Temperature dependence of frog spontaneous otoacoustic emissions

Spontaneous otoacoustic emissions are recorded from frog ears at various body temperatures. Frequency spectra and amplitude distributions of emissions are determined. Both power and frequency of emission signals show a strong temperature dependence.     

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.     

Prevalence of spontaneous otoacoustic emissions in neonates.

The prevalence of spontaneous otoacoustic emissions (SOAEs) was measured in a group of 100 neonates and in a group of 50 normal-hearing young adults. The prevalence of SOAEs in the adult group (0.62) is at the high end of the range of prevalences reported in other surveys of adult SOAEs based on measurements using similar microphones. The prevalence of SOAEs in neonates (0.64) is not significantly different from that in adults. The various tendencies that have been found to be significant in the pooled results of other surveys are also evident in our adult group: more SOAEs in right ears, a higher prevalence of SOAEs in females, and a dependence between ears for the occurrence of SOAEs. The above-mentioned tendencies are also significant in the infant data. The major differences between the infant and adult results are the predominant SOAE frequency range and the average levels of SOAEs. The majority of adult SOAEs are between 1.0 and 2.0 kHz, whereas the majority of neonatal SOAEs are between 2.5 and 5.0 kHz. The average SOAE level is -2.6 dB SPL for adults and 8.5 dB SPL for infants.     

Mammalian spontaneous otoacoustic emissions are amplitude-stabilized cochlear standing waves

Mammalian spontaneous otoacoustic emissions (SOAEs) have been suggested to arise by three different mechanisms. The local-oscillator model, dating back to the work of Thomas Gold, supposes that SOAEs arise through the local, autonomous oscillation of some cellular constituent of the organ of Corti (e.g., the "active process" underlying the cochlear amplifier). Two other models, by contrast, both suppose that SOAEs are a global collective phenomenon--cochlear standing waves created by multiple internal reflection--but differ on the nature of the proposed power source: Whereas the "passive" standing-wave model supposes that SOAEs are biological noise, passively amplified by cochlear standing-wave resonances acting as narrow-band nonlinear filters, the "active" standing-wave model supposes that standing-wave amplitudes are actively maintained by coherent wave amplification within the cochlea. Quantitative tests of key predictions that distinguish the local-oscillator and global standing-wave models are presented and shown to support the global standing-wave model. In addition to predicting the existence of multiple emissions with a characteristic minimum frequency spacing, the global standing-wave model accurately predicts the mean value of this spacing, its standard deviation, and its power-law dependence on SOAE frequency. Furthermore, the global standing-wave model accounts for the magnitude, sign, and frequency dependence of changes in SOAE frequency that result from modulations in middle-ear stiffness. Although some of these SOAE characteristics may be replicable through artful ad hoc adjustment of local-oscillator models, they all arise quite naturally in the standing-wave framework. Finally, the statistics of SOAE time waveforms demonstrate that SOAEs are coherent, amplitude-stabilized signals, as predicted by the active standing-wave model. Taken together, the results imply that SOAEs are amplitude-stabilized standing waves produced by the cochlea acting as a biological, hydromechanical analog of a laser oscillator. Contrary to recent claims, spontaneous emission of sound from the ear does not require the autonomous mechanical oscillation of its cellular constituents.     

The behavior of spontaneous otoacoustic emissions during and after postural changes

Spontaneous otoacoustic emissions (SOAEs) were studied in humans during and after postural changes. The subjects were tilted from upright to a recumbent position (head down 30 degrees) and upright again in a 6-min period. The SOAEs were recorded continuously and analyzed off-line. The tilting caused a change in the SOAE spectrum for all subjects. Frequency shifts of 10 Hz, together with changes of amplitude (5 dB) and width (5 Hz), were typically observed. However, these changes were observed in both directions (including the appearance and disappearance of emission peaks). The most substantial changes occurred in the frequency region below 2 kHz. An increase of the intracranial pressure, and consequently of the intracochlear fluid pressure, is thought to result in an increased stiffness of the cochlear windows, which is probably mainly responsible for the SOAE changes observed after the downward turn. The time for the spectrum to regain stability after a postural change differed between the two maneuvers: 1 min for the downward and less than 10 s for the upward turn.     

