Spontaneous Otoacoustic Emissions Research Articles

Conditions Underlying the Appearance of Spontaneous Otoacoustic Emissions in Mammals.

Across the wide range of land vertebrate species, spontaneous otoacoustic emissions (SOAE) are common, but not always found. The reasons for the differences between species of the various groups in their emission patterns are often not well understood, particularly within mammals. This review examines the question as to what determines in mammals whether SOAE are emitted or not, and suggests that the coupling between hair-cell regions diminishes when the space constant of frequency distribution becomes larger. The reduced coupling is assumed to result in a greater likelihood of SOAE being emitted.

Something in Our Ears Is Oscillating, but What? A Modeller's View of Efforts to Model Spontaneous Emissions.

When David Kemp discovered "spontaneous ear noise" in 1978, it opened up a whole new perspective on how the cochlea works. The continuous tonal sound emerging from most healthy human ears, now called spontaneous otoacoustic emissions or SOAEs, was an unmistakable sign that our hearing organ must be considered an active detector, not just a passive microphone, just as Thomas Gold had speculated some 30 years earlier. Clearly, something is oscillating as a byproduct of that sensitive inbuilt detector, but what exactly is it? Here, we give a chronological account of efforts to model SOAEs as some form of oscillator, and at intervals, we illustrate key concepts with numerical simulations. We find that after many decades there is still no consensus, and the debate extends to whether the oscillator is local, confined to discrete local sources on the basilar membrane, or global, in which an assembly of micro-mechanical elements and basilar membrane sections, coupled by inner ear fluid, interact over a wide region. It is also undecided whether the cochlear oscillator is best described in terms of the well-known Van der Pol oscillator or the less familiar Duffing or Hopf oscillators. We find that irregularities play a key role in generating the emissions. This paper is not a systematic review of SOAEs and their properties but more a historical survey of the way in which various oscillator configurations have been applied to modelling human ears. The conclusion is that the difference between the local and global approaches is not clear-cut, and they are probably not mutually exclusive concepts. Nevertheless, when one sees how closely human SOAEs can be matched to certain arrangements of oscillators, Gold would no doubt say we are on the right track.

Robust estimation of synchronized spontaneous otoacoustic emission via singular value decomposition and optimal shrinkage.

We investigate matrix signal processing techniques for estimating synchronized spontaneous otoacoustic emission (OAE) in noise. Responses to repeated clicks are first stored in a matrix, and singular value decomposition is either applied in the time domain or the frequency domain after constructing a Hankel matrix at every frequency. The singular values are subject to optimal shrinkage (OS) which maximizes the signal-to-noise ratio. Human OAE data were analyzed, and the Hankel matrix method outperforms the time-domain OS method in synchronized spontaneous otoacoustic emission estimation, but not in the estimation of transient-evoked otoacoustic emission. Reasons for the performance discrepancy are discussed.

Extended High Frequency Thresholds and Their Relationship to Distortion Product Otoacoustic Emissions, Hearing Acuity, Age, Gender, Presence of Spontaneous Otoacoustic Emissions, and Side of Measurement

Hearing is normally evaluated up to 8 kHz, even though testing can easily be performed at higher frequencies (up to 16 or 20 kHz). The range beyond 8 kHz is often referred to as the extended high frequency (EHF) range. This study aimed to explore the relationship between EHF hearing thresholds (HTs) and distortion product otoacoustic emissions (DPOAEs) in adult subjects. Also of interest were the effects of the presence of spontaneous otoacoustic emissions (SOAEs), gender, ear side, and age. The main finding was that DPOAEs, both within the standard frequency (SF) range (0.125–8 kHz) and the EHF range (10–16 kHz), decrease as thresholds deteriorate. For both ranges, DPOAEs and HTs depend on age, even for those with normal hearing, although EHFs seem to be especially affected by age. The presence of SOAEs was the only other factor that significantly influenced DPOAE level. For both DPOAEs and HTs, only minor and non-significant effects were related to gender and ear side. It was concluded that DPOAEs in the EHF range appear to be good predictors of EHF HTs. Moreover, since DPOAEs and HTs in the EHF range both correlate with age, these two measures may be suitable markers for incipient presbycusis.

