Julien Meaud

The remarkable cochlear amplifier

This composite article is intended to give the experts in the field of cochlear mechanics an opportunity to voice their personal opinion on the one mechanism they believe dominates cochlear amplification in mammals. A collection of these ideas are presented here for the auditory community and others interested in the cochlear amplifier. Each expert has given their own personal view on the topic and at the end of their commentary they have suggested several experiments that would be required for the decisive mechanism underlying the cochlear amplifier. These experiments are presently lacking but if successfully performed would have an enormous impact on our understanding of the cochlear amplifier.

The effect of tectorial membrane and basilar membrane longitudinal coupling in cochlear mechanics

Most mathematical models of the mammalian cochlea neglect structural longitudinal coupling. However, recent experimental data suggest that viscoelastic longitudinal coupling, in the basilar membrane (BM) and the tectorial membrane (TM), is non-negligible. In this paper, mathematical models for BM and TM longitudinal coupling are presented to determine the influence of such a coupling on the tuning of the BM. The longitudinal coupling models are added to a macroscopic linear model of the guinea pig cochlea that includes the micromechanics of the organ of Corti and outer hair cell (OHC) somatic motility. The predictions of the BM response to acoustic stimulus show that the characteristic frequency is controlled by a TM radial resonance and that TM longitudinal coupling has a more significant effect than BM longitudinal coupling. TM viscoelasticity controls the sharpness of the BM frequency response and the duration of the impulse response. The results with realistic TM longitudinal coupling are more consistent with experiments. The model predicts that OHC somatic electromotility is able to supply power to the BM at frequencies well above the cutoff of the OHC basolateral membrane. Moreover, TM longitudinal coupling is predicted to stabilize the cochlea and enable a higher BM sensitivity to acoustic stimulation.

Coupling Active Hair Bundle Mechanics, Fast Adaptation, and Somatic Motility in a Cochlear Model

One of the central questions in the biophysics of the mammalian cochlea is determining the contributions of the two active processes, prestin-based somatic motility and hair bundle (HB) motility, to cochlear amplification. HB force generation is linked to fast adaptation of the transduction current via a calcium-dependent process and somatic force generation is driven by the depolarization caused by the transduction current. In this article, we construct a global mechanical-electrical-acoustical mathematical model of the cochlea based on a three-dimensional fluid representation. The global cochlear model is coupled to linearizations of nonlinear somatic motility and HB activity as well as to the micromechanics of the passive structural and electrical elements of the cochlea. We find that the active HB force alone is not sufficient to power high frequency cochlear amplification. However, somatic motility can overcome resistor-capacitor filtering by the basolateral membrane and deliver sufficient mechanical energy for amplification at basal locations. The results suggest a new theory for high frequency active cochlear mechanics, in which fast adaptation controls the transduction channel sensitivity and thereby the magnitude of the energy delivered by somatic motility.

Simultaneously High Stiffness and Damping in Nanoengineered Microtruss Composites

Materials combining high stiffness and mechanical energy dissipation are needed in automotive, aviation, construction, and other technologies where structural elements are exposed to dynamic loads. In this paper we demonstrate that a judicious combination of carbon nanotube engineered trusses held in a dissipative polymer can lead to a composite material that simultaneously exhibits both high stiffness and damping. Indeed, the combination of stiffness and damping that is reported is quite high in any single monolithic material. Carbon nanotube (CNT) microstructures grown in a novel 3D truss topology form the backbone of these nanocomposites. The CNT trusses are coated by ceramics and by a nanostructured polymer film assembled using the layer-by-layer technique. The crevices of the trusses are then filled with soft polyurethane. Each constituent of the composite is accurately modeled, and these models are used to guide the manufacturing process, in particular the choice of the backbone topology and the optimization of the mechanical properties of the constituent materials. The resulting composite exhibits much higher stiffness (80 times) and similar damping (specific damping capacity of 0.8) compared to the polymer. Our work is a step forward in implementing the concept of materials by design across multiple length scales.

Three-Dimensional-Printed Multistable Mechanical Metamaterials With a Deterministic Deformation Sequence

Multistable mechanical metamaterials are materials that have multiple stable configurations. The geometrical changes caused by the transition of the metamaterial from one stable state to another, could be exploited to obtain multifunctional and programmable materials. As the stimulus amplitude is varied, a multistable metamaterial goes through a sequence of stable configurations. However, this sequence (which we will call the deformation sequence) is unpredictable if the metamaterial consists of identical unit cells. This paper proposes to use small variations in the unit cell geometry to obtain a deterministic deformation sequence for one type of multistable metamaterial that consists of bistable unit cells. Based on an analytical model for a single unit cell and on the minimization of the total strain energy, a rigorous theoretical model is proposed to analyze the nonlinear mechanics of this type of metamaterials and to inform the designs. The proposed theoretical model is able to accurately predict the deformation sequence and the stress–strain curves that are observed in the finite-element simulations with an elastic constitutive model. A deterministic deformation sequence that matches the sequence predicted by the theory and finite-element simulations is obtained in experiments with 3D-printed samples. Furthermore, an excellent quantitative agreement between simulations and experiments is obtained once a viscoelastic constitutive model is introduced in the finite-element model. [DOI: 10.1115/1.4034706]

Effect of the Attachment of the Tectorial Membrane on Cochlear Micromechanics and Two-Tone Suppression

