The impact of size on middle-ear sound transmission in elephants, the largest terrestrial mammal.

Elephants have a unique auditory system that is larger than any other terrestrial mammal. To quantify the impact of larger middle ear (ME) structures, we measured 3D ossicular motion and ME sound transmission in cadaveric temporal bones from both African and Asian elephants in response to air-conducted (AC) tonal pressure stimuli presented in the ear canal (PEC). Results were compared to similar measurements in humans. Velocities of the umbo (VU) and stapes (VST) were measured using a 3D laser Doppler vibrometer in the 7-13,000 Hz frequency range, stapes velocity serving as a measure of energy entering the cochlea-a proxy for hearing sensitivity. Below the elephant ME resonance frequency of about 300 Hz, the magnitude of VU/PEC was an order of magnitude greater than in human, and the magnitude of VST/PEC was 5x greater. Phase of VST/PEC above ME resonance indicated that the group delay in elephant was approximately double that of human, which may be related to the unexpectedly high magnitudes at high frequencies. A boost in sound transmission across the incus long process and stapes near 9 kHz was also observed. We discuss factors that contribute to differences in sound transmission between these two large mammals.

Measurements of bone-conducted sound in the chinchilla external ear.

We measure bone-conduction (BC) induced skull velocity, sound pressure at the tympanic membrane (TM) and inner-ear compound-action potentials (CAP) before and after manipulating the ear canal, ossicles, and the jaw to investigate the generation of BC induced ear-canal sound pressures and their contribution to inner-ear BC response in the ears of chinchillas. These measurements suggest that in chinchilla: i.) Vibrations of the bony ear canal walls contribute significantly to BC-induced ear canal sound pressures, as occluding the ear canal at the bone-cartilaginous border causes a 10 dB increase in sound pressure at the TM (PTM) at frequencies below 2 kHz. ii.) The contributions to PTM of ossicular and TM motions when driven in reverse by BC-induced inner-ear sound pressures are small. iii.) The contribution of relative motions of the jaw and ear canal to PTM is small. iv.) Comparison of the effect of canal occlusion on PTM and CAP thresholds point out that BC-induced ear canal sound pressures contribute significantly to bone-conduction stimulation of the inner ear when the ear canal is occluded.

The impact of size on middle-ear sound transmission in elephants, the largest terrestrial mammal.

Elephants have a unique auditory system that is larger than any other terrestrial mammal. To quantify the impact of larger middle ear (ME) structures, we measured 3D ossicular motion and ME sound transmission in cadaveric temporal bones from both African and Asian elephants in response to air-conducted (AC) tonal pressure stimuli presented in the ear canal (P EC ). Results were compared to similar measurements in humans. Velocities of the umbo (V U ) and stapes (V ST ) were measured using a 3D laser Doppler vibrometer in the 7-13,000 Hz frequency range, stapes velocity serving as a measure of energy entering the cochlea-a proxy for hearing sensitivity. Below the elephant ME resonance frequency of about 300 Hz, the magnitude of V U /P EC was an order of magnitude greater than in human, and the magnitude of V ST /P EC was 5x greater. Phase of V ST /P EC above ME resonance indicated that the group delay in elephant was approximately double that of human, which may be related to the unexpectedly high magnitudes at high frequencies. A boost in sound transmission across the incus long process and stapes near 9 kHz was also observed. We discuss factors that contribute to differences in sound transmission between these two large mammals.

Human middle-ear muscle pulls change tympanic-membrane shape and low-frequency middle-ear transmission magnitudes and delays.

The three-bone flexible ossicular chain in mammals may allow independent alterations of middle-ear (ME) sound transmission via its two attached muscles, for both acoustic and non-acoustic stimuli. The tensor tympani (TT) muscle, which has its insertion on the malleus neck, is thought to increase tension of the tympanic membrane (TM). The stapedius (St) muscle, which has its insertion on the stapes posterior crus, is known to stiffen the stapes annular ligament. We produced ME changes in human cadaveric temporal bones by statically pulling on the TT and St muscles. The 3D static TM shape and sound-induced umbo motions from 20 Hz to 10 kHz were measured with optical coherence tomography (OCT); stapes motion was measured using laser-Doppler vibrometry (LDV). TT pulls made the TM shape more conical and moved the umbo medially, while St pulls moved the umbo laterally. In response to sound below about 1 kHz, stapes-velocity magnitudes generally decreased by about 10 dB due to TT pulls and 5 dB due to St pulls. In the 250 to 500 Hz region, the group delay calculated from stapes-velocity phase showed a decrease in transmission delay of about 150 µs by TT pulls and 60 µs by St pulls. Our interpretation of these results is that ME-muscle activity may provide a way of mechanically changing interaural time- and level-difference cues. These effects could help the brain align head-centered auditory and ocular-centered visual representations of the environment.

