The mechanical waveform of the basilar membrane. II. From data to models--and back

Mechanical responses in the basal turn of the guinea-pig cochlea are measured with low-level broad-band noise as the acoustical stimulus [for details see de Boer and Nuttall, J. Acoust. Soc. Am. 101, 3583-3592 (1997)]. Results are interpreted within the framework of a classical three-dimensional model of the cochlea that belongs to a very wide class of nonlinear models. The use of linear-systems analysis for this class of nonlinear models has been justified earlier [de Boer, Audit. Neurosci. 3, 377-388 (1997)]. The data are subjected to inverse analysis with the aim to recover the "effective basilar-membrane impedance." This is a parameter function that, when inserted into the model, produces a model response, the "resynthesized" response, that is similar to the measured response. With present-day solution methods, resynthesis leads back to an almost perfect replica of the original response in the spatial domain. It is demonstrated in this paper that this also applies to the response in the frequency domain and in the time domain. This paper further reports details with regard to geometrical properties of the model employed. Two three-dimensional models are studied; one has its dimensions close to that of the real cochlea, the other is a stylized model which has homogeneous geometry over its length. In spite of the geometric differences the recovered impedance functions are very similar. An impedance function computed for one model can be used in resynthesis of the response in the other one, and this leads to global amplitude deviations between original and resynthesized response functions not exceeding 8 dB. Discrepancies are much larger (particularly in the phase) when a two-dimensional model is compared with a three-dimensional model. It is concluded that a stylized three-dimensional model with homogeneous geometric parameters will give sufficient information in further work on unraveling cochlear function via inverse analysis. In all cases of a sensitive cochlea stimulated by a signal with a stimulus level of 50 dB SPL per octave or less, the resulting basilar-membrane impedance is found to be locally active, that is, the impedance function shows a region where the basilar membrane is able to amplify acoustic power or to reduce dissipation of power by the organ of Corti. Finally, the influence of deliberate errors added to the data is discussed in order to judge the accuracy of the results.     

The mechanical waveform of the basilar membrane. III. Intensity effects

Mechanical responses in the basal turn of the guinea-pig cochlea were measured with broad-band noise stimuli and expressed as input-output cross-correlation functions. The experiments were performed over the full range of stimulus intensities in order to try to understand the influence of cochlear nonlinearity on frequency selectivity, tuning, signal compression and the impulse response. The results are interpreted within the framework of a nonlinear, locally active, three-dimensional model of the cochlea. The data have been subjected to inverse analysis in order to recover the basilar-membrane (BM) impedance, a parameter function that, when inserted into the (linearized version of that) model, produces a model response that is similar to the measured response. This paper reports details about intensity effects for noise stimulation, in particular, the way the BM impedance varies with stimulus intensity. In terms of the underlying cochlear model, the decrease of the "activity component" in the BM impedance with increasing stimulus level is attributed to saturation of transduction in the outer hair cells. In the present paper this property is brought into a quantitative form. According to the theory [the EQ-NL theorem, de Boer, Audit. Neurosci. 3, 377-388 (1997)], the BM impedance is composed of two components, both intrinsically independent of stimulus level. One is the passive impedance Zpass and the other one is the "extra" impedance Zextra. The latter impedance is to be multiplied by a real factor gamma (0 < or = gamma < or = 1) that depends on stimulus level. This concept about the composition of the BM impedance is termed the "two-component theory of the BM impedance." In this work both impedances are entirely derived from experimental data. The dependence of the factor gamma on stimulus level can be derived by using a unified form of the outer-hair-cell transducer function. From an individual experiment, the two functions Zpass and Zextra are determined, and an approximation (Zpass + gamma Zextra) to the BM impedance constructed. Next, the model response (the "resynthesized" response) corresponding to this "artificial" impedance is computed. The same procedure is executed for several stimulus-level values. For all levels, the results show a close correspondence with the original experimental data; this includes correct prediction of the compression of response amplitudes, the reduction of frequency selectivity, the shift in peak frequency and, most importantly, the preservation of timing in the impulse response. All these findings illustrate the predictive power of the underlying model.     

