Anabantoid Fish Research Articles

Expansion and Functional Diversification of Long-Wavelength-Sensitive Opsin in Anabantoid Fishes

Gene duplication is one of the most important sources of novel genotypic diversity and the subsequent evolution of phenotypic diversity. Determining the evolutionary history and functional changes of duplicated genes is crucial for a comprehensive understanding of adaptive evolution. The evolutionary history of visual opsin genes is very dynamic, with repeated duplication events followed by sub- or neofunctionalization. While duplication of the green-sensitive opsins rh2 is common in teleost fish, fewer cases of multiple duplication events of the red-sensitive opsin lws are known. In this study, we investigate the visual opsin gene repertoire of the anabantoid fishes, focusing on the five lws opsin genes found in the genus Betta. We determine the evolutionary history of the lws opsin gene by taking advantage of whole-genome sequences of nine anabantoid species, including the newly assembled genome of Betta imbellis. Our results show that at least two independent duplications of lws occurred in the Betta lineage. The analysis of amino acid sequences of the lws paralogs of Betta revealed high levels of diversification in four of the seven transmembrane regions of the lws protein. Amino acid substitutions at two key-tuning sites are predicted to lead to differentiation of absorption maxima (λmax) between the paralogs within Betta. Finally, eye transcriptomics of B. splendens at different developmental stages revealed expression shifts between paralogs for all cone opsin classes. The lws genes are expressed according to their relative position in the lws opsin cluster throughout ontogeny. We conclude that temporal collinearity of lws expression might have facilitated subfunctionalization of lws in Betta and teleost opsins in general.

Association of predation risk with a heterospecific vocalization by an anabantoid fish.

Honey gouramis (Trichogaster chuna) received chemical alarm cues derived from conspecific epidermal tissue and, simultaneously, the vocalization produced by a heterospecific gourami species, the sparkling pygmy gourami (Trichopsis pumila). Control trials paired water with the vocalization. In trials that received alarm cues, honey gouramis significantly increased activity relative to control trials that received water, suggesting an attempt to flee and search for refuge. When the recording of the vocalization was later replayed to test fish without any additional chemical cue, fish that had previously experienced the alarm cue froze while those that had received water with the vocalization did not change their behaviour. These data indicate that honey gouramis recognize and respond to chemical alarm cues, making this report the second anabantoid species to be recorded with this response. Second, these data indicate that honey gouramis can associate risk of predation with a novel auditory stimulus, including vocalizations from other species. These data suggest the potential for vocalizations to evolve into alarm signals in this group of fishes.

Histological Study of Suprabranchial Chamber Membranes in Anabantoidei and Clariidae Fishes.

Simple SummaryAir-breathing fish constitute a broad evolutionary group of fish, which are generally characterized by distinctive phenotypical plasticity. These fishes usually inhabit waters where oxygen deficiency occurs periodically, which is why they have developed a variety of accessory respiratory organs (AROs) that may be used in an obligatory or a facultative manner. Knowledge of the structure of these organs is important for both the breeding and the conservation of these fish species. The aim of this study was to conduct a comparative histological analysis of two types of AROs found in the Anabantoidei suborder and the Clariidae family, both of which are freshwater fish taxa of high ecological and commercial importance.Accessory respiratory organs (AROs) are a group of anatomical structures found in fish, which support the gills and skin in the process of oxygen uptake. AROs are found in many fish taxa and differ significantly, but in the suborder Anabantoidei, which has a labyrinth organ (LO), and the family Clariidae, which has a dendritic organ (DO), these structures are found in the suprabranchial cavity (SBC). In this study, the SBC walls, AROs, and gills were studied in anabantoid (Betta splendens, Ctenopoma acutirostre, Helostoma temminckii) and clariid (Clarias angolensis, Clarias batrachus) fishes. The histological structure of the investigated organs was partially similar, especially in relation to their connective tissue core; however, there were noticeable differences in the epithelial layer. There were no significant species-specific differences in the structure of the AROs within the two taxa, but the SBC walls had diversified structures, depending on the observed location. The observed differences between species suggest that the remarkable physiological and morphological plasticity of the five investigated species can be associated with structural variety within their AROs. Furthermore, based on the observed histology of the SBC walls, it is reasonable to conclude that this structure participates in the process of gas exchange, not only in clariid fish but also in anabantoids.

Parental Care in the Climbing Perch (Anabas testudineus): Confusion or Lost Data?

