Auditory Frequency Selectivity Research Articles

Effects of ipsilateral, contralateral, and bilateral noise precursors on psychoacoustical tuning curves in humans

Cochlear tuning and hence auditory frequency selectivity are thought to change in noisy environments by activation of the medial olivocochlear reflex (MOCR). In humans, auditory frequency selectivity is often assessed using psychoacoustical tuning curves (PTCs), a plot of the level required for pure-tone maskers to just mask a fixed-level pure-tone probe as a function of masker frequency. Sometimes, however, the stimuli used to measure a PTC are long enough that they can activate the MOCR by themselves and thus affect the PTC. Here, PTCs for probe frequencies of 500 Hz and 4 kHz were measured in forward masking using short maskers (30 ms) and probes (10 ms) to minimize the activation of the MOCR by the maskers or the probes. PTCs were also measured in the presence of long (300 ms) ipsilateral, contralateral, and bilateral broadband noise precursors to investigate the effect of the ipsilateral, contralateral, and bilateral MOCR on PTC tuning. Four listeners with normal hearing participated in the experiments. At 500 Hz, ipsilateral and bilateral precursors sharpened the PTCs by decreasing the thresholds for maskers with frequencies at or near the probe frequency with minimal effects on thresholds for maskers remote in frequency from the probe. At 4 kHz, by contrast, ipsilateral and bilateral precursors barely affected thresholds for maskers near the probe frequency but broadened PTCs by reducing thresholds for maskers far from the probe. Contralateral precursors barely affected PTCs. An existing computational model was used to interpret the results. The model suggested that despite the apparent differences, the pattern of results is consistent with the ipsilateral and bilateral MOCR inhibiting the cochlear gain similarly at the two probe frequencies and more strongly than the contralateral MOCR.

Responses in area hMT+ reflect tuning for both auditory frequency and motion after blindness early in life

Previous studies report that human middle temporal complex (hMT+) is sensitive to auditory motion in early-blind individuals. Here, we show that hMT+ also develops selectivity for auditory frequency after early blindness, and that this selectivity is maintained after sight recovery in adulthood. Frequency selectivity was assessed using both moving band-pass and stationary pure-tone stimuli. As expected, within primary auditory cortex, both moving and stationary stimuli successfully elicited frequency-selective responses, organized in a tonotopic map, for all subjects. In early-blind and sight-recovery subjects, we saw evidence for frequency selectivity within hMT+ for the auditory stimulus that contained motion. We did not find frequency-tuned responses within hMT+ when using the stationary stimulus in either early-blind or sight-recovery subjects. We saw no evidence for auditory frequency selectivity in hMT+ in sighted subjects using either stimulus. Thus, after early blindness, hMT+ can exhibit selectivity for auditory frequency. Remarkably, this auditory frequency tuning persists in two adult sight-recovery subjects, showing that, in these subjects, auditory frequency-tuned responses can coexist with visually driven responses in hMT+.

A fast algorithm for the measurement of stimulus frequency otoacoustic emission suppression tuning curves

Frequency selectivity is an important indicator of auditory function in the human ear. Stimulus frequency otoacoustic emission (SFOAE) suppression tuning curves (STCs) have great potential in the objective analysis of human auditory frequency selectivity but are costly to measure by the traditional algorithm using pure-tone adaptive method. To improve the measurement efficiency of the SFOAE STCs, a faster algorithm based on a suppressor with gradiently changing intensity and interpolation is proposed in this paper. Twelve subjects participated in this study by measuring their SFOAE STCs through the traditional and fast algorithm. The results shows that the average correlation coefficients of the SFOAE STCs measured using the fast and the traditional algorithms at different probe frequencies is 0.94. And the measurement speed of the fast algorithm was approximately 2.10 times higher than the traditional one. Also, the fast algorithm is more efficient than the algorithms of Keefe et al. (2008) and Charaziak et al. (2013). Experimental evidence is provided that the proposed fast algorithm greatly improves the measurement speed and reduces the processing time, exhibiting good accuracy and reliability.

