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The topographic distribution of retinal ganglion cells was investigated in a retinal wholemount of an Irrawaddy dolphin (Orcaella brevirostris). Two zones of increased concentration of ganglion cells were observed—one in the temporal segment and the other in the nasal segment of the retina. The maximal cell concentration was 250 cell/mm2 in the temporal area and 194 cell/mm2 in the nasal area. Based on the posterior nodal distance of 11 mm, the resolution was evaluated as 19.8 arc min in the temporal retinal area (the frontal visual field) and 22.4 arc min in the nasal retinal area (the temporal visual field). Among retinal ganglion cells, a group of giant cells that ranged from 42 to 52 µm in diameter were observed; these cells represented 8% of the total population. The obtained data allowed us to suggest that the presence of two areas of concentrated ganglion cells is characteristic of many delphinids inhabiting both clear oceanic waters or turbid river waters, whereas low ganglion cell density is associated with optic properties of the media.

Sound stimuli with rippled spectra are usable for testing the frequency resolving power of hearing. Estimates of the resolution depend on the center frequency and bandwidth of the test stimulus. In the present study, ripple density resolution was measured at various center frequencies and various bandwidths in two experimental paradigms: with the use of rippled or flat reference signals. With rippled reference signals, there was no statistically significant resolution dependence on either the center frequency or bandwidth. In all cases, the resolution was near 10 ripples/oct. With flat (nonrippled) reference signals, the ripple density resolution depended on both the center frequency and bandwidth: the higher the center frequency and the wider the frequency band, the higher the resolution was. The difference between resolution estimates obtained with different reference signals may be explained by different contributions of the spectral and temporal mechanisms of frequency analysis. The spectral (excitation pattern) mechanism is effective for discriminating between a rippled test and a rippled reference signal. The temporal processing mechanism is effective for discriminating between a rippled test and a flat reference signal.

Morphological investigation of the retinal ganglion layer in the rough-toothed dolphin Steno bredanensis was performed using the retinal wholemount technique. Mapping of the density of ganglion cells revealed two areas of their concentration: in the nasal (544 cells/mm2) and temporal (594 cells/mm2) quadrants. Ganglion cell diameters ranged from 12 to 50 µm, with mean values varying from 20 to 29 µm in different areas. The obtained cell densities corresponded to a retinal resolution of 11'.

The current levels of development of diagnostic methods for the sensitivity and frequency resolving power (FRP) of hearing are significantly different. Devices for measuring FRP have not entered clinical practice. Laboratory methods for measuring FRP are not suitable for practical applications. Signals with rippled spectra may provide an effective tool for testing the FRP of hearing both in studies of the basic mechanisms of hearing and in practical audiology. The use of such signals for testing allows measurement not of the acuteness of individual frequency-selective filter channels but directly assesses their ability to analyze sound signals with complex time-spectral patterns. In addition, measurement of FRP using rippled test signals is convenient for practical application. Measurements using rippled test signals yield data on the real ability of the auditory system to discriminate complex sound signals. Furthermore, these measurements demonstrate the roles of a number of basic auditory mechanisms – compressive nonlinearity, lateral suppression, and the frequency- and time-based mechanisms of frequency analysis – in the perception of complex auditory signals. Depending on the discrimination task, either frequency or time analysis is activated, giving fundamentally different evaluations of FRP. There is successful experience of the use of test signals with rippled spectra for assessment of auditory acuity in users of cochlear implants.

Forward masking was investigated by the auditory evoked potentials (AEP) method in a bottlenose dolphin Tursiops truncatus using stimulation by two successive acoustic pulses (the masker and test) projected from spatially separated sources. The positions of the two sound sources either coincided with or were symmetrical relative to the head axis at azimuths from 0 to ± 90°. AEPs were recorded either from the vertex or from the lateral head surface next to the auditory meatus. In the last case, the test source was ipsilateral to the recording side, whereas the masker source was either ipsi- or contralateral. For lateral recording, AEP release from masking (recovery) was slower for the ipsi- than for the contralateral masker source position. For vertex recording, AEP recovery was equal both for the coinciding positions of the masker and test sources and for their symmetrical positions relative to the head axis. The data indicate that at higher levels of the auditory system of the dolphin, binaural convergence makes the forward masking nearly equal for ipsi- and contralateral positions of the masker and test.

