Basilar membrane vibration is not involved in the reverse propagation of otoacoustic emissions.
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ABSTRACT:
SUBMITTER: He W
PROVIDER: S-EPMC3660718 | biostudies-other | 2013
REPOSITORIES: biostudies-other
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Project description:Sounds originating from within the inner ear, known as otoacoustic emissions (OAEs), are widely exploited in clinical practice but the mechanisms underlying their generation are not entirely clear. Here we present simulation results and theoretical considerations based on a hydrodynamic model of the human inner ear. Simulations show that, if the cochlear amplifier (CA) gain is a smooth function of position within the active cochlea, filtering performed by a middle ear with an irregular, i.e., nonsmooth, forward transfer function suffices to produce irregular and long-lasting residual oscillations of cochlear basilar membrane (BM) at selected frequencies. Feeding back to the middle ear through hydrodynamic coupling afforded by the cochlear fluid, these oscillations are detected as transient evoked OAEs in the ear canal. If, in addition, the CA gain profile is affected by irregularities, residual BM oscillations are even more irregular and tend to evolve towards self-sustaining oscillations at the loci of gain irregularities. Correspondingly, the spectrum of transient evoked OAEs exhibits sharp peaks. If both the CA gain and the middle-ear forward transfer function are smooth, residual BM oscillations have regular waveforms and extinguish rapidly. In this case no emissions are produced. Finally, and paradoxically albeit consistent with observations, simulating localized damage to the CA results in self-sustaining BM oscillations at the characteristic frequencies (CFs) of the sites adjacent to the damage region, accompanied by generation of spontaneous OAEs. Under these conditions, stimulus-frequency OAEs, with typical modulation patterns, are also observed for inputs near hearing threshold. This approach can be exploited to provide novel diagnostic tools and a better understanding of key phenomena relevant for hearing science.
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Project description:Sound is detected and converted into electrical signals within the ear. The cochlea not only acts as a passive detector of sound, however, but can also produce tones itself. These otoacoustic emissions are a striking manifestation of the cochlea's mechanical active process. A controversy remains of how these mechanical signals propagate back to the middle ear, from which they are emitted as sound. Here, we combine theoretical and experimental studies to show that mechanical signals can be transmitted by waves on Reissner's membrane, an elastic structure within the cochlea. We develop a theory for wave propagation on Reissner's membrane and its role in otoacoustic emissions. Employing a scanning laser interferometer, we measure traveling waves on Reissner's membrane in the gerbil, guinea pig, and chinchilla. The results are in accord with the theory and thus support a role for Reissner's membrane in otoacoustic emissions.
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Project description:It has previously not been possible to measure eardrum vibration of human subjects in the region of auditory threshold. It is proposed that such measurements should provide information about the status of the mechanical amplifier in the cochlea. It is this amplifier that is responsible for our extraordinary hearing sensitivity. Here, we present results from a laser Doppler vibrometer that we designed to noninvasively probe cochlear mechanics near auditory threshold. This device enables picometer-sized vibration measurements of the human eardrum in vivo. With this sensitivity, we found the eardrum frequency response to be linear down to at least a 20-dB sound pressure level (SPL). Nonlinear cochlear amplification was evaluated with the cubic distortion product of the otoacoustic emissions (DPOAEs) in response to sound stimulation with two tones. DPOAEs originate from mechanical nonlinearity in the cochlea. For stimulus frequencies, f1 and f2, with f2/f1 = 1.2 and f2 = 4-9.5 kHz, and intensities L1 and L2, with L1 = 0.4L(2) + 39 dB and L2 = 20-65 dB SPL, the DPOAE displacement amplitudes were no more than 8 pm across subjects (n = 20), with hearing loss up to 16 dB. DPOAE vibration was nonlinearly dependent on vibration at f2. The dependence allowed the hearing threshold to be estimated objectively with high accuracy; the standard deviation of the threshold estimate was only 8.6 dB SPL. This device promises to be a powerful tool for differentially characterizing the mechanical condition of the cochlea and middle ear with high accuracy.
