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1.  Mechanics of the Mammalian Cochlea 
Physiological reviews  2001;81(3):1305-1352.
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.
PMCID: PMC3590856  PMID: 11427697
2.  Two-Tone Distortion on the Basilar Membrane of the Chinchilla Cochlea 
Journal of neurophysiology  1997;77(5):2385-2399.
Basilar membrane responses to pairs of tones were measured, with the use of a laser velocimeter, in the basal turn of the cochlea in anesthetized chinchillas. Frequency spectra of basilar membrane responses to primary tones with frequencies (f1, f2) close to the characteristic frequency (CF) contain prominent odd-order two-tone distortion products (DPs) at frequencies both higher and lower than CF (such as 2f1 − f2, 3f − 2f2, 2f2 − f1 and 3f2 − 2f1). For equal-level primaries with frequencies such that 2f1 − f2 equals CF, the magnitude of the 2f1 − f2 DP grows with primary level at linear or faster rates at low stimulus levels, but it saturates or decreases slightly at higher levels. For a fixed level of one of the primary tones, the magnitude of the 2f1 − f2 DP is a nonmonotonic function of the level of the other primary tone. For low intensities of the variable tone, the grows at a rate of 2f1 − f2 DP grows at a rate of ~2 dB/dB with f1 level and 1 dB/dB with f2 level. DP magnitudes decrease rapidly with increasing primary frequency ratio (f2/f1) at low stimulus levels. For more intense stimuli, DP magnitudes remain constant or decrease slowly over a wide range of frequency ratios until a critical value is reached, at which DP magnitudes fall with slopes as steep as −300 dB/octave. As stimulus level grows, DP phases increasingly lag large f2/f1 ratios, but exhibit leads for small f2/f1 ratios. Cochlear exposure to an intense tone that produces large sensitivity losses for the primary frequencies (but only small losses for tones with frequency equal to 2f1 − f2) causes a substantial decrease in magnitude of the 2f1 − f2 DP. This result demonstrates that the 2f1 − f2 DP originates at the basilar membrane region with CFs corresponding to the primary frequencies and propagates to the location with CF equal to the DP frequency. 2f1 − f2 DPs on the basilar membrane resemble those measured in human psychophysics in most respects. However, the magnitude of basilar membrane DPs does not show the nonmonotonic dependence on f2/f1 ratio evident in DP otoacoustic emissions.
PMCID: PMC3582226  PMID: 9163365
3.  Basilar-membrane responses to clicks at the base of the chinchilla cochlea 
Basilar-membrane responses to clicks were measured, using laser velocimetry, at a site of the chinchilla cochlea located about 3.5 mm from the oval window (characteristic frequency or CF: typically 8–10 kHz). They consisted of relatively undamped oscillations with instantaneous frequency that increased rapidly (time constant: 200 µs) from a few kHz to CF. Such frequency modulation was evident regardless of stimulus level and was also present post-mortem. Responses grew linearly at low stimulus levels, but exhibited a compressive nonlinearity at higher levels. Velocity-intensity functions were almost linear near response onset but became nonlinear within 100 µs. Slopes could be as low as 0.1–0.2 dB/dB at later times. Hence, the response envelopes became increasingly skewed at higher stimulus levels, with their center of gravity shifting to earlier times. The phases of near-CF response components changed by nearly 180 degrees as a function of time. At high stimulus levels, this generated cancellation notches and phase jumps in the frequency spectra. With increases in click level, sharpness of tuning deteriorated and the spectral maximum shifted to lower frequencies. Response phases also changed as a function of increasing stimulus intensity, exhibiting relative lags and leads at frequencies somewhat lower and higher than CF, respectively. In most respects, the magnitude and phase frequency spectra of responses to clicks closely resembled those of responses to tones. Post-mortem responses were similar to in vivo responses to very intense clicks.
