Cochlear nonlinearity between 500 and 8000 Hz in listeners with normal hearing

Cochlear nonlinearity was estimated over a wide range of center frequencies and levels in listeners with normal hearing, using a forward-masking method. For a fixed low-level probe, the masker level required to mask the probe was measured as a function of the masker-probe interval, to produce a temporal masking curve (TMC). TMCs were measured for probe frequencies of 500, 1000, 2000, 4000, and 8000 Hz, and for masker frequencies 0.5, 0.7, 0.9, 1.0 (on frequency), 1.1, and 1.6 times the probe frequency. Across the range of probe frequencies, the TMCs for on-frequency maskers showed two or three segments with clearly distinct slopes. If it is assumed that the rate of decay of the internal effect of the masker is constant across level and frequency, the variations in the slopes of the TMCs can be attributed to variations in cochlear compression. Compression-ratio estimates for on-frequency maskers were between 3:1 and 5:1 across the range of probe frequencies. Compression did not decrease at low frequencies. The slopes of the TMCs for the lowest frequency probe (500 Hz) did not change with masker frequency. This suggests that compression extends over a wide range of stimulus frequencies relative to characteristic frequency in the apical region of the cochlea.     

A new procedure for measuring peripheral compression in normal-hearing and hearing-impaired listeners.

Forward-masking growth functions for on-frequency (6-kHz) and off-frequency (3-kHz) sinusoidal maskers were measured in quiet and in a high-pass noise just above the 6-kHz probe frequency. The data show that estimates of response-growth rates obtained from those functions in quiet, which have been used to infer cochlear compression, are strongly dependent on the spread of probe excitation toward higher frequency regions. Therefore, an alternative procedure for measuring response-growth rates was proposed, one that employs a fixed low-level probe and avoids level-dependent spread of probe excitation. Fixed-probe-level temporal masking curves (TMCs) were obtained from normal-hearing listeners at a test frequency of 1 kHz, where the short 1-kHz probe was fixed in level at about 10 dB SL. The level of the preceding forward masker was adjusted to obtain masked threshold as a function of the time delay between masker and probe. The TMCs were obtained for an on-frequency masker (1 kHz) and for other maskers with frequencies both below and above the probe frequency. From these measurements, input/output response-growth curves were derived for individual ears. Response-growth slopes varied from >1.0 at low masker levels to <0.2 at mid masker levels. In three subjects, response growth increased again at high masker levels (>80 dB SPL). For the fixed-level probe, the TMC slopes changed very little in the presence of a high-pass noise masking upward spread of probe excitation. A greater effect on the TMCs was observed when a high-frequency cueing tone was used with the masking tone. In both cases, however, the net effects on the estimated rate of response growth were minimal.     

Inferred basilar-membrane response functions for listeners with mild to moderate sensorineural hearing loss

Psychophysical estimates of cochlear function suggest that normal-hearing listeners exhibit a compressive basilar-membrane (BM) response. Listeners with moderate to severe sensorineural hearing loss may exhibit a linearized BM response along with reduced gain, suggesting the loss of an active cochlear mechanism. This study investigated how the BM response changes with increasing hearing loss by comparing psychophysical measures of BM compression and gain for normal-hearing listeners with those for listeners who have mild to moderate sensorineural hearing loss. Data were collected from 16 normal-hearing listeners and 12 ears from 9 hearing-impaired listeners. The forward masker level required to mask a fixed low-level, 4000-Hz signal was measured as a function of the masker-signal interval using a masker frequency of either 2200 or 4000 Hz. These plots are known as temporal masking curves (TMCs). BM response functions derived from the TMCs showed a systematic reduction in gain with degree of hearing loss. Contrary to current thinking, however, no clear relationship was found between maximum compression and absolute threshold.     

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

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

The effect of basilar-membrane nonlinearity on the shapes of masking period patterns in normal and impaired hearing

Masking period patterns (MPPs) were measured in listeners with normal and impaired hearing using amplitude-modulated tonal maskers and short tonal probes. The frequency of the masker was either the same as the frequency of the probe (on-frequency masking) or was one octave below the frequency of the probe (off-frequency masking). In experiment 1, MPPs were measured for listeners with normal hearing using different masker levels. Carrier frequencies of 3 and 6 kHz were used for the masker. The probe had a frequency of 6 kHz. For all masker levels, the off-frequency MPPs exhibited deeper and longer valleys compared with the on-frequency MPPs. Hearing-impaired listeners were tested in experiment 2. For some hearing-impaired subjects, masker frequencies of 1.5 kHz and 3 kHz were paired with a probe frequency of 3 kHz. MPPs measured for listeners with hearing loss had similar shapes for on- and off-frequency maskers. It was hypothesized that the shapes of MPPs reflect nonlinear processing at the level of the basilar membrane in normal hearing and more linear processing in impaired hearing. A model assuming different cochlear gains for normal versus impaired hearing and similar parameters of the temporal integrator for both groups of listeners successfully predicted the MPPs.     

