Low-frequency characteristics of human and guinea pig cochleae

Previous physiological studies investigating the transfer of low-frequency sound into the cochlea have been invasive. Predictions about the human cochlea are based on anatomical similarities with animal cochleae but no direct comparison has been possible. This paper presents a noninvasive method of observing low frequency cochlear vibration using distortion product otoacoustic emissions (DPOAE) modulated by low-frequency tones. For various frequencies (15-480 Hz), the level was adjusted to maintain an equal DPOAE-modulation depth, interpreted as a constant basilar membrane displacement amplitude. The resulting modulator level curves from four human ears match equal-loudness contours (ISO226:2003) except for an irregularity consisting of a notch and a peak at 45 Hz and 60 Hz, respectively, suggesting a cochlear resonance. This resonator interacts with the middle ear stiffness. The irregularity separates two regions of the middle ear transfer function in humans: A slope of 12 dB/octave below the irregularity suggests mass-controlled impedance resulting from perilymph movement through the helicotrema; a 6-dB/octave slope above the irregularity suggests resistive cochlear impedance and the existence of a traveling wave. The results from four guinea pig ears showed a 6-dB/octave slope on either side of an irregularity around 120 Hz, and agree with published data.     

The temporal representation of speech in a nonlinear model of the guinea pig cochlea

The temporal representation of speechlike stimuli in the auditory-nerve output of a guinea pig cochlea model is described. The model consists of a bank of dual resonance nonlinear filters that simulate the vibratory response of the basilar membrane followed by a model of the inner hair cell/auditory nerve complex. The model is evaluated by comparing its output with published physiological auditory nerve data in response to single and double vowels. The evaluation includes analyses of individual fibers, as well as ensemble responses over a wide range of best frequencies. In all cases the model response closely follows the patterns in the physiological data, particularly the tendency for the temporal firing pattern of each fiber to represent the frequency of a nearby formant of the speech sound. In the model this behavior is largely a consequence of filter shapes; nonlinear filtering has only a small contribution at low frequencies. The guinea pig cochlear model produces a useful simulation of the measured physiological response to simple speech sounds and is therefore suitable for use in more advanced applications including attempts to generalize these principles to the response of human auditory system, both normal and impaired.     

A macro-mechanical model of the guinea pig cochlea with realistic parameters

The post-mortem transfer function of the cochlea of the guinea pig was compared to the transfer function generated by a model with parameters derived from physical measurements of the guinea pig cochlea. Both the formulation and parameters of the model were carefully chosen to be realistic using evidence from published measurements. The fit between the transfer function of the model and recent mechanical measurements of the passive guinea pig cochlear response was good, with a root mean square ratio of 6.3 dB in amplitude and 0.33 pi rad in phase. The model was used to explore the effect of cochlear partition mode factor and duct geometry upon the mechanical response of the cochlea. Possible inadequacies of the model which could explain the remaining differences between the output of the model and measurements are discussed.     

Perceptually optimized gain function for cochlear implant signal-to-noise ratio based noise reduction

Noise reduction in cochlear implants has achieved significant speech perception improvements through spectral subtraction and signal-to-noise ratio based noise reduction techniques. Current methods use gain functions derived through mathematical optimization or motivated by normal listening psychoacoustic experiments. Although these gain functions have been able to improve speech perception, recent studies have indicated that they are not optimal for cochlear implant noise reduction. This study systematically investigates cochlear implant recipients' speech perception and listening preference of noise reduction with a range of gain functions. Results suggest an advantageous gain function and show that gain functions currently used for noise reduction are not optimal for cochlear implant recipients. Using the cochlear implant optimised gain function, a 27% improvement over the current advanced combination encoder (ACE) stimulation strategy in speech weighted noise and a 7% improvement over current noise reduction strategies were observed in babble noise conditions. The optimized gain function was also most preferred by cochlear implant recipients. The CI specific gain function derived from this study can be easily incorporated into existing noise reduction strategies, to further improve listening performance for CI recipients in challenging environments.     

