Behaviors of cubic distortion product otoacoustic emissions evoked by amplitude modulated tones

Distortion product otoacoustic emissions (DPOAEs) were measured using sinusoidal amplitude modulation (AM) tones. When one of the primary stimuli (f(1) or f(2), f(1)?< f(2)) was amplitude modulated, a series of changes in the cubic difference tone (CDT) were observed. In the frequency domain, multiple sidebands were present around the CDT and their sizes grew with the modulation depth of the AM stimulus. In the time domain, the CDT showed different modulation patterns between two major signal conditions: the AM tone was used as the f(1) or the f(2). The CDT amplitude followed the AM tone when the f(1) was amplitude modulated. However, when the AM tone acted as the f(2), the CDT showed a more complex modulation pattern with a notch present at the AM tone peak. The relatively linear dependence of CDT on f(1) and the nonlinear relation with f(2) can be explained with a variable gain-control model representing hair cell functions at the DPOAE generation site. It is likely that processing of AM signals at a particular cochlear location depends on whether the hair cells are tuned to the frequency of the carrier. Nonlinear modulation is related to on-frequency carriers and off-frequency carriers are processed relatively linearly.     

Effects of low-frequency biasing on spontaneous otoacoustic emissions: amplitude modulation

The dynamic effects of low-frequency biasing on spontaneous otoacoustic emissions (SOAEs) were studied in human subjects under various signal conditions. Results showed a combined suppression and modulation of the SOAE amplitudes at high bias tone levels. Ear-canal acoustic spectra demonstrated a reduction in SOAE amplitude and growths of sidebands while increasing the bias tone level. These effects varied depending on the relative strength of the bias tone to a particular SOAE. The SOAE magnitudes were suppressed when the cochlear partition was biased in both directions. This quasi-static modulation pattern showed a shape consistent with the first derivative of a sigmoid-shaped nonlinear function. In the time domain, the SOAE amplitudes were modulated with the instantaneous phase of the bias tone. For each biasing cycle, the SOAE envelope showed two peaks each corresponded to a zero crossing of the bias tone. The temporal modulation patterns varied systematically with the level and frequency of the bias tone. These dynamic behaviors of the SOAEs are consistent with the shifting of the operating point along the nonlinear transducer function of the cochlea. The results suggest that the nonlinearity in cochlear hair cell transduction may be involved in the generation of SOAEs.     

Cochlear compression: effects of low-frequency biasing on quadratic distortion product otoacoustic emission

Distortion product otoacoustic emissions (DPOAEs) are generated from the nonlinear transduction n cochlear outer hair cells. The transducer function demonstrating a compressive nonlinearity can be estimated from low-frequency modulation of DPOAEs. Experimental results from the gerbils showed that the magnitude of quadratic difference tone (QDT, f2-f1) was either enhanced or suppressed depending on the phase of the low-frequency bias tone. Within one period of the bias tone, QDT magnitudes exhibited two similar modulation patterns, each resembling the absolute value of the second derivative of the transducer function. In the time domain, the center notches of the modulation patterns occurred around the zero crossings of the bias pressure, whereas peaks corresponded to the increase or decrease in bias pressure. Evaluated with respect to the bias pressure, modulated QDT magnitude displayed a double-modulation pattern marked by a separation of the center notches. Loading/unloading of the cochlear transducer or rise/fall in bias pressure shifted the center notch to positive or negative sound pressures, indicating a mechanical hysteresis. These results suggest that QDT arises from the compression that coexists with the active hysteresis in cochlear transduction. Modulation of QDT magnitude reflects the dynamic regulation of cochlear transducer gain and compression.     

