RESUMEN
Mathematical modeling suggests that relatively large values of otolith mass asymmetry in fishes can alter acoustic functionality and may be responsible for abnormal fish behavior when subjected to weightlessness during parabolic or space flight [D.V. Lychakov, Y.T. Rebane, Otolith mass asymmetry in 18 species of fish and pigeon, J. Grav. Physiol. 11 (3) (2004) 17-34; D.V. Lychakov, Y.T. Rebane, Fish otolith mass asymmetry: morphometry and influence on acoustic functionality, Hear. Res. 201 (2005) 55-69]. The results of morphometric studies of otolith mass asymmetry suppose that the absolute value and the sign of the otolith mass asymmetry can change many times during the growth of individual fish within the range +/-20% [D.V. Lychakov, Y.T. Rebane, Otolith mass asymmetry in 18 species of fish and pigeon, J. Grav. Physiol. 11 (3) (2004) 17-34; D.V. Lychakov, Y.T. Rebane, Fish otolith mass asymmetry: morphometry and influence on acoustic functionality, Hear. Res. 201 (2005) 55-69]. This implies that the adverse effects of otolith asymmetry on acoustic and vestibular functionality could change during the lifetime of an individual fish. The aims of the present article were to examine the nature of otolith mass asymmetry fluctuation and to quantify otolith mass asymmetry in a large number of teleost fishes to verify our previous measurements. A dimensionless measure of otolith mass asymmetry, chi, was calculated as the difference between the masses of the right and left paired otoliths divided by average otolith mass. Saccular otolith mass asymmetry was studied in 59 Mediterranean teleost species (395 otolith pairs), 14 Black Sea teleost species (42 otolith pairs), red drum (196 otolith pairs) and guppy (30 otolith pairs). Utricular otolith mass asymmetry was studied in carp (103 otolith pairs) and goldfish (45 otolith pairs). In accordance with our previous results the value of chi did not depend on fish size (length or mass), systematic or ecological position of the fish, or otolith growth rate. In the great majority of the fishes studied, the saccular otolith chi was small /chi/ <0.05 (or <5%). Mathematical modeling indicates that values of chi vary among individual fish, but that the value is probably stable during a fish's lifetime.
Asunto(s)
Peces/anatomía & histología , Modelos Biológicos , Membrana Otolítica/anatomía & histología , Animales , Distribución de Chi-Cuadrado , Microscopía Electrónica de Rastreo , Membrana Otolítica/ultraestructura , Análisis de Regresión , Sáculo y Utrículo/anatomía & histologíaRESUMEN
The role of the fish otolith mass asymmetry in acoustic functionality is studied. The saccular, lagenar and utricular otoliths are weighted in two species of the Black Sea rays, 15 species of the Black Sea teleost fish and guppy fish. The dimensionless otolith mass asymmetry chi is calculated as ratio of the difference between masses of the right and left paired otoliths to average otolith mass. In the most fish studied the otolith mass asymmetry is within the range of -0.2 < chi < +0.2 (< 20%). We do not find specific fish species with extremely large or extremely small otolith asymmetry. The large otoliths do not belong solely to any particular side, left or right. The heavier otoliths of different otolithic organs can be located in different labyrinths. No relationship has been found between the magnitude of the otolith mass asymmetry and the length (mass, age) of the animal. The suggested fluctuation model of the otolith growth can interpret these results. The model supposes that the otolith growth rate varies slightly hither and thither during lifetime of the individual fish. Therefore, the sign of the relative otolith mass asymmetry can change several times in the process of the individual fish growth but within the range outlined above. Mathematical modeling shows that acoustic functionality (sensitivity, temporal processing, sound localization) of the fish can be disturbed by the otolith mass asymmetry. But this is valid only for the fish with largest otolith masses, characteristic of the bottom and littoral fish, and with highest otolith asymmetry. For most fish the values of otolith mass asymmetry is well below critical values. Thus, the most fish get around the troubles related to the otolith mass asymmetry. We suggest that a specific physicochemical mechanism of the paired otolith growth that maintains the otolith mass asymmetry at the lowest possible level should exist. However, the principle and details of this mechanism are still far from being understood.
Asunto(s)
Percepción Auditiva/fisiología , Peces/anatomía & histología , Membrana Otolítica/anatomía & histología , Animales , Peces/fisiología , Microscopía Electrónica de Rastreo/veterinaria , Modelos Biológicos , Membrana Otolítica/fisiología , Membrana Otolítica/ultraestructura , Poecilia/anatomía & histología , Poecilia/fisiología , Rajidae/anatomía & histología , Rajidae/fisiologíaRESUMEN
The masses and the area sizes of the otoliths for the utriculus, sacculus and lagena of 15 species of the Black Sea fish are analyzed. Morphometrical otolith regularities are derived and their functional and ecomorphological explanations are suggested. The otolith regularities are summarized in four otolith rules: (1) the masses of the otoliths gradually increase with the fish growth. (2) The mass ratio of the sacculus and utriculus or the sacculus and lagena otoliths does not change with the fish growth. (3) The ratio between the otolith area s and the otolith mass m is described by the exponential equation s=alpham(2/3). (4) The ratio between the otolith and macula sizes does not change with fish growth. Mathematical modeling of the otolith displacement responses to the acoustic and the instant force stimuli is performed. Based on the modeling the functional and ecomorphological explanations of the otolith regularities are suggested: (1) the greater the otolith mass, the higher the acoustic sensitivity at low frequencies and the sharper the frequency-response curve at its maximum. (2) The separation between maxima of the frequency-response curves for the saccular and lagenar otoliths remains virtually constant with the fish growth. (3) The bottom and littoral fish have better auditory capabilities than the pelagic fish. (4) The sensitivity to vestibular stimuli for greater otoliths is higher but the response is slower. The corresponding acceleration resolution for greater otoliths is higher and the range of accelerations in which the otolith organ can operate is narrower. (5) The relative vestibular sensitivities of the utriculus, sacculus and lagena otolith organs remain constant with fish growth. (6) The otolith organs of the bottom and littoral fish are tuned to different accelerations and possess different functional properties. The otolith organs of pelagic fish are adapted to a limited range of accelerations and are less sensitive to low accelerations as compared to the bottom and littoral fish.