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Project description:The tectorial membrane (TM) is widely believed to play an important role in determining the ear's ability to detect and resolve incoming acoustic information. While it is still unclear precisely what that role is, the TM has been hypothesized to help overcome viscous forces and thereby sharpen mechanical tuning of the sensory cells. Lizards present a unique opportunity to further study the role of the TM given the diverse inner-ear morphological differences across species. Furthermore, stimulus-frequency otoacoustic emissions (SFOAEs), sounds emitted by the ear in response to a tone, noninvasively probe the frequency selectivity of the ear. We report estimates of auditory tuning derived from SFOAEs for 12 different species of lizards with widely varying TM morphology. Despite gross anatomical differences across the species examined herein, low-level SFOAEs were readily measurable in all ears tested, even in non-TM species whose basilar papilla contained as few as 50-60 hair cells. Our measurements generally support theoretical predictions: longer delays/sharper tuning features are found in species with a TM relative to those without. However, SFOAEs from at least one non-TM species (Anolis) with long delays suggest there are likely additional micromechanical factors at play that can directly affect tuning. Additionally, in the one species examined with a continuous TM (Aspidoscelis) where cell-to-cell coupling is presumably relatively stronger, delays were intermediate. This observation appears consistent with recent reports that suggest the TM may play a more complex macromechanical role in the mammalian cochlea via longitudinal energy distribution (and thereby affect tuning). Although significant differences exist between reptilian and mammalian auditory biophysics, understanding lizard OAE generation mechanisms yields significant insight into fundamental principles at work in all vertebrate ears.
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Project description:Auditory sensory outer hair cells are thought to amplify sound-induced basilar membrane vibration through a feedback mechanism to enhance hearing sensitivity. For optimal amplification, the outer hair cell-generated force must act on the basilar membrane at an appropriate time at every cycle. However, the temporal relationship between the outer hair cell-driven reticular lamina vibration and the basilar membrane vibration remains unclear. By measuring sub-nanometer vibrations directly from outer hair cells using a custom-built heterodyne low-coherence interferometer, we demonstrate in living gerbil cochleae that the reticular lamina vibration occurs after, not before, the basilar membrane vibration. Both tone- and click-induced responses indicate that the reticular lamina and basilar membrane vibrate in opposite directions at the cochlear base and they oscillate in phase near the best-frequency location. Our results suggest that outer hair cells enhance hearing sensitivity through a global hydromechanical mechanism, rather than through a local mechanical feedback as commonly supposed.
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Project description:The mammalian cochlea is able to detect faint sounds due to the presence of an active nonlinear feedback mechanism that boosts cochlear vibrations of low amplitude. Because of this feedback, self-sustained oscillations called spontaneous otoacoustic emissions (SOAEs) can often be measured in the ear canal. Recent experiments in genetically modified mice have demonstrated that mutations of the genes expressed in the tectorial membrane (TM), an extracellular matrix located in the cochlea, can significantly enhance the generation of SOAEs. Multiple untested mechanisms have been proposed to explain these unexpected results. In this work, a physiologically motivated computational model of a mammalian species commonly studied in auditory research, the gerbil, is used to demonstrate that altering the viscoelastic properties of the TM tends to affect the linear stability of the cochlea, SOAE generation and the cochlear response to low amplitude stimuli. These results suggest that changes in TM properties might be the underlying cause for SOAE enhancement in some mutant mice. Furthermore, these theoretical findings imply that the TM contributes to keeping the mammalian cochlea near an oscillatory instability, which promotes high sensitivity and the detection of low level stimuli.
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Project description:Long-delay components showing a symmetrical pattern with positive and negative delays can be observed in the time-frequency representation (or in the inverse Fourier transform) of distortion product otoacoustic emissions. Positive-only phase-gradient delays are predicted by place-fixed backscattering mechanisms, such as coherent reflection due to roughness, whereas the nonlinear distortion wave-fixed mechanism should generate an almost null-delay component. The symmetrical delay pattern arises whenever spectral amplitude fluctuations are not fully correlated to phase fluctuations. An interpretation of this phenomenon is proposed, involving place-fixed modulation of the spectral strength of the wave-fixed nonlinear generator. Experimental data are shown in which these negative-delay sources are particularly strong, and further amplified by contralateral stimulation, suggesting that this effect could be dynamically enhanced. Analytical solutions of a linear 1-d transmission line model, in which cubic nonlinearity and roughness were added as small perturbations, have been used to test this hypothesis.
