Basilar Membrane Motion Research Articles

Tuning and Timing of Organ of Corti Vibrations at the Apex of the Intact Chinchilla Cochlea.

The goal of this research was to record sound-evoked vibrations in the organ of Corti at the apex of the intact cochlea of the chinchilla, an animal with a frequency hearing range similar to that of humans. Twelve adult anesthetized chinchillas of either sex were used. Measurements of sound-evoked vibrations of the intact organ of Corti (OoC) were performed using optical coherence tomography (OCT). Acoustic stimuli consisted of single tones of 1-s duration. OoC vibrations were recorded using a Telesto Spectral Domain OCT system (Thorlabs GmbH, Germany) and ThorImage® OCT version 5.4.2 (Thorlabs GmbH, Germany). Further analysis of the output of the ThorImage software was performed by ad hoc programs written using Matlab® R2020b. Recordings were performed at several OoC sites extending from the Hensen's cell region to the vicinity of the basilar membrane (BM). The measurement angle between the optical axis of the OCT system and the BM was approximately 45°. Under that experimental condition, delays among the different OoC locations indicate that BM motion occurs earlier than at other sites. At all OoC sites, sound-evoked vibrations grow nonlinearly with stimulus level at compressive rates. The sharpness of tuning of OoC vibrations increases with stimulus level and death. Iso-velocity curves as a function of frequency are well-tuned around 500 Hz and closely resemble threshold tuning curves of chinchilla auditory-nerve fibers with similar characteristic frequencies. The nonlinear processing of sounds at the apex of the chinchilla cochlea differs significantly from the processing of sounds at the base of the cochlea in the same species.

Neural Coincidence Detection Strategies during Perception of Multi-Pitch Musical Tones

Multi-pitch perception is investigated in a listening test using 30 recordings of musical sounds with two tones played simultaneously, except for two gong sounds with inharmonic overtone spectra, judging roughness and separateness as the ability to tell the two tones in each recording apart. Of the sounds, 13 were from a Western guitar playing all 13 intervals in one octave, the other sounds were mainly from non-Western instruments, comparing familiar with unfamiliar instrument sounds for Western listeners. Additionally the sounds were processed in a cochlea model, transferring the mechanical basilar membrane motion into neural spikes followed by post-processing simulating different degrees of coincidence detection. Separateness perception showed a clear distinction between familiar and unfamiliar sounds, while roughness perception did not. By correlating perception with simulation different perception strategies were found. Familiar sounds correlated strongly positively with high degrees of coincidence detection, where only 3–5 periodicities were left, while unfamiliar sounds correlated with low coincidence levels. This corresponds to an attention to pitch and timbre, respectively. Additionally, separateness perception showed an opposite correlation between perception and neural correlates between familiar and unfamiliar sounds. This correlates with the perceptional finding of the distinction between familiar and unfamiliar sounds with separateness.

Regional differences in cochlear nonlinearity across the basal organ of Corti of gerbil: Regional differences in cochlear nonlinearity

Auditory sensation is based in nanoscale vibration of the sensory tissue of the cochlea, the organ of Corti complex (OCC). Motion within the OCC is now observable due to optical coherence tomography. In a previous study (Cooper et al., 2018), the region that includes the electro-motile outer hair cells (OHC) and Deiters cells (DC) was observed to move with larger amplitude than the basilar membrane (BM) and surrounding regions and was termed the "hotspot." In addition to this quantitative distinction, the hotspot moved qualitatively differently than the BM, in that its motion scaled nonlinearly with stimulus level at all frequencies, evincing sub-BF activity. Sub-BF activity enhances non-BF motion; thus the frequency tuning of the OHC/DC region was reduced relative to the BM. In this work we further explore the motion of the gerbil basal OCC and find that regions that lack significant sub-BF activity include the BM, the medial and lateral OCC, and the reticular lamina (RL) region. The observation that the RL region does not move actively sub-BF (already observed in Cho and Puria 2022), suggests that hair cell stereocilia are not exposed to sub-BF activity in the cochlear base. The observation that the lateral and RL regions move approximately linearly sub-BF indicates that linear forces dominate non-linear OHC-based forces on these components at sub-BF frequencies. A complex difference analysis was performed to reveal the internal motion of the OHC/DC region and showed that amplitude structure and phase shifts in the directly measured OHC/DC motion emerge due to the internal OHC/DC motion destructively interfering with BM motion.

