Maturation of the traveling-wave delay in the human cochlea

The maturation of the traveling-wave delay in the human cochlea was investigated in 227 subjects ranging in age from 29 weeks conceptional age to 49 years by using frequency specific auditory brain-stem responses (ABRs). The derived response technique was applied to ABRs obtained with click stimuli (presented at a fixed level equal to 60-dB sensation level in normal hearing adults) in the presence of high-pass noise masking (slope 96 dB/oct) to obtain frequency specific responses from octave-wide bands. The estimate of traveling-wave delay was obtained by taking the difference between wave I latencies from adjacent derived bands. It was found that the traveling-wave delay between the octave band with center frequency (CF) of 11.3 kHz and that with CF of 5.7 kHz decreased (about 0.4 ms on average) in exponential fashion with age to reach adult values at 3-6 months of age. This decrease was in agreement with reported data in kitten auditory-nerve fibers. The traveling-wave delays between adjacent octave bands with successive lower CF did not change with age.     

Latency of auditory brain-stem responses and otoacoustic emissions using tone-burst stimuli

A comparison of the latency of auditory brain-stem responses (ABR) and evoked otoacoustic emissions (EOAE) has led to an interpretation for the travel of transients in the peripheral auditory system that is consistent with both sets of data. The "cochlear echo" theory for the origin of the EOAE indicates that the latency of a particular frequency component back to the ear canal should be twice the forward latency of its characteristic place in the cochlea. The latency of wave V of the ABR to tone-burst stimuli can be described as the sum of two components: (1) a component that varies with intensity and frequency in an orderly and predictable manner and (2) a component that is independent of both intensity and frequency. Because the EOAE data can be predicted by taking twice the value of component (1) of the ABR latency, this component is interpreted to be due to mechanical travel through the cochlea. A consequence of this interpretation is that the remaining neural component of the ABR latency must be relatively independent of frequency and intensity.     

The effects of sensory hearing loss on cochlear filter times estimated from auditory brainstem response latencies.

Derived-band auditory brainstem responses (ABRs) were obtained in 43 normal-hearing and 80 cochlear hearing-impaired individuals using clicks and high-pass noise masking. The response times across the cochlea [the latency difference between wave V's of the 5.7- and 1.4-kHz center frequency (CF) derived bands] were calculated for five levels of click stimulation ranging from 53 to 93 dB p.-p.e. SPL (23 to 63 dB nHL) in 10-dB steps. Cochlear response times appeared to shorten significantly with hearing loss, especially when the average pure tone (1 to 8 kHz) hearing loss exceeded 30 dB. Examination of derived-band latencies indicates that this shortening is due to a dramatic decrease of wave V latency in the lower CF derived band. Estimates of cochlear filter times in terms of the number of periods to maximum response (Nmax) were calculated from derived-band latencies corrected for gender-dependent cochlear transport and neural conduction times. Nmax decreased as a function of hearing loss, especially for the low CF derived bands. The functions were similar for both males and females. These results are consistent with broader cochlear tuning due to peripheral hearing loss. Estimating filter response times from ABR latencies enhances objective noninvasive diagnosis and allows delineation of the differential effects of pathology on the underlying cochlear mechanisms involved in cochlear transport and filter build-up times.     

Modeling auditory evoked brainstem responses to transient stimuli

A quantitative model is presented that describes the formation of auditory brainstem responses (ABRs) to tone pulses, clicks, and rising chirps as a function of stimulation level. The model computes the convolution of the instantaneous discharge rates using the "humanized" nonlinear auditory-nerve model of Zilany and Bruce [J. Acoust. Soc. Am. 122, 402-417 (2007)] and an empirically derived unitary response function which is assumed to reflect contributions from different cell populations within the auditory brainstem, recorded at a given pair of electrodes on the scalp. It is shown that the model accounts for the decrease of tone-pulse evoked wave-V latency with frequency but underestimates the level dependency of the tone-pulse as well as click-evoked latency values. Furthermore, the model correctly predicts the nonlinear wave-V amplitude behavior in response to the chirp stimulation both as a function of chirp sweeping rate and level. Overall, the results support the hypothesis that the pattern of ABR generation is strongly affected by the nonlinear and dispersive processes in the cochlea.     

