this approach to identify an abrupt change in the best frequency of estimated STRFs along posteromedial-to-anterolateral recording locations along the long axis of Heschl’s gyrus. This change correlates well with a proposed transition from core to non-core auditory fields previously identified using the temporal response properties of Heschl’s gyrus recordings elicited by click-train stimuli.

Monaural and binaural contributions to interaural-level-difference sensitivity in human auditory cortex

Whole-brain functional magnetic resonance imaging was used to measure blood-oxygenation-level-dependent (BOLD) responses in human auditory cortex (AC) to sounds with intensity varying independently in the left and right ears. Echoplanar images were acquired at 3 Tesla with sparse image acquisition once per 12-second block of sound stimulation. Combinations of binaural intensity and stimulus presentation rate were varied between blocks, and selected to allow measurement of response-intensity functions in three configurations: monaural 55–85 dB SPL, binaural 55–85 dB SPL with intensity equal in both ears, and binaural with average binaural level of 70 dB SPL and interaural level differences (ILD) ranging ±30 dB (i.e., favoring the left or right ear). Comparison of response functions equated for contralateral intensity revealed that BOLD-response magnitudes (1) generally increased with contralateral intensity, consistent with positive drive of the BOLD response by the contralateral ear, (2) were larger for contralateral monaural stimulation than for binaural stimulation, consistent with negative effects (e.g., inhibition) of ipsilateral input, which were strongest in the left hemisphere, and (3) also increased with ipsilateral intensity when contralateral input was weak, consistent with additional, positive, effects of ipsilateral stimulation. Hemispheric asymmetries in the spatial extent and overall magnitude of BOLD responses were generally consistent with previous studies demonstrating greater bilaterality of responses in the right hemisphere and stricter contralaterality in the left hemisphere. Finally, comparison of responses to fast (40/s) and slow (5/s) stimulus presentation rates revealed significant rate-dependent adaptation of the BOLD response that varied across ILD values.

Sustained BOLD and theta activity in auditory cortex are related to slow stimulus fluctuations rather than to pitch

Human functional MRI (fMRI) and magnetoencephalography (MEG) studies indicate a pitch-specific area in lateral Heschl's gyrus. Single-cell recordings in monkey suggest that sustained-firing, pitch-specific neurons are located lateral to primary auditory cortex. We reevaluated whether pitch strength contrasts reveal sustained pitch-specific responses in human auditory cortex. Sustained BOLD activity in auditory cortex was found for iterated rippled noise (vs. noise or silence) but not for regular click trains (vs. jittered click trains or silence). In contrast, iterated rippled noise and click trains produced similar pitch responses in MEG. Subsequently performed time-frequency analysis of the MEG data suggested that the dissociation of cortical BOLD activity between iterated rippled noise and click trains is related to theta band activity. It appears that both sustained BOLD and theta activity are associated with slow non-pitch-specific stimulus fluctuations. BOLD activity in the inferior colliculus was sustained for both stimulus types and varied neither with pitch strength nor with the presence of slow stimulus fluctuations. These results suggest that BOLD activity in auditory cortex is much more sensitive to slow stimulus fluctuations than to constant pitch, compromising the accessibility of the latter. In contrast, pitch-related activity in MEG can easily be separated from theta band activity related to slow stimulus fluctuations.

Auditory evoked responses in human auditory cortex to the variation of sound intensity in an ongoing tone

In daily life, variations of sound intensity, frequency, and other auditory parameters, can be perceived as transitions from one sound to another. The neural mechanisms underlying the processing of intensity change are currently unclear. The present study sought to clarify the effects of frequency and initial sound pressure level (SPL) on the auditory evoked response elicited by sounds of different SPL. We examined responses approximately 100 ms after an SPL change (the N1m′). Experiment 1 examined the effects of frequency on the N1m′. Experiment 2 examined the effects of initial SPL on the N1m′. The results revealed that N1m′ amplitude increased with greater SPL changes. The increase in N1m′ amplitude with increasing SPL was almost constant for low frequency sounds (250 and 1000 Hz); however, this increase was reduced for high frequency sounds (4000 Hz). The increase in N1m′ amplitude was reduced with high initial SPL. The pattern of amplitude change may reflect a difference in activation in the auditory nerve and/or primary auditory cortex.