Brain-Specific Angiogenesis Inhibitor 3 Is Expressed in the Cochlea and Is Necessary for Hearing Function in Mice.

Mammalian auditory hair cells transduce sound-evoked traveling waves in the cochlea into nerve stimuli, which are essential for hearing function. Pillar cells located between the inner and outer hair cells are involved in the formation of the tunnel of Corti, which incorporates outer-hair-cell-driven fluid oscillation and basilar membrane movement, leading to the fine-tuned frequency-specific perception of sounds by the inner hair cells. However, the detailed molecular mechanism underlying the development and maintenance of pillar cells remains to be elucidated. In this study, we examined the expression and function of brain-specific angiogenesis inhibitor 3 (Bai3), an adhesion G-protein-coupled receptor, in the cochlea. We found that Bai3 was expressed in hair cells in neonatal mice and pillar cells in adult mice, and, interestingly, Bai3 knockout mice revealed the abnormal formation of pillar cells, with the elevation of the hearing threshold in a frequency-dependent manner. Furthermore, old Bai3 knockout mice showed the degeneration of hair cells and spiral ganglion neurons in the basal turn. The results suggest that Bai3 plays a crucial role in the development and/or maintenance of pillar cells, which, in turn, are necessary for normal hearing function. Our results may contribute to understanding the mechanisms of hearing loss in human patients.

Intracochlear overdrive: Characterizing nonlinear wave amplification in the mouse apex.

In this study, we explore nonlinear cochlear amplification by analyzing basilar membrane (BM) motion in the mouse apex. Through in vivo, postmortem, and mechanical suppression recordings, we estimate how the cochlear amplifier nonlinearly shapes the wavenumber of the BM traveling wave, specifically within a frequency range where the short-wave approximation holds. Our findings demonstrate that a straightforward mathematical model, depicting the cochlear amplifier as a wavenumber modifier with strength diminishing monotonically as BM displacement increases, effectively accounts for the various experimental observations. This empirically derived model is subsequently incorporated into a physics-based "overturned" framework of cochlear amplification [see Altoè, Dewey, Charaziak, Oghalai, and Shera (2022), J. Acoust. Soc. Am. 152, 2227-2239] and tested against additional experimental data. Our results demonstrate that the relationships established within the short-wave region remain valid over a much broader frequency range. Furthermore, the model, now exclusively calibrated to BM data, predicts the behavior of the opposing side of the cochlear partition, aligning well with recent experimental observations. The success in reproducing key features of the experimental data and the mathematical simplicity of the resulting model provide strong support for the "overturned" theory of cochlear amplification.

High level sound transmission through cadaver human ears—On the influence of bone conduction and hearing protective devices

High level sound exposure can cause substantial injury to the auditory system, motivating efforts to predict and prevent this injury. Measurement techniques using acoustic manikins are effective for low and moderate sound levels, but nonlinear effects in the middle ear and alternate sound transmission pathways to the inner ear limit their utility at higher sound pressure levels. Here, we describe results from a series of measurements made in cadaveric human ears conducted in our laboratory over the last several years. We quantified sound transmission to the inner ear by measuring the difference in sound pressure level across the cochlear partition near the base of the cochlea, which drives basilar membrane motion. We describe measurements quantifying sound transmission through the human middle ear, which is limited by suspensory ligaments for sounds above approximately 130 dB SPL, resulting in nonlinear (harmonic) distortion of sounds in the cochlea at higher sound pressure levels. Similarly, we describe observations of sound transmission to the cochlea via bone conduction from vibratory transducers, as well as high level sound and impulse noise/blast exposures. The implications of these measurements for hearing loss predictions, effectiveness of hearing protective devices, and injury mechanisms are discussed.