Preserving Wideband Tympanometry Information With Artifact Mitigation.

OBJECTIVE: Absorbance measured using wideband tympanometry (WBT) has been shown to be sensitive to changes in middle and inner ear mechanics, with potential to diagnose various mechanical ear pathologies. However, artifacts in absorbance due to measurement noise can obscure information related to pathologies and increase intermeasurement variability. Published reports frequently present absorbance that has undergone smoothing to minimize artifact; however, smoothing changes the true absorbance and can destroy important narrow-band characteristics such as peaks and notches at different frequencies. Because these characteristics can be unique to specific pathologies, preserving them is important for diagnostic purposes. Here, we identify the cause of artifacts in absorbance and develop a technique to mitigate artifacts while preserving the underlying WBT information. DESIGN: A newly developed Research Platform for the Interacoustics Titan device allowed us to study raw microphone recordings and corresponding absorbances obtained by WBT measurements. We investigated WBT measurements from normal hearing ears and ears with middle and inner ear pathologies for the presence of artifact and noise. Furthermore, it was used to develop an artifact mitigation procedure and to evaluate its effectiveness in mitigating artifacts without distorting the true WBT information. RESULTS: We observed various types of noise that can plague WBT measurements and that contribute to artifacts in computed absorbances, particularly intermittent low-frequency noise. We developed an artifact mitigation procedure that incorporates a high-pass filter and a Tukey window. This artifact mitigation resolved the artifacts from low-frequency noise while preserving characteristics in absorbance in both normal hearing ears and ears with pathology. Furthermore, the artifact mitigation reduced intermeasurement variability. CONCLUSIONS: Unlike smoothing algorithms used in the past, our artifact mitigation specifically removes artifacts caused by noise. It does not change frequency response characteristics, such as narrow-band peaks and notches in absorbance at different frequencies that can be important for diagnosis. Also, by reducing intermeasurement variability, the artifact mitigation can improve the test-retest reliability of these measurements.

Mechanical overstimulation causes acute injury and synapse loss followed by fast recovery in lateral-line neuromasts of larval zebrafish.

Excess noise damages sensory hair cells, resulting in loss of synaptic connections with auditory nerves and, in some cases, hair-cell death. The cellular mechanisms underlying mechanically induced hair-cell damage and subsequent repair are not completely understood. Hair cells in neuromasts of larval zebrafish are structurally and functionally comparable to mammalian hair cells but undergo robust regeneration following ototoxic damage. We therefore developed a model for mechanically induced hair-cell damage in this highly tractable system. Free swimming larvae exposed to strong water wave stimulus for 2 hr displayed mechanical injury to neuromasts, including afferent neurite retraction, damaged hair bundles, and reduced mechanotransduction. Synapse loss was observed in apparently intact exposed neuromasts, and this loss was exacerbated by inhibiting glutamate uptake. Mechanical damage also elicited an inflammatory response and macrophage recruitment. Remarkably, neuromast hair-cell morphology and mechanotransduction recovered within hours following exposure, suggesting severely damaged neuromasts undergo repair. Our results indicate functional changes and synapse loss in mechanically damaged lateral-line neuromasts that share key features of damage observed in noise-exposed mammalian ear. Yet, unlike the mammalian ear, mechanical damage to neuromasts is rapidly reversible.

Sound pressure distribution within human ear canals: II. Reverse mechanical stimulation.

