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1.  Response Characteristics in the Apex of the Gerbil Cochlea Studied Through Auditory Nerve Recordings 
In this study, we analyze the processing of low-frequency sounds in the cochlear apex through responses of auditory nerve fibers (ANFs) that innervate the apex. Single tones and irregularly spaced tone complexes were used to evoke ANF responses in Mongolian gerbil. The spike arrival times were analyzed in terms of phase locking, peripheral frequency selectivity, group delays, and the nonlinear effects of sound pressure level (SPL). Phase locking to single tones was similar to that in cat. Vector strength was maximal for stimulus frequencies around 500 Hz, decreased above 1 kHz, and became insignificant above 4 to 5 kHz. We used the responses to tone complexes to determine amplitude and phase curves of ANFs having a characteristic frequency (CF) below 5 kHz. With increasing CF, amplitude curves gradually changed from broadly tuned and asymmetric with a steep low-frequency flank to more sharply tuned and asymmetric with a steep high-frequency flank. Over the same CF range, phase curves gradually changed from a concave-upward shape to a concave-downward shape. Phase curves consisted of two or three approximately straight segments. Group delay was analyzed separately for these segments. Generally, the largest group delay was observed near CF. With increasing SPL, most amplitude curves broadened, sometimes accompanied by a downward shift of best frequency, and group delay changed along the entire range of stimulus frequencies. We observed considerable across-ANF variation in the effects of SPL on both amplitude and phase. Overall, our data suggest that mechanical responses in the apex of the cochlea are considerably nonlinear and that these nonlinearities are of a different character than those known from the base of the cochlea.
doi:10.1007/s10162-010-0255-y
PMCID: PMC3085685  PMID: 21213012
cochlear mechanics; cochlear apex; phase locking; Meriones unguiculatus
2.  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
3.  The Group Delay and Suppression Pattern of the Cochlear Microphonic Potential Recorded at the Round Window 
PLoS ONE  2012;7(3):e34356.
Background
It is commonly assumed that the cochlear microphonic potential (CM) recorded from the round window (RW) is generated at the cochlear base. Based on this assumption, the low-frequency RW CM has been measured for evaluating the integrity of mechanoelectrical transduction of outer hair cells at the cochlear base and for studying sound propagation inside the cochlea. However, the group delay and the origin of the low-frequency RW CM have not been demonstrated experimentally.
Methodology/Principal Findings
This study quantified the intra-cochlear group delay of the RW CM by measuring RW CM and vibrations at the stapes and basilar membrane in gerbils. At low sound levels, the RW CM showed a significant group delay and a nonlinear growth at frequencies below 2 kHz. However, at high sound levels or at frequencies above 2 kHz, the RW CM magnitude increased proportionally with sound pressure, and the CM phase in respect to the stapes showed no significant group delay. After the local application of tetrodotoxin the RW CM below 2 kHz became linear and showed a negligible group delay. In contrast to RW CM phase, the BM vibration measured at location ∼2.5 mm from the base showed high sensitivity, sharp tuning, and nonlinearity with a frequency-dependent group delay. At low or intermediate sound levels, low-frequency RW CMs were suppressed by an additional tone near the probe-tone frequency while, at high sound levels, they were partially suppressed only at high frequencies.
Conclusions/Significance
We conclude that the group delay of the RW CM provides no temporal information on the wave propagation inside the cochlea, and that significant group delay of low-frequency CMs results from the auditory nerve neurophonic potential. Suppression data demonstrate that the generation site of the low-frequency RW CM shifts from apex to base as the probe-tone level increases.