Cochlear amplification in the short-wave region by outer hair cells changing organ-of-Corti area to amplify the fluid traveling wave

Many details of the operation of the mammalian cochlea are known, but how they all work together to produce cochlear amplification is not understood. Outer-hair-cell (OHC) motility produces two kinds of amplification: non-propagating amplification (NPA) that is from local OHCs, and traveling-wave amplification (TWA) that increases basilar-membrane (BM) motion. Proposed here are a series of hypotheses that provide a new explanation, the “OoC-area-pump”, for TWA: (1) In the short-wave region OHC vibrations cause cyclic longitudinal motion of fluid in the organ of Corti (OoC) and peri‑Deiters-cell tissue, (2) the longitudinal motion changes the local OoC area, which (3) by reticular-lamina (RL) movement drives the fluid in scala media in a way that amplifies the fluid-pressure traveling wave. (4) At the NPA-TWA changeover frequency, an abrupt change in the OoC frequency-wavenumber relationship is due to positive feedback between TWA and the mode of cochlear motion that is dominant, aided by focusing of the pressure traveling wave. It is hypothesized that OoC radial expansion and radial force from the Deiters-cell phalangeal process act to advance RL and/or lateral-compartment phase. Finally, it is hypothesized that human and lab-animal frequency tuning have similar bandwidths in distance along the cochlea because their traveling-wave wavelengths are similar in the corresponding short-wave regions. Experiments are needed to test these hypotheses and to determine for TWA whether the OoC-area-pump hypothesis replaces or supplements the “OHCs-act-on-BM” hypothesis. Several tests are outlined that can be done with current methodology. A key step in the evolution of mammalian hearing was the development of the complex OoC anatomy, including Deiters cells and OoC fluid spaces that allow local wide-band NPA to produce TWA that enables small local increments of gain to accumulate in the traveling wave and sharpen tuning.

Revealing the actions of the human cochlear basilar membrane at low frequency

The basilar membrane (BM) is the key infrastructure that supports the microstructure of cochlear acoustic function, and its interaction with lymph in the cochlea involves a complex, highly nonlinear coupling motion and energy conversion. In 1928, Nobel Laureate Gordon von Békésy experimented and first discovered the traveling wave motion of the BM, thus uncovering the mystery of BM motion. However, with the further development of experimental technology in recent years, scientists have discovered that the traveling wave motion does not explain many experimental observations and phenomena associated with the detection of low-frequency sounds by the cochlea. Because the cochlea is very small and complex, Békésy could only obtain medium- and high-frequency motion data for the BM but not low-frequency motion data. Based on a theory of mathematical physics and biology combined with data from medical and modern light source imaging experiments, a theoretical model of the spiral cochlea and a numerical model that conforms to the real human ear were established. The results reproduce the known traveling wave motion of the BM. Meanwhile, an exciting new finding has revealed a standing wave motion pattern at low frequencies. This newly discovered motion pattern intrinsically explains many experimental observations that could not be explained by the traveling wave theory. These results not only complement the low-frequency motion characteristics of the BM, but also probably reveal the mechanism of active phonoreceptive amplification in the cochlea, which has been a difficult problem to unravel in otology medicine.

유체-구조 연성을 고려한 기저막 운동에서 와우공 크기의 영향

유체-구조 연성을 고려한 기저막 운동에서 와우공 크기의 영향

Significance of the Microfluidic Flow Inside the Organ of Corti.

We study the vibration modes of a short section in the middle turn of the gerbil cochlea including both longitudinal and radial interstitial fluid spaces between the pillar cells (PC) and the sensory hair cells to determine the role of the interstitial fluid flow within the organ of corti (OoC). Three detailed finite element (FE) models of the cochlear short section (CSS) are studied. In model 1, the CSS is without fluids; model 2 includes the OoC fluid, but not the exterior scalae fluids; and model 3 is the CSS with both scalae and OoC fluids. We find that: (1) the fundamental mode shape of models 1 or 3 is similar to the classical basilar membrane (BM) bending mode that includes pivoting of the arch of corti, and hence determines the low frequency vibrational mode shape of the cochlea in the presence of the cochlear wave. (2) The fundamental mode shape of model 2 is characterized by a cross-sectional shape change similar to the passive response of the cochlea. This mode shape includes a tilting motion of the inner hair cell (IHC) region, a fluid motion within the tunnel of corti (ToC) in the radial direction and along the OoC, and a bulging motion of the reticular lamina (RL) above the outer hair cell (OHC). Each of these motions provides a plausible mode of excitation of the sensory hair cells. (3) The higher vibrational modes of model 1 are similar to the electrically evoked response within the OoC and suggests that the higher vibrational modes are responsible for the active response of the cochlea. We also observed that the fluid flow through the OoC interstitial space is significant, and the model comparison suggests that the OoC fluid contributes to the biphasic BM motion seen in electrical stimulation experiments. The effect of fluid viscosity on cilium deflection was assessed by performing a transient analysis to calculate the cilium shearing gain. The gain values are found to be within the range of experimentally measured values reported by Dallos et al. (1996, The Cochlea, Springer-Verlag, New York).