Eye sizes and retinal organization were studied in newborn and adult bottlenose dolphins Tursiops truncatus. In the newborn animals, the eye optics and retinal organization were completely formed, including the presence of two zones of increased concentration of retinal ganglion cells. Adult animals had larger eye sizes as compared to the newborn ones, larger retinal thickness, larger retinal area, and more ganglion cells. The peak ganglion cell density in the newborn animals was higher than that in the adult ones; however, the retinal resolution differed insignificantly. It is shown that the postnatal development does not principally modify the eye organization but results in a quantitative shift of the main parameters.

Hearing is considered the primary sensory modality of cetaceans and enables their vital life functions. Information on the hearing sensitivity variability within a species obtained in a biologically relevant wild context is fundamental to evaluating potential noise impact and population-relevant management. Here, non-invasive auditory evoked-potential methods were adopted to describe the audiograms (11.2–152 kHz) of a group of four wild Yangtze finless porpoises (Neophocaena asiaeorientalis asiaeorientalis) during a capture-and-release health assessment project in Poyang Lake, China. All audiograms presented a U shape, generally similar to those of other delphinids and phocoenids. The lowest auditory threshold (51–55 dB re 1 µPa) was identified at a test frequency of 76 kHz, which was higher than that observed in aquarium porpoises (54 kHz). The good hearing range (within 20 dB of the best hearing sensitivity) was from approximately 20 to 145 kHz, and the low- and high-frequency hearing cut-offs (threshold > 120 dB re l μPa) were 5.6 and 170 kHz, respectively. Compared with aquarium porpoises, wild porpoises have significantly better hearing sensitivity at 32 and 76 kHz and worse sensitivity at 54, 108 and 140 kHz. The audiograms of this group can provide a basis for better understanding the potential impact of anthropogenic noise.

The impact of maskers on the receiving beam of a bottlenose dolphin, Tursiops truncatus, was investigated using the auditory evoked potential (AEP) method. The test signal was a train of tone pips with a 64 kHz carrier frequency. The stimulus produced AEPs as a sequence of waves replicating the pip rate: the rate following response (RFR). The masker was band-limited noise, with a passband of 45 to 90 kHz and a level of 105 dB re 1 μPa. Masker azimuths were at 0°, ±30°, ±60°, and ±90° relative to the head midline. The receiving beam was evaluated in terms of the RFR threshold dependence on the signal azimuth. The masked thresholds were higher than the baseline thresholds, which appeared mostly as a shift rather than a deformation in the receiving beam. The largest threshold shift appeared when the masker source was located in the most sensitive direction (zero azimuth); at lateral masker source positions, the threshold shift decreased. When the masker source was not at the head midline, the masked thresholds were higher at signal positions ipsilateral to the masker source than at positions contralateral to the masker source. The largest asymmetry was observed at the 30° masker azimuth in conjunction with the ±30° and ±120° signal azimuths; the asymmetries were 5.6 and 8.1 dB, respectively. This masking asymmetry was lower than expected from the previously found interaural intensity difference, which may be explained by the conflict between the test signal and the masker when it appeared at a binaural level of the auditory system.

The correlation between ripple spectrum resolution and speech recognition in prelingual and postlingual cochlear implant users was analysed. The test signal had a band-limited rippled spectrum with a bandwidth of 2 octaves. The central frequency of the spectrum was 1, 2, or 4 kHz. The ripple phase reversion test of was used for estimation of ripple spectrum discrimination. A balanced test set of Russian worlds by Greenberg–Zinder was used for estimating the speech recognition. The average ripple discrimination thresholds were approximately 2 ripples/oct in both pre- and postlingual patients. The thresholds did not depend on the central frequency. The best correlation between speech recognition and rippled spectrum resolution was found in the prelingual patients with a perimodiolar electrode array (r = 0.85).

Forward masking was investigated by the auditory evoked potentials (AEP) method in a bottlenose dolphin Tursiops truncatus using stimulation by two successive acoustic pulses (the masker and test) projected from spatially separated sources. The positions of the two sound sources either coincided with or were symmetrical relative to the head axis at azimuths from 0 to ±90°. AEPs were recorded either from the vertex or from the lateral head surface next to the auditory meatus. In the last case, the test source was ipsilateral to the recording side, whereas the masker source was either ipsi- or contralateral. For lateral recording, AEP release from masking (recovery) was slower for the ipsi- than for the contralateral masker source position. For vertex recording, AEP recovery was equal both for the coinciding positions of the masker and test sources and for their symmetrical positions relative to the head axis. The data indicate that at higher levels of the auditory system of the dolphin, binaural convergence makes the forward masking nearly equal for ipsi- and contralateral positions of the masker and test.