The representation of steady-state vowel sounds in the temporal discharge patterns of the guinea pig cochlear nerve and primarylike cochlear nucleus neurons

We have recorded the responses of fibers in the cochlear nerve and cells in the cochlear nucleus of the anesthetized guinea pig to synthetic vowels [i], [a], and [u] at 60 and 80 dB SPL. Histograms synchronized to the pitch period of the vowel were constructed, and locking of the discharge to individual harmonics was estimated from these by Fourier transformation. In cochlear nerve fibers from the guinea pig, the responses were similar in all respects to those previously described for the cat. In particular, the average-localized-synchronized-rate functions (ALSR), computed from pooled data, had well-defined peaks corresponding to the formant frequencies of the three vowels at both sound levels. Analysis of the components dominating the discharge could also be used to determine the voice pitch and the frequency of the first formants. We have computed similar population measures over a sample of primarylike cochlear nucleus neurons. In these primarylike cochlear nucleus cell responses, the locking to the higher-frequency formants of the vowels is weaker than in the nerve. This results in a severe degradation of the peaks in the ALSR function at the second and third formant frequencies at least for [i] and [u]. This result is somewhat surprising in light of the reports that primarylike cochlear nucleus cells phaselock, as well as do cochlear nerve fibers.     

Narrow-band AP latencies in normal and recruiting human ears.

Derived narrow-band action potential latencies increase monotonically with decreasing central frequency, and can be interpreted as reflecting the traveling wave delay in the cochlea. It was found that, for recruiting human ears with average flat hearing losses around 40 dB, this accumulating latency increase was smaller than for normal ears. A comparison of 15 normal ears and 37 recruiting ears showed, however, that in only half of the recruiting ears this difference was significant. These recruiting ears were therefore divided in two groups based on the waveform of the narrow-band action potential AP, which correlated well with the subdivision according to latency. The findings have been explained on the basis that latency of the narrow-band APs is not determined solely by the mechanical traveling-wave delay, but also by the response time of the (second?) cochlear filter. When this filter broadens, one expects a decrease in its impulse response time. Since this impulse response time. Since this impulse response depends on the sum of the high- and low-frequency slope values of the cochlear filter, one expects only a latency decrease when the steep high-frequency slope also becomes more shallow. A support for the influence of the response times of the cochlear filter is found in the narrow-band AP latencies for restricted cochlear losses (e.g., in a 4-kHz noise dip). It appears that the latency in that area actually is shorter than for the higher central frequencies, a fact which cannot be explained solely on the basis of a traveling wave phenomenon.     

Distortion product otoacoustic emission input/output functions in normal-hearing and hearing-impaired human ears.

DPOAE input/output (I/O) functions were measured at 7f2 frequencies (1 to 8 kHz; f2/f1 = 1.22) over a range of levels (-5 to 95 dB SPL) in normal-hearing and hearing-impaired human ears. L1-L2 was level dependent in order to produce the largest 2f1-f2 responses in normal ears. System distortion was determined by collecting DP data in six different acoustic cavities. These data were used to derive a multiple linear regression model to predict system distortion levels. The model was tested on cochlear-implant users and used to estimate system distortion in all other ears. At most but not all f2's, measurements in cochlear implant ears were consistent with model predictions. At all f2 frequencies, the ears with normal auditory thresholds produced I/O functions characterized by compressive nonlinear regions at moderate levels, with more rapid growth at low and high stimulus levels. As auditory threshold increased, DPOAE threshold increased, accompanied by DPOAE amplitude reductions, notably over the range of levels where normal ears showed compression. The slope of the I/O function was steeper in impaired ears. The data from normal-hearing ears resembled direct measurements of basilar membrane displacement in lower animals. Data from ears with hearing loss showed that the compressive region was affected by cochlear damage; however, responses at high levels of stimulation resembled those observed in normal ears.     

Effects of middle-ear immaturity on distortion product otoacoustic emission suppression tuning in infant ears

Distortion product otoacoustic emission (DPOAE) measures of cochlear function, including DPOAE suppression tuning curves and input/output (I/O) functions, are not adultlike in human infants. These findings suggest the cochlear amplifier might be functionally immature in newborns. However, many noncochlear factors influence DPOAEs and must be considered. This study examines whether age differences in DPOAE I/O functions recorded from infant and adult ears reflect maturation of ear-canal/middle-ear function or cochlear mechanics. A model based on linear middle-ear transmission and nonlinear cochlear generation was developed to fit the adult DPOAE I/O data. By varying only those model parameters related to middle-ear transmission (and holding cochlear parameters at adult values), the model successfully fitted I/O data from infants at birth through age 6 months. This suggests that cochlear mechanics are mature at birth. The model predicted an attenuation of stimulus energy through the immature ear canal and middle ear, and evaluated whether immaturities in forward transmission could explain the differences consistently observed between infant and adult DPOAE suppression. Results show that once the immaturity was compensated for by providing infants with a relative increase in primary tone level, DPOAE suppression tuning at f2= 6000 Hz was similar in adults and infants.     