PMCID: PMC3582372  PMID: 9566320
4.  Middle-ear response in the chinchilla and its relationship to mechanics at the base of the cochleaa) 
The responses of the malleus and the stapes to sinusoidal acoustic stimulation have been measured in the middle ears of anesthetized chinchillas using the Mössbauer technique. With “intact” bullas (i.e., closed except for venting via capillary tubing), the vibrations of the tip of the malleus reach a maximal peak velocity of about 2 mm/s in responses to 100-dB SPL tones in the frequency range 500–6000 Hz; vibration velocity diminishes toward lower frequencies with a slope of about 6 dB/oct. Opening the bulla widely increases the responses to low-frequency stimuli by as much as 16 dB. At low frequencies, malleus response sensitivity with either open or intact bullas far exceeds all previous measurements in cats and matches or exceeds such measurements in guinea pigs. Whether measured in open or intact bullas, phase-versus-frequency curves closely approximate those predicted from the magnitude-versus-frequency curves by minimum phase theory. The stapes responses are similar to those of the malleus, except that stapes response magnitude is lower, on the average, by 7.5 dB at frequencies below 2 kHz and 10.7 dB at 2 kHz and above. Comparison of the responses of the middle ear with those of the basilar membrane at a site 3.5 mm from the stapes indicates that, at frequencies below 150 Hz, the basilar membrane displacement is proportional to stapes acceleration. At frequencies between 150 and 2000 Hz, basilar membrane displacement is proportional to stapes velocity.
PMCID: PMC3580955  PMID: 2341666
5.  Furosemide Alters Organ of Corti Mechanics: Evidence for Feedback of Outer Hair Cells upon the Basilar Membrane 
A widely held hypothesis of mammalian cochlear function is that the mechanical responses to sound of the basilar membrane depend on transduction by the outer hair cells. We have tested this hypothesis by studying the effect upon basilar membrane vibrations (measured by means of either the Mössbauer technique or Doppler-shift laser velocimetry) of systemic injection of furosemide, a loop diuretic that decreases transduction currents in hair cells. Furosemide reversibly altered the responses to tones and clicks of the chinchilla basilar membrane, causing response-magnitude reductions that were largest (up to 61 dB, averaging 25-30 dB) at low stimulus intensities at the characteristic frequency (CF) and small or nonexistent at high intensities and at frequencies far removed from CF. Furosemide also induced response-phase lags that were largest at low stimulus intensities (averaging 77°) and were confined to frequencies close to CF. These results constitute the most definitive demonstration to date that mechanical responses of the basilar membrane are dependent on the normal function of the organ of Corti and strongly implicate the outer hair cells as being responsible for the high sensitivity and frequency selectivity of basilar membrane responses. A corollary of these findings is that sensorineural hearing deficits in humans due to outer hair cell loss reflect pathologically diminished vibrations of the basilar membrane.
PMCID: PMC3580957  PMID: 2010805
6.  Responses to sound of the basilar membrane of the mammalian cochlea 
Current opinion in neurobiology  1992;2(4):449-456.
Recent evidence shows that the frequency-specific non-linear properties of auditory nerve and inner hair cell responses to sound, including their sharp frequency tuning, are fully established in the vibration of the basilar membrane. In turn, the sensitivity, frequency selectivity and non-linear properties of basilar membrane responses probably result from an influence of the outer hair cells.
PMCID: PMC3579517  PMID: 1525542
7.  Two-tone distortion in the basilar membrane of the cochlea 
Nature  1991;349(6308):413-414.
When humans listen to pairs of thnes they hear additional tones, or distortion products, that are not present in the stimulus1. Two-tone distortion products are also known as combination tones, because their pitches match combinations of the primary frequencies (f1 and f2, f2 > f1), such as f2 – f1, (n + 1)f1 – nf2 and (n + 1)f2 – nf1 (n = 1,2,3 …) (refs 2–4). Physiological correlates of the perceived distortion products exist in responses of auditory-nerve fibres5–8 and inner hair cells9 and in otoacoustic emissions (sounds generated by the cochlea, recordable at the ear canal)7,10–12. Because the middle ear responds linearly to sound13,14 and neural responses to distortion products can be abolished by damage to hair cells at cochlear sites preferentially tuned to the frequencies of the primary tones8, it was hypothesized that distortion products are generated at these sites and propagate mechanically along the basilar membrane to the location tuned to the distortion-product frequency7,8. But until now, efforts to confirm this hypothesis have failed15,16. Here we report the use of a new laser-velocimetry technique17 to demonstrate two-tone distortion in basilar-membrane motion at low and moderate stimulus intensities.
doi:10.1038/349413a0
PMCID: PMC3579518  PMID: 1992342
8.  Application of a commercially-manufactured Doppler-shift laser velocimeter to the measurement of basilar-membrane vibration* 
Hearing research  1991;51(2):215-230.
