Relating the variability of tone-burst otoacoustic emission and auditory brainstem response latencies to the underlying cochlear mechanics

Forward and reverse cochlear latency and its relation to the frequency tuning of the auditory filters can be assessed using tone bursts (TBs). Otoacoustic emissions (TBOAEs) estimate the cochlear roundtrip time, while auditory brainstem responses (ABRs) to the same stimuli aim at measuring the auditory filter buildup time. Latency ratios are generally close to two and controversy exists about the relationship of this ratio to cochlear mechanics. We explored why the two methods provide different estimates of filter buildup time, and ratios with large inter-subject variability, using a time-domain model for OAEs and ABRs. We compared latencies for twenty models, in which all parameters but the cochlear irregularities responsible for reflection-source OAEs were identical, and found that TBOAE latencies were much more variable than ABR latencies. Multiple reflection-sources generated within the evoking stimulus bandwidth were found to shape the TBOAE envelope and complicate the interpretation of TBOAE latency and TBOAE/ABR ratios in terms of auditory filter tuning

The Noise Within: Signal-to-Noise Enhancement via Coherent Wave Amplification in the Mammalian Cochlea

The mammalian inner ear's extraordinary sensitivity has captivated scientists for decades, largely due to the crucial role played by outer hair cells (OHCs) and their unique piezoelectric properties. These specialized cells, arranged in three rows along the cochlea's sensory tissue, work in concert to amplify the faintest sounds. Referred to as the "cochlear amplifier", this mechanism poses a fascinating question: How does it effectively enhance ear sensitivity in real-world scenarios? While simplistic views attribute this enhancement solely to increased cochlear gain, the presence of internal noise in practical settings necessitates a more nuanced approach. Achieving a genuine boost in sensitivity through amplification requires that the signals are amplified more than the internal noise, thus presenting an intriguing challenge. In this study, we analyze the effects of coherent amplification on both signals and internal noise, employing a simple yet powerful mathematical framework and a simplified model of cochlear physics. Our findings not only generalize and expand upon previous discoveries concerning the impact of spatially coherent amplification on signal degradation in active gain media, but also unveil the elegant and efficient wave-based strategy employed by the cochlea to boost ear sensitivity

The cochlea as a smart structure

The cochlea is part of the inner ear and its mechanical response provides us with many aspects of our amazingly sensitive and selective hearing. The human cochlea is a coiled tube, with two main fluid chambers running along its length, separated by a 35 mm-long flexible partition that has its own internal dynamics. A dispersive wave can propagate along the cochlea due to the interaction between the inertia of the fluid and the dynamics of the partition. This partition includes about 12 000 outer hair cells, which have different structures, on a micrometre and a nanometre scale, and act both as motional sensors and as motional actuators. The local feedback action of all these cells amplifies the motion inside the inner ear by more than 40 dB at low sound pressure levels. The feedback loops become saturated at higher sound pressure levels, however, so that the feedback gain is reduced, leading to a compression of the dynamic range in the cochlear amplifier. This helps the sensory cells, with a dynamic range of only about 30 dB, to respond to sounds with a dynamic range of more than 120 dB. The active and nonlinear nature of the dynamics within the cochlea give rise to a number of other phenomena, such as otoacoustic emissions, which can be used as a diagnostic test for hearing problems in newborn children, for example. In this paper we view the mechanical action of the cochlea as a smart structure. In particular a simplified wave model of the cochlear dynamics is reviewed that represents its essential features. This can be used to predict the motion along the cochlea when the cochlea is passive, at high levels, and also the effect of the cochlear amplifier, at low level

Simultaneous Measurement of Middle-Ear Input Impedance and Forward/Reverse Transmission in Cat