Anabantoid fishes comprise a unique group, most of whose representatives are characterized by diverse forms of parental care. Evolutional reconstruction of the reproductive strategies of anabantoids is challenging, partly due to the lack of reliable evidence of parental care in a number of its representatives, the most famous of which is the climbing perch. Moreover, apparent contradictions between literary sources frequently occur on the subject. This brief overview attempts to analyze the situation and identify the possible causes of disagreement. The further development in understanding the phylogeny of life history strategies in the group, as well as the concept of evolutionary transitions among parental care patterns in fishes in general, depend on the clarification of this situation.

Hypoxia‐induced developmental plasticity of larval growth, gill and labyrinth organ morphometrics in two anabantoid fish: The facultative air‐breather Siamese fighting fish (Betta splendens) and the obligate air‐breather the blue gourami (Trichopodus trichopterus)

Larval and juvenile air breathing fish may experience nocturnal and/or seasonal aquatic hypoxia. Yet, whether hypoxia induces respiratory developmental plasticity in larval air breathing fish is uncertain. This study predicted that larvae of two closely related anabantid fish-the facultative air breather the Siamese fighting fish (Betta splendens) and the obligate air breathing blue gourami (Trichopodus trichopterus)-show distinct differences in developmental changes in body, gill, and labyrinth morphology because of their differences in levels of dependency upon air breathing and habitat. Larval populations of both species were reared in normoxia or chronic nocturnal hypoxia from hatching through 35-38 days postfertilization. Gill and labyrinth variables were measured at the onset of air breathing. Betta splendens reared in normoxia possessed larger, more developed gills (~3× greater area) than T. trichopterus at comparable stages. Surface area of the emerging labyrinth, the air breathing organ, was ~ 85% larger in normoxic B. splendens compared to T. trichopterus. Rearing in mild hypoxia stimulated body growth in B. splendens, but neither mild nor severe hypoxia affected growth in T. trichopterus. Condition factor, K (~ 1.3 in B. splendens, 0.7 in T. trichopterus) was unaffected by mild hypoxia in either species, but was reduced by severe hypoxia to <0.9 only in B. splendens. Severe, but not mild, hypoxia decreased branchial surface area in B. splendens by ~40%, but neither hypoxia level affected Trichopodus branchial surface. Mild, but not severe, hypoxia increased labyrinth surface area by 30% in B. splendens. However, as for branchial surface area, labyrinth surface area was not affected in Trichopodus. These differential larval responses to hypoxic rearing suggest that different larval habitats and activity levels are greater factors influencing developmental plasticity than genetic closeness of the two species.

Very high blood oxygen affinity and large Bohr shift differentiates the air-breathing siamese fighting fish (Betta splendens) from the closely related anabantoid the blue gourami (Trichopodus trichopterus)

The Siamese fighting fish, Betta splendens, and the blue gourami, Trichopodus trichopterus, are two closely related air-breathing anabantoid fishes. B. splendens is a sedentary facultative air breather frequenting often hypoxic waters, while T. trichopterus is a more active obligatory air-breather inhabiting better oxygenated waters. Despite their close taxonomic relationship, previous studies have shown inter-specific differences in both physiological and morphological plasticity. Consequently, we hypothesized that B. splendens would have the higher blood oxygen affinity characteristics typical of more hypoxia-tolerant fishes. Whole blood oxygen equilibrium curves were determined at 27 °C and pHs of 7.62, 7.44 and 7.25. At a pH of 7.62, the blood O2 affinity (P50) of B. splendens was just 2.9 mmHg, while that of T. trichopterus was ~5 times higher at 14.7 mmHg. There were no significant differences in P50 between males and females in either species. The Bohr coefficient in B. splendens and T. trichopterus was −1.79 and − 0.83, respectively. B. splendens, unlike T. trichopterus, showed a large Root effect. Hills cooperatively coefficient, n, was ~2 in both species, indicating a significant binding cooperative between oxygen and hemoglobin. Collectively, these differences in blood O2 transport characteristics in these two closely related species are likely correlated with the differing habitats in which they breed and inhabit as adults, as well as different activity levels. Finally, the very high blood O2 affinity of B. splendens is not extraordinary among air-breathing fish, as revealed by a review of the literature of blood oxygen affinity in air-breathing fishes.

The Air-Breathing Paradise Fish (Macropodus opercularis) Differs From Aquatic Breathers in Strategies to Maintain Energy Homeostasis Under Hypoxic and Thermal Stresses.

