Effects of temperature change on cortisol release by common carp Cyprinus carpio.

Common carp Cyprinus carpio, stressed by fish handling practices, responded with a decrease in cortisol secretion when temperature was lowered from 20 to 14° C within 3·5 h compared to those kept at 20° C.

How do albino fish hear?

Pigmentation disorders such as albinism are occasionally associated with hearing impairments in mammals. Therefore, we wanted to investigate whether such a phenomenon also exists in non-mammalian vertebrates. We measured the hearing abilities of normally pigmented and albinotic specimens of two catfish species, the European wels Silurus glanis (Siluridae) and the South American bronze catfish Corydoras aeneus (Callichthyidae). The non-invasive auditory evoked potential (AEP) recording technique was utilized to determine hearing thresholds at 10 frequencies from 0.05 to 5 kHz. Neither auditory sensitivity nor shape of AEP waveforms differed between normally pigmented and albinotic specimens at any frequency tested in both species. Silurus glanis and C. aeneus showed the best hearing between 0.3 and 1 kHz; the lowest thresholds were 78.4 dB at 0.5 kHz in S. glanis (pigmented), 75 dB at 1 kHz in S. glanis (albinotic), 77.6 dB at 0.5 kHz in C. aeneus (pigmented) and 76.9 dB at 1 kHz in C. aeneus (albinotic). This study indicates no association between albinism and hearing ability. Perhaps because of the lack of melanin in the fish inner ear, hearing in fishes is less likely to be affected by albinism than in mammals.

Sound-generating and -detecting motor system in catfish: design of swimbladder muscles in doradids and pimelodids.

Catfishes have evolved a diversity of swimbladder muscles serving in the generation of different sounds and probably other acoustic functions. In order to find out if anatomical and acoustical differences are parallelled by fine structural differences, I examined the sonic muscles of the doradid Platydoras and the pimelodid Pimelodus by gross dissections and ultrastructural methods. In Platydoras, the sound-generating (drumming) muscle (DM) inserts on a dorsal bony plate that vibrates the swimbladder. In pimelodids, the large DM attaches directly on the ventral surface of the swimbladder, whereas the small tensor tripodis muscle (TT) inserts on the rostral surface near the tripus, the most caudal Weberian ossicle. Fibers of all three muscles possess an extensive development of sarcoplasmatic reticulum (SR) in association with very thin myofibrils (MF) but differed widely in their arrangement. In Platydoras, ribbons of MFs are arranged radially around a central core. Mitochondria were found within the core and the peripheral sarcoplasm. Pimelodus does not have a differentiated core and the cross-sectional area of DM-MFs is about 15% larger as determined by stereological measurements. The TT possesses shorter sarcomeres and more mitochondria than DMs, which were primarily found between MFs. This suggests faster contraction properties and greater resistance to fatigue compared with sonic muscles. Data indicate that the higher amount of DM-myofibrils in pimelodids might result in stronger muscle contractions and, presumably, in higher sound intensities. The fine structure of the TT reveals that contractions most likely prevent transmission of swimbladder vibrations to the inner ear via the Weberian ossicles during vocalization.

Comparison of the inner ear ultrastructure between teleost fishes using different channels for communication.

The anatomy and ultrastructure of the inner ear of three species of gouramis which differ widely in acoustic behavior were studied using scanning electron microscopy. Of the three species. Trichopsis possess a pectoral sound-producing mechanism while Macropodus and Betta lack sonic organs. The general structure of the inner ear and the shapes of the sensory epithelia are very similar, although they do differ on the posterior part of the saccular macula which is more S-shaped in Trichopsis and Macropodus than in Betta. The maculae on the three species do not differ either in ciliary bundle type (cells with long kinocilia on the periphery of the maculae and cells with short kinocilia in the central region) or in hair cell orientation pattern. Quantitative measurements of hair cell densities and the size of the sensory epithelia of the saccule did not show significant differences between species. Data presented correlate with physiological results from other investigators showing similar auditory sensitivity in Trichopsis and Macropodus. The similarity in structure and function of the inner ears of gouramis on one hand, and the occurrence of sound-generating organs in just one genus, suggests that hearing evolved prior to vocalization and thus acoustic communication in this taxon.