doi:10.1371/journal.pone.0034356
PMCID: PMC3314608  PMID: 22470560
4.  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
5.  Tuning of SFOAEs Evoked by Low-Frequency Tones Is Not Compatible with Localized Emission Generation 
Stimulus-frequency otoacoustic emissions (SFOAEs) appear to be well suited for assessing frequency selectivity because, at least on theoretical grounds, they originate over a restricted region of the cochlea near the characteristic place of the evoking tone. In support of this view, we previously found good agreement between SFOAE suppression tuning curves (SF-STCs) and a control measure of frequency selectivity (compound action potential suppression tuning curves (CAP-STC)) for frequencies above 3 kHz in chinchillas. For lower frequencies, however, SF-STCs and were over five times broader than the CAP-STCs and demonstrated more high-pass rather than narrow band-pass filter characteristics. Here, we test the hypothesis that the broad tuning of low-frequency SF-STCs is because emissions originate over a broad region of the cochlea extending basal to the characteristic place of the evoking tone. We removed contributions of the hypothesized basally located SFOAE sources by either pre-suppressing them with a high-frequency interference tone (IT; 4.2, 6.2, or 9.2 kHz at 75 dB sound pressure level (SPL)) or by inducing acoustic trauma at high frequencies (exposures to 8, 5, and lastly 3-kHz tones at 110–115 dB SPL). The 1-kHz SF-STCs and CAP-STCs were measured for baseline, IT present and following the acoustic trauma conditions in anesthetized chinchillas. The IT and acoustic trauma affected SF-STCs in an almost indistinguishable way. The SF-STCs changed progressively from a broad high-pass to narrow band-pass shape as the frequency of the IT was lowered and for subsequent exposures to lower-frequency tones. Both results were in agreement with the “basal sources” hypothesis. In contrast, CAP-STCs were not changed by either manipulation, indicating that neither the IT nor acoustic trauma affected the 1-kHz characteristic place. Thus, unlike CAPs, SFOAEs cannot be considered as a place-specific measure of cochlear function at low frequencies, at least in chinchillas.
doi:10.1007/s10162-015-0513-0
PMCID: PMC4417092  PMID: 25813430
acoustic trauma; two-tone suppression; compound action potential; suppression tuning curve; chinchilla
6.  Wiener Kernels of Chinchilla Auditory-Nerve Fibers: Verification Using Responses to Tones, Clicks, and Noise and Comparison With Basilar-Membrane Vibrations 
Journal of neurophysiology  2005;93(6):3635-3648.
Responses to tones, clicks, and noise were recorded from chinchilla auditory-nerve fibers (ANFs). The responses to noise were analyzed by computing the zeroth-, first-, and second-order Wiener kernels (h0, h1, and h2). The h1s correctly predicted the frequency tuning and phases of responses to tones of ANFs with low characteristic frequency (CF). The h2s correctly predicted the frequency tuning and phases of responses to tones of all ANFs, regardless of CF. Also regardless of CF, the kernels jointly predicted about 77% of the features of ANF responses to “frozen” samples of noise. Near-CF group delays of kernels and signal-front delays of responses to intense rarefaction clicks exceeded by 1 ms the corresponding basilar-membrane delays at both apical and basal sites of the chinchilla cochlea. This result, confirming that synaptic and neural processes amount to 1 ms regardless of CF, permitted drawing a map of basilar-membrane delay as a function of position for the entire length of the chinchilla cochlea, a first for amniotic species.
doi:10.1152/jn.00885.2004
PMCID: PMC1876724  PMID: 15659530
7.  Auditory brainstem responses predict auditory nerve fiber thresholds and frequency selectivity in hearing impaired chinchillas 
Hearing research  2011;280(1-2):236-244.
Non-invasive auditory brainstem responses (ABRs) are commonly used to assess cochlear pathology in both clinical and research environments. In the current study, we evaluated the relationship between ABR characteristics and more direct measures of cochlear function. We recorded ABRs and auditory nerve (AN) single-unit responses in seven chinchillas with noise induced hearing loss. ABRs were recorded for 1–8 kHz tone burst stimuli both before and several weeks after four hours of exposure to a 115 dB SPL, 50 Hz band of noise with a center frequency of 2 kHz. Shifts in ABR characteristics (threshold, wave I amplitude, and wave I latency) following hearing loss were compared to AN-fiber tuning curve properties (threshold and frequency selectivity) in the same animals. As expected, noise exposure generally resulted in an increase in ABR threshold and decrease in wave I amplitude at equal SPL. Wave I amplitude at equal sensation level (SL), however, was similar before and after noise exposure. In addition, noise exposure resulted in decreases in ABR wave I latency at equal SL and, to a lesser extent, at equal SPL. The shifts in ABR characteristics were significantly related to AN-fiber tuning curve properties in the same animal at the same frequency. Larger shifts in ABR thresholds and ABR wave I amplitude at equal SPL were associated with greater AN threshold elevation. Larger reductions in ABR wave I latency at equal SL, on the other hand, were associated with greater loss of AN frequency selectivity. This result is consistent with linear systems theory, which predicts shorter time delays for broader peripheral frequency tuning. Taken together with other studies, our results affirm that ABR thresholds and wave I amplitude provide useful estimates of cochlear sensitivity. Furthermore, comparisons of ABR wave I latency to normative data at the same SL may prove useful for detecting and characterizing loss of cochlear frequency selectivity.
