Backward Propagation of Otoacoustic Emissions

IntroductionsSounds impinging the eardrum are transmitted viamiddle ear ossicles to the oval window. Stapes vibra-tion creates pressure difference between the scala tym-pani and the scala vestibuli. This pressure differencecauses a movement of cochlear partition and adjacentcochlear fluids. Basilar membrane vibrations result indeflection of hair cell stereocilia, which gate ion chan-nels on their tips. This mechanical-to-electrical trans-duction process converts mechanical vibrations intoelec…     

Stimulus Frequency Otoacoustic Emission Delays and Generating Mechanisms in Guinea Pigs,Chinchillas, and Simulations

According to coherent reflection theory (CRT), stimulus frequency otoacoustic emissions (SFOAEs) arise from cochlear irregularities coherently reflecting energy from basilar membrane motion within the traveling-wave peak. This reflected energy arrives in the ear canal predominantly with a single delay at each frequency. However, data from humans and animals indicate that (1) SFOAEs can have multiple delay components, (2) low-frequency SFOAE delays are too short to be accounted for by CRT, and (3) “SFOAEs” obtained with a 2nd (“suppressor”) tone ≥2 octaves above the probe tone have been interpreted as arising from the area basal to the region of cochlear amplification. To explore these issues, we collected SFOAEs by the suppression method in guinea pigs and time-frequency analyzed these data, simulated SFOAEs, and published chinchilla SFOAEs. Time-frequency analysis revealed that most frequencies showed only one SFOAE delay component while other frequencies had multiple components including some with short delays. We found no systematic patterns in the occurrence of multiple delay components. Using a cochlear model that had significant basilar membrane motion only in the peak region of the traveling wave, simulated SFOAEs had single and multiple delay components similar to the animal SFOAEs. This result indicates that multiple components (including ones with short delays) can originate from cochlear mechanical irregularities in the SFOAE peak region and are not necessarily indicative of SFOAE sources in regions ≥2 octaves basal of the SFOAE peak region. We conclude that SFOAEs obtained with suppressors close to the probe frequency provide information primarily about the mechanical response in the region that receives amplification, and we attribute the too-short SFOAE delays at low frequencies to distortion-source SFOAEs and coherent reflection from multiple cochlear motions. Our findings suggest that CRT needs revision to include reflections from multiple motions in the cochlear apex.     

On Cochlear Impedances and the Miscomputation of Power Gain

In their article, “Measurement of cochlear power gain in the sensitive gerbil ear,” Ren et al. (Nat Commun 2:216, 2011) claim to provide “the first direct experimental evidence of power amplification in the sensitive living cochlea.” While we recognize the technical challenges of the experiments and appreciate the beauty of the data, the authors’ analysis and interpretation of the measurements are invalid. We review the concept of impedance (i.e., the ratio of pressure to velocity) as it applies to cochlear mechanics and show that Ren et al. mistakenly equate the impedances near the basilar membrane and stapes with the impedance characteristic of an infinite, uniform tube of fluid. As a consequence of this error, Ren et al.’s measurements and analysis provide no evidence for power amplification in the cochlea. Compelling evidence for power amplification has, however, been previously provided by others.     

Spatial representation of basilar membrane movement with 3D computer graphics

A mathematical model of the cochlea was implemented on a computer. The basilar membrane motion was computed for single, two, and multi-tone stimuli as well as for musical sounds and vowels. The pattern of the travelling waves were presented in three-dimensional color computer graphics. The high performance 3D graphics system performs local hidden surface removal, 3D geometric transformations and supports local lighting models to generate truly realistic shading for complex 3D objects. An addressable 1280 by 1024 pixel matrix assures crisp, precise resolution of the finest detail in the graphic images. The spatial pattern of basilar membrane motion conveys an impression of the image of acoustic stimuli on the basilar membrane. Firstly, the motion pattern of the travelling wave to a single tone is presented. The superposition of several tones (two-tone, multi-tone) causes a superposition of the travelling waves along the basilar membrane whereby the place principle in the cochlear partition becomes more clearly recognizable. Sounds (flute and violin) and vowels (German "u" and "i") evoke a complex motion pattern on the basilar membrane. The realization of the chronological order of movements on the basilar membrane can be made by computer animation. This enables the analysis of the space-time patterns of complex acoustic stimuli.     