The ontogenetic development of auditory sensitivity, vocalization and acoustic communication in the labyrinth fish Trichopsis vittata.

The anabantoid fish Trichopsis vittata starts vocalizing as 8-week-old juveniles. In order to determine whether juveniles are able to detect conspecific sounds, hearing sensitivities were measured in six size groups utilizing the auditory brainstem response-recording technique. Results were compared to sound pressure levels and spectra of sounds recorded during fighting. Auditory evoked potentials were present in all size groups and complete audiograms were obtained starting with 0.18 to 0.30 g juveniles. Auditory sensitivity during development primarily increased between 0.8 kHz and 3.0 kHz. The most sensitive frequency within this range shifted from 2.5 kHz to 1.5 kHz, whereas thresholds decreased by 14 dB. Sound production, on the other hand, started at 0.1 g and sound power spectra at dominant frequencies increased by 43 dB, while dominant frequencies shifted from 3 kHz to 1.5 kHz. Comparisons between audiograms and sound power spectra in similar-sized juveniles revealed no clear match between most sensitive frequencies and dominant frequencies of sounds. This also revealed that juveniles cannot detect conspecific sounds below the 0.31 to 0.65 g size class. These results indicate that auditory sensitivity develops prior to the ability to vocalize and that vocalization occurs prior to the ability to communicate acoustically.

Acoustic communication and the evolution of hearing in fishes.

Fishes have evolved a diversity of sound-generating organs and acoustic signals of various temporal and spectral content. Additionally, representatives of many teleost families such as otophysines, anabantoids, mormyrids and holocentrids possess accessory structures that enhance hearing abilities by acoustically coupling air-filled cavities to the inner ear. Contrary to the accessory hearing structures such as Weberian ossicles in otophysines and suprabranchial chambers in anabantoids, sonic organs do not occur in all members of these taxa. Comparison of audiograms among nine representatives of seven otophysan families from four orders revealed major differences in auditory sensitivity, especially at higher frequencies (> 1 kHz) where thresholds differed by up to 50 dB. These differences showed no apparent correspondence to the ability to produce sounds (vocal versus non-vocal species) or to the spectral content of species-specific sounds. In anabantoids, the lowest auditory thresholds were found in the blue gourami Trichogaster trichopterus, a species not thought to be vocal. Dominant frequencies of sounds corresponded with optimal hearing bandwidth in two out of three vocalizing species. Based on these results, it is concluded that the selective pressures involved in the evolution of accessory hearing structures and in the design of vocal signals were other than those serving to optimize acoustic communication.

Auditory role of lateral trunk channels in cobitid fishes.

In cobitid fishes the anterior part of the swimbladder is encapsulated by bone to varying extent. This might diminish the auditory sensitivity of these otophysine fishes by reducing the vibrations of the swimbladder wall in the sound field. However, according to prior studies the auditory thresholds of the cobitid Botia modesta is similar to that of other otophysine fishes. According to anatomical investigation B. modesta has a cranial encapsulation of the anterior part of the swimbladder (camera aerea Weberiana) as expected and in addition special channels stretching laterally from the swimbladder to the outer body wall. These lateral trunk channels are filled with fat and lymph. They form a muscle-free acoustic window beneath the skin, which could be demonstrated by measuring the auditory brainstem response at 400 Hz, 800 Hz, 1500 Hz, and 3000 Hz. Filling the lateral trunk channels with wettex (cotton/rayon staple) resulted in an increase of the auditory thresholds by 13.6-17.6 dB, indicating mechanical damping of the swimbladder. Our experiments demonstrate that the intact lateral trunk channels enhance the hearing sensitivity of cobitid fishes.

Did auditory sensitivity and vocalization evolve independently in otophysan fishes?

