Acoustic signalling in female fish: factors influencing sound characteristics in croaking gouramis

The characteristics of sounds produced by fishes are influenced by several factors such as size. The current study analyses factors affecting structural properties of acoustic signals produced by female croaking gouramis Trichopsis vittata during agonistic interactions. Female sounds (although seldom analysed separately from male sounds) can equally be used to investigate factors affecting the sound characteristics in fish. Sound structure, dominant frequency and sound pressure levels (SPL) were determined and correlated to body size and the order in which sounds were emitted. Croaking sounds consisted of series of single-pulsed or double-pulsed bursts, each burst produced by one pectoral fin. Main energies were concentrated between 1.3 and 1.5 kHz. The dominant frequency decreased with size, as did the percentage of single-pulsed bursts within croaking sounds. The SPL and the number of bursts within a sound were independent of size but decreased significantly with the order of their production. Thus, acoustic signals produced at the beginning of agonistic interactions were louder and consisted of more bursts than subsequent ones. Our data indicate that body size affects the dominant frequency and structure of sounds. The increase in the percentage of double-pulsed bursts with size may be due to stronger pectoral muscles in larger fish. In contrast, ongoing fights apparently result in muscle fatigue and subsequently in a decline in the number of bursts and SPL. The factor ‘order of sound production’ points to an intra-individual variability of sounds and should be considered in future studies.     

Bluegill Lepomis macrochirus synchronize pectoral fin motion and opercular pumping

The relative timing between operculum and pectoral fin motion was examined in swimming bluegill Lepomis macrochirus to determine if respiratory fluid flows from the operculum might have an effect on flow over the pectoral fin. Five bluegill were filmed swimming at speeds from 0·5 to 1·5 body (total) lengths s⁻¹. The timing of opercular pumping and pectoral fin beating was noted and analysed using circular statistics. Fish tended to ventilate their gills every second or third pectoral fin beat. While locomotion and ventilation had different frequencies, however, they were synchronized: fish maintained a consistent phase relationship between them. Thus, within pectoral fin beats when the operculum pumps, the jet consistently occurred during pectoral fin abduction, ending just after the fin was fully abducted and beginning adduction. Based on the distance between the opercular slit and the pectoral fin base, the jet was estimated to reach the fin during maximum abduction. Dye flow visualization confirmed this estimate, revealing that the opercular flow wraps around the base of the fin during peak abduction, when it is likely to have little hydrodynamic effect.     

Sound production and reproductive behaviour of the armoured catfish Corydoras paleatus (Callichthyidae)

Sound production during reproductive behaviour, dyadic encounters and distress situations was investigated in the callichthyid catfish Corydoras paleatus. Sounds were broad-band, pulsed, acoustic signals produced during abduction of the pectoral spines. Only males emitted trains of sounds during courting and trains of sounds of shorter duration during dyadic encounters. Several males, which are usually smaller than females, courted one gravid female without obvious cooperation or competition between them. During mating, one previously vocalizing male clasped the female's barbels with one pectoral spine and inseminated the eggs. The number of successful spawnings, days until spawning, and number of eggs laid was not related to the number of males (one, two or three) combined with one female. Males did not behave aggressively towards each other during courting or in dyadic encounters. In distress situations, when fish were hand held, both sexes and juveniles produced single sounds. The dominant frequency was negatively correlated with body size and the sound duration was positively correlated with relative length of pectoral spines (standardized to body length). This acoustical behaviour in C. paleatus differs considerably from Hoplosternum thoracatum, a representative of the callichthyine subfamily, in which vocalization was observed during territorial behaviour in males and aggressive behaviour in both sexes. This is the first report of a major difference in vocalizing behaviour within one teleost family.     

A Three-Dimensional Kinematics Analysis of a Koi Carp Pectoral Fin by Digital Image Processing

Pectoral fins fascinate researchers for their important role in fish maneuvers. By possessing a complicated flexible structure with several fin rays made by a thin film, the fin exhibits a three-dimensional (3D) motion. The complex 3D fin kinematics makes it challenging to study the performance of pectoral fin. Nevertheless, a detailed study on the 3D motion pattern of pectoral fins is necessary to the design and control of a bio-inspired fin rays. Therefore, a highspeed photography system is introduced in this paper to study the 3D motion of a Koi Carp by analyzing the two views of its pectoral fin simultaneously. The key motions of the pectoral fins are first captured in both hovering and retreating. Next, the 3D configuration of the pectoral fins is reconstructed by digital image processing, in which the movement of fin rays during fish retreating and hovering is obtained. Furthermore, the method of Singular Value Decomposition (SVD) is adopted to extract the basic motion patterns of pectoral fins from extensive image sequences, i.e. expansion, bending, cupping, and undulation. It is believed that the movement of the fin rays and the basic patterns of the pectoral fins obtained in the present work can provide a good foundation for the development and control of bionic flexible pectoral fins for underwater propeller.     