Suppression tuning curves for spontaneous otoacoustic emissions in infants and adults

Suppression tuning curves (STCs) for spontaneous otoacoustic emissions (SOAEs) were longitudinally obtained in seven infants (at 3 weeks, 2 months, and 3, 4, or 6 months of age) and in five adults. Excellent reproducibility was obtained for adult STCs. Reproducibility for infant STCs was poorer, but the curves at each age resemble those of adults both qualitatively and quantitatively as measured by slopes of the lower and upper segments of the STCs and by Q10's. Evidence from two subjects suggests that developmental changes in the fine tuning of the system may occur postnatally. Results are discussed with respect to the development of cochlear frequency selectivity.     

Are spontaneous otoacoustic emissions generated by self-sustained cochlear oscillators?

Theoretical analyses supporting the assumption that spontaneous otoacoustic emissions (SOAEs) can be described as self-sustained oscillations (requiring a power source) are reviewed and extended. Spectral and statistical properties of spontaneous otoacoustic emissions are examined and shown to be consistent with this assumption. Several alternative models of spontaneous emissions (noise-driven saturating memoryless nonlinearity, noise-driven nonlinear-stiffness oscillator) are examined. Although some of these models are able to produce the types of statistical distributions of amplitude and displacement similar to those observed in the experimental data, this similarity is destroyed upon narrow-band filtering.     

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.     

Modification of spontaneous and evoked otoacoustic emissions and associated psychoacoustic microstructure by aspirin consumption

The discovery that aspirin consumption can abolish spontaneous otoacoustic emissions [D. McFadden and H.S. Plattsmier, J. Acoust. Soc. Am. 76, 443-448 (1984)] provides a technique for further exploring the relation between otoacoustic emissions (spontaneous and evoked) and psychoacoustic threshold microstructure. Spontaneous emissions, delayed evoked emissions, synchronous evoked emissions, and threshold microstructure in four subjects were monitored before, during, and after consumption of 3.9 g of aspirin per day (three 325-mg tablets every 6 h) for 3 or 4 days. The changes in spontaneous emissions are consistent with the findings of McFadden and Plattsmier except that one spontaneous emission appeared to plateau at a reduced level above the noise floor during the last day and a half of the 3-day period of aspirin consumption. Evoked emissions and threshold microstructure were also reduced by aspirin consumption but persisted longer and recovered sooner. In most instances, the initial change in threshold microstructure was a trend to increased sensitivity (reduced thresholds), with a greater increase near threshold maxima than at threshold minima. Further reduction in the levels of the evoked emissions was accompanied by the eventual decrease in sensitivity (elevation of all thresholds).     

A parametric model of the spectral periodicity of stimulus frequency otoacoustic emissions

A model for estimating the spectral period of stimulus frequency otoacoustic emissions (SFOAEs) is presented. The model characterizes the frequency spectrum of an SFOAE in terms of four parameters which can be directly related to cochlear mechanical quantities featuring in the theory of SFOAE generation proposed by Zweig and Shera [J. Acoust. Soc. Am. 98, 2018-2047 (1995)]. The results of applying the parametric model to SFOAEs generated by cochlear models suggest that it gives a sensitive measure of spectral period. It is concluded that the parametric model may be a useful tool for detecting small changes in cochlear function using SFOAE measurements.     

Partial dissociation of spontaneous otoacoustic emissions and distortion products during aspirin use in humans

Otoacoustic emissions (OAEs) of two types--spontaneous and evoked distortion products--were studied before, during, and following a period of aspirin use. As previously reported, aspirin consumption uniformly reduced the spontaneous OAEs (SOAEs) to unmeasurable or extremely low levels. Aspirin consumption also reduced the amplitude of the evoked distortion products (EDPs) but did not eliminate them entirely. The amplitude of the EDP and its change with aspirin consumption were related to both the proximity of the EDP to the frequency of the SOAE and to the level of the primaries producing the EDP. At low primary levels, even with the SOAE absent (due to aspirin consumption, or suppression), EDPs near the SOAE frequency were 10-20 dB higher than when they were 100 Hz away from the SOAE frequency.     