Gamma band pitch responses in human auditory cortex measured with magnetoencephalography

We have previously used direct electrode recordings in two human subjects to identify neural correlates of the perception of pitch (Griffiths, Kumar, Sedley et al., Direct recordings of pitch responses from human auditory cortex, Curr. Biol. 22 (2010), pp. 1128–1132). The present study was carried out to assess virtual-electrode measures of pitch perception based on non-invasive magnetoencephalography (MEG). We recorded pitch responses in 13 healthy volunteers using a passive listening paradigm and the same pitch-evoking stimuli (regular interval noise; RIN) as in the previous study. Source activity was reconstructed using a beamformer approach, which was used to place virtual electrodes in auditory cortex. Time-frequency decomposition of these data revealed oscillatory responses to pitch in the gamma frequency band to occur, in Heschl's gyrus, from 60 Hz upwards. Direct comparison of these pitch responses to the previous depth electrode recordings shows a striking congruence in terms of spectrotemporal profile and anatomical distribution. These findings provide further support that auditory high gamma oscillations occur in association with RIN pitch stimuli, and validate the use of MEG to assess neural correlates of normal and abnormal pitch perception.

Representation of temporal sound features in the human auditory cortex

Temporal information in acoustic signals is important for the perception of environmental sounds, including speech. This review focuses on several aspects of temporal processing within human auditory cortex and its relevance for the processing of speech sounds. Periodic non-speech sounds, such as trains of acoustic clicks and bursts of amplitude-modulated noise or tones, can elicit different percepts depending on the pulse repetition rate or modulation frequency. Such sounds provide convenient methodological tools to study representation of timing information in the auditory system. At low repetition rates of up to 8-10 Hz, each individual stimulus (a single click or a sinusoidal amplitude modulation cycle) within the sequence is perceived as a separate event. As repetition rates increase up to and above approximately 40 Hz, these events blend together, giving rise first to the percept of flutter and then to pitch. The extent to which neural responses of human auditory cortex encode temporal features of acoustic stimuli is discussed within the context of these perceptual classes of periodic stimuli and their relationship to speech sounds. Evidence for neural coding of temporal information at the level of the core auditory cortex in humans suggests possible physiological counterparts to perceptual categorical boundaries for periodic acoustic stimuli. Temporal coding is less evident in auditory cortical fields beyond the core. Finally, data suggest hemispheric asymmetry in temporal cortical processing.

BOLD Responses in Human Auditory Cortex Are More Closely Related to Transient MEG Responses Than to Sustained Ones

Blood oxygen level dependent-functional magnetic resonance imaging (BOLD-fMRI) and magnetoencephalographic (MEG) signals are both coupled to postsynaptic potentials, although their relationship is incompletely understood. Here, the wide range of BOLD-fMRI and MEG responses produced by auditory cortex was exploited to better understand the BOLD-fMRI/MEG relationship. Measurements of BOLD and MEG responses were made in the same subjects using the same stimuli for both modalities. The stimuli, 24-s sequences of click trains, had duty cycles of 2.5, 25, 72, and 100%. For the 2.5% sequence, the BOLD response was elevated throughout the sequence, whereas for 100%, it peaked after sequence onset and offset and showed a diminished elevation in between. On the finer timescale of MEG, responses at 2.5% consisted of a complex of transients, including N(1)m, to each click train of the sequence, whereas for 100% the only transients occurred at sequence onset and offset between which there was a sustained elevation in the MEG signal (a sustained field). A model that separately estimated the contributions of transient and sustained MEG signals to the BOLD response best fit BOLD measurements when the transient contribution was weighted 8- to 10-fold more than the sustained one. The findings suggest that BOLD responses in the auditory cortex are tightly coupled to the neural activity underlying transient, not sustained, MEG signals.

Stream segregation based on interaural time differences: differential effects on transient and sustained components of the fMRI BOLD response in human auditory cortex

Stream segregation based on interaural time differences: differential effects on transient and sustained components of the fMRI BOLD response in human auditory cortex

Dynamic movement of N100m current sources in auditory evoked fields: Comparison of ipsilateral versus contralateral responses in human auditory cortex

We recorded auditory evoked magnetic fields (AEFs) to monaural 400 Hz tone bursts and investigated spatio-temporal features of the N100m current sources in the both hemispheres during the time before the N100m reaches at the peak strength and 5 ms after the peak. A hemispheric asymmetry was evaluated as the asymmetry index based on the ratio of N100m peak dipole strength between right and left hemispheres for either ear stimulation. The results of asymmetry indices showed right-hemispheric dominance for left ear stimulation but no hemispheric dominance for right ear stimulation. The current sources for N100m in both hemispheres in response to monaural 400 Hz stimulation moved toward anterolateral direction along the long axis of the Heschl gyri during the time before it reaches the peak strength; the ipsilateral N100m sources were located slightly posterior to the contralateral N100m ones. The onset and peak latencies of the right hemispheric N100m in response to right ear stimulation are shorter than those of the left hemispheric N100m to left ear stimulation. The traveling distance of the right hemispheric N100m sources following right ear stimulation was longer than that for the left hemispheric ones following left ear stimulation. These results suggest the right-dominant hemispheric asymmetry in pure tone processing.

