Ethological analysis of social and environmental effects on the distribution of the behavioural elements of the paradise fish (Macropodus opercularis L.)

The behaviour of the paradise fish in a new environment was examined. The influence of space, the number of conspecifics, the number of each sex in the various groups and the time allowed for habituation on the distribution of various behavioural elements were revealed. By means of principal component analysis five biologically meaningful factors were extracted. Using the scores of the various factors as indices of settlement, fear passive behaviours and exploration, special effects of the environment were assessed. In small tanks the fish are in an emotionally low but stable continuously trying to escape. Being prevented from escaping they develop various passive behaviour forms which alternate with escape. In a large tank the main occupation of a female fish on its own is exploration at the start; this is accompanied by a fearful state. In groups of four the activity of exploration is less and the fearful state is not so pronounced. The presence of males seems to secure the most stable state in a new environment though after about two days the males start breeding activities and this involves a new spectrum of activities.     

Variable learning performance: the levels of behaviour organization

Our experiments were focused on some special aspects of learning in the paradise fish. Passive avoidance conditioning method was used with different success depending on the complexity of the learning tasks. In the case of simple behavioural elements various "constrains" on avoidance learning were found. In a small, covered place the fish were ready to perform freezing reaction and mild punishment increased the frequency and duration of the freezing bouts very substantially. However, it was very difficult to enhance the frequency of freezing by punishment in a tank with transparent walls, where the main response to punishment was escape. The most easily learned tasks were the complex ones which had several different solutions. The fish learned to avoid either side of an aquarium very easily because they could use various behavioural elements to solve the problem. These findings could be interpreted within the framework of different organizational levels of behaviour.     

Long-term memory and recognition of another species in the paradise fish

The habituation of exploration of a goldfish, Carassius auratus, by paradise fish, Macropodus opercularis, was examined. A first encounter of at least 1 min was necessary for habituation to be found in a second encounter 3 h later. When the paradise fish were allowed 5 min to explore the goldfish and the second encounters were staged between 3 h and 3 months later, persistent habituation was found. The relation between memories formed during encounters with other species and modelling of the environment by the paradise fish is discussed.     

Behavioural elements of the paradise fish (Macropodus opercularis). II. A functional analysis

The effects of various environmental influences, such as presence of a living predator, dominant conspecific or frightening artificial stimulus, on particular behavioural elements of the paradise fish (Macropodus opercularis) were studied. It has been found that certain sets of the elements are manifested as organizational complexes which characterize a certain mood or form of defensive behaviour. The compound elements of these complexes seemed to be partly interchangeable, partly under environmental control. The main behavioural complexes of the paradise fish were the Territorial complex characterizing the peaceful, home-living animal, Emotionality complex appearing in case of immediate and past danger, Active and Passive defense complexes which represent alternatives in defense, depending on particular stimulus settings. Some supporting evidence for these behavioural complexes are presented and discussed.     

