Nonlinear active force generation by cochlear outer hair cell

We analyze the nonlinear behavior of the longitudinal and circumferential components of the active force generated by the outer hair cell wall in response to changes of its transmembrane potential. We treat the material of the wall as electroelastic, linear orthotropic in terms of strains and as nonlinear in terms of the transmembrane potential. To describe the nonlinear behavior of the active force versus the transmembrane potential, we use two (Boltzmann and simple exponential) types of approximation. We estimate free parameters of these approximations by combining the previously reported passive stiffnesses with the active strains measured in the microchamber experiment. We analyze the sensitivity of the estimated parameters corresponding to changes of the cell axial stiffness, a characteristic independently measured by several groups. We also study the effect of combining the active strains measured in the microchamber experiment with those measured in the whole cell recording experiment. We show agreement between our prediction of the active force and measurements in the whole cochlea and in isolated cells.     

Effect of voltage-dependent membrane properties on active force generation in cochlear outer hair cell

A computational model is proposed to analyze the active force production in an individual outer hair cell (OHC) under high-frequency conditions. The model takes into account important biophysical properties of the cell as well as constraints imposed by the surrounding environment. The biophysical properties include the elastic, piezoelectric, and viscous characteristics of the cell wall. The effect of the environment is associated with the stiffness of the constraint and the drag forces acting on the cell due to the interaction with the external and internal viscous fluids. The study concentrated on a combined effect of the transmembrane potential, frequency, and stiffness of the constraints. The effect of the voltage-dependent stiffness of the cell was particularly investigated and it was found to be twofold. First, it results in higher sensitivity and nonlinearity of the OHC active force production in the physiological range. Second, it determines smaller active forces in the hyperpolarization range. The resonant properties of the active force as functions of voltage and the constraint stiffness were also analyzed. The obtained results can be important for a better understanding of the OHC active force production and the contribution of cell electromotility to the cochlear amplification, sensitivity, and nonlinearity.     

Elasticity and active force generation of cochlear outer hair cells.

The cochlear outer hair cell is described by a cylindrical membrane model, characterized by area and shear moduli for a passive elastic element and an active tension element dependent on the membrane potential. In passive experiments, these moduli are determined from the pressure-strain relations. The area modulus obtained is 0.07 N m-1, similar to a lipid bilayer and the shear modulus is 0.007 N m-1. These moduli combined with previous active experiments show that the active tension is nearly isotropic and is about 1.6 x 10(-2) N m-1 V-1, resulting in a 0.5 nN/mV force per cell. This implies that the receptor potential for acoustical stimulation produces an active force comparable to the acoustic force applied to the basilar membrane per outer hair cell. This finding supports the hypothesis that the outer hair cell acts as feedback motor in the fine tuning mechanism of the mammalian ear.     

Electrokinetic model of cochlear hair cell motility

Recent experiments have shown that isolated outer hair cells of the cochlea can vibrate under the influence of a transcellular oscillating electric field. Since high voltages have been measured in the cochlea, this result might suggest a basis for electromechanical feedback. A mechanical model of the hair cell has been developed and adapted to test the electrokinetic theory of motility, a postulate of which is that cochlear voltage gradients act on charged proteins embedded in the cell membrane to deform the cell. From the model it was deduced that the amount of charge density required is within the physiologic range. The significant result is that the amplitude of cell elongation for a fixed voltage amplitude is virtually constant for frequency less than a certain cutoff. The value of this frequency depends on the various physical parameters of the system and especially on the spacing between cells. Power transfer to the basilar membrane appears to peak near the cutoff frequency, and the amount is not very dependent on cell length, but is highly dependent on cell spacing.     

Outer hair cell active force generation in the cochlear environment

Outer hair cells are critical to the amplification and frequency selectivity of the mammalian ear acting via a fine mechanism called the cochlear amplifier, which is especially effective in the high-frequency region of the cochlea. How this mechanism works under physiological conditions and how these cells overcome the viscous (mechanical) and electrical (membrane) filtering has yet to be fully understood. Outer hair cells are electromotile, and they are strategically located in the cochlea to generate an active force amplifying basilar membrane vibration. To investigate the mechanism of this cell's active force production under physiological conditions, a model that takes into account the mechanical, electrical, and mechanoelectrical properties of the cell wall (membrane) and cochlear environment is proposed. It is shown that, despite the mechanical and electrical filtering, the cell is capable of generating a frequency-tuned force with a maximal value of about 40 pN. It is also found that the force per unit basilar membrane displacement stays essentially the same (40 pNnm) for the entire linear range of the basilar membrane responses, including sound pressure levels close to hearing threshold. Our findings can provide a better understanding of the outer hair cell's role in the cochlear amplifier.     