Auditory physics. Physical principles in hearing theory. II

In the first part of this series of papers [Phys. Reports 62 (1980) 87–174] several physical problems are described that are relevant for the study of the auditory system. The present paper extends this treatment, it describes more facts upon which auditory theory is to be based and it delves considerably deeper into the mechanics of the cochlea (inner ear). The first two chapters treat nonlinear phenomena that are found in physiological and mechanical responses of the cochlea and in psychophysical experiments (listening tests carried out in human subjects). The main part of the paper is devoted to the mechanics of the cochlea. This part is preceded by an overview of the results of mechanical measurements on the cochlea. As it turns out, the newest experimental data present a specific challenge for cochlear mechanics.The central three chapters of the paper describe the development of a linear three-dimensional model of the cochlea. The main intention is to describe this model in a step-by-step fashion (hence the chapter headings borrowed from the field of architecture). Even in this simplified case, the solution for the response of the cochlear model is far from easy. Therefore, a few excursions are made into fields of physics and engineering in which related problems are worked out in analytical form. Just as in Part I of this series of papers, this extended treatment considerably deepens insight into the physical factors involved. Confrontation of the requirements for modelling described in the first few chapters with what has been achieved in the final part reveals how much study in the field of auditory physics remains to be done.     

Wave propagation patterns in a "classical" three-dimensional model of the cochlea

The generation mechanisms of cochlear waves, in particular those that give rise to otoacoustic emissions (OAEs), are often complex. This makes it difficult to analyze wave propagation. In this paper two unusual excitation methods are applied to a three-dimensional stylized classical nonlinear model of the cochlea. The model used is constructed on the basis of data from an experimental animal selected to yield a smooth basilar-membrane impedance function. Waves going in two directions can be elicited by exciting the model locally instead of via the stapes. Production of DPOAEs was simulated by presenting the model with two relatively strong primary tones, with frequencies f1 and f2, estimating the driving pressure for the distortion product (DP) with frequency 2f1 - f2, and computing the resulting DP response pattern - as a function of distance along the basilar membrane. For wide as well as narrow frequency separations the resulting DP wave pattern in the model invariably showed that a reverse wave is dominant in nearly the entire region from the peak of the f2-tone to the stapes. The computed DP wave pattern was further analyzed as to its constituent components with the aim to isolate their properties.     

一种基于特征理论模拟凝聚炸药爆轰的单元中心型Lagrange方法

提出一种数值模拟凝聚炸药爆轰问题的单元中心型Lagrange方法.利用有限体积离散爆轰反应流动方程组,基于双曲型偏微分方程组的特征理论获得离散网格节点的速度与压力,获得的网格节点速度与压力用于更新网格节点位置以及计算网格单元边的数值通量.以这种方式获得的网格节点解是一种"真正多维"的理论解,是一维Godunov格式在二维Riemann问题的推广.有限体积离散得到的爆轰反应流动的半离散系统使用一种显-隐Runge-Kutta格式来离散求解：显式格式处理对流项,隐式格式处理化学反应刚性源项.算例表明,提出的单元中心型Lagrange方法能够较好地模拟凝聚炸药的爆轰反应流动.     

A macro-mechanical model of the guinea pig cochlea with realistic parameters

The post-mortem transfer function of the cochlea of the guinea pig was compared to the transfer function generated by a model with parameters derived from physical measurements of the guinea pig cochlea. Both the formulation and parameters of the model were carefully chosen to be realistic using evidence from published measurements. The fit between the transfer function of the model and recent mechanical measurements of the passive guinea pig cochlear response was good, with a root mean square ratio of 6.3 dB in amplitude and 0.33 pi rad in phase. The model was used to explore the effect of cochlear partition mode factor and duct geometry upon the mechanical response of the cochlea. Possible inadequacies of the model which could explain the remaining differences between the output of the model and measurements are discussed.     

Comments on "Earphones in Audiometry" [Zwislocki et al., J. Acoust. Soc. Am. 83, 1688-1689 (1988)

The Zwislocki et al. ["Earphones in Audiometry," J. Acoust. Soc. Am. 83, 1688-1689 (1988)] Letter to the Editor states that insert earphones have some unresolved technical problems, such as limited frequency response, limited dynamic range and power handling capability, intersubject variability, and hygiene safety. In evaluating circumaural earphones, Zwislocki et al. say that the lack of a standard coupler disqualifies them for audiometry. Since this letter carries the weight of a CHABA committee recommendation, these issues are commented on herein. Section I was written primarily by Mead Killion and Sec. II primarily by Edgar Villchur. For brevity throughout, the authors of the Zwislocki et al. letter will be referred to as "the authors."     

Comparison of WKB calculations and experimental results for three-dimensional cochlear models.

