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PubMed2002

Sound-level-dependent representation of frequency modulations in human auditory cortex: a low-noise fMRI study.

Recognition of sound patterns must be largely independent of level and of masking or jamming background sounds. Auditory patterns of relevance in numerous environmental sounds, species-specific vocalizations and speech are frequency modulations (FM). Level-dependent activation of the human auditory cortex (AC) in response to a large set of upward and downward FM tones was studied with low-noise (48 dB) functional magnetic resonance imaging at 3 Tesla. Separate analysis in four territories of AC was performed in each individual brain using a combination of anatomical landmarks and spatial activation criteria for their distinction. Activation of territory T1b (including primary AC) showed the most robust level dependence over the large range of 48-102 dB in terms of activated volume and blood oxygen level dependent contrast (BOLD) signal intensity. The left nonprimary territory T2 also showed a good correlation of level with activated volume but, in contrast to T1b, not with BOLD signal intensity. These findings are compatible with level coding mechanisms observed in animal AC. A systematic increase of activation with level was not observed for T1a (anterior of Heschl's gyrus) and T3 (on the planum temporale). Thus these areas might not be specifically involved in processing of the overall intensity of FM. The rostral territory T1a of the left hemisphere exhibited highest activation when the FM sound level fell 12 dB below scanner noise. This supports the previously suggested special involvement of this territory in foreground-background decomposition tasks. Overall, AC of the left hemisphere showed a stronger level-dependence of signal intensity and activated volume than the right hemisphere. But any side differences of signal intensity at given levels were lateralized to right AC. This might point to an involvement of the right hemisphere in more specific aspects of FM processing than level coding.

Acoustic Stimulation↗
PubMed2002

Auditory cortical images of cochlear-implant stimuli: coding of stimulus channel and current level.

This study quantified the accuracy with which populations of neurons in the auditory cortex can represent aspects of electrical cochlear stimuli presented through a cochlear implant. We tested the accuracy of coding of the place of stimulation (i.e., identification of the active stimulation channel) and of the stimulus current level. Physiological data came from the companion study, which recorded spike activity of neurons simultaneously from 16 sites along the tonotopic axis of the guinea pig's auditory cortex. In that study, cochlear electrical stimuli were presented to acutely deafened animals through a 6-electrode animal version of the 22-electrode Nucleus banded electrode array (Cochlear). Cochlear electrode configurations consisted of monopolar (MP), bipolar (BP + N) with N inactive electrodes between the active and return electrodes (0 < or = N < or = 3), tripolar (TP) with one active electrode and two flanking return electrodes, and common ground (CG) with one active electrode and as many as five return electrodes. In the present analysis, an artificial neural network was trained to recognize spatiotemporal patterns of cortical activity in response to single presentations of particular stimuli and, thereby, to identify those stimuli. The accuracy of pair-wise discrimination of stimulation channels or of current levels was represented by the discrimination index, d', where d' = 1 was taken as threshold. In many cases, the threshold for discrimination of place of cochlear stimulation was < 0.75 mm, and the threshold for discrimination of current levels was < 1 dB. Cochlear electrode configurations varied in the accuracy with which they signaled to the auditory cortex the place of cochlear stimulation. The BP + N and TP configurations provided considerably greater sensitivity to place of stimulation than did the MP configuration. The TP configuration maintained accurate signaling of place of stimulation up to the highest current levels, whereas sensitivity was degraded at high current levels in BP + N configurations. Electrode configurations also varied in the dynamic range over which they signaled stimulus current level. Dynamic ranges were widest for the BP + 0 configuration and narrowest for the TP configuration. That is, the configuration that showed the most accurate signaling of cochlear place of stimulation (TP) showed the most restricted dynamic range for signaling of current level. These results suggest that the choice of the optimal electrode configuration for use by human cochlear-prosthesis users would depend on the particular demands of the speech-processing strategy that is to be employed.

Action Potentials↗
PubMed2002

Auditory cortical images of cochlear-implant stimuli: dependence on electrode configuration.