Hypoxia and the antipredator behaviours of fishes

Hypoxia is a phenomenon occurring in marine coastal areas with increasing frequency. While hypoxia has been documented to affect fish activity and metabolism, recent evidence shows that hypoxia can also have a detrimental effect on various antipredator behaviours. Here, we review such evidence with a focus on the effect of hypoxia on fish escape responses, its modulation by aquatic surface respiration (ASR) and schooling behaviour. The main effect of hypoxia on escape behaviour was found in responsiveness and directionality. Locomotor performance in escapes was expected to be relatively independent of hypoxia, since escape responses are fuelled anaerobically. However, hypoxia decreased locomotor performance in some species (Mugilidae) although only in the absence of ASR in severe hypoxia. ASR allows fish to show higher escape performance than fish staying in the water column where hypoxia occurs. This situation provides a trade-off whereby fish may perform ASR in order to avoid the detrimental effects of hypoxia, although they would be subjected to higher exposure to aerial predation. As a result of this trade-off, fishes appear to minimize surfacing behaviour in the presence of aerial predators and to surface near shelters, where possible. For many fish species, schooling can be an effective antipredator behaviour. Severe hypoxia may lead to the disruption of the school unit. At moderate levels, hypoxia can increase school volume and can change the shuffling behaviour of individuals. By altering school structure and dynamics, hypoxia may affect the well functioning of schooling in terms of synchronization and execution of antipredator manoeuvres. School structure and volume appear to be the results of numerous trade-offs, where school shape may be dictated by the presence of predators, the need for energy saving via hydrodynamic advantages and oxygen level. The effects of hypoxia on aquatic organisms can be taxon specific. While hypoxia may not necessarily increase the vulnerability of fish subject to predation by other fish (since feeding in fish also decreases in hypoxia), predators from other taxa such as birds, jellyfish or aquatic mammals may take advantage of the detrimental effects of hypoxia on fish escape ability. Therefore, the effect of hypoxia on fish antipredator behaviours may have major consequences for the composition of aquatic communities.     

Behavioural elements of the paradise fish (Macropodus opercularis). I. Regularities of defensive behaviour

Elements of behaviour were described for the paradise fish on the basis of the topography, location and orientation of the animal observed in various seminatural and laboratory environments. Correlations and repeatability of the parameters of observation were analyzed. Environmental influences, such as periods of the day, housing, feeding, raising conditions, presence of various frightening stimuli and species, and strain differences, were studied. Several functional connections were found among the behavioural elements.     

The ontogeny of antipredator behaviour in paradise fish larvae III. Size-related avoidance of predator models

A series of laboratory experiments examined the effect of the size of predator models on the behaviour of larval paradise fish Macropodus opercularis L., of different age (15 and 20 days old). In a small runway, individually tested larval paradise fish could approach a model predator of different size, with or without eyespots. The duration of approach, the number of fleeings and backings were measured. Fifteen-day-old larvae showed similar moderate level avoidance to all models but fled more in the presence of larger models. The approach behaviour in 20-day-old larvae depended on the size of the model but only if it had eyespots, and larvae fled and backed more with increasing size of the model. However, manipulation of larval length within age category had no effect on predator avoidance. Since the recognition of the two eyespots seems to be age dependent, it is hypothesized that only larvae older than 20 days will behave in a 'threat-sensitive' manner in the presence of predator models by avoiding larger ones.     

The influence of olfaction on exploratory behaviour in the paradise fish (Macropodus opercularis L.)

We studied the role of olfaction on the exploratory behaviour of the paradise fish. Intact animals showed habituation of exploratory behaviour toward a heterospecific fish after five consecutive encounters. Fish with olfactory nerves destroyed spent significantly longer time with exploration compared to the control even at the fifth encounter. We suggest that olfactory inputs have a strong influence on exploratory behaviour.     

The behaviour of the paradise fish (Macropodus opercularis) in two different open-fields. A correlation study

The behaviour of the paradise fish in a traditional "closed" and in a new "transparent" open-field was investigated. The traditional way of measuring ambulation scores was extended by recording ethologically defined behaviour units. The correlations found between the scores measured in the "closed" field and those measured in the "transparent" field are discussed in this paper.     

The Role of Eyespots in Predator Recognition and Antipredatory Behaviour of the Paradise Fish,Macropodus opercularis L.

The behaviour of the paradise fish in the presence of either a living pike or catfish or various dummies or a catfish equipped with a pair of artificial big eyes was studied. Analyzing the first encounter two separate periods of the predator recognition process could be identified. The primary form of the antipredatory reaction was orientation (looking at the object while keeping a constant eye and body position) which was elicited by any fish-like bodies but not by bare eyes without a body. During orientation the presence of pike or catfish equipped with artificial eyes or a dummy with horizontal eyes elicited fin erecting display. This reaction was sporadic in the presence of a normal catfish which has very small eyes. The biological significance of this separate organization of exploration is discussed.     