Two major strategies are used by most fish to maintain energy homeostasis under hypoxia. One is to utilize alternative metabolic pathways to increase energy production, and the other is to limit energy expenditure by suppressing energy-consuming processes, especially ionoregulation. Some anabantoid fishes live in tropical rivers, where hypoxic environments occur frequently. We previously found that under ambient hypoxia, anabantoid fishes do not downregulate Na+/K+-ATPase (NKA) activity to conserve energy in gills but instead increase the frequency of air-breathing respiration (ABR). In addition to the hypoxic condition, another factor that may cause cellular hypoxia in fish is abnormally high environmental temperatures. The frequency of such extreme thermal events has increased due to global climate change. In the present study, we examined whether the anabantoid fish, Macropodus opercularis employs the two strategies mentioned above to resist both ambient hypoxic and elevated thermal (cellular hypoxic) conditions. Results indicate that neither glucose metabolism nor gill NKA activity were altered by hypoxia (DO = 1.5 ± 1 mg/L), but glucose metabolism was increased by thermal stress (34 ± 1°C). NH4+ excretion and ABR frequency were both increased under hypoxia, thermal or hypoxic-and-thermal treatments. In fish that were restricted from breathing air, increased mortality and glucose metabolism were observed under hypoxic or thermal treatments. These results suggest that for M. opercularis, increasing ABR is an important strategy for coping with unmet oxygen demand under hypoxic or thermal stress. This behavioral compensation allows anabantoid fish to physiologically withstand hypoxic and thermal stresses, and constitutes a mechanism of stress resistance that is unavailable to water-breathing fishes.

Phylogenesis of reproductive strategies in labyrinth fishes (Anabantoidei) and their sister groups

Clado- and semogenetic approaches, when used in concert, make it possible to resolve questions concerning the phylogenetic relationships between group representatives, as well as the phylogenesis of those representatives’ characters. Parental care patterns and other forms of reproductive behavior, along with the reproductive strategy as a whole, can be the subject of semogenetic analysis to no less an extent than morphological structures sensu stricto. One highly specialized form of parental care in fish, including representatives of the suborder of labyrinth fishes and their sister groups, appears to be parental food provisioning. In our view, the evolutionary origin of postembryonic brood provisioning in bony fishes is characterized by three main features: (1) in fish, different forms of postembryonic food provisioning are convergent in their origin; (2) any kind of brood provisioning is realized through exploitation of the already existing character and is maintained by selection due to an enhancement of offspring fitness; (3) the main evolutionary path of the emergence and development of this phenomenon consists in function expansion and replacement. The hypothesis does have heuristic power, since it enables prediction of the presence of the reproductive strategy component in question through the identification of adequate basic adaptations. Although parental care occurs in the majority of anabantoid fishes, there are still several species for which such care is not known. On the cladogram, these species by no means take the basal position but are surrounded by fishes that provide care for their eggs, or even their hatchlings. The parsimony principle leads to the suggestion that parental care is a plesiomorphic character in the suborder Anabantoidei (or in the order Anabantiformes). It seems that the ancestors of present day noncarrying species that take various positions within this phylogenetic group were fishes showing parental care. Their reproductive strategy later changed as a result of r selection. If this hypothesis is correct, the absence of parental care should be considered a case of reproductive strategy degradation. It is quite probable that parental food provisioning was a component of the ancestral reproductive strategies. It is also possible that the reproductive strategy of present-day anabantoids that supposedly do not care for their offspring actually includes some optional forms of parental care.

A fossil gourami (Teleostei, Anabantoidei) from probable Eocene deposits of the Ombilin Basin, Sumatra, Indonesia

ABSTRACT—Fossil fishes were first collected from deposits of the Sangkarewang Formation of the Ombilin Basin in Sumatra, Indonesia, in the 1870s, but a comprehensive study of these fishes was not published until almost 50 years later. New material from these deposits was collected in 2009, which included a small anabantoid fish. This fish is not conspecific with any of the material described previously and is here named as a new genus and species. A phylogenetic analysis indicates that the new anabantoid is closely related to Osphronemus among the taxa studied, and we place it in the family Osphronemidae. However, the new fish does not appear to belong to any of the named modern subfamilies, so we leave it incertae sedis in the family. Additionally, the osphronemid subfamilies are not recovered as a monophyletic group. The anabantoid named here and another previously described from the same locality are the only fossil anabantoids known. However, the age of the Sangkarewang Formation is not confirmed and has been variously attributed to Cretaceous, Palaeocene, Eocene, and Miocene, with Eocene being the presently favored estimate. Based on the presence of an anabantoid in these deposits, an Eocene age is reasonable and is not contradicted by the known fish fauna.http://zoobank.org/lsid:urn.lsid.zoobank.org:pub:53281C25-8E47-4C67-9952-A994B47F6656SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP

Morphological and Biochemical Variations in the Gills of 12 Aquatic Air-Breathing Anabantoid Fish

All fish species in the Anabantoidei suborder are aquatic air-breathing fish. These species have an accessory air-breathing organ, called the labyrinth organ, in the branchial cavity and can engulf air at the surface of the water to assist in gas exchange. It is therefore necessary to examine the extent of gill modification among anabantoid fish species and the potential trade-offs in their function. The experimental hypothesis that we aimed to test is whether anabantoid fishes have both morphological and functional variations in the gills among different species. We examined the gills of 12 species from three families and nine genera of Anabantoidei. Though the sizes of the fourth gill arch in three species of Trichogaster were reduced significantly, not all anabantoid species had morphological and functional variations in the gills. In these three species, the specific enzyme activity and relative protein abundance of Na(+)/K(+)-ATPase were significantly higher in the anterior gills as compared with the posterior gills and the labyrinth organ. The relative abundance of cytosolic carbonic anhydrase, an indicator of gas exchange, was found to be highest in the labyrinth organ. The phylogenetic distribution of the fourth gill's morphological differentiation suggests that these variations are lineage specific, which may imply a phylogenetic influence on gill morphology in anabantoid species.

Cerebral Lateralization and Its Relationship to Phylogeny and Aggression in Anabantoid Fishes

We examined the relationship between aggression and cerebral lateralization, as expressed by eye-use preference in a mirror-response paradigm, in six species of anabantoid fishes in the teleost family Belontiidae. Our objective was to test the hypothesis that frequency-dependent selection should decrease lateralization at the population level in more aggressive species. The mean laterality index did not differ significantly among the six species, nor did any of the species differ from zero, which suggests that populations of these fishes are neither left- nor right-eye biased. In spite of species differences in aggressive behavior, there was no relationship between population-level laterality and aggression as we had originally predicted. The degree of individual lateralization did differ significantly among some species. A phylogenetic generalized least squares (PGLS) analysis showed that these species differences were not due to phylogenetic distance. Individual laterality was also unrelated to aggression. Fishes in the genus Betta were the most individually lateralized, but varied in the intensity of aggressive behavior they displayed toward the mirrors. Taxa such as paradise fish (Macropodus opercularis and Pseudosphromenus dayi) were intermediate in eye-use preferences and gourami (Trichogaster trichopterus) displayed little cerebral lateralization. We also used the PGLS method to reconstruct ancestral values for individual lateralization in this group of fishes, from which we conclude that the ancestral condition was one of low variance in eye-use preference.

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.

Auditory role of the suprabranchial chamber in gourami fish.

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.

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.

Ontogenetic features of Luciocephalus (Perciformes, Anabantoidei) with a revised hypothesis of anabantoid intrarelationships

Abstract The anatomy of the anabantoid fish Luciocephalus is reinvestigated and the ontogeny of certain character complexes is described for the first time. Luciocephalus possesses the following previously unrecognized or equivocally interpreted anatomical characters: a clearly separate ectopterygoid bone, pharyngeal processes of the basioccipital, a toothless pharyngeal process of the parasphenoid, an infrapharyngobranchial I and an endoskeletal mental ossification. In addition, aspects of the ontogeny of the hyopalatine arch in the belontiids Belontia, Trichogaster and Ctenops are described. Species of these three genera all possess an ectopterygoid bone previously thought to be lacking in the family Belontiidae. The significance of the results for reconstructing interrelationships of the anabantoid families and the phylogenetic position of Luciocephalus is discussed. A revised hypothesis of anabantoid intrarelationships is presented. Four synapomorphic characters indicate that Luciocephalus is more closely related to the Beontiidae (including Osphronemus ) rather than forming the sister group of all other anabantoids.

Ontogenetic features of Luciocephalus (Perciformes, Anabantoidei) with a revised hypothesis of anabantoid intrarelationships

The anatomy of the anabantoid fish Luciocephalus is reinvestigated and the ontogeny of certain character complexes is described for the first time. Luciocephalus possesses the following previously unrecognized or equivocally interpreted anatomical characters: a clearly separate ectopterygoid bone, pharyngeal processes of the basioccipital, a toothless pharyngeal process of the parasphenoid, an infrapharyngobranchial I and an endoskeletal mental ossification. In addition, aspects of the ontogeny of the hyopalatine arch in the belontiids Belontia, Trichogaster and Ctenops are described. Species of these three genera all possess an ectopterygoid bone previously thought to be lacking in the family Belontiidae. The significance of the results for reconstructing interrelationships of the anabantoid families and the phylogenetic position of Luciocephalus is discussed. A revised hypothesis of anabantoid intrarelationships is presented. Four synapomorphic characters indicate that Luciocephalus is more closely related to the Beontiidae (including Osphronemus) rather than forming the sister group of all other anabantoids.

