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PubMed2003

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.

Animals↗
PubMed2003

Enhancing interaural-delay-based extents of laterality at high frequencies by using "transposed stimuli".

An acoustic pointing task was used to determine whether interaural temporal disparities (ITDs) conveyed by high-frequency "transposed" stimuli would produce larger extents of laterality than ITDs conveyed by bands of high-frequency Gaussian noise. The envelopes of transposed stimuli are designed to provide high-frequency channels with information similar to that conveyed by the waveforms of low-frequency stimuli. Lateralization was measured for low-frequency Gaussian noises, the same noises transposed to 4 kHz, and high-frequency Gaussian bands of noise centered at 4 kHz. Extents of laterality obtained with the transposed stimuli were greater than those obtained with bands of Gaussian noise centered at 4 kHz and, in some cases, were equivalent to those obtained with low-frequency stimuli. In a second experiment, the general effects on lateral position produced by imposed combinations of bandwidth, ITD, and interaural phase disparities (IPDs) on low-frequency stimuli remained when those stimuli were transposed to 4 kHz. Overall, the data were fairly well accounted for by a model that computes the cross-correlation subsequent to known stages of peripheral auditory processing augmented by low-pass filtering of the envelopes within the high-frequency channels of each ear.

Adult↗
PubMed2003

Perceptual weights in auditory level discrimination.

Perceptual weights in level discrimination (also called intensity discrimination) were determined for 3-, 7-, 15-, and 24-component tone complexes with flat spectral envelopes using a correlational paradigm. Each frequency component was randomly and independently perturbed in level oneach presentation. For the target interval, frequency-component levels were additionally increased by the level increment to be detected, deltaL [= 201og10((p + deltap)/p), where p is pressure]. Weights were calculated from the across-trial correlation between the level perturbations for each frequency component and the interval chosen by the listener. Two conditions were investigated: (1) deltaL was equal across frequency components, and (2) deltaL increased progressively across frequency components. For both conditions, data for four listeners usually showed the greatest weight for the highest frequency component. The two-to-four highest frequency components generally were most important for level discrimination. The effect of increasing deltaL progressively with frequency was small and inconsistent. Additional measurements showed that flanking noise maskers designed to mask spread of excitation caused only small and generally unsystematic changes to the weights. Overall, these results indicate that listeners combine information across a wide range of auditory channels to arrive at a decision for level discrimination, but the weighting of channels appears to be suboptimal.

Adult↗
PubMed2003

A comparison of psychophysical procedures for level-discrimination thresholds.

Five different psychophysical procedures were used to measure level-discrimination (also called intensity discrimination) thresholds for 1-kHz tones at two levels (30 and 90 dB SPL) and two durations (10 and 500 ms). The procedures were the classic transformed up-down staircase method with a two-alternative forced-choice (2AFC) paradigm (UPD), 15- and 50-trial implementations of the method of maximum likelihood (MML) with a cued yes-no paradigm, and 18-trial implementations of ZEST using both cued yes-no and 2AFC paradigms. Results obtained from nine normal listeners show that estimates of level-discrimination thresholds for the four conditions are similar across all five procedures when different points of convergence are accounted for. The variance of threshold estimates within listener and condition was smallest for UPD, largest for the MML with 15 trials, and statistically indistinguishable among the others. The sweat factors ranged from 5.5 for MML with 50 trials to about 1.4 for UPD and ZEST. Simulations show that ideal performance of procedures may be far from real-life experience and that these deviations are likely to depend on complex interactions between listener behavior and parameter choices used for implementing the procedures. Therefore, empirical verification is important for judging the effectiveness of psychophysical procedures.

Adolescent↗
PubMed2003

Distortion product otoacoustic emission suppression tuning curves in normal-hearing and hearing-impaired human ears.