This work is part of a study of the interactions of ear canal (EC) sound with tympanic membrane (TM) surface displacements. In human temporal bones, the ossicles were stimulated mechanically "in reverse" to mimic otoacoustic emissions (OAEs), and the sound field within the ear canal was sampled with 0.5-2 mm spacing near the TM surface and at more distal locations within the EC, including along the longitudinal EC axis. Sound fields were measured with the EC open or occluded. The reverse-driven sound field near the TM had larger and more irregular spatial variations below 10 kHz than with forward sound stimulation, consistent with a significant contribution of nonuniform sound modes. These variations generally did not propagate more than ∼4 mm laterally from the TM. Longitudinal sound field variations with the EC open or blocked were consistent with standing-wave patterns in tubes with open or closed ends. Relative contributions of the nonuniform components to the total sound pressure near the TM were largest at EC natural frequencies where the longitudinal component was small. Transverse variations in EC sound pressure can be reduced by reducing longitudinal EC sound pressure variations, e.g., via reducing reflections from occluding earplugs.

Tympanic membrane surface motions in forward and reverse middle ear transmissions.

Characterization of Tympanic Membrane (TM) surface motions with forward and reverse stimulation is important to understanding how the TM transduces acoustical and mechanical energy in both directions. In this paper, stroboscopic opto-electronic holography is used to quantify motions of the entire TM surface induced by forward sound and reverse mechanical stimulation in human cadaveric ears from 0.25 to 18.4 kHz. The forward sound stimulus was coupled to an anatomically realistic artificial ear canal that allowed optical access to the entire TM surface, and the reverse mechanical stimulus was applied to the body of the incus by a piezo-electric stimulator. The results show clear differences in TM surface motions evoked by the two stimuli. In the forward case, TM motion is dominated by standing-wave-like modal motions that are consistent with a relatively uniform sound-pressure load over the entire TM surface. With reverse mechanical stimulation, the TM surface shows more traveling waves, consistent with a localized mechanical drive applied to the manubrium embedded in the TM. With both stimuli, the manubrium moves less than the rest of the TM, consistent with the TM acting like a compliant membrane rather than a stiff diaphragm, and also consistent with catenary behavior due to the TM's curved shape.

Mapping the phase and amplitude of ossicular chain motion using sound-synchronous optical coherence vibrography.

The sound-driven vibration of the tympanic membrane and ossicular chain of middle-ear bones is fundamental to hearing. Here we show that optical coherence tomography in phase synchrony with a sound stimulus is well suited for volumetric, vibrational imaging of the ossicles and tympanic membrane. This imaging tool - OCT vibrography - provides intuitive motion pictures of the ossicular chain and how they vary with frequency. Using the chinchilla ear as a model, we investigated the vibrational snapshots and phase delays of the manubrium, incus, and stapes over 100 Hz to 15 kHz. The vibrography images reveal a previously undescribed mode of motion of the chinchilla ossicles at high frequencies.

Chinchilla middle ear transmission matrix model and middle-ear flexibility.

The function of the middle ear (ME) in transforming ME acoustic inputs and outputs (sound pressures and volume velocities) can be described with an acoustic two-port transmission matrix. This description is independent of the load on the ME (cochlea or ear canal) and holds in either direction: forward (from ear canal to cochlea) or reverse (from cochlea to ear canal). A transmission matrix describing ME function in chinchilla, an animal commonly used in auditory research, is presented, computed from measurements of forward ME function: input admittance YTM, ME pressure gain GMEP, ME velocity transfer function HV, and cochlear input admittance YC, in the same set of ears [Ravicz and Rosowski (2012b). J. Acoust. Soc. Am. 132, 2437-2454; (2013a). J. Acoust. Soc. Am. 133, 2208-2223; (2013b). J. Acoust. Soc. Am. 134, 2852-2865]. Unlike previous estimates, these computations require no assumptions about the state of the inner ear, effectiveness of ME manipulations, or measurements of sound transmission in the reverse direction. These element values are generally consistent with physical constraints and the anatomical ME "transformer ratio." Differences from a previous estimate in chinchilla [Songer and Rosowski (2007). J. Acoust. Soc. Am. 122, 932-942] may be due to a difference in ME flexibility between the two subject groups.

Response of the human tympanic membrane to transient acoustic and mechanical stimuli: Preliminary results.

