The dynamic range of inner hair cell and organ of Corti responses

Inner hair cell (IHC) and organ of Corti (OC) responses are measured from the apical three turns of the guinea pig cochlea, allowing access to regions with best, or most sensitive, frequencies at approximately 250, 1000, and 4000 Hz. In addition to measuring both ac and dc receptor potentials, the average value of the half-wave rectified response (AVEHR) is computed to better reflect the signal that induces transmitter release. This measure facilitates comparisons with single-unit responses in the auditory nerve. Although IHC ac responses exhibit compressive growth, response magnitudes at high levels depend on stimulus frequency. For example, IHCs with moderate and high best frequencies (BF) exhibit more linear responses below the BF of the cell, where higher sound-pressure levels are required to approach saturation. Because a similar frequency dependence is observed in extracellular OC responses, this phenomenon may originate in cochlear mechanics. At the most apical recording location, however, the pattern documented at the base of the cochlea is not seen in IHCs with low BFs around 250 Hz. In fact, more linear behavior is measured above the BF of the cell. These frequency-dependent features require modification of cochlear models that do not provide for longitudinal variations and generally depend on a single stage of saturation located at the synapse. Finally, behavior of dc and AVEHR responses suggests that a single IHC is capable of coding intensity over a large dynamic range [Patuzzi and Sellick, J. Acoust. Soc. Am. 74, 1734-1741 (1983); Smith et al., in Hearing--Physiological Bases and Psychophysics (Springer, Berlin, 1983); Smith, in Auditory Function (Wiley, New York, 1988)] and that information compiled over wide areas along the cochlear partition is not essential for loudness perception, consistent with psychophysical results [Viemeister, Hearing Res. 34, 267-274 (1988)].     

Contribution of low-frequency acoustic information to Chinese speech recognition in cochlear implant simulations

Chinese sentence recognition strongly relates to the reception of tonal information. For cochlear implant (CI) users with residual acoustic hearing, tonal information may be enhanced by restoring low-frequency acoustic cues in the nonimplanted ear. The present study investigated the contribution of low-frequency acoustic information to Chinese speech recognition in Mandarin-speaking normal-hearing subjects listening to acoustic simulations of bilaterally combined electric and acoustic hearing. Subjects listened to a 6-channel CI simulation in one ear and low-pass filtered speech in the other ear. Chinese tone, phoneme, and sentence recognition were measured in steady-state, speech-shaped noise, as a function of the cutoff frequency for low-pass filtered speech. Results showed that low-frequency acoustic information below 500 Hz contributed most strongly to tone recognition, while low-frequency acoustic information above 500 Hz contributed most strongly to phoneme recognition. For Chinese sentences, speech reception thresholds (SRTs) improved with increasing amounts of low-frequency acoustic information, and significantly improved when low-frequency acoustic information above 500 Hz was preserved. SRTs were not significantly affected by the degree of spectral overlap between the CI simulation and low-pass filtered speech. These results suggest that, for CI patients with residual acoustic hearing, preserving low-frequency acoustic information can improve Chinese speech recognition in noise.     

Response dynamics of goldfish saccular fibers: effects of stimulus frequency and intensity on fibers with different tuning, sensitivity, and spontaneous activity

The effects of stimulus frequency and intensity on response patterns (PST histograms) to tone burst stimulation were examined in differently tuned saccular fibers of the goldfish. In addition, the sensitivity of these fibers to amplitude-modulated (AM) signals of different carrier frequencies was measured. The response patterns evoked by unmodulated signals were a complex function of tuning, spontaneous activity and sensitivity of the fiber, and the frequency and intensity of the signal. Frequency-dependent response patterns were found in low-frequency fibers with best frequencies (BF) below 200 Hz. Responses in these fibers ranged from tonic to phasic in nonspontaneous fibers and included more complex patterns in spontaneously active fibers, such as suppression of evoked activity below spontaneous levels. Midfrequency fibers (BF = 500-600 Hz) showed responses similar to those in low-frequency fibers, but with less dependence on frequency. In contrast, both high-frequency (BF = 800-1000 Hz) and wideband, untuned fibers showed frequency-invariant patterns of adaptation. High-frequency fibers were equally sensitive to AM signals at all frequencies tested. The sensitivity of low-frequency fibers to AM, however, increased as a function of carrier frequency and corresponded to the degree of adaptation in response to unmodulated tones. In general, the AM sensitivity of a fiber could be predicted more by its pattern of response to unmodulated signals than by its tuning characteristics.     