Significance of Agonistic Vocalization in the Croaking Gourami (Trichopsis vittatus,Teleostei)

Trichopsis vittatus emits high amplitude sounds during agonistic encounters with conspecifics. The sound producing organ is derived from the structural components of the pectoral fins. The study involved muting a sample of subjects by removing two pectoral fin tendons without any further restriction in movements and behaviour. Mute and unaltered males were then placed together in pairs and the following agonistic behavioural elements were determined: attacks, lateral displays, sound production and frontal displays. Soniferous males had a higher probability of winning contests when size differences were small. In pairs with big size ratios, the larger males were more often successful. Besides visual and acoustical signals, lateral line stimuli seem to play no role in threatening displays. These results demonstrate that vocalization during agonistic encounters is important for becoming dominant in specific circumstances. Because of correlation between main frequencies and size, sound emission might be a very effective method of assessing the physical strength of an opponent.     

Lycodes nishimurai, a new deep-sea eelpout (Teleostei: Zoarcidae) from the Sea of Japan

A new zoarcid fish, Lycodes nishimuari, is described on the basis of four specimens from the Yamato Bank, the Sea of Japan, at depths of 642–669 m. Although this species resembles Lycodes diapterus, L. hubbsi, L. pectoralis, L. macrochir, and L. nakamurae in having a deep notch in the pectoral fin and a single lateral line on the body, it is distinguished from them by the following combination of characters: no scales on pectoral fin base; scales on dorsum before dorsal fin origin; suborbital, preopercular, and last postorbital pores with risen rim; whitish lips; no whitish transverse bars on body; pectoral fin membrane and anterior part of dorsal fin membrane without black spots; basal color of vertical fins black. Supplementary material to this paper is available in electronic format at .     

西藏双须叶须鱼早期发育特征

通过双须叶须鱼(Ptychobarbus dipogon Regan)早期发育特征研究, 旨在为该鱼的科学保护和合理开发提供技术支撑。结果显示: 双须叶须鱼卵径3.7—3.9 mm, 吸水后的卵径可达5.1—5.3 mm。在水温10℃左右的条件下, 经历336.02h孵化出膜。根据胚胎的外部形态特征可将胚胎发育分为准备卵裂阶段、卵裂阶段、囊胚阶段、原肠阶段、神经胚阶段、器官分化阶段、孵化阶段共7个阶段34个时期。初孵仔鱼全长12.4 mm, 第1天体色素出现, 胸鳍上翘, 鳃盖骨出现, 下颌原基出现; 第2天鳃弓原基出现; 第3天消化道出现, 肝胰脏原基出现; 第4天鳃耙出现, 体表色素细胞带出现; 第5天口凹形成, 鳃丝形成; 第6天胸鳍褶, 背鳍褶, 腹鳍褶出现; 第7天鼻凹出现, 星芒状色素团出现; 第9天鳔前原基出现; 第11天尾鳍鳍条开始出现, 胸鳍开始颤动; 第13天鳔1室出现, 半规管形成; 第17天背鳍分化出来; 第21天腹部鳍褶变大, 舌颌骨清晰可见; 第28天脾脏出现; 第33天出现腹鳍鳍条; 第34天鳞片出现; 第85天稚鱼的形态与成鱼无异。双须叶须鱼是已报道裂腹鱼类卵径最大, 较四大家鱼卵周隙小, 是对高原隆起所导致的高寒自然环境的一种适应。     

Functional Morphology of Aquatic Flight in Fishes: Kinematics, Electromyography, and Mechanical Modeling of Labriform Locomotion