doi:10.1016/j.heares.2011.06.002
PMCID: PMC3179834  PMID: 21699970
Auditory brainstem response (ABR); auditory nerve; auditory threshold; frequency selectivity; hearing loss
8.  Maintaining acoustic communication at a cocktail party: heterospecific masking noise improves signal detection through frequency separation 
The Journal of experimental biology  2013;216(0 24):4655-4665.
SUMMARY
We examined acoustic masking in a chirping katydid species of the Mecopoda elongata complex due to interference with a sympatric Mecopoda species where males produce continuous trills at high amplitudes. Frequency spectra of both calling songs range from 1 to 80 kHz; the chirper species has more energy in a narrow frequency band at 2 kHz and above 40 kHz. Behaviourally, chirper males successfully phase-locked their chirps to playbacks of conspecific chirps under masking conditions at signal-to-noise ratios (SNRs) of −8 dB. After the 2 kHz band in the chirp had been equalised to the level in the masking trill, the breakdown of phase-locked synchrony occurred at a SNR of +7 dB. The remarkable receiver performance is partially mirrored in the selective response of a first-order auditory interneuron (TN1) to conspecific chirps under these masking conditions. However, the selective response is only maintained for a stimulus including the 2 kHz component, although this frequency band has no influence on the unmasked TN1 response. Remarkably, the addition of masking noise at 65 dB sound pressure level (SPL) to threshold response levels of TN1 for pure tones of 2 kHz enhanced the sensitivity of the response by 10 dB. Thus, the spectral dissimilarity between masker and signal at a rather low frequency appears to be of crucial importance for the ability of the chirping species to communicate under strong masking by the trilling species. We discuss the possible properties underlying the cellular/synaptic mechanisms of the ‘novelty detector’.
doi:10.1242/jeb.089888
PMCID: PMC3971153  PMID: 24307713
selective encoding; insect; auditory interneuron; katydid; ambient noise; novelty detection
9.  The Auditory Nerve Overlapped Waveform (ANOW) Originates in the Cochlear Apex 
Measurements of cochlear function with compound action potentials (CAPs), auditory brainstem responses, and otoacoustic emissions work well with high-frequency sounds but are problematic at low frequencies. We have recently shown that the auditory nerve overlapped waveform (ANOW) can objectively quantify low-frequency (<1 kHz) auditory sensitivity, as thresholds for ANOW at low frequencies and for CAP at high frequencies relate similarly to single auditory nerve fiber thresholds. This favorable relationship, however, does not necessarily mean that ANOW originates from auditory nerve fibers innervating low-frequency regions of the cochlear apex. In the present study, we recorded the cochlear response to tone bursts of low frequency (353, 500, and 707 Hz) and high frequency (2 to 16 kHz) during administration of tetrodotoxin (TTX) to block neural function. TTX was injected using a novel method of slow administration from a pipette sealed into the cochlear apex, allowing real-time measurements of systematic neural blocking from apex to base. The amplitude of phase-locked (ANOW) and onset (CAP) neural firing to moderate-level, low-frequency sounds were markedly suppressed before thresholds and responses to moderate-level, high-frequency sounds were affected. These results demonstrate that the ANOW originates from responses of auditory nerve fibers innervating cochlear apex, confirming that ANOW provides a valid physiological measure of low-frequency auditory nerve function.
doi:10.1007/s10162-014-0447-y
PMCID: PMC4010591  PMID: 24515339
auditory nerve neurophonic; compound action potential; low-frequency hearing; neural synchrony; phase locking
10.  A new auditory threshold estimation technique for low frequencies: Proof of concept 
Ear and hearing  2013;34(1):42-51.
Objectives
Presently available non-behavioral methods to estimate auditory thresholds perform less well at frequencies below 1 kHz than at 1 kHz and above. For many uses, such as providing accurate infant hearing aid amplification for low-frequency vowels, we need an accurate non-behavioral method to estimate low-frequency thresholds. Here we develop a novel technique to estimate low-frequency cochlear thresholds based on the use of a previously-reported waveform. We determine how well the method works by comparing the resulting thresholds to thresholds from onset-response compound action potentials (CAPs) and single auditory-nerve (AN) fibers in cats. A long-term goal is to translate this technique for use in humans.