This study examines patterns of auditory cortical activity elicited by single-pulse cochlear implant stimuli that vary in electrode configuration, cochlear place of stimulation, and stimulus level. Recordings were made from the primary auditory cortex (area A1) of ketamine-anesthetized guinea pigs. The spatiotemporal pattern of neural spike activity was measured simultaneously across 16 cortical locations spanning approximately 2-3 octaves of the tonotopic axis. Such a pattern, averaged over 40 presentations of any particular stimulus, was defined as the "cortical image" of that stimulus. Acutely deafened guinea pigs were implanted with a 6-electrode animal version of the 22-electrode Nucleus banded electrode array (Cochlear). Cochlear electrode configurations consisted of monopolar (MP), bipolar (BP + N) with N inactive electrodes between the active and return electrodes (0 < or = N < or = 4), tripolar (TP) with one active electrode and two flanking return electrodes, and common ground (CG) with one active electrode and as many as five return electrodes. Cortical images typically showed a focus of maximum spike probability and minimum latency. Spike probabilities tended to decrease, and latencies tended to increase, with increasing cortical distance from that focus. Cortical images of TP stimuli were the most spatially compact, followed by BP + N images, and then MP images, which were the broadest. Images of CG stimuli were rather variable across animals and stimulus channels. The locations of cortical images shifted systematically from caudal to rostral as the cochlear place of stimulation changed from basal to apical. At the most sensitive cortical site for each condition, the dynamic ranges over which spike rates increased with increased current level were restricted to about 1-2 dB, regardless of configuration. Dynamic ranges tended to increase with increasing cortical distance from the most sensitive site. Electrode configurations that produced compact cortical images (e.g., TP and BP + 0) showed the greatest range of thresholds within each cortical image and the largest dynamic range at cortical sites removed from the most sensitive site.

Action Potentials↗
PubMed2003

Effects of amplitude modulation on the coding of interaural time differences of low-frequency sounds in the inferior colliculus. I. Response properties.

Most sounds in the natural environment are amplitude-modulated (AM). To determine if AM alters the neuronal sensitivity to interaural time differences (ITDs) in low-frequency sounds, we tested neuronal responses to a binaural beat stimulus with and without modulation. We recorded from single units in the inferior colliculus of the unanesthetized rabbit. We primarily used low frequency ( approximately 25 Hz) modulation that was identical at both ears. We found that modulation could enhance, suppress, or not affect the discharge rate. In extreme cases, a neuron that showed no response to the unmodulated binaural beat did so when modulation was added to both ears. At the other extreme, a neuron that showed sensitivity to the unmodulated binaural beat ceased firing with modulation. Modulation could also affect the frequency range of ITD sensitivity, best ITD, and ITD tuning width. Despite these changes in individual neurons, averaging across all neurons, the peak and width of the population ITD function remained unchanged. Because ITD-sensitive neurons also time-locked to the modulation frequency, the location and sound attributes are processed simultaneously by these neurons.

Acoustic Stimulation↗
PubMed2003

Effects of amplitude modulation on the coding of interaural time differences of low-frequency sounds in the inferior colliculus. II. Neural mechanisms.

In our companion paper, we reported on interaural time difference (ITD)-sensitive neurons that enhanced, suppressed, or did not change their response when identical AM was added to both ears. Here, we first examined physical factors such as the difference in the interaural correlation, spectrum, or energy between the modulated and unmodulated signals. These were insufficient to explain the observed enhancement and suppression. We then examined neural mechanisms by selectively modulating the signal to each ear, varying modulation depth, and adding background noise to the unmodulated signal. These experiments implicated excitatory and inhibitory monaural inputs to the inferior colliculus (IC). These monaural inputs are postulated to adapt to an unmodulated signal and adapt less to a modulated signal. Thus enhancement or suppression is created by the convergence of these excitatory or inhibitory inputs with the inputs from the binaural comparators. Under modulation, the role of the monaural input is to shift the threshold of the IC neuron. Consistent with this role, background noise mimicked the effect of modulation. Functionally, enhancement and suppression may serve in detecting the degree of modulation in a sound source while preserving ITD information.