Cochlear Function in Individuals with and without Spontaneous Otoacoustic Emissions.

This study investigated the status of spontaneous otoacoustic emissions (SOAEs) on cochlear function in a cohort of male/female participants with a wide age range. It examined whether there was a correlation between the presence of SOAEs and measurements of transient evoked otoacoustic emissions (TEOAEs), distortion product otoacoustic emissions (DPOAEs), SOAEs and extended high-frequency (EHF) hearing thresholds. 463 participants (222 male, 241 female; age range 20-59 years) with pure-tone thresholds ≤25 dB HL for octave frequencies of 500-8000 Hz were included in the study, divided into three age groups (20-29, 30-39, and 40-59 years). Evaluations included EHF (9000-16,000 Hz) hearing thresholds and TEOAE, DPOAE and SOAE measures. Multiple regression models showed that participants with SOAEs had larger expected amplitudes and signal-to-noise ratios (SNRs) for TEOAE and DPOAE responses than participants without SOAEs, holding gender and age variables constant. Spearman correlation tests identified deterioration in TEOAE and DPOAE amplitudes and SNRs, and EHF hearing thresholds with age in participants without SOAEs. Among participants with SOAEs, no significant decreases in TEOAE and DPOAE measures were shown in participants with older age. Nonetheless, as expected, EHF hearing thresholds did become worse with age, with or without SOAEs. Participants with identifiable SOAEs had greater TEOAE and DPOAE amplitudes and SNRs than participants without SOAEs. SOAEs appear to be a useful marker of cochlear health in adults.

Distortion Product Otoacoustic Emissions in Mice Above and Below the Eliciting Primaries.

Normal hearing is associated with cochlear nonlinearity. When two tones (f1 and f2) are presented, the intracochlear response contains additional components that can be recorded from the ear canal as distortion product otoacoustic emissions (DPOAEs). Although the most prominent intermodulation distortion component is at 2f1-f2, other cubic distortion products are also generated. Because these measurements are noninvasive, they are used in humans and in animal models to detect hearing loss. This study evaluated how loss of sensitivity affects DPOAEs with frequencies above and below the stimulating primaries, i.e., for upper sideband (USB) components like 2f2-f1 and for lower sideband (LSB) components like 2f1-f2. DPOAEs were recorded in several mouse mutants with varying degrees of hearing loss associated with structural changes to the tectorial membrane (TM), or with loss of outer hair cell (OHC) somatic electromotility due to lack of prestin or to the expression of a non-functional prestin. In mice with changes in sensitivity, magnitude reductions were observed for 2f1-f2 relative to controls with mice lacking prestin showing the greatest changes. In contrast, 2f2-f1 was minimally affected by reductions in cochlear gain due to changes in the TM or by the loss of OHC somatic electromotility. In addition, TM mutants with spontaneous otoacoustic emissions (SOAEs) generated larger responses than controls at 2f2-f1 when its frequency was similar to that for the SOAEs. Although cochlear pathologies appear to affect USB and LSB DPOAEs in different ways, both 2f1-f2 and 2f2-f1 reflect nonlinearities associated with the transducer channels. However, in mice, the component at 2f2-f1 does not appear to receive enhancement due to prestin's motor action.

ABO blood group and cochlear function: evidence from a large sample size study

Objective The present study investigated the effect of blood group on cochlear function in a large participant sample across different age groups. The study hypothesis was that participants with blood group O would show relatively reduced cochlear function as reflected in otoacoustic emission (OAE) measures. Design Data were collected from transient evoked otoacoustic emission (TEOAE), distortion product otoacoustic emission (DPOAE), DPOAE input/output (I/O) function, and spontaneous otoacoustic emission (SOAE) recordings. Study sample Four hundred and sixty-three normal hearing adults aged 20–59 years among the four ABO blood groups participated in the study. Results TEOAE and DPOAE amplitudes did not reveal significant differences for participants with blood group O compared with participants with non-O blood groups. No significant differences in I/O function categories were found among participants with different blood groups. SOAE prevalence was also not significantly different across blood groups. However, previously reported age and gender differences for OAE variables were confirmed. Conclusions Participants with blood group O were not found to have significantly reduced cochlear function, based on OAE measures. Results from the current study do not support the hypothesis that normal hearing individuals with different ABO blood groups differ in level of cochlear function.