FLIM-FRET Measurements of Voltage Dependent Conformational Changes in the Outer Hair Cell Motor Protein Prestin

Cochlear sensory cells, outer hairs cells (OHCs), elongate and contract in response to changes in transmembrane potential. This electromotility is important for auditory frequency selectivity and sensitivity and is believed to be driven by a motor protein, prestin, which is highly expressed within the membrane of OHCs. Experiments have shown that prestin undergoes self-association leading to the observation that oligomerization might be important for proper function. We have previously utilized acceptor photobleach fluorescence resonance energy transfer (FRET) to study pretin-prestin interactions in living cells. However, traditional FRET, based upon detection of fluorescence intensity, is inherently subjected to artifacts that arise from variations in excitation intensity, photobleaching, and fluorophore concentration. We have recently implemented fluorescence lifetime imaging microscopy (FLIM) to perform robust FLIM-FRET measurements in HEK cells cotransfected with prestin-TFP and prestin-YFP. FRET efficiency was calculated by fitting the lifetime decay of prestin-TFP in cells clamped at various holding potentials. High FRET efficiencies were measured at hyperpolarized potentials, and low FRET efficiencies were measured at depolarized. The results indicated that changes in the transmembrane potential induce a conformational change in prestin that can be detected by FLIM-FRET. Thus, FLIM-FRET can provide a sensitive assay for the mechanical correlate of prestin function.

Time-efficient measures of auditory frequency selectivity

Objective: The objective of this study was to compare two recently proposed methods for fast measurements of psychophysical tuning curves (fast-PTCs) in terms of resulting tuning curve features and training effects. Design: Fast-PTCs with swept-noise (SN) and gated-noise (GN) maskers were measured at signal frequencies of 500, 1000, 2000, and 4000 Hz. The effect of amplitude modulating the signal in the GN condition was evaluated. Two PTC runs were obtained for each condition to assess training effects. Study sample: Eight normally-hearing young adults participated in the study. Results: The SN and GN methods resulted in similar estimates of frequency selectivity when training effects were considered. Amplitude modulating the tone in the GN method reduced the effect of training. On average, SN-PTCs were most repeatable compared to the two other methods and they were not affected by training. Estimation of the shift in the PTC tip frequency was not affected by the measurement method or training effects. Fast-PTC methods resulted in similar estimates of tuning as compared to published notched-noise data. Conclusions: The SN method and the GN procedure with amplitude modulated signals allowed for time-efficient estimation of frequency selectivity that was unaffected by training.

Comodulation masking release for regular and irregular modulators

The present study investigates whether the difference in comodulation masking release (CMR) for different modulator types is due to the different degrees of modulator regularity, as suggested in the literature, or results from different envelope distributions. Thresholds of a sinusoidal signal are measured in the presence of a noise masker which was either broadband or narrow band. A square-wave modulator with different degrees of regularity is used that preserves the envelope distribution. The measured CMR does not decrease as the regularity decreases. This finding argues against the hypothesis that the difference in CMR for different modulator types reported in the literature is due to differences in regularity. The data for the narrow-band conditions which either mimic the auditory frequency selectivity or preserve the modulation spectrum indicate that most of the CMR of the present study is due to within-channel cues. In agreement with this finding, within-channel models using either a peripheral nonlinearity or a modulation filterbank predict a CMR of a similar size. In contrast to the model predictions and the findings for the narrow-band conditions, the CMR for the broadband masker increases as the regularity decreases. This suggests that the CMR is not solely determined by the envelope distributions.

