A cochlea with three parts? Evidence from otoacoustic emission phase in humans.

The apical and basal regions of the cochlea appear functionally distinct. In humans, compelling evidence for an apical-basal transition derives from the phase of otoacoustic emissions (OAEs), whose frequency dependence differs at low and high frequencies. Although OAEs arising from the two major source mechanisms (distortion and reflection) both support the existence of an apical-basal transition-as identified via a prominent bend (or "break") in OAE phase slope-the two OAE types disagree about its precise location along the cochlea. Whereas distortion OAEs at frequency 2f1-f2 suggest that the apical-basal transition occurs near the 2.5 kHz place, reflection OAEs locate the transition closer to 1 kHz. To address this discrepancy, distortion and reflection OAEs were measured and analyzed in 20 young human adults from 0.25-8 kHz and at eight primary-frequency ratios f2/f1 in the range 1-1.5. Break frequencies and OAE phase-gradient delays were estimated by fitting segmented linear models to the unwrapped phase. When distortion- and reflection-OAE phase are considered as functions of ln f2-that is, as linear functions of the location of their putative site of generation within the cochlea-the analysis identifies not just two but three main cochlear segments, meeting at transition frequencies of approximately 0.9 and 2.6 kHz, whose locations are largely independent both of primary-frequency ratio and emission type. A simple model incorporating an abrupt transition from wave- to place-fixed behavior near the middle of the cochlea accounts for key features of distortion-OAE phase.

Variable-rate frequency sweeps and their application to the measurement of otoacoustic emissions.

Swept tones allow the efficient measurement of otoacoustic emissions (OAEs) with fine frequency resolution. Although previous studies have explored the influence of different sweep parameters on the measured OAE, none have directly considered their effects on the measurement noise floor. The present study demonstrates that parameters such as sweep type (e.g., linear or logarithmic), sweep rate, and analysis bandwidth affect the measurement noise and can be manipulated to control the noise floor in individual subjects. Although responses to discrete-tone stimuli can be averaged until the uncertainty of the measurement meets a specified criterion at each frequency, linear or logarithmic sweeps offer no such flexibility. However, measurements of the power spectral density of the ambient noise can be used to construct variable-rate sweeps that yield a prescribed (e.g., constant) noise floor across frequency; in effect, they implement a form of frequency-dependent averaging. The use of noise-compensating frequency sweeps is illustrated by the measurement of distortion-product OAEs at low frequencies, where the ear-canal noise is known to vary significantly.

Stimulus ratio dependence of low-frequency distortion-product otoacoustic emissions in humans.

Active amplifiers within the cochlea generate, as a by-product of their function, distortion-product otoacoustic emissions (DPOAEs) in response to specific two-tone stimuli. Focus has been on invoking emissions in a mid-frequency range from ∼0.5 to 4 kHz. The present study investigates stimulus parameters of the DPOAE at 2f1-f2 frequencies below 0.5 kHz. Eighteen out of 21 young human adults screened had audiometrically normal hearing for inclusion in the experiment. DPOAEs were measured with pure-tone stimuli in four configurations: f2 fixed around 2.13 kHz, f2 fixed around 0.53 kHz, 2f1-f2 fixed at 1.23 kHz and 0.25 kHz. Eight stimulus ratios, f2/f1, and three stimulus sound pressure levels, L1/L2, were measured in each configuration. Trends in ratio-magnitude responses for the mid-frequency DPOAE agree with those reported in previous literature. DPOAEs are not limited to distortion frequencies >0.5 kHz, but the stimulus ratio invoking the largest DPOAE in the mid-frequency range does not do so in the low-frequency range. Guiding the ratio according to the equivalent rectangular bandwidth of auditory filters maintains the DPOAE level.

Extended low-frequency phase of the distortion-product otoacoustic emission in human newborns.

At constant f 2 / f 1 ratios, the phase of the nonlinear distortion component of the 2 f 1 - f 2 distortion-product otoacoustic emission (DPOAE) has a steep low-frequency segment and a flat high-frequency segment in adults and newborns. In adults, recent work found that a third segment characterizes the phase at even lower frequencies. The present study tests whether the same is true of the newborn DPOAE phase. Newborn and adult phase curves are generally similar. However, as previously reported, phase-gradient delays at mid frequencies (the region of steepest phase slope) are 50% longer in newborns.

Probing Apical-Basal Differences in the Human Cochlea Using Distortion-Product Otoacoustic Emission Phase.

Distortion-product otoacoustic emission (DPOAE) phase is shaped by interaction between the evoking stimulus waves. Near-invariant at high frequencies, DPOAE phase-vs-frequency functions measured at fixed ratios bend into sloping functions at low frequencies. The different phase behaviors observed suggest that the mechanics underlying the generation of OAEs differ in the halves of the cochlea. To map out the phenomenological extent of low-to-mid frequency phase bends, this study recorded DPOAE responses from 20 normal-hearing human adult ears for a wide range of stimulus frequencies, f1 and f2, where f2 frequency sweeps from 0.25 to 8 kHz, and the f2/ f1 ratio varies from 1.05 to 1.49. Our preliminary results show two transitions in the phase slopes. One near 2.6 kHz in agreement with the literature, and another of opposite polarity near 0.75 kHz which has not been reported before. We find that the f2 frequencies marking these defining phase features are invariant with stimulus ratio. Even as the underlying mechanics remain unknown, the invariance opens the door for DPOAE phase to reliably characterize apical-basal differences across age groups and species.

Distortion-Product Otoacoustic Emission Measured Below 300 Hz in Normal-Hearing Human Subjects.

Physiological noise levels in the human ear canal often exceed naturally low levels of otoacoustic emissions (OAEs) near the threshold of hearing. Low-frequency noise, and electronic filtering to cope with it, has effectively limited the study of OAE to frequencies above about 500 Hz. Presently, a custom-built low-frequency acoustic probe was put to use in 21 normal-hearing human subjects (of 34 recruited). Distortion-product otoacoustic emission (DPOAE) was measured in the enclosed ear canal volume as the response to two simultaneously presented tones with frequencies f 1 and f 2. The stimulus-frequency ratio f 2/f 1 was varied systematically to find the "optimal" ratio evoking the largest level at 2 f 1-f 2 frequencies 87.9, 176, and 264 Hz. No reference data exist in this frequency region. Results show that DPOAE exists down to at least 87.9 Hz, maintaining the bell-shaped dependence on the f 2/f 1 ratio known from higher frequencies. Toward low frequencies, however, the bell broadens and the optimal ratio increases proportionally to the bandwidth of an auditory filter as defined by the equivalent rectangular bandwidth. The DPOAE phase rotates monotonously as a function of the stimulus ratio, and its slope trend supports the notion of a lack of scaling symmetry in the apex of the cochlea.