A commercially-available laser Doppler-shift velocimeter has been coupled to a compound microscope equipped with ultra-long-working-distance objectives for the purpose of measuring basilar membrane vibrations in the chinchilla. The animal preparation is nearly identical to that used in our laboratory for similar measurements using the Mössbauer technique. The vibrometer head is mounted on the third tube of the microscope’s trinocular head and its laser beam is focused on high-refractive-index glass microbeads (10–30 µm) previously dropped, through the perilymph of Scala tympani, on the basilar membrane. For equal sampling times, overall sensitivity of the laser velocimetry system is at least one order of magnitude greater than usually attained using the Mössbauer technique. However, the most important advantage of laser velocimetry vis-à-vis the Mössbauer technique is its linearity, which permits undistorted recording of signals over a wide velocity range. Thus, for example, we have measured basilar-membrane responses to clicks whose waveforms have dynamic ranges exceeding 60 dB.
PMCID: PMC3579526  PMID: 1827787
Laser Doppler-shift velocimetry; Laser vibrometry; Laser heterodyne interferometry; Basilar membrane; Cochlear mechanics
9.  Cochlear Delays and Traveling Waves: Comments on ‘Experimental Look at Cochlear Mechanics’ 
A. Dancer argued that direct and indirect measures of basilar membrane motion are more consistent with theories of cochlear resonance than with the traveling-wave theory. The present communication reviews empirical evidence that contradicts Dancer’s argument. Such evidence – recordings of mechanical responses of the basilar and Reissner’s membranes to sound – strongly supports the existence of displacement waves that propagate on the basilar membrane from the base of the cochlea toward its apex.
PMCID: PMC3579543  PMID: 8042934
Cochlear mechanics; Basilar membrane; Propagation delay
10.  Basilar membrane responses to two-tone and broadband stimuli 
SUMMARY
The responses to sound of mammalian cochlear neurons exhibit many nonlinearities, some of which (such as two-tone rate suppression and intermodulation distortion) are highly frequency specific, being strongly tuned to the characteristic frequency (cf) of the neuron. With the goal of establishing the cochlear origin of these auditory-nerve nonlinearities, mechanical responses to clicks and to pairs of tones were studied in relatively healthy chinchilla cochleae at a basal site of the basilar membrane with cf of 8–10 kHz. Responses were also obtained in cochleae in which hair cell receptor potentials were reduced by systemic furosemide injection. Vibrations were recorded using either the Mössbauer technique or laser Doppler-shift velocimetry. Responses to tone pairs contained intermodulation distortion products whose magnitudes as a function of stimulus frequency and intensity were comparable to those of distortion products in cochlear afferent responses. Responses to cf tones could be selectively suppressed by tones with frequency either higher or lower than cf; in most respects, mechanical two-tone suppression resembled rate suppression in cochlear afferents. Responses to clicks displayed a cf-specific compressive nonlinearity, similar to that present in responses to single tones, which could be profoundly and selectively reduced by furosemide. The present findings firmly support the hypothesis that all cf-specific nonlinearities present in the auditory nerve originate in analogous phenomena of basilar membrane vibration. However, because of their lability, it is almost certain that the mechanical nonlinearities themselves originate in outer hair cells.
doi:10.1098/rstb.1992.0063
PMCID: PMC3578387  PMID: 1354369
11.  Basilar-membrane responses to tones at the base of the chinchilla cochlea 
Basilar-membrane responses to single tones were measured, using laser velocimetry, at a site of the chinchilla cochlea located 3.5 mm from its basal end. Responses to low-level (<10–20 dB SPL) characteristic-frequency (CF) tones (9–10 kHz) grow linearly with stimulus intensity and exhibit gains of 66–76 dB relative to stapes motion. At higher levels, CF responses grow monotonically at compressive rates, with input–output slopes as low as 0.2 dB/dB in the intensity range 40–80 dB. Compressive growth, which is significantly correlated with response sensitivity, is evident even at stimulus levels higher than 100 dB. Responses become rapidly linear as stimulus frequency departs from CF. As a result, at stimulus levels >80 dB the largest responses are elicited by tones with frequency about 0.4–0.5 octave below CF. For stimulus frequencies well above CF, responses stop decreasing with increasing frequency: A plateau is reached. The compressive growth of responses to tones with frequency near CF is accompanied by intensity-dependent phase shifts. Death abolishes all nonlinearities, reduces sensitivity at CF by as much as 60–81 dB, and causes a relative phase lead at CF.