Acoustic Stimulation↗
PubMed2003

Directional sensitivity of neurons in the primary auditory (AI) cortex: effects of sound-source intensity level.

Transient sounds were delivered from different directions in virtual acoustic space while recording from single neurons in primary auditory cortex (AI) of cats under general anesthesia. The intensity level of the sound source was varied parametrically to determine the operating characteristics of the spatial receptive field. The spatial receptive field was constructed from the onset latency of the response to a sound at each sampled direction. Spatial gradients of response latency composing a receptive field are due partially to a systematic co-dependence on sound-source direction and intensity level. Typically, at any given intensity level, the distribution of response latency within the receptive field was unimodal with a range of approximately 3-4 ms, although for some cells and some levels, the spread could be as much as 20 or as little as 2 ms. Response latency, averaged across directions, differed among neurons for the same intensity level, and also differed among intensity levels for the same neuron. Generally, increases in intensity level resulted in decreases in the mean and variance, which follows an inverse Gaussian distribution. Receptive field models, based on response latency, are developed using multiple parameters (azimuth, elevation, intensity), validated with Monte Carlo simulation, and their spatial filtering described using spherical harmonic analysis. Observations from an ensemble of modeled receptive fields are obtained by linking the inverse Gaussian density to the probabilistic inverse problem of estimating sound-source direction and intensity. Upper bounds on acuity is derived from the ensemble using Fisher information, and the predicted patterns of estimation errors are related to psychophysical performance.

Acoustic Stimulation↗
PubMed2003

Neural model for physiological responses to frequency and amplitude transitions uncovers topographical order in the auditory cortex.

We characterize primary auditory cortex (AI) units using a neural model for the detection of frequency and amplitude transitions. The model is a generalization of a model for the detection of amplitude transition. A set of neurons, tuned in the spectrotemporal domain, is created by means of neural delays and frequency filtering. The sensitivity of the model to frequency and amplitude transitions is achieved by applying a 2-dimensional rotatable receptive field to the set of spectrotemporally tuned neurons. We evaluated the model using data recorded in AI of anesthetized ferrets. We show that the model is able to fit the responses of AI units to variety of stimuli, including single tones, delayed 2-tone stimuli and various frequency-modulated tones, using only a small number of parameters. Furthermore, we show that the topographical order in maps of the model parameters is higher than in maps created from response indices extracted directly from the responses to any single stimulus. These results suggest a possible ordered organization of a simple rotatable spectrotemporal receptive field in the mammalian AI.

Acoustic Stimulation↗
PubMed2005

Contextual modulation of olivocochlear pathway effects on loud sound-induced cochlear hearing desensitization.

This study shows that the cochlear hearing losses [temporary threshold shifts (TTSs)] induced by traumatic sound and the effect of olivocochlear (OC) pathways to the cochlea on these hearing losses depend on the context of the sound. Background atraumatic white noise (WN) has been shown to 1) exacerbate loud-pure-tone-induced TTSs, and 2) promote the modulation of TTSs by the uncrossed OC (UOC) pathways additional to the action on TTSs, elicited by binaural loud tones themselves, by the crossed OC (COC) pathway. Here the same atraumatic WN reduced TTSs caused by loud narrow band sound. It also reduced TTS modulation by OC pathways. The UOC no longer exerted any effects on TTSs, and COC effects were significantly reduced in two discrete frequency bands: low frequencies within the narrow band ("within-band" frequencies) and high frequencies outside the band ("high-side" frequencies). COC effects were unchanged at high frequencies within the band. Despite these reductions in OC effects, because the WN itself reduced TTSs, the total effect of OC pathways and background WN now produced larger TTS reductions, especially at higher frequencies. Thus the modulatory effects of the OC pathways on TTSs depend on how background WN modulates cochlear state. It is postulated that the WN background and the OC pathways both modulate TTSs by acting on the outer hair cells, in a way that promotes the reduction of TTSs caused by the narrow band sound trauma. This joint promotion of a protective end-effect on TTSs to narrow band sound trauma contrasts against the effects seen with pure tone trauma where the same background WN exacerbated TTSs at high-side frequencies.