Nonlinear dynamics of spontaneous otoacoustic emissions in the presence of internal noise in models of the inner ear

Spontaneous otoacoustic emissions (SOAEs) are sounds generated by the inner ear in the absence of an external stimulus and are a direct consequence of the inner ear nonlinearity that arises from hair cell electromechanics. SOAEs with similar spectral characteristics have been observed in both mammals and non-mammalian species such as lizards, suggesting a common mechanism in their generation, despite striking differences in inner ear physiology between mammals and non-mammals. In this work, a model based on coupled limit-cycle oscillators (Vilfan and Duke, Biophys. J, vol. 95, pp. 4622–4630, 2008) and a model based on standing waves (Bowling etal., Sci. Rep., vol. 11, pp. 1–14, 2021) are implemented to analyze their ability to predict key SOAE characteristics. As in experiments, the models predict discrete peaks in the SOAEs with quasi-periodic spacing. However, if noise is neglected, the SOAE peaks are unrealistically sharp. In this work, the effect of noise on SOAE spectral properties (e.g., bandwidth of SOAE peaks) and statistical properties (e.g., interpeak cross-correlations) is evaluated to assess whether either model is a better representation of SOAE generation. This is a key step in understanding the theoretical underpinnings of SOAE generation. [Research funded by NIH grant R01 DC016114.]

Frequency Shifts in a Local Oscillator Model for the Generation of Spontaneous Otoacoustic Emissions by the Lizard Ear

Introduction: In order to understand human hearing, it helps to understand how the ears of lower vertebrates, like, for instance, lizards, function. A key feature in common is that the ears of both humans and lizards emit faint, pure tones known as spontaneous otoacoustic emissions (SOAEs). More than four decades after their discovery, the mechanism underlying these emissions is still imperfectly understood, although it is known that they are important for improving the sensitivity and sharpness of hearing. In both humans and lizards, the frequencies of SOAEs change by a few percent when static pressure is applied to the tympanic membrane. For the human ear, this observation is normally explained by a so-called global oscillator model (such as with Shera’s coherent reflection model), in which the emissions result from standing waves, and external pressure changes the boundary conditions – the stiffness of the oval and round windows – which then has a global effect on the SOAE frequencies. Methods: Here we investigate how changing parameters of an earlier developed local oscillator model for the lizard ear can change the frequencies of the SOAEs. A major feature of the model is that each oscillator is coupled only to its immediate neighbours. The oscillators then cluster into groups of identical frequency, and each of these so-called frequency plateaus can be taken to represent an SOAE. Results: Even though the natural (unperturbed) frequencies of all the oscillators remain fixed, here we find for several model parameters that by slightly changing their value the frequency plateaus – the SOAEs – shift by a few percent. Plots of how these changes alter SOAE frequencies are given, and their magnitude corresponds well with observations of SOAE changes in lizards. Discussion: Investigation of the influence of the change of parameters in an earlier developed local oscillator model for the lizard ear shows that a local oscillator model can explain small SOAE frequency changes as well as a global oscillator model.

The effects of mechanically gated ion channels of the inner ear on thermal sensitivity of spontaneous otoacoustic emissions

In quiet environments, the inner ears of vertebrates can produce low intensity sounds that are detectable in the ear canal, termed spontaneous otoacoustic emissions (SOAEs). This background activity of the inner ear has been regarded as an epiphenomenon of the active processes performed by hair cells – the sensory receptors of the auditory system. Experimental measurements of SOAEs from the ears of tokay geckos reveal a linear increase of SOAE frequency with body temperature, with the emission at higher frequencies displaying greater thermal sensitivity. In this work, we elucidated the cellular mechanism underlying the thermal sensitivity of SOAE frequency using a mathematical model of hair cell’s transduction process performed by mechanically gated ion channels. We employed the previously proposed gating-spring model which described an individual ion channel by a two-state system, whose activation energy associated with channel gating depended on the level of temperature. Our results from numerical simulations revealed that a rise in temperature elicited an increase in the frequency of spontaneous oscillations displayed by a single hair cell. The magnitude of the frequency shift increased with the oscillation frequency following a quadratic polynomial, a characteristic observed in the physiological recordings of SOAEs from tokay geckos. The model further suggested that the intrinsic energy difference between the open and closed state of the ion channels greatly controlled the temperature sensitivity of a hair bundle.

