H. Y. Yan

A comparative study of hearing ability in fishes: the auditory brainstem response approach

Auditory brainstem response (ABR) techniques, an electrophysiological far-field recording method widely used in clinical evaluation of human hearing, were adapted for fishes to overcome the major limitations of traditional behavioral and electrophysiological methods (e.g., invasive surgery, lengthy training of fishes, etc.) used for fish hearing research. Responses to clicks and tone bursts of different frequencies and amplitudes were recorded with cutaneous electrodes. To evaluate the effectiveness of this method, the auditory sensitivity of a hearing specialist (goldfish, Carassius auratus) and a hearing generalist (oscar, Astronotus ocellatus) was investigated and compared to audiograms obtained through psychophysical methods. The ABRs could be obtained between 100 Hz and 2000 Hz (oscar), and up to 5000 Hz (goldfish). The ABR audiograms are similar to those obtained by behavioral methods in both species. The ABR audiogram of curarized (i.e., Flaxedil-treated) goldfish did not differ significantly from two previously published behavioral curves but was lower than that obtained from uncurarized fish. In the oscar, ABR audiometry resulted in lower thresholds and a larger bandwidth than observed in behavioral tests. Comparison between methods revealed the advantages of this technique: rapid evaluation of hearing in untrained fishes, and no limitations on repeated testing of animals.

Empirical refinements applicable to the recording of fish sounds in small tanks

Many underwater bioacoustical recording experiments (e.g., fish sound production during courtship or agonistic encounters) are usually conducted in a controlled laboratory environment of small-sized tanks. The effects of reverberation, resonance, and tank size on the characteristics of sound recorded inside small tanks have never been fully addressed, although these factors are known to influence the recordings. In this work, 5-cycle tone bursts of 1-kHz sound were used as a test signal to investigate the sound recorded in a 170-l rectangular glass tank at various depths and distances from a transducer. The dominant frequency, sound-pressure level, and power spectrum recorded in small tanks were significantly distorted compared to the original tone bursts. Due to resonance, the dominant frequency varied with water depth, and power spectrum level of the projected frequency decreased exponentially with increased distance between the hydrophone and the sound source; however, the resonant component was nearly uniform throughout the tank. Based on the empirical findings and theoretical calculation, a working protocol is presented that minimizes distortion in fish sound recordings in small tanks. To validate this approach, sounds produced by the croaking gourami (Trichopsis vittata) during staged agonistic encounters were recorded according to the proposed protocol in an 1800-l circular tank and in a 37-l rectangular tank to compare differences in acoustic characteristics associated with tank size and recording position. The findings underscore pitfalls associated with recording fish sounds in small tanks. Herein, an empirical solution to correct these distortions is provided.

Variability in the role of the gasbladder in fish audition

Abstract The teleost gasbladder is believed to aid in fish audition by transferring pressure components of incoming sound to the inner ears. This idea is primarily based on both anatomical observations of the mechanical connection between the gasbladder and the ear, followed by physiological experiments by various researchers. The gasbladder movement has been modeled mathematically as a pulsating bubble. This study is extending the previous work on fish with a physical coupling of the gasbladder and ear by investigating hearing in two species (the blue gourami Trichogaster trichopterus, and the oyster toadfish Opsanus tau) without a mechanical linkage. An otophysan specialist (the goldfish Carassius auratus) with mechanical coupling, is used as the control. Audiograms were obtained with acoustically evoked potentials (e.g., auditory brainstem response) from intact fish and from the same individuals with their gasbladders deflated. In blue gourami and oyster toadfish, removal of gas did not significantly change thresholds, and evoked potentials had similar waveforms. In goldfish thresholds increased by 33–55 dB (frequency dependent) after deflation, and major changes in evoked potentials were observed. These results suggest that the gasbladder may not serve an auditory enhancement function in teleost fishes that lack mechanical coupling between the gasbladder and the inner ear.

Correlation between auditory sensitivity and vocalization in anabantoid fishes

Abstract Several anabantoid species produce broad-band sounds with high-pitched dominant frequencies (0.8–2.5 kHz), which contrast with generally low-frequency hearing abilities in (perciform) fishes. Utilizing a recently developed auditory brainstem response recording-technique, auditory sensitivities of the gouramis Trichopsis vittata, T. pumila, Colisa lalia, Macropodus opercularis and Trichogaster trichopterus were investigated and compared with the sound characteristics of the respective species. All five species exhibited enhanced sound-detecting abilities and perceived tone bursts up to 5 kHz, which qualifies this group as hearing specialists. All fishes possessed a high-frequency sensitivity maximum between 800 Hz and 1500 Hz. Lowest hearing thresholds were found in T. trichopterus (76 dB re 1 μPa at 800 Hz). Dominant frequencies of sounds correspond with the best hearing bandwidth in T. vittata (1–2 kHz) and C. lalia (0.8–1 kHz). In the smallest species, T. pumila, dominant frequencies of acoustic signals (1.5–2.5 kHz) do not match lowest thresholds, which were below 1.5 kHz. However, of all species studied, T. pumila had best hearing sensitivity at frequencies above 2 kHz. The association between high-pitched sounds and hearing may be caused by the suprabranchial air-breathing chamber, which, lying close to the hearing and sonic organs, enhances both sound perception and emission at its resonant frequency.