PMCID: PMC3578390  PMID: 9104018
12.  Frequency Tuning of Basilar Membrane and Auditory Nerve Fibers in the Same Cochleae 
Science (New York, N.Y.)  1998;282(5395):1882-1884.
Responses to tones of a basilar membrane site and of auditory nerve fibers innervating neighboring inner hair cells were recorded in the same cochleae in chinchillas. At near-threshold stimulus levels, the frequency tuning of auditory nerve fibers closely paralleled that of basilar membrane displacement modified by high-pass filtering, indicating that only relatively minor signal transformations intervene between mechanical vibration and auditory nerve excitation. This finding establishes that cochlear frequency selectivity in chinchillas (and probably in mammals in general) is fully expressed in the vibrations of the basilar membrane and renders unnecessary additional (“second”) filters, such as those present in the hair cells of the cochleae of reptiles.
PMCID: PMC3578392  PMID: 9836636
13.  Distortion in those good vibrations 
Current biology : CB  1993;3(11):755-758.
The features that make inner ear hair cells so sensitive to vibrations may also be responsible for the introduction of surprisingly large distortions.
PMCID: PMC3578393  PMID: 15335839
14.  Low-frequency suppression of auditory nerve responses to characteristic frequency tones 
Hearing research  1997;113(1-2):29-56.
The effects of low-frequency (50, 100, 200 and 400 Hz) ‘suppressor’ tones on responses to moderate-level characteristic frequency (CF) tones were measured in chinchilla auditory nerve fibers. Two-tone interactions were evident at suppressor intensities of 70–100 dB SPL. In this range, the average response rate decreased as a function of increasing suppressor level and the instantaneous response rate was modulated periodically. At suppression threshold, the phase of suppression typically coincided with basilar membrane displacement toward scala tympani, regardless of CF. At higher suppressor levels, two suppression maxima coexisted, synchronous with peak basilar membrane displacement toward scala tympani and scala vestibuli. Modulation and rate-suppression thresholds did not vary as a function of spontaneous activity and were only minimally correlated with fiber sensitivity. Except for fibers with CF < 1 kHz, modulation and rate-suppression thresholds were lower than rate and phase-locking thresholds for the suppressor tones presented alone. In the case of high-CF fibers with low spontaneous activity, excitation thresholds could exceed suppression thresholds by more than 30 dB. The strength of modulation decreased systematically with increasing suppressor frequency. For a given suppressor frequency, modulation was strongest in high-CF fibers and weakest in low-CF fibers. The present findings strongly support the notion that low-frequency suppression in auditory nerve fibers largely reflects an underlying basilar membrane phenomenon closely related to compressive non-linearity.
PMCID: PMC3578423  PMID: 9387984
Auditory nerve; Biasing; Modulation; Rate suppression; Basilar membrane; Inner hair cells; Cochlea; Chinchilla
15.  Basilar-Membrane Responses to Broadband Noise Modeled Using Linear Filters With Rational Transfer Functions 
Basilar-membrane responses to white Gaussian noise were recorded using laser velocimetry at basal sites of the chinchilla cochlea with characteristic frequencies near 10 kHz and first-order Wiener kernels were computed by cross correlation of the stimuli and the responses. The presence or absence of minimum-phase behavior was explored by fitting the kernels with discrete linear filters with rational transfer functions. Excellent fits to the kernels were obtained with filters with transfer functions including zeroes located outside the unit circle, implying nonminimum-phase behavior. These filters accurately predicted basilar-membrane responses to other noise stimuli presented at the same level as the stimulus for the kernel computation. Fits with all-pole and other minimum-phase discrete filters were inferior to fits with nonminimum-phase filters. Minimum-phase functions predicted from the amplitude functions of the Wiener kernels by Hilbert transforms were different from the measured phase curves. These results, which suggest that basilar-membrane responses do not have the minimum-phase property, challenge the validity of models of cochlear processing, which incorporate minimum-phase behavior.