Hyperbaric oxygen upregulates cochlear constitutive nitric oxide synthase

Background  

Hyperbaric oxygen therapy (HBOT) is a known adjuvant for treating ischemia-related inner ear diseases. Controversies still exist in the role of HBOT in cochlear diseases. Few studies to date have investigated the cellular changes that occur in inner ears after HBOT. Nitric oxide, which is synthesized by nitric oxide synthase (NOS), is an important signaling molecule in cochlear physiology and pathology. Here we investigated the effects of hyperbaric oxygen on eardrum morphology, cochlear function and expression of NOS isoforms in cochlear substructures after repetitive HBOT in guinea pigs.     

Consonant identification under maskers with sinusoidal modulation: masking release or modulation interference?

The present study investigated the effect of envelope modulations in a background masker on consonant recognition by normal hearing listeners. It is well known that listeners understand speech better under a temporally modulated masker than under a steady masker at the same level, due to masking release. The possibility of an opposite phenomenon, modulation interference, whereby speech recognition could be degraded by a modulated masker due to interference with auditory processing of the speech envelope, was hypothesized and tested under various speech and masker conditions. It was of interest whether modulation interference for speech perception, if it were observed, could be predicted by modulation masking, as found in psychoacoustic studies using nonspeech stimuli. Results revealed that masking release measurably occurred under a variety of conditions, especially when the speech signal maintained a high degree of redundancy across several frequency bands. Modulation interference was also clearly observed under several circumstances when the speech signal did not contain a high redundancy. However, the effect of modulation interference did not follow the expected pattern from psychoacoustic modulation masking results. In conclusion, (1) both factors, modulation interference and masking release, should be accounted for whenever a background masker contains temporal fluctuations, and (2) caution needs to be taken when psychoacoustic theory on modulation masking is applied to speech recognition.     

Growth of masking as a measure of response growth in hearing-impaired listeners

Growth-of-masking functions were obtained from 19 normal and 5 hearing-impaired listeners using a simultaneous-masking paradigm. When masker and probe frequency are identical, the slope of masking approximates 1.0 for both normal-hearing and impaired listeners. For masker frequencies less than or greater than probe frequency, the slopes for impaired listeners are shallower than those of normals. These findings are consistent with previously reported physiological data (single-fiber rate versus level and AP masking functions) for animals with induced cochlear lesions. Results are discussed in terms of a potential masking technique to estimate the growth of response in normal and impaired ears.     

Characterizing auditory processing and perception in individual listeners with sensorineural hearing loss

This study considered consequences of sensorineural hearing loss in ten listeners. The characterization of individual hearing loss was based on psychoacoustic data addressing audiometric pure-tone sensitivity, cochlear compression, frequency selectivity, temporal resolution, and intensity discrimination. In the experiments it was found that listeners with comparable audiograms can show very different results in the supra-threshold measures. In an attempt to account for the observed individual data, a model of auditory signal processing and perception [Jepsen et al., J. Acoust. Soc. Am. 124, 422-438 (2008)] was used as a framework. The parameters of the cochlear processing stage of the model were adjusted to account for behaviorally estimated individual basilar-membrane input-output functions and the audiogram, from which the amounts of inner hair-cell and outer hair-cell losses were estimated as a function of frequency. All other model parameters were left unchanged. The predictions showed a reasonably good agreement with the measured individual data in the frequency selectivity and forward masking conditions while the variation of intensity discrimination thresholds across listeners was underestimated by the model. The model and the associated parameters for individual hearing-impaired listeners might be useful for investigating effects of individual hearing impairment in more complex conditions, such as speech intelligibility in noise.     

Cochlear compression estimates from measurements of distortion-product otoacoustic emissions