The Volume of Air Within the Swimbladder and Breathing Cavities of the Anabantoid Fish Colisa Laua (Perciformes, Belonthdae)

ABSTRACT The swimbladder volume and air volume within the breathing chambers of the anabantoid fish Colisa lalia have been measured. These data help in the understanding of some of the functions of these organs and are necessary for an analysis of their role in hearing and sound production. By means of a simple trick (based on new data) it was possible to analyse the time course of air volume changes in the breathing chambers at different temperatures. The results are well described by a simple diffusion model. The temperature-dependence of the time course suggests an interesting increase of the diffusion constant with temperature. Under constant conditions the chambers were always filled with about the same volume of air. No excess pressure was found. Typical values of a single chamber’s air content range from 34 to 58 μl. Air content increases with about the third power of fish length. By using the present data, the time course of air volume changes in the chambers of a given fish can be estimated. Swimbladder volumes, determined using Boyle’s law, ranged from 70 to 220 μ1 and were also found to increase with about the third power of fish length, in accordance with a simple estimation. The data are discussed in relation to buoyancy, diffusion processes, blood circulation, hearing and sound production and suggest some interesting new work.

A possible relationship between aspects of dentition and feeding in the centrarchid and anabantoid fishes

Certain components of dentition — teeth on the third basibranchial in the Centrarchidae and on the parasphenoid in the anabantoids (sensu lato) — are very rare elsewhere in higher teleostean fishes. Though these basibranchial and parasphenoid teeth in the two fish groups are on opposite sides of the oral cavity, it is hypothesized that they both developed as adaptations for gripping a particular category of food items, namely strong-clawed, hard-shelled, active animals that, once within the oral cavity, would try to crawl out again. A corollary to this hypothesis is that higher teleosts with extensive dentition in the central part of the oral cavity have a grasping jaw bite, which, unlike a piercing, shearing, or crushing jaw bite, does not necessarily kill the prey that is taken into the oral cavity.

Comparative analysis of the hyomandibula during respiration in anabantoid teleost fishes:Macropodus opercularis in relation toCtenopoma acutirostre andAnabas testudineus

The hyomandibula is an important element of the head and has been influenced in its structural features by different functions. It is roughly T-shaped with vertical and horizontal ridges. In fish, such asMacropodus opercularis, the deep curve at the antero-dorsal end of this ridge may be correlated with the concentrated force applied by the adductor mandibulae muscles. InMacropodus opercularis, the hyomandibula is shorter and broader than that ofAnabas testudineus andCtenopoma acutirostre. In the latter two genera, almost the entire anterior surface of the vertical ridge is uniformly curved, which may be explained by the distributive force applied by the adductor muscles perpendicularly. InMacropodus, the hyomandibulo-interhyal articulation is not straight along the axis of the interhyal as inCtenopoma, but the interhyal makes an angle of 45‡ to the hyomandibula. This difference might be functional as well as due to the oblique orientation of the hyomandibula. InCtenopoma, the articulating facet of the symplectic is located slightly below the interhyal articulating facet. This might be caused by the location of the hyomandibulo-interhyal articulation. The hyomandibulo-interhyal articulation also helps in the depression of the lower jaw when the latter is depressed by the hyoid bar. Thus, a structural analysis of the hyomandibula and its relationships with functions depict the interrelation between form and function.

Defensive behaviour and its inheritance in the anabantoid fish, Macropodus opercularis and Macropodus opercularis concolor

In this preliminary study defense behaviour patterns (fear responses) are described in two closely related, behaviourally different inbred labyrinth fish subspecies and in their F 1 generation. The subspecies M. opercularis (characterized briefly by “active escape”) and M. opercularis concolor (characterized by “passive escape”) showed specific differences in the manifestation of certain defense behaviour patterns. In the F 1 hybrid generation dominance and overdominance of M. opercularis was found in most defense behaviour patterns. Analysing the frequencies and sequences of movement patterns it could be shown that defensive behaviour is not a random or entirely “plastic” process but that there is sequential linkage between the patterns and they form characteristic clusters. Our results suggest that manifestations of different patterns are under genetic control and presumably, genetic determination of certain patterns is not very complex. Attempts were made to determine whole brain noradrenaline, serotonine and dopamine levels of the two subspecies and a significant difference was found in the noradrenaline content.