The response of the tympanic membrane (TM) to transient environmental sounds and the contributions of different parts of the TM to middle-ear sound transmission were investigated by measuring the TM response to global transients (acoustic clicks) and to local transients (mechanical impulses) applied to the umbo and various locations on the TM. A lightly-fixed human temporal bone was prepared by removing the ear canal, inner ear, and stapes, leaving the incus, malleus, and TM intact. Motion of nearly the entire TM was measured by a digital holography system with a high speed camera at a rate of 42 000 frames per second, giving a temporal resolution of <24 µs for the duration of the TM response. The entire TM responded nearly instantaneously to acoustic transient stimuli, though the peak displacement and decay time constant varied with location. With local mechanical transients, the TM responded first locally at the site of stimulation, and the response spread approximately symmetrically and circumferentially around the umbo and manubrium. Acoustic and mechanical transients provide distinct and complementary stimuli for the study of TM response. Spatial variations in decay and rate of spread of response imply local variations in TM stiffness, mass, and damping.

Restoration of middle-ear input in fluid-filled middle ears by controlled introduction of air or a novel air-filled implant.

The effect of small amounts of air on sound-induced umbo velocity in an otherwise saline-filled middle ear (ME) was investigated to examine the efficacy of a novel balloon-like air-filled ME implant suitable for patients with chronically non-aerated MEs. In this study, air bubbles or air-filled implants were introduced into saline-filled human cadaveric MEs. Umbo velocity, a convenient measure of ME response, served as an indicator of hearing sensitivity. Filling the ME with saline reduced umbo velocity by 25-30 dB at low frequencies and more at high frequencies, consistent with earlier work (Ravicz et al., Hear. Res. 195: 103-130 (2004)). Small amounts of air (∼30 µl) in the otherwise saline-filled ME increased umbo velocity substantially, to levels only 10-15 dB lower than in the dry ME, in a frequency- and location-dependent manner: air in contact with the tympanic membrane (TM) increased umbo velocity at all frequencies, while air located away from the TM increased umbo velocity only below about 500 Hz. The air-filled implant also affected umbo velocity in a manner similar to an air bubble of equivalent compliance. Inserting additional implants into the ME had the same effect as increasing air volume. These results suggest these middle-ear implants would significantly reduce conductive hearing loss in patients with chronically fluid-filled MEs.

Sound pressure distribution within natural and artificial human ear canals: forward stimulation.

This work is part of a study of the interaction of sound pressure in the ear canal (EC) with tympanic membrane (TM) surface displacement. Sound pressures were measured with 0.5-2 mm spacing at three locations within the shortened natural EC or an artificial EC in human temporal bones: near the TM surface, within the tympanic ring plane, and in a plane transverse to the long axis of the EC. Sound pressure was also measured at 2-mm intervals along the long EC axis. The sound field is described well by the size and direction of planar sound pressure gradients, the location and orientation of standing-wave nodal lines, and the location of longitudinal standing waves along the EC axis. Standing-wave nodal lines perpendicular to the long EC axis are present on the TM surface >11-16 kHz in the natural or artificial EC. The range of sound pressures was larger in the tympanic ring plane than at the TM surface or in the transverse EC plane. Longitudinal standing-wave patterns were stretched. The tympanic-ring sound field is a useful approximation of the TM sound field, and the artificial EC approximates the natural EC.

Middle-ear velocity transfer function, cochlear input immittance, and middle-ear efficiency in chinchilla.

The transfer function H(V) between stapes velocity V(S) and sound pressure near the tympanic membrane P(TM) is a descriptor of sound transmission through the middle ear (ME). The ME power transmission efficiency (MEE), the ratio of sound power entering the cochlea to power entering the middle ear, was computed from H(V) measured in seven chinchilla ears and previously reported measurements of ME input admittance Y(TM) and ME pressure gain G(MEP) [Ravicz and Rosowski, J. Acoust. Soc. Am. 132, 2437-2454 (2012); J. Acoust. Soc. Am. 133, 2208-2223 (2013)] in the same ears. The ME was open, and a pressure sensor was inserted into the cochlear vestibule for most measurements. The cochlear input admittance Y(C) computed from H(V) and G(MEP) is controlled by a combination of mass and resistance and is consistent with a minimum-phase system up to 27 kHz. The real part Re{Y(C)}, which relates cochlear sound power to inner-ear sound pressure, decreased gradually with frequency up to 25 kHz and more rapidly above that. MEE was about 0.5 between 0.1 and 8 kHz, higher than previous estimates in this species, and decreased sharply at higher frequencies.

