The Active Traveling Wave in the Cochlea

A sound stimulus entering the inner ear excites a deformation of the basilar membrane which travels along the cochlea towards the apex. It is well established that this wave-like disturbance is amplified by an active system. Recently, it has been proposed that the active system consists of a set of self-tuned critical oscillators which automatically operate at an oscillatory instability. Here, we show how the concepts of a traveling wave and of self-tuned critical oscillators can be combined to describe the nonlinear wave in the cochlea.Comment: 5 pages, 2 figure

Transfer function for vital infrasound pressures between the carotid artery and the tympanic membrane

While occupational injury is associated with numerous individual and work-related risk factors, including long working hours and short sleep duration, the complex mechanisms causing such injuries are not yet fully understood. The relationship between the infrasound pressures of the tympanic membrane [ear canal pressure (ECP)], detected using an earplug embedded with a low-frequency microphone, and the carotid artery [carotid artery pressure (CAP)], detected using a stethoscope fitted with the same microphone, can be quantitatively characterized using systems analysis. The transfer functions of 40 normal workers (19 to 57 years old) were characterized, involving the analysis of 446 data points. The ECP waveform exhibits a pulsatile character with a slow respiratory component, which is superimposed on a biphasic recording that is synchronous with the cardiac cycle. The respiratory ECP waveform correlates with the instantaneous heart rate. The results also revealed that various fatigue-related risk factors may affect the mean magnitudes of the measured pressures and the delay transfer functions between CAP and ECP in the study population; these factors include systolic blood pressure, salivary amylase activity, age, sleep duration, postural changes, chronic fatigue, and pulse rate. [http://dx.doi.org/10.1121/1.4773270

Non-invasive biophysical measurement of travelling waves in the insect inner ear

Frequency analysis in the mammalian cochlea depends on the propagation of frequency information in the form of a travelling wave (TW) across tonotopically arranged auditory sensilla. TWs have been directly observed in the basilar papilla of birds and the ears of bush-crickets (Insecta: Orthoptera) and have also been indirectly inferred in the hearing organs of some reptiles and frogs. Existing experimental approaches to measure TW function in tetrapods and bushcrickets are inherently invasive, compromising the fine-scale mechanics of each system. Located in the forelegs, the bushcricket ear exhibits outer, middle and inner components; the inner ear containing tonotopically arranged auditory sensilla within a fluid-filled cavity, and externally protected by the leg cuticle. Here, we report bush-crickets with transparent ear cuticles as potential model species for direct, non-invasive measuring of TWs and tonotopy. Using laser Doppler vibrometry and spectroscopy, we show that increased transmittance of light through the ear cuticle allows for effective non-invasive measurements of TWs and frequency mapping. More transparent cuticles allow several properties of TWs to be precisely recovered and measured in vivo from intact specimens. Our approach provides an innovative, noninvasive alternative to measure the natural motion of the sensillia-bearing surface embedded in the intact inner ear fluid

Optics-less smart sensors and a possible mechanism of cutaneous vision in nature

Optics-less cutaneous (skin) vision is not rare among living organisms, though its mechanisms and capabilities have not been thoroughly investigated. This paper demonstrates, using methods from statistical parameter estimation theory and numerical simulations, that an array of bare sensors with a natural cosine-law angular sensitivity arranged on a flat or curved surface has the ability to perform imaging tasks without any optics at all. The working principle of this type of optics-less sensor and the model developed here for determining sensor performance may be used to shed light upon possible mechanisms and capabilities of cutaneous vision in nature

Rippling pattern of distortion product otoacoustic emissions evoked by high-frequency primaries in guinea pigs

The origin of ripples in distortion product otoacoustic emission (DPOAE) amplitude which appear at specific DPOAE frequencies during f1 tone sweeps using fixed high frequency f2 (>20 kHz) in guinea pigs is investigated. The peaks of the ripples, or local DPOAE amplitude maxima, are separated by approximately half octave intervals and are accompanied by phase oscillations. The local maxima appear at the same frequencies in DPOAEs of different order and velocity responses of the stapes and do not shift with increasing levels of the primaries. A suppressor tone had little effect on the frequencies of the maxima, but partially suppressed DPOAE amplitude when it was placed close to the f2 frequencies. These findings agree with earlier observations that the maxima occur at the same DPOAE frequencies, which are independent of the f2 and the primary ratio, and thus are likely to be associated with DPOAE propagation mechanisms. Furthermore, the separation of the local maxima by approximately half an octave may suggest that the maxima are due to interference of the travelling waves along the basilar membrane at the frequency of the DPOAE. It is suggested that the rippling pattern appears because of interaction between DPOAE reverse travelling waves with standing waves formed in the cochlea