Effects of Efferent Activity on Hair Bundle Mechanics.

Hair cells in both the auditory and vestibular systems receive efferent innervation. A number of prior studies have indicated that efferent regulation serves to diminish the overall sensitivity of the auditory system. The efferent pathway is believed to affect the sensitivity and frequency selectivity of the hair cell by modulating its membrane potential. However, its effect on the mechanical response of the hair cell has not been established. We explored how stimulation of the efferent neurons affects the mechanical responsiveness of an individual hair bundle. We tested this effect on in vitro preparations of hair cells in the sacculi of American bullfrogs of both genders. Efferent stimulation routinely resulted in an immediate increase of the frequency of hair bundle spontaneous oscillations for the duration of the stimulus. Enlarging the stimulus amplitude and pulse length, or conversely, decreasing the interpulse interval led to oscillation suppression. Additionally, we tested the effects of efference on the hair bundle response to mechanical stimulation. The receptive field maps of hair cells undergoing efferent actuation demonstrated an overall desensitization with respect to those of unstimulated cells.SIGNIFICANCE STATEMENT The efferent system is an important aide for the performance of the auditory system. It has been seen to contribute to sound detection and localization, ototoxicity prevention, and speech comprehension. Although measurements have demonstrated that efference suppresses basilar membrane movement, there is still much unknown about how efferent activity affects hearing mechanics. Here, we explore the mechanical basis for the efferent system's capabilities at the level of the hair bundle. We present optical recordings, receptive field maps, and sensitivity curves that show a hair bundle is desensitized by efferent stimulation. This supports the hypothesis that efferent regulation may be a biological control parameter for tuning the hair bundle's mechanical sensitivity.

Nonlinear cochlear mechanics without direct vibration-amplification feedback.

Recent in vivo recordings from the mammalian cochlea indicate that although the motion of the basilar membrane appears actively amplified and nonlinear only at frequencies relatively close to the peak of the response, the internal motions of the organ of Corti display these same features over a much wider range of frequencies. These experimental findings are not easily explained by the textbook view of cochlear mechanics, in which cochlear amplification is controlled by the motion of the basilar membrane (BM) in a tight, closed-loop feedback configuration. This study shows that a simple phenomenological model of the cochlea inspired by the work of Zweig [J. Acoust. Soc. Am. 138, 1102 (2015)] can account for recent data in mouse and gerbil. In this model, the active forces are regulated indirectly, through the effect of BM motion on the pressure field across the cochlear partition, rather than via direct coupling between active-force generation and BM vibration. The absence of strong vibration-amplification feedback in the cochlea also provides a compelling explanation for the observed intensity invariance of fine time structure in the BM response to acoustic clicks.

Nonlinearity and amplification in cochlear responses to single and multi-tone stimuli

Mechanical displacements of the basilar membrane (BM) and the electrophysiological responses of the auditory outer hair cells (OHCs) are key components of the frequency tuning and cochlear amplification in the mammalian cochlea. In the work presented here, we measured the responses of (1) the extracellular voltage generated by OHCs (VOHC) and (2) displacements within the organ of Corti complex (OCC) to a multi-tone stimulus, and to single tones. Using optical coherence tomography (OCT), we were able to measure displacements of different layers in the OCC simultaneously, in the base of the gerbil cochlea. We explored the effect of the two types of sound stimuli to the nonlinear behavior of voltage and displacement in two frequency regions: a frequency region below the BM nonlinearity (sub-BF region: f < ∼0.7 BF), and in the best frequency (BF) region. In the sub-BF region, BM motion (XBM) had linear growth for both stimulus types, and the motion in the OHC region (XOHC) was mildly nonlinear for single tones, and relatively strongly nonlinear for multi-tones. Sub-BF, the nonlinear character of VOHC was similar to that of XOHC. In the BF region XBM, VOHC and XOHC all possessed the now-classic nonlinearity of the BF peak. Coupling these observations with previous findings on phasing between OHC force and traveling wave motions, we propose the following framework for cochlear nonlinearity: The BF-region nonlinearity is an amplifying nonlinearity, in which OHC forces input power into the traveling wave, allowing it to travel further apical to the region where it peaks. The sub-BF nonlinearity is a non-amplifying nonlinearity; it represents OHC electromotility, and saturates due to OHC current saturation, but the OHC forces do not possess the proper phasing to feed power into the traveling wave.