Distortion product otoacoustic emission (DPOAE) suppression measurements were made in 20 subjects with normal hearing and 21 subjects with mild-to-moderate hearing loss. The probe consisted of two primary tones (f2, f1), with f2 held constant at 4 kHz and f2/f1 = 1.22. Primary levels (L1, L2) were set according to the equation L1 = 0.4 L2 + 39 dB [Kummer et al., J. Acoust. Soc. Am. 103, 3431-3444 (1998)], with L2 ranging from 20 to 70 dB SPL (normal-hearing subjects) and 50-70 dB SPL (subjects with hearing loss). Responses elicited by the probe were suppressed by a third tone (f3), varying in frequency from 1 octave below to 1/2 octave above f2. Suppressor level (L3) varied from 5 to 85 dB SPL. Responses in the presence of the suppressor were subtracted from the unsuppressed condition in order to convert the data into decrements (amount of suppression). The slopes of the decrement versus L3 functions were less steep for lower frequency suppressors and more steep for higher frequency suppressors in impaired ears. Suppression tuning curves, constructed by selecting the L3 that resulted in 3 dB of suppression as a function of f3, resulted in tuning curves that were similar in appearance for normal and impaired ears. Although variable, Q10 and Q(ERB) were slightly larger in impaired ears regardless of whether the comparisons were made at equivalent SPL or equivalent sensation levels (SL). Larger tip-to-tail differences were observed in ears with normal hearing when compared at either the same SPL or the same SL, with a much larger effect at similar SL. These results are consistent with the view that subjects with normal hearing and mild-to-moderate hearing loss have similar tuning around a frequency for which the hearing loss exists, but reduced cochlear-amplifier gain.

Adolescent↗
PubMed2003

Perceived naturalness of spectrally distorted speech and music.

We determined how the perceived naturalness of music and speech (male and female talkers) signals was affected by various forms of linear filtering, some of which were intended to mimic the spectral "distortions" introduced by transducers such as microphones, loudspeakers, and earphones. The filters introduced spectral tilts and ripples of various types, variations in upper and lower cutoff frequency, and combinations of these. All of the differently filtered signals (168 conditions) were intermixed in random order within one block of trials. Levels were adjusted to give approximately equal loudness in all conditions. Listeners were required to judge the perceptual quality (naturalness) of the filtered signals on a scale from 1 to 10. For spectral ripples, perceived quality decreased with increasing ripple density up to 0.2 ripple/ERB(N) and with increasing ripple depth. Spectral tilts also degraded quality, and the effects were similar for positive and negative tilts. Ripples and/or tilts degraded quality more when they extended over a wide frequency range (87-6981 Hz) than when they extended over subranges. Low- and mid-frequency ranges were roughly equally important for music, but the mid-range was most important for speech. For music, the highest quality was obtained for the broadband signal (55-16,854 Hz). Increasing the lower cutoff frequency from 55 Hz resulted in a clear degradation of quality. There was also a distinct degradation as the upper cutoff frequency was decreased from 16,845 Hz. For speech, there was a marked degradation when the lower cutoff frequency was increased from 123 to 208 Hz and when the upper cutoff frequency was decreased from 10,869 Hz. Typical telephone bandwidth (313 to 3547 Hz) gave very poor quality.

Adolescent↗
PubMed2003

A diminished rate of "physiological decay" at noise offset contributes to age-related changes in temporal acuity in the CBA mouse model of presbycusis.

The persistence of afferent activity at stimulus offset was examined in mice as a function of age (experiment 1) and noise level (experiment 2), using a procedure devised by von Bekesy to study the physiological decay of afferent activity. Noise offset was presented from 1 to 10 ms (the temporal gap) prior to an acoustic startle stimulus, with an abrupt edge or a linear ramp having the duration of the gap. Noise offset inhibited the startle reflex, this effect declining with age. For young (2-3 months old) and "young-old" mice (18-19 months old), the abrupt offset was always more inhibitory than the ramp, even for the one millisecond gap. In "old-old" mice (24-26 months old) abrupt and ramped offsets were not different until the gap exceeded 4 ms. The behavioral difference between abrupt and ramped decay times did not depend on noise level in young mice (4-5 months old), though overall inhibition increased with level. These data indicate that temporal acuity as measured by this gap detection method is limited in very old mice by their reduced ability to follow the abrupt change in noise level at the beginning of the gap.

Acoustic Stimulation↗
PubMed2003

The effect of speech presentation level on measurement of auditory acclimatization to amplified speech.

A systematic improvement in auditory performance over time, following a change in the acoustic information available to the listener (that cannot be attributed to task, procedural or training effects) is known as auditory acclimatization. However, there is conflicting evidence concerning the existence of auditory acclimatization; some studies show an improvement in performance over time while other studies show no change. In an attempt to resolve this conflict, speech recognition abilities of 16 subjects with bilateral sensorineural hearing impairments were measured over a 12-week period following provision of a monaural hearing instrument for the first time. The not-fitted ear was used as the control. Three presentation levels were used representing quiet, normal, and raised speech. The results confirm the presence of acclimatization. In addition, the results show that acclimatization is evident at the higher presentation levels but not at the lowest.