Labriform locomotion is the primary swimming mode for many fishesthat use the pectoral fins to generate thrust across a broadrange of speeds. A review of the literature on hydrodynamics,kinematics, and morphology of pectoral fin mechanisms in fishesreveals that we lack several kinds of morphological and kinematicdata that are critical for understanding thrust generation inthis mode, particularly at higher velocities. Several needsinclude detailed three-dimensional kinematic data on speciesthat are pectoral fin swimmers across a broad range of speeds,data on the motor patterns of pectoral fin muscles, and thedevelopment of a mechanical model of pectoral fin functionalmorphology. New data are presented here on pectoral fin locomotionin Gomphosus varius, a labrid fish that uses the pectoral finsat speeds of 1 –6 total body lengths per second. Three-dimensionalkinematic data for the pectoral fins of G. varius show thata typical "drag-based" mechanism is not used in this species.Instead, the thrust mechanics of this fish are dominated bylift forces and acceleration reaction forces. The fin is twistedlike a propeller during the fin stroke, so that angles of attackare variable along the fin length. Electromyographic data onsix fin muscles indicate the sequence of muscle activity thatproduces antagonistic fin abduction and adduction and controlsthe leading edge of the fin. EMG activity in abductors and adductorsis synchronous with the start of abduction and adduction, respectively,so that muscle mechanics actuate the fin with positive work.A mechanical model of the pectoral fin is proposed in whichfin morphometrics and computer simulations allow predictionsof fin kinematics in three dimensions. The transmission of forceand motion to the leading edge of the fin depends on the mechanicaladvantage of fin ray levers. An integrative program of researchis suggested that will synthesize data on morphology, physiology,kinematics, and hydrodynamics to understand the mechanics ofpectoral fin swimming.     

Outgrowth by fin motor axons in wildtype and a finless mutant of the Japanese medaka fish

The outgrowth of motor axons to the developing pectoral fin of the Japanese medaka fish (Oryzias latipes) was investigated both in wildtype embryos and in the pectoral finless (pl) mutants in which adults are missing pectoral fins. Late in embryogenesis the pectoral fin is a simple limb which contains two antagonist muscles which are innervated by presumptive motor neurons from the first four spinal segments (S1-4). The pectoral fin develops from a fin bud located in S1 and S2 centered on the border between S1 and S2 and, as with other limbs, one of the earliest signs of differentiation is the apical ectodermal ridge (AER). By the time the AER is well formed the growth cones of the presumptive motor neurons have reached the base of the fin bud and formed a plexus by extending toward the fin bud upon emergence from the spinal cord. This is especially evident on the ventral surface of the metamerically arranged axial muscles. For example, growth cones from S2 extend in a diagonal direction (both anterior and lateral) towards the fin bud. One hypothesis which can account for the pattern of motor outgrowth is that growth cones are attracted to the base of the fin bud, perhaps via a long distance cue. This hypothesis was tested by examining outgrowth of segmental nerves in pl embryos in which the fin buds arrest early in development following the initial appearance of the AER. In pl, nerves from S1-4 converged to form a plexus at the base of the abnormal fin bud, but the pattern of outgrowth varied from wildtype in a way consistent with a diminished capacity of the fin bud to attract segmental nerves to it.     

Acoustic and visual adaptations to predation risk: a predator affects communication in vocal female fish

Predation is an important ecological constraint that influences communication in animals. Fish respond to predators by adjusting their visual signaling behavior, but the responses in calling behavior in the presence of a visually detected predator are largely unknown. We hypothesize that fish will reduce visual and acoustic signaling including sound levels and avoid escalating fights in the presence of a predator. To test this we investigated dyadic contests in female croaking gouramis (Trichopsis vittata, Osphronemidae) in the presence and absence of a predator (Astronotus ocellatus, Cichlidae) in an adjoining tank. Agonistic behavior in T. vittata consists of lateral (visual) displays, antiparallel circling, and production of croaking sounds and may escalate to frontal displays. We analyzed the number and duration of lateral display bouts, the number, duration, sound pressure level, and dominant frequency of croaking sounds as well as contest outcomes. The number and duration of lateral displays decreased significantly in predator when compared with no-predator trials. Total number of sounds per contest dropped in parallel but no significant changes were observed in sound characteristics. In the presence of a predator, dyadic contests were decided or terminated during lateral displays and never escalated to frontal displays. The gouramis showed approaching behavior toward the predator between lateral displays. This is the first study supporting the hypothesis that predators reduce visual and acoustic signaling in a vocal fish. Sound properties, in contrast, did not change. Decreased signaling and the lack of escalating contests reduce the fish’s conspicuousness and thus predation threat.     

Externally attached radio transmitters do not affect the parental care behaviour of rock bass

Turning, pectoral fin and caudal fin rates and time spent on the nest of male rock bass Ambloplites rupestris , engaged in parental care, were not affected after the attachment of external radio transmitters. Reproductive success was similar between treatment and control fish. Micro external radio transmitters can be used on small fishes for studying parental care duration and post-care movement without altering their behaviour.     