Nonlinearities in cochlear receptor potentials and their origins

Using intracellular recording methods in vivo [P. Dallos, J. Neurosci. 5, 1591-1608 (1985)], various nonlinear characteristics of receptor potentials from hair cells located in the low-frequency region of the guinea pig cochlea have been examined. Patterns of saturation for ac and dc response components obtained from Fourier analysis and directly from averaged waveforms are studied. Growth patterns of lower harmonic components are investigated and the interesting nonmonotonic properties of even harmonics noted. The latter are seen in both inner and outer hair cell responses, primarily with stimuli near the cells' best frequency. Fundamental ac and the dc potentials occasionally exhibit nonmonotonic growth. These patterns are studied and their occurrence in inner and outer hair cell responses considered.     

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.     

Auditory filter shapes at 8 and 10 kHz

Auditory filter shapes were derived from notched-noise masking data at center frequencies of 8 kHz (for three spectrum levels, N0 = 20, 35, and 50 dB) and 10 kHz (N0 = 50 dB). In order to minimize variability due to earphone placement, insert earphones (Etymotic Research ER2) were used and individual earmolds were made for each subject. These earphones were designed to give a flat frequency response at the eardrum for frequencies up to 14 kHz. The filter shapes were derived under the assumption that a frequency-dependent attenuation was applied to all stimuli before reaching the filter; this attenuation function was estimated from the variation of absolute threshold with frequency for the three youngest normally hearing subjects in our experiments. At 8 kHz, the mean equivalent rectangular bandwidths (ERBs) of the filters derived from the individual data for three subjects were 677, 637, and 1011 Hz for N0 = 20, 35, and 50 dB, respectively. The filters at N0 = 50 dB were roughly symmetrical, while, at the lower spectrum levels, the low-frequency skirt was steeper than the high-frequency skirt. The mean ERB at 10 kHz was 957 Hz. At this frequency, the filters for two subjects were steeper on the high-frequency side than the low-frequency side, while the third subject showed a slight asymmetry in the opposite direction.     

Speech recognition in noise as a function of highpass-filter cutoff frequency for people with and without low-frequency cochlear dead regions

Regions in the cochlea with very few functioning inner hair cells and/or neurons are called "dead regions" (DRs). Previously, we measured the recognition of highpass-filtered nonsense syllables as a function of filter cutoff frequency for hearing-impaired people with and without low-frequency (apical) DRs [J. Acoust. Soc. Am. 122, 542-553 (2007)]. DRs were diagnosed using the TEN(HL) test, and psychophysical tuning curves were used to define the edge frequency (fe) more precisely. Stimuli were amplified differently for each ear, using the "Cambridge formula." The present study was similar, but the speech was presented in speech-shaped noise at a signal-to-noise ratio of 3 dB. For subjects with low-frequency hearing loss but without DRs, scores were high (65-80%) for low cutoff frequencies and worsened with increasing cutoff frequency above about 430 Hz. For subjects with low-frequency DRs, performance was poor (20-40%) for the lowest cutoff frequency, improved with increasing cutoff frequency up to about 0.56fe, and then worsened. As for speech in quiet, these results indicate that people with low-frequency DRs are able to make effective use of frequency components that fall in the range 0.56fe to fe, but that frequency components below 0.56fe have deleterious effects.     