Reported here is a technique for measuring forward and reverse middle-ear transmission that exploits distortion-product otoacoustic emissions (DPOAEs) to drive the middle ear in reverse without opening the inner ear. The technique allows measurement of DPOAEs, middle-ear input impedance, and forward and reverse middle-ear transfer functions in the same animal. Intermodulation distortion in the cochlea generates a DPOAE at frequency 2f1-f 2 measurable in both ear-canal pressure and the velocity of the stapes. The forward transfer function is computed from stapes velocities and corresponding ear-canal pressures measured at the two primary frequencies; the reverse transfer function is computed from velocity and pressure measurements at the DPOAE frequency. Middle-ear input impedance is computed from ear-canal pressure measurements and the measured Thévenin equivalent of the sound-delivery system. The technique was applied to measure middle-ear characteristics in anesthetized cats with widely opened middle-ear cavities (0.2-10 kHz). Stapes velocity was measured at the incudo-stapedial joint. Results on five animals are reported and compared with a published middle-ear model. The measured forward transfer functions and input impedances generally agree with previous measurements, and all measurements agree qualitatively with model predictions. The reverse transfer function is shown to depend on the acoustic load in the ear canal, and the measurements are used to compute the round-trip middle-ear gain and delay. Finally, the measurements are used to estimate the parameters of a two-port transfer-matrix description of the cat middle ear

Listening to the Ear

Otoacoustic emissions demonstrate that the ear creates sound while listening to sound, offering a promising acoustic window on the mechanics of hearing in awake, listening human beings. That window is clouded, however, by an incomplete knowledge of wave reflection and transmission, both forth and back within the cochlea and through the middle ear. This thesis "does windows," addressing wave propagation and scattering on both sides of the middle ear. A summary of highlights follows. Measurements of the cochlear input impedance in cat are used to identify a new symmetry in cochlear mechanics-termed "tapering symmetry" after its geometric interpretation in simple models-that guarantees that the wavelength of the traveling wave changes slowly with position near the stapes. Waves therefore propagate without reflection through the basal turns of the cochlea. Analytic methods for solving the cochlear wave equations using a perturbative scattering series are given and used to demonstrate that, contrary to common belief, conventional cochlear models exhibit negligible internal reflection whether or not they accurately represent the tapering symmetries of the inner ear. Frameworks for the systematic "deconstruction" of eardrum and middle-ear transduction characteristics are developed and applied to the analysis of noninvasive measurements of middle-ear and cochlear mechanics. A simple phenomenological model of inner-ear compressibility that correctly predicts hearing thresholds in patients with missing or disarticulated middle-ear ossicles is developed and used to establish an upper bound on cochlear compressibility several orders of magnitude smaller than that provided by direct measurements. Accurate measurements of stimulus frequency evoked otoacoustic emissions are performed and used to determine the form and frequency variation of the cochlear traveling-wave ratio noninvasively. Those measurements are inverted to obtain the spatial distribution of mechanical inhomogeneities responsible for evoked emission. Although current models require that the periodicities found in emission spectra and threshold hearing curves originate in a corresponding corrugation in the mechanics of the cochlea, it is shown that the observed spectral periodicities can arise spontaneously through the dynamics of wave propagation and reflection and that the organ of Corti, as suggested by the anatomy, need manifest no particular translational symmetries

Acoustic Mechanisms that Determine the Ear-Canal Sound Pressures Generated by Earphones