Swept-tone transient-evoked otoacoustic emissions

Transient-evoked otoacoustic emissions (TEOAE) are responses generated within the inner ear in response to acoustic stimuli and are indicative of normal cochlear function. They are commonly acquired by averaging post-stimulus acoustic responses recorded near the eardrum in response to brief stimuli such as clicks or tone pips. In this study a new long duration stimulus consisting of a frequency swept tone is introduced for the acquisition of TEOAEs. Like stimulus frequency generated OAEs, swept-tone responses contain embedded OAEs. With swept-tone analysis, OAEs can be recovered by convolving it with a time reversed swept-tone signal resulting in time-compression. In addition, higher order nonlinear OAE responses were removed from the linear TEOAE. The results show comparable phase and time-frequency properties between the click and swept-tone evoked OAEs. Swept-tone acquisition of TEOAEs has beneficial noise properties, improving the signal to noise ratio by 6 dB compared to click evoked responses thus offering testing time savings. Additionally, swept-tone analysis removed synchronized spontaneous OAE activity from the recordings of subjects exhibiting such responses in conventional click TEOAEs. Since swept-tone stimulus consists of a single frequency component at any instantaneous moment, its analysis also provides for direct comparison with stimulus-frequency OAEs and click evoked OAEs.     

Modeling the effect of suppression on the periodicity of stimulus frequency otoacoustic emissions

The distributed roughness theory of the origins of spectral periodicity in stimulus frequency otoacoustic emissions (SFOAEs) predicts that the spectral period will be altered by suppression of the traveling wave (TW) [Zweig and Shera, J. Acoust. Soc. Am. 98, 2018-2047 (1995)]. In order to investigate this effect in more detail, simulations of the variation of the spectral period under conditions of self-suppression and two-tone suppression are obtained from nonlinear cochlear models based on this theory. The results show that during self-suppression the spectral period is increased, while during high-side two-tone suppression, the period is reduced, indicating that the detailed pattern of disruption of the cochlear amplifier must be examined if the nonlinear behavior of SFOAEs is to be understood. The model results suggest that the SFOAE spectral period may be sensitive to changes in the state of the cochlear amplifier. A companion paper [Lineton and Lutman, J. Acoust. Soc. Am. 114, 871-882 (2003)] presents experimental data which are compared with the results of the above models with a view to testing the underlying theory of Zweig and Shera.     

A review of otoacoustic emissions

Otoacoustic emissions measured in the external ear canal describe responses that the cochlea generates in the form of acoustic energy. For the convenience of discussing their principal features, emitted responses can be classified into several categories according to the type of stimulation used to evoke them. On this basis, four distinct but interrelated classes can be distinguished including spontaneous, transiently evoked, stimulus-frequency, and distortion-product otoacoustic emissions. The present review details the findings that have been described for each emission type according to this classification schema. Additionally, the known features of emitted responses are discussed for both normally hearing and hearing-impaired humans and experimental animals, and with respect to their potential clinical applications. The findings reviewed here clearly indicate that future studies of otoacoustic emissions will significantly increase our understanding of the basic mechanisms of cochlear function while, at the same time, provide a new and important clinical tool.     

Synchronization of spontaneous otoacoustic emissions to a 2f1-f2 distortion product

Synchronization of spontaneous otoacoustic emissions to a cubic distortion frequency fs = 2f1-f2 has been studied. Stimulus, consisting of two primary tones at frequency f1 and f2, could easily be filtered out of the microphone signal. This enabled us to monitor emission phase with respect to synchronization frequency fs, by recording zero-crossing moments of the microphone signal. When primaries were sufficiently loud (typically 30 dB SPL), phase fluctuated around a constant value: The emission was constantly synchronized to fs. Lowering primary levels (to typically 20 dB SPL) resulted in 2 pi-phase jumps at random moments: The emission occasionally slipped out of synchronization, trying to maintain its own natural frequency f0. This behavior can be described as synchronization of an oscillator (frequency f0) to a sinusoidal force (frequency fs) in the presence of noise.     