doi:10.1109/TBME.2010.2052254
PMCID: PMC3572753  PMID: 20542757
Autoregressive moving-average (ARMA) modeling; basilar membrane (BM); cochlea; Hilbert transform; minimum phase; Wiener kernels
16.  Traveling waves on the organ of Corti of the chinchilla cochlea: spatial trajectories of inner hair cell depolarization inferred from responses of auditory-nerve fibers 
Spatial magnitude and phase profiles for inner hair cell depolarization throughout the chinchilla cochlea were inferred from responses of auditory-nerve fibers to threshold- and moderate-level tones and tone complexes. Firing-rate profiles for frequencies ≤ 2 kHz are bimodal, with the major peak at the characteristic place and a secondary peak at 3–5 mm from the extreme base. Response-phase trajectories are synchronous with peak outward stapes displacement at the extreme cochlear base and accumulate 1.5-period lags at the characteristic places. High-frequency phase trajectories are very similar to the trajectories of basilar-membrane peak velocity toward scala tympani. Low-frequency phase trajectories undergo a polarity flip in a region, 6.5–9 mm from the cochlear base, where traveling-wave phase velocity attains a local minimum and a local maximum and where the onset latencies of near-threshold impulse responses computed from responses to near-threshold white noise exhibit a local minimum. That region is the same where frequency-threshold tuning curves of auditory-nerve fibers undergo a shape transition. Since depolarization of inner hair cells presumably indicates the mechanical stimulus to their stereocilia, the present results suggest that distinct low-frequency forward waves of organ of Corti vibration are launched simultaneously at the extreme base of the cochlea and at the 6.5–9 mm transition region, from where antiphasic reflections arise.
doi:10.1523/JNEUROSCI.1138-12.2012
PMCID: PMC3436599  PMID: 22855802
17.  Timing of cochlear responses inferred from frequency-threshold tuning curves of auditory-nerve fibers 
Hearing research  2010;272(1-2):178-186.
Links between frequency tuning and timing were explored in the responses to sound of auditory-nerve fibers. Synthetic transfer functions were constructed by combining filter functions, derived via minimum-phase computations from average frequency-threshold tuning curves of chinchilla auditory-nerve fibers with high spontaneous activity (A. N. Temchin et al., J. Neurophysiol. 100: 2889–2898, 2008), and signal-front delays specified by the latencies of basilar-membrane and auditory-nerve fiber responses to intense clicks (A. N. Temchin et al., J. Neurophysiol. 93: 3635–3648, 2005). The transfer functions predict several features of the phase-frequency curves of cochlear responses to tones, including their shape transitions in the regions with characteristic frequencies of 1 kHz and 3–4 kHz (A. N. Temchin and M. A. Ruggero, JARO 11: 297–318, 2010). The transfer functions also predict the shapes of cochlear impulse responses, including the polarities of their frequency sweeps and their transition at characteristic frequencies around 1 kHz. Predictions are especially accurate for characteristic frequencies < 1 kHz.
doi:10.1016/j.heares.2010.10.002
PMCID: PMC3039049  PMID: 20951191
18.  Phase-Locked Responses to Tones of Chinchilla Auditory Nerve Fibers: Implications for Apical Cochlear Mechanics 
Responses to tones with frequency ≤ 5 kHz were recorded from auditory nerve fibers (ANFs) of anesthetized chinchillas. With increasing stimulus level, discharge rate–frequency functions shift toward higher and lower frequencies, respectively, for ANFs with characteristic frequencies (CFs) lower and higher than ∼0.9 kHz. With increasing frequency separation from CF, rate–level functions are less steep and/or saturate at lower rates than at CF, indicating a CF-specific nonlinearity. The strength of phase locking has lower high-frequency cutoffs for CFs >4 kHz than for CFs < 3 kHz. Phase–frequency functions of ANFs with CFs lower and higher than ∼0.9 kHz have inflections, respectively, at frequencies higher and lower than CF. For CFs >2 kHz, the inflections coincide with the tip-tail transitions of threshold tuning curves. ANF responses to CF tones exhibit cumulative phase lags of 1.5 periods for CFs 0.7–3 kHz and lesser amounts for lower CFs. With increases of stimulus level, responses increasingly lag (lead) lower-level responses at frequencies lower (higher) than CF, so that group delays are maximal at, or slightly above, CF. The CF-specific magnitude and phase nonlinearities of ANFs with CFs < 2.5 kHz span their entire response bandwidths. Several properties of ANFs undergo sharp transitions in the cochlear region with CFs 2–5 kHz. Overall, the responses of chinchilla ANFs resemble those in other mammalian species but contrast with available measurements of apical cochlear vibrations in chinchilla, implying that either the latter are flawed or that a nonlinear “second filter” is interposed between vibrations and ANF excitation.