Frequency specificity of chirp-evoked auditory brainstem responses

This study examines the usefulness of the upward chirp stimulus developed by Dau et al. [J. Acoust. Soc. Am. 107, 1530-1540 (2000)] for retrieving frequency-specific information. The chirp was designed to produce simultaneous displacement maxima along the cochlear partition by compensating for frequency-dependent traveling-time differences. In the first experiment, auditory brainstem responses (ABR) elicited by the click and the broadband chirp were obtained in the presence of high-pass masking noise, with cutoff frequencies of 0.5, 1, 2, 4, and 8 kHz. Results revealed a larger wave-V amplitude for chirp than for click stimulation in all masking conditions. Wave-V amplitude for the chirp increased continuously with increasing high-pass cutoff frequency while it remains nearly constant for the click for cutoff frequencies greater than 1 kHz. The same two stimuli were tested in the presence of a notched-noise masker with one-octave wide spectral notches corresponding to the cutoff frequencies used in the first experiment. The recordings were compared with derived responses, calculated offline, from the high-pass masking conditions. No significant difference in response amplitude between click and chirp stimulation was found for the notched-noise responses as well as for the derived responses. In the second experiment, responses were obtained using narrow-band stimuli. A low-frequency chirp and a 250-Hz tone pulse with comparable duration and magnitude spectrum were used as stimuli. The narrow-band chirp elicited a larger response amplitude than the tone pulse at low and medium stimulation levels. Overall, the results of the present study further demonstrate the importance of considering peripheral processing for the formation of ABR. The chirp might be of particular interest for assessing low-frequency information.     

Place specific influences on the wave I to V interpeak latency of the auditory brain-stem response.

There is controversy over whether the wave I to V interpeak latency (I-V IPL) of the auditory brain-stem response can be manipulated by cochlear processing. In this study, a forward masking paradigm was used to test the predictions of two contrasting models of I-V IPL generation. The paradigm was designed to determine if the I-V IPL can be affected by masking selected portions of the cochlear response region. The results from ten normal hearing subjects suggest that: (1) waves I and V can be masked semi-independent of each other, and (2) the I-V IPL can be shortened or prolonged by masking the basal or apical portion of the cochlear response region respectively. These findings support the hypothesis that, at least in normal hearing subjects, wave V is biased to reflect more apical cochlear events than wave I. Additionally, they offer tentative support for anecdotal reports of shortened I-V IPLs in the presence of high-frequency hearing loss.     

The effect of broadband noise on the human brain-stem auditory evoked response. III. Anatomic locus

The effects of broadband noise on the brain-stem auditory evoked response (BAER) are reported for two experiments. Experiment 1 used a high-pass subtractive-masking technique and covaried derived bandwidth and continuous broadband noise level. Comparison of responses to half-octave wide derived bands in the presence of within-band noise showed that wave V latency changes were greater than could be explained on the basis of shifts in the cochlear region responsible for generating the response. The magnitude of within-band noise-induced wave V latency shift was independent of the frequency separation of the masker cutoffs. In experiment 2 the effects of noise level and rate on waves I, III, and V of the BAER were evaluated. Peak latencies increased and peak amplitudes decreased with increasing noise level and rate. Higher noise levels and rates produced an increased central (I-V) conduction time in which the wave III-V increase was greater than the wave I-III increase. Together, these results are most consistent with the hypothesis that a nonplace, central auditory mechanism produces most of the noise-induced latency shifts in normal-hearing adults.     

Automated threshold detection for auditory brainstem responses: comparison with visual estimation in a stem cell transplantation study

Background  

Auditory brainstem responses (ABRs) are used to study auditory acuity in animal-based medical research. ABRs are evoked by acoustic stimuli, and consist of an electrical signal resulting from summated activity in the auditory nerve and brainstem nuclei. ABR analysis determines the sound intensity at which a neural response first appears (hearing threshold). Traditionally, threshold has been assessed by visual estimation of a series of ABRs evoked by different sound intensities. Here we develop an automated threshold detection method that eliminates the variability and subjectivity associated with visual estimation.     