Evidence of the compressive growth of basilar-membrane displacement can be seen in distortion-product otoacoustic emission (DPOAE) levels measured as a function of stimulus level. When the levels of the two stimulus tones (f1 and f2) are related by the formula L1 = 39 dB + 0.4 x L2 [Kummer et al., J. Acoust. Soc. Am. 103, 3431-3444 (1998)] the shape of the function relating DPOAE level to L2 is similar (up to an L2 of 70 dB SPL) to the classic Fletcher and Munson [J. Acoust. Soc. Am. 9, 1-10 (1933)] loudness function when plotted on a logarithmic scale. Explicit estimates of compression have been derived based on recent DPOAE measurements from the laboratory. If DPOAE growth rate is defined as the slope of the DPOAE I/O function (in dB/dB), then a cogent definition of compression is the reciprocal of the growth rate. In humans with normal hearing, compression varies from about 1 at threshold to about 4 at 70 dB SPL. With hearing loss, compression is still about 1 at threshold, but grows more slowly above threshold. Median DPOAE I/O data from ears with normal hearing, mild loss, and moderate loss are each well fit by log functions. When the I/O function is logarithmic, then the corresponding compression is a linear function of stimulus level. Evidence of cochlear compression also exists in DPOAE suppression tuning curves, which indicate the level of a third stimulus tone (f3) that reduces DPOAE level by 3 dB. All three stimulus tones generate compressive growth within the cochlea; however, only the relative compression (RC) of the primary and suppressor responses is observable in DPOAE suppression data. An RC value of 1 indicates that the cochlear responses to the primary and suppressor components grow at the same rate. In normal ears, RC rises to 4, when f3 is an octave below f2. The similarities between DPOAE and loudness compression estimates suggest the possibility of predicting loudness growth from DPOAEs; however, intersubject variability makes such predictions difficult at this time.     

Objective evaluation of speech signal quality by the prediction of multiple foreground diagnostic acceptability measure attributes

A methodology is described to objectively diagnose the quality of speech signals by predicting the perceptual detectability of a selected set of distortions. The distortions are a statistically selected subset of the broad number of distortions used in diagnostic acceptability measure (DAM) testing. The justification for such a methodology is established from the analysis of a set of speech signals representing a broad set of distortions and their respective DAM scores. At the heart of the ability to isolate and diagnose the perceptibility of the individual distortions is a physiologically motivated cochlear model. The philosophy and methodology is thus distinct from traditional objective measures that are typically designed to predict mean opinion scores (MOS) using well versed functional psychoacoustic models. Even so, a weighted sum of these objectively predicted set of distortions is able to predict accurate and robust MOS scores, even when the reference speech signals have been subject to the Lombard effect.     

The effects of binaural spectral resolution mismatch on Mandarin speech perception in simulated electric hearing

This study assessed the effects of binaural spectral resolution mismatch on the intelligibility of Mandarin speech in noise using bilateral cochlear implant simulations. Noise-vocoded Mandarin speech, corrupted by speech-shaped noise at 0 and 5?dB signal-to-noise ratios, were presented unilaterally or bilaterally to normal-hearing listeners with mismatched spectral resolution between ears. Significant binaural benefits for Mandarin speech recognition were observed only with matched spectral resolution between ears. In addition, the performance of tone identification was more robust to noise than that of sentence recognition, suggesting factors other than tone identification might account more for the degraded sentence recognition in noise.     

A cochlear model for acoustic emissions

Variability in cochlear emission properties among different species, particularly humans and small mammals, and within individuals in the same species, is modeled by a cochlear nonlinear transmission line. The difference between humans and animals is largely explained by a lower cochlear input impedance in human ears than in cats, gerbils, or chinchillas. Inconstancy in emission properties among individual human or animal subjects is related to structural variability among ears, which can be the result of a nonuniform connection between the outer hair cells cilia and the tectorial membrane. These structural differences are modeled by a nonuniform cochlear partition resistance along the cochlear length. The model predicts that an ear which has a uniform cochlear partition resistance and an adequate cochlear input impedance will emit acoustic distortion products (ADP), but not spontaneous acoustic emission (SAE), nor click-evoked emission (CE). Only a nonuniform cochlea emits SAE and CE in addition to enhanced ADPs. The model predictions agree quantitatively with cochlear emission data from humans and animals.     

Statistical bias in the assessment of binaural benefit relative to the better ear

The comparison of measured binaural performance with the better of two monaural measures (one from each ear) may lead to underestimated binaural benefit due to statistical sampling bias that favors the monaural condition. The mathematical basis of such bias is reviewed and applied to speech reception thresholds measured in 32 bilateral cochlear implant (CI) users for coincident and spatially separated speech and noise. It is shown that the bias increases with test-retest variation and is maximal for uncorrelated samples of identical underlying performance in each ear. When measured differences between ears were assumed to reflect actual underlying performance differences, the bias averaged across the CI users was about 0.2 dB for coincident target and noise, and 0.1 dB for spatially separated conditions. An upper-bound estimate of the bias, based on the assumption that both ears have the same underlying performance and observed differences were due to test-retest variation, was about 0.7 dB regardless of noise location. To the extent that the test-retest variation in these data is comparable to other studies, the results indicate that binaural benefits in bilateral cochlear implant users are not substantially underestimated (on for average) when binaural performance is compared with the better ear in each listening configuration.     