Normal growth of the "see-through" medaka

The see-through stock in the medaka Oryzias latipes, causes pigments to be absent from the whole body and has a transparent body in the adult stage as well as during embryonic stages. To establish a standard table of growth stages for this model fish, morphological features were examined during the growing period from hatching to adulthood. The main observations were performed on morphological changes in external and internal organs that could be seen through the body wall of the living fish during growth. Finally, five growth stages from just after hatching to the adult stage were defined on the basis of synchronized or definite changes in morphology as follows: (1) stage 40 in which the nodes (joints) in bony rays of the caudal and pectoral fins first appear, (2) the stage 41 in which the ribs and the anal, dorsal and ventral fins are formed by degeneration of the membrane fin folds, as recognized by the first appearance of nodes in the fin rays of the anal, pectoral and dorsal fins, and the parallel distribution of the dorsal artery and ventral vein of the tail, (3) stage 42 in which the 2-spiral pattern of the gut, the ray nodes in the ventral fins, and the scales first appear, (4) stage 43 in which early secondary sexual characters such as urinogenital protruberances (female) and papillar processes (male) appear, (5) stage 44 in which the 3-spiral pattern of the gut and the papillar process on the 2nd ray of pectral fins (male) appear.     

Functional morphology of the pectoral fins in bamboo sharks, Chiloscyllium plagiosum: benthic vs. pelagic station-holding

Bamboo sharks (Chiloscyllium plagiosum) are primarily benthic and use their relatively flexible pectoral and pelvic fins to rest on and move about the substrate. We examined the morphology of the pectoral fins and investigated their locomotory function to determine if pectoral fin function during both benthic station-holding and pelagic swimming differs from fin function described previously in leopard sharks, Triakis semifasciata. We used three-dimensional kinematics and digital particle image velocimetry (DPIV) to quantify pectoral fin function in five white-spotted bamboo sharks, C. plagiosum, during four behaviors: holding station on the substrate, steady horizontal swimming, and rising and sinking during swimming. During benthic station-holding in current flow, bamboo sharks decrease body angle and adjust pectoral fin angle to shed a clockwise fluid vortex. This vortex generates negative lift more than eight times that produced during open water vertical maneuvering and also results in an upstream flow that pushes against the posterior surface of the pectoral fin to oppose drag. In contrast, there is no evidence of significant lift force in the wake of the pectoral fin during steady horizontal swimming. The pectoral fin is held concave downward and at a negative dihedral angle during steady horizontal swimming, promoting maneuverability rather than stability, although this negative dihedral angle is much less than that observed previously in sturgeon and leopard sharks. During sinking, the pectoral fins are held concave upward and shed a clockwise vortex with a negative lift force, while in rising the pectoral fin is held concave downward and sheds a counterclockwise vortex with a positive lift force. Bamboo sharks appear to sacrifice maneuverability for stability when locomoting in the water column and use their relatively flexible fins to generate strong negative lift forces when holding position on the substrate and to enhance stability when swimming in the water column.     

Fish fingers: digit homologues in sarcopterygian fish fins

A defining feature of tetrapod evolutionary origins is the transition from fish fins to tetrapod limbs. A major change during this transition is the appearance of the autopod (hands, feet), which comprises two distinct regions, the wrist/ankle and the digits. When the autopod first appeared in Late Devonian fossil tetrapods, it was incomplete: digits evolved before the full complement of wrist/ankle bones. Early tetrapod wrists/ankles, including those with a full complement of bones, also show a sharp pattern discontinuity between proximal elements and distal elements. This suggests the presence of a discontinuity in the proximal-distal sequence of development. Such a discontinuity occurs in living urodeles, where digits form before completion of the wrist/ankle, implying developmental independence of the digits from wrist/ankle elements. We have observed comparable independent development of pectoral fin radials in the lungfish Neoceratodus (Osteichthyes: Sarcopterygii), relative to homologues of the tetrapod limb and proximal wrist elements in the main fin axis. Moreover, in the Neoceratodus fin, expression of Hoxd13 closely matches late expression patterns observed in the tetrapod autopod. This evidence suggests that Neoceratodus fin radials and tetrapod digits may be patterned by shared mechanisms distinct from those patterning the proximal fin/limb elements, and in that sense are homologous. The presence of independently developing radials in the distal part of the pectoral (and pelvic) fin may be a general feature of the Sarcopterygii.     