Anomalous phase relations in threshold-level responses from gerbil auditory nerve fibers

Phase-locked responses at near-threshold levels obtained from single units in the auditory nerve of gerbils show that the relation between phase lag and linear frequency often contains an unexpected microstructure. In frequency regions below 1 kHz that are also an octave or more below CF, phase curves often have multiple straight-line segments, rapid slope changes, or other major inflections. These detailed features of the phase curves are not accounted for by any identified artifact. For example, the anomalous features cannot be removed simply by lowering the stimulus level; they remain down to levels where phase locking first occurs. Phase curves for single units with the same CF recorded from different animals can have similar microstructures, suggesting that the form of the phase curves may reflect some joint effect of stimulus frequency and CF or longitudinal position.     

Mapping auditory nerve firing density using high-level compound action potentials and high-pass noise masking

Future implementation of regenerative treatments for sensorineural hearing loss may be hindered by the lack of diagnostic tools that specify the target(s) within the cochlea and auditory nerve for delivery of therapeutic agents. Recent research has indicated that the amplitude of high-level compound action potentials (CAPs) is a good predictor of overall auditory nerve survival, but does not pinpoint the location of neural damage. A location-specific estimate of nerve pathology may be possible by using a masking paradigm and high-level CAPs to map auditory nerve firing density throughout the cochlea. This initial study in gerbil utilized a high-pass masking paradigm to determine normative ranges for CAP-derived neural firing density functions using broadband chirp stimuli and low-frequency tonebursts, and to determine if cochlear outer hair cell (OHC) pathology alters the distribution of neural firing in the cochlea. Neural firing distributions for moderate-intensity (60 dB pSPL) chirps were affected by OHC pathology whereas those derived with high-level (90 dB pSPL) chirps were not. These results suggest that CAP-derived neural firing distributions for high-level chirps may provide an estimate of auditory nerve survival that is independent of OHC pathology.     

The effects of external- and middle-ear filtering on auditory threshold and noise-induced hearing loss

A model of external- and middle-ear function is described that uses existing data to quantify the flow of sound power from the environment to the cochlea of humans, cats, and chinchillas. This model estimates the sound power produced at the entrance of the cochlea by an environmental sound stimulus, and can be used to predict the shape of the auditory threshold function and the relative potency of various traumatic acoustic stimuli. The shapes of the predicted and measured threshold functions in the three species are similar in best frequency, bandwidth, and low-frequency slope, and the model accurately predicts the hypersensitivity of the middle-frequency regions of the cochlea to acoustic trauma. The model assumes that the mechanics of the middle-ear system are linear even at high stimulus levels and does not include the effects of either middle-ear or cochlear efferent loops. The effects of these simplifications on the model are discussed as are the implications of the model results for hearing protection and damage risk criteria.     

The possible role of the cochlear frequency-position map in auditory signal coding

The frequency-position map in the cochlea is considered as part of a signal decomposition mechanism whose purpose is redundancy reduction and information compression of auditory stimuli. A logarithmic-type frequency distribution is analytically derived from a heuristic model of the autocorrelation function of auditory signals and confronted with empirical data from the cat cochlea. It is argued that the logarithmic organization of frequency along the cochlea is designed to extract the maximum amount of information with the minimum number of hair cells.     

Ear asymmetries in middle-ear, cochlear, and brainstem responses in human infants