Design
An electrode near the cochlea records a combination of cochlear microphonic (CM) and neural responses. In response to low-frequency, near threshold-level tones, the CM is almost sinusoidal while the neural responses occur preferentially at one phase of the tone. If the tone is presented again but with its polarity reversed, the neural response keeps the same shape, but shifts ½ cycle in time. Averaging responses to tones presented separately at opposite polarities overlaps and interleaves the neural responses and yields a waveform in which the CM is cancelled and the neural response appears twice each tone cycle, i.e. the resulting neural response is mostly at twice the tone frequency. We call the resultant waveform “the auditory nerve overlapped waveform” (ANOW). ANOW level functions were measured in anesthetized cats from 10 to 80 dB SPL in 10 dB steps using tones between 0.3 and 1 kHz. As a response metric, we calculated the magnitude of the ANOW component at twice the tone frequency (ANOW2f). The ANOW threshold was the sound level where the interpolated ANOW2f crossed a statistical criterion that was higher than 95% of the noise floor distribution. ANOW thresholds were compared to onset-CAP thresholds from the same recordings and single-AN-fiber thresholds from the same animals.
Results
We obtained ANOW and onset-CAP level functions for 0.3 to 1 kHz tones, and single-AN-fiber responses from cats. Except at 1 kHz, typical ANOW thresholds were mostly 10-20 dB more sensitive than onset-CAP thresholds and 10-20 dB less sensitive than the most sensitive single-AN-fiber thresholds.
Conclusions
ANOW provides frequency-specific estimates of cochlear neural thresholds over a frequency range that is important for hearing but is not well accessed by non-behavioral, non-invasive methods. Our results suggest that, with further targeted development, the ANOW low-frequency threshold estimation technique can be useful both clinically in humans and in basic-science animal experiments.
doi:10.1097/AUD.0b013e31825f9bd3
PMCID: PMC3495092  PMID: 22874644
audiogram; auditory nerve neurophonic; compound action potential; neural synchrony; phase locking
11.  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
12.  Membrane properties specialize mammalian inner hair cells for frequency or intensity encoding 
eLife  null;4:e08177.
The auditory pathway faithfully encodes and relays auditory information to the brain with remarkable speed and precision. The inner hair cells (IHCs) are the primary sensory receptors adapted for rapid auditory signaling, but they are not thought to be intrinsically tuned to encode particular sound frequencies. Here I found that under experimental conditions mimicking those in vivo, mammalian IHCs are intrinsically specialized. Low-frequency gerbil IHCs (~0.3 kHz) have significantly more depolarized resting membrane potentials, faster kinetics, and shorter membrane time constants than high-frequency cells (~30 kHz). The faster kinetics of low-frequency IHCs allow them to follow the phasic component of sound (frequency-following), which is not required for high-frequency cells that are instead optimally configured to encode sustained, graded responses (intensity-following). The intrinsic membrane filtering of IHCs ensures accurate encoding of the phasic or sustained components of the cell’s in vivo receptor potential, crucial for sound localization and ultimately survival.
DOI: http://dx.doi.org/10.7554/eLife.08177.001
eLife digest
Many animals’ survival depends on them accurately and quickly identifying sounds in their environment. In animals with backbones, cells with hair-like projections (called hair cells) inside the ear convert information collected from sound waves into electrical signals. These signals are then transmitted to the brain, which processes the information further.
Animals like bullfrogs are adapted to hearing low frequency sounds, like their own mating calls. These frog’s hair cells are individually tuned so that they can capture sounds in this low frequency range. Mammals, on the other hand, have evolved to hear a much wider range of sounds from loud and low frequency sounds, such as thunder, to soft and high frequency sounds, like the cries of their young. In mammals, the part of inner ear involved in hearing (called the cochlea) has an elaborate spiral-like shape. The structure of the cochlea results in different frequencies of sound being transformed by the hair cells into electrical signals at different points around the spiral. Because of this, most researchers didn’t think that hair cells in mammals were individually tuned like those in bullfrogs.