Sexual dimorphism of the Weberian apparatus and pectoral girdle in Sundadanio axelrodi (Ostariophysi: Cyprinidae)

The miniature cyprinid fish, Sundadanio axelrodi , exhibits extreme sexual dimorphism in the skeleton of the Weberian apparatus, the fifth rib and pectoral girdle. Musculature associated with the fifth rib and Weberian apparatus also shows a high degree of sexual dimorphism. It is suggested that these modifications are responsible for the production of a croaking sound that seems to be restricted to males of the species, based on the lack of any corresponding anatomical specializations in females.     

Sound production by the river bullhead, Cottus gobio L. (Cottidae, Teleostei)

Acoustic signals of Cottus gobio consist of knocking sounds produced as single pulses (48 ms) or as trains of 4–6 pulses (230 ms). Frequencies extend up to 3 kHz, but most sound energy is concentrated between 50 and 500 Hz in both sound types. Cottus gobio is solitary, maintains territories, and defends them by threat display, seldom by biting and fighting. Threatening consists of spreading gill covers and fins, darkening, and sound production. Calling is accompanied by a nodding movement of the head, during which the pectoral girdle and the skull are moved rapidly against each other. No difference in ability of sound production was observed between sexes, but males emitted significantly more sounds than females. In the laboratory an increase in vocalization activity was observed between night and day. An increase in the number of encounters and calls was noted when temperatures were raised from 8°C to 13°C. Sound production was registered throughout the year.     

Fin duplications and deletions induced by disruption of retinoic acid signaling

 Retinoic acid (RA), a derivative of vitamin A, plays a critical role as a signaling molecule in axial patterning of vertebrates. Here we report that RA exposure of zebrafish (Danio rerio) and mummichog (Fundulus heteroclitus) embryos during gastrulation results in homeotic duplications of the pectoral fins in up to 94% of fish. We have observed three to four pairs of fins in an individual fish. Although some duplications are partial, many represent complete axial duplications of the pectoral girdle and fin and include coracoscapulae, proximal radials, and dermal fin elements. Fin duplications are observed only at a defined dose of RA. Inhibition of RA synthesis by exposure to citral during a narrow developmental window leads to fish which lack pectoral fins but can be rescued by addition of exogenous RA, suggesting that RA signaling is critical to fin specification during early development. The ability to consistently induce multiple fins in a large number of vertebrate embryos should contribute to the understanding of genetic regulation of the normal positioning of limbs during embryogenesis. Received: 30 August 1997 / Accepted: 6 December 1997     

Comparison of ageing methods and growth rates for largemouth bass,Micropterus salmoides Lacépède,from northern latitudes

Synopsis Ages determined by counts of apparent annuli on scales, sagittae, vertebrae, pectoral fin ray and dorsal fin spine cross sections of largemouth bass from northern populations, which are older and slower growing fish than in the southern parts of its native range, were compared to establish the accuracy of each method. Linear regression techniques indicated strong agreement (r> 0.9) among ages assigned from the examination of scales, sagittae, and vertebrae. The pattern of growth zones on pectoral fin ray and dorsal fm spine cross sections proved too variable for accurate age determination. Limited data suggest that ages greater than 7+ assigned from scales were more likely to underestimate true ages than the other body parts used, although none of these methods gave satisfactory results. Examination of scales from recovered tagged fish, and the similarity between back-calculated lengths of fish through age 7+ to annulus I and observed lengths of juvenile largemouth bass near the end of their first growing season, support the validity of ages determined from scales. Despite a very limited amount of habitat suitable for largemouth bass and severe climatic conditions, growth of this species in Tadenac Lake was similar to growth in other waters north of the Great Lakes. Differences in physical characteristics among these waters does not appear to influence growth rates of largemouth bass, but probably affects production and biomass.     

Agonistic behaviour of age 0, age 1 and non-breeding adult Arctic grayling Thymallus arcticus (Pallas)

Age 0 Arctic grayling begin to show agonistic behaviour 3 weeks after emergence and are territorial by their fourth week. Age 1 fish are territorial throughout the summer feeding period. In adults, territoriality is restricted to larger males during the early spring breeding season while after the breeding season both smaller adult males and adult females also hold territories.
In sub-adults, larger fish and resident fish tend to win most fights. In adults, males usually win against females. In fights between adults of the same sex, larger and resident fish usually win.
The lateral display becomes more important in the agonistic repertoire of Arctic grayling as they mature and the characteristic large dorsal fin develops. Arctic grayling appear to lack an appeasement display.