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Project description:Otoacoustic emissions are an indicator of a normally functioning cochlea and as such are a useful tool for non-invasive diagnosis as well as for understanding cochlear function. While these emitted waves are hypothesized to arise from active processes and exit through the cochlear fluids, neither the precise mechanism by which these emissions are generated nor the transmission pathway is completely known. With regard to the acoustic pathway, two competing hypotheses exist to explain the dominant mode of emission. One hypothesis, the backward-traveling wave hypothesis, posits that the emitted wave propagates as a coupled fluid-structure wave while the alternate hypothesis implicates a fast, compressional wave in the fluid as the main mechanism of energy transfer. In this paper, we study the acoustic pathway for transmission of energy from the inside of the cochlea to the outside through a physiologically-based theoretical model. Using a well-defined, compact source of internal excitation, we predict that the emission is dominated by a backward traveling fluid-structure wave. However, in an active model of the cochlea, a forward traveling wave basal to the location of the force is possible in a limited region around the best place. Finally, the model does predict the dominance of compressional waves under a different excitation, such as an apical excitation.
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Project description:When two pure tones (or primaries) of slightly different frequencies (f (1) and f (2)) are presented to the ear, new frequency components are generated by nonlinear interaction of the primaries within the cochlea. These new components can be recorded in the ear canal as otoacoustic emissions (OAE). The level of the 2f (1)-f (2) OAE component is known as the distortion product otoacoustic emission (DPOAE) and is regarded as an indicator of the physiological state of the cochlea. The current view is that maximal level DPOAEs occur for primaries that produce equal excitation at the f (2) cochlear region, but this notion cannot be directly tested in living humans because it is impossible to record their cochlear responses while monitoring their ear canal DPOAE levels. On the other hand, it has been claimed that the temporal masking curve (TMC) method of inferring human basilar membrane responses allows measurement of the levels of equally effective pure tones at any given cochlear site. The assumptions of this behavioral method, however, lack firm physiological support in humans. Here, the TMC method was applied to test the current notion on the conditions that maximize DPOAE levels in humans. DPOAE and TMC results were mutually consistent for frequencies of 1 and 4 kHz and for levels below around 65 dB sound pressure level. This match supports the current view on the generation of maximal level DPOAEs as well as the assumptions of the behavioral TMC method.
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Project description:OBJECTIVES:The purpose of this study was to analyze distortion product otoacoustic emission (DPOAE) level and signal to noise ratio in a group of infants from birth to 4 months of age to optimize prediction of hearing status. DPOAEs from infants with normal hearing (NH) and hearing loss (HL) were used to predict the presence of conductive HL (CHL), sensorineural HL (SNHL), and mixed HL (MHL). Wideband ambient absorbance was also measured and compared among the HL types. DESIGN:This is a prospective, longitudinal study of 279 infants with verified NH and HL, including conductive, sensorineural, and mixed types that were enrolled from a well-baby nursery and two neonatal intensive care units in Cincinnati, Ohio. At approximately 1 month of age, DPOAEs (1-8?kHz), wideband absorbance (0.25-8?kHz), and air and bone conduction diagnostic tone burst auditory brainstem response (0.5-4?kHz) thresholds were measured. Hearing status was verified at approximately 9 months of age with visual reinforcement audiometry (0.5-4?kHz). Auditory brainstem response air conduction thresholds were used to assign infants to an NH or HL group, and the efficacy of DPOAE data to classify ears as NH or HL was analyzed using receiver operating characteristic (ROC) curves. Two summary statistics of the ROC curve were calculated: the area under the ROC curve and the point of symmetry on the curve at which the sensitivity and specificity were equal. DPOAE level and signal to noise ratio cutoff values were defined at each frequency as the symmetry point on their respective ROC curve, and DPOAE results were combined across frequency in a multifrequency analysis to predict the presence of HL. RESULTS:Single-frequency test performance of DPOAEs was best at mid to high frequencies (3-8?kHz) with intermediate performance at 1.5 and 2?kHz and chance performance at 1?kHz. Infants with a conductive component to their HL (CHL and MHL combined) displayed significantly lower ambient absorbance values than the NH group. No differences in ambient absorbance were found between the NH and SNHL groups. Multifrequency analysis resulted in the best prediction of HL for the SNHL/MHL group with poorer sensitivity values when infants with CHL were included. CONCLUSIONS:Clinical interpretation of DPOAEs in infants can be improved by using age-appropriate normative ranges and optimized cutoff values. DPOAE interpretation is most predictive at higher F2 test frequencies in young infants (2-8?kHz) due to poor test performance at 1 to 1.5?kHz. Multifrequency rules can be used to improve sensitivity while balancing specificity. Last, a sensitive middle ear measure such as wideband absorbance should be included in the test battery to assess possibility of a conductive component to the HL.
| S-EPMC6127008 | biostudies-literature