Spectral fine-structures of low-frequency modulated distortion product otoacoustic emissions

Biasing of the cochlear partition with a low-frequency tone can produce an amplitude modulation of distortion product otoacoustic emissions (DPOAEs) in gerbils. In the time domain, odd- versus even-order DPOAEs demonstrated different modulation patterns depending on the bias tone phase. In the frequency domain, multiple sidebands are presented on either side of each DPOAE component. These sidebands were located at harmonic multiples of the biasing frequency from the DPOAE component. For odd-order DPOAEs, sidebands at the even-multiples of the biasing frequency were enhanced, while for even-order DPOAEs, the sidebands at the odd-multiples were elevated. When a modulation in DPOAE magnitude was presented, the magnitudes of the sidebands were enhanced and even greater than the DPOAEs. The amplitudes of these sidebands varied with the levels of the bias tone and two primary tones. The results indicate that the maximal amplitude modulations of DPOAEs occur at a confined bias and primary level space. This can provide a guide for optimal selections of signal conditions for better recordings of low-frequency modulated DPOAEs in future research and applications. Spectral fine-structure and its unique relation to the DPOAE modulation pattern may be useful for direct acquisition of cochlear transducer nonlinearity from a simple spectral analysis.     

Deriving a cochlear transducer function from low-frequency modulation of distortion product otoacoustic emissions

In this paper, a new method is introduced to derive a cochlear transducer function from measuring distortion product otoacoustic emissions (DPOAEs). It is shown that the cubic difference tone (CDT, 2f1-f2) is produced from the odd-order terms of a power series that approximates a nonlinear function characterizing cochlear transduction. Exploring the underlying mathematical formulation, it is found that the CDT is proportional to the third derivative of the transduction function when the primary levels are sufficiently small. DPOAEs were measured from nine gerbils in response to two-tone signals biased by a low-frequency tone with different amplitudes. The CDT magnitude was obtained at the peak regions of the bias tone. The results of the experiment demonstrated that the shape of the CDT magnitudes as a function of bias levels was similar to the absolute value of the third derivative of a sigmoidal function. A second-order Boltzmann function was derived from curve fitting the CDT data with an equation that represents the third derivative of the Boltzmann function. Both the CDT-bias function and the derived nonlinear transducer function showed effects of primary levels. The results of the study indicate that the low-frequency modulated DPOAEs can be used to estimate the cochlear transducer function.     

Constructing a cochlear transducer function from the summating potential using a low-frequency bias tone

A new method is developed to construct a cochlear transducer function using modulation of the summating potential (SP), a dc component of the electrical response of the cochlea to a sinusoid. It is mathematically shown that the magnitude of the SP is determined by the even-order terms of the power series representing a nonlinear function. The relationship between the SP magnitudes and the second derivative of the transducer function was determined by using a low-frequency bias tone to position a high-frequency probe tone at different places along the cochlear transducer function. Two probe tones (6 kHz and 12 kHz) ranging from 70 to 90 dB SPL and a 25-Hz bias tone at 130 dB SPL were simultaneously presented. Electric responses from the cochlea were recorded by an electrode placed at the round window to obtain the SP magnitudes. The experimental results from eight animals demonstrated that the SP magnitudes as a function of bias levels are essentially proportional to the second derivative of a sigmoidal Boltzmann function. This suggests that the low-frequency modulated SP amplitude can be used to construct a cochlear transducer function.     

Maturation of cochlear nonlinearity as measured by distortion product otoacoustic emission suppression growth in humans

The growth of distortion product otoacoustic emission (DPOAE) suppression follows a systematic, frequency-dependent pattern. The pattern is consistent with direct measures of basilar-membrane response growth, psychoacoustic measures of masking growth, and measures of neural rate growth. This pattern has its basis in the recognized nonlinear properties of basilar-membrane motion and, as such, the DPOAE suppression growth paradigm can be applied to human neonates to study the maturation of cochlear nonlinearity. The objective of this experiment was to investigate the maturation of human cochlear nonlinearity and define the time course for this maturational process. Normal-hearing adults, children, term-born neonates, and premature neonates, plus a small number of children with sensorineural hearing loss, were included in this experiment. DPOAE suppression growth was measured at two f2 frequencies (1500 and 6000 Hz) and three primary tone levels (55-45, 65-55, and 75-65 dB SPL). Slope of DPOAE suppression growth, as well as an asymmetry ratio (to compare slope for suppressor tones below and above f2 frequency), were generated. Suppression threshold was also measured in all subjects. Findings indicate that both term-born neonates and premature neonates who have attained term-like age, show non-adult-like DPOAE suppression growth for low-frequency suppressor tones. These age effects are most evident at f2 = 6000 Hz. In neonates, suppression growth is shallower and suppression thresholds are elevated for suppressor tones lower in frequency than f2. Additionally, the asymmetry ratio is smaller in neonates, indicating that the typical frequency-dependent pattern of suppression growth is not present. These findings suggest that an immaturity of cochlear nonlinearity persists into the first months of postnatal life. DPOAE suppression growth examined for a small group of hearing-impaired children also showed abnormalities.     