Acoustic Stimulation↗
PubMed2005

Wiener-kernel analysis of responses to noise of chinchilla auditory-nerve fibers.

Responses to broadband Gaussian white noise were recorded in auditory-nerve fibers of deeply anesthetized chinchillas and analyzed by computation of zeroth-, first-, and second-order Wiener kernels. The first-order kernels (similar to reverse correlations or "revcors") of fibers with characteristic frequency (CF) <2 kHz consisted of lightly damped transient oscillations with frequency equal to CF. Because of the decay of phase locking strength as a function of frequency, the signal-to-noise ratio of first-order kernels of fibers with CFs >2 kHz decreased with increasing CF at a rate of about -18 dB per octave. However, residual first-order kernels could be detected in fibers with CF as high as 12 kHz. Second-order kernels, 2-dimensional matrices, reveal prominent periodicity at the CF frequency, regardless of CF. Thus onset delays, frequency glides, and near-CF group delays could be estimated for auditory-nerve fibers innervating the entire length of the chinchilla cochlea.

Acoustic Stimulation↗
PubMed2006

Concurrent encoding of frequency and amplitude modulation in human auditory cortex: MEG evidence.

A natural sound can be described by dynamic changes in envelope (amplitude) and carrier (frequency), corresponding to amplitude modulation (AM) and frequency modulation (FM), respectively. Although the neural responses to both AM and FM sounds are extensively studied in both animals and humans, it is uncertain how they are corepresented when changed simultaneously but independently, as is typical for ecologically natural signals. This study elucidates the neural coding of such sounds in human auditory cortex using magnetoencephalography (MEG). Using stimuli with both sinusoidal modulated envelope (f(AM), 37 Hz) and carrier frequency (f(FM), 0.3-8 Hz), it is demonstrated that AM and FM stimulus dynamics are corepresented in the neural code of human auditory cortex. The stimulus AM dynamics are represented neurally with AM encoding, by the auditory steady-state response (aSSR) at f(AM). For sounds with slowly changing carrier frequency (f(FM) <5 Hz), it is shown that the stimulus FM dynamics are tracked by the phase of the aSSR, demonstrating neural phase modulation (PM) encoding of the stimulus carrier frequency. For sounds with faster carrier frequency change (f(FM) > or = 5 Hz), it is shown that modulation encoding of stimulus FM dynamics persists, but the neural encoding is no longer purely PM. This result is consistent with the recruitment of additional neural AM encoding over and above the original neural PM encoding, indicating that both the amplitude and phase of the aSSR at f(AM) track the stimulus FM dynamics. A neural model is suggested to account for these observations.

Acoustic Stimulation↗
PubMed1983

Some features of binaural input to single neurons in physiologically defined area AI of cat cerebral cortex.

1. In the ectosylvian cortex of 24 barbiturate-anesthetized cats, area AI was identified by its frequency organization and the responses to tonal stimuli of single neurons in that field were examined using sealed stimulating systems incorporating calibrated probe microphone assemblies. 2. The responsiveness to monaural and binaural best-frequency stimuli was examined quantitatively for 282 single units in AI. One hundred thirty-nine cells (49%) were excited by independent stimulation of only one ear and were classified as EO cells. In general, the effective monaural excitatory input was derived from the contralateral ear. One hundred ten (39%) neurons were excited by independent stimulation of each ear and were classified as EE units. For these neurons, the contralateral responses were generally stronger, shorter in latency, and lower in threshold than were their ipsilateral responses. Thirty-three cells (12%) gave weak or no responses to monaural stimuli but responded securely to binaural stimuli. These cells were classified as predominantly binaural (PB). 3. Binaural interactions were examined by comparison of the response to binaural, equally intense stimuli to the stronger monaural response. Among EO cells suppression was the most common form of interaction, while for EE cells summation was the more common. Less than 8% of cells were found to be monaural. 4. In electrode penetrations radial to the cortex surface, cells received their stronger or sole monaural excitatory input from a common ear, generally the contralateral. Within such penetrations, however, cells commonly differed with regard to the nature of their input from the other ear and/or in their binaural interactions. 5. Comparison of these data with data previously reported for subcortical auditory nuclei revealed that AI preserves many of the stimulus specificity characteristics of the lower nuclei. The reasons for the preservation of these characteristics at the cortex and the implications of the present data for the binaural column hypothesis are discussed.