Gender differences in contralateral suppression of spontaneous otoacoustic emissions in individuals with auditory neuropathy spectrum disorders.

The current study aims to evaluate gender variations in the efferent auditory system functioning in the ANSD population and compare them to normal hearing persons using contralateral suppression of spontaneous OAE (SOAEs). For the current study, a total of 54 ears were considered prospectively. 27 ears with normal hearing (NH) sensitivity were in group I, while 27 ears with ANSD were in group II. All of the subjects had a regular audiological examination followed by synchronised SOAE recordings with and without broadband noise (in the contralateral ear) at 50dB SPL and statistical analysis was carried out. Wilcoxon signed rank test revealed a significant decrease in SOAE amplitude with contralateral noise stimulation in NH, while no such trend was observed in ANSD. However, contralateral SOAE stimulation resulted in significant high-frequency shifts for both the ANSD and NH groups in the noise condition. In the comparison of gender effects on SOAE amplitude and frequency shifts using Mann-Whitney, no significant gender differences were seen. The lack of gender differences in SOAE suppression is explained based on global standing wave theory and local oscillator theory. Contralateral effect of noise on SOAE shows frequency shifts, but not amplitude shifts both of which are not affected by gender effects.

Medial olivocochlear reflex reliability: The effects of averaging and presence of synchronized spontaneous otoacoustic emissions.

The medial olivocochlear reflex (MOCR), usually assessed by the inhibition of transiently evoked otoacoustic emissions (TEOAEs) with contralateral noise, is a very small effect. In understanding the origin of the MOCR, it is crucial to obtain data of the highest accuracy, i.e., with a high signal-to-noise ratio (SNR), which in turn largely depends on the number of signal averages. This study investigates how the reliability of MOCR measures is affected by the number of averages. At the same time, the effect of the presence of synchronized spontaneous otoacoustic emissions (SSOAEs) is taken into account, as it is known that this factor significantly affects TEOAE amplitudes and SNRs. Each recording session consisted of two series of four measurements, allowing comparison of MOCR magnitude based on 250, 500, 750, and 1000 averages. Reliability was based on comparing the two series. The results show that, for a good quality MOCR measure (i.e., intraclass correlation above 0.9), the required number of averages is at least double that obtainable from a standard TEOAE test (i.e., 500 compared to 250). Ears without SSOAEs needed a higher number of averages to reach a correlation of 0.9 than ears with SSOAEs.

OD PĘCZKA KOMÓREK SŁUCHOWYCH DO BŁONY BĘBENKOWEJ

BackgroundAn earlier oscillator model for the generation of spontaneous otoacoustic emissions (SOAEs) from the lizard ear is extended with a connection of the oscillators to the basilar papilla, to make it possible that these SOAEs can be transported to the tympanic membrane, to be emitted.Material and methodsThe generators of spontaneous otoacoustic emissions are modelled as a one-dimensional array of Hopf-resonators. The resonators (or oscillators) are coupled to their neighbours, and to the basilar papilla. The papilla is modelled as a rigid structure, that is flexibly connected to its surroundings.ResultsFrequency spectra are given for different sets of coupling parameters, both for nearest neighbour coupling of the oscillators, and for coupling to the papilla, and also after the introduction of irregularities in the damping of the oscillators. Waterfall and density plots show clustering of the oscillators in frequency plateaus, and entrainment of a cluster of oscillators by an externally applied sinusoidal force. All these model outcomes correspond with characteristics of SOAEs emitted by real lizard ears.ConclusionsThe present model is a useful extension of an earlier model. Because its characteristics differ from that of a model that is used to describe the generation of SOAEs by mammalian ears, it revives the discussion whether different models are needed for SOAE generation in different animal species.