Highly concentrated live mass stranding events of dolphins and whales happened in the eastern coast of China between June and October 2021. The current study adopted the non-invasive auditory evoked-potential technique to investigate the hearing threshold of a stranded melon headed whale (Peponocephala electra) at a frequency range of between 9.5 and 181 kHz. It was found that, at the frequency range of from 10 to 100 kHz, hearing thresholds for the animal were between 20 and 65 dB higher than those of its phylogenetically closest species (Pygmy killer whale). The severe hearing loss in the melon headed whale was probably caused by transient intense anthropogenic sonar or chronic shipping noise exposures. The hearing loss could have been the cause for the observed temporal and spatial clustered stranding events. Therefore, there is need for noise mitigation strategies to reduce noise exposure levels for marine mammals in the coastal areas of China.

Auditory adaptation manifests as temporary change in the hearing sensitivity under influence of sound stimuli, and little attention has been paid to hearing adaptation in cetaceans. In a Yangtze fi...

The objective of the study was to better understand of contribution of excitation-pattern and temporal-processing mechanisms of frequency analysis to discrimination of complex-spectrum signals in various discrimination tasks. Using rippled-spectrum signals, the ripple depth thresholds were measured as functions of ripple density under conditions of rippled or non-rippled reference signals. With rippled reference signals, the ripple depth thresholds were as low as 0.11 at low ripple densities (2–3 cycles/oct) and rose to 1.0 at a ripple density of 8.9 cycles/oct. For non-rippled reference signals, ripple depth thresholds were nearly the same as for rippled reference signals at ripple densities of up to 7 cycles/oct; at ripple densities of 10 cycles/oct and higher, ripple depth thresholds rose slowly and reached 1.0 at a ripple density of 26 cycles/oct. The results hypothetically suggest contributions of the excitation-pattern processing and temporal-processing mechanisms of frequency analysis to discrimination of rippled signals. The excitation-pattern mechanism featured low depth thresholds at low ripple densities but could not function at ripple densities above 10 cycles/oct. The temporal-processing mechanism manifested at higher ripple densities and non-rippled reference stimuli.

At suprathreshold sound levels, interactions between masking noise and sound signals are liable to compressive nonlinearity in the auditory system. The compressive nonlinearity is a property of the "active" cochlear mechanism. It is not known whether this mechanism is capable to function at frequencies close to or above 100 kHz that are available to odontocetes (toothed whales, dolphins, and porpoises). This question may be answered by the use of the frequency-specific masking. Auditory evoked potentials to sound stimuli in a bottlenose dolphin, Tursiops truncatus, were recorded in the presence of simultaneous maskers. Stimulus frequencies were 45, 64, or 90 kHz. Maskers were on-frequency bandlimited noise or low-frequency noise of frequencies 0.25-1 oct below the stimulus frequency. The stimuli provoked responses as a series of brain-potential waves following the pip-train rate. For the on-frequency masker, the masker level at threshold dependence on the signal level was 1.1 dB/dB. For maskers of 1 oct below the stimulus, the dependence was 0.53-0.57 dB/dB. The data considered evidence for the compressive nonlinearity of responses to stimuli, and therefore, are indicative of the functioning of the active mechanism at frequencies up to 90 kHz.

The effects of adapting sounds (pip trains or pure tones) on auditory evoked potentials (the rate following response, RFR) were investigated in a beluga whale. During RFR acquisition, adapting signals lasting 128 ms each were alternated with test signals lasting 16 ms each; the test signal levels varied randomly. Adapting signals were trains of cosine-enveloped tone pips or pure tones. Pip rate varied with the envelope cosine cycle maintained at 0.125 of pip intervals and the cosine rise-fall time maintained at 0.0625 of pip intervals. Adapting signals shifted the amplitude-level function upward compared to the baseline (no adapting signal) function. The higher the adapting signal level was, the bigger the shift in the amplitude-level function was. The slower the pips were in the adapting signal, the smaller the adaptation effect was. A train of pips with a 0.0625-ms rise-fall time and 125 dB SPL shifted the function by 35-40 dB, whereas a train of pips with a 1-ms rise-fall time or a pure tone with the same SPL shifted the function by approximately 15 dB. The difference between the "fast" and "slow" adapting signals is supposed to be associated with their abilities to stimulate the auditory system in odontocetes.