Otophysine fishes have a series of bones, the Weberian ossicles, which acoustically couple the swimbladder to the inner ear. These fishes have evolved a diversity of sound-generating organs and acoustic signals, although some species, such as the goldfish, are not known to be vocal. Utilizing a recently developed auditory brainstem response (ABR)-recording technique, the auditory sensitivities of representatives of seven families from all four otophysine orders were investigated and compared to the spectral content of their vocalizations. All species examined detect tone bursts from 100 Hz to 5 kHz, but ABR-audiograms revealed major differences in auditory sensitivities, especially at higher frequencies (>1 kHz) where thresholds differed by up to 50 dB. These differences showed no apparent correspondence to the ability to produce sounds (vocal versus non-vocal species) or to the spectral content of species-specific sounds. All fishes have maximum sensitivity between 400 Hz and 1,500 Hz, whereas the major portion of the energy of acoustic signals was in the frequency range of 100-400 Hz (swimbladder drumming sounds) and of 1-3 kHz (stridulatory sounds). Species producing stridulatory sounds exhibited better high-frequency hearing sensitivity (pimelodids, doradids), except for callichthyids, which had poorest hearing ability in this range. Furthermore, fishes emitting both low- and high-frequency sounds, such as pimelodid and doradid catfishes, did not possess two corresponding auditory sensitivity maxima. Based on these results it is concluded that selective pressures involved in the evolution of the Weberian apparatus and the design of vocal signals in otophysines were others (primarily predator or prey detection in quiet freshwater habitats) than those serving to optimize acoustical communication.

Sonic/vocal motor pathways in catfishes: comparisons with other teleosts.

Among teleost fishes, representatives of several distantly related groups have sound-producing (sonic/vocal) muscles associated with the swimbladder or pectoral girdle/fin. Here, the diversity of vocal organs and central motor pathways in four families of catfish, order Siluriformes, is compared to that in families from two distantly related orders, the Scorpaeniformes and Batrachoidiformes. Several catfish families have two sonic mechanisms--a swimbladder vibration established by 'drumming muscles' that differ in origin and insertion between families, and a pectoral spine stridulatory apparatus. In ariids, mochokids and doradids, sonic swimbladder muscles originate at various cranial or postcranial elements and insert onto an 'elastic spring' that vibrates the swimbladder, while in pimelodids the muscles insert ventrally at the swimbladder. Sonic motoneurons are located along the midline, ventral to the fourth ventricle/central canal in doradids and mochokids but lateral to the medial longitudinal fasciculus in ariids; pimelodids have motoneurons in both locations. The axonal trajectory for the lateral motoneurons in pimelodids and ariids implies that they are a migrated, midline population of sonic motoneurons. Pectoral spine-associated motoneurons are located in the ventral motor column. Unlike catfishes, a diversity of sonic mechanisms in Scorpaeniformes is not associated with different positions for sonic motoneurons. Cottids (sculpin) lack a swimbladder but have sonic muscles that originate at the occipital cranium and insert at the pectoral girdle; sonic motoneurons are located within the ventral motor column. Some triglids have intrinsic swimbladder muscles, although ontogenetic data indicate a transient association with the pectoral girdle; sonic motoneurons are in the same location as in cottids. Among Batrachoidiformes, all known representatives have intrinsic swimbladder muscles that are never associated with the pectoral girdle and are innervated by midline sonic motoneurons. The results suggest two patterns of organization for sound-producing systems in teleost fishes: pectoral fin/girdle-associated muscles are innervated by sonic motoneurons positioned within the ventral motor column, adjacent to the ventral fasciculus; non-pectoral associated muscles are innervated by sonic motoneurons located on or close to the midline, adjacent to the medial longitudinal fasciculus.

Correlation between auditory sensitivity and vocalization in anabantoid fishes.

Several anabantoid species produce broadband 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 I microPa 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 airbreathing chamber, which, lying close to the hearing and sonic organs, enhances both sound perception and emission at its resonant frequency.

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.

Sonic/vocal-acousticolateralis pathways in teleost fishes: a transneuronal biocytin study in mochokid catfish.

