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.     

Allen-Fahey and related experiments support the predominance of cochlear slow-wave otoacoustic emissions

Originally proposed as a method for measuring the power gain of the cochlear amplifier, Allen-Fahey experiments compare intracochlear distortion products and ear-canal otoacoustic emissions (OAEs) under tightly controlled conditions. In this paper Allen-Fahey experiments are shown to place significant constraints on the dominant mode of reverse energy propagation within the cochlea. Existing Allen-Fahey experiments are reviewed and shown to contradict the predictions of compression-wave OAE models recently proposed in the literature. In compression-wave models, distortion products propagate from their site of generation to the stapes via longitudinal compression waves in the cochlear fluids (fast waves); in transverse traveling-wave models, by contrast, distortion products propagate primarily via pressure-difference waves whose velocity and other characteristics depend on the mechanical properties of the cochlear partition (slow waves). Compression-wave models predict that the distortion-product OAEs (DPOAEs) measured in the Allen-Fahey paradigm increase at close primary-frequency ratios (or remain constant in the hypothetical absence of tuned suppression). The behavior observed experimentally is just the opposite-a pronounced decrease in DPOAE amplitude at close ratios. Since neither compression-wave nor simple conceptual "hybrid-wave" models can account for the experimental results--whereas slow-wave models can, via systematic changes in distortion-source directionality arising from wave-interference effects--Allen-Fahey and related experiments provide compelling evidence against the predominance of compression-wave OAEs in mammalian cochlear mechanics.     

Maturation of the traveling-wave delay in the human cochlea

The maturation of the traveling-wave delay in the human cochlea was investigated in 227 subjects ranging in age from 29 weeks conceptional age to 49 years by using frequency specific auditory brain-stem responses (ABRs). The derived response technique was applied to ABRs obtained with click stimuli (presented at a fixed level equal to 60-dB sensation level in normal hearing adults) in the presence of high-pass noise masking (slope 96 dB/oct) to obtain frequency specific responses from octave-wide bands. The estimate of traveling-wave delay was obtained by taking the difference between wave I latencies from adjacent derived bands. It was found that the traveling-wave delay between the octave band with center frequency (CF) of 11.3 kHz and that with CF of 5.7 kHz decreased (about 0.4 ms on average) in exponential fashion with age to reach adult values at 3-6 months of age. This decrease was in agreement with reported data in kitten auditory-nerve fibers. The traveling-wave delays between adjacent octave bands with successive lower CF did not change with age.     

Medial olivocochlear efferent inhibition of basilar-membrane responses to clicks: evidence for two modes of cochlear mechanical excitation

Conceptualizations of mammalian cochlear mechanics are based on basilar-membrane (BM) traveling waves that scale with frequency along the length of the cochlea, are amplified by outer hair cells (OHCs), and excite inner hair cells and auditory-nerve (AN) fibers in a simple way. However, recent experimental work has shown medial-olivocochlear (MOC) inhibition of AN responses to clicks that do not fit with this picture. To test whether this AN-initial-peak (ANIP) inhibition might result from hitherto unrecognized aspects of the traveling-wave or MOC-evoked inhibition, MOC effects on BM responses to clicks in the basal turns of guinea pig and chinchilla cochleae were measured. MOC stimulation inhibited BM click responses in a time and level dependent manner. Inhibition was not seen during the first half-cycle of the responses, but built up gradually, and ultimately increased the responses' decay rates. MOC stimulation also produced small phase leads in the response wave forms, but had little effect on the instantaneous frequency or the waxing and waning of the responses. These data, plus recent AN data, support the hypothesis that the MOC-evoked inhibitions of the traveling wave and of the ANIP response are separate phenomena, and indicate that the OHCs can affect at least two separate modes of excitation in the mammalian cochlea.     