In clinical measurements of hearing sensitivity, a given earphone is assumed to produce essentially the same sound-pressure level in all ears. However, recent measurements [Voss et al., Ear and Hearing (in press)] show that with some middle-ear pathologies, ear-canal sound pressures can deviate by as much as 35 dB from the normal-ear value; the deviations depend on the earphone, the middle-ear pathology, and frequency. These pressure variations cause errors in the results of hearing tests. Models developed here identify acoustic mechanisms that cause pressure variations in certain pathological conditions. The models combine measurement-based Thevenin equivalents for insert and supra-aural earphones with lumped-element models for both the normal ear and ears with pathologies that alter the ear\u27s impedance (mastoid bowl, tympanostomy tube, tympanic-membrane perforation, and a \u27high- impedance\u27 ear). Comparison of the earphones\u27 Thevenin impedances to the ear\u27s input impedance with these middle-ear conditions shows that neither class of earphone acts as an ideal pressure source; with some middle-ear pathologies, the ear\u27s input impedance deviates substantially from normal and thereby causes abnormal ear-canal pressure levels. In general, for the three conditions that make the ear\u27s impedance magnitude lower than normal, the model predicts a reduced ear-canal pressure (as much as 35 dB), with a greater pressure reduction with an insert earphone than with a supra-aural earphone. In contrast, the model predicts that ear-canal pressure levels increase only a few dB when the ear has an increased impedance magnitude; the compliance of the air-space between the tympanic membrane and the earphone determines an upper limit on the effect of the middle-ear\u27s impedance increase. Acoustic leaks at the earphone-to-ear connection can also cause uncontrolled pressure variations during hearing tests. From measurements at the supra-aural earphone-to-ear connection, we conclude that it is unusual for the connection between the earphone cushion and the pinna to seal effectively for frequencies below 250 Hz. The models developed here explain the measured pressure variations with several pathologic ears. Understanding these mechanisms should inform the design of more accurate audiometric systems which might include a microphone that monitors the ear-canal pressure and corrects deviations from normal

Mammalian behavior and physiology converge to confirm sharper cochlear tuning in humans

Frequency analysis of sound by the cochlea is the most fundamental property of the auditory system. Despite its importance, the resolution of this frequency analysis in humans remains controversial. The controversy persists because the methods used to estimate tuning in humans are indirect and have not all been independently validated in other species. Some data suggest that human cochlear tuning is considerably sharper than that of laboratory animals, while others suggest little or no difference between species. We show here in a single species (ferret) that behavioral estimates of tuning bandwidths obtained using perceptual masking methods, and objective estimates obtained using otoacoustic emissions, both also employed in humans, agree closely with direct physiological measurements from single auditory-nerve fibers. Combined with human behavioral data, this outcome indicates that the frequency analysis performed by the human cochlea is of significantly higher resolution than found in common laboratory animals. This finding raises important questions about the evolutionary origins of human cochlear tuning, its role in the emergence of speech communication, and the mechanisms underlying our ability to separate and process natural sounds in complex acoustic environments

Across-Channel Timing Differences as a Potential Code for the Frequency of Pure Tones

When a pure tone or low-numbered harmonic is presented to a listener, the resulting travelling wave in the cochlea slows down at the portion of the basilar membrane (BM) tuned to the input frequency due to the filtering properties of the BM. This slowing is reflected in the phase of the response of neurons across the auditory nerve (AN) array. It has been suggested that the auditory system exploits these across-channel timing differences to encode the pitch of both pure tones and resolved harmonics in complex tones. Here, we report a quantitative analysis of previously published data on the response of guinea pig AN fibres, of a range of characteristic frequencies, to pure tones of different frequencies and levels. We conclude that although the use of across-channel timing cues provides an a priori attractive and plausible means of encoding pitch, many of the most obvious metrics for using that cue produce pitch estimates that are strongly influenced by the overall level and therefore are unlikely to provide a straightforward means for encoding the pitch of pure tones

Signal Transmission in the Auditory System

Contains table of contents for Section 3, an introduction and reports on six research projects.National Institutes of Health Grant RO1-DC-00194-11National Institutes of Health Grant PO1-DC00119 Sub-Project 1National Institutes of Health Grant F32-DC00073-3National Institutes of Health Contract P01-DC00119National Institutes of Health Grant R01 DC00238National Institutes of Health Grant P01-DC00119National Institutes of Health Grant T32-DC00038National Institutes of Health Contract P01-DC00361National Institutes of Health Grant R01-DC00235National Institutes of Health Contract NO1-DC2240