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1.  Mechanical Overstimulation of Hair Bundles: Suppression and Recovery of Active Motility 
PLoS ONE  2013;8(3):e58143.
We explore the effects of high-amplitude mechanical stimuli on hair bundles of the bullfrog sacculus. Under in vitro conditions, these bundles exhibit spontaneous limit cycle oscillations. Prolonged deflection exerted two effects. First, it induced an offset in the position of the bundle. Recovery to the original position displayed two distinct time scales, suggesting the existence of two adaptive mechanisms. Second, the stimulus suppressed spontaneous oscillations, indicating a change in the hair bundle’s dynamic state. After cessation of the stimulus, active bundle motility recovered with time. Both effects were dependent on the duration of the imposed stimulus. External calcium concentration also affected the recovery to the oscillatory state. Our results indicate that both offset in the bundle position and calcium concentration control the dynamic state of the bundle.
doi:10.1371/journal.pone.0058143
PMCID: PMC3591416  PMID: 23505461
2.  Mechanics of the frog ear 
Hearing research  2010;273(1-2):46-58.
The frog inner ear contains three regions that are sensitive to airborne sound and which are functionally distinct. (1) The responses of nerve fibres innervating the low-frequency, rostral part of the amphibian papilla (AP) are complex. Electrical tuning of hair cells presumably contributes to the frequency selectivity of these responses. (2) The caudal part of the AP covers the mid-frequency portion of the frog's auditory range. It shares the ability to generate both evoked and spontaneous otoacoustic emissions with the mammalian cochlea and other vertebrate ears. (3) The basilar papilla functions mainly as a single auditory filter. Its simple anatomy and function provide a model system for testing hypotheses concerning emission generation. Group delays of stimulus frequency otoacoustic emissions (SFOAEs) from the basilar papilla are accounted for by assuming that they result from forward and reverse transmission through the middle ear, a mechanical delay due to tectorial membrane filtering and a rapid forward and reverse propagation through the inner ear fluids, with negligible delay.
doi:10.1016/j.heares.2010.02.004
PMCID: PMC3023005  PMID: 20149854
Frog; amphibian papilla; basilar papilla; frequency selectivity; tectorial membrane; otoacoustic emission
3.  Frequency matching of vocalizations to inner-ear sensitivity along an altitudinal gradient in the coqui frog 
Biology Letters  2009;6(2):278-281.
Acoustic communication involves both the generation and the detection of a signal. In the coqui frog (Eleutherodactylus coqui), it is known that the spectral contents of its calls systematically change with altitude above sea level. Here, distortion product otoacoustic emissions are used to assess the frequency range over which the inner ear is sensitive. It is found that both the spectral contents of the calls and the inner-ear sensitivity change in a similar fashion along an altitudinal gradient. As a result, the call frequencies and the auditory tuning are closely matched at all altitudes. We suggest that the animal's body size determines the frequency particulars of the call apparatus and the inner ear.
doi:10.1098/rsbl.2009.0763
PMCID: PMC2865046  PMID: 19939848
coqui frog; Eleutherodactylus; otoacoustic emission; vocalization
4.  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
5.  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
6.  Distortion Product Otoacoustic Emissions Evoked by Tone Complexes 
Distortion product otoacoustic emissions (DPOAEs) are traditionally evoked by two-tone stimuli. In this study, emission data from Mongolian gerbils are reported that were obtained with stimuli consisting of six to 10 tones. The stimuli were constructed by replacing one of the tones of a tone pair by a narrowband multitone complex. This produced rich spectra of the ear canal sound pressure in which many of the third-order DPOAEs originated from the interaction of triplets of stimulus components. A careful choice of the stimulus frequencies ensured that none of these DPOAE components coincided. Three groups of DPOAEs are reported, two of which are closely related to DPOAEs evoked by tone pairs. The third group has no two-tone equivalent and only arises when using a multitone stimulus. We analyzed the relation between multitone-evoked DPOAEs and DPOAEs evoked by tone pairs, and explored the new degrees of freedom offered by the multitone paradigm.