In a beluga whale, the positions of sound receiving areas on the head surface were determined by comparing the acoustic delays from different sound source positions. For this investigation, auditory evoked potentials (AEPs) in response to short tone pips were recorded. Latencies of the first AEP wave that presumably reflected the activity of the auditory nerve were measured at different sound source azimuths. For AEPs of equal amplitudes, the difference in AEP latencies was attributed to the difference in the acoustic delays. These delay differences were used to compute the azimuths of sound receiving points. Measurements were conducted at frequencies from 22.5 to 90 kHz in half-octave steps. At all stimulus frequencies, the receiving points were located 24–38 cm caudal of the melon tip, which is near a proximal part of the lower jaw. Thus, the results indicated the latero-mandibular acoustic window. Possible causes for not finding a lateral or ventro-mandibular window are discussed.In a beluga whale, the positions of sound receiving areas on the head surface were determined by comparing the acoustic delays from different sound source positions. For this investigation, auditory evoked potentials (AEPs) in response to short tone pips were recorded. Latencies of the first AEP wave that presumably reflected the activity of the auditory nerve were measured at different sound source azimuths. For AEPs of equal amplitudes, the difference in AEP latencies was attributed to the difference in the acoustic delays. These delay differences were used to compute the azimuths of sound receiving points. Measurements were conducted at frequencies from 22.5 to 90 kHz in half-octave steps. At all stimulus frequencies, the receiving points were located 24–38 cm caudal of the melon tip, which is near a proximal part of the lower jaw. Thus, the results indicated the latero-mandibular acoustic window. Possible causes for not finding a lateral or ventro-mandibular window are discussed.

Psychophysical experiments on listeners with normal hearing were conducted to discriminate the rippled spectra of an acoustic signal against maskers with different positions of the spectral band relative to the signal band. As the signal level changed from 50 to 80 dB SPL, the on-frequency masker level changed by 29 dB, whereas the low-frequency masker level (the position of the center of the spectral band was from –1.25 to –1 octave relative to the signal) changed by 8.7–9.8 dB. These results are interpreted as 0.3 dB/dB compression of responses to the signal and no compression of the effect of low-frequency maskers. If the spectral bands of the signal and masker partially overlap, discrimination of the spectral structure occurs predominantly in the part of the spectrum that does not overlap the masker spectrum and is subjected to low-frequency masking that is not compressed.

Two paradigms have been compared, which were aimed to investigate an effect of an additional signal pattern on ripple density thresholds. The discrimination between an extreme ripple density reference and rippled test pattern and the discrimination between a rippled reference and rippled test stimuli were measured. Stimuli contained or did not contain an additional signal pattern. The additional signal had the same length, bandwidth, and level as the signal. A ripple density of the additional signal was varied from 0 to 7 ripples/oct. For the stimuli with an additional rippled signal and non-rippled reference signal, the mean resolution was 11 ripples/oct with 16,47 ripples/oct for a non-rippled additional signal, and 52.4 ripples/oct with no additional signal. For a rippled additional signal, the mean resolution was 8.86 with the additional signal ripples/oct and 8.5 ripples/oct with no additional signal. For non-rippled reference signals results could be explained by the temporal mechanism when spectral pattern of the stimulus was unresolvable. For rippled reference signals, the data can be explained by similar cochlear excitation patterns.

The “active” cochlear mechanism of hearing manifests in the cochlear compression. Investigations of compression in odontocetes help to determine the frequency limit of the active mechanism. The compression may be evaluated by comparison of low- and on-frequency masking. In a bottlenose dolphin, forward masking of auditory evoked potentials to tonal pips was investigated. Measurements were performed for test frequencies of 45 and 90 kHz. The low-frequency maskers were − 0.25 to − 0.75 oct relative the test. Masking efficiency was varied by masker-to-test delay variation from 2 to 20 ms, and masker levels at threshold (MLTs) were evaluated at each of the delays. It was assumed that low-frequency maskers were not subjected or little subjected to compression whereas on-frequency maskers were subjected equally to the test. Therefore, the compression rate was assessed as the slope of low-frequency MLT dependence on on-frequency MLT. For the 90-kHz test, the slopes were 0.63 and 0.18 dB/dB for masker of − 0.25 and − 0.5 oct, respectively. For the 45 kHz test, the slopes were 0.69 and 0.39 dB/dB for maskers of − 0.25 and − 0.5 oct. So, compression did not decay at the upper boundary of the hearing frequency range in the dolphin.