Aged↗
PubMed2003

Precedence-effect thresholds for a population of untrained listeners as a function of stimulus intensity and interclick interval.

Data are reported from 127 untrained individuals under lag- and single-click conditions in a precedence-effect task. In experiment I, each subject completed ten runs in a two-interval forced-choice design under a lag-click condition and three runs under a single-click condition. The cue to be discriminated was an interaural time difference (ITD). Stimuli were 125-micros rectangular pulses and the interclick interval (ICI) was 2 ms. Subjects were randomly assigned to three groups of approximately 30. Each group was tested at one stimulus intensity (43, 58, or 73 dB). Mean threshold within each group was greater than 500 micros for lag-click ITD conditions, although substantial intersubject variability and a clear effect of stimulus intensity on lag-click ITD thresholds were observed, with lower thresholds for higher intensities. In experiment II, the ICI was varied from 0.3 to 10 ms, and thresholds were obtained from groups of approximately 20 untrained subjects. Data were also collected from three highly experienced observers as a function of ICI. The best naive subject produced mean thresholds near, but not as low as those obtained from experienced subjects. Analysis of adaptive-track patterns revealed abrupt irregularities in threshold tracking, consistent with either losing the cue or listening to the wrong cue in an ambiguous stimulus.

Acoustic Stimulation↗
PubMed2003

The effects of a high-frequency suppressor on tuning curves and derived basilar-membrane response functions.

Forward-masked psychophysical tuning curves were obtained using a fixed, low-level signal at a frequency of 4 kHz, and masker frequencies of 2.0, 2.5, 3.0, 3.5, 3.75, 4.0, 4.25, 4.5, 4.75, 5.0, and 5.5 kHz, at masker-signal gaps of 20, 30, 40, 60, 80, and 100 ms. An adaptive two-interval, two alternative forced-choice (21-2AFC) procedure was used to obtain the masker level at threshold. This procedure was repeated with the addition of a 4.75-kHz suppressor at 50 or 60 dB SPL, gated with the masker. Tuning curves were broader, and estimates of compression and gain from derived input/output functions were decreased in the presence of a suppressor as compared to the no-suppressor condition. The results are consistent with physiological results, which show that suppression leads to a broadening of tuning curves and a partial linearization of the midlevel portion of the basilar-membrane input/output function.

Acoustic Stimulation↗
PubMed2003

Intensity discrimination and increment detection in cochlear-implant users.

Intensity difference limens (DLs) were measured in users of the Nucleus 22 and Clarion v1.2 cochlear implants and in normal-hearing listeners to better understand mechanisms of intensity discrimination in electric and acoustic hearing and to evaluate the possible role of neural adaptation. Intensity DLs were measured for three modes of presentation: gated (intensity increments gated synchronously with the pedestal), fringe (intensity increments delayed 250 or 650 ms relative to the onset of the pedestal), and continuous (intensity increments occur in the presence of a pedestal that is played throughout the experimental run). Stimuli for cochlear-implant listeners were trains of biphasic pulses; stimuli for normal-hearing listeners were a 1-kHz tone and a wideband noise. Clarion cochlear-implant listeners showed level-dependent effects of presentation mode. At low pedestal levels, gated thresholds were generally similar to thresholds obtained in the fringe and continuous conditions. At higher pedestal levels, however, the fringe and continuous conditions produced smaller intensity DLs than the gated condition, similar to the gated-continuous difference in intensity DLs observed in acoustic hearing. Nucleus cochlear-implant listeners did not show consistent threshold differences for the gated and fringe conditions, and were not tested in the continuous condition. It is not clear why a difference between gated and fringe thresholds occurred for the Clarion but not the Nucleus subjects. Normal-hearing listeners showed improved thresholds for the continuous condition relative to the gated condition, but the effect was larger for the 1-kHz tonal carrier than for the noise carrier. Findings suggest that adaptation occurring central to the inner hair cell synapse mediates the gated-continuous difference observed in Clarion cochlear-implant listeners and may also contribute to the gated-continuous difference in acoustic hearing.

Acoustic Stimulation↗
PubMed2003

Basilar-membrane responses to multicomponent (Schroeder-phase) signals: understanding intensity effects.