doi:10.1007/s10162-009-0197-4
PMCID: PMC2862913  PMID: 19921334
basilar membrane; cochlear apex; phase–frequency functions; rate–frequency functions
19.  Basilar Membrane Responses to Noise at a Basal Site of the Chinchilla Cochlea: Quasi-Linear Filtering 
Basilar membrane responses to clicks and to white noise were recorded using laser velocimetry at basal sites of the chinchilla cochlea with characteristic frequencies near 10 kHz. Responses to noise grew at compressive rates and their instantaneous frequencies decreased with increasing stimulus level. First-order Wiener kernels were computed by cross-correlation of the noise stimuli and the responses. For linear systems, first-order Wiener kernels are identical to unit impulse responses. In the case of basilar membrane responses, first-order Wiener kernels and responses to clicks measured at the same sites were similar but not identical. Both consisted of transient oscillations with onset frequencies which increased rapidly, over about 0.5 ms, from 4–5 kHz to the characteristic frequency. Both first-order Wiener kernels and responses to clicks were more highly damped, exhibited slower frequency modulation, and grew at compressive rates with increasing stimulus levels. Responses to clicks had longer durations than the Wiener kernels. The statistical distribution of basilar membrane responses to Gaussian white noise is also Gaussian and the envelopes of the responses are Rayleigh distributed, as they should be for Gaussian noise passing through a linear band-pass filter. Accordingly, basilar membrane responses were accurately predicted by linear filters specified by the first-order Wiener kernels of responses to noise presented at the same level. Overall, the results indicate that cochlear nonlinearity is not instantaneous and resembles automatic gain control.
doi:10.1007/s10162-009-0172-0
PMCID: PMC2774406  PMID: 19495878
Wiener kernels; clicks; laser velocimetry; frequency glides; impulse responses
20.  Basilar Membrane Vibrations Near the Round Window of the Gerbil Cochlea 
Using a laser velocimeter, responses to tones were measured at a basilar membrane site located about 1.2 mm from the extreme basal end of the gerbil cochlea. In two exceptional cochleae in which function was only moderately disrupted by surgical preparations, basilar membrane responses had characteristic frequencies (CFs) of 34–37 kHz and exhibited a CF-specific compressive nonlinearity: Sensitivity near the CF decreased systematically and the response peaks shifted toward lower frequencies with increasing stimulus level. Response phases also changed with increases in stimulus level, exhibiting small relative lags and leads at frequencies just lower and higher than CF, respectively. Basilar membrane responses to low-level CF tones exceeded the magnitude of stapes vibrations by 54–56 dB. Response phases led stapes vibrations by about 90° at low stimulus frequencies; at higher frequencies, basilar membrane responses increasingly lagged stapes vibration, accumulating 1.5 periods of phase lag at CF. Postmortem, nonlinearities were abolished and responses peaked at ~0.5 octave below CF, with phases which lagged and led in vivo responses at frequencies lower and higher than CF, respectively. In conclusion, basilar membrane responses near the round window of the gerbil cochlea closely resemble those for other basal cochlear sites in gerbil and other species.
doi:10.1007/sl01620020023
PMCID: PMC1868570  PMID: 12382108
inner ear; auditory; basilar membrane; cochlear mechanics; compressive nonlinearity
21.  High-Frequency Sensitivity of the Mature Gerbil Cochlea and Its Development 
Audiology & neuro-otology  2003;8(1):19-27.
The thresholds of compound action potentials evoked by tone pips were measured in the cochleae of anesthetized gerbils, both in adults and in neonates aged 14, 16, 18, 20 and 30 days, using round-window electrodes. Stapes vibrations were also measured, using a laser velocimeter, in many of the same ears of adults and neonates aged 14, 16, 18 and 20 days to assess cochlear sensitivity in isolation from middle ear effects and to circumvent problems associated with calibration of acoustic stimuli at high frequencies. Whether referenced to sound pressure level in the ear canal or stapes vibration velocity, thresholds in adults were roughly uniform in the entire range of tested frequencies, 1.25–38.5 kHz. In neonates, thresholds decreased systematically as a function of age, with the largest reductions occurring at the highest frequencies. Thresholds remained slightly immature at all frequencies 30 days after birth. The results for adult gerbils are consistent with the recent finding that basilar-membrane responses to characteristic frequency tones normalized to stapes vibrations are as sensitive at sites near the round window as at more apical sites. The results for neonates confirm that the extreme basal region of the cochlea is the last to approach maturity, with substantial development occurring between 20 and 30 days after birth.