Two-tone suppression in auditory nerve of the cat: rate-intensity and temporal analyses

Responses to two-tone stimuli were recorded from auditory-nerve fibers in anesthetized cats. One tone, the suppressor, was set at a frequency above characteristic frequency and was fixed in intensity. A second tone was set at an excitatory frequency and was varied in intensity. The suppressor tone, when set at a sufficient level, always reduced the response to the excitatory tone by an amount equivalent to a fixed number of decibels, regardless of the excitatory tone's intensity. Estimates of suppression magnitude were derived from shifts in rate-intensity function obtained when the suppressor tone was present relative to the functions obtained for the excitatory tone alone. When suppressor-tone intensity was increased, suppression magnitude likewise increased. When the two tones were increasingly separated in frequency, either by varying the excitor or by varying the suppressor, suppression magnitude decreased monotonically. Suppression behaved in the same manner regardless of whether suppresor tone was excitatory or nonexcitatory. When frequency separation was small enough and when both tones were above the neuron's characteristic frequency, responses synchronized to low-order combination tones could be elicited. These responses usually possessed different rate-intensity characteristics and resulted in estimates of suppression magnitude which were spuriously low. When frequency separation is normalized with regard to position of traveling wave maxima within the cochlear duct, the magnitude of two-tone suppression for a given suppressor-tone intensity is seen to be frequency independent.     

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.     

Transient-evoked stimulus-frequency and distortion-product otoacoustic emissions in normal and impaired ears

Transient-evoked stimulus-frequency otoacoustic emissions (SFOAEs), recorded using a nonlinear differential technique, and distortion-product otoacoustic emissions (DPOAEs) were measured in 17 normal-hearing and 10 hearing-impaired subjects using pairs of tone pips (pp), gated tones (gg), and for DPOAEs, continuous and gated tones (cg). Temporal envelopes of stimulus and OAE waveforms were obtained by narrow-band filtering at the stimulus or DP frequency. Mean SFOAE latencies in normal ears at 2.7 and 4.0 kHz decreased with increasing stimulus level and were larger at 4.0 kHz than latencies in impaired ears. Equivalent auditory filter bandwidths were calculated as a function of stimulus level from SFOAE latencies by assuming that cochlear transmission is minimum phase. DPOAE latencies varied less with level than SFOAE latencies. The ppDPOAEs often had two (or more) peaks separated in time with latencies consistent with model predictions for distortion and reflection components. Changes in ppDPOAE latency with level were sometimes explained by a shift in relative amplitudes of distortion and reflection components. The pp SFOAE SPL within the main spectral lobe of the pip stimulus was higher for normal ears in the higher-frequency half of the pip than the lower-frequency half, which is likely an effect of basilar membrane two-tone suppression.     

Ipsilateral distortion product otoacoustic emission (2f1-f2) suppression in children with sensorineural hearing loss