Animals↗
PubMed1983

Auditory nerve representation of vowels in background noise.

Responses of auditory nerve fibers to steady-state vowels presented alone and in the presence of background noise were obtained from anesthetized cats. Representation of vowels based on average discharge rate and representation based primarily on phase-locked properties of responses are considered. Profiles of average discharge rate versus characteristic frequency (CF) ("rate-place" representation) can show peaks of discharge rate in the vicinity of formant frequencies when vowels are presented alone. These profiles change drastically in the presence of background noise, however. At moderate vowel and noise levels and signal/noise ratios of +9 dB, there are not peaks of rate near the second and third formant frequencies. In fact, because of two-tone suppression, rate to vowels plus noise is less than rate to noise alone for fibers with CFs above the first formant. Rate profiles measured over 5-ms intervals near stimulus onset show clear formant-related peaks at higher sound levels than do profiles measured over intervals later in the stimulus (i.e., in the steady state). However, in background noise, rate profiles at onset are similar to those in the steady state. Specifically, for fibers with CFs above the first formant, response rates to the noise are suppressed by the addition of the vowel at both vowel onset and steady state. When rate profiles are plotted for low spontaneous rate fibers, formant-related peaks appear at stimulus levels higher than those at which peaks disappear for high spontaneous fibers. In the presence of background noise, however, the low spontaneous fibers do not preserve formant peaks better than do the high spontaneous fibers. In fact, the suppression of noise-evoked rate mentioned above is greater for the low spontaneous fibers than for high. Representations that reflect phase-locked properties as well as discharge rate ("temporal-place" representations) are much less affected by background noise. We have used synchronized discharge rate averaged over fibers with CFs near (+/- 0.25 octave) a stimulus component as a measure of the population temporal response to that component. Plots of this average localized synchronized rate (ALSR) versus frequency show clear first and second formant peaks at all vowel and noise levels used. Except at the highest level (vowel at 85 dB sound pressure level (SPL), signal/noise = +9 dB), there is also a clear third formant peak. At signal-to-noise ratios where there are no second formant peaks in rate profiles, human observers are able to discriminate second formant shifts of less than 112 Hz. ALSR plots show clear second formant peaks at these signal/noise ratios.

Animals↗
PubMed1984

Responses of single neurons in posterior field of cat auditory cortex to tonal stimulation.

In the auditory cortex of barbiturate-anesthetized cats, the posterior auditory field (field P) was identified by its tonotopic organization, and single neurons in that field were studied quantitatively for their sensitivity to the frequency and intensity of tonal stimuli presented via calibrated, sealed stimulating systems. Field P neurons had narrow, V-shaped, threshold frequency tuning curves. At suprathreshold levels, spike counts were generally greatest at frequencies at or close to the neuron's threshold best frequency (BF). Eighty-six percent of posterior-field neurons displayed spike counts that were a nonmonotonic function of the intensity of a BF tone. Of these, over 90% showed at least a 50% reduction in spike count at high stimulus levels, and almost 20% of nonmonotonic cells ceased responding entirely at high stimulus intensities. The nonmonotonic shape of spike count-versus-intensity profiles was typically preserved across the range of frequencies to which any given neuron was responsive. For some neurons, this had the consequence of generating a completely circumscribed frequency-intensity response area. That is, these neurons responded to a tonal stimulus only if the stimulus was within a restricted range of both frequency and intensity. These response areas showed internal organizations that appeared to reflect one or both of two processes. For some neurons, the optimal sound pressure level for spike counts varied with tone frequency, roughly paralleling the threshold tuning curve. For other neurons, the optimal sound pressure level tended to be constant across frequency despite threshold variations of up to 20 dB. The minimum response latencies of posterior-field neurons were generally in the range of 20-50 ms, while cells in the primary auditory cortex (AI) in the same animals generally had minimum latent periods of less than 20 ms. Comparison of these data with those previously presented for neurons in two other cortical auditory fields suggests that the cat's auditory cortex might show an interfield segregation of neurons according to their coding properties.