In 2004, Sininger and Cone-Wesson examined asymmetries in the signal-to-noise ratio (SNR) of otoacoustic emissions (OAE) in infants, reporting that distortion-product (DP)OAE SNR was larger in the left ear, whereas transient-evoked (TE)OAE SNR was larger in the right. They proposed that cochlear and brainstem asymmetries facilitate development of brain-hemispheric specialization for sound processing. Similarly, in 2006 Sininger and Cone-Wesson described ear asymmetries mainly favoring the right ear in infant auditory brainstem responses (ABRs). The present study analyzed 2640 infant responses to further explore these effects. Ear differences in OAE SNR, signal, and noise were evaluated separately and across frequencies (1.5, 2, 3, and 4 kHz), and ABR asymmetries were compared with cochlear asymmetries. Analyses of ear-canal reflectance and admittance showed that asymmetries in middle-ear functioning did not explain cochlear and brainstem asymmetries. Current results are consistent with earlier studies showing right-ear dominance for TEOAE and ABR. Noise levels were higher in the right ear for OAEs and ABRs, causing ear asymmetries in SNR to differ from those in signal level. No left-ear dominance for DPOAE signal was observed. These results do not support a theory that ear asymmetries in cochlear processing mimic hemispheric brain specialization for auditory processing.     

A nonlinear filter-bank model of the guinea-pig cochlear nerve: rate responses

The aim of this study is to produce a functional model of the auditory nerve (AN) response of the guinea-pig that reproduces a wide range of important responses to auditory stimulation. The model is intended for use as an input to larger scale models of auditory processing in the brain-stem. A dual-resonance nonlinear filter architecture is used to reproduce the mechanical tuning of the cochlea. Transduction to the activity on the AN is accomplished with a recently proposed model of the inner-hair-cell. Together, these models have been shown to be able to reproduce the response of high-, medium-, and low-spontaneous rate fibers from the guinea-pig AN at high best frequencies (BFs). In this study we generate parameters that allow us to fit the AN model to data from a wide range of BFs. By varying the characteristics of the mechanical filtering as a function of the BF it was possible to reproduce the BF dependence of frequency-threshold tuning curves, AN rate-intensity functions at and away from BF, compression of the basilar membrane at BF as inferred from AN responses, and AN iso-intensity functions. The model is a convenient computational tool for the simulation of the range of nonlinear tuning and rate-responses found across the length of the guinea-pig cochlear nerve.     

Binaural interference and auditory grouping

The phenomenon of binaural interference, where binaural judgments of a high-frequency target stimulus are disrupted by the presence of a simultaneous low-frequency interferer, can largely be explained using principles of auditory grouping and segregation. Evidence for this relationship comes from a number of previous studies showing that the manipulation of simultaneous grouping cues such as harmonicity and onset synchrony can influence the strength of the phenomenon. In this study, it is shown that sequential grouping cues can also influence whether binaural interference occurs. Subjects indicated the lateral position of a high-frequency sinusoidally amplitude-modulated (SAM) tone containing an interaural time difference. Perceived lateral positions were reduced by the presence of a simultaneous diotic low-frequency SAM tone, but were largely restored when the interferer was "captured" in a stream of identical tones. A control condition confirmed that the effect was not due to peripheral adaptation. The data lend further support to the idea that binaural interference is affected by processes related to the perceptual organization of auditory information. Modifications to existing grouping-based models are proposed that may help account for binaural interference effects more successfully.     

Speech coding in the auditory nerve: V. Vowels in background noise

Responses of auditory-nerve fibers to steady-state, two-formant vowels in low-pass background noise (S/N = 10 dB) were obtained in anesthetized cats. For fibers over a wide range of characteristic frequencies (CFs), the peaks in discharge rate at the onset of the vowel stimuli were nearly eliminated in the presence of noise. In contrast, strong effects of noise on fine time patterns of discharge were limited to CF regions that are far from the formant frequencies. One effect is a reduction in the amplitude of the response component at the fundamental frequency in the high-CF regions and for CFs between F1 and F2 when the formants are widely separated. A reduction in the amplitude of the response components at the formant frequencies, with concomitant increase in components near CF or low-frequency components occurs in CF regions where the signal-to-noise ratio is particularly low. The processing schemes that were effective for estimating the formant frequencies and fundamental frequency of vowels in quiet generally remain adequate in moderate-level background noise. Overall, the discharge patterns contain many cues for distinctions among the vowel stimuli, so that the central processor should be able to identify the different vowels, consistent with psychophysical performance at moderate signal-to-noise ratios.     