The effect of temporal context on the sustained pitch response in human auditory cortex.

Recent neuroimaging studies have shown that activity in lateral Heschl's gyrus covaries specifically with the strength of musical pitch. Pitch strength is important for the perceptual distinctiveness of an acoustic event, but in complex auditory scenes, the distinctiveness of an event also depends on its context. In this magnetoencephalography study, we evaluate how temporal context influences the sustained pitch response (SPR) in lateral Heschl's gyrus. In 2 sequences of continuously alternating, periodic target intervals and a more irregular baseline interval, the distinctiveness of the target was decreased in 1 of 2 ways--either by increasing the pitch strength of the baseline or by decreasing the pitch strength of the target. The results show that the amplitude of the SPR increases monotonically with the distinctiveness of the target. Moreover, SPR amplitude is greater for the sequence, where the pitch strength of the target is varied, compared with the condition, where the baseline is varied. Two subsequent experiments show that the amplitude of the SPR increases as duty cycle decreases, in a pitch "strength" contrast and in a pitch "value" contrast. These results indicate that the SPR adapts to recent stimulus history, enhancing the response to rare and brief events.

Nonlinearity of FMRI responses in human auditory cortex.

An investigation was made into the nature of the nonlinearity observed in auditory functional magnetic resonance imaging (fMRI) experiments associated with increases in total duration of acoustic imaging noise [e.g., Edmister et al., 1999; Shah et al., 1999]. A two-stimulus, four-condition paradigm was used to evaluate four acoustic conditions involving: (1) the presence or absence of a desired broadband music stimulus; and (2) two possible durations of trains of acoustic noise associated with image acquisition. Responses observed while increasing the duration of acoustic imaging noise were consistent with previous work (Talavage et al. [1999]: Hum Brain Mapp7:79-88) but the response to combined stimulation did not exhibit variation as a function of the acoustic imaging noise duration. These results suggest that spectral overlap of the stimuli produced colocalized responses that did not add linearly. This conclusion has implications for conducting both blocked and rapid-presentation event-related auditory fMRI experiments. The cortical activity induced by the stimulus may not reflect the activation, in spatial extent or magnitude of signal change, occurring in the absence of other acoustic noise.

Effects of cholinergic enhancement on conditioning-related responses in human auditory cortex.

It has previously been shown that cholinergic blockade attenuates conditioning-related neuronal responses in human auditory cortex. The present study was conducted to investigate the effect of cholinergic enhancement on such experience-dependent cortical responses. The cholinesterase inhibitor physostigmine, or a placebo control, were continuously infused into healthy young volunteers, during differential aversive conditioning whilst brain activity was measured using event-related functional magnetic resonance imaging (fMRI). Volunteers were presented with two tones, one of which (CS+) was conditioned by pairing with an electrical shock whereas the other was always presented without the shock (CS-). Conditioning-related activations, expressed as an enhanced blood oxygenation level dependent (BOLD) response to the salient CS+, were evident in left auditory cortex under placebo but not under physostigmine. This absence of conditioning-related activations under physostigmine was due to enhanced responses to the CS- under physostigmine as compared to placebo. We suggest that an overactive cholinergic system leads to increased processing of behaviourally irrelevant stimuli and thus attenuates differential conditioning-related cortical activations.

Human Cortical Dynamics Determined by Speech Fundamental Frequency

Evidence for speech-specific brain processes has been searched for through the manipulation of formant frequencies which mediate phonetic content and which are, in evolutionary terms, relatively “new” aspects of speech. Here we used whole-head magnetoencephalography and advanced stimulus reproduction methodology to examine the contribution of the fundamental frequency F0 and its harmonic integer multiples in cortical processing. The subjects were presented with a vowel, a frequency-matched counterpart of the vowel lacking in phonetic contents, and a pure tone. The F0 of the stimuli was set at that of a typical male (i.e., 100 Hz), female (200 Hz), or infant (270 Hz) speaker. We found that speech sounds, both with and without phonetic content, elicited the N1m response in human auditory cortex at a constant latency of 120 ms, whereas pure tones matching the speech sounds in frequency, intensity, and duration gave rise to N1m responses whose latency varied between 120 and 160 ms. Thus, it seems that the fundamental frequency F0 and its harmonics determine the temporal dynamics of speech processing in human auditory cortex and that speech specificity arises out of cortical sensitivity to the complex acoustic structure determined by the human sound production apparatus.

Learning-related Modulation of Tonotopic Responses in Human Auditory Cortex

Learning-related Modulation of Tonotopic Responses in Human Auditory Cortex