Effects of mobile phone exposure on time frequency fine structure of transiently evoked otoacoustic emissions

Mobile phones have become very commonly used worldwide within a short period of time. To date there is only limited knowledge about interaction between electromagnetic fields (EMFs) emitted by mobile phones and the auditory function. Moreover, there is widespread concern that there may be potential for harm. The aim of this study was to assess potential subtle changes in cochlear function by measuring the temporal and spectral fine structure of transiently evoked otoacoustic emissions (TEOAE) in normal hearing subjects after exposure to EMFs emitted by Global System for Mobile Communication (GSM) mobile phones. TEOAEs were recorded in 27 healthy young adults before and after 10 min of real or sham exposure in a double-blind design. TEOAE data were analyzed both globally (broadband analysis) and using the Wavelet Transform (analysis of the time-frequency fine structure). The broadband analysis revealed no significant effect on TEOAEs related to exposure, confirming results of previous studies; in addition, no significant change was detected in the analysis of the elementary wavelet components, suggesting that the temporal and spectral fine structure of TEOAEs is not affected by 10 min exposure to low-intensity EMFs emitted by GSM mobile phones.     

The effect of suppression on the periodicity of stimulus frequency otoacoustic emissions: experimental data

In a companion paper [Lineton and Lutman, J. Acoust. Soc. Am. 114, 859-870 (2003)], changes in the spectral period of stimulus frequency otoacoustic emissions (SFOAEs) during self-suppression and two-tone suppression were simulated using a nonlinear cochlear model based on the distributed roughness theory of otoacoustic emission generation [Zweig and Shera, J. Acoust. Soc. Am. 98, 2018-2047 (1995)1. The current paper presents the results of an experimental investigation of SFOAE suppression obtained from 20 human subjects. It was found that, in most subjects, the spectral period increased during self-suppression, but reduced during high-side two-tone suppression. This pattern of results is in close agreement with the predictions of the cochlear model, and therefore strongly supports the distributed roughness theory of Zweig and Shera. In addition, the results suggest that the SFOAE spectral period is sensitive to changes in the state of the cochlear amplifier.     

Tone-burst-evoked otoacoustic emissions from normal-hearing subjects

Tone-burst-evoked otoacoustic emissions were measured as a function of tone-burst sound pressure level and frequency in normally hearing ears. Although the spectral and temporal properties varied across individual ears, there was a close correspondence between stimulus and response spectra. Both the spectral and latency characteristics of tone-burst-evoked emissions are consistent with the hypothesis that they are generated at sites along the cochlear partition corresponding to their frequency.     

Modeling synchronization and suppression of spontaneous otoacoustic emissions using Van der Pol oscillators: effects of aspirin administration

Many of the aspects of the interaction of spontaneous otoacoustic emissions with external tones (suppression and synchronization) can be qualitatively simulated by the behavior of a single driven Van der Pol oscillator. Analytical and numerical investigations of a model of spontaneous otoacoustic emissions based on such an oscillator (with appropriate parametric changes in the nonlinear and negative damping components) lead to predictions of the nature of the changes in suppression and synchronization (frequency-locking) tuning curves when the levels of spontaneous otoacoustic emissions are modified. Observations of the suppression and synchronization of spontaneous otoacoustic emissions by external tones of different frequencies and levels were obtained while the levels of spontaneous emissions were altered by aspirin administration. Modeling an emission as a single Van der Pol oscillator qualitatively accounts for: (1) the reduction of the level of an external tone required to suppress the emission by a decibel amount equivalent to the level reduction induced by aspirin administration; (2) the broadening of the frequency-locking tuning curve of an emission whose level is reduced; and (3) the pulling of the emission frequency by an external tone. It does not account for: (1) the observed asymmetry in the slopes of the external-tone suppression curves (more gradual for frequencies of the suppressor tone higher, rather than lower, than that of the emission); and (2) the frequency pushing of the emission by an external tone.