A comparison of OAEs arising from different generation mechanisms in guinea pig

Otoacoustic emissions provide unambiguous evidence that the cochlea supports energy propagation both towards, and away from, the stapes. The standard wave model for energy transport and cochlear mechanical amplification provides for compressional and inertial waves to transport this energy, the compressional wave through the fluids and the inertial wave along the basilar membrane via fluid coupling. It is generally accepted that energy propagation away from the stapes is dominated by a traveling wave mechanism along the basilar membrane. The mechanism by which energy is predominantly transported back to the stapes remains controversial. Here, we compared signal onset delay measurements and rise/steady-state/fall times for SFOAEs and 2f1-f2 OAEs (f2/f1=1.2) obtained using a pulsed-tone paradigm in guinea pig. Comparison of 2f1-f2 OAE signal onset delay for the OAE arising from the f2 region with SFOAE signal onset delay (matched to the f2 stimulus frequency) based on signal onset occurring at 10% of the peak signal amplitude was suggestive of a bi-directional traveling wave mechanism. However, significant variability in signal onset delay and signal rise, steady-state duration, and fall times for both the 2f1-f2 OAE and SFOAE was found, qualifying this interpretation. Such variability requires explanation, awaiting further studies.     

Implications from cochlear implant insertion for cochlear mechanics

It is usually thought that the displacements of the two inner ear windows induced by sound stimuli lead to pressure differences across the basilar membrane and to a passive mechanical traveling wave progressing along the membrane. However, opening a hole in the sealed inner ear wall in experimental animals is surprisingly not accompanied by auditory threshold elevations. It has also been shown that even in patients undergoing cochlear implantation, elevation of threshold to low-frequency acoustic stimulation is often not seen accompanying the making of a hole in the wall of the cochlea for insertion of the implant. Such threshold elevations would be expected to result from opening the cochlea, reducing cochlear impedance, altering hydrodynamics. These considerations can be taken as additional evidence that it may not be the passive basilar membrane traveling wave which elicits hearing at low sound intensities, but rather factors connected with cochlear fluid pressures and fluid mechanics.     

Effect of bimodal hearing in Korean children with profound hearing loss

Conclusion. Bimodal hearing with combined acoustic stimulation and electric stimulation could enhance speech performance in deaf patients by residual hearing even though the amount of residual hearing is not enough to be used for communication by amplification. Objectives. The cochlear implant (CI) is a well-known therapeutic option for patients with profound hearing loss. However, deaf patients with a CI still have trouble in localization of sounds and understanding speech in a noisy environment. The aim of this study was to evaluate the benefits of bimodal hearing with a CI in one ear and a hearing aid in the contralateral ear in Korean children with profound hearing loss. Subjects and methods. Fourteen deaf children with residual hearing participated in this study. There were eight male and six female patients, with an age range of 4.6–13.8 years at the time of testing. The test was conducted between 3 months and 4.2 years after cochlear implantation. Speech performance was examined in a noisy environment using Korean word lists. A speech sound and the noise were presented to the child from the front loudspeaker. Results. The results showed that speech performance in a noisy environment was significantly better with bimodal hearing than with a CI alone.     