Mochokid catfish have two sound producing (sonic) organs--a pectoral spine stridulatory apparatus and a swimbladder whose vibration is established by nearby "drumming" muscles. Dextran-biotin or biocytin application to sonic nerves or muscles identified topographically separated motoneuron pools. Pectoral spine-related motoneurons are located within the ventral motor column whereas swimbladder motoneurons lie just ventral to the central canal or fourth ventricle. Axons of both groups of motoneurons exit the brain and spinal cord via ventral roots of occipital (swimbladder and pectoral) and spinal (swimbladder only) nerves. Transneuronal biocytin transport identified an extensive premotor network only for the swimbladder motor nuclei. Premotoneuron somata are located ipsilaterally in 1) a dorsolateral region of the sonic motor nucleus (SMN); motoneurons were clustered in the ventromedial region of the SMN and 2) the ventromedial medulla at the rostral pole of the SMN. Biocytin-filled fibers and less frequently premotoneuron somata were also found in the contralateral SMN. Biocytin-labeled fibers were continuous farther rostrally with 1) a commissural bundle that terminated bilaterally in the medial reticular formation near the caudal pole of the descending octaval nucleus and 2) a lateral brainstem bundle that terminated ipsilaterally in regions of the medulla and cerebellum considered to subserve acoustic and lateral line functions. Together with other data in distantly related teleost fishes, the results support the hypotheses that 1) central pathways linking sound-generating (sonic or vocal) and acoustic regions of the brain are traits common to both teleosts fishes and tetrapods that actively generate sounds, and 2) sonic/vocal pathways in teleosts have a conserved pattern of organization suggestive of common developmental origins.

Localization of swimbladder and pectoral motoneurons involved in sound production in pimelodid catfish.

We localized the motoneurons and occipital and true spinal innervation of sound-producing organs in pimelodid catfish. Pimelodids have a stridulatory organ composed of the pectoral girdle and the first pectoral fin ray, a swimbladder with extrinsic muscles to produce drumming sounds, and a tensor tripodis (TT) muscle that inserts on the swimbladder. Sonic muscles are innervated by three branches (rostral, dorsal and caudal) of the occipital nerve (Oc) and the first two true spinal nerves (S1 and 2): pectoral spine muscles (abductor, adductor and ventral rotator) by the rostral branch of Oc and S1 and 2, drumming muscle by the caudal branch of Oc and twigs of the S1 and 2, and TT by the dorsal branch of Oc. Sonic nuclei from ipsilateral medial, intermediate and ventrolateral columns in the caudal medulla and spinal cord. Pectoral neurons form a ventrolateral motor column, and neurons for the first spine occupy the rostral part of the column. The medial division of the swimbladder drumming motor nucleus (DMm) is situated on the midline between the central canal and the medial longitudinal fasciculus. The rostral pole of the DM nucleus expands ventrolaterally to include a population of neurons of intermediate position (DMi). The TT nucleus also assumes an intermediate position ventrolateral to DMm. The pectoral, TT, and DMi have a restricted rostrocaudal extent, whereas DMm extends further caudally. These data demonstrate that fish can evolve multiple sonic motor nuclei and that sound producing organs can be innervated in parallel by occipital and spinal nerves.

Localization of pectoral fin motoneurons (sonic and hovering) in the croaking gourami Trichopsis vittatus.

The pectoral fin of the croaking gourami, Trichopsis vittatus, has become modified as a sound-producing organ and retains its original function in locomotion and hovering. We used retrograde transport of horseradish peroxidase to localize sonic motoneurons in Trichopsis. Betta splendens, a related nonsonic gourami with unspecialized pectoral fins, served as a control. A single injection into Trichopsis epaxial muscle labeled a dorsal motor column of large cells (mean of 16.3 microns) ventrolateral to the central canal. Pectoral motoneurons formed a ventrolateral spinal motor column of smaller neurons (means from 7.7 to 11.9 microns, depending upon fish size), of about 2 mm in rostrocaudal extent, starting in the caudal medulla. Our data suggest that motoneurons for different pectoral muscles are segregated into rostrocaudal pools within the column. Distribution, morphology and size of motoneurons were similar between Trichopsis and Betta, and there was no evidence of a distinct population of neurons which might be specialized exclusively for sound production. These data suggest that a fish can evolve a specialized end organ without major reorganization of the central nervous system.