In mammals, environmental sounds stimulate the auditory receptor, the cochlea, via vibrations of the stapes, the innermost of the middle ear ossicles. These vibrations produce displacement waves that travel on the elongated and spirally wound basilar membrane (BM). As they travel, waves grow in amplitude, reaching a maximum and then dying out. The location of maximum BM motion is a function of stimulus frequency, with high-frequency waves being localized to the “base” of the cochlea (near the stapes) and low-frequency waves approaching the “apex” of the cochlea. Thus each cochlear site has a characteristic frequency (CF), to which it responds maximally. BM vibrations produce motion of hair cell stereocilia, which gates stereociliar transduction channels leading to the generation of hair cell receptor potentials and the excitation of afferent auditory nerve fibers. At the base of the cochlea, BM motion exhibits a CF-specific and level-dependent compressive nonlinearity such that responses to low-level, near-CF stimuli are sensitive and sharply frequency-tuned and responses to intense stimuli are insensitive and poorly tuned. The high sensitivity and sharp-frequency tuning, as well as compression and other nonlinearities (two-tone suppression and intermodulation distortion), are highly labile, indicating the presence in normal cochleae of a positive feedback from the organ of Corti, the “cochlear amplifier.” This mechanism involves forces generated by the outer hair cells and controlled, directly or indirectly, by their transduction currents. At the apex of the cochlea, nonlinearities appear to be less prominent than at the base, perhaps implying that the cochlear amplifier plays a lesser role in determining apical mechanical responses to sound. Whether at the base or the apex, the properties of BM vibration adequately account for most frequency-specific properties of the responses to sound of auditory nerve fibers.

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Mechanics of the Mammalian Cochlea

Phantom auditory perception (tinnitus): mechanisms of generation and perception

The Evolution of Language

Otoacoustic emissions (OAEs) of all types are widely assumed to arise by a common mechanism: nonlinear electromechanical distortion within the cochlea. In this view, both stimulus-frequency (SFOAEs) and distortion-product emissions (DPOAEs) arise because nonlinearities in the mechanics act as "sources" of backward-traveling waves. This unified picture is tested by analyzing measurements of emission phase using a simple phenomenological description of the nonlinear re-emission process. The analysis framework is independent of the detailed form of the emission sources and the nonlinearities that produce them. The analysis demonstrates that the common assumption that SFOAEs originate by nonlinear distortion requires that SFOAE phase be essentially independent of frequency, in striking contradiction with experiment. This contradiction implies that evoked otoacoustic emissions arise by two fundamentally different mechanisms within the cochlea. These two mechanisms (linear reflection versus nonlinear distortion) are described and two broad classes of emissions--reflection-source and distortion-source emissions--are distinguished based on the mechanisms of their generation. The implications of this OAE taxonomy for the measurement, interpretation, and clinical use of otoacoustic emissions as noninvasive probes of cochlear function are discussed.

http://web.mit.edu/apg/papers/shera-guinan-taxonomy-JASA99.pdf

Evoked otoacoustic emissions arise by two fundamentally different mechanisms: A taxonomy for mammalian OAEs

Our movements can hinder our ability to sense the world. Movements can induce sensory input (for example, when you hit something) that is indistinguishable from the input that is caused by external agents (for example, when something hits you). It is critical for nervous systems to be able to differentiate between these two scenarios. A ubiquitous strategy is to route copies of movement commands to sensory structures. These signals, which are referred to as corollary discharge (CD), influence sensory processing in myriad ways. Here we review the CD circuits that have been uncovered by neurophysiological studies and suggest a functional taxonomic classification of CD across the animal kingdom. This broad understanding of CD circuits lays the groundwork for more challenging studies that combine neurophysiology and psychophysics to probe the role of CD in perception.

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Corollary discharge across the animal kingdom

Evolutionary perspectives invertebrates aspects of hearing among vertebrates anamniotes non-mammalian amniotes mammals epilogue.

The Evolutionary biology of hearing

A class of cochlear models which account for much of the characteristic variation with frequency of human otoacoustic emissions and hearing threshold microstructure is presented. The models are based upon wave reflections via distributed spatial cochlear inhomogeneities and tall and broad cochlear activity patterns, as suggested by Zweig and Shera [J. Acoust. Soc. Am. 98, 2018–2047 (1995)]. They successfully describe in particular the following features: (1) the characteristic quasiperiodic frequency variations (fine structures) of the hearing threshold, synchronous and click-evoked emissions, distortion-product emissions, and spontaneous emissions; (2) the relationships between these fine structures; and (3) the distortion product emission filter shape. All of the characteristic frequency spacings are approximately the same (0.4 bark) and are mainly determined by the phase behavior of the apical reflection function. The frequency spacings for spontaneous emissions and threshold microstructure are predicted to be the same, but some deviations from these values are predicted for synchronous and click-evoked and distortion-product emissions. The analysis of models is aided considerably by the use of the solutions of apical, and basal, moving solutions (basis functions) of the cochlear wave equation in the absence of inhomogeneities.

Modeling otoacoustic emission and hearing threshold fine structures

High-resolution measurements of distortion product otoacoustic emissions (DPOAEs) from three different experimental paradigms are shown to be in agreement with the implications of a realistic “two-source” cochlear model of DPOAE fine structure. The measurements of DPOAE amplitude and phase imply an interference phenomenon involving one source in the region of strong nonlinear interaction of the primary waves (the strong “overlap” or generation region), and the other source region around the DPOAE tonotopic place. The component from the DPOAE place can be larger than the one from the generator region. These findings are supported by the analysis of the onset and offset of the DPOAE when the higher-frequency primary is pulsed on and off. The two-source hypothesis was further tested by adding a third tone closer in frequency to the DPOAE which modifies the amplitude of the component from the DPOAE place and leaves the one from the generator region unchanged. The results agree well with the model prediction that the variation with frequency, and implied latency, of the phase of the DPOAE tonotopic-place component are greater than the corresponding quantities for the component from the generation region.

Experimental confirmation of the two-source interference model for the fine structure of distortion product otoacoustic emissions

Distortion product otoacoustic emission (DPOAE) level from normal hearing individuals can vary by as much as 30dB with small frequency changes (a phenomenon known as DPOAE fine structure). This fine structure is hypothesized to stem from the interaction of components from two different regions of the cochlea (the nonlinear generator region and the reflection component from the DP region). An efficient procedure to separate these two components would improve the clinical and research utility of DPOAE by permitting separate evaluation of different cochlea regions. In this paper, two procedures for evaluating DPOAE fine structure are compared: DPOAE generated by fixed-frequency primaries versus continuously sweeping primaries. The sweep DPOAE data are analyzed with a least squares fit filter. Sweep rates of greater than 8s per octave permit rapid evaluation of the cochlear fine structure. A higher sweep rate of 2s per octave provided DPOAE without fine structure. Under these conditions, the longer latency reflection component falls outside the range of the filter. Consequently, DPOAE obtained with sweeping tones can be used either to get more rapid estimates of DPOAE fine structure or to obtain estimates of DPOAE from the generator region uncontaminated by energy from the reflection region.

Glenis R. Long

Papers

The Evolutionary biology of hearing

Modeling otoacoustic emission and hearing threshold fine structures

Experimental confirmation of the two-source interference model for the fine structure of distortion product otoacoustic emissions

Measuring distortion product otoacoustic emissions using continuously sweeping primaries.

New off-line method for detecting spontaneous otoacoustic emissions in human subjects.