AUDITORY ROLE OF THE SUPRABRANCHIAL CHAMBER IN GOURAMI FISH

Abstract Fish hearing specialists (e.g., goldfish, holocentrids, clupeoids, mormyrids) have evolved specialized structures (e.g., Weberian ossicles, swimbladder diverticulae, gas-filled bullae) to enhance their auditory frequency range and threshold sensitivity. The inner ears of anabantoid fish are encased in membranous cranial bones and are protruded into air-filled suprabranchial chambers. This research was intended to test the hypothesis that the gas bubbles inside the suprabranchial chambers may modulate the hearing abilities of anabantoid fish because of their proximity to the membranous bone-encased inner ears. Three species of gourami (blue gourami Trichogaster trichopterus; kissing gourami Helostoma temminckii; dwarf gourami Colisa lalia) were examined. Using the auditory brainstem response recording technique, baseline audiograms tested at 300, 500, 800, 1500, 2500, 4000 Hz were obtained. The air bubbles in the suprabranchial chambers were replaced by water, and the audiograms were remeasured. Thresholds were elevated in all three species. When three blue gouramis were allowed to replenish air into the suprabranchial chambers their hearing abilities returned to baseline levels. These results support the hypothesis that air bubbles in the suprabranchial chambers can affect hearing abilities of gouramis by lowering the thresholds.

The otic gasbladder as an ancillary auditory structure in a mormyrid fish.

Abstract Mormyrid fishes use acoustic signals for long-distance communication and a weakly electric field for short-distance interaction. Mormyrids are unique in having an otic gasbladder attached directly to the saccule on each side of the inner ear. Karl von Frisch (1938) hypothesized that the tightly coupled otic gasbladder might aid mormyrid hearing. Using the mormyrid fish (Brienomyrus brachyistius), this study manipulated gas in the otic gasbladder to test this hypothesis and histological sections were made to examine the anatomical relationship between the gasbladder and inner ear. The hearing sensitivity curves (audiograms) were obtained with the auditory brainstem response protocol. Audiograms were obtained from normal fish and from fish in which gas was withdrawn from either one or two otic gasbladders. Removal of gas from one otic gasbladder did not result in a significant change in either hearing ability or acoustically evoked brainwaves as compared to the control fish. Bilateral deflation of the otic gasbladders led to significant threshold changes. Histological sections revealed a particularly close coupling between the otic gasbladder and the saccule chamber. These results support von Frischs hypothesis that the otic gasbladders of mormyrids assist in underwater sound detection.

The hearing sensitivity of the little skate, Raja erinacea: A comparison of two methods

We determined the hearing sensitivity of the little skate, Raja erinacea using two methods: Behavioral conditioning and the auditory brainstem response (ABR). This marks the first time that the hearing in any member of the Rajiformes has been examined and the first time that the ABR method has been used with an elasmobranch. We obtained audiograms of R. erinacea using each method and were found to be statistically similar. The best hearing sensitivity for R. erinacea was between 100 and 300 Hz. We compared the audiograms to audiograms obtained from other species of elasmobranchs. This analysis showed that R. erinacea, a bottom-dwelling elasmobranch, has less sensitive hearing than the lemon shark, Negaprion brevirostris, and the bull shark, Carcharhinus leucas, a free-swimming, raptorial elasmobranch. However, R. erinacea showed sensitivity comparable to that of the horn shark, Heterodontus francisi, another bottom-dwelling elasmobranch; both species feed primarily on benthic prey. These findings are in agreement with Corwins hypothesis (1978) that hearing sensitivity is correlated with feeding behavior. An examination of the macula neglecta of R. erinacea found a total count of 10 000 hair cells, which is within the range of other bottom-dwelling elasmobranchs.

ANALYSES OF SMALL TANK ACOUSTICS: EMPIRICAL AND THEORETICAL APPROACHES

Due to practical constraints, most of fish sound recordings are made in small tanks. Because of the boundary effects, serious distortion of recorded sound is expected (Parvulescu 1967). Thus there is a considerable need for a basis for interpreting the fish sound spectrum recorded in small tanks and providing practical ways of extracting useful information. Here we present the mechanism of spectrum distortion observed in small tanks both in empirical and theoretical ways. The minimum resonant frequency of small rectangular tanks with coordinates of X, Y, and Z, respectively, is provided by solving the wave equation (Kinsler et al 2000) as follows:

Acoustic intensity discrimination by the cichlid fish Astronotus ocellatus (Cuvier)

The acoustic intensity discrimination ability of the oscar (Astronotus ocellatus), a cichlid fish, was investigated using an automated positive reward method. Intensity discrimination thresholds (ΔI, in dB) for 7-s continuous pure tone signals were measured both as functions of sound intensity above thresholds, i.e., sensation levels, (SL)(+10 dB, +20 dB and +30 dB) and frequency (200 Hz, 500 Hz, and 800 Hz). ΔI at 500 Hz for +10 dB, +20 dB, and +30 dB SLs are 8.9, 5.5, and 3.3 dB, respectively. ΔI (at+20 dB SL) for 200 Hz, 500 Hz, and 800 Hz are 4.5, 5.5, and 9.3 dB, respectively. Despite having poor auditory sensitivity (narrow frequency range and high thresholds), the intensity discrimination ability of the oscar follows the general trends of previously studied fish species, however, with higher thresholds.

CORRELATION OF SOUND PRODUCTION WITH HEARING SENSITIVITY IN THE LAKE MALAWI CICHLID TRAMITICHROMIS INTERMEDIUS

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