Cochlear compound action potentials from high-level tone bursts originate from wide cochlear regions that are offset toward the most sensitive cochlear region.

Little is known about the spatial origins of auditory nerve (AN) compound action potentials (CAPs) evoked by moderate to intense sounds. We studied the spatial origins of AN CAPs evoked by 2- to 16-kHz tone bursts at several sound levels by slowly injecting kainic acid solution into the cochlear apex of anesthetized guinea pigs. As the solution flowed from apex to base, it sequentially reduced CAP responses from low- to high-frequency cochlear regions. The times at which CAPs were reduced, combined with the cochlear location traversed by the solution at that time, showed the cochlear origin of the removed CAP component. For low-level tone bursts, the CAP origin along the cochlea was centered at the characteristic frequency (CF). As sound level increased, the CAP center shifted basally for low-frequency tone bursts but apically for high-frequency tone bursts. The apical shift was surprising because it is opposite the shift expected from AN tuning curve and basilar membrane motion asymmetries. For almost all high-level tone bursts, CAP spatial origins extended over 2 octaves along the cochlea. Surprisingly, CAPs evoked by high-level low-frequency (including 2 kHz) tone bursts showed little CAP contribution from CF regions ≤ 2 kHz. Our results can be mostly explained by spectral splatter from the tone-burst rise times, excitation in AN tuning-curve "tails," and asynchronous AN responses to high-level energy ≤ 2 kHz. This is the first time CAP origins have been identified by a spatially specific technique. Our results show the need for revising the interpretation of the cochlear origins of high-level CAPs-ABR wave 1. NEW & NOTEWORTHY Cochlear compound action potentials (CAPs) and auditory brain stem responses (ABRs) are routinely used in laboratories and clinics. They are typically interpreted as arising from the cochlear region tuned to the stimulus frequency. However, as sound level is increased, the cochlear origins of CAPs from tone bursts of all frequencies become very wide and their centers shift toward the most sensitive cochlear region. The standard interpretation of CAPs and ABRs from moderate to intense stimuli needs revision.

Amplification and Suppression of Traveling Waves along the Mouse Organ of Corti: Evidence for Spatial Variation in the Longitudinal Coupling of Outer Hair Cell-Generated Forces.

Mammalian hearing sensitivity and frequency selectivity depend on a mechanical amplification process mediated by outer hair cells (OHCs). OHCs are situated within the organ of Corti atop the basilar membrane (BM), which supports sound-evoked traveling waves. It is well established that OHCs generate force to selectively amplify BM traveling waves where they peak, and that amplification accumulates from one location to the next over this narrow cochlear region. However, recent measurements demonstrate that traveling waves along the apical surface of the organ of Corti, the reticular lamina (RL), are amplified over a much broader region. Whether OHC forces accumulate along the length of the RL traveling wave to provide a form of "global" cochlear amplification is unclear. Here we examined the spatial accumulation of RL amplification. In mice of either sex, we used tones to suppress amplification from different cochlear regions and examined the effect on RL vibrations near and far from the traveling-wave peak. We found that although OHC forces amplify the entire RL traveling wave, amplification only accumulates near the peak, over the same region where BM motion is amplified. This contradicts the notion that RL motion is involved in a global amplification mechanism and reveals that the mechanical properties of the BM and organ of Corti tune how OHC forces accumulate spatially. Restricting the spatial buildup of amplification enhances frequency selectivity by sharpening the peaks of cochlear traveling waves and constrains the number of OHCs responsible for mechanical sensitivity at each location.SIGNIFICANCE STATEMENT Outer hair cells generate force to amplify traveling waves within the mammalian cochlea. This force generation is critical to the ability to detect and discriminate sounds. Nevertheless, how these forces couple to the motions of the surrounding structures and integrate along the cochlear length remains poorly understood. Here we demonstrate that outer hair cell-generated forces amplify traveling-wave motion on the organ of Corti throughout the wave's extent, but that these forces only accumulate longitudinally over a region near the wave's peak. The longitudinal coupling of outer hair cell-generated forces is therefore spatially tuned, likely by the mechanical properties of the basilar membrane and organ of Corti. Our findings provide new insight into the mechanical processes that underlie sensitive hearing.