Testing the concept of softness imperception: loudness near threshold for hearing-impaired ears

Buus and Florentine [J. Assoc. Res. Otolaryngol. 3, 120-139 (2002)] have proposed that loudness recruitment in cases of cochlear hearing loss is caused partly by an abnormally large loudness at absolute threshold. This has been called "softness imperception." To evaluate this idea, loudness-matching functions were obtained using tones at very low sensation levels. For subjects with asymmetrical hearing loss, matches were obtained for a single frequency across ears. For subjects with sloping hearing loss, matches were obtained between tones at two frequencies, one where the absolute threshold was nearly normal and one where there was a moderate hearing loss. Loudness matching was possible for sensation levels (SLs) as low as 2 dB. When the fixed tone was presented at a very low SL in an ear (or at a frequency) where there was hearing impairment, it was matched by a tone with approximately the same SL in an ear (or at a frequency) where hearing was normal (e.g., 2 dB SL matched 2 dB SL). This relationship held for SLs up to 4-10 dB, depending on the subject. These results are not consistent with the concept of softness imperception.     

Discrimination of interaural temporal disparities by normal-hearing listeners and listeners with high-frequency sensorineural hearing loss

Thresholds of ongoing interaural time difference (ITD) were obtained from normal-hearing and hearing-impaired listeners who had high-frequency, sensorineural hearing loss. Several stimuli (a 500-Hz sinusoid, a narrow-band noise centered at 500 Hz, a sinusoidally amplitude-modulated 4000-Hz tone, and a narrow-band noise centered at 4000 Hz) and two criteria [equal sound-pressure level (Eq SPL) and equal sensation level (Eq SL)] for determining the level of stimuli presented to each listener were employed. The ITD thresholds and slopes of the psychometric functions were elevated for hearing-impaired listeners for the two high-frequency stimuli in comparison to: the listener's own low-frequency thresholds; and data obtained from normal-hearing listeners for stimuli presented with Eq SPL interaurally. The two groups of listeners required similar ITDs to reach threshold when stimuli were presented at Eq SLs to each ear. For low-frequency stimuli, the ITD thresholds of the hearing-impaired listener were generally slightly greater than those obtained from the normal-hearing listeners. Whether these stimuli were presented at either Eq SPL or Eq SL did not differentially affect the ITD thresholds across groups.     

Acoustic response and tuning in saccular nerve fibers of the goldfish (Carassius auratus)

The acoustic frequency selectivity of over 500 saccular nerve fibers of the goldfish was studied using automated threshold tracking based on spike rate increments defined statistically. Saccular fibers of the goldfish show great variation in (1) best sensitivity (-26 to + 35 dB re: 1 dyn/cm2), (2) best frequency (below 100 to 1770 Hz), (3) spontaneous rate (0 to over 200 spikes/s), (4) spontaneous type (silent, regular, irregular, burst), and (5) degree of tuning (Q 10 dB from less than 0.1 to 2). Saccular fibers may be grouped into four nonoverlapping categories based on tuning and best frequency: (1) untuned (less than 10-dB variation in sensitivity between 100 and 1000 Hz), (2) low frequency (BF from below 120 to 290 Hz), (3) midfrequency (BF between 330 and 670 Hz), and (4) high frequency (BF between 790 and 1770 Hz). Within each category, all spontaneous rates and types, and all degrees of tuning can be observed. The least sensitive fibers within each group have zero spontaneous rates. The goldfish is like all other vertebrates studied in that the peripheral auditory system is adapted for frequency selectivity throughout the animal's entire frequency range of hearing. Peripheral tuning most likely accounts for behavioral determinations of the "auditory filter" and for the detectability of signals masked by noise. The signal-to-noise ratio enhancement provided by these peripheral filters is likely to be of primary biological significance. A "place principle" of sound quality analysis based on lines "labeled" according to best frequency in the brain cannot be ruled out on the basis of the peripheral physiology.     