doi:10.1007/s10162-010-0233-4
PMCID: PMC3015028  PMID: 20838846
DPOAEs; otoacoustic; gerbil; Meriones unguiculatus; tone complex; zwuis
7.  Distortion Product Otoacoustic Emissions Evoked by Tone Complexes 
Distortion product otoacoustic emissions (DPOAEs) are traditionally evoked by two-tone stimuli. In this study, emission data from Mongolian gerbils are reported that were obtained with stimuli consisting of six to 10 tones. The stimuli were constructed by replacing one of the tones of a tone pair by a narrowband multitone complex. This produced rich spectra of the ear canal sound pressure in which many of the third-order DPOAEs originated from the interaction of triplets of stimulus components. A careful choice of the stimulus frequencies ensured that none of these DPOAE components coincided. Three groups of DPOAEs are reported, two of which are closely related to DPOAEs evoked by tone pairs. The third group has no two-tone equivalent and only arises when using a multitone stimulus. We analyzed the relation between multitone-evoked DPOAEs and DPOAEs evoked by tone pairs, and explored the new degrees of freedom offered by the multitone paradigm.
doi:10.1007/s10162-010-0233-4
PMCID: PMC3015028  PMID: 20838846
DPOAEs; otoacoustic; gerbil; Meriones unguiculatus; tone complex; zwuis
8.  Temperature Dependence of Anuran Distortion Product Otoacoustic Emissions 
To study the possible involvement of energy-dependent mechanisms in the transduction of sound within the anuran ear, distortion product otoacoustic emissions (DPOAEs) were recorded in the northern leopard frog over a range of body temperatures. The effect of body temperature depended on the stimulus levels used and on the hearing organ under investigation. Low-level DPOAEs from the amphibian papilla (AP) were reversibly depressed for decreased body temperatures. Apparently, DPOAE generation in the AP depends on metabolic rate, indicating the involvement of active processes in the transduction of sound. In contrast, in the other hearing organ, the basilar papilla (BP), the effects of body temperature on DPOAEs were less pronounced, irrespective of the stimulus levels used. Apparently, metabolic rate is less influencing DPOAE generation. We interpret these results as evidence that no amplifier is involved in sound transduction in the BP. The passive functioning of the anuran BP would place this hearing organ in a unique position within tetrapod hearing, but may actually be beneficial to ectothermic species because it will provide the animal with a consistent spectral window, regardless of ambient or body temperature.
doi:10.1007/s10162-006-0039-6
PMCID: PMC2504610  PMID: 16724291
distortion product otoacoustic emissions; anuran; temperature dependence; passive hearing
9.  Detailed f1, f2 Area Study of Distortion Product Otoacoustic Emissions in the Frog 
Distortion product otoacoustic emissions (DPOAEs) are weak sounds emitted from the ear when it is stimulated with two tones. They are a manifestation of the nonlinear mechanics of the inner ear. As such, they provide a noninvasive tool for the study of the inner ear mechanics involved in the transduction of sound into nerve fiber activity. Based on the DPOAE phase behavior as a function of frequency, it is currently believed that mammalian DPOAEs are the combination of two components, each generated by a different mechanism located at a different location in the cochlea. In frogs, instead of a cochlea, two separate hearing papillae are present. Of these, the basilar papilla (BP) is a relatively simple structure that essentially functions as a single auditory filter. A two-mechanism model of DPOAE generation is not expected to apply to the BP. In contrast, the other hearing organ, the amphibian papilla (AP), exhibits a tonotopic organization. In the past it has been suggested that this papilla supports a traveling wave in its tectorial membrane. Therefore, a two-mechanism model of DPOAE generation may be applicable for DPOAEs from the AP. In the present study we report on the amplitude and phase of DPOAEs in the frog ear in a detailed f1, f2 area study. The result is markedly different from that in the mammalian cochlea. It indicates that DPOAEs generated by neither papilla agree with the two-mechanism traveling wave model. This confirms our expectation for the BP and does not support the hypothesized presence of a mechanical traveling wave in the AP.
doi:10.1007/s10162-004-5019-0
PMCID: PMC2504638  PMID: 15735935
distortion product otoacoustic emissions; amphibian; frog; traveling wave; two-mechanism DPOAE model

Results 1-9 (9)