Rapid Increase in Neural Conduction Time in the Adult Human Auditory Brainstem Following Sudden Unilateral Deafness

Individuals with sudden unilateral deafness offer a unique opportunity to study plasticity of the binaural auditory system in adult humans. Stimulation of the intact ear results in increased activity in the auditory cortex. However, there are no reports of changes at sub-cortical levels in humans. Therefore, the aim of the present study was to investigate changes in sub-cortical activity immediately before and after the onset of surgically induced unilateral deafness in adult humans. Click-evoked auditory brainstem responses (ABRs) to stimulation of the healthy ear were recorded from ten adults during the course of translabyrinthine surgery for the removal of a unilateral acoustic neuroma. This surgical technique always results in abrupt deafferentation of the affected ear. The results revealed a rapid (within minutes) reduction in latency of wave V (mean pre = 6.55 ms; mean post = 6.15 ms; p < 0.001). A latency reduction was also observed for wave III (mean pre = 4.40 ms; mean post = 4.13 ms; p < 0.001). These reductions in response latency are consistent with functional changes including disinhibition or/and more rapid intra-cellular signalling affecting binaurally sensitive neurons in the central auditory system. The results are highly relevant for improved understanding of putative physiological mechanisms underlying perceptual disorders such as tinnitus and hyperacusis.     

Wave propagation and dispersion in the cochlea

Insight into cochlear mechanics can be obtained from semi-analytical and asymptotic solution methods of which the Liouville-Green (LG) method — in another context known as the WKB method — is the most important one. This paper describes the dispersion properties of fluid waves in general terms and develops the LG formulation on that basis. The eikonal equation of the LG method is shown to be identical to the dispersion relation in dispersive-wave theory. Consideration of the group velocity then leads to the derivation of the central LG formula as it has been used in an earlier paper on the LG method (Boer E. and Viergever M.A. (1982): Hearing Res. 8, 131–155). The formulation appears to apply as well to dissipative and active (i.e., energy-producing) systems. Of the many possible collateral subjects two are selected for a deeper discussion: amplification, concentration and expansion of energy, and the problem of reflection of cochlear waves. In the latter context, it is shown why — and under which conditions — cochlear waves are not reflected, despite the large degree of dispersion that they show. The analysis brings to light a fundamental asymmetry of the model regarding the direction of wave travel: waves travelling in the direction opposite to the normal one are likely to undergo reflection, while waves in the normal direction are not reflected.     

Monopolar Intracochlear Pulse Trains Selectively Activate the Inferior Colliculus

Previous cochlear implant studies using isolated electrical stimulus pulses in animal models have reported that intracochlear monopolar stimulus configurations elicit broad extents of neuronal activation within the central auditory system—much broader than the activation patterns produced by bipolar electrode pairs or acoustic tones. However, psychophysical and speech reception studies that use sustained pulse trains do not show clear performance differences for monopolar versus bipolar configurations. To test whether monopolar intracochlear stimulation can produce selective activation of the inferior colliculus, we measured activation widths along the tonotopic axis of the inferior colliculus for acoustic tones and 1,000-pulse/s electrical pulse trains in guinea pigs and cats. Electrical pulse trains were presented using an array of 6–12 stimulating electrodes distributed longitudinally on a space-filling silicone carrier positioned in the scala tympani of the cochlea. We found that for monopolar, bipolar, and acoustic stimuli, activation widths were significantly narrower for sustained responses than for the transient response to the stimulus onset. Furthermore, monopolar and bipolar stimuli elicited similar activation widths when compared at stimulus levels that produced similar peak spike rates. Surprisingly, we found that in guinea pigs, monopolar and bipolar stimuli produced narrower sustained activation than 60 dB sound pressure level acoustic tones when compared at stimulus levels that produced similar peak spike rates. Therefore, we conclude that intracochlear electrical stimulation using monopolar pulse trains can produce activation patterns that are at least as selective as bipolar or acoustic stimulation.     