Middle-ear response in the chinchilla and its relationship to mechanics at the base of the cochlea

The responses of the malleus and the stapes to sinusoidal acoustic stimulation have been measured in the middle ears of anesthetized chinchillas using the M?ssbauer technique. With "intact" bullas (i.e., closed except for venting via capillary tubing), the vibrations of the tip of the malleus reach a maximal peak velocity of about 2 mm/s in responses to 100-dB SPL tones in the frequency range 500-6000 Hz; vibration velocity diminishes toward lower frequencies with a slope of about 6 dB/oct. Opening the bulla widely increases the responses to low-frequency stimuli by as much as 16 dB. At low frequencies, malleus response sensitivity with either open or intact bullas far exceeds all previous measurements in cats and matches or exceeds such measurements in guinea pigs. Whether measured in open or intact bullas, phase-versus-frequency curves closely approximate those predicted from the magnitude-versus-frequency curves by minimum phase theory. The stapes responses are similar to those of the malleus, except that stapes response magnitude is lower, on the average, by 7.5 dB at frequencies below 2 kHz and 10.7 dB at 2 kHz and above. Comparison of the responses of the middle ear with those of the basilar membrane at a site 3.5 mm from the stapes indicates that, at frequencies below 150 Hz, the basilar membrane displacement is proportional to stapes acceleration. At frequencies between 150 and 2000 Hz, basilar membrane displacement is proportional to stapes velocity.     

Group delay measurement from spiral ganglion cells in the basal turn of the guinea pig cochlea

Measurements of group delay were made extracellularly from spiral ganglion cells in the 3.7 to 5.0-mm region of the guinea pig cochlea, using sinusoidally amplitude modulated tones with constant modulating frequency (100 Hz) and depth of modulation (0.19). Threshold cochlear tuning was accompanied by frequency-dependent group delays. The group delay on the low-frequency tail was independent of carrier frequency; the interunit variation was 0.28-1.28 ms. The difference in group delay between CF and the low-frequency tail decreased as the CF threshold increased (-0.09 +/- 0.02 ms per 10 dB, beginning at 0.62 +/- 0.07 ms at 0 dB SPL). The group delay decreased above CF; at the units' maximum frequency it was less than the low-frequency tail value, and was sometimes negative. Following arterial injections of furosemide the CF threshold increased and the group delay peak decreased; the low-frequency tail was unaffected. The group delay decreased with increasing intensity; the reduction near and above CF was not only larger than that on the low-frequency tail, but also the change at 5-10 dB above threshold was far greater than expected from the Q10dB of the suprathreshold iso-rate tuning curves. A minimum-phase analysis suggested that the group delay response above CF, together with its nonlinear behavior, can be accounted for by a high-frequency, level-independent, amplitude plateau, in combination with the single unit, amplitude nonlinearity which is known to exist above CF.     

Temporal masking functions for listeners with real and simulated hearing loss

A functional simulation of hearing loss was evaluated in its ability to reproduce the temporal masking functions for eight listeners with mild to severe sensorineural hearing loss. Each audiometric loss was simulated in a group of age-matched normal-hearing listeners through a combination of spectrally-shaped masking noise and multi-band expansion. Temporal-masking functions were obtained in both groups of listeners using a forward-masking paradigm in which the level of a 110-ms masker required to just mask a 10-ms fixed-level probe (5-10 dB SL) was measured as a function of the time delay between the masker offset and probe onset. At each of four probe frequencies (500, 1000, 2000, and 4000 Hz), temporal-masking functions were obtained using maskers that were 0.55, 1.0, and 1.15 times the probe frequency. The slopes and y-intercepts of the masking functions were not significantly different for listeners with real and simulated hearing loss. The y-intercepts were positively correlated with level of hearing loss while the slopes were negatively correlated. The ratio of the slopes obtained with the low-frequency maskers relative to the on-frequency maskers was similar for both groups of listeners and indicated a smaller compressive effect than that observed in normal-hearing listeners.     