The mechanical stimulation of the outer hair cell hair bundle (HB) is a key step in nonlinear cochlear amplification. We show how two-tone suppression (TTS), a hallmark of cochlear nonlinearity, can be used as an indirect measure of HB stimulation. Using two different nonlinear computational models of the cochlea, we investigate the effect of altering the mechanical load applied by the tectorial membrane (TM) on the outer hair cell HB. In the first model (TM-A model), the TM is attached to the spiral limbus (as in wild-type animals); in the second model (TM-D model), the TM is detached from the spiral limbus (mimicking the cochlea of Otoa(EGFP/EGFP) mutant mice). As in recent experiments, model simulations demonstrate that the absence of the TM attachment does not preclude cochlear amplification. However, detaching the TM alters the mechanical load applied by the TM on the HB at low frequencies and therefore affects TTS by low-frequency suppressors. For low-frequency suppressors, the suppression threshold obtained with the TM-A model corresponds to a constant suppressor displacement on the basilar membrane (as in experiments with wild-type animals), whereas it corresponds to a constant suppressor velocity with the TM-D model. The predictions with the TM-D model could be tested by measuring TTS on the basilar membrane of the Otoa(EGFP/EGFP) mice to improve our understanding of the fundamental workings of the cochlea.

Nonlinear response to a click in a time-domain model of the mammalian ear

In this paper, a state-space implementation of a previously developed frequency-domain model of the cochlea is coupled to a lumped parameter model of the middle ear. After validation of the time-domain model by comparison of its steady-state response to results obtained with a frequency-domain formulation, the nonlinear response of the cochlea to clicks is investigated. As observed experimentally, a compressive nonlinearity progressively develops within the first few cycles of the response of the basilar membrane (BM). Furthermore, a time-frequency analysis shows that the instantaneous frequency of the BM response to a click progressively approaches the characteristic frequency. This phenomenon, called glide, is predicted at all stimulus intensities, as in experiments. In typical experiments with sensitive animals, the click response is characterized by a long ringing and the response envelope includes several lobes. In order to achieve similar results, inhomogeneities are introduced in the cochlear model. Simulations demonstrate the strong link between characteristics of the frequency response, such as dispersion and frequency-dependent nonlinearity, and characteristics of the time-domain response, such as the glide and a time-dependent nonlinearity. The progressive buildup of cochlear nonlinearity in response to a click is shown to be a consequence of the glide and of frequency-dependent nonlinearity.

Dependence of the dynamic properties of Voigt and Reuss composites on the Poisson's ratios and bulk loss factors of the constituent materials

Materials possessing both high stiffness and high damping would be beneficial in many structural applications. Composites that combine a stiff material, which usually has low damping, with a soft and lossy material have been proposed to engender both high dynamic modulus and high loss factor. In this article, we investigate the effective dynamic moduli and loss factors of Reuss and Voigt composites in response to a uniaxial harmonic load. The constituent materials are characterized by multiaxial viscoelastic models in the frequency domain. Using the viscoelastic correspondence principle, we derive formulae for Reuss and Voigt composites of infinite dimensions taking into account Poisson effects. We show that the effective loss factor of a Reuss composite is sensitive to the values of the Poissons ratio and bulk loss factors of the constituent materials. Finally we simulate, using finite element analysis, the response of cylindrical Reuss composite rods of finite radius to an axial load. We demonstrate that the effective dynamic properties of these rods is highly sensitive to the ratio of the radius to the layer thickness of the composite.

Middle-ear function in the chinchilla: Circuit models and comparison with other mammalian species

The middle ear efficiently transmits sound from the ear canal into the inner ear through a broad range of frequencies. Thus, understanding middle-ear transmission characteristics is essential in the study of hearing mechanics. Two models of the chinchilla middle ear are presented. In the first model, the middle ear is modeled as a lumped parameter system with elements that represent the ossicular chain and the middle-ear cavity. Parameters of this model are fit using available experimental data of two-port transmission matrix parameters. In an effort to improve agreement between model simulations and the phase of published experimental measurements for the forward pressure transfer function at high frequencies, a second model in which a lossless transmission line model of the tympanic membrane is appended to the original model is proposed. Two-port transmission matrix parameter results from this second model were compared with results from previously developed models of the guinea pig, cat, and human middle ears. Model results and published experimental data for the two-port transmission matrix parameters are found to be qualitatively similar between species. Quantitative differences in the two-port transmission matrix parameters suggest that the ossicular chains of chinchillas, cats, and guinea pigs are less flexible than in humans.

Coupling the Subtectorial Fluid with the Tectorial Membrane and Hair Bundles of the Cochlea

Two different kinds of flow—(i) shearing of fluid between the reticular lamina (RL) and tectorial membrane (TM) and (ii) so‐called pulsating flow in the RL‐TM gap—have been implicated as the dominant source of fluidic stimulation of the inner hair cell (IHC) hair bundle (HB). However, the frequency and spatial dependence of these flows for IHC stimulation is unresolved in vivo and estimates of the effect of the cochlear amplifier on these flows has not been quantified. Indeed, the relative importance these flow modalities and active processes likely varies with tonotopic location. In this paper, a microfluidic model is developed which features the interaction of the subtectorial fluid with the TM, IHC HBs, and the outer hair cell HBs. The framework of the model allows for incorporation into active macroscopic models as well as for comparison of experiments performed on excised sections of the cochlea.