Now, Stuart Johnson demonstrates that hair cells in different parts of the gerbil’s cochlea are specialized for encoding sounds of specific frequencies. In conditions that mimic the environment inside the ear, a very precise jet of fluid was used to stimulate single hair cells in a similar way to a sound wave. The experiments then compared how hair cells from the upper and lower parts of the cochlea’s spiral responded. Johnson found that hair cells from the upper portion of the gerbils’ cochlea are specialized to capture low frequency sounds. They have electrical properties that allow them to quickly transmit information to the brain about low frequency sounds. In the lower portion of the cochlea, hair cells are specialized to capture high frequency sounds. That is, their electrical properties make it easier for these hair cells to transmit detailed information to the brain about the volume of high frequency sounds. Together, these findings help explain how these animals are able to localize sounds, which requires capturing both the timing and intensity of different types of sounds.
DOI: http://dx.doi.org/10.7554/eLife.08177.002
doi:10.7554/eLife.08177
PMCID: PMC4709266  PMID: 26544545
gerbil; cochlea; hair cell; Other
13.  Development of wide-band middle ear transmission in the Mongolian gerbila) 
Stapes vibrations were measured in deeply anesthetized adult and neonatal (ages: 14 to 20 days) Mongolian gerbils. In adult gerbils, the velocity magnitude of stapes responses to tones was approximately constant over the entire frequency range of measurements, 1 to 40 kHz. Response phases referred to pressure near the tympanic membrane varied approximately linearly as a function of increasing stimulus frequency, with a slope corresponding to a group delay of 30 μs. In neonatal gerbils, the sensitivity of stapes responses to tones was lower than in adults, especially at mid-frequencies (e.g., by about 15 dB at 10–20 kHz in gerbils aged 14 days). The input impedance of the adult gerbil cochlea, calculated from stapes vibrations and published measurements of pressure in scala vestibuli near the oval window [E. Olson, J. Acoust. Soc. Am. 103, 3445–3463 (1998)], is principally dissipative at frequencies lower than 10 kHz.
Conclusions
(a) middle-ear vibrations in adult gerbils do not limit the input to the cochlea up to at least 40 kHz, i.e., within 0.5 oct of the high-frequency cutoff of the behavioral audiogram; and (b) the results in both adult and neonatal gerbils are inconsistent with the hypothesis that mass reactance controls high-frequency ossicular vibrations and support the idea that the middle ear functions as a transmission line.
doi:10.1121/1.1420382
PMCID: PMC1868569  PMID: 11831800
14.  Distortion Product Otoacoustic Emission and Auditory Brainstem Responses in the Echidna (Tachyglossus aculeatus)  
The auditory function of four wild-caught echidnas was measured using distortion product otoacoustic emissions (DPOAEs) and auditory brainstem responses (ABRs). Emission audiograms were constructed by finding the stimulus levels required to produce a criterion emission amplitude at a given stimulus frequency. For an emission amplitude of -10 dB SPL, the median "best threshold" was 28 dB SPL, and this minimum threshold occurred between 4 and 8 kHz for all animals. The relative effective range of auditory function was defined by the frequencies at which the audiogram was 30 dB above its best threshold. For the emission audiograms, the median lower-frequency limit was 2.3 kHz, the upper limit was 18.4 kHz, and the effective range was 2.7 octaves. The audiogram as measured by ABR was also found to be strongly "U" shaped with similar low- and high-frequency limits, i.e., from 1.6 to 13.9 kHz, with an effective range of 3.1 octaves. These results suggest that the echidna has a behavioral hearing sensitivity comparable to that of typical therian mammals (e.g., rabbits and gerbils) but with a significantly narrower frequency range. DPOAE responses were also measured in selected animals as a function of the variation of all four stimulus parameters (frequencies and intensities of both stimulus tones). Overall, the measured emission responses establish that the echidna does have a cochlear amplifier, and that it could be the same type as in therian mammals. The amplification mechanism in the echidna, currently unidentified, clearly operates to frequencies above 20 kHz, higher than the hearing function observed in any birds or reptiles but lower than for typical therian mammals. This raises the possibility that at least some aspects of the mammalian cochlear amplifier developed early in evolution, before the divergence of the monotremes (echidna and platypus) from the mainstream therian mammals (marsupials and placentals). In this respect, the presence or absence of outer hair cell electromotility in monotremes would have important consequences for understanding the function and evolution of the vertebrate inner ear.