Efferent Inhibition of the Cochlea.

Cholinergic efferent neurons originating in the brainstem innervate the acoustico-lateralis organs (inner ear, lateral line) of vertebrates. These release acetylcholine (ACh) to inhibit hair cells through activation of calcium-dependent potassium channels. In the mammalian cochlea, ACh shunts and suppresses outer hair cell (OHC) electromotility, reducing the essential amplification of basilar membrane motion. Consequently, medial olivocochlear neurons that inhibit OHCs reduce the sensitivity and frequency selectivity of afferent neurons driven by cochlear vibration of inner hair cells (IHCs). The cholinergic synapse on hair cells involves an unusual ionotropic ACh receptor, and a near-membrane postsynaptic cistern. Lateral olivocochlear (LOC) neurons modulate type I afferents by still-to-be-defined synaptic mechanisms. Olivocochlear neurons can be activated by a reflex arc that includes the auditory nerve and projections from the cochlear nucleus. They are also subject to modulation by higher-order central auditory interneurons. Through its actions on cochlear hair cells, afferent neurons, and higher centers, the olivocochlear system protects against age-related and noise-induced hearing loss, improves signal coding in noise under certain conditions, modulates selective attention to sensory stimuli, and influences sound localization.

Cochlear amplification and tuning depend on the cellular arrangement within the organ of Corti

The field of cochlear mechanics has been undergoing a revolution due to recent findings made possible by advancements in measurement techniques. While it has long been assumed that basilar-membrane (BM) motion is the most important determinant of sound transduction by the inner hair cells (IHCs), it turns out that other parts of the sensory epithelium closer to the IHCs, such as the reticular lamina (RL), move with significantly greater amplitude for weaker sounds. It has not been established how these findings are related to the complex cytoarchitecture of the organ of Corti between the BM and RL, which is composed of a lattice of asymmetric Y-shaped elements, each consisting of a basally slanted outer hair cell (OHC), an apically slanted phalangeal process (PhP), and a supporting Deiters' cell (DC). Here, a computational model of the mouse cochlea supports the hypothesis that the OHC micromotors require this Y-shaped geometry for their contribution to the exquisite sensitivity and frequency selectivity of the mammalian cochlea. By varying only the OHC gain parameter, the model can reproduce measurements of BM and RL gain and tuning for a variety of input sound levels. Malformations such as reversing the orientations of the OHCs and PhPs or removing the PhPs altogether greatly reduce the effectiveness of the OHC motors. These results imply that the DCs and PhPs must be properly accounted for in emerging OHC regeneration therapies.

Real-time intracochlear imaging of automated cochlear implant insertions in whole decalcified cadaver cochleas using ultrasound

Objectives: This study aimed to determine the feasibility of combining high-frequency ultrasound imaging, automated insertion, and force sensing to yield more information about cochlear implant insertion dynamics.Methods: An apparatus was developed combining these aspects along with software to control implant and imaging probe positions. Decalcified unfixed human cochleas were implanted at various speeds, insertion sites, and implant models while imaging near the implant tip throughout insertion and recording force data from the cochlea mounting stage. Ultrasound video data were also captured.Results: The basilar membrane (BM) was frequently penetrated by the implant in either the mid-basal or lower middle turn. Measurements were also performed of apical BM motion in response to upstream implant movement at varying insertion speeds. Increasing insertion speed resulted in greater BM displacement.Discussion: Multiple insertions per cochlea increase the volume of data per specimen while also reducing variability due to differences between cochleas. However, to image inside the cochlea with ultrasound, the bone had to be decalcified, which likely had a significant effect upon the response of tissue to contact by the implant. As calcified bone strongly reflects ultrasound, we also found ultrasound imaging to be an excellent method for easily assessing bone decalcification progress.Conclusion: This technique may be very useful for some studies, although the confounding effects of bone decalcification may make results of other studies too difficult to generalize. The approach could be adapted to other real-time imaging modalities, such as optical coherence tomography.