Signals with rippled frequency spectra are used to investigate capabilities of hearing to discriminate complex sounds. The maximum resolvable ripple density (FRP). FRP estimates depend on the bandwidth and central frequency of the signals as well as in the measurement paradigm, specifically, the type of the used reference signal in the discrimination task. In order to find how these parameters influence FRP, the measurements were performed at various signal center frequencies and bandwidths in two experimental designs: with the use of rippled or “flat” reference signals. For rippled reference signals, FRP was about 10 cycles/oct with negligible FRP dependence on signal frequency or bandwidth. For flat reference signals, substantial positive FRP dependence on signal frequency and bandwidth occurred. Models predicting the difference RFP dependence on the type of reference signals are suggested. The models imply different contributions of the excitation-pattern and temporal-processing mechanisms of frequency analysis in rippled-spectrum discrimination. The excitation-pattern mechanism predominates in discrimination of a rippled test and a rippled reference signal whereas the temporal-processing mechanism predominates in discrimination between a rippled test and a “flat” signal.

Frequency resolving power (FRP) was measured in normal listeners. FRP was estimated on the basis of the maximum resolvable ripple density in rippled-spectrum signals. Two measurement procedures were compared: detection of ripple-pattern change and comparison of rippled-spectrum signals. In the change detection method, two successive sound signals were presented to the listener: a test signal and a reference signal. The test signal contained ripple phase reversals every 400 ms; in the reference signal, the ripple phase was constant. The listener’s task was to identify the test signal. In the comparison method, three signals were presented to the listener. The ripple phase in one of the three signals was opposite to that in other two signals. The listener’s task was to identify the signal different from the other two signals. The signal frequency bands varied from 0.5 to 5 oct at a level of 0.5 of the maximum. At all frequency bands, the change-detection method yielded, on average, 1.75 oct–1 higher FRP estimates compared to the comparison method. This difference between the two methods is supposed to be due to the greater involvement of cognitive processes (short-term memory) in the comparison method. The change-detection method is more preferable for measuring the sensory component of FRP.

The goal of the study was to assess cochlear compression when rippled-spectrum signals are perceived in noise assuming that the noise might produce both masking and confounding effects. In normal listeners, discrimination between rippled signals with and without ripple phase reversals was assessed in background noise. The signals were band-limited (0.5 oct at a − 6-dB level) rippled noise centered at 2 kHz, with a ripple density of 3.5 oct(−1). The noise (masker) was band-limited nonrippled noise centered at either 2 kHz (on-frequency masker) or 1 kHz (low-frequency masker). The masker was simultaneously presented with the signals. Masker levels at the discrimination threshold were measured as a function of the signal level using the adaptive (staircase) two-alternative forced-choice procedure. For the on-frequency masker, the searched-for function had a slope of 0.98 dB/dB. For the low-frequency masker, the function had a slope of 1.19 dB/dB within a signal level range of 30 to 40 dB sound pressure level (SPL) and as low as 0.15 dB/dB within a signal level range of 70 to 80 dB SPL. These results were interpreted as indicating compression of responses to both the signal and on-frequency masker and no compression of the effect of the low-frequency masker. In conditions when above-threshold signals are presented in simultaneous noise (the masker), cochlear compression manifests to a substantial degree despite possible confounding effects.

The resolution of spectral patterns in adaptation background noise was investigated in a beluga whale, Delphinapterus leucas, using the evoked-potential technique. The resolution of spectral patterns was investigated using rippled-spectrum test stimuli of various levels and ripple densities and recording the rhythmic evoked responses (the rate following response, RFR) to ripple phase reversals. In baseline (no adaptation background noise) experiments, the highest RFR magnitude was observed at signal sound pressure levels (SPLs) of 100-110 dB re 1 μPa; at SPLs both below the optimum (down to 80 dB re 1 μPa) and above the optimum (up to 140 dB re 1 μPa), the RFR magnitude decreased. For high signal levels (above 110 dB re 1 μPa), low-level adaptation background noise (from -10 to -20 dB re signal level) increased RFR magnitude compared to baseline. This effect is considered to be a result of the optimization of the sensation level of the high-SPL signals due to decreasing hearing sensitivity caused by the adaptation background noise.