Frequency characteristics of sound transmission in middle ears from Norwegian cattle, and the effect of static pressure differences across the tympanic membrane and the footplate

For 23 cadaver ears from Norwegian cattle, frequency characteristics for the round-window volume displacement relative to the sound pressure at the eardrum have been measured, and are compared to earlier results for human ears [M. Kringlebotn and T. Gundersen, J. Acoust. Soc. Am. 77(1), 159-164 (1985)]. For human as well as for cattle ears, mean amplitude curves have peaks at about 0.7 kHz. At lower frequencies, the mean amplitude for cattle ears is about 5 dB smaller than for human ears. The amplitude curves cross at about 2 kHz, and toward higher frequencies the amplitude for cattle ears becomes increasingly larger. If amplitude curves are roughly approximated by straight lines above 1 kHz, the slope for cattle ears is about -5 dB/octave as compared to about -15 dB/octave for human ears. The phase of the round-window volume displacement lags behind the phase of the sound pressure at the tympanic membrane. The phase lag is close to zero below 0.2 kHz, but increases to about 3.5 pi at 20 kHz for cattle ears, as compared to less than 2 pi for human ears. Further investigations are needed in order to explain the observed differences. Sound transmission in the ear decreases with an increasing static pressure difference across the tympanic membrane, especially at frequencies below 1 kHz, where pressure differences of 10 and 60 cm water cause mean transmission losses of about 10 and 26 dB, respectively, the losses being somewhat larger for overpressures than for underpressures in the ear canal. At higher frequencies, the transmission losses are smaller. For small overpressures, and in a limited frequency range near 3 kHz, even some transmission enhancement may occur. Static pressure variations in the inner ear have only a minor influence on sound transmission. Static pressures relative to the middle ear in the range 0-60 cm water cause mean sound transmission losses less than 5 dB below 1 kHz, and negligible losses at higher frequencies.     

膜迷路积水对豚鼠听觉外周系统振动特性的影响

为探究梅尼埃病引起的膜迷路积水对豚鼠听觉系统振动特性的作用机制,通过药物注射建立了内耳膜迷路积水豚鼠模型模拟梅尼埃病病理,搭建了多普勒激光测振系统对健康和膜迷路积水豚鼠听觉系统的振动特性进行测试研究,得到了健康与膜迷路积水豚鼠的听觉系统振动特性,并通过对比明确了内耳膜迷路积水对豚鼠听觉系统振动特性的影响。实验结果表明,在测试频率范围内,离体豚鼠镫骨及圆窗膜的振动特性与活体豚鼠无明显差异,离体豚鼠听觉系统振动特性与输入声压无关,镫骨及圆窗膜的振动位移与输入声压具有明显的线性关系。膜迷路积水显著降低了豚鼠听觉系统镫骨及圆窗膜的振动幅度,在低频及高频区间尤为明显,在500 Hz处镫骨最高由11.64 nm降低至2.27 nm,圆窗膜由24.97 nm降低至5.05 nm。在10 kHz处,圆窗膜最高由0.45 nm降低至0.03 nm。同时膜迷路积水导致豚鼠耳蜗传递比普遍下降,最高在1 kHz和10 kHz处分别由2.82,2.91下降至1.46,0.28。      

Modeling sensorineural hearing loss. I. Model and retrospective evaluation

The present article describes an approach to the evaluation of psychoacoustic data from the hearing impaired. The results obtained from the hearing impaired in several studies of frequency resolution, temporal resolution, and speech recognition are compared to the results expected for noise-masked normal listeners. It is presumed in this approach that the hypothetical noise-masked normal listeners have masked thresholds that agree perfectly with the quiet thresholds of the hearing-impaired subjects. Using this approach, most of the results obtained from impaired ears on spectral-resolution and speech-recognition tasks could be accurately predicted, an exception being results from spectral-resolution paradigms using fixed-level signals. Some of the data from hearing-impaired listeners on temporal-resolution tasks, on the other hand, could not be adequately described with this approach. The latter data, however, were much more limited. Additional data are needed to better evaluate the adequacy of this approach in describing the performance of the hearing impaired on temporal-resolution tasks.