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.     

Forward masking patterns produced by intracochlear electrical stimulation of one and two electrode pairs in the human cochlea

Three psychophysical forward masking studies were conducted on a multichannel cochlear implant patient. The first study investigated the masking pattern produced by a bipolar electrode pair at different stimulus currents. It was found that the masking pattern for a single-masker bipolar electrode pair had a maximum located at an electrode position where the masker and probe coincided. The spread of the masking pattern was not symmetrical about the maximum. The amount of masking decreased very rapidly toward the apical direction and less rapidly toward the basal direction from the position of the maximum. As the stimulus current increased, the amount of masking at the maximum increased and the masking pattern broadened toward the base. The second study investigated the masking pattern produced by the activation of single bipolar electrode pairs with different spatial extents. The spatial extent of a bipolar electrode pair is defined as the distance between the apical and basal electrode members of the bipolar pair. With a small spatial extent (1.5 mm), the more basal electrode pairs (higher threshold and smaller dynamic range) produced broader masking patterns than the more apical electrode pairs (lower threshold and wider dynamic range), suggesting that there was more current spread at the basal region. With a larger spatial extent (4.5 mm), an additional secondary masking maximum was observed in the vicinity of the apical electrode member of the masker; this was observed only when the apical electrode member lay within the low-threshold apical region. The third study investigated the masking patterns produced by two loudness balanced bipolar masker electrode pairs activated within a stimulus period (inverse of the pulse repetition rate). The biphasic current pulses delivered to the two electrode pairs were nonoverlapping in time. It was found that, at any probe electrode position, the amount of masking produced by the two combined bipolar electrode pairs approximately followed the greater of the two maskings produced respectively by the two individual bipolar masker electrode pairs.     

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.     

Enhancements of the edges of temporal masking functions by complex patterns of overshoot and undershoot

The purpose of this report is to present new data that provide a novel perspective on temporal masking, different from that found in the classical auditory literature on this topic. Specifically, measurement conditions are presented that minimize rather than maximize temporal spread of masking for a gated (200-ms) narrow-band (405-Hz-wide) noise masker logarithmically centered at 2500 Hz. Masked detection thresholds were measured for brief sinusoids in a two-interval, forced-choice (21FC) task. Detection was measured at each of 43 temporal positions within the signal observation interval for the sinusoidal signal presented either preceding, during, or following the gating of the masker, which was centered temporally within each 500-ms observation interval. Results are presented for three listeners; first, for detection of a 1900-Hz signal across a range of masker component levels (0-70 dB SPL) and, second, for masked detection as a function of signal frequency (fs = 500-5000 Hz) for a fixed masker component level (40 dB SPL). For signals presented off-frequency from the masker, and at low-to-moderate masker levels, the resulting temporal masking functions are characterized by sharp temporal edges. The sharpness of the edges is accentuated by complex patterns of temporal overshoot and undershoot, corresponding with diminished and enhanced detection, respectively, at both masker onset and offset. This information about the onset and offset timing of the gated masker is faithfully represented in the temporal masking functions over the full decade range of signal frequencies (except for fs=2500 Hz presented at the center frequency of the masker). The precise representation of the timing information is remarkable considering that the temporal envelope characteristics of the gated masker are evident in the remote masking response at least two octaves below the frequencies of the masker at a cochlear place where little or no masker activity would be expected. This general enhancement of the temporal edges of the masking response is reminiscent of spectral edge enhancement by lateral suppression/inhibition.     

Excess masking among listeners with a sensorineural hearing loss

Three experiments were conducted to determine whether listeners with a sensorineural hearing loss exhibited greater than normal amounts of masking at frequencies above the frequency of the masker. Excess masking was defined as the difference (in dB) between the masked thresholds actually obtained from a hearing-impaired listener and the expected thresholds calculated for the same individual. The expected thresholds were the power sum of the listener's thresholds in quiet and the average masked thresholds obtained from a group of normal-hearing subjects at the test frequency. Hearing-impaired listeners, with thresholds in quiet ranging from approximately 35-70 dB SPL (at test frequencies between 500-3000 Hz), displayed approximately 12-15 dB of maximum excess masking. The maximum amount of excess masking occurred in the region where the threshold in quiet of the hearing-impaired listener and the average normal masked threshold were equal. These findings indicate that listeners with a sensorineural hearing loss display one form of reduced frequency selectivity (i.e., abnormal upward spread of masking) even when their thresholds in quiet are taken into account.     