Distortion product otoacoustic emission (DPOAE) ipsilateral suppression has been applied to study cochlear function and maturation in laboratory animals and humans. Although DPOAE suppression appears to be sensitive to regions of specialized cochlear function and to cochlear immaturity, it is not known whether it reflects permanent cochlear damage, i.e., sensorineural hearing loss (SNHL), in a reliable and systematic manner in humans. Eight school-aged children with mild-moderate SNHL and 20 normal-hearing children served as subjects in this study. DPOAE (2f1-f2) suppression data were collected at four f2 frequencies (1500, 3000, 4000, and 6000 Hz) using moderate-level primary tones. Features of the DPOAE iso-suppression tuning curves and suppression growth were analyzed for both subject groups. Results show that DPOAE suppression tuning curves from hearing-impaired subjects can be reliably recorded. DPOAE suppression tuning curves were generally normal in appearance and shape for six out of eight hearing-impaired subjects but showed subtle abnormalities in at least one feature. There was not one single trend or pattern of abnormality that characterized all hearing-impaired subjects. The most prominent patterns of abnormality included: broadened tuning, elevated tip, and downward shift of tip frequency. The unique patterns of atypical DPOAE suppression in subjects with similar audiograms may suggest different patterns of underlying sensory cell damage. This speculation warrants further investigation.     

Moderate cochlear hearing loss leads to a reduced ability to use temporal fine structure information

The ability of normally hearing and hearing-impaired subjects to use temporal fine structure information in complex tones was measured. Subjects were required to discriminate a harmonic complex tone from a tone in which all components were shifted upwards by the same amount in Hz, in a three-alternative, forced-choice task. The tones either contained five equal-amplitude components (non-shaped stimuli) or contained many components, but were passed through a fixed bandpass filter to reduce excitation pattern changes (shaped stimuli). Components were centered at nominal harmonic numbers (N) 7, 11, and 18. For the shaped stimuli, hearing-impaired subjects performed much more poorly than normally hearing subjects, with most of the former scoring no better than chance when N=11 or 18, suggesting that they could not access the temporal fine structure information. Performance for the hearing-impaired subjects was significantly improved for the non-shaped stimuli, presumably because they could benefit from spectral cues. It is proposed that normal-hearing subjects can use temporal fine structure information provided the spacing between fine structure peaks is not too small relative to the envelope period, but subjects with moderate cochlear hearing loss make little use of temporal fine structure information for unresolved components.     

Comodulation masking release using SAM tonal complex maskers: effects of modulation depth and signal position

The purpose of this investigation was to examine two stimulus parameters that were reasoned to be of importance to comodulation masking release (CMR). The first was the degree of fluctuation, or depth of modulation, in the masker bands, and the second was the temporal position of the signal with respect to the modulations of the masker. The investigation began by demonstrating the efficacy of sinusoidally amplitude-modulated (SAM) tonal complex maskers in eliciting CMR. "Nine-band" maskers, 650 ms in duration, were constructed by adding together nine SAM tones spaced at 100-Hz intervals from 300 to 1100 Hz. The rate of modulation for each SAM tone was 10 Hz, and the depth of modulation was 100%. Using such maskers, it was shown that when the on-frequency SAM tone had a modulation depth of 100%, the threshold for a 250-ms, 700-Hz tone improved monotonically as the modulation depths of the flanking SAM tones increased from 0% to 100%. When the on-frequency SAM tone had a modulation depth of 63%, some listeners performed optimally when the flanking SAM tones also exhibited a modulation depth of 63%, whereas others performed best when the flankers had modulation depths of 100%. With regard to signal position, a typical CMR effect was observed when the signal, consisting of a train of three 50-ms, 700-Hz tone bursts, was placed in the dips of the on-frequency masker. However, when the signal was placed at the peaks of the envelope, an increase in masking was observed for a comodulated masker.(ABSTRACT TRUNCATED AT 250 WORDS)     

Distortion product otoacoustic emission (2f1-f2) suppression in 3-month-old infants: evidence for postnatal maturation of human cochlear function?