Cochlear traveling-wave amplification, suppression, and beamforming probed using noninvasive calibration of intracochlear distortion sources

Originally developed to estimate the power gain of the cochlear amplifier, so-called "Allen-Fahey" and related experiments have proved invaluable for probing the mechanisms of wave generation and propagation within the cochlea. The experimental protocol requires simultaneous measurement of intracochlear distortion products (DPs) and ear-canal otoacoustic emissions (DPOAEs) under tightly controlled conditions. To calibrate the intracochlear response to the DP, Allen-Fahey experiments traditionally employ invasive procedures such as recording from auditory-nerve fibers or measuring basilar-membrane velocity. This paper describes an alternative method that allows the intracochlear distortion source to be calibrated noninvasively. In addition to the standard pair of primary tones used to generate the principal DP the noninvasive method employs a third, fixed tone to create a secondary DPOAE whose amplitude and phase provide a sensitive assay of the intracochlear value of the principal DP near its characteristic place. The method is used to perform noninvasive Allen-Fahey experiments in cat and shown to yield results in quantitative agreement with the original, auditory-nerve-based paradigm performed in the same animal. Data obtained using a suppression-compensated variation of the noninvasive method demonstrate that neither traveling-wave amplification nor two-tone suppression constitutes the controlling influence in DPOAE generation at close frequency ratios. Rather, the dominant factor governing the emission magnitude appears to be the variable directionality of the waves radiated by the distortion-source region, which acts as a distortion beamformer tuned by the primary frequency ratio.     

Medial-olivocochlear-efferent inhibition of the first peak of auditory-nerve responses: evidence for a new motion within the cochlea

Despite the insights obtained from click responses, the effects of medial-olivocochlear (MOC) efferents on click responses from single-auditory-nerve (AN) fibers have not been reported. We recorded responses of cat single AN fibers to randomized click level series with and without electrical stimulation of MOC efferents. MOC stimulation inhibited (1) the whole response at low sound levels, (2) the decaying part of the response at all sound levels, and (3) the first peak of the response at moderate to high sound levels. The first two effects were expected from previous reports using tones and are consistent with a MOC-induced reduction of cochlear amplification. The inhibition of the AN first peak, which was strongest in the apex and middle of the cochlea, was unexpected because the first peak of the classic basilar-membrane (BM) traveling wave receives little or no amplification. In the cochlear base, the click data were ambiguous, but tone data showed particularly short group delays in the tail-frequency region that is strongly inhibited by MOC efferents. Overall, the data support the hypothesis that there is a motion that bends inner-hair-cell stereocilia and can be inhibited by MOC efferents, a motion that is present through most, or all, of the cochlea and for which there is no counterpart in the classic BM traveling wave.     

Investigation of the photon statistics of parametric fluorescence in a traveling-wave parametric amplifier by means of self-homodyne tomography

Photon-number distributions for parametric fluorescence from a nondegenerate optical parametric amplifier are measured with a novel self-homodyne technique. These distributions exhibit the thermal-state character predicted by theory. However, a difference between the fluorescence gain and the signal gain of the parametric amplifier is observed. We attribute this difference to a change in the signal-beam profile during the traveling-wave pulsed amplification process.     

Nonlinear effects in optical fibers

The basic differential equations necessary to represent nonlinear propagation of short and ultrashort optical pulses in dielectric waveguides are derived. After introducing fundamental and higher-order soliton solutions of the nonlinear Schroedinger equations, the possibilities of realizing soliton-based communication systems are discussed. Higher-order nonlinear effects associated with selj-Raman gain and optical amplification in connection with the use of active fibers are also described.     

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.     

Testing coherent reflection in chinchilla: Auditory-nerve responses predict stimulus-frequency emissions

Coherent-reflection theory explains the generation of stimulus-frequency and transient-evoked otoacoustic emissions by showing how they emerge from the coherent "backscattering" of forward-traveling waves by mechanical irregularities in the cochlear partition. Recent published measurements of stimulus-frequency otoacoustic emissions (SFOAEs) and estimates of near-threshold basilar-membrane (BM) responses derived from Wiener-kernel analysis of auditory-nerve responses allow for comprehensive tests of the theory in chinchilla. Model predictions are based on (1) an approximate analytic expression for the SFOAE signal in terms of the BM traveling wave and its complex wave number, (2) an inversion procedure that derives the wave number from BM traveling waves, and (3) estimates of BM traveling waves obtained from the Wiener-kernel data and local scaling assumptions. At frequencies above 4 kHz, predicted median SFOAE phase-gradient delays and the general shapes of SFOAE magnitude-versus-frequency curves are in excellent agreement with the measurements. At frequencies below 4 kHz, both the magnitude and the phase of chinchilla SFOAEs show strong evidence of interference between short- and long-latency components. Approximate unmixing of these components, and association of the long-latency component with the predicted SFOAE, yields close agreement throughout the cochlea. Possible candidates for the short-latency SFOAE component, including wave-fixed distortion, are considered. Both empirical and predicted delay ratios (long-latency SFOAE delay/BM delay) are significantly less than 2 but greater than 1. Although these delay ratios contradict models in which SFOAE generators couple primarily into cochlear compression waves, they are consistent with the notion that forward and reverse energy propagation in the cochlea occurs predominantly by means of traveling pressure-difference waves. The compelling overall agreement between measured and predicted delays suggests that the coherent-reflection model captures the dominant mechanisms responsible for the generation of reflection-source otoacoustic emissions.     