Improved speech recognition in noise in simulated binaurally combined acoustic and electric stimulation

Speech recognition in noise improves with combined acoustic and electric stimulation compared to electric stimulation alone [Kong et al., J. Acoust. Soc. Am. 117, 1351-1361 (2005)]. Here the contribution of fundamental frequency (F0) and low-frequency phonetic cues to speech recognition in combined hearing was investigated. Normal-hearing listeners heard vocoded speech in one ear and low-pass (LP) filtered speech in the other. Three listening conditions (vocode-alone, LP-alone, combined) were investigated. Target speech (average F0=120 Hz) was mixed with a time-reversed masker (average F0=172 Hz) at three signal-to-noise ratios (SNRs). LP speech aided performance at all SNRs. Low-frequency phonetic cues were then removed by replacing the LP speech with a LP equal-amplitude harmonic complex, frequency and amplitude modulated by the F0 and temporal envelope of voiced segments of the target. The combined hearing advantage disappeared at 10 and 15 dB SNR, but persisted at 5 dB SNR. A similar finding occurred when, additionally, F0 contour cues were removed. These results are consistent with a role for low-frequency phonetic cues, but not with a combination of F0 information between the two ears. The enhanced performance at 5 dB SNR with F0 contour cues absent suggests that voicing or glimpsing cues may be responsible for the combined hearing benefit.     

Development of form and function in peripheral auditory structures of the zebrafish (Danio rerio)

Investigations of the development of auditory form and function have, with a few exceptions, thus far been largely restricted to birds and mammals, making it difficult to postulate evolutionary hypotheses. Teleost fishes represent useful models for developmental investigations of the auditory system due to their often extensive period of posthatching development and the diversity of auditory specializations in this group. Using the auditory brainstem response and morphological techniques we investigated the development of auditory form and function in zebrafish (Danio rerio) ranging in size from 10 to 45 mm total length. We found no difference in auditory sensitivity, response latency, or response amplitude with development, but we did find an expansion of maximum detectable frequency from 200 Hz at 10 mm to 4000 Hz at 45 mm TL. The expansion of frequency range coincided with the development of Weberian ossicles in zebrafish, suggesting that changes in hearing ability in this species are driven more by development of auxiliary specializations than by the ear itself. We propose a model for the development of zebrafish hearing wherein the Weberian ossicles gradually increase the range of frequencies available to the inner ear, much as middle ear development increases frequency range in mammals.     

Measurement of acoustic distortion reveals underlying similarities between human and rodent mechanical responses

The level of 2f1-f2 acoustic distortion product (ADP) measured in the meatus during two-tone stimulation was compared with N 1 thresholds measured at the round window for the guinea pig. A significant inverse relation was found between distortion level and N 1 threshold. A similar relationship has also been reported for ADP level and subjective thresholds in half the human ears measured [S.A. Gaskill and A.M. Brown, J. Acoust. Soc. Am. 88, 821-839 (1990)]. Guinea pig and human ADP levels behave similarly in response to varying stimulus parameters. The ADP levels grow to a maximum and decline with increasing stimulus separation. The decline is steeper in the human ear. In both species, ADP growth as a function of stimulus level is approximately 1 with covaried stimuli; more gradual with the level of f2 (L 2) alone increasing and steeper when the level of f1 (L 1) alone is increased. The latter slopes are strongly influenced by the level of the stationary L 2 and are less steep in the human ear. A link has been proposed between differences in ADP behavior and differences in auditory filter bandwidth in the two species. Guinea pigs show little intersubject variability in ADP level. They do not show the fine structure in distortion level across frequency or the variation in growth rate seen in human responses. Differences in organ of Corti fine structure may underly these differences.     

Psychophysical estimates of level-dependent best-frequency shifts in the apical region of the human basilar membrane

It is now undisputed that the best frequency (BF) of basal basilar-membrane (BM) sites shifts downwards as the stimulus level increases. The direction of the shift for apical sites is, by contrast, less well established. Auditory nerve studies suggest that the BF shifts in opposite directions for apical and basal BM sites with increasing stimulus level. This study attempts to determine if this is the case in humans. Psychophysical tuning curves (PTCs) were measured using forward masking for probe frequencies of 125, 250, 500, and 6000 Hz. The level of a masker tone required to just mask a fixed low-level probe tone was measured for different masker-probe time intervals. The duration of the intervals was adjusted as necessary to obtain PTCs for the widest possible range of masker levels. The BF was identified from function fits to the measured PTCs and it almost always decreased with increasing level. This result is inconsistent with most auditory-nerve observations obtained from other mammals. Several explanations are discussed, including that it may be erroneous to assume that low-frequency PTCs reflect the tuning of apical BM sites exclusively and that the inherent frequency response of the inner hair cell may account for the discrepancy.     