The WKB asymptotic method is applied to the calculation of cochlear models with square scala cross section, for which the fluid motion is fully three dimensional. The analysis begins with the exact solution for wave propagation in a duct with constant properties. This solution is somewhat tedious but straightforward, since it requires a Fourier series expansion across the duct. Then with the formulation of Whitham [Linear and Nonlinear Waves (Wiley, New York, 1974)], the approximate solution is readily generated for the duct with properties which vary slowly along the length. Numerical calculations are carried out for the experimental models of Cannel [Ph.D. thesis, Univ. of Warwick (1969)] and Helle [Dr.-Ing. disser., Technische Univ., Müchen (1974)] who furnish quantitative details of both "basilar membrane" response and model parameters. Without any free parameters for adjusting, the present WKB solution shows quite satisfactory agreement with the experimental model results. Computer time is reasonable; the calculation of displacement envelope and phase at a number of stations along the cochlea for a given frequency requires only one second of CPU time. Thus the credibility and practically of the approach is established for the investigation of yet more realistic and more elaborate cochlear models.     

Solutions of the Lévy type for the free vibration analysis of diagonally supported rectangular plates

A Lévy type solution is developed for the vibratory response of a simply supported rectangular plate subjected to a harmonic force distributed along the diagonal. The solution is then extended to determine the free vibration response of the same rectangular plate with inelastic lateral support on the diagonal. It is found that there is an excellent agreement between computed eigenvalues obtained here and those obtained by the author in an earlier paper in which a Navier type solution was utilized. The significant advantages inherent in the present Lévy type solution are discussed.     

Coherent reflection in a two-dimensional cochlea: Short-wave versus long-wave scattering in the generation of reflection-source otoacoustic emissions

The theory of coherent reflection filtering explains the empirical form of the cochlear reflectance by showing how it emerges from the coherent "backscattering" of forward-traveling waves by impedance perturbations in the mechanics of the cochlear partition. Since the theory was developed using the one-dimensional (1-D) transmission-line model of the cochlea, an obvious logical shortcoming is the failure of the long-wavelength approximation near the peak of the traveling wave, where coherent backscattering is purported to occur. Indeed, existing theory suggests that wave reflection may be strongly suppressed in the short-wave regime. To understand how short-wave behavior near the peak modifies the predictions of the long-wave theory, this paper solves the scattering problem in the 2-D cochlear model. The 2-D problem is reduced to a 1-D wave equation and the solution expressed as an infinite series in which successive terms arise via multiple scattering within the cochlea. The cochlear reflectance is computed in response-matched models constructed by solving the inverse problem to control for variations in mechanical tuning among models of different heights and dimensionality. Reflection from the peak region is significantly enhanced by the short-wave hydrodynamics, but other conclusions of the 1-D analysis--such as the predicted relation between emission group delay and the wavelength of the traveling wave--carry over with only minor modifications. The results illustrate the important role of passive hydromechanical effects in shaping otoacoustic emissions and cochlear tuning.     

A computer model of medial efferent suppression in the mammalian auditory system

Stimulation of the olivocochlear bundle reduces basilar membrane displacement, driven auditory nerve activity, and compound action potential (CAP) response to acoustic stimulation. These effects were simulated using a computer model of the auditory periphery. The model simulates the medial efferent activity by attenuating the basilar membrane response. The model was evaluated against three animal studies reporting measurements at three levels of the auditory system; basilar membrane, single auditory nerve fibers and whole auditory nerve CAP. The CAP data included conditions where tones were masked by noise and "unmasked" by stimulation of the olivocochlear bundle. The model was able to simulate the data both qualitatively and quantitatively. As a consequence, it may be a suitable platform for studying the contribution of the efferent system to auditory processing of more complex auditory sounds in distracting backgrounds.     

Basilar-membrane responses to multicomponent (Schroeder-phase) signals: understanding intensity effects