Harmonic complexes comprised of the same spectral components in either positive-Schroeder (+Schr) or negative-Schroeder (-Schr) phase [see Schroeder, IEEE Trans. Inf. Theory 16, 85-89 (1970)] have identical long-term spectra and similar waveform envelopes. However, localized patterns of basilar-membrane (BM) excitation can be quite different in response to these two stimuli. Measurements in chinchillas showed more modulated (peakier) BM excitation for +Schr than -Schr complexes [Recio and Rhode, J. Acoust. Soc. Am. 108, 2281-2298 (2000)]. In the current study, laser velocimetry was used to examine BM responses at a location tuned to approximately 17 kHz in the basal turn of the guinea-pig cochlea, for +Schr and -Schr complexes with a 203-Hz fundamental frequency and including 101 equal-amplitude components from 2031 to 22,344 Hz. At 35-dB SPL, +Schr response waveforms showed greater amplitude modulation than -Schr responses. With increasing stimulation level, internal modulation decreased for both complexes. To understand the observed phenomena quantitatively, responses were predicted on the basis of a linearized model of the cochlea. Prediction was based on an "indirect impulse response" measured in the same animal. Response waveforms for Schroeder-phase signals were accurately predicted, provided that the level of the indirect impulse used in prediction closely matched the level of the Schroeder-phase stimulus. This result confirms that the underlying model, which originally was developed for noise stimuli, is valid for stimuli that produce completely different response waveforms. Moreover, it justifies explanation of cochlear filtering (i.e., differential treatment of different frequencies) in terms of a linear system.

Animals↗
PubMed2003

Vertical-plane sound localization probed with ripple-spectrum noise.

Ripple-spectrum stimuli were used to investigate the scale of spectral detail used by listeners in interpreting spectral cues for vertical-plane localization. In three experiments, free-field localization judgments were obtained for 250-ms, 0.6-16-kHz noise bursts with log-ripple spectra that varied in ripple density, peak-to-trough depth, and phase. When ripple density was varied and depth was held constant at 40 dB, listeners' localization error rates increased most (relative to rates for flat-spectrum targets) for densities of 0.5-2 ripples/oct. When depth was varied and density was held constant at 1 ripple/oct, localization accuracy was degraded only for ripple depths > or = 20 dB. When phase was varied and density was held constant at 1 ripple/oct and depth at 40 dB, three of five listeners made errors at consistent locations unrelated to the ripple phase, whereas two listeners made errors at locations systematically modulated by ripple phase. Although the reported upper limit for ripple discrimination is 10 ripples/oct [Supin et al., J. Acoust. Soc. Am. 106, 2800-2804 (1999)], present results indicate that details finer than 2 ripples/oct or coarser than 0.5 ripples/oct do not strongly influence processing of spectral cues for sound localization. The low spectral-frequency limit suggests that broad-scale spectral variation is discounted, even though components at this scale are among those contributing the most to the shapes of directional transfer functions.

Acoustic Stimulation↗
PubMed2003

Effectiveness of narrow-band versus tonal off-frequency maskers.

The present study was a follow-up to a pilot study in which it was found that a 500-Hz-wide narrow-band noise (NBN) masker produced more masking than a tonal (T) masker for signal frequencies both above and below the masker frequency. The aim of the present study was to determine to what extent these results were influenced by an interaction of the relatively rapid temporal envelope fluctuations of the NBN and the short (10-ms) duration of the signal. In the first experiment, the masking produced by a regular NBN, a low-noise noise (LNN), and a T was compared. The LNN produced less masking than the NBN, and about as much as the T, suggesting that the inherent amplitude fluctuations in the NBN were largely responsible for the greater masking produced by that masker. In the second experiment, the masking produced by a regular NBN was compared with that by a T for a signal duration of 10 or 200 ms. The difference in masking between the two maskers was reduced or eliminated when the signal duration was 200 ms, because the threshold in the presence of the NBN masker decreased more with increasing signal duration. This could reflect a decreased "confusion" between the signal and the inherent fluctuations of the NBN masker.

Acoustic Stimulation↗
PubMed2003

Asymmetry of masking between complex tones and noise: partial loudness.