The complete timeline for maturation of human cochlear function has not been defined. Distortion product otoacoustic emission (DPOAE)-based measures of cochlear function show non-adult-like responses from premature and term-born neonates at high f2 frequencies; however, older infants were not included in these studies. In the present experiment, previously collected DPOAE ipsilateral suppression data from premature neonates were combined with new data collected from adults, term-born neonates, and 3-month-old infants to further examine the time course for maturation of cochlear function. DPOAE suppression tuning curves (STC) and suppression growth patterns were measured in the three age groups at f2 = 6000 Hz, L1 = 65, L2 = 55 dB SPL, with an f2/f1 of 1.2. Results indicate that term-born neonates and 3-month-old infants have non-adult-like STC width, slope on the low-frequency flank, and tip features. However, the two infant groups are not significantly different from one another. Suppression growth patterns for low-frequency suppressor tones show a clear developmental progression. In general, the younger the infant, the more shallow and compressive the suppression growth for the lowest suppressor frequencies. These findings suggest a high-frequency postnatal immaturity in cochlear function as measured by DPOAE suppression. Results may have been influenced by noncochlear factors, such as middle-ear immaturity. These factors are reviewed and considered.     

Relationship of neural and otoacoustic emission thresholds during endocochlear potential development in the gerbil

Distortion product otoacoustic emissions and auditory brainstem responses (ABRs) were measured in neonatal gerbils at three ages: at 15-16 days after birth (dab), near the onset of hearing when the endocochlear potential (EP) is known to be still immature; at 22 dab, when the EP first reaches mature levels; and at 30 dab. Comparing individual 15-16 dab animals to the 22 dab group, ABR threshold changes were typically larger than those for cubic distortion tone (CDT, 2f1-f2) emission thresholds which were, in turn, larger than those for the simple difference tone (DT, f2-f1). In contrast, from 22 to 30 dab there were no important changes in CDT or DT emission thresholds. Observed threshold-change relationships were very similar to those found in differential diagnosis investigations, where the EP was experimentally decreased using a chronic furosemide application. Therefore, most of the change in cochlear function over the two week period studied could be attributed to the maturation of EP during the first week. Model calculations further show that relative changes in CDT and DT emission thresholds are compatible with a movement of the operating point of the cochlear amplifier toward its symmetrical "central" point as the EP reaches mature levels.     

Distortion-product otoacoustic emission suppression growth in normal and noise-exposed rabbits

This study investigated noise-induced changes in suppression growth (SG) of distortion product otoacoustic emissions (DPOAEs). Detailed measurements of SG were obtained in rabbits as a function of f2 frequencies at four primary-tone levels. SG measures were produced by using suppressor tones (STs) presented at two fixed distances from f2. The magnitude of suppression was calculated for each ST level and depicted as contour plots showing the amount of suppression as a function of the f2 frequency. At each f2, SG indices included slope, suppression threshold, and an estimate of the tip-to-tail value. All suppression measures were obtained before and after producing a cochlear dysfunction using a monaural exposure to a 2-h, 110-dB SPL octave-band noise centered at 2 kHz. The noise exposure produced varying amounts of cochlear damage as revealed by changes in DP-grams and auditory brainstem responses. However, average measures of SG slopes, suppression thresholds, and tip-to-tail values failed to mirror the mean DP-gram loss patterns. When suppression-based parameters were correlated with the amount of DPOAE loss, small but significant correlations were observed for some measures. Overall, the findings suggest that measures derived from DPOAE SG are limited in their ability to detect noise-induced cochlear damage.     

Interaction between adenosine triphosphate and mechanically induced modulation of electrically evoked otoacoustic emissions

It was shown previously that electrically evoked otoacoustic emissions (EEOAEs) can be amplitude modulated by low-frequency bias tones and enhanced by application of adenosine triphosphate (ATP) to scala media. These effects were attributed, respectively, to the mechano-electrical transduction (MET) channels and ATP-gated ion channels on outer hair cell (OHC) stereocilia, two conductance pathways that appear to be functionally independent and additive in their effects on ionic current through the OHC. In the experiments described here, the separate influences of ATP and MET channel bias on EEOAEs did not combine linearly. Modulated EEOAEs increased in amplitude, but lost modulation at the phase and frequency of the bias tone (except at very high sound levels) after application of ATP to scala media, even though spectral components at the modulation sideband frequencies were still present. Some sidebands underwent phase shifts after ATP. In EEOAEs modulated by tones at lower sound levels, substitution of the original phase values restored modulation to the waveform, which then resembled a linear summation of the separate effects of ATP and low-frequency bias. While the physiological meaning of this procedure is not clear, the result raises the possibility that a secondary effect of ATP on one or more nonlinear stages in the transduction process, which may have caused the phase shifts, obscured linear summation at lower sound levels. In addition, "acoustic enhancement" of the EEOAE may have introduced nonlinear interaction at higher levels of the bias tones.     