doi:10.1159/000067892
PMCID: PMC1868568  PMID: 12566689
Cochlea; Basilar membrane; Development; Middle ear; Stapes; Gerbil
22.  Middle-ear transmission in humans: wide-band, not frequency-tuned? 
Postmortem and in vivo vibration responses to sound of the stapes and the umbo of human ears are surveyed. The magnitudes of umbo velocity responses recorded postmortem decay between 1 and 5 or 10 kHz at rates between 0 and −3 dB/octave. In contrast, the magnitudes of in vivo umbo vibration are relatively invariant over a wide frequency range, amply exceeding the bandwidth of the audiogram according to one report. Similarly, most studies of postmortem stapes vibration report velocities tuned to about 1 kHz, with magnitudes that decay at a rate of about −6 dB/octave at higher frequencies. In contrast, in vivo stapes responses are apparently only mildly tuned. We conjecture that the bandwidth of stapes vibration velocity in humans will eventually be shown to exceed the bandwidth of the audiogram, in line with findings in other amniotic vertebrates.
doi:10.1121/1.1566924
PMCID: PMC1868690  PMID: 17505561
23.  Similarity of Traveling-Wave Delays in the Hearing Organs of Humans and Other Tetrapods 
Transduction of sound in mammalian ears is mediated by basilar-membrane waves exhibiting delays that increase systematically with distance from the cochlear base. Most contemporary accounts of such “traveling-wave” delays in humans have ignored postmortem basilar-membrane measurements in favor of indirect in vivo estimates derived from brainstem-evoked responses, compound action potentials, and otoacoustic emissions. Here, we show that those indirect delay estimates are either flawed or inadequately calibrated. In particular, we argue against assertions based on indirect estimates that basilar-membrane delays are much longer in humans than in experimental animals. We also estimate in vivo basilar-membrane delays in humans by correcting postmortem measurements in humans according to the effects of death on basilar-membrane vibrations in other mammalian species. The estimated in vivo basilar-membrane delays in humans are similar to delays in the hearing organs of other tetrapods, including those in which basilar membranes do not sustain traveling waves or that lack basilar membranes altogether.
doi:10.1007/s10162-007-0081-z
PMCID: PMC1868567  PMID: 17401604
cochlea; basilar membrane; auditory; otoacoustic emissions; compound action potentials; brainstem evoked responses
24.  Comparison of group delays of 2f1–f2 distortion product otoacoustic emissions and cochlear travel times 
To help elucidate how distortion-product otoacoustic emissions propagate from their cochlear sites of origin to the middle ear, their group delays were compared with basilar-membrane and organ of Corti travel times measured in guinea pig, gerbil, and chinchilla.
doi:10.1121/1.1771711
PMCID: PMC2196440  PMID: 18196179
25.  Similarity of Traveling-Wave Delays in the Hearing Organs of Humans and Other Tetrapods 
Transduction of sound in mammalian ears is mediated by basilar-membrane waves exhibiting delays that increase systematically with distance from the cochlear base. Most contemporary accounts of such “traveling-wave” delays in humans have ignored postmortem basilar-membrane measurements in favor of indirect in vivo estimates derived from brainstem-evoked responses, compound action potentials, and otoacoustic emissions. Here, we show that those indirect delay estimates are either flawed or inadequately calibrated. In particular, we argue against assertions based on indirect estimates that basilar-membrane delays are much longer in humans than in experimental animals. We also estimate in vivo basilar-membrane delays in humans by correcting postmortem measurements in humans according to the effects of death on basilar-membrane vibrations in other mammalian species. The estimated in vivo basilar-membrane delays in humans are similar to delays in the hearing organs of other tetrapods, including those in which basilar membranes do not sustain traveling waves or that lack basilar membranes altogether.
doi:10.1007/s10162-007-0081-z
PMCID: PMC1868567  PMID: 17401604
cochlea; basilar membrane; auditory; otoacoustic emissions; compound action potentials; brainstem evoked responses

Results 1-25 (29)