Intra- and interspecific social and aggressive behaviour in the Siamese fighting fish, Betta splendens

Siamese fighting fish and paradise fish were operantly conditioned to swim through a cylinder to open a door allowing them to explore visually conspecifics, non-conspecifics, or inanimate objects. All stimuli significantly elevated response levels compared to control conditions, but response rate for conspecifics was not higher than for other stimuli. In experiments 2 and 3, fighting fish threatened and attacked non-conspecifics as well as conspecifics, and using a blind procedure, displays to different fish could not be easily differentiated. A final experiment with models failed to reveal any specific shape critical to the elicitation of the threat display. Discussion considers the possibility that the operant behaviour of the fighting fish may be motivated by visual exploration as well as by aggression, and that in actual fighting the social behaviour of the opponent may be more crucial than any particular visual stimulus.     

Differences in Timidity and Escape Responses between Predator-naive and Predator-sympatric Rainbowfish Populations

Responses of rainbowfish ( Melanotaenia duboulayi ) from two populations towards a) an active and a passive predatory fish and b) a novel trawl apparatus, were compared. Predator-sympatric fish avoided the fish predators and showed stronger avoidance behaviour in response to the active predator. These fish used predator inspection excursions to rapidly assess the potential risk and their escape responses were consistently effective. In contrast the predator-naive fish ignored the passive predator but were continually drawn towards the active predator possibly due to generalized curiosity and the absence of significant negative feedback from the predator, which was restrained by a clear Perspex partition. Despite this attraction, the predator-naive fish did not display typical predator inspection behaviour and showed very poor escape performance when initially confronted by the trawl apparatus. Many of these fish, however, showed rapid improvement in their escape performance through learning. These results suggest that predator-sympatric rainbowfish have the capacity to assess the level of threat posed by a predator and predator-naive rainbowfish learn to implement appropriate escape strategies when forced to evade a threat.     

Different Duration of Memory for Conspecific and Heterospecific Fish in the Paradise Fish (Macropodus opercularis L.)

During an encounter with a conspecific male paradise fish memorize some features of their opponent. This is revealed by the shorter contest and significant changes in their behaviour at a second encounter with the same opponent. Memory of an opponent lasts for one day but is undetectable after a week. In contrast memorization of other species could easily be preserved for several months. A possible explanation of these different durations is discussed in relation to their functions in the life of the fish.     

Conditions for recognition of a physical space as two compartments by the paradise fish (Macropodus opercularis, Pisces, Anabantidae).

By using a series of aquaria which were more or less separated into two compartments by a "gate" made of plastic or glass strips, we studied how paradise fish changed their patterns of movement relative to the nature of the gates. We found that opaque plastic gates were well recognized by the paradise fish (Macropodus opercularis) even if the strips were very thin, but that they only changed their pattern of movement when the strips exceeded a certain width. Thus, paradise fish appear to be able to distinguish a physical space of the same size as consisting of either one or two compartments, depending on the material (transparency) and width of the dividing strips.     

Effects of Emergence Time and Early Social Rearing Environment on Behaviour of Atlantic Salmon: Consequences for Juvenile Fitness and Smolt Migration

Consistent individual differences in behaviour have been well documented in a variety of animal taxa, but surprisingly little is known about the fitness and life-history consequences of such individual variation. In wild salmonids, the timing of fry emergence from gravel spawning nests has been suggested to be coupled with individual behavioural traits. Here, we further investigate the link between timing of spawning nest emergence and behaviour of Atlantic salmon (Salmo salar), test effects of social rearing environment on behavioural traits in fish with different emergence times, and assess whether behavioural traits measured in the laboratory predict growth, survival, and migration status in the wild. Atlantic salmon fry were sorted with respect to emergence time from artificial spawning nest into three groups: early, intermediate, and late. These emergence groups were hatchery-reared separately or in co-culture for four months to test effects of social rearing environment on behavioural traits