Suppression and (2f1-f2)-difference tones in a nonlinear cochlear preprocessing model with active feedback

The two nonlinear effects of two-tone suppression and of (2f1-f2)-difference tone creation are measured in a hardware model which consists of 90 sections containing nonlinear feedback loops. The basic data are the level and phase distributions along the 90 sections produced by single tones in the linear passive system which are almost identical to those produced in the nonlinear active system at high levels. Enhancement is created at medium and low input levels resulting in more strongly peaked level-place patterns. Two-tone suppression is, therefore, described as a "de-enhancement" which is produced by the gain reduction in the saturating nonlinearity of the feedback loop in consequence of increasing input levels (that of the feedback loop in consequence of increasing input levels (that of the suppressor as well!). Characteristics of suppression are given in normalized form. The creation of (2f1-f2)-difference tones is based on the same nonlinear effects. In each section, difference-tone wavelets are created which travel--changing level and phase thereby--to their characteristic place, where they add up to a vector sum corresponding to the audible difference tone. In case of cancellation, the vector sum has to be compensated by an additional tone of the same frequency and amount but opposite phase. Based on this strategy of (2f1-f2)-difference tone development, the relevant relations are measured on the model and averaged either in normalized graphs or in equations in order to offer the possibility to simulate the hardware model on the computer. Psychoacoustically measured cancellation data are compared with data measured using the model. The two data sets agree not only in general but also in many details indicating that the model describes cochlear nonlinear preprocessing to a useful approximation.     

Effects of flanking noise bands on the rate of growth of loudness of tones in normal and recruiting ears

Five subjects with unilateral cochlear hearing impairments and three normally hearing subjects made loudness matches between tones presented alternately to two ears, as a function of the intensity of the tone in the impaired ear (or the left ear of the normal subjects). The impaired ears showed recruitment; the rate of growth of loudness with increasing intensity was more rapid in the impaired ear than the normal ear. Presenting the tone in the impaired ear with two noise bands on either side of the tone frequency, at a fixed signal-to-noise ratio, did not abolish the recruitment. This suggests that recruitment is not caused by an abnormally rapid spread of excitation in the peripheral auditory system. At low signal-to-noise ratios, a continuous background noise reduced the loudness of the tone more than a noise gated with the tone, suggesting that the continuous noise induces adaptation to the tone. The noise had a greater effect on the loudness of the tone in normal ears than in impaired ears. It is possible that the loudness reduction of the tone in noise is mediated by suppression; suppression is weak or absent in impaired ears, and so the loudness reduction is smaller.     

Perception of across-frequency asynchrony and the role of cochlear delays

Cochlear filtering results in earlier responses to high than to low frequencies. This study examined potential perceptual correlates of cochlear delays by measuring the perception of relative timing between tones of different frequencies. A brief 250-Hz tone was combined with a brief 1-, 2-, 4-, or 6-kHz tone. Two experiments were performed, one involving subjective judgments of perceived synchrony, the other involving asynchrony detection and discrimination. The functions relating the proportion of "synchronous" responses to the delay between the tones were similar for all tone pairs. Perceived synchrony was maximal when the tones in a pair were gated synchronously. The perceived-synchrony function slopes were asymmetric, being steeper on the low-frequency-leading side. In the second experiment, asynchrony-detection thresholds were lower for low-frequency rather than for high-frequency leading pairs. In contrast with previous studies, but consistent with the first experiment, thresholds did not depend on frequency separation between the tones, perhaps because of the elimination of within-channel cues. The results of the two experiments were related quantitatively using a decision-theoretic model, and were found to be highly correlated. Overall the results suggest that frequency-dependent cochlear group delays are compensated for at higher processing stages, resulting in veridical perception of timing relationships across frequency.