Carbamazepine induces upward frequency shifts of spontaneous otoacoustic emissions

In three subjects, we measured spontaneous otoacoustic emissions (SOAEs) when they were using carbamazepine (CBZ), and compared this to the SOAE measurement when they were not using CBZ. We observed 14 SOAEs showing a consistent upward shift of center frequency, related to CBZ intake. On average, the magnitude of the frequency shift increased with increasing frequency. The magnitude of the shift was 30–104 Hz, at frequencies ranging from 1.3 to 2.3 kHz, corresponding to a shift between 2.3 and 4.5%. Compared to other causes and manipulations known to change SOAE frequency, these shifts are relatively large. The underlying mechanism is most likely an increased stiffness of the cochlear partition. This would also explain the downward pitch shift due to CBZ, which has been reported by subjects with absolute pitch.

Effects of Adding Noise to Circuit Model and Numerical Model of Spontaneous Otoacoustic Emissions

Spontaneous otoacoustic emissions have long been modeled using self-excited, nonlinear oscillators. The van der Pol oscillator is a common choice, as many of its properties reflect those of SOAEs: both can begin oscillation in the absence of a direct stimulus, both have narrow frequency bands, and both become stable over time, to name a few. Yet such an idealized equation cannot have a one-to-one correspondence with SOAEs in all factors. Many previously used mathematical and circuit models lack the addition of noise to more accurately show how real world SOAEs operate in an organism’s ear, where noise from the environment is almost entirely unavoidable. The inclusion of uniformly distributed noise in both numerical and circuit models of the van der Pol oscillator was studied to determine whether these models can still accurately explicate SOAEs when modified to be more realistic. In both cases, both models retained the attributes of real world SOAEs despite the addition of noise, allowing them to serve as more useful and accurate models of the phenomenon.

Relationship between irregularities in spontaneous otoacoustic emissions suppression and psychophysical tuning curves.

The suppression of spontaneous otoacoustic emissions (SOAEs) allows the objective evaluation of cochlear frequency selectivity by determining the suppression tuning curve (STC). Interestingly, some STCs have additional sidelobes at the high frequency flank, which are thought to result from interaction between the probe tone and the cochlear standing wave corresponding to the SOAE being suppressed. Sidelobes are often in regions of other neighboring SOAEs but can also occur in the absence of any other SOAE. The aim of this study was to compare STCs and psychoacoustic tuning curves (PTCs). Therefore, STCs and PTCs were measured in: (1) subjects in which the STC had a sidelobe, and (2) subjects without STC sidelobes. Additionally, PTCs were measured in subjects without SOAEs. Across participant groups, the quality factor Q10dB of the PTCs was similar, independently from whether SOAEs were present or absent. Thus, the presence of an SOAE does not provide enhanced frequency selectivity at the emission frequency. Moreover, both PTC and STC show irregularities, but these are not related in a straightforward way. This suggests that different mechanisms cause these irregularities.

Whistling While it Works: Spontaneous Otoacoustic Emissions and the Cochlear Amplifier.

Perhaps the most striking evidence for active processes operating within the inner ears of mammals and non-mammals alike is their ability to spontaneously produce sound. Predicted by Thomas Gold in 1948, some 30 years prior to their discovery, the narrow-band sounds now known as spontaneous otoacoustic emissions (SOAEs) remain incompletely understood, their origins controversial. Without a single equation in the main text, we review the essential concepts underlying the "local-" and "global-oscillator" frameworks for understanding SOAE generation. Comparing their key assumptions and predictions, we relate the two frameworks to unresolved questions about the biophysical mechanisms of cochlear amplification.