doi:10.1007/s101620010059
PMCID: PMC3201180  PMID: 11550523
15.  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
16.  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/s101620020023
PMCID: PMC1868570  PMID: 12382108
17.  Estimating Cochlear Frequency Selectivity with Stimulus-frequency Otoacoustic Emissions in Chinchillas 
It has been suggested that the tuning of the cochlear filters can be derived from measures of otoacoustic emissions (OAEs). Two approaches have been proposed to estimate cochlear frequency selectivity using OAEs evoked with a single tone (stimulus-frequency (SF)) OAEs: based on SFOAE group delays (SF-GDs) and on SFOAE suppression tuning curves (SF-STCs). The aim of this study was to evaluate whether either SF-GDs or SF-STCs obtained with low probe levels (30 dB SPL) correlate with more direct measures of cochlear tuning (compound action potential suppression tuning curves (CAP-STCs)) in chinchillas. The SFOAE-based estimates of tuning covaried with CAP-STCs tuning for >3 kHz probe frequencies, indicating that these measures are related to cochlear frequency selectivity. However, the relationship may be too weak to predict tuning with either SFOAE method in an individual. The SF-GD prediction of tuning was sharper than CAP-STC tuning. On the other hand, SF-STCs were consistently broader than CAP-STCs implying that SFOAEs may have less restricted region of generation in the cochlea than CAPs. Inclusion of <3 kHz data in a statistical model resulted in no significant or borderline significant covariation among the three methods: neither SFOAE test appears to reliably estimate an individual’s CAP-STC tuning at low-frequencies. At the group level, SF-GDs and CAP-STCs showed similar tuning at low frequencies, while SF-STCs were over five times broader than the CAP-STCs indicating that low-frequency SFOAE may originate over a very broad region of the cochlea extending ≥5 mm basal to the tonotopic place of the probe.
doi:10.1007/s10162-014-0487-3
PMCID: PMC4389964  PMID: 25230801
frequency selectivity; SFOAE; CAP; tuning curve
18.  Enhancement and Distortion in the Temporal Representation of Sounds in the Ventral Cochlear Nucleus of Chinchillas and Cats 
PLoS ONE  2012;7(9):e44286.
A subset of neurons in the cochlear nucleus (CN) of the auditory brainstem has the ability to enhance the auditory nerve's temporal representation of stimulating sounds. These neurons reside in the ventral region of the CN (VCN) and are usually known as highly synchronized, or high-sync, neurons. Most published reports about the existence and properties of high-sync neurons are based on recordings performed on a VCN output tract—not the VCN itself—of cats. In other species, comprehensive studies detailing the properties of high-sync neurons, or even acknowledging their existence, are missing.
Examination of the responses of a population of VCN neurons in chinchillas revealed that a subset of those neurons have temporal properties similar to high-sync neurons in the cat. Phase locking and entrainment—the ability of a neuron to fire action potentials at a certain stimulus phase and at almost every stimulus period, respectively—have similar maximum values in cats and chinchillas. Ranges of characteristic frequencies for high-sync neurons in chinchillas and cats extend up to 600 and 1000 Hz, respectively. Enhancement of temporal processing relative to auditory nerve fibers (ANFs), which has been shown previously in cats using tonal and white-noise stimuli, is also demonstrated here in the responses of VCN neurons to synthetic and spoken vowel sounds.
Along with the large amount of phase locking displayed by some VCN neurons there occurs a deterioration in the spectral representation of the stimuli (tones or vowels). High-sync neurons exhibit a greater distortion in their responses to tones or vowels than do other types of VCN neurons and auditory nerve fibers.
Standard deviations of first-spike latency measured in responses of high-sync neurons are lower than similar values measured in ANFs' responses. This might indicate a role of high-sync neurons in other tasks beyond sound localization.
doi:10.1371/journal.pone.0044286
PMCID: PMC3445608  PMID: 23028514
19.  Dynamics of Infant Cortical Auditory Evoked Potentials (CAEPs) for Tone and Speech Tokens 
Objectives
Cortical auditory evoked potentials (CAEPs) to tones and speech sounds were obtained in infants to: 1) further knowledge of auditory development above the level of the brainstem during the first year of life; 2) establish CAEP input-output functions for tonal and speech stimuli as a function of stimulus level and to 3) elaborate the data-base that establishes CAEP in infants tested while awake using clinically relevant stimuli, thus providing methodology that would have translation to pediatric audiological assessment. Hypotheses concerning CAEP development were that the latency and amplitude input-output functions would reflect immaturity in encoding stimulus level.