Psychophysical tuning curves at very high frequencies

For most normal-hearing listeners, absolute thresholds increase rapidly above about 16 kHz. One hypothesis is that the high-frequency limit of the hearing-threshold curve is imposed by the transmission characteristics of the middle ear, which attenuates the sound input [Masterton et al., J. Acoust. Soc. Am. 45, 966-985 (1969)]. An alternative hypothesis is that the high-frequency limit of hearing is imposed by the tonotopicity of the cochlea [Ruggero and Temchin, Proc. Nat. Acad. Sci. U.S.A. 99, 13206-13210 (2002)]. The aim of this study was to test these hypotheses. Forward-masked psychophysical tuning curves (PTCs) were derived for signal frequencies of 12-17.5 kHz. For the highest signal frequencies, the high-frequency slopes of some PTCs were steeper than the slope of the hearing-threshold curve. The results also show that the human auditory system displays frequency selectivity for characteristic frequencies (CFs) as high as 17 kHz, above the frequency at which absolute thresholds begin to increase rapidly. The findings suggest that, for CFs up to 17 kHz, the high-frequency limitation in humans is imposed in part by the middle-ear attenuation, and not by the tonotopicity of the cochlea.     

Risk factors for hearing loss at different frequencies in a population of 47,388 noise-exposed workers.

Weighted regression analysis was applied to determine the dependence of the hearing thresholds of 47,388 noise-exposed workers on age, sex, noise immission level, ear disease, head injury, tinnitus, hearing protector usage, and audiometric frequency in the range from 0.5 to 6 kHz. It could be shown that the hearing thresholds at any frequency are dominated by the age of the worker and that women, after equivalent exposure conditions, hear better than men. The relative effects of sex, noise immission level, ear diseases, tinnitus, and hearing protector usage are related to the audiometric frequency. Users of hearing protectors at the last audiometric investigation hear worse than nonusers. Hearing protector usage is strongly related with the hearing threshold in the low-frequency range. The noise immission level does not noticeably affect the hearing threshold below 3 kHz. The most important frequency of the noise immission level is as expected 4 kHz. For 4 kHz, it was shown that the variables age, noise immission level, tinnitus, head injuries, and ear diseases act in a good approximation additively on the pure-tone hearing threshold.     

Speech recognition as a function of high-pass filter cutoff frequency for people with and without low-frequency cochlear dead regions

Regions in the cochlea with no (or very few) functioning inner hair cells and/or neurons are called "dead regions" (DRs). The recognition of high-pass filtered nonsense syllables was measured as a function of filter cutoff frequency for hearing-impaired people with and without low-frequency (apical) cochlear DRs. The diagnosis of any DR was made using the TEN(HL) test, and psychophysical tuning curves were used to define the edge frequency (f(e)) more precisely. Stimuli were amplified differently for each ear, using the "Cambridge formula." For subjects with low-frequency hearing loss but without DRs, scores were high (about 78%) for low cutoff frequencies, remained approximately constant for cutoff frequencies up to 862 Hz, and then worsened with increasing cutoff frequency. For subjects with low-frequency DRs, performance was typically poor for the lowest cutoff frequency (100 Hz), improved as the cutoff frequency was increased to about 0.57f(e), and worsened with further increases. These results indicate that people with low-frequency DRs are able to make effective use of frequency components that fall in the range 0.57f(e) to f(e), but that frequency components below 0.57f(e) have deleterious effects. The results have implications for the fitting of hearing aids to people with low-frequency DRs.