Frequency- and level-dependent changes in auditory brainstem responses (ABRS) in developing mice

The development of the auditory brainstem response was studied to quantitatively assess its dependence on stimulus frequency and level. Responses were not observed to stimuli > or =16 kHz on P12, however, the full range of responsive frequencies included in the study was observed by P14. Response thresholds were high on P12, exceeding 100 dB SPL for all stimuli tested. The rate of threshold development increased progressively for stimulus frequencies between -2 and 10 kHz, with the most rapid changes occurring at frequencies >10 kHz. Adultlike thresholds were observed by P18. Response latencies and interpeak intervals matured rapidly over the course of the second and third postnatal weeks and did not achieve adultlike characteristics until after P18. Latencies of higher-order peaks were progressively and sequentially delayed relative to wave I. Wave I amplitudes developed nonmonotonically, growing during the first 24 days and stabilizing at adult values by approximately P36. Slopes of wave I amplitude-and latency-level curves were significantly steeper than those of adults during the neonatal period and the outcome of input-output analyses, as well as frequency-specific maturational profiles, support developmental models in which function initially matures in the mid-frequency range and proceeds, simultaneously, in both apical and basal directions.     

Responses to amplitude-modulated tones in the auditory nerve of the cat.

Sinusoidally amplitude-modulated (AM) tones are frequently used in psychophysical and physiological studies, yet a comprehensive study on the coding of AM tones in the auditory nerve is lacking. AM responses of single auditory-nerve fibers of the cat are studied, systematically varying modulation depth, frequency, and sound level. Synchrony-level functions were nonmonotonic with maximum values that were inversely correlated with spontaneous rate (SR). In most fibers, envelope phase-locking showed a positive gain. Modulation transfer functions were uniformly low pass. Their corner frequency increased with characteristic frequency (CF), but changed little for CFs above 10 kHz. The highest modulation frequencies to which phase locking occurred were more than 0.8 oct lower than the highest frequencies to which phase locking to pure tones occurs. Cumulative, or unwrapped, phase increased linearly with modulation frequency: The slope was inversely related to CF, and slightly higher than group delays reported for pure tones. High SR, low CF fibers showed the poorest envelope phase locking. In some low CF fibers, phase locking increased at high levels, associated with "peak-splitting" phenomena. Changes in average rate due to modulation were small, and could be enhancement or suppression.     

听觉模型输出谱特征在声目标识别中的应用

利用模拟人耳声信号处理过程的CcGC滤波器组模型,研究了听觉特征应用于声目标识别相比传统特征的优势。结果表明:当信号的信噪比下降时,听觉特征逐渐表现出更好的性能,体现了听觉系统优异的抗噪声能力。随后,本文从CcGC滤波器组模型所反映的听觉系统四个主要特性入手,通过仿真实验研究了耳蜗抑制噪声的机理,结果表明听觉系统的临界带划分和非线性压缩在耳蜗抑制噪声中起着关键的作用。      

Killer whale (Orcinus orca) hearing: auditory brainstem response and behavioral audiograms.

Killer whale (Orcinus orca) audiograms were measured using behavioral responses and auditory evoked potentials (AEPs) from two trained adult females. The mean auditory brainstem response (ABR) audiogram to tones between 1 and 100 kHz was 12 dB (re 1 mu Pa) less sensitive than behavioral audiograms from the same individuals (+/- 8 dB). The ABR and behavioral audiogram curves had shapes that were generally consistent and had the best threshold agreement (5 dB) in the most sensitive range 18-42 kHz, and the least (22 dB) at higher frequencies 60-100 kHz. The most sensitive frequency in the mean Orcinus audiogram was 20 kHz (36 dB), a frequency lower than many other odontocetes, but one that matches peak spectral energy reported for wild killer whale echolocation clicks. A previously reported audiogram of a male Orcinus had greatest sensitivity in this range (15 kHz, approximately 35 dB). Both whales reliably responded to 100-kHz tones (95 dB), and one whale to a 120-kHz tone, a variation from an earlier reported high-frequency limit of 32 kHz for a male Orcinus. Despite smaller amplitude ABRs than smaller delphinids, the results demonstrated that ABR audiometry can provide a useful suprathreshold estimate of hearing range in toothed whales.