A filterbank model of human nonlinear auditory frequency selectivity

Filterbank models of human nonlinear auditory frequency selectivity are useful in a variety of applications. Existing filterbanks are almost certainly flawed because they do not account for the recent observation that compression extends to a wider range of stimulus frequencies in apical than in basal cochlear sites [Lopez-Poveda et al. (2003), J. Acoust. Soc. Am. 113, 951-960]. Here, we present a filterbank based on the forward-masking data from which this observation was made. A forward-masking model was constructed by cascading a linear “outer/middle-ear” filter, followed by a dual-resonance nonlinear (DRNL) filter, followed by a linear temporal window. The temporal window was assumed to be identical across listeners and cochlear places. Only the parameters of the DRNL filter were adjusted to simulate the absolute threshold and the masking data of three listeners for probe frequencies of 0.5, 1, 2, 4, and 8 kHz. A bank of DRNL filters was then constructed by linear or quadratic regression of the optimum parameters. It is discussed that this filterbank is the best possible approximation of human level-dependent auditory frequency selectivity to date. Additionally, the results support the linearity of the temporal window. [Work supported by IMSERSO 131/06, PROFIT CIT-390000-2005-4, MEC BFU-2006-07536.]

Auditory frequency selectivity is better for expected than for unexpected sound duration

Previous studies show that insectivorous bats prepare their auditory system to analyze expected returning echoes within a time window to extract target features after pulse emission. These studies suggest that the bat's auditory system must be highly sensitive to signal parameters within this time window. In the current study, we show that most neurons in the central nucleus of the inferior colliculus discharge maximally to a best duration and they have better echo frequency selectivity when the duration of both echo and pulse matches the best duration. This finding complements previous studies that show listeners can better detect a sound when its duration or frequency is expected than unexpected.

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Abstract

Interaural Time Sensitivity Dominated by Cochlea-Induced Envelope Patterns

To localize sounds in space, humans heavily depend on minute interaural time differences (ITDs) generated by path-length differences to the two ears. Physiological studies of ITD sensitivity have mostly used deterministic, periodic sounds, in which either the waveform fine structure or a sinusoidal envelope is delayed interaurally. For natural broadband stimuli, however, auditory frequency selectivity causes individual channels to have their own envelopes; the temporal code in these channels is thus a mixture of fine structure and envelope. This study introduces a method to disentangle the contributions of fine structure and envelope in both binaural and monaural responses to broadband noise. In the inferior colliculus (IC) of the cat, a population of neurons was found in which envelope fluctuations dominate ITD sensitivity. This population extends over a surprisingly wide range of frequencies, including low frequencies for which fine-structure information is also available. A comparison with the auditory nerve suggests that an elaboration of envelope coding occurs between the nerve and the IC. These results suggest that internally generated envelopes play a more important role in binaural hearing than is commonly thought.

Reconciling frequency selectivity and phase effects in masking.

The effects of auditory frequency selectivity and phase response on masking were studied using harmonic tone complex maskers with a 100-Hz fundamental frequency. Positive and negative Schroeder-phase complexes (m+ and m-), were used as maskers and the signal was a long-duration sinusoid. In the first experiment, thresholds for signal frequencies of 1 and 4 kHz were measured as a function of masker bandwidth and number of components. A large difference in thresholds between the m+ and m- complexes was found only when masker components were presented ipsilateral to the signal over a frequency range wider than the traditional critical band, regardless of the absolute number of components. In the second experiment, frequency selectivity was measured in harmonic tone complexes with fixed or random phases as well as in noise, using a variant of the notched-noise method with a fixed masker level. The data showed that frequency selectivity is not affected by masker type, indicating that the wide listening bandwidth suggested by the first experiment cannot be ascribed to broader effective filters in complex-tone maskers than in noise maskers. The third experiment employed a novel method of measuring frequency selectivity, which has the advantage that the overall level at the input and the output of the auditory filter remains roughly constant across all conditions. The auditory filter bandwidth measured using this method was wider than that measured in the second experiment, but may still be an underestimate, due to the effects of off-frequency listening. The data were modeled using a single-channel model with various initial filters. The main findings from the simulations were: (1) the magnitude response of the Gammatone filter is too narrow to account for the phase effects observed in the data; (2) none of the other filters currently used in auditory models can account for both frequency selectivity and phase effects in masking; (3) the Gammachirp filter can be made to provide a good account of the data by altering its phase response. The final conclusion suggests that masker phase effects can be accounted for with a single-channel model, while still remaining consistent with measures of frequency selectivity: effects that appear to involve broadband processing do not necessarily require across-channel mechanisms.