Hearing sensitivity change was investigated when a warning sound preceded a loud sound in the false killer whale (Pseudorca crassidens), the bottlenose dolphin (Tursiops truncatus), the beluga whale (Delphinaperus leucas) and the harbor porpoise (Phocoena phocoena). Hearing sensitivity was measured using pip-train test stimuli and auditory evoked potential recording. When the test/warning stimuli preceded a loud sound, hearing thresholds before the loud sound increased relative to the baseline by 13 to 17 dB. Experiments with multiple frequencies of exposure and shift provided evidence of different amounts of hearing change depending on frequency, indicating that the hearing sensation level changes were not likely due to a simple stapedial reflex.

In psychophysical experiments on humans, a cochlear compression was revealed by comparing onfrequency and low-frequency masking in the conditions of discrimination of complex spectra of sound signals. It has been established that, with an increase in the level of the masker, the limit of the discriminable density of the spectrum ripples decreases. It was noted that when the signal level increased, the on-frequency masker level at threshold grew proportionally to the level of the signal, whereas the low-frequency masker level grew much more slowly, indicating the compression of the signal, which was not affected by lateral suppression and off-frequency listening.

In the psychophysical experiments reported here, cochlear compression function was derived by comparing on-frequency and off-frequency masking. The signal was rippled spectrum noise. The ripple density discrimination threshold was measured in the ripple phase reversion test. An increase in masker intensity led to a decrease in a resolvable ripple density threshold. The on-frequency masker level at threshold increased proportionally to the signal intensity. The off-frequency masker level at threshold also increased proportionally to the signal at signal intensity levels below 50 dB, whereas at signal levels above 60 dB SPL, the ratio of the masker level at threshold gradient to signal level gradient was 1 : 5 dB/dB, revealing cochlear compression.

The eye optics and topographic distribution of ganglion cells were studied using whole mount preparations from European beaver Castor fiber L. The beaver eye optics provides emmetropia in air and hypermetropia in water. The optometrical measurements predict retinal resolution of the beaver eye around 17' in air and 9' in water. In air, retinal resolution corresponds to the real visual acuity, whereas in water, visual acuity is below the retinal resolution because of the non-precise focusing.

The resolution of spectral ripples is a useful test for the spectral resolution of hearing. However, the use of different measurement paradigms might yield diverging results because of a paradigm-dependent contribution of excitation-pattern and temporal-processing mechanisms. In the present study, ripple-density resolution was measured in normal-hearing listeners for several frequency bands (centered at 0.5, 1, 2, and 4 kHz), using two paradigms: (i) discrimination of a rippled-spectrum test signal from a rippled reference signal differing by the ripple phase pattern, and (ii) discrimination of a rippled-spectrum test signal from a nonrippled reference signal. For the rippled reference signals, the resolution slightly depended on signal frequency. For the nonrippled reference signals, the resolution depended on the signal frequency; it varied from 8.8 ripples/oct at 0.5 kHz to 34.2 ripples/oct at 4 kHz. Excitation-pattern and temporal-processing models of spectral analysis were considered. Predictions of the excitation-pattern model agreed with the data obtained with the rippled reference signals. In contrast, predictions of the temporal-processing model agreed with the data obtained with the nonrippled reference signals. Thus, depending on the used reference signal type, the ripple-density resolution estimates characterize the discrimination abilities of the corresponding mechanisms.

The topography of ganglion cells in the retina of three neonatal bottlenose dolphins Tursiops truncatus has been studied in retinal wholemounts. Two areas of high ganglion cell density have been identified in the form of local spots in the temporal and nasal quadrants of the retina, near a horizontal diameter at a distance of 10–15 mm from the optical disk. The maximum density of ganglion cells in these areas on average for five preparations is 657 and 636 cells/mm2 in the temporal and nasal quadrants, respectively. The retinal resolution, estimated by the maximum density of ganglion cells and a posterior nodal distance of 13 mm, was 0.17° for the temporal quadrant and 0.18° for the nasal one. These data are comparable with the results for the previously studied adult dolphin and whale species.