Temporal Suppression of Clicked-Evoked Otoacoustic Emissions and Basilar-Membrane Motion in Gerbils.

Otoacoustic emissions evoked by a click (CEOAEs) can be reduced if the evoking sound is preceded or followed by another ("suppressor") click. Studying the temporal suppression of CEOAEs can provide insights into the dynamics of cochlear nonlinearity. However, temporal suppression has never been measured in basilar-membrane (BM) motion. Thus, it remains unclear whether the characteristics of CEOAE temporal suppression are reflected in BM responses. Here we simultaneously measured ear-canal pressure and BM velocity (laser Doppler vibrometry) in response to clicks in gerbil. When the suppressor click preceded the evoking click, CEOAEs were maximally suppressed for interclick intervals (ICIs) equivalent to ~2 periods of the analyzed frequency (9-14 kHz). Maximal temporal suppression at nonzero ICIs has been previously observed in human CEOAEs. BM responses to clicks were maximally reduced when the suppressor click preceded the evoking one by ~1 period of the characteristic frequency (CF ~14 kHz). Thus, the "delayed" characteristics of CEOAE temporal suppression are reflected in BM motion, although on a different time scale. When the suppressor click followed the evoking click, CEOAEs were augmented rather than suppressed, while enhancement was not observed in BM motion at the CF. This result indicates that some aspects of CEOAE temporal suppression are intrinsic to CEOAE generation mechanisms and/or to places that are not reflected in a BM motion at a single-location.

Non-tip auditory-nerve responses that are suppressed by low-frequency bias tones originate from reticular lamina motion

Recent cochlear mechanical measurements show that active processes increase the motion response of the reticular lamina (RL) at frequencies more than an octave below the local characteristic frequency (CF) for CFs above 5 kHz. A possible correlate is that in high-CF (>5 kHz) auditory-nerve (AN) fibers, responses to frequencies 1-3 octaves below CF (“tail” frequencies) can be inhibited by medial olivocochlear (MOC) efferents. These results indicate that active processes enhance the sensitivity of tail-frequency RL and AN responses. Perhaps related is that some apical low-CF AN fibers have tuning-curve (TC) “side-lobe” response areas at frequencies above and below the TC-tip that are MOC inhibited. We hypothesized that the tail and side-lobe responses are enhanced by the same active mechanisms as CF cochlear amplification. If responses to CF, tail-frequency, and TC-side-lobe tones are all enhanced by prestin motility controlled by outer-hair-cell (OHC) transmembrane voltage, then they should depend on OHC stereocilia position in the same way. To test this, we cyclically changed the OHC-stereocilia mechano-electric-transduction (MET) operating point with low-frequency “bias” tones (BTs) and increased the BT level until the BT caused quasi-static OHC MET saturation that reduced or “suppressed” the gain of OHC active processes. While measuring cat AN-fiber responses, 50 Hz BT level series, 70–120 dB SPL, were run alone and with CF tones, or 2.5 kHz tail-frequency tones, or side-lobe tones. BT-tone-alone responses were used to exclude BT sound levels that produced AN responses that might obscure BT suppression. Data were analyzed to show the BT phase that suppressed the tone responses at the lowest sound level. We found that AN responses to CF, tail-frequency, and side-lobe tones were suppressed at the same BT phase in almost all cases. The data are consistent with the enhancement of responses to CF, tail-frequency, and side-lobe tones all being due to the same OHC-stereocilia MET-dependent active process. Thus, OHC active processes enhance AN responses at frequencies outside of the cochlear-amplified TC-tip region in both high- and low-frequency cochlear regions. The data are consistent with the AN response enhancements being due to enhanced RL motion that drives IHC-stereocilia deflection by traditional RL-TM shear and/or by changing the RL-TM gap. Since tail-frequency basilar membrane (BM) motion is not actively enhanced, the tail-frequency IHC drive is from a vibrational mode little present on the BM, not a “second filter” of BM motion.