In a second experiment, infants were tested with the same stimuli used to evoke the CAEPs. Thresholds for these stimuli were determined using observer-based psychophysical techniques. The hypothesis was that the behavioral thresholds would be correlated with CAEP input-output functions because of shared cortical response areas known to be active in sound detection.
Design
36 infants, between the ages of 4-12 months (mean= 8 months, s.d.=1.8 months) and 9 young adults (mean age 21 years) with normal hearing were tested. First, CAEPs amplitude and latency input-output functions were obtained for 4 tone bursts and 7 speech tokens. The tone bursts stimuli were 50 ms tokens of pure tones at 0.5, 1.0, 2.0 and 4.0 kHz. The speech sound tokens, /a/, /i/, /o/, /u/, /m/, /s/, and /∫/, were created from natural speech samples and were also 50 ms in duration. CAEPs were obtained for tone burst and speech token stimuli at 10 dB level decrements in descending order from 70 dB SPL. All CAEP tests were completed while the infants were awake and engaged in quiet play.
For the second experiment, observer-based psychophysical methods were used to establish perceptual threshold for the same speech sound and tone tokens.
Results
Infant CAEP component latencies were prolonged by 100-150 ms in comparison to adults. CAEP latency-intensity input output functions were steeper in infants compared to adults. CAEP amplitude growth functions with respect to stimulus SPL are adult-like at this age, particularly for the earliest component, P1-N1.
Infant perceptual thresholds were elevated with respect to those found in adults. Furthermore, perceptual thresholds were higher, on average, than levels at which CAEPs could be obtained. When CAEP amplitudes were plotted with respect to perceptual threshold (dB SL), the infant CAEP amplitude growth slopes were steeper than in adults.
Conclusions
Although CAEP latencies indicate immaturity in neural transmission at the level of the cortex, amplitude growth with respect to stimulus SPL is adult-like at this age, particularly for the earliest component, P1-N1. The latency and amplitude input-output functions may provide additional information as to how infants perceive stimulus level. The reasons for the discrepancy between electrophysiologic and perceptual threshold may be due to immaturity in perceptual temporal resolution abilities and the broad-band listening strategy employed by infants.
The findings from the current study can be translated to the clinical setting. It is possible to use tonal or speech sound tokens to evoke CAEPs in an awake, passively alert infant, and thus determine whether these sounds activate the auditory cortex. This could be beneficial in the verification of hearing aid or cochlear implant benefit.
doi:10.1016/j.ijporl.2013.04.030
PMCID: PMC3700622  PMID: 23722003
infant; evoked potential; perception
20.  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
21.  Phase Locking of Auditory-Nerve Fibers to the Envelopes of High-Frequency Sounds: Implications for Sound Localization 
Journal of neurophysiology  2006;96(5):2327-2341.
Although listeners are sensitive to interaural time differences (ITDs) in the envelope of high-frequency sounds, both ITD discrimination performance and the extent of lateralization are poorer for high-frequency sinusoidally amplitude-modulated (SAM) tones than for low-frequency pure tones. Psychophysical studies have shown that ITD discrimination at high frequencies can be improved by using novel transposed-tone stimuli, formed by modulating a high-frequency carrier by a half-wave–rectified sinusoid. Transposed tones are designed to produce the same temporal discharge patterns in high-characteristic frequency (CF) neurons as occur in low-CF neurons for pure-tone stimuli. To directly test this hypothesis, we compared responses of auditory-nerve fibers in anesthetized cats to pure tones, SAM tones, and transposed tones. Phase locking was characterized using both the synchronization index and autocorrelograms. With both measures, phase locking was better for transposed tones than for SAM tones, consistent with the rationale for using transposed tones. However, phase locking to transposed tones and that to pure tones were comparable only when all three conditions were met: stimulus levels near thresholds, low modulation frequencies (<250 Hz), and low spontaneous discharge rates. In particular, phase locking to both SAM tones and transposed tones substantially degraded with increasing stimulus level, while remaining more stable for pure tones. These results suggest caution in assuming a close similarity between temporal patterns of peripheral activity produced by transposed tones and pure tones in both psychophysical studies and neurophysiological studies of central neurons.