Harmonic complexes comprised of the same spectral components in either positive-Schroeder (+Schr) or negative-Schroeder (-Schr) phase [see Schroeder, IEEE Trans. Inf. Theory 16, 85-89 (1970)] have identical long-term spectra and similar waveform envelopes. However, localized patterns of basilar-membrane (BM) excitation can be quite different in response to these two stimuli. Measurements in chinchillas showed more modulated (peakier) BM excitation for +Schr than -Schr complexes [Recio and Rhode, J. Acoust. Soc. Am. 108, 2281-2298 (2000)]. In the current study, laser velocimetry was used to examine BM responses at a location tuned to approximately 17 kHz in the basal turn of the guinea-pig cochlea, for +Schr and -Schr complexes with a 203-Hz fundamental frequency and including 101 equal-amplitude components from 2031 to 22,344 Hz. At 35-dB SPL, +Schr response waveforms showed greater amplitude modulation than -Schr responses. With increasing stimulation level, internal modulation decreased for both complexes. To understand the observed phenomena quantitatively, responses were predicted on the basis of a linearized model of the cochlea. Prediction was based on an "indirect impulse response" measured in the same animal. Response waveforms for Schroeder-phase signals were accurately predicted, provided that the level of the indirect impulse used in prediction closely matched the level of the Schroeder-phase stimulus. This result confirms that the underlying model, which originally was developed for noise stimuli, is valid for stimuli that produce completely different response waveforms. Moreover, it justifies explanation of cochlear filtering (i.e., differential treatment of different frequencies) in terms of a linear system.     

The mechanical waveform of the basilar membrane. IV. Tone and noise stimuli

Analysis of mechanical cochlear responses to wide bands of random noise clarifies many effects of cochlear nonlinearity. The previous paper [de Boer and Nuttall, J. Acoust. Soc. Am. 107, 1497-1507 (2000)] illustrates how closely results of computations in a nonlinear cochlear model agree with responses from physiological experiments. In the present paper results for tone stimuli are reported. It was found that the measured frequency response for pure tones differs little from the frequency response associated with a noise signal. For strong stimuli, well into the nonlinear region, tones have to be presented at a specific level with respect to the noise for this to be true. In this report the nonlinear cochlear model originally developed for noise analysis was modified to accommodate pure tones. For this purpose the efficiency with which outer hair cells modify the basilar-membrane response was made into a function of cochlear location based on local excitation. For each experiment, the modified model is able to account for the experimental findings, within 1 or 2 dB. Therefore, the model explains why the type of filtering that tones undergo in the cochlea is essentially the same as that for noise signals (provided the tones are presented at the appropriate level).     

Electromotile hearing: acoustic tones mask psychophysical response to high-frequency electrical stimulation of intact guinea pig cochleae

When sinusoidal electric stimulation is applied to the intact cochlea, a frequency-specific acoustic emission can be recorded in the ear canal. Acoustic emissions are produced by basilar membrane motion, and have been used to suggest a corresponding acoustic sensation termed "electromotile hearing." Electromotile hearing has been specifically attributed to electric stimulation of outer hair cells in the intact organ of Corti. To determine the nature of the auditory perception produced by electric stimulation of a cochlea with intact outer hair cells, guinea pigs were tested in a psychophysical task. First, subjects were trained to report detection of sinusoidal acoustic stimuli and dynamic range was assessed using response latency. Subjects were then implanted with a ball electrode placed into scala tympani. Following the surgical implant procedure, subjects were transferred to a task in which acoustic signals were replaced by sinusoidal electric stimulation, and dynamic range was assessed again. Finally, the ability of acoustic pure-tone stimuli to mask the detection of the electric signals was assessed. Based on the masking effects, it is concluded that sinusoidal electric stimulation of the intact cochlea results in perception of a tonal (rather than a broadband or noisy) sound at a frequency of 8 kHz or above.     

Finite element cochlear models and their steady state response

Numerical cochlear models are constructed by means of a finite element approach and their frequency and spatial responses are calculated. The cochlea is modelled as a coupled fluid-membrane system, for which both two- and three-dimensional models are considered. The fluid in the scala canals is assumed to be incompressible and the basilar membrane is assumed to be a locally reactive impedance wall or a lossy elastic membrane. With the three-dimensional models, the effects are examined of the spiral configuration of the cochlea, of the presence of the lamina and the ligament that narrows the coupling area between the two fluid canals (scala vestibuli and scala tympani), and of the extended reaction of the basilar membrane which cannot be included in case of the two-dimensional models. The conclusion is that these effects on the cochlear response and the inherent mechanism governing the cochlear behaviour are found to be rather secondary.     