This experiment examined the partial masking of periodic complex tones by a background of noise, and vice versa. The tones had a fundamental frequency (F0) of 62.5 or 250 Hz, and components were added in either cosine phase (CPH) or random phase (RPH). The tones and the noise were bandpass filtered into the same frequency region, from the tenth harmonic up to 5 kHz. The target alone was alternated with the target and the background; for the mixture, the background and target were either gated together, or the background was turned on 400 ms before, and off 200 ms after, the target. Subjects had to adjust the level of either the target alone or the target in the background so as to match the loudness of the target in the two intervals. The overall level of the background was 50 dB SPL, and loudness matches were obtained for several fixed levels of the target alone or in the background. The resulting loudness-matching functions showed clear asymmetry of partial masking. For a given target-to-background ratio, the partial loudness of a complex tone in a noise background was lower than the partial loudness of a noise in a complex tone background. Expressed as the target-to-background ratio required to achieve a given loudness, the asymmetry typically amounted to 12-16 dB. When the F0 of the complex tone was 62.5 Hz, the asymmetry of partial masking was greater for CPH than for RPH. When the F0 was 250 Hz, the asymmetry was greater for RPH than for CPH. Masked thresholds showed the same pattern as for partial masking for both F0's. Onset asynchrony had some effect on the loudness matching data when the target was just above its masked threshold, but did not significantly affect the level at which the target in the background reached its unmasked loudness. The results are interpreted in terms of the temporal structure of the stimuli.

Adult↗
PubMed2003

Louder sounds can produce less forward masking: effects of component phase in complex tones.

The influence of the degree of envelope modulation and periodicity on the loudness and effectiveness of sounds as forward maskers was investigated. In the first experiment, listeners matched the loudness of complex tones and noise. The tones had a fundamental frequency (F0) of 62.5 or 250 Hz and were filtered into a frequency range from the 10th harmonic to 5000 Hz. The Gaussian noise was filtered in the same way. The components of the complex tones were added either in cosine phase (CPH), giving a large crest factor, or in random phase (RPH), giving a smaller crest factor. For each F0, subjects matched the loudness between all possible stimulus pairs. Six different levels of the fixed stimulus were used, ranging from about 30 dB SPL to about 80 dB SPL in 10-dB steps. Results showed that, at a given overall level, the CPH and the RPH tones were louder than the noise, and that the CPH tone was louder than the RPH tone. The difference in loudness was larger at medium than at low levels and was only slightly reduced by the addition of a noise intended to mask combination tones. The differences in loudness were slightly smaller for the higher than for the lower F0. In the second experiment, the stimuli with the lower F0s were used as forward maskers of a 20-ms sinusoid, presented at various frequencies within the spectral range of the maskers. Results showed that the CPH tone was the least effective forward masker, even though it was the loudest. The differences in effectiveness as forward maskers depended on masker level and signal frequency; in order to produce equal masking, the level of the CPH tone had to be up to 35 dB above that of the RPH tone and the noise. The implications of these results for models of loudness are discussed and a model is presented based on neural activity patterns in the auditory nerve; this predicts the general pattern of loudness matches. It is suggested that the effects observed in the experiments may have been influenced by two factors: cochlear compression and suppression.

Adult↗
PubMed2003

Evaluation of an auditory model for echo delay accuracy in wideband biosonar.

In a psychophysical task with echoes that jitter in delay, big brown bats can detect changes as small as 10-20 ns at an echo signal-to-noise ratio of approximately 49 dB and 40 ns at approximately 36 dB. This performance is possible to achieve with ideal coherent processing of the wideband echoes, but it is widely assumed that the bat's peripheral auditory system is incapable of encoding signal waveforms to represent delay with the requisite precision or phase at ultrasonic frequencies. This assumption was examined by modeling inner-ear transduction with a bank of parallel bandpass filters followed by low-pass smoothing. Several versions of the filterbank model were tested to learn how the smoothing filters, which are the most critical parameter for controlling the coherence of the representation, affect replication of the bat's performance. When tested at a signal-to-noise ratio of 36 dB, the model achieved a delay acuity of 83 ns using a second-order smoothing filter with a cutoff frequency of 8 kHz. The same model achieved a delay acuity of 17 ns when tested with a signal-to-noise ratio of 50 dB. Jitter detection thresholds were an order of magnitude worse than the bat for fifth-order smoothing or for lower cutoff frequencies. Most surprising is that effectively coherent reception is possible with filter cutoff frequencies well below any of the ultrasonic frequencies contained in the bat's sonar sounds. The results suggest that only a modest rise in the frequency response of smoothing in the bat's inner ear can confer full phase sensitivity on subsequent processing and account for the bat's fine acuity or delay.

Animals↗