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.

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.

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.

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.

In this article, a nonlinear mathematical model is developed based on the physiology of the cochlea of the guinea pig. The three-dimensional intracochlear fluid dynamics are coupled to a micromechanical model of the organ of Corti and to electrical potentials in the cochlear ducts and outer hair cells (OHC). OHC somatic electromotility is modeled by linearized piezoelectric relations whereas the OHC hair-bundle mechanoelectrical transduction current is modeled as a nonlinear function of the hair-bundle deflection. The steady-state response of the cochlea to a single tone is simulated in the frequency domain using an alternating frequency time scheme. Compressive nonlinearity, harmonic distortion, and DC shift on the basilar membrane (BM), tectorial membrane (TM), and OHC potentials are predicted using a single set of parameters. The predictions of the model are verified by comparing simulations to available in vivo experimental data for basal cochlear mechanics. In particular, the model predicts more amplification on the reticular lamina (RL) side of the cochlear partition than on the BM, which replicates recent measurements. Moreover, small harmonic distortion and DC shifts are predicted on the BM, whereas more significant harmonic distortion and DC shifts are predicted in the RL and TM displacements and in the OHC potentials.

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.

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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.