Auditory chain reaction: Effects of sound pressure and particle motion on auditory structures in fishes.

Despite the diversity in fish auditory structures, it remains elusive how otolith morphology and swim bladder-inner ear (= otophysic) connections affect otolith motion and inner ear stimulation. A recent study visualized sound-induced otolith motion; but tank acoustics revealed a complex mixture of sound pressure and particle motion. To separate sound pressure and sound-induced particle motion, we constructed a transparent standing wave tube-like tank equipped with an inertial shaker at each end while using X-ray phase contrast imaging. Driving the shakers in phase resulted in maximised sound pressure at the tank centre, whereas particle motion was maximised when shakers were driven out of phase (180°). We studied the effects of two types of otophysic connections-i.e. the Weberian apparatus (Carassius auratus) and anterior swim bladder extensions contacting the inner ears (Etroplus canarensis)-on otolith motion when fish were subjected to a 200 Hz stimulus. Saccular otolith motion was more pronounced when the swim bladder walls oscillated under the maximised sound pressure condition. The otolith motion patterns mainly matched the orientation patterns of ciliary bundles on the sensory epithelia. Our setup enabled the characterization of the interplay between the auditory structures and provided first experimental evidence of how different types of otophysic connections affect otolith motion.

Sound pressure and particle acceleration audiograms in three marine fish species from the Adriatic Sea.

Fishes show great variability in hearing sensitivity, bandwidth, and the appropriate stimulus component for the inner ear (particle motion or pressure). Here, hearing sensitivities in three vocal marine species belonging to different families were described in terms of sound pressure and particle acceleration. In particular, hearing sensitivity to tone bursts of varying frequencies were measured in the red-mouthed goby Gobius cruentatus, the Mediterranean damselfish Chromis chromis, and the brown meagre Sciaena umbra using the non-invasive auditory evoked potential-recording technique. Hearing thresholds were measured in terms of sound pressure level and particle acceleration level in the three Cartesian directions using a newly developed miniature pressure-acceleration sensor. The brown meagre showed the broadest hearing range (up to 3000 Hz) and the best hearing sensitivity, both in terms of sound pressure and particle acceleration. The red-mouthed goby and the damselfish were less sensitive, with upper frequency limits of 700 and 600 Hz, respectively. The low auditory thresholds and the large hearing bandwidth of S. umbra indicate that sound pressure may play a role in S. umbra's hearing, even though pronounced connections between the swim bladder and the inner ears are lacking.

Potential mechanism of sound production in Oreochromis niloticus (Cichlidae).

Although acoustic communication is an integral part of cichlid behaviour, its mechanism has never been identified before. In the present study, a combination of approaches was used to investigate the sound-producing mechanism of Oreochromis niloticus. Synchronisation of high-speed video data (500 frames s(-1)) and cineradiographies (250 frames s(-1)) with the sound recordings made it possible to locate the different body parts involved in sound production in territorial males. Sounds are made during a backward movement of the pelvic and pectoral girdles and a forward movement of the second pterygiophore of the anal fin. Various electrostimulation experiments, dissections and observation of histological cross-sections revealed a set of bundles (that we call the vesica longitudinalis) situated in the hypaxial musculature, ventro-laterally to the swimbladder. Contraction of these bundles should result in compression of the rib cage and also of the swimbladder, because of its close association with the serosa and ribs. Deflation of the swimbladder resulted in a reduced sound intensity.

Acoustic pressure and particle motion thresholds in six sciaenid fishes.

Sciaenid fishes are important models of fish sound production, but investigations into their auditory abilities are limited to acoustic pressure measurements on five species. In this study, we used auditory brainstem response (ABR) to assess the pressure and particle acceleration thresholds of six sciaenid fishes commonly found in Chesapeake Bay, eastern USA: weakfish (Cynoscion regalis), spotted seatrout (Cynoscion nebulosus), Atlantic croaker (Micropogonias undulatus), red drum (Sciaenops ocellatus), spot (Leiostomus xanthurus) and northern kingfish (Menticirrhus saxatilis). Experimental subjects were presented with pure 10 ms tone bursts in 100 Hz steps from 100 Hz to 1.2 kHz using an airborne speaker. Sound stimuli, monitored with a hydrophone and geophone, contained both pressure and particle motion components. Sound pressure and particle acceleration thresholds varied significantly among species and between frequencies; audiograms were notably flatter for acceleration than pressure at low frequencies. Thresholds of species with diverticulae projecting anteriorly from their swim bladders (weakfish, spotted seatrout, and Atlantic croaker) were typically but not significantly lower than those of species lacking such projections (red drum, spot, northern kingfish). Sciaenids were most sensitive at low frequencies that overlap the peak frequencies of their vocalizations. Auditory thresholds of these species were used to estimate idealized propagation distances of sciaenid vocalizations in coastal and estuarine environments.

Electrical and mechanical properties and mode of innervation in scorpionfish sound-producing muscle fibres.