Rippled-spectrum stimuli are used to evaluate the resolution of the spectro-temporal structure of sounds. Measurements of spectrum-pattern resolution imply the discrimination between the test and reference stimuli. Therefore, estimates of rippled-pattern resolution could depend on both the test stimulus and the reference stimulus type. In this study, the ripple-density resolution was measured using combinations of two test stimuli and two reference stimuli. The test stimuli were rippled-spectrum signals with constant phase or rippled-spectrum signals with ripple-phase reversals. The reference stimuli were rippled-spectrum signals with opposite ripple phase to the test or nonrippled signals. The spectra were centered at 2 kHz and had an equivalent rectangular bandwidth of 1 oct and a level of 70 dB sound pressure level. A three-alternative forced-choice procedure was combined with an adaptive procedure. With rippled reference stimuli, the mean ripple-density resolution limits were 8.9 ripples/oct (phase-reversals test stimulus) or 7.7 ripples/oct (constant-phase test stimulus). With nonrippled reference stimuli, the mean resolution limits were 26.1 ripples/oct (phase-reversals test stimulus) or 22.2 ripples/oct (constant-phase test stimulus). Different contributions of excitation-pattern and temporal-processing mechanisms are assumed for measurements with rippled and nonrippled reference stimuli: The excitation-pattern mechanism is more effective for the discrimination of rippled stimuli that differ in their ripple-phase patterns, whereas the temporal-processing mechanism is more effective for the discrimination of rippled and nonrippled stimuli.

Rippled-spectrum signals are used to measure signal resolution in hearing-impaired listeners and cochlear implant users. Two mechanisms are believed to be responsible for rippled-spectrum resolution. The excitation-pattern mechanism determines the ripple density resolution (ripples/oct). The temporal processing mechanism determines the ripple frequency spacing limit (kHz). The contributions of these two mechanisms can be assessed by comparing the resolutions of band-limited rippled spectra with different center frequencies. Ripple-density resolutions of half-octave rippled spectra were measured at center frequencies from 0.5 to 4 kHz. Measurements were performed either by discrimination between the rippled-spectrum test and reference signals differing by ripple phases or by discrimination between the rippled-spectrum test and a non-rippled reference signal. For discrimination between the rippled-spectrum test and reference signals, resolutions specified in ripples/oct minimally depended on the center frequency, as predicted by the excitation-pattern model. For discrimination between the rippled test and non-rippled reference signals, the resolution specified by the ripple frequency spacing minimally depended on the center frequency, as predicted by the temporal processing model. It was concluded that the contributions of the excitation-pattern and temporal processing mechanisms depended on the discrimination task.

Rippled-spectrum signals are used for signal-resolution measurements in hearing-impaired listeners and cochlear implant users. Two mechanisms are presumably responsible for rippled-spectrum resolution. The excitation-pattern mechanism determines the ripple-density resolution (ripples/oct). The temporal-processing mechanism determines the ripple frequency spacing limit (kHz). The contributions of these two mechanisms can be assessed by comparing the resolutions of band-limited rippled spectra with different center frequencies. Ripple-density resolutions of half-octave rippled spectra were measured at center frequencies ranging from 0.5 to 4 kHz. The measurements were performed either by discrimination between a rippled-spectrum test and reference signals differing by ripple phases or by discrimination between a rippled-spectrum test and a nonrippled reference signal. For discrimination between rippled-spectrum tests and reference signals, the resolution specified in ripples/oct depended very little on the center frequency, as predicted by the excitation-pattern model. For discrimination between rippled tests and nonrippled reference signals, the resolution specified in the ripple frequency spacing depended very little on the center frequency, as predicted by the temporal-processing model. This study concluded that contributions of the excitation-pattern and temporal-processing mechanisms depend on the discrimination task.Rippled-spectrum signals are used for signal-resolution measurements in hearing-impaired listeners and cochlear implant users. Two mechanisms are presumably responsible for rippled-spectrum resolution. The excitation-pattern mechanism determines the ripple-density resolution (ripples/oct). The temporal-processing mechanism determines the ripple frequency spacing limit (kHz). The contributions of these two mechanisms can be assessed by comparing the resolutions of band-limited rippled spectra with different center frequencies. Ripple-density resolutions of half-octave rippled spectra were measured at center frequencies ranging from 0.5 to 4 kHz. The measurements were performed either by discrimination between a rippled-spectrum test and reference signals differing by ripple phases or by discrimination between a rippled-spectrum test and a nonrippled reference signal. For discrimination between rippled-spectrum tests and reference signals, the resolution specified in ripples/oct depended very little on the ...