doi:10.1152/jn.00326.2006
PMCID: PMC2013745  PMID: 16807349
22.  Age-Related Hearing Loss in C57BL/6J Mice has both Frequency-Specific and Non-Frequency-Specific Components that Produce a Hyperacusis-Like Exaggeration of the Acoustic Startle Reflex 
Auditory brainstem-evoked response (ABR) thresholds were obtained in a longitudinal study of C57BL/6J mice between 10 and 53 weeks old, with repeated testing every 2 weeks. On alternate weeks, acoustic startle reflex (ASR) amplitudes were measured, elicited by tone pips with stimulus frequencies of 3, 6, 12, and 24 kHz, and intensities from subthreshold up to 110 dB sound pressure level. The increase in ABR thresholds for 3 and 6 kHz test stimuli followed a linear time course with increasing age from 10 to 53 weeks, with a slope of about 0.7 dB/week, and for 48 kHz a second linear time course, but beginning at 10 weeks with a slope of about 2.3 dB/week. ABR thresholds for 12, 24, and 32 kHz increased after one linear segment with a 0.7 dB slope, then after a variable delay related to the test frequency, shifted to a second segment having slopes of 3–5 dB/week. Hearing loss initially reduced the ASR for all eliciting stimuli, but at about 6 months of age, the response elicited by intense 3 and 6 kHz stimuli began to increase to reach values about three times above normal, and previously subthreshold stimuli came to elicit vigorous responses seen at first only for the intense stimuli. This hyperacusis-like effect appeared in all mice but was especially pronounced in mice with more serious hearing loss. These ABR data, together with a review of histopathological data in the C57BL/6 literature, suggest that the non-frequency-specific slow time course of hearing loss results from pathology in the lateral wall of the cochlea, whereas the stimulus-specific hearing loss with a rapid time course results from hair cell loss. Delayed exaggeration of the ASR with hearing loss reveals a deficit in centrifugal inhibitory control over the afferent reflex pathways after central neural reorganization, suggesting that this mouse may provide a useful model of age-related tinnitus and associated hyperacusis.
doi:10.1007/s10162-007-0098-3
PMCID: PMC2538342  PMID: 17952509
aging; hearing loss; startle; plasticity; mixed strial/sensory presbycusis; tinnitus/hyperacusis
23.  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
24.  Immediate and Delayed Cochlear Neuropathy after Noise Exposure in Pubescent Mice 
PLoS ONE  2015;10(5):e0125160.
Moderate acoustic overexposure in adult rodents is known to cause acute loss of synapses on sensory inner hair cells (IHCs) and delayed degeneration of the auditory nerve, despite the completely reversible temporary threshold shift (TTS) and morphologically intact hair cells. Our objective was to determine whether a cochlear synaptopathy followed by neuropathy occurs after noise exposure in pubescence, and to define neuropathic versus non-neuropathic noise levels for pubescent mice. While exposing 6 week old CBA/CaJ mice to 8-16 kHz bandpass noise for 2 hrs, we defined 97 dB sound pressure level (SPL) as the threshold for this particular type of neuropathic exposure associated with TTS, and 94 dB SPL as the highest non-neuropathic noise level associated with TTS. Exposure to 100 dB SPL caused permanent threshold shift although exposure of 16 week old mice to the same noise is reported to cause only TTS. Amplitude of wave I of the auditory brainstem response, which reflects the summed activity of the cochlear nerve, was complemented by synaptic ribbon counts in IHCs using confocal microscopy, and by stereological counts of peripheral axons and cell bodies of the cochlear nerve from 24 hours to 16 months post exposure. Mice exposed to neuropathic noise demonstrated immediate cochlear synaptopathy by 24 hours post exposure, and delayed neurodegeneration characterized by axonal retraction at 8 months, and spiral ganglion cell loss at 8-16 months post exposure. Although the damage was initially limited to the cochlear base, it progressed to also involve the cochlear apex by 8 months post exposure. Our data demonstrate a fine line between neuropathic and non-neuropathic noise levels associated with TTS in the pubescent cochlea.
doi:10.1371/journal.pone.0125160
PMCID: PMC4425526  PMID: 25955832
25.  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

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