Inner-ear sound pressures near the base of the cochlea in chinchilla: further investigation.

The middle-ear pressure gain GMEP, the ratio of sound pressure in the cochlear vestibule PV to sound pressure at the tympanic membrane PTM, is a descriptor of middle-ear sound transfer and the cochlear input for a given stimulus in the ear canal. GMEP and the cochlear partition differential pressure near the cochlear base ΔPCP, which determines the stimulus for cochlear partition motion and has been linked to hearing ability, were computed from simultaneous measurements of PV, PTM, and the sound pressure in scala tympani near the round window PST in chinchilla. GMEP magnitude was approximately 30 dB between 0.1 and 10 kHz and decreased sharply above 20 kHz, which is not consistent with an ideal transformer or a lossless transmission line. The GMEP phase was consistent with a roughly 50-µs delay between PV and PTM. GMEP was little affected by the inner-ear modifications necessary to measure PST. GMEP is a good predictor of ΔPCP at low and moderate frequencies where PV >> PST but overestimates ΔPCP above a few kilohertz where PV ≈ PST. The ratio of PST to PV provides insight into the distribution of sound pressure within the cochlear scalae.

Chinchilla middle-ear admittance and sound power: high-frequency estimates and effects of inner-ear modifications.

The middle-ear input admittance relates sound power into the middle ear (ME) and sound pressure at the tympanic membrane (TM). ME input admittance was measured in the chinchilla ear canal as part of a larger study of sound power transmission through the ME into the inner ear. The middle ear was open, and the inner ear was intact or modified with small sensors inserted into the vestibule near the cochlear base. A simple model of the chinchilla ear canal, based on ear canal sound pressure measurements at two points along the canal and an assumption of plane-wave propagation, enables reliable estimates of Y(TM,) the ME input admittance at the TM, from the admittance measured relatively far from the TM. Y(TM) appears valid at frequencies as high as 17 kHz, a much higher frequency than previously reported. The real part of Y(TM) decreases with frequency above 2 kHz. Effects of the inner-ear sensors (necessary for inner ear power computation) were small and generally limited to frequencies below 3 kHz. Computed power reflectance was ~0.1 below 3.5 kHz, lower than with an intact ME below 2.5 kHz, and nearly 1 above 16 kHz.

Middle ear function and cochlear input impedance in chinchilla.

Simultaneous measurements of middle ear-conducted sound pressure in the cochlear vestibule P(V) and stapes velocity V(S) have been performed in only a few individuals from a few mammalian species. In this paper, simultaneous measurements of P(V) and V(S) in six chinchillas are reported, enabling computation of the middle ear pressure gain G(ME) (ratio of P(V) to the sound pressure in the ear canal P(TM)), the stapes velocity transfer function SVTF (ratio of the product of V(S) and area of the stapes footplate A(FP) to P(TM)), and, for the first time, the cochlear input impedance Z(C) (ratio of P(V) to the product of V(S) and A(FP)) in individuals. mid R:G(ME)mid R: ranged from 25 to 35 dB over 125 Hz-8 kHz; the average group delay between 200 Hz and 10 kHz was about 52 mus. SVTF was comparable to that of previous studies. Z(C) was resistive from the lowest frequencies up to at least 10 kHz, with a magnitude on the order of 10(11) acoustic ohms. P(V), V(S), and the acoustic power entering the cochlea were good predictors of the shape of the audiogram at frequencies between 125 Hz and 2 kHz.

Middle-ear pressure gain and cochlear partition differential pressure in chinchilla.

An important step to describe the effects of inner-ear impedance and pathologies on middle- and inner-ear mechanics is to quantify middle- and inner-ear function in the normal ear. We present middle-ear pressure gain G(MEP) and trans-cochlear-partition differential sound pressure DeltaP(CP) in chinchilla from 100 Hz to 30 kHz derived from measurements of intracochlear sound pressures in scala vestibuli P(SV) and scala tympani P(ST) and ear-canal sound pressure near the tympanic membrane P(TM). These measurements span the chinchilla's auditory range. G(MEP) had constant magnitude of about 20 dB between 300 Hz and 20 kHz and phase that implies a 40-micros delay, values with some similarities to previous measurements in chinchilla and other species. DeltaP(CP) was similar to G(MEP) below about 10 kHz and lower in magnitude at higher frequencies, decreasing to 0 dB at 20 kHz. The high-frequency rolloff correlates with the audiogram and supports the idea that middle-ear transmission limits high-frequency hearing, providing a stronger link between inner-ear macromechanics and hearing. We estimate the cochlear partition impedance Z(CP) from these and previous data. The chinchilla may be a useful animal model for exploring the effects of non-acoustic inner-ear stimulation such as "bone conduction" on cochlear mechanics.