To obtain information about the neural mechanism underlying sound production in teleost fish, we studied the electrical and mechanical properties and mode of innervation in the swimbladder muscle (SBM) fibres of scorpionfish Sebastiscus marmoratus. Action potentials of the SBM fibres in response to direct electrical stimulation neither exhibited overshoot nor propagated along the fibre. Stimulation of the motor nerve, however, uniformly evoked action potentials along the fibre. When neuromuscular transmission was blocked by curare, motor nerve stimulation uniformly evoked endplate potentials along the fibre. These results indicate that action potentials propagate along the nerve branches but not along the SBM fibre membrane. In accordance with the above results, histochemical studies showed that motor nerve branches run along the SBM fibres to form many endplates with cholinesterase activity, indicating multiterminal innervation. The SBM consisted of about 600 fibres, while its motor nerve contained about 100 axons, giving an innervation ratio of about 1:6. Like mammalian fast muscle fibres, the SBM fibres exhibited a low succinic dehydrogenase activity and a high ATPase activity. These results are discussed in connection with the function of the SBM fibres in producing sound.

Acoustic communication in two freshwater gobies: the relationship between ambient noise, hearing thresholds and sound spectrum.

Two freshwater gobies Padogobius martensii and Gobius nigricans live in shallow (5-70 cm) stony streams, and males of both species produce courtship sounds. A previous study demonstrated high noise levels near waterfalls, a quiet window in the noise around 100 Hz at noisy locations, and extremely short-range propagation of noise and goby signals. To investigate the relationship of this acoustic environment to communication, we determined audiograms for both species and measured parameters of courtship sounds produced in the streams. We also deflated the swimbladder in P. martensii to determine its effect on frequency utilization in sound production and hearing. Both species are maximally sensitive at 100 Hz and produce low-frequency sounds with main energy from 70 to 100-150 Hz. Swimbladder deflation does not affect auditory threshold or dominant frequency of courtship sounds and has no or minor effects on sound amplitude. Therefore, both species utilize frequencies for hearing and sound production that fall within the low-frequency quiet region, and the equivalent relationship between auditory sensitivity and maximum ambient noise levels in both species further suggests that ambient noise shapes hearing sensitivity.

Genetic analysis of vertebrate sensory hair cell mechanosensation: the zebrafish circler mutants.

The molecular basis of sensory hair cell mechanotransduction is largely unknown. In order to identify genes that are essential for mechanosensory hair cell function, we characterized a group of recently isolated zebrafish motility mutants. These mutants are defective in balance and swim in circles but have no obvious morphological defects. We examined the mutants using calcium imaging of acoustic-vibrational and tactile escape responses, high resolution microscopy of sensory neuroepithelia in live larvae, and recordings of extracellular hair cell potentials (microphonics). Based on the analyses, we have identified several classes of genes. Mutations in sputnik and mariner affect hair bundle integrity. Mutant astronaut and cosmonaut hair cells have relatively normal microphonics and thus appear to affect events downstream of mechanotransduction. Mutant orbiter, mercury, and gemini larvae have normal hair cell morphology and yet do not respond to acoustic-vibrational stimuli. The microphonics of lateral line hair cells of orbiter, mercury, and gemini larvae are absent or strongly reduced. Therefore, these genes may encode components of the transduction apparatus.

Respiratory responses of domestic fowl to hyperthermia following selective air sac occlusions.

Ventilation together with blood and respiratory gas tensions were measured in adult domestic fowl under normothermic and hyperthermic conditions, following bilateral occlusion of the cranial and caudal thoracic air sacs (series I) or the cranial and caudal thoracic plus the abdominal air sacs (series II). Series I birds showed no significant differences from controls. Both control and experimental animals displayed a typical thermal polypnoea combined with mild hypocapnaemia. A larger drop in PCO2 was demonstrated in the clavicular sac than in the blood, possibly indicating partial failure of inspiratory valving at the ventrobronchi. However, there was no evidence of any effect of thoracic air sac occlusion on inspiratory airflow valving in the palaeopulmo. Series II birds were strongly hypercapnaemic/hypoxaemic in normothermic conditions, with a normal minute volume, but a faster, shallower breathing pattern. During hyperthermia they increased minute ventilation 3-fold, as in control animals, and blood gas tensions were almost restored to normal. Again, there was no evidence that experimental reduction in air sac capacity, in this case up to 70% of the total, had any effect on inspiratory airflow valving in the palaeopulmo, although inevitably in this case airflow in the neopulmo was abolished.

Responses of the swimbladder of the carp to sound stimulation.

The oscillations of the swimbladder anterior chamber of the carp (Cyprinus carpio) following stimulation with tones of 300-1500 Hz were studied by the method of holographic interferometry. The oscillation amplitude appeared to be maximal at frequencies close to the resonance frequency of an air bladder of equivalent volume as well as at frequencies corresponding approximately to the second and third harmonics of the resonance frequency. A change in the frequency of the sound signal or in the instantaneous pressure amplitude could result in spatial displacement of the oscillation centers on the swimbladder wall. The interference picture which resulted from recording the swimbladder oscillations over the tested frequency range was not observed on the holograms recorded within 20-24 h after the fish had been killed.

Hyperbaric oxygen: toxicity to fish at pressures present in their swimbladders.

When juvenile Pacific rock-fish, Sebastodes miniatus, are exposed to oxygen tensions equal to those in their swimbladders, they exhibit symptoms characteristic of oxygen poisoning in mammals and ultimately die. Thus their central nervous system appears to be as sensitive to elevated oxygen pressure as that of higher vertebrates, whereas the cells of the gas gland tissue inside the swimbladder must be insensitive to the partial pressure of oxygen which they help to produce.