Animals↗
PubMed1985

Responses of neurons in the cat's superior colliculus to acoustic stimuli. II. A model of interaural intensity sensitivity.

Most neurons in the deep and intermediate layers of the superior colliculus (SC) that respond to acoustic stimuli are sensitive to interaural intensity disparities (IIDs). We examine a model for the generation of sensitivity to IIDs that depends upon temporal coincidence of the inputs from each ear at a given binaural neuron. Because the neural response latency decreases with increasing stimulus intensity, IIDs affect the relative timing of arrival of the inputs. If this model were true, the neurons sensitive to IIDs should also respond to interaural time differences (ITDs) of isointensive stimuli, provided that the magnitude of the delays reflect the neural latency-intensity relationship. For both major classes of binaural cells in the SC, namely those that exhibit binaural inhibition (BI) and binaural facilitation (BF), our results support the model in that the detection of IIDs is largely due to their sensitivity to the temporal overlap of inputs from each ear. The shapes of the IID and ITD functions for each class are similar. The summation of inputs includes inhibitory as well as facilitatory interactions. Estimates of the durations of the subliminal excitatory events in BF cells using the model indicate that they are relatively short (1-4 ms), whereas the durations of the inhibitory processes in BI cells are much longer. The model specifies a common neuronal mechanism for comparison of interaural disparities of time and intensity and does not separate the processing of IIDs and ITDs, as the classic duplex theory suggests. The model provides a physiological explanation for certain features of the psychophysical phenomenon of time-intensity trading. It is also consistent with recent experiments that have shown that the auditory system is sensitive to behaviorally significant ITDs of high-frequency complex signals. The model applies only to the processing of transient stimuli and does not address neural sensitivity to IIDs of continuous high-frequency tones.

Animals↗
PubMed1986

Generator potentials and spike initiation in auditory fibers of goldfish.

Responses were recorded extra- or intracellularly from single, small afferent fibers of the goldfish saccule, that is, S2 fibers, to clarify how the spontaneous and sound-evoked firings in these fibers are initiated from generator potentials. Spontaneously active units randomly selected from the saccular nerve of five goldfish (total 78 units) were classified by the coefficient of variation (CV) of interspike intervals into irregular (59 units; CV greater than 0.3), intermediate (10 units), and regular types (9 units; CV less than 0.23). The irregular type showed a burst (38 units) or random (21 units) pattern of firing in spontaneous activity. In cases where a clear generator potential could be recorded in response to each sound wave, spontaneous generator potentials could also be observed in the absence of stimulus sound. These spontaneous potentials were irregular in amplitude and time course, and often contained components much slower than the sound-evoked generator potentials. The sound-evoked generator potentials in S2 fibers were produced with a delay of 0.4-0.85 ms following each sound wave, and had a time course comparable to the sound-evoked excitatory postsynaptic potentials (EPSPs) of large S1 fibers. They also shared other properties such as adaptive decline in amplitude, incremental and decremental responses, and off-suppression. The mean amplitude of the sound-evoked generator potentials in S2 fibers was linearly related to the rate of afferent firing. There was no apparent difference in the action potential threshold amplitude for spontaneous and sound-evoked generator potentials. It may be concluded that the generator potentials that underlie the spontaneous and sound-evoked firing of S2 fibers are produced by the release of transmitter by hair cells.

Adaptation, Physiological↗