Spontaneous otoacoustic emissions are biomarkers for mice with tectorial membrane defects

Cochlear function depends on the operation of a coupled feedback loop, incorporating outer hair cells (OHCs), and structured to assure that inner hair cells (IHCs) convey frequency specific acoustic information to the brain, even at very low sound levels. Although our knowledge of OHC function and its contribution to cochlear amplification has expanded, the importance of the tectorial membrane (TM) to the processing of mechanical inputs has not been fully elucidated. In addition, there are a surprising number of genetic mutations that affect TM structure and that produce hearing loss in humans. By synthesizing old and new results obtained on several mouse mutants, we learned that animals with abnormal TMs are prone to generate spontaneous otoacoustic emissions (SOAE), which are uncommon in most wildtype laboratory animals. Because SOAEs are not produced in TM mutants or in humans when threshold shifts exceed approximately 25 dB, some degree of cochlear amplification is required. However, amplification by itself is not sufficient because normal mice are rarely spontaneous emitters. Since SOAEs reflect active cochlear operation, TM mutants are valuable for studying the oscillatory nature of the amplification process and the structures associated with its stabilization. Inasmuch as the mouse models were selected to mirror human auditory disorders, using SOAEs as a noninvasive clinical tool may assist the classification of individuals with genetic defects that influence the active mechanisms responsible for sensitivity and frequency selectivity, the hallmarks of mammalian hearing.

Effect of Musical Experience on Cochlear Frequency Resolution: An Estimation of PTCs, DLF and SOAEs.

Background: There is a lot of debate on whether musical experience/training can affect cochlear filter characteristics and thereby enhance frequency resolution skills in musicians (Ms). Thus, the objective of the study was to compare between Ms and non-musicians (NMs) and correlate the frequency resolution skills with years of musical experience, the difference limen for frequency (DLF), and the Q10 values of psychophysical tuning curves (PTCs) measured at 2 characteristic frequencies (CFs), 1000 and 4000 Hz. The secondary objective was to find out whether spontaneous otoacoustic emissions (SOAEs) can be recorded among Ms more frequently than among NMs. Methods:Thirty-six listeners with normal hearing participated in the study. They were 18 Ms with a minimum of 8 years of musical experience, and 18 NMs. Forward-masked PTCs and DLF tests were assessed for each listener in the right ear at 2 CFs, 1 and 4 kHz, and SOAEs were measured using standard protocol. Results:The results obtained indicated that values of the psychophysical measures, DLF and Q10, were better among the Ms group. Statistically significant differences were seen when measurements were done at 1 KHz and 4 KHz. SOAEs were recorded in a higher number of Ms than NMs. Musical experience had a moderate positive correlation with PTC Q10 values at both 1 and 4 kHz, but not DLF values. Conclusion:This study indicates that musical experience enhances peripheral filtering, and thereby betters cochlear frequency selectivity in Ms.

Synchronization and chaos in systems of coupled inner-ear hair cells

Hair cells of the auditory and vestibular systems display astonishing sensitivity, frequency selectivity, and temporal resolution to external signals. These specialized cells utilize an internal active amplifier to achieve highly sensitive mechanical detection. One of the manifestations of this active process is the occurrence of spontaneous limit-cycle motion of the hair cell bundle. As hair bundles under in vivo conditions are typically coupled to each other by overlying structures, we explore the role of this coupling on the dynamics of the system, using a combination of theoretical and experimental approaches. Our numerical model suggests that the presence of chaotic dynamics in the response of individual bundles enhances their ability to synchronize when coupled, resulting in significant improvement in the system's ability to detect weak signals. This synchronization persists even for a large frequency dispersion and a large number of oscillators comprising the system. Further, the amplitude and coherence of the active motion is not reduced upon increasing the number of oscillators. Using artificial membranes, we impose mechanical coupling on groups of live and functional hair bundles, selected from in vitro preparations of the sensory epithelium, allowing us to explore the role of coupling experimentally. Consistent with the numerical simulations of the chaotic system, synchronization occurs even for large frequency dispersion and a large number of hair cells. Further, the amplitude and coherence of the spontaneous oscillations are independent of the number of hair cells in the network. We therefore propose that hair cells utilize their chaotic dynamics to stabilize the synchronized state and avoid the amplitude death regime, resulting in collective coherent motion that could play a role in generating spontaneous otoacoustic emissions and an enhanced ability to detect weak signals.