Basilar Membrane and Tectorial Membrane Stiffness in the CBA/CaJ Mouse

The mouse has become an important animal model in understanding cochlear function. Structures, such as the tectorial membrane or hair cells, have been changed by gene manipulation, and the resulting effect on cochlear function has been studied. To contrast those findings, physical properties of the basilar membrane (BM) and tectorial membrane (TM) in mice without gene mutation are of great importance. Using the hemicochlea of CBA/CaJ mice, we have demonstrated that tectorial membrane (TM) and basilar membrane (BM) revealed a stiffness gradient along the cochlea. While a simple spring mass resonator predicts the change in the characteristic frequency of the BM, the spring mass model does not predict the frequency change along the TM. Plateau stiffness values of the TM were 0.6 ± 0.5, 0.2 ± 0.1, and 0.09 ± 0.09 N/m for the basal, middle, and upper turns, respectively. The BM plateau stiffness values were 3.7 ± 2.2, 1.2 ± 1.2, and 0.5 ± 0.5 N/m for the basal, middle, and upper turns, respectively. Estimations of the TM Young’s modulus (in kPa) revealed 24.3 ± 25.2 for the basal turns, 5.1 ± 4.5 for the middle turns, and 1.9 ± 1.6 for the apical turns. Young’s modulus determined at the BM pectinate zone was 76.8 ± 72, 23.9 ± 30.6, and 9.4 ± 6.2 kPa for the basal, middle, and apical turns, respectively. The reported stiffness values of the CBA/CaJ mouse TM and BM provide basic data for the physical properties of its organ of Corti.     

In vivo imaging of mammalian cochlear blood flow using fluorescence microendoscopy.

AIMS: We sought to develop techniques for visualizing cochlear blood flow in live mammalian subjects using fluorescence microendoscopy. BACKGROUND: Inner ear microcirculation appears to be intimately involved in cochlear function. Blood velocity measurements suggest that intense sounds can alter cochlear blood flow. Disruption of cochlear blood flow may be a significant cause of hearing impairment, including sudden sensorineural hearing loss. However, inability to image cochlear blood flow in a nondestructive manner has limited investigation of the role of inner ear microcirculation in hearing function. Present techniques for imaging cochlear microcirculation using intravital light microscopy involve extensive perturbations to cochlear structure, precluding application in human patients. The few previous endoscopy studies of the cochlea have suffered from optical resolution insufficient for visualizing cochlear microvasculature. Fluorescence microendoscopy is an emerging minimally invasive imaging modality that provides micron-scale resolution in tissues inaccessible to light microscopy. In this article, we describe the use of fluorescence microendoscopy in live guinea pigs to image capillary blood flow and movements of individual red blood cells within the basal turn of the cochlea. METHODS: We anesthetized eight adult guinea pigs and accessed the inner ear through the mastoid bulla. After intravenous injection of fluorescein dye, we made a limited cochleostomy and introduced a compound doublet gradient refractive index endoscope probe 1 mm in diameter into the inner ear. We then imaged cochlear blood flow within individual vessels in an epifluorescence configuration using one-photon fluorescence microendoscopy. RESULTS: We observed single red blood cells passing through individual capillaries in several cochlear structures, including the round window membrane, spiral ligament, osseous spiral lamina, and basilar membrane. Blood flow velocities within inner ear capillaries varied widely, with observed speeds reaching up to approximately 500 microm/s. CONCLUSION: Fluorescence microendoscopy permits visualization of cochlear microcirculation with micron-scale optical resolution and determination of blood flow velocities through analysis of video sequences.     

Pitch Comparisons between Electrical Stimulation of a Cochlear Implant and Acoustic Stimuli Presented to a Normal-hearing Contralateral Ear

Four cochlear implant users, having normal hearing in the unimplanted ear, compared the pitches of electrical and acoustic stimuli presented to the two ears. Comparisons were between 1,031-pps pulse trains and pure tones or between 12 and 25-pps electric pulse trains and bandpass-filtered acoustic pulse trains of the same rate. Three methods—pitch adjustment, constant stimuli, and interleaved adaptive procedures—were used. For all methods, we showed that the results can be strongly influenced by non-sensory biases arising from the range of acoustic stimuli presented, and proposed a series of checks that should be made to alert the experimenter to those biases. We then showed that the results of comparisons that survived these checks do not deviate consistently from the predictions of a widely-used cochlear frequency-to-place formula or of a computational cochlear model. We also demonstrate that substantial range effects occur with other widely used experimental methods, even for normal-hearing listeners.     