Essential nonlinearities in hearing

Our hearing organ, the cochlea, evidently poises itself at a Hopf bifurcation to maximize tuning and amplification. We show that in this condition several effects are expected to be generic: compression of the dynamic range, infinitely shrap tuning at zero input, and generation of combination tones. These effects are "essentially" nonlinear in that they become more marked the smaller the forcing: there is no audible sound soft enough not to evoke them. All the well-documented nonlinear aspects of hearing therefore appear to be consequences of the same underlying mechanism.Comment: 4 pages, 3 figure

Travelling Ultrasonic Wave Enhances Keyclick Sensation

International audienceA realistic keyclick sensation is a serious challenge for haptic feedback since vibrotactile rendering faces the limitation of the absence of contact force as experienced on physical buttons. It has been shown that creating a keyclick sensation is possible with stepwise ultrasonic friction modulation. However, the intensity of the sensation is limited by the impedance of the fingertip and by the absence of a lateral force component external to the finger. In our study, we compare this technique to rendering with an ultrasonic travelling wave, which exerts a lateral force on the fingertip. For both techniques, participants were asked to report the detection (or not) of a keyclick during a forced choice one interval procedure. In experiment 1, participants could press the surface as many time as they wanted for a given trial. In experiment 2, they were constrained to press only once. The results show a lower perceptual threshold for travelling waves. Moreover, participants pressed less times per trial and exerted smaller normal force on the surface. The subjective quality of the sensation was found similar for both techniques. In general, haptic feedback based on travelling ultrasonic waves is promising for applications without lateral motion of the finger

Localization of the Cochlear Amplifier in Living Sensitive Ears

BACKGROUND: To detect soft sounds, the mammalian cochlea increases its sensitivity by amplifying incoming sounds up to one thousand times. Although the cochlear amplifier is thought to be a local cellular process at an area basal to the response peak on the spiral basilar membrane, its location has not been demonstrated experimentally. METHODOLOGY AND PRINCIPAL FINDINGS: Using a sensitive laser interferometer to measure sub-nanometer vibrations at two locations along the basilar membrane in sensitive gerbil cochleae, here we show that the cochlea can boost soft sound-induced vibrations as much as 50 dB/mm at an area proximal to the response peak on the basilar membrane. The observed amplification works maximally at low sound levels and at frequencies immediately below the peak-response frequency of the measured apical location. The amplification decreases more than 65 dB/mm as sound levels increases. CONCLUSIONS AND SIGNIFICANCE: We conclude that the cochlea amplifier resides at a small longitudinal region basal to the response peak in the sensitive cochlea. These data provides critical information for advancing our knowledge on cochlear mechanisms responsible for the remarkable hearing sensitivity, frequency selectivity and dynamic range

Resultant pressure distribution pattern along the basilar membrane in the spiral shaped cochlea

Cochlea is an important auditory organ in the inner ear. In most mammals, it is coiled as a spiral. Whether this specific shape influences hearing is still an open problem. By employing a three dimensional fluid model of the cochlea with an idealized geometry, the influence of the spiral geometry of the cochlea is examined. We obtain solutions of the model through a conformal transformation in a long-wave approximation. Our results show that the net pressure acting on the basilar membrane is not uniform along its spanwise direction. Also, it is shown that the location of the maximum of the spanwise pressure difference in the axial direction has a mode dependence. In the simplest pattern, the present result is consistent with the previous theory based on the WKB-like approximation [D. Manoussaki, Phys. Rev. Lett. 96, 088701(2006)]. In this mode, the pressure difference in the spanwise direction is a monotonic function of the distance from the apex and the normal velocity across the channel width is zero. Thus in the lowest order approximation, we can neglect the existance of the Reissner's membrane in the upper channel. However, higher responsive modes show different behavior and, thus, the real maximum is expected to be located not exactly at the apex, but at a position determined by the spiral geometry of the cochlea and the width of the cochlear duct. In these modes, the spanwise normal velocities are not zero. Thus, it indicates that one should take into account of the detailed geometry of the cochlear duct for a more quantitative result. The present result clearly demonstrates that not only the spiral geometry, but also the geometry of the cochlear duct play decisive roles in distributing the wave energy.Comment: 21 pages. (to appear in J. Biol. Phys.