Motion of the tympanic membrane after cartilage tympanoplasty determined by stroboscopic holography.

Stroboscopic holography was used to quantify dynamic deformations of the tympanic membrane (TM) of the entire surface of the TM before and after cartilage tympanoplasty of the posterior or posterior-superior part of the TM. Cartilage is widely used in tympanoplasties to provide mechanical stability for the TM. Three human cadaveric temporal bones were used. A 6 mm x 3 mm oval cartilage graft was placed through the widely opened facial recess onto the medial surface of the posterior or posterior-superior part of the TM. The graft was either in contact with the bony tympanic rim and manubrium or not. Graft thickness was either 0.5 or 1.0mm. Stroboscopic holography produced displacement amplitude and phase maps of the TM surface in response to stimulus sound. Sound stimuli were 0.5, 1, 4 and 7 (or 8)kHz tones. Middle-ear impedance was measured from the motion of the entire TM. Cartilage placement generally produced reductions in the motion of the TM apposed to the cartilage, especially at 4 kHz and 7 or 8 kHz. Some parts of the TM showed altered motion compared to the control in all three cases. In general, middle-ear impedance was either unchanged or increased somewhat after cartilage reconstruction both at low (0.5 and 1 kHz) and high (4 and 7 kHz) frequencies. At 4 kHz, with the 1.0mm thick graft that was in contact with the bony tympanic rim, the impedance slightly decreased. While our earlier work with time-averaged holography allowed us to observe differences in the pattern of TM motion caused by application of cartilage to the TM, stroboscopic holography is more sensitive to TM motions and allowed us to quantify the magnitude and phase of motion of each point on the TM surface. Nonetheless, our results are similar to those of our earlier work: The placement of cartilage on the medial surface of TM reduces the motion of the TM that apposes the cartilage. These obvious local changes occur even though the cartilage had little effect on the sound-induced motion of the stapes.

Evaluation of round window stimulation using the floating mass transducer by intracochlear sound pressure measurements in human temporal bones.

HYPOTHESIS: Round window (RW) stimulation with a floating mass transducer (FMT) can be studied experimentally and optimized to enhance auditory transduction. BACKGROUND: The FMT (MED-EL Vibrant Soundbridge) has been recently implanted in patients with refractory conductive or mixed hearing loss to stimulate the RW with varying degrees of success. The mechanics of RW stimulation with the FMT have not been studied in a systematic manner. METHODS: In cadaveric human temporal bones, measurements of stapes velocity with laser vibrometry in response to FMT-RW stimulation were used to optimize FMT insertion. The effect of RW stimulation on hearing was estimated using simultaneous measurements of intracochlear pressures in both perilymphatic scalae with micro-optical pressure transducers. This enabled calculation of the differential pressure across the cochlear partition, which is directly tied to auditory transduction. RESULTS: The best coupling between the FMT and RW was achieved with a piece of fascia placed between the RW and the FMT, and by "bracing" the free end of the FMT against the hypotympanic wall with dental impression material. FMT-RW stimulation provided differential pressures comparable with sound-induced oval window stimulation greater than 1 kHz. However, less than 1 kHz, the FMT was less capable. CONCLUSION: Measurements of stapes velocity and intracochlear sound pressures in scala vestibuli and scala tympani enabled experimental evaluation of FMT stimulation of the RW. The efficacy of FMT-RW coupling was influenced significantly by technical and surgical factors, which can be optimized. This temporal bone preparation also lays the foundation for future studies to investigate multiple issues of relevance to both basic and clinical science such as RW stimulation in stapes fixation, nonaerated middle ears, and third-window lesions, and to answer basic questions regarding bone conduction.