Noise exposure of the inner ear during drilling a cochleostomy for cochlear implantation

OBJECTIVES: Inserting an electrode array into the cochlea may cause inner ear trauma, which has to be minimized, particularly in cochlear implant patients with substantial residual hearing. Another potential inner ear trauma has, to a large extent, been neglected so far: the acoustic trauma that can occur during cochleostomy using different techniques. In this study, the noise exposure of the inner ear during the drilling procedure was re-evaluated. In experiments on temporal bones, quantitative measurements of sound pressure level (SPL) were carried out while a cochleostomy for cochlear implantation was drilled. STUDY DESIGN: Experimental study. MATERIALS AND METHODS: Acoustic measurements during different drilling procedures were carried out on four human temporal bone preparations equipped with microphones attached to the round window. Special calibrations were carried out, which allowed determination of SPLs affecting the cochlea during the drilling procedure. RESULTS: The highest SPLs measured on the cochlea were recorded when a still-intact endosteal membrane was touched by the burr. The SPL exceeded 130 dB and reached a level almost comparable with the situation when the ossicular chain is touched by a running burr. CONCLUSIONS: In the drilling procedure for a cochleostomy, the inner ear may be affected by very high SPLs, particularly if the endosteal membrane is left intact and comes into contact with the running burr. Of course, the resulting SPLs depend on the drilling speed and the size and characteristics of the burr (larger burrs cause higher SPLs); however, we are of the opinion that the cochlear function is at risk, anyway, if special precaution is not exercised. Even when working with reduced drilling speed, the surgeon should be aware of the high risk in the form of an acoustic trauma, which may endanger residual hearing. Recommendations in terms of "soft surgery" are given in the paper (e.g., the use of microhooks instead of a drill to remove the very last shell of bone covering the cochlea).     

Intermodulation distortion in the cochlea: could basal vibration be the major cause of round window CM distortion?

Acoustic and cochlear microphonic (CM) distortion products (DPs) have been measured at the meatus and round window, respectively, in the gerbil. It was hoped to deduce the relationship between round window CM distortion and vibrational energy of cochlear origin recorded as acoustic DP emissions in the meatus. Five different distortion components have been studied. For moderate intensities of stimulation, correlation between ear canal acoustic and CM distortion signals in respect of level and delay provides evidence of a common transmission path. One explanation of this is that the CM is being generated as a direct result of cochlea-generated vibration which reaches the base of the cochlea. The basalward propagation of these signals is discussed.     

Speech Perception in Noise with a Harmonic Complex Excited Vocoder

A cochlear implant (CI) presents band-pass-filtered acoustic envelope information by modulating current pulse train levels. Similarly, a vocoder presents envelope information by modulating an acoustic carrier. By studying how normal hearing (NH) listeners are able to understand degraded speech signals with a vocoder, the parameters that best simulate electric hearing and factors that might contribute to the NH-CI performance difference may be better understood. A vocoder with harmonic complex carriers (fundamental frequency, f₀ = 100 Hz) was used to study the effect of carrier phase dispersion on speech envelopes and intelligibility. The starting phases of the harmonic components were randomly dispersed to varying degrees prior to carrier filtering and modulation. NH listeners were tested on recognition of a closed set of vocoded words in background noise. Two sets of synthesis filters simulated different amounts of current spread in CIs. Results showed that the speech vocoded with carriers whose starting phases were maximally dispersed was the most intelligible. Superior speech understanding may have been a result of the flattening of the dispersed-phase carrier’s intrinsic temporal envelopes produced by the large number of interacting components in the high-frequency channels. Cross-correlogram analyses of auditory nerve model simulations confirmed that randomly dispersing the carrier’s component starting phases resulted in better neural envelope representation. However, neural metrics extracted from these analyses were not found to accurately predict speech recognition scores for all vocoded speech conditions. It is possible that central speech understanding mechanisms are insensitive to the envelope-fine structure dichotomy exploited by vocoders.     