The goal of the study was to investigate the role of combination products in the higher ripple-density resolution estimates obtained by discrimination between a spectrally rippled and a nonrippled noise signal than that obtained by discrimination between two rippled signals. To attain this goal, a noise band was used to mask the frequency band of expected low-frequency combination products. A three-alternative forced-choice procedure with adaptive ripple-density variation was used. The mean background (unmasked) ripple-density resolution was 9.8 ripples/oct for rippled reference signals and 21.8 ripples/oct for nonrippled reference signals. Low-frequency maskers reduced the ripple-density resolution. For masker levels from −10 to 10 dB re. signal, the ripple-density resolution for nonrippled reference signals was approximately twice as high as that for rippled reference signals. At a masker level as high as 20 dB re. signal, the ripple-density resolution decreased in both discrimination tasks. This result leads to the conclusion that low-frequency combination products are not responsible for the task-dependent difference in ripple-density resolution estimates.

The effects of prolonged (up to 1500 s) sound stimuli (tone pip trains) on evoked potentials (the rate following response, RFR) were investigated in a beluga whale. The stimuli (rhythmic tone pips) were of frequencies of 45, 64, and 90 kHz at levels from 20 to 60 dB above threshold. Two experimental protocols were used: short- and long-duration. For the short-duration protocol, the stimuli were 500-ms-long pip trains that repeated at a rate of 0.4 trains/s. For the long-duration protocol, the stimuli were continuous pip successions lasting up to 1500 s. The RFR amplitude gradually decreased by three to seven times from 10 ms to 1500 s of stimulation. Decrease of response amplitude during stimulation was approximately proportional to initial (at the start of stimulation) response amplitude. Therefore, even for low stimulus level (down to 20 dB above the baseline threshold) the response was never suppressed completely. The RFR amplitude decay that occurred during stimulation could be satisfactorily approximated by a combination of two exponents with time constants of 30-80 ms and 3.1-17.6 s. The role of adaptation in the described effects and the impact of noise on the acoustic orientation of odontocetes are discussed.

Short-latency auditory evoked potentials (SLAEP) produced in response to sound impulses with strictly specifi ed 0.5-oct frequency bands, from 11.2–16 to 90–128 kHz, were recorded in bottle-nose dolphins. Using stimuli of constant intensity, SLAEP amplitude depended on the spectral frequency of the stimulus: the higher the frequency, the greater the amplitude. Response shape showed little dependence on spectral band, though the response delay changed with frequency: the higher the frequency, the shorter the delay. The differences in delays between the responses to low-frequency (11.2–16 kHz) and high-frequency (90–128 kHz) stimuli was 0.3 msec. This difference in response delays could be attributed to the propagation time of the traveling wave in the cochlea; the results obtained here were used to calculate the velocity of the traveling wave. These assessments of velocity were from 38.2 oct/msec at the proximal (high-frequency) end of the basilar membrane to 2.8 oct/msec at the distal (low-frequency) end. Comparison of traveling wave velocities in humans and dolphins showed that traveling wave velocity was not linked to the location on the cochlea, but to the characteristic frequency.

The total number, size, topographic distribution, and cell density of ganglion cells were studied in retinal wholemounts of Baikal seals (Pusa sibirica). The ganglion cell size varied from 10 to 38 μm. A distinct cell group consisted of large ganglion cells of more than 30 μm in diameter. The topographic distribution of ganglion cells showed a definite area of high cell density similar to the area centralis of terrestrial carnivores. This area was located approximately 6-7 mm dorsotemporally of the geometric center of the wholemount. In this area, the peak cell densities in two wholemounts were 3,800 and 3,400 cells/mm2 (mean 3,600 cells/mm2). With a posterior nodal distance of 24 mm (underwater), this density corresponds to 631 cells/square degree. These values predict a retinal resolution of 2.4′ in water and 3.0′ in air. The topographic distribution of large cells featured the highest density in the same location as the total ganglion cell population.