Near equivalence of human click-evoked and stimulus-frequency otoacoustic emissions

Otoacoustic emissions (OAEs) evoked by broadband clicks and by single tones are widely regarded as originating via different mechanisms within the cochlea. Whereas the properties of stimulus-frequency OAEs (SFOAEs) evoked by tones are consistent with an origin via linear mechanisms involving coherent wave scattering by preexisting perturbations in the mechanics, OAEs evoked by broadband clicks (CEOAEs) have been suggested to originate via nonlinear interactions among the different frequency components of the stimulus (e.g., intermodulation distortion). The experiments reported here test for bandwidth-dependent differences in mechanisms of OAE generation. Click-evoked and stimulus-frequency OAE input/output transfer functions were obtained and compared as a function of stimulus frequency and intensity. At low and moderate intensities human CEOAE and SFOAE transfer functions are nearly identical. When stimulus intensity is measured in "bandwidth-compensated" sound-pressure level (cSPL), CEOAE and SFOAE transfer functions have equivalent growth functions at fixed frequency and equivalent spectral characteristics at fixed intensity. This equivalence suggests that CEOAEs and SFOAEs are generated by the same mechanism. Although CEOAEs and SFOAEs are known by different names because of the different stimuli used to evoke them, the two OAE "types" are evidently best understood as members of the same emission family.     

Laser amplification with a twist: traveling-wave propagation and gain functions from throughout the cochlea

Except at the handful of sites explored by the inverse method, the characteristics-indeed, the very existence-of traveling-wave amplification in the mammalian cochlea remain largely unknown. Uncertainties are especially pronounced in the apex, where mechanical and electrical measurements lack the independent controls necessary for assessing damage to the preparation. At a functional level, the form and amplification of cochlear traveling waves are described by quantities known as propagation and gain functions. A method for deriving propagation and gain functions from basilar-membrane mechanical transfer functions is presented and validated by response reconstruction. Empirical propagation and gain functions from locations throughout the cochlea are obtained in mechanically undamaged preparations by applying the method to published estimates of near-threshold basilar membrane responses derived from Wiener-kernel (chinchilla) and zwuis analysis (cat) of auditory-nerve responses to broadband stimuli. The properties of these functions, and their variation along the length of the cochlea, are described. In both species, and at all locations examined, the gain functions reveal a region of positive power gain basal to the wave peak. The results establish the existence of traveling-wave amplification throughout the cochlea, including the apex. The derived propagation and gain functions resemble those characteristic of an active optical medium but rotated by 90 degrees in the complex plane. Rotation of the propagation and gain functions enables the mammalian cochlea to operate as a wideband, hydromechanical laser analyzer.     

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.     

Nuclear levels in152Eu

The transitional nucleus¹⁵²Eu has been studied using the (n, e), (n, γ), (n _res,γ), (n, γγ), (d, p), (d, t) and (p, d) reactions. The experiments have been performed at nine different laboratories. A model independent level scheme was established including 95 levels below 510 keV and nearly 900 transitions by combination of low energy transitions and reaction data. More than 20 additional levels result from gamma rays and/or charged particle reactions. The level scheme is interpreted in terms of the Nilsson model indicating that¹⁵²Eu is a deformed nucleus. Seven rotational bands and Nilsson configurations are established. An additional 27 rotational bands are tentatively or speculatively assugned. Gallagher-Moszkowski splittings are discussed. The neutron binding energy was determined as 6305.2±0.5 keV. The energy of the 9.3 h 0^? isomer is 45.599 keV. The lifetimes of four levels were measured. Nuclear Reactions¹⁵¹Eu(n,γ),E _n =thermal and resonance; measuredE _γ ,I _γ ,E _c.e.,I _c.e.,γγ Coinc.,γγΔt coinc.;¹⁵¹Eu(d, p),E=12MeV and 14MeV;¹⁵³Eu(d, t),E=12MeV;¹⁵³Eu(p, d),E =18MeV; deduced level scheme of¹⁵²Eu,J, π, T _1/2,cc, Nilsson configurations. Magnetic electron spectrometer, curved crystal spectrometer, Ge(Li) and Si(Li) detectors, magnetic spectrographs. Enriched targets.     

Reflection of retrograde waves within the cochlea and at the stapes

A number of authors [de Boer and Viergever, Hear. Res. 13, 101-112 (1984); de Boer et al., in Peripheral Auditory Mechanisms (Springer-Verlag, Berlin, 1986); Hear. Res. 23, 1-7 (1986); Viergever, in Auditory Frequency Selectivity (Plenum, New York, 1986), pp. 31-38; Kaernbach et al., J. Acoust. Soc. Am. 81, 408-411 (1987)] have argued that backward-traveling waves, in striking contrast to waves traveling forward towards the helicotrema, suffer appreciable reflection as they move through the basal turns of the cochlea. Such reflection, if present, would have important consequences for understanding the nature and strength of otoacoustic emissions. The apparent asymmetry in reflection of cochlear waves is shown, however, to be an artifact of the boundary condition those authors impose at the stapes: conventional cochlear models are found not to generate reflections of waves traveling in either direction even when the wavelength changes rapidly and the WKB approximation breaks down. Although backward-traveling waves are not reflected by the secular variation of the geometrical and mechanical characteristics of the cochlea, they are reflected when they reach the stapes. The magnitude of that boundary reflection is computed for the cat and shown to be a large, rapidly varying function of frequency.     

Intensity-invariance of fine time structure in basilar-membrane click responses: implications for cochlear mechanics

Basilar-membrane and auditory-nerve responses to impulsive acoustic stimuli, whether measured directly in response to clicks or obtained indirectly using cross- or reverse-correlation and/or Fourier analysis, manifest a striking symmetry: near-invariance with stimulus intensity of the fine time structure of the response over almost the entire dynamic range of hearing. This paper explores the origin and implications of this symmetry for cochlear mechanics. Intensity-invariance is investigated by applying the EQ-NL theorem [de Boer, Aud. Neurosci. 3, 377-388 (1997)] to define a family of linear cochlear models in which the strength of the active force generators is controlled by a real-valued, intensity-dependent parameter, gamma (with 0 < or = gamma < or = 1). The invariance of fine time structure is conjectured to imply that as gamma is varied the poles of the admittance of the cochlear partition remain within relatively narrow bands of the complex plane oriented perpendicular to the real frequency axis. Physically, the conjecture implies that the local resonant frequencies of the cochlear partition are nearly independent of intensity. Cochlear-model responses, computed by extending the model obtained by solution of the inverse problem in squirrel monkey at low sound levels [Zweig, J. Acoust. Soc. Am. 89, 1229-1254 (1991)] with three different forms of the intensity dependence of the partition admittance, support the conjecture. Intensity-invariance of cochlear resonant frequencies is shown to be consistent with the well-known "half-octave shift," describing the shift with intensity in the peak (or best) frequency of the basilar-membrane frequency response. Shifts in best frequency do not arise locally, via changes in the underlying resonant frequencies of the partition, but globally through the intensity dependence of the driving pressure. Near-invariance of fine time structure places strong constraints on the mechanical effects of force generation by outer hair cells. In particular, the symmetry requires that the feedback forces generated by outer hair cells (OHCs) not significantly affect the natural resonant frequencies of the cochlear partition. These results contradict many, if not most, cochlear models, in which OHC forces produce significant changes in the reactance and resonant frequencies of the partition.     

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.