Comparing different estimates of cochlear compression in listeners with normal and impaired hearing

A loss of cochlear compression may underlie many of the difficulties experienced by hearing-impaired listeners. Two behavioral forward-masking paradigms that have been used to estimate the magnitude of cochlear compression are growth of masking (GOM) and temporal masking (TM). The aim of this study was to determine whether these two measures produce within-subjects results that are consistent across a range of signal frequencies and, if so, to compare them in terms of reliability or efficiency. GOM and TM functions were measured in a group of five normal-hearing and five hearing-impaired listeners at signal frequencies of 1000, 2000, and 4000 Hz. Compression values were derived from the masking data and confidence intervals were constructed around these estimates. Both measures produced comparable estimates of compression, but both measures have distinct advantages and disadvantages, so that the more appropriate measure depends on factors such as the frequency region of interest and the degree of hearing loss. Because of the long testing times needed, neither measure is suitable for clinical use in its current form.     

Relations among some psychoacoustic parameters in normal and cochlearly impaired listeners

Frequency resolution (viz., masking by low-pass-filtered noise and broadband noise) and temporal resolution (viz., masking by interrupted noise) were compared with hearing thresholds and acoustic reflex thresholds for four normally hearing and 13 cochlearly impaired subjects. Two models, one for frequency resolution (model I) and one for temporal resolution (model II), were introduced, and these provided a means of predicting individual frequency and temporal resolution from hearing thresholds for both normal-hearing and hearing-impaired listeners. Model I is based on the assumption that the upward spread of masking increases in cochlearly impaired hearing with an amount proportional to the hearing threshold in dB HL. Model II is based on the assumption that the poststimulatory masked thresholds return to the level of the hearing threshold within a duration of 200 ms, independent of the level of the masker and the amount of cochlear hearing loss. Model parameters were determined from results from other studies. Although some discrepancies between measured and predicted values were observed, the model predictions generally agree with measurements. Thus, to a first-order approximation, it seems possible to predict individual frequency and temporal resolution of cochlearly hearing-impaired listeners solely on the basis of their hearing thresholds.     

Temporal integration and compression near absolute threshold in normal and impaired ears

The decrease in absolute threshold with increasing stimulus duration (often referred to as "temporal integration") is greater for listeners with normal hearing than for listeners with sensorineural hearing loss. It has been suggested that the difference is related to reduced basilar-membrane (BM) compression in the impaired group. The present experiment tested this hypothesis by comparing temporal integration and BM compression in normal and impaired ears at low levels. Absolute thresholds were measured for 4, 24, and 44 ms pure-tone signals, with frequencies (f(s)) of 2 and 4 kHz. The difference between the absolute thresholds for the 4 and 24 ms signals was used as a measure of temporal integration. Compression near threshold was estimated by measuring the level of a 100 ms off-frequency (0.45f(s)) pure-tone forward masker required to mask a 44 ms pure-tone signal presented at sensation levels of 5 and 10 dB. There was a significant negative correlation between amount of temporal integration and absolute threshold. However, there was no correlation between absolute threshold and compression at low levels; both normal and impaired ears showed a nearly linear response. The results suggest that the differences in integration between normal and impaired ears cannot be explained by differences in BM compression.     

The effects of cochlear hypothermia on compound action potential tuning

The effects of lowered cochlear temperature on eighth-nerve tuning were assessed by using forward masking of whole nerve action potential (AP) responses to generate AP tuning curves (APTCs) at cochlear temperatures ranging from 38.5 degrees to 30 degrees C for probe frequencies from 8 to 36 kHz. The data indicate that subnormal cochlear temperatures result in: broadened APTCs for probe frequencies above 10 kHz which are interpreted as resulting from reduced hair-cell frequency selectivity, lowered or more sensitive APTC tips where tone-burst thresholds are unchanged, and raised or less sensitive tips where thresholds to tone bursts were elevated. Increased tip sensitivity is explained in terms of enhanced eighth-nerve adaptation which occurred during hypothermia. Experiments directly addressing adaptation were performed, in which the masker-probe interval (delta t) was systematically lengthened. The normalized AP decrement versus delta t functions indicate an enhancement of both the amount and duration of adaptation during hypothermia. Functions relating the growth of response to the masker (AP decrement versus masker intensity functions) were reduced at low temperatures.     

Basilar-membrane nonlinearity estimated by pulsation threshold

The pulsation threshold technique was used to estimate the basilar-membrane (BM) response to a tone at characteristic frequency (CF). A pure-tone signal was alternated with a pure-tone masker. The frequency of the masker was 0.6 times that of the signal. For signal levels from around 20 dB above absolute threshold to 85 dB SPL, the masker level was varied to find the level at which a transition occurred between the signal being perceived as "pulsed" or "continuous" (the pulsation threshold). The transition is assumed to occur when the masker excitation is somewhat greater than the signal excitation at the place on the BM tuned to the signal. If it is assumed further that the response at this place to the lower-frequency masker is linear, then the shape of the masking function provides an estimate of the BM response to the signal. Signal frequencies of 0.25, 0.5, 1, 2, 4, and 8 kHz were tested. The mean slopes of the masking functions for signal levels between 50 and 80 dB SPL were 0.76, 0.50, 0.34, 0.32, 0.35, and 0.41, respectively. The results suggest that compression on the BM increases between CFs of 0.25 and 1 kHz and is roughly constant for frequencies of 1 kHz and above. Despite requiring a subjective criterion, the pulsation threshold measurements had a reasonably low variability. However, the estimated compression was less than in an earlier study using forward masking. The smaller amount of compression observed here may be due to the effects of off-frequency listening.     