Can shape deformations of the organ of Corti influence the travelling wave in the cochlea?

Via their motile reactions outer hair cells may produce oscillatory power or amplify the fluid waves in the cochlea. When the cells move or change their lengths, the organ of Corti (OC) will change its shape. This paper describes the consequences of such shape changes for the physics of cochlear waves. The assertions posed are based on a mathematical derivation but the major conclusions can be grasped without following the mathematics in detail. There are two basic types of OC deformations. In the first type the net cross-sectional area of the OC is periodically varying, in the second type--the central subject of this paper--is is not. It is shown that for the latter type of OC shape deformation there is no interaction with the cochlear fluids, at least for long waves. Hence, in this case the outer hair cells cannot amplify cochlear waves. The other type of OC deformation has a better coupling with the cochlear waves. However, much of the pressure developed by oscillating hair cells is spent in the 'wrong' way, namely, by squeezing fluid lengthwise through the narrow channel of the OC. Therefore, in this mode power transfer from cochlear hair cells to cochlear fluids (and from there to the basilar membrane) would be very inefficient.     

How does the inner ear generate distortion product otoacoustic emissions?. Results from a realistic model of the human cochlea

There is general agreement that distortion product (DP) otoacoustic emissions elicited by stimuli up to 80-90 dB SPL originate from the saturating nonlinearity of the cochlear amplifier at the basilar membrane site, S, where the responses to the two primary tones overlap. There are, however, different interpretations of how the inner ear transmits the effects of this process to the stapes. The supporters of transmission line models assert that the phenomenon depends upon two main mechanisms: (1) the generation of forward and backward traveling waves (TWs) by DP oscillations at S; (2) the backward propagation of wave components reflected by 'micromechanical impedance perturbations' at the sites where the DP TWs peak. However, quantitative predictions based on this view are still lacking. In contrast, here we show, using a nonlinear hydrodynamic model, that the emissions are propagated almost instantaneously through the fluid.     

Representation of acoustic signals in the human cochlea in presence of a cochlear implant electrode

BACKGROUND: In subjects with remaining low frequency hearing, combined electric-acoustic stimulation (EAS) of the auditory system is a new therapeutic perspective. Intracochlear introduction of a cochlear implant electrode, however, may alter the biomechanical properties of the inner ear and thus affect perception of acoustic stimuli. STUDY DESIGN: Based on histological observations of morphologic changes after cochlear implantation in cadaveric and post mortem studies the effects of basilar membrane (BM) stiffening in the ascending basal and middle turns of the cochlea due to close contact of the BM with the electrode were simulated in a 3D-computational finite element model of the inner ear. To verify our simulated results, pre- and postoperative pure-tone audiograms of 13 subjects with substantial residual hearing, who underwent cochlear implantation, were evaluated. RESULTS: In the scenario of partial BM-fixation, acoustic energy of middle (2 kHz) and high (6 kHz) frequency was focused basally and apically to the fixed section, increasing BM displacement amplitudes up to 6 dB at a stimulation level of 94 dB (SPL). Lower frequencies were not affected by fixation in the basal and middle turn of the cochlea. In implanted subjects, a small but significant decrease of thresholds was observed at 1.5 kHz, a place in tonotopy adjacent to the tip region of the implanted electrode. CONCLUSION: Our model suggests that stiffening of the basilar membrane adjacent to an implanted electrode into the basal and middle cochlear turn did not affect BM movement in the low frequency area. Focussing of acoustic energy may increase perception in regions adjacent to the fixed section. Observations in implanted subjects were concordant with our model predictions. High frequencies, however, should not be amplified in patients using EAS to avoid disturbances in discrimination due to tonotopically incorrect frequency representation.