Frequency-specific maturation of the eighth nerve and brain-stem auditory pathway: evidence from derived auditory brain-stem responses (ABRs).

Previous studies of human auditory development using frequency-specific auditory brain-stem responses (ABRs) have reported that maturation for both peak and interpeak latencies occurs earlier for responses generated by low-frequency stimuli. In two of these studies, low-frequency ABRs presumed to originate from apical locations in the cochlea were likely dominated by activity from higher frequency regions closer to the base. In the present study, the high-pass noise-masking technique was used to generate derived ABRs that represent activity from isolated place specific regions along the basilar membrane. Analysis of auditory brain-stem maturation based on I-V interpeak latency differences with adult means revealed a frequency-specific pattern of development. Developmental changes occurred faster and mature function was attained earlier for ABRs from the mid-center-frequency (CF) derived conditions than from either the highest or lowest CF derived conditions. The differential maturation of mid-CF derived ABRs may reflect the delayed effects of the pattern of development that occurs in the cochlea.     

The application of a capacitive probe technique for direct observation of electromechanical processes in the guinea pig cochlea

The demonstration of evoked mechanical responses of the outer hair cells in the mammalian cochlea by indirect measurements introduces a new range of problems into direct mechanical measurements. Direct and indirect measurements indicate that the frequency spectra of evoked electromechanical responses may extend well into the range of audio frequencies, revealing a need to develop terminology and protocols for distinguishing evoked mechanical responses from the traditional traveling wave when both are apparently superimposed on the motion of the basilar membrane in the normally functioning cochlea. Details are presented of a frequency-modulation capacitive probe technique for measurement of vibrating structures of the guinea pig ear. Considerations include the design of the transducer, calibration, sensitivity, linearity, and sources of noise, as well as the influence of the technique upon the animal preparation, and in particular the issues associated with draining scala tympani for the measurement. Relative advantages and disadvantages of the technique are compared with salient features of other techniques currently available. In view of the apparent complexity of cochlear mechanics some preliminary experiments are required to elucidate some of the key questions about reverse-transduction processes in general. A "simple" first experiment is to test existence of any rectifying or motile response.     

New Evidence of Active Tuning in Cochlea

The wonderful performance of hearing systems is mainly attributed to the tuning tiltering of basilar membrane （BM）. Although theory of the cochlear mechanism has been greatly developed since the 1970s and the amplification or sensitivity of the cochlea has been concluded due to the out hair cells, the mechanics underlying the sharp-tuning or frequency selectivity of cochlea remains a puzzle. We use the cochlear translation function derived from the data of an experiment of the BM in vivo to calculate basilar responses to tone bursts, and find that there are resonant peaks with the characteristic frequency at the corresponding place in the initial and terminal part of the responses. However, when the translation function is shallower, there will be no resonant peaks in the responses. The result indicates that the sharp tuning is due to existence of the active resonant tuning mechanism.     

Analysis of waveguiding properties of traveling-wave semiconductor laser amplifiers using perturbation technique

The polarization sensitivities of traveling-wave amplifiers (TWAs) can be analyzed by determining the modal gain coefficients for different propagating modes. We applied the perturbation method to introduce a new technique for the calculation of these gain coefficients, which avoids the use of the optical confinement factor. It was found that this technique was particularly useful in deducing the gain dependence on polarization for hybrid mode propagation. Using the proposed technique, we discovered that the refractive index distribution has a significant effect on the polarization sensitivities of hybrid modes in TWAs, especially in buried heterostructures (BHs). This is a promising method in designing polarization-insensitive TWAs.     