Effect of outer hair cell piezoelectricity on high-frequency receptor potentials

The low-pass voltage response of outer hair cells predicted by conventional equivalent circuit analysis would preclude the active force production at high frequencies. We have found that the band pass characteristics can be improved by introducing the piezoelectric properties of the cell wall. In contrast to the conventional analysis, the receptor potential does not tend to zero and at any frequency is greater than a limiting value. In addition, the phase shift between the transduction current and receptor potential tends to zero. The piezoelectric properties cause an additional, strain-dependent, displacement current in the cell wall. The wall strain is estimated on the basis of a model of the cell deformation in the organ of Corti. The limiting value of the receptor potential depends on the ratio of a parameter determined by the piezoelectric coefficients and the strain to the membrane capacitance. In short cells, we have found that for the low-frequency value of about 2-3 mV and the strain level of 0.1% the receptor potential can reach 0.4 mV throughout the whole frequency range. In long cells, we have found that the effect of the piezoelectric properties is much weaker. These results are consistent with major features of the cochlear amplifier.     

A diffusion model of the transient response of the cochlear inner hair cell synapse

One class of models of hair cell synaptic function that has been investigated in recent years consists of one or more reservoirs of synaptic material connected to other reservoirs and/or the synaptic cleft by means of diffusion paths. One such general model is considered here, comprising two reservoirs and a global source of synaptic material connected in series and releasing material into the synaptic cleft by a diffusion path characterized by an intensity-dependent permeability. The explicit form of the solution of the model for a sudden onset of stimulation is derived: The solution comprises two exponentially decaying terms plus a constant. This solution is shown to have a unique inverse. This allows the determination of the parameters of the model directly from experimental data from auditory-nerve fibers in the Mongolian gerbil. The behavior of the derived model parameters with variation of stimulus intensity is demonstrated, and implications for synaptic function are discussed.     

Multiple reservoir model of neurotransmitter release by a cochlear inner hair cell

A probabilistic model is described for transmitter release from hair cells, auditory neuron EPSP's, and discharge patterns. The present model assumes that several reservoirs of neurotransmitter exist, having individual probability-of-release functions centered at successively higher intensities. The model accurately mimics the adaptation of successive EPSP amplitudes of the afferent neuron of the goldfish sacculus and, for mammalian auditory-nerve fibers, the adaptation of neural discharge rate, the saturation of onset and steady-state neural rate versus intensity, and the change in neural rate in response to incremental stimuli. The model also produces realistic interval and period histograms. The data shown support the hypothesis that multiple populations of neurotransmitter are involved in the afferent hair-cell synapses.     

On one mechanism of mechanoelectrical transformations in impact-excited composites based on the cement binder

A mechanism of mechanoelectrical transformations produced by impact excitation in cement-binder-based composites is suggested and experimentally verified. The transformations proceed in two stages: material electrification at the point of impact and mechanoelectrical processes at the matrix-binder interface.     

Discharges and mechanoelectrical phenomena in charged glasses

Conclusions In high-resistance solid dielectrics (with resistances of 10¹² · cm or higher, when they are irradiated with electron beams, spontaneous electric discharges are observed while the irradiation is occurring. The discharges occur at defects in the surface and develop in the layer within which the charged particles are distributed.When the charged particles are distributed in the surface or near-surface layer of the dielectric, the discharge may be initiated close to the charged surface by a grounded metal needle, even some time after the irradiation. After long storage and self-dis charge of the surface layer of the irradiated dielectric or compensation of the surface charge, discharge of the charge stored in the depth of the dielectric can occur by scratching the surface or pricking it with a needle. When the charge lies deep inside the dielectric, a brush discharge occurs when a strong shock is applied to the surface of the dielectric using a grounded needle.In all the above cases initiation of an electric discharge in the solid dielectric occurred by mechanoelectrical phenomena of direct conversion of mechanical energy into electrical energy with a high field strength. These phenomena occur in solid dielectrics with a defect structure.Translated from Izvestiya Vysshikh Uchebnykh Zavedenii, Fizika, No. 2, pp. 40–46, February, 1977.     

Effect of humidity on the parameters of mechanoelectrical transformations in concretes

An increase in the humidity of a cement solution from 0 to 8% is found to result in an exponential decrease in the principal spectral maximum of its electric response to an impact excitation according to the change in the electrical resistivity. A physical model is proposed for the mechanoelectrical transformations occurring in a wet heterogeneous material, and this model agrees well with the experimental data.     