Two-tone suppression of stimulus frequency otoacoustic emissions

Stimulus frequency otoacoustic emissions (SFOAEs) measured using a suppressor tone in human ears are analogous to two-tone suppression responses measured mechanically and neurally in mammalian cochleae. SFOAE suppression was measured in 24 normal-hearing adults at octave frequencies (f(p)=0.5-8.0 kHz) over a 40 dB range of probe levels (L(p)). Suppressor frequencies (f(s)) ranged from -2.0 to 0.7 octaves re: f(p), and suppressor levels ranged from just detectable suppression to full suppression. The lowest suppression thresholds occurred for "best" f(s) slightly higher than f(p). SFOAE growth of suppression (GOS) had slopes close to one at frequencies much lower than best f(s), and shallow slopes near best f(s), which indicated compressive growth close to 0.3 dBdB. Suppression tuning curves constructed from GOS functions were well defined at 1, 2, and 4 kHz, but less so at 0.5 and 8.0 kHz. Tuning was sharper at lower L(p) with an equivalent rectangular bandwidth similar to that reported behaviorally for simultaneous masking. The tip-to-tail difference assessed cochlear gain, increasing with decreasing L(p) and increasing f(p) at the lowest L(p) from 32 to 45 dB for f(p) from 1 to 4 kHz. SFOAE suppression provides a noninvasive measure of the saturating nonlinearities associated with cochlear amplification on the basilar membrane.     

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.     

Ipsilateral masking between acoustic and electric stimulations

Residual acoustic hearing can be preserved in the same ear following cochlear implantation with minimally traumatic surgical techniques and short-electrode arrays. The combined electric-acoustic stimulation significantly improves cochlear implant performance, particularly speech recognition in noise. The present study measures simultaneous masking by electric pulses on acoustic pure tones, or vice versa, to investigate electric-acoustic interactions and their underlying psychophysical mechanisms. Six subjects, with acoustic hearing preserved at low frequencies in their implanted ear, participated in the study. One subject had a fully inserted 24 mm Nucleus Freedom array and five subjects had Iowa/Nucleus hybrid implants that were only 10 mm in length. Electric masking data of the long-electrode subject showed that stimulation from the most apical electrodes produced threshold elevations over 10 dB for 500, 625, and 750 Hz probe tones, but no elevation for 125 and 250 Hz tones. On the contrary, electric stimulation did not produce any electric masking in the short-electrode subjects. In the acoustic masking experiment, 125-750 Hz pure tones were used to acoustically mask electric stimulation. The acoustic masking results showed that, independent of pure tone frequency, both long- and short-electrode subjects showed threshold elevations at apical and basal electrodes. The present results can be interpreted in terms of underlying physiological mechanisms related to either place-dependent peripheral masking or place-independent central masking.     

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.     

Forward masking among infant and adult listeners

Psychophysical forward-masked thresholds were estimated for 3- and 6-month-old infants and for adults. Listeners detected a repeated 1000-Hz probe, with 16-ms rise time, no steady-state duration, and 16-ms fall time. Unmasked thresholds were determined for one group of listeners who were trained to respond when they heard the probe but not at other times. In the masking conditions, each tone burst was preceded by a 100-ms broadband noise masker at 65 dB SPL. Listeners were trained to respond when they heard the probe and masker, but not when they heard the masker alone. The masker-probe interval, delta t, was either 5, 10, 25, or 200 ms. Four groups of subjects listened in the masked conditions, each at one value of delta t. Each listener attempted to complete a block of 32 trials including four probe levels chosen to span the range of expected thresholds. "Group" thresholds, based on average psychometric functions, as well as thresholds for individual listeners, were estimated. Both group and individual thresholds declined with delta t, as expected, for both infants and adults. Infants' masked thresholds were higher than those of adults, and comparison of masked to unmasked thresholds suggested that infants demonstrate more forward masking than adults, particularly at short delta t. Forward masking appeared to have greater effects on 3-month-olds' detection than on either 6-month-olds' or adults'. Compared to adults, 6-month-olds demonstrated more forward masking only for delta t of 5 ms. Thus, susceptibility to forward masking may be nearly mature by 6 months of age.