Stimulus-frequency-emission group delay: a test of coherent reflection filtering and a window on cochlear tuning

This paper tests and applies a key prediction of the theory of coherent reflection filtering for the generation of reflection-source otoacoustic emissions. The theory predicts that reflection-source-emission group delay is determined by the group delay of the basilar-membrane (BM) transfer function at its peak. This prediction is tested over a seven-octave frequency range in cats and guinea pigs using measurements of stimulus-frequency-emission (SFOAE) group delay. A comparison with group delays calculated from published measurements of BM mechanical transfer functions supports the theory at the basal end of the cochlea. A comparison across the whole frequency range based on variations in the sharpness of neural tuning with characteristic frequency (CF) suggests that the predicted relation holds in the basal-most 60% of the cochlea. At the apical end of the cochlea, however, the measurements disagree with neural and mechanical group delays. This disagreement suggests that there are important differences in cochlear mechanics and/or mechanisms of emission generation between the base and apex of the cochlea. Measurements in humans over a four-octave range indicate that human SFOAE group delays are roughly a factor of 3 longer than their counterparts in cat and guinea pig but manifest similar trends across CF. The measurements thus reveal global deviations from scaling whose form appears quantitatively similar in all three species. Interpreted using the theory of coherent reflection filtering, the group delay measurements indicate that the wavelength at the peak of the traveling wave decreases with increasing CF at a rate of roughly 25% per octave in the base of the cochlea. The measurements and analysis reported here illustrate the rich potential inherent in OAE measurements for obtaining valuable information about basic cochlear properties such as tuning.     

A model for active elements in cochlear biomechanics

A linear, mathematical model of cochlear biomechanics is presented in this paper. In this model, active elements are essential for simulating the high sensitivity and sharp tuning characteristic of the mammalian cochlea. The active elements are intended to represent the motile action of outer hair cells; they are postulated to be mechanical force generators that are powered by electrochemical energy of the cochlear endolymph, controlled by the bending of outer hair cell stereocilia, and bidirectionally coupled to cochlear partition mechanics. The active elements are spatially distributed and function collectively as a cochlear amplifier. Excessive gain in the cochlear amplifier causes spontaneous oscillations and thereby generates spontaneous otoacoustic emissions.     

Propagation characteristics of biphotons in cold atomic vapor

We investigate the propagation characteristics of the narrowband Stokes/anti-Stokes photons in cold atomic vapor. The four-wave mixing process results from parametric amplification of the anti-Stokes photons. We find that the process of parametric amplification is very similar to the light pulse propagating through an anomalous dispersion gain medium. Finally, we obtain the general solutions of the Glauber biphoton correlation functions, which are in good agreement with the experiment results.     

Quantum noise measurements in a continuous-wave laser-diode-pumped Nd:YAG saturated amplifier

We present measurements of the power noise due to optical amplification in a Nd:YAG free-space traveling-wave amplifier as the amplifier transitions from the linear regime into the heavily saturated regime. The quantum noise behavior is demonstrated by saturating the gain of a 100-W class zigzag slab amplifier with a high-power beam and measuring the power noise detected by a single-spatial-mode probe beam traversing the same optical path through the amplifier.     

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.     

Vibration responses of the organ of Corti and the tectorial membrane to electrical stimulation

Coupling of somatic electromechanical force from the outer hair cells (OHCs) into the organ of Corti is investigated by measuring transverse vibration patterns of the organ of Cori and tectorial membrane (TM) in response to intracochlear electrical stimulation. Measurement places at the organ of Corti extend from the inner sulcus cells to Hensen's cells and at the lower (and upper) surface of the TM from the inner sulcus to the OHC region. These locations are in the neighborhood of where electromechanical force is coupled into (1) the mechanoelectrical transducers of the stereocilia and (2) fluids of the organ of Corti. Experiments are conducted in the first, second, and third cochlear turns of an in vitro preparation of the adult guinea pig cochlea. Vibration measurements are made at functionally relevant stimulus frequencies (0.48-68 kHz) and response amplitudes (<15 nm). The experiments provide phase relations between the different structures, which, dependent on frequency range and longitudinal cochlear position, include in-phase transverse motions of the TM, counterphasic transverse motions between the inner hair cell and OHCs, as well as traveling-wave motion of Hensen's cells in the radial direction. Mechanics of sound processing in the cochlea are discussed based on these phase relationships.