Influence of direct current on dc receptor potentials from cochlear inner hair cells in the guinea pig

Inner hair cell responses to sound were monitored while direct current was applied across the membranous labyrinth in the first turn of the guinea pig cochlea. The current injection electrodes were positioned in the scala vestibuli and on the round window membrane. Positive and negative current (less than 100 microA) caused changes in the sound-evoked dc receptor potentials which were dependent on the sound frequency and intensity. The frequencies most affected by this extracellular current were those comprising the "tip" portion of the inner hair cell frequency tuning characteristic (FTC). The influence of current increased with increasing frequency. Positive current increased the amount of dc receptor potential for the affected frequencies while negative current decreased the potential. Current-induced changes (on a percentage basis) were greater for low intensity sounds and the negative current direction. These frequency specific changes are evidenced as a loss in sensitivity for the tip area of the FTC and a downward shift of the inner hair cell characteristic frequency. Larger current levels (greater than 160 microA) cause more complex changes including unrecoverable loss of cell performance. In separate experiments positive and negative currents (less than 1.1 microA) were injected into the inner hair cell from the recording electrode during simultaneous measurement of the sound-evoked dc receptor potential. This condition caused a shift in IHC sensitivity that was independent of sound frequency and intensity. Positive current decreased the sensitivity of the level of the cell while negative current increased the responses. The effect of current level on sound-evoked dc receptor potential was nonlinear, as comparatively greater increases in cell response were observed for negative than decreases for positive current. The intracellular current injection results are accounted for by the mechano-resistive model of hair cell transduction, where nonlinear responses with current level may reflect outward rectification. Response changes induced by extracellular current are evidence of current effects on both inner and outer hair cells. The frequency and intensity dependences are hypothesized to represent voltage mediated control of inner hair cell response by the outer hair cells.     

On S-transformation in the strong coupling theory

S-transformation is a unitary transformation accomplishing the main part of the diagonalization of the interaction Hamiltonian in strong coupling theories. For a compact symmetry group the action of the S-transformation on the integrals of motion is derived.Dedicated to Professor V. Votruba on his sixtieth birthday.     

On nonlinear cascades and resonances in the outer magnetosphere

The paper addresses nonlinear phenomena that control the interaction between plasma flow (solar wind) and magnetic barrier (magnetosphere). For the first time we demonstrate that the dominant solar wind kinetic energy: (i) excites boundary resonances and their harmonics which modulate plasma jets under the bow shock; (ii) produces discrete three-wave cascades, which could merge into a turbulent-like one; (iii) jet produced cascades provide the effective anomalous plasma transport inside and out of the magnetosphere; (iv) intermittency and multifractality characteristics for the statistic properties of jets result in a super-ballistic turbulent transport regime. Our results could be considered as suggestive for the space weather predictions, for turbulent cascades in different media and for the laboratory plasma confinement (e.g., for fusion devices).     

Peculiarities of the mechanoelectrical response in dielectric materials in the region of structural phase transformations

The influence of thermally induced structural phase transformations on the electromagnetic response of gypsum upon impact excitation is investigated. It is found that the electromagnetic response upon impact excitation is the structure-sensitive characteristics of dielectric materials. Data on the character of phase transformations in gypsum heated above 150°C are obtained using X-ray diffraction techniques.     

Mixed outer synchronization of coupled complex networks with time-varying coupling delay

In this paper, the problem of outer synchronization between two complex networks with the same topological structure and time-varying coupling delay is investigated. In particular, we introduce a new type of outer synchronization behavior, i.e., mixed outer synchronization (MOS), in which different state variables of the corresponding nodes can evolve into complete synchronization, antisynchronization, and even amplitude death simultaneously for an appropriate choice of the scaling matrix. A novel nonfragile linear state feedback controller is designed to realize the MOS between two networks and proved analytically by using Lyapunov-Krasovskii stability theory. Finally, numerical simulations are provided to demonstrate the feasibility and efficacy of our proposed control approach.     

On nondegenerate coupling forms

The aim of the present paper is to investigate new classes of symplectically fat fibre bundles. We prove a general existence theorem for fat vectors with respect to the canonical invariant connections. Based on this result we give new proofs of some constructions of symplectic structures. This includes twistor bundles and locally homogeneous complex manifolds. The proofs are conceptually simpler and allow for obtaining more general results.