Psychophysical evidence for auditory compression at low characteristic frequencies

Psychophysical estimates of compression often assume that the basilar-membrane response to frequencies well below characteristic frequency (CF) is linear. Two techniques for estimating compression are described here that do not depend on this assumption at low CFs. In experiment 1, growth of forward masking was measured for both on- and off-frequency pure-tone maskers for pure-tone signals at 250, 500, and 4000 Hz. The on- and off-frequency masking functions at 250 and 500 Hz were just as shallow as the on-frequency masking function at 4000 Hz. In experiment 2, the forward masker level required to mask a fixed low-level signal was measured as a function of the masker-signal interval. The slopes of these functions did not differ between signal frequencies of 250 and 4000 Hz for the on-frequency maskers. At 250 Hz, the slope for the 150-Hz masker was almost as steep as that for the on-frequency masker, whereas at 4000 Hz the slope for the 2400-Hz masker was much shallower than that for the on-frequency masker. The results suggest that there is substantial compression, of around 0.2-0.3 dB/dB, at low CFs in the human auditory system. Furthermore, the results suggest that at low CFs compression does not vary greatly with stimulation frequency relative to CF.     

Carrier-dependent temporal processing in an auditory interneuron

Signal processing in the auditory interneuron Omega Neuron 1 (ON1) of the cricket Teleogryllus oceanicus was compared at high- and low-carrier frequencies in three different experimental paradigms. First, integration time, which corresponds to the time it takes for a neuron to reach threshold when stimulated at the minimum effective intensity, was found to be significantly shorter at high-carrier frequency than at low-carrier frequency. Second, phase locking to sinusoidally amplitude modulated signals was more efficient at high frequency, especially at high modulation rates and low modulation depths. Finally, we examined the efficiency with which ON1 detects gaps in a constant tone. As reflected by the decrease in firing rate in the vicinity of the gap, ON1 is better at detecting gaps at low-carrier frequency. Following a gap, firing rate increases beyond the pre-gap level. This "rebound" phenomenon is similar for low- and high-carrier frequencies.     

Measurement of auditory temporal processing using modified masking period patterns

A common metric of auditory temporal processing is the difference in the threshold for a pure-tone signal masked by either unmodulated or amplitude-modulated noise. This technique may be viewed as a modification of the masking period pattern technique. Such measurements have been proposed as an efficient means of estimating auditory temporal resolution in a clinical setting, although in many cases threshold differences may reflect additional spectro-temporal processes. The primary purpose of the present experiment was to examine interactions among signal frequency and masker bandwidth and the effects of modulation frequency on modified masking period patterns. The results revealed unmodulated-modulated threshold differences that increased with increasing masker bandwidth and decreased with increasing modulation frequency. There was little effect of signal frequency for narrow-band noise maskers that were equal in absolute bandwidth across frequency. However, unmodulated-modulated threshold differences increased substantially with increasing signal frequency for bandwidths proportional to the signal frequency and for wideband maskers. Although the results are interpreted in terms of a combination of both within-channel and across-channel cues, the specific contributions of these cues in particular conditions are difficult to ascertain. Because modified masking period patterns depend strongly upon a number of specific stimulus parameters, and because it is difficult to determine with any precision the underlying perceptual processes, this technique is not recommended for use as a clinical measure of auditory temporal processing.     

Effects of acoustic overstimulation on spectral and temporal processing in the amphibian auditory nerve

Activity of isolated auditory-nerve fibers in tree frogs (Eleutherodactylus coqui) exposed to continuous 3-min tones of different intensities at their characteristic frequencies (CFs) was recorded. Period histograms show a retardation in the preferred phase of discharge during and after the cessation of the exposure. Postexposure phase shift is concomitant with an elevation in CF thresholds and related to the level of tone exposure above threshold. Vector strength does not show comparable trends of change; postexposure shifts are related to preexposure CF thresholds. Recovery of phase retardation is rapid; units exposed to successive 3-min tones of the same intensities with intervals of 10-14 min between exposures experienced similar changes in their patterns of temporal discharge. Micromechanical changes affecting stereocilia stiffness or structural alterations in the tectorial membrane of the amphibian papilla may underly the transitory phase shifts observed in traumatized anuran auditory fibers.     

Investigating temporal asymmetry using masking period patterns and models of peripheral auditory processing

Two experiments were conducted in conjunction with modeling to evaluate the role of peripheral nonlinearity and neural adaptation in the perception of temporally asymmetric sounds. In both experiments, maskers were broadband noises amplitude modulated with ramped and damped exponential modulators that repeated at 40 Hz. Masking period patterns (MPPs) were constructed by measuring detection threshold of a 5-ms, 1000-Hz tone burst as function of the signal's onset delay. Experiment I showed that varying modulator half-life from 1 to 16 ms led to differences in the damped and the ramped MPPs that were largest at the short half-lives and diminished at the longer half-lives. When masker level was varied (experiment II), the largest difference between ramped and damped MPPs occurred at moderate stimulus levels. Two peripheral auditory models were evaluated, one a simple auditory filter followed by a power-law nonlinearity and another, a model of auditory nerve processing [J. Acoust. Soc. Am. 126, 2390-2412 (2009)] that includes neural adaptation. Neither models predicted differences between the ramped and damped MPPs, providing indirect support that the central auditory system has a role in perceptual temporal asymmetry.     

Temporal processing in the aging auditory system.

Measures of monaural temporal processing and binaural sensitivity were obtained from 12 young (mean age = 26.1 years) and 12 elderly (mean age = 70.9 years) adults with clinically normal hearing (pure-tone thresholds < or = 20 dB HL from 250 to 6000 Hz). Monaural temporal processing was measured by gap detection thresholds. Binaural sensitivity was measured by interaural time difference (ITD) thresholds. Gap and ITD thresholds were obtained at three sound levels (4, 8, or 16 dB above individual threshold). Subjects were also tested on two measures of speech perception, a masking level difference (MLD) task, and a syllable identification/discrimination task that included phonemes varying in voice onset time (VOT). Elderly listeners displayed poorer monaural temporal analysis (higher gap detection thresholds) and poorer binaural processing (higher ITD thresholds) at all sound levels. There were significant interactions between age and sound level, indicating that the age difference was larger at lower stimulus levels. Gap detection performance was found to correlate significantly with performance on the ITD task for young, but not elderly adult listeners. Elderly listeners also performed more poorly than younger listeners on both speech measures; however, there was no significant correlation between psychoacoustic and speech measures of temporal processing. Findings suggest that age-related factors other than peripheral hearing loss contribute to temporal processing deficits of elderly listeners.     

Aging and temporal discrimination in auditory sequences.

This study examined age-related changes in temporal sensitivity to increments in the inter-onset intervals (IOI) of successive components in tonal sequences. Temporal discrimination was examined using reference stimulus patterns consisting of five 50-ms, 4000-Hz components with equal tonal IOIs selected from the range 100-600 ms. Discrimination was examined in separate conditions by measuring the relative difference limen (DL) for increments of tonal IOI in comparison sequences. In some conditions, comparison sequences featured equal increments of all tonal lOIs to examined listener sensitivity to uniform changes of sequence rate, or tempo. Other conditions measured the DL for increments of a single target IOI within otherwise uniform-rate comparison sequences. For these measurements, the single target IOI was either fixed in sequence location, or randomized in location across listening trials. Listeners in the study included four groups of young and elderly adults with and without high-frequency hearing loss. The results for all listeners showed the relative DL for rate discrimination to decrease from a maximum at the 100-ms IOI to a smaller stable value across the range of longer sequence IOI. All listeners also exhibited larger relative DLs for discrimination of single target intervals compared to rate discrimination for equivalent reference IOI values. Older listeners showed poorer performance than younger listeners in all conditions, with the largest age differences observed for discrimination of brief single intervals that were varied randomly in sequence location. None of the results revealed significant effects of hearing loss on performance of younger and older listeners.     

Asymmetry of masking between noise and iterated rippled noise: evidence for time-interval processing in the auditory system.

This study describes the masking asymmetry between noise and iterated rippled noise (IRN) as a function of spectral region and the IRN delay. Masking asymmetry refers to the fact that noise masks IRN much more effectively than IRN masks noise, even when the stimuli occupy the same spectral region. Detection thresholds for IRN masked by noise and for noise masked by IRN were measured with an adaptive two-alternative, forced choice (2AFC) procedure with signal level as the adaptive parameter. Masker level was randomly varied within a 10-dB range in order to reduce the salience of loudness as a cue for detection. The stimuli were filtered into frequency bands, 2.2-kHz wide, with lower cutoff frequencies ranging from 0.8 to 6.4 kHz. IRN was generated with 16 iterations and with varying delays. The reciprocal of the delay was 16, 32, 64, or 128 Hz. When the reciprocal of the IRN delay was within the pitch range, i.e., above 30 Hz, there was a substantial masking asymmetry between IRN and noise for all filter cutoff frequencies; threshold for IRN masked by noise was about 10 dB larger than threshold for noise masked by IRN. For the 16-Hz IRN, the masking asymmetry decreased progressively with increasing filter cutoff frequency, from about 9 dB for the lowest cutoff frequency to less than 1 dB for the highest cutoff frequency. This suggests that masking asymmetry may be determined by different cues for delays within and below the pitch range. The fact that masking asymmetry exists for conditions that combine very long IRN delays with very high filter cutoff frequencies means that it is unlikely that models based on the excitation patterns of the stimuli would be successful in explaining the threshold data. A range of time-domain models of auditory processing that focus on the time intervals in phase-locked neural activity patterns is reviewed. Most of these models were successful in accounting for the basic masking asymmetry between IRN and noise for conditions within the pitch range, and one of the models produced an exceptionally good fit to the data.     

Psychophysical tuning curves and auditory thresholds after hair cell damage in the chinchilla

Chinchillas were treated with kanamycin sulfate (150--200 mg/kg/day) to produce high-frequency hearing loss extending to about 4.0 kHz. Thresholds and psychophysical tuning curves (PTCs) were obtained before and after treatment, utilizing a shuttlebox avoidance procedure, and cochlear hair cells were evaluated under phase contrast microscopy. Hair cell loss resulting from kanamycin treatment varied from restricted lesions of the outer hair cells (OHCs) in the cochlear base, with no loss of inner hair cells (IHCs), to more extensive lesions involving both OHCs and IHCs. Threshold shift of at least 40 dB was always associated with OHC loss. PTCs obtained from frequency regions exhibiting 40--50 dB of threshold shift were normal in shape. With threshold shift in excess of 50 dB, PTCs were progressively distorted, with truncation of the tip segment and in some cases increased sensitivity of the tail segment. The results suggest that the threshold of optimally functional IHCs after kanamycin-induced OHC loss is about 40 dB higher than normal. Threshold shift in excess of 40 dB may represent IHC damage. IHCs are capable of transducing the fine-frequency information necessary for generating normally sharp PTCs in the absence of OHCs. However, with threshold shift in excess of approximately 50 dB, this frequency resolution is increasingly compromised.     

Psychophysical assessment of the level-dependent representation of high-frequency spectral notches in the peripheral auditory system

To discriminate between broadband noises with and without a high-frequency spectral notch is more difficult at 70-80 dB sound pressure level than at lower or higher levels [Alves-Pinto, A. and Lopez-Poveda, E. A. (2005). "Detection of high-frequency spectral notches as a function of level," J. Acoust. Soc. Am. 118, 2458-2469]. One possible explanation is that the notch is less clearly represented internally at 70-80 dB SPL than at any other level. To test this hypothesis, forward-masking patterns were measured for flat-spectrum and notched noise maskers for masker levels of 50, 70, 80, and 90 dB SPL. Masking patterns were measured in two conditions: (1) fixing the masker-probe time interval at 2 ms and (2) varying the interval to achieve similar masked thresholds for different masker levels. The depth of the spectral notch remained approximately constant in the fixed-interval masking patterns and gradually decreased with increasing masker level in the variable-interval masking patterns. This difference probably reflects the effects of peripheral compression. These results are inconsistent with the nonmonotonic level-dependent performance in spectral discrimination. Assuming that a forward-masking pattern is a reasonable psychoacoustical correlate of the auditory-nerve rate-profile representation of the stimulus spectrum, these results undermine the common view that high-frequency spectral notches must be encoded in the rate-profile of auditory-nerve fibers.     

Speech processing in the auditory system. II: Lateral inhibition and the central processing of speech evoked activity in the auditory nerve

A biologically realistic model of a uniform lateral inhibitory network (LIN) is shown capable of extracting from the complex spatio-temporal firing patterns of the cat's auditory nerve the formants and low-order harmonics of synthetic voiced speech stimuli. The model provides a realistic mechanism to utilize the temporal aspects of the firing and thus supports the hypothesis that the neural coding of complex sounds in terms of average rates can be supplemented by the information coded in the synchronous firing. At low levels of intensity the LIN can sharpen the average rate profiles. At moderate and high levels the LIN uses the cues available in the distribution of phases of the synchronous activity which exhibit rapid relative phase shifts at specific characteristic frequency (CF) locations (corresponding to the frequencies of the low-order harmonics in the stimulus). These temporal phase shifts manifest themselves at the input of the LIN as steep and localized spatial discontinuities in the instantaneous pattern of activity across the fiber array. The LIN enhances its output from these spatially steep input regions while suppressing its output from spatially smooth input regions (where little phase shifts occur). In this manner the LIN recreates from the response patterns a representation of the stimulus spectrum using the temporal cues as spatial markers of the stimulus components rather than as absolute measures of their frequencies. Similar results are obtained with various lateral inhibitory topologies, e.g., recurrent versus nonrecurrent, single versus double layer, and linear versus nonlinear.     

Psychophysical estimates of nonlinear cochlear processing in younger and older listeners

The primary goal of this project was to compare the performance of younger and older listeners on a number of psychophysical measures thought to be influenced by nonlinear cochlear processing. Younger (mean of 25.6 years) and older (mean of 63.8 years) listeners with normal hearing were matched (within 5 dB) according to their quiet thresholds at the two test frequencies of 1200 and 2400 Hz. They were similarly matched at the adjacent octave frequencies of 600 and 4800 Hz (within 5 dB at one and 9 dB at the other). Performance was compared on measures of auditory filter shape, psychophysical suppression, and growth of forward masking. There was no difference between the two age groups on these psychophysical estimates reflecting nonlinear processing, suggesting that aging per se does not affect the cochlear nonlinearity, at least for the ages sampled here. The results did, however, consistently demonstrate an age-related increase in the susceptibility to forward masking.     

反映时序特征的听神经自发发放模型

听神经发放时序在听信息编码中起重要作用,要认识和运用听觉系统对听神经发放时序特征的利用机制必须对听神经发放时序特征有认识,但是已有的随机过程数学模型均不能很好地反映听神经发放时序特征。而关于发放的生理模型,如Hodgkin-Huxley方程、Meddis模型对发放过程的描述均是决定性的、连续性的,不能反映生理过程的离散性和发放过程的随机性。本文基于神经发放生理过程,通过对神经发放条件以及自发发放时神经递质释放时序的描述,建立了一个简单自发发放模型。模型在很好地反映听神经自发发放时序外部统计特征的同时,所得到的神经递质的释放速率在生理上也是合理的。模型还能对纯音激励时听神经的锁相发放做出定性解释,可作为进一步研究反映时序特征的有激励发放模型的基础。     

Speech processing in the auditory system. I: The representation of speech sounds in the responses of the auditory nerve

In a previous paper the speech evoked spatio-temporal response patterns recorded in large populations of auditory-nerve fibers in the cat were examined [M.I. Miller and M.B. Sachs, J. Acoust. Soc. Am. 74, 502-517 (1983)]. The distribution of the relative phases of synchronized activity emerges as an important response feature reflecting the stimulus spectral parameters. Specifically, each strong low-order harmonic of the stimulus (less than or equal to 1.5-2 kHz) dominates the synchrony of a relatively broad segment of fibers near its corresponding characteristic frequency (CF) location in a pattern which mirrors the underlying traveling wave component. Each such fiber segment can be roughly subdivided into two regions: (1) a region basal to the point of resonance of the harmonic where the fiber PST histograms accumulate only small delays (or phase shifts) relative to each other reflecting the fast speed of propagation of the traveling wave, and (2) a region at or very near the point of resonance where the responses exhibit drastic relative phase shifts owing to the sudden slow down of the traveling wave and the consequent rapid accumulation of phase shifts. These rapid phase shifts thus manifest themselves as steep and localized spatial discontinuities in an otherwise relatively uniform instantaneous pattern of activity across the fiber array, all occurring at the CF locations corresponding to the low-order harmonics of the stimulus.