Dynamic characterization of hysteresis elements in mechanical systems. I. Theoretical analysis

The pre-sliding-pre-rolling phase of friction behavior is dominated by rate-independent hysteresis. Many machine elements in common engineering use exhibit, therefore, the characteristic of "hysteresis springs," for small displacements at least. Plain and rolling element bearings that are widely used in motion guidance of machine tools are typical examples. While the presence of a hysteresis element may mark the character of the resulting dynamics, little is to be found about this topic in the literature. The study of the nonlinear dynamics caused by such elements becomes imperative if we wish to achieve accurate control of such machines. In this Part I of the investigation, we examine a single-degree-of-freedom mass-hysteresis-spring system and show that, while the free response case is amenable to an exact solution, the more important case of forced response has no closed form solution and requires other methods of treatment. We consider harmonic-balance analysis methods (which are common analysis tools in engineering) suitable for frequency-domain treatment, in particular the approximate describing function (DF) method, and compare those results with "exact" numerical simulations. The DF method yields basically a linear equation with amplitude-dependent modal parameters. We find that agreement in the frequency response function, between DF and exact solution, is good for small excitation amplitudes and for very large amplitudes. Intermediate values, however, show high sensitivity to amplitude variations and, consequently, no regular solution is obtainable by either approach. This appears to be an inherent property of the system pointing to the need for developing further analysis methods. Experimental verification of the analysis outlined in this Part I is given in Part II of the paper.     

Solution of the three-dimensional problem of wave diffraction by an ensemble of objects

The pattern equations method is extended to solving three-dimensional problems of wave diffraction by an ensemble of bodies. The method is based on the reduction of the initial problem to a system of N (N is the number of scatterers in the ensemble) integro-operator equations of the second kind for the scattering patterns of scatterers. With the use of the series expansions of the scattering patterns in angular spherical harmonics, the problem is reduced to an algebraic system of equations in the expansion coefficients. An explicit (asymptotic) solution to the problems is obtained in the case when the scattering bodies are separated by sufficiently long distances. It is shown that the method can be used to model the characteristics of wave scattering by complex-shaped bodies.     

Using acoustic distortion products to measure the cochlear amplifier gain on the basilar membrane.

Most models of the cochlea developed during the last decade have explained frequency selectivity and sensitivity of the cochlea at threshold by the use of power amplification of the acoustic wave on the basilar membrane. This power amplification has been referred to as the cochlear amplifier (CA). In this paper, a method to measure the cochlear amplifier gain as a function of position along the basilar membrane is derived from a simple model. Next, experimental evidence is presented that strongly restricts the properties of these proposed cochlear amplifier models. Specifically, it is shown that small signals generated by mechanical nonlinearities in the basilar membrane motion are not amplified during basilar membrane propagation, contrary to what would be expected from the cochlear amplifier hypotheses. This paper describes a method of measuring the cochlear power gain as a function of frequency and position, from the stapes to within 2 mm of the place corresponding to the frequency being measured. Experimental results in the cat indicate that the total gain of the cochlear amplifier, over the range of positions measured, must be less than 10 dB. The simplest interpretation of the experimental results is that there is no cochlear amplifier. The results suggest that the cochlea must achieve its frequency selectivity by some other means.     

基于有限差分法刚性圆柱体涡激振动响应特性数值研究

基于尾流振子模型对刚性圆柱体涡激振动响应特性进行数值研究.建立圆柱体结构振子及尾流振子之间的耦合方程,基于二阶精度中心差分格式对耦合模型先离散后迭代进行求解.对不同质量比及不同阻尼比圆柱体涡激振动响应的无量纲位移、无量纲升力、频率比及锁定区间等参数进行分析.结果表明数值方法可以很好地模拟刚性圆柱体的涡激振动响应特性.随着质量比的增加,锁定开始点逐渐延后,锁定结束点逐渐提前,锁定区间宽度逐渐变窄.     

Nitrate removal from groundwater using Amberlite IRN-78: Modelling the system

A laboratory study was conducted to investigate the ability of Amberlite IRN-78 ion-exchange resin for the removal of nitrate from different waters. Aim of the present study is to propose a simple model that simulates the equilibrium data for this ion-exchange system.Tests were performed using two synthetic waters and two groundwaters. In the case of synthetic water, experimental data are reported for the ion-exchange equilibria determination in the absence or presence of chloride. For both groundwater types, “Salvatierra city” and “Miajadas city”, an ion-exchange Langmuir-type model is proposed, taking into account the presence of other competitive anions in solution, specifically chloride, sulphate and bicarbonate. Although the model is very simple, it is consistent with the experimental results. Finally, a general Langmuir-type expression is proposed for all water types, synthetic or natural. This equation considers the presence of other anions in solution by means of the parameter C_0T.