A computational algorithm for computing nonlinear auditory frequency selectivity.

Computational algorithms that mimic the response of the basilar membrane must be capable of reproducing a range of complex features that are characteristic of the animal observations. These include complex input output functions that are nonlinear near the site's best frequency, but linear elsewhere. This nonlinearity is critical when using the output of the algorithm as the input to models of inner hair cell function and subsequent auditory-nerve models of low- and high-spontaneous rate fibers. We present an algorithm that uses two processing units operating in parallel: one linear and the other compressively nonlinear. The output from the algorithm is the sum of the outputs of the linear and nonlinear processing units. Input to the algorithm is stapes motion and output represents basilar membrane motion. The algorithm is evaluated against published chinchilla and guinea pig observations of basilar membrane and Reissner's membrane motion made using laser velocimetry. The algorithm simulates both quantitatively and qualitatively, differences in input/output functions among three different sites along the cochlear partition. It also simulates quantitatively and qualitatively a range of phenomena including isovelocity functions, phase response, two-tone suppression, impulse response, and distortion products. The algorithm is potentially suitable for development as a bank of filters, for use in more comprehensive models of the peripheral auditory system.

Spectro-temporal modulation transfer functions and speech intelligibility.

Detection thresholds for spectral and temporal modulations are measured using broadband spectra with sinusoidally rippled profiles that drift up or down the log-frequency axis at constant velocities. Spectro-temporal modulation transfer functions (MTFs) are derived as a function of ripple peak density (omega cycles/octave) and drifting velocity (omega Hz). The MTFs exhibit a low-pass function with respect to both dimensions, with 50% bandwidths of about 16 Hz and 2 cycles/octave. The data replicate (as special cases) previously measured purely temporal MTFs (omega = 0) [Viemeister, J. Acoust. Soc. Am. 66, 1364-1380 (1979)] and purely spectral MTFs (omega = 0) [Green, in Auditory Frequency Selectivity (Plenum, Cambridge, 1986), pp. 351-359]. A computational auditory model is presented that exhibits spectro-temporal MTFs consistent with the salient trends in the data. The model is used to demonstrate the potential relevance of these MTFs to the assessment of speech intelligibility in noise and reverberant conditions.

Auditory frequency selectivity and masked hearing capabilities in an Atlantic bottlenose dolphin

Frequency selectivity capabilities of an Atlantic bottlenose dolphin were examined by calculating critical ratios from masked hearing data. Absolute sensitivity to pure tones from 40–140 kHz was measured as a base line, and masked sensitivity was determined for the same signals masked by three levels of white noise (52, 42, and 32 dB re:1 μ Pa2/Hz). Absolute and masked sensitivity were essentially constant between 40 and 120 kHz at each of the masking conditions. Sensitivity decreased approximately 100 dB per octave between 120 and 140 kHz. Critical ratios averaged across noise levels were constant between 40 and 120 kHz, averaging 26 dB. The critical ratio at 140 kHz was 42 dB. This flat trend in critical ratios between 40–120 kHz does not agree with the constant-Q filter-bank models used to account for earlier critical ratio and bandwidth measurements for the species [W. W. L. Au and P. W. B. Moore, J. Acoust. Soc. Am. 88, 1635–1638 (1990)], but comparison to other cetacean masked hearing work [Johnson et al., J. Acoust. Soc. Am. 85, 2651–2654 (1989)] indicates that the trend has been recorded before, but not explicitly reported.

Effects of Salicylate and Lanthanides on Outer Hair Cell Motility and Associated Gating Charge

Salicylate, one of the most widely used drugs, is known to induce reversible tinnitus and hearing loss. Salicylate interferes with outer hair cells (OHCs), which are believed to underlie normal auditory frequency selectivity and sensitivity. In the present experiments, the effects of salicylate and lanthanides on OHC motility and nonlinear capacitance were investigated by using isolated guinea-pig OHCs while attempting to avoid inadvertent intracellular pressure change, which itself can affect OHC motility and capacitance. Either extracellularly or intracellularly applied salicylate reduced nonlinear peak capacitance (Cmpk) and shifted the voltage at peak capacitance to depolarized levels. Concentration-response curves for reduction in Cmpk by salicylate and GdCl3 revealed a half-maximal concentration and Hill coefficient of 1.6 mM and 1.0, and 0.6 mM and 1.2, respectively. In comparable groups of OHCs, the normal Cmpk values of which were near 40 pF, average Cmpk decreased to 28 and 36 pF for intracellularly and extracellularly applied salicylate, respectively. Salicylate reduced, but did not completely block, the voltage-induced length change. Extracellularly, but not intracellularly, applied lanthanide blocked voltage-induced movement and capacitance almost completely. After intracellular trypsin treatment, salicylate reduced voltage-dependent capacitance reversibly, suggesting that salicylate directly acts on the sensor/motor and not via effects on intracellular structures, such as the subsurface cisternae. The results are consistent with the hypothesis that the dissociated, charged form of salicylate directly interacts with the sensor/motor on the inner aspect of the OHC plasma, whereas lanthanides interact on the outer aspect.

Tuned hair cells for hearing, but tuned basilar membrane for overload protection: Evidence from dolphins, bats, and desert rodents

A cochlear model is presented suggesting that the organ of Corti (OC) and the basilar membrane (BM) are both tuned resonant systems, but have different functions. The OC provides frequency filtering and amplification by means of tuned outer hair cells. The BM provides resonant absorption of excessive vibrational energy as an overload protection for vulnerable elements in the OC. Evidence supporting this model is demonstrated in dolphins, bats, and desert rodents. Specialized auditory capabilities correlate with cochlear deviations, some of them dramatically changing BM compliance. In characteristic regions along the cochlea there are BM thickenings and, on both sides of the OC, hypertrophied supporting cells. Structures of striking similarity have evolved independently across orders or families, revealing multiple events of convergent evolution. In all cases, the locations of deviating structures rule out a BM function in auditory frequency selectivity but support one in resonant absorption. Cochlear microphonics and BM responses demonstrate strongest high-level absorption in the frequency bands most vital for the tested species. The assumed cause is increased internal damping in the enlarged structures during BM motion. Species with intermediate specializations supply further evidence that resonant absorption is universally the genuine function of BM mechanics in mammals, providing complementary high-level protection of low-level sensitivity.

Effects of aspirin on human auditory filters

Auditory filter shapes were measured in eight male volunteers with normal hearing, using a notched-noise forward-masking paradigm and a signal frequency of 4 kHz. The measurements were made under three conditions: after listeners had taken eight doses of three 320 mg aspirin tablets every six hours; after an identical schedule of placebo ingestion; and one week after testing in the first two conditions had been completed. Half of the listeners did the placebo condition first, and half did the aspirin condition first. Aspirin and placebo were administered double-blind, and testing took place approximately one hour after the last dose. Filter shapes were significantly broader in the aspirin condition than in the placebo and post-test conditions, indicating that even a modest dose affects auditory frequency selectivity. Two-point measures of growth of masking did not differ significantly between conditions.

Auditory processing of complex sounds: an overview.

The past 30 years has seen a remarkable development in our understanding of how the auditory system--particularly the peripheral system--processes complex sounds. Perhaps the most significant has been our understanding of the mechanisms underlying auditory frequency selectivity and their importance for normal and impaired auditory processing. Physiologically vulnerable cochlear filtering can account for many aspects of our normal and impaired psychophysical frequency selectivity with important consequences for the perception of complex sounds. For normal hearing, remarkable mechanisms in the organ of Corti, involving enhancement of mechanical tuning (in mammals probably by feedback of electro-mechanically generated energy from the hair cells), produce exquisite tuning, reflected in the tuning properties of cochlear nerve fibres. Recent comparisons of physiological (cochlear nerve) and psychophysical frequency selectivity in the same species indicate that the ear's overall frequency selectivity can be accounted for by this cochlear filtering, at least in bandwidth terms. Because this cochlear filtering is physiologically vulnerable, it deteriorates in deleterious conditions of the cochlea--hypoxia, disease, drugs, noise overexposure, mechanical disturbance--and is reflected in impaired psychophysical frequency selectivity. This is a fundamental feature of sensorineural hearing loss of cochlear origin, and is of diagnostic value. This cochlear filtering, particularly as reflected in the temporal patterns of cochlear fibres to complex sounds, is remarkably robust over a wide range of stimulus levels. Furthermore, cochlear filtering properties are a prime determinant of the 'place' and 'time' coding of frequency at the cochlear nerve level, both of which appear to be involved in pitch perception. The problem of how the place and time coding of complex sounds is effected over the ear's remarkably wide dynamic range is briefly addressed. In the auditory brainstem, particularly the dorsal cochlear nucleus, are inhibitory mechanisms responsible for enhancing the spectral and temporal contrasts in complex sounds. These mechanisms are now being dissected neuropharmacologically. At the cortical level, mechanisms are evident that are capable of abstracting biologically relevant features of complex sounds. Fundamental studies of how the auditory system encodes and processes complex sounds are vital to promising recent applications in the diagnosis and rehabilitation of the hearing impaired.

Frequency selectivity as a potential measure of noise damage susceptibility.

The notched-noise procedure of Patterson (1974, Journal of the Accoustical Society of America, 55, 802-809; 1976, Journal of the Acoustical Society of America, 59, 640-654) was employed in revealing changes in auditory frequency selectivity (FS) in noise-exposed industrial workers in manufacturing plants in which noise levels in the working environment exceeded 90 dB(A). With 4 kHz as the test frequency, both threshold and FS measures were applied to three groups of non-exposed young adults (total N = 61) for reference values and to workers in three different factories, some ('noise-exposed') working in noise levels over 90 dB(A) (total N = 222). In another study involving 185 workers only the FS test was repeated after a work session, and the results are reported on 31 who worked in high noise levels. Pre-work-shift data showed that when thresholds were better than 30-dB HL most FS values were 10 dB or more, while subjects with hearing threshold levels beyond that showed reduced FS. Two post-work-shift studies of 144 ears of the noise-exposed workers showed significant deleterious changes in both thresholds and FS but the relationships were different in the two studies. The findings indicate that the notched-noise test is sensitive to noise-induced auditory damage, even after a single work-shift. Attractive features of the procedure are that it reduces the need for a quiet test environment and that it can be more resistant to testee manipulation or threshold 'learning' effects, often seen in industrial audiometric testing.

Reflection of retrograde waves within the cochlea and at the stapes

A number of authors [de Boer and Viergever, Hear. Res. 13, 101-112 (1984); de Boer et al., in Peripheral Auditory Mechanisms (Springer-Verlag, Berlin, 1986); Hear. Res. 23, 1-7 (1986); Viergever, in Auditory Frequency Selectivity (Plenum, New York, 1986), pp. 31-38; Kaernbach et al., J. Acoust. Soc. Am. 81, 408-411 (1987)] have argued that backward-traveling waves, in striking contrast to waves traveling forward towards the helicotrema, suffer appreciable reflection as they move through the basal turns of the cochlea. Such reflection, if present, would have important consequences for understanding the nature and strength of otoacoustic emissions. The apparent asymmetry in reflection of cochlear waves is shown, however, to be an artifact of the boundary condition those authors impose at the stapes: conventional cochlear models are found not to generate reflections of waves traveling in either direction even when the wavelength changes rapidly and the WKB approximation breaks down. Although backward-traveling waves are not reflected by the secular variation of the geometrical and mechanical characteristics of the cochlea, they are reflected when they reach the stapes. The magnitude of that boundary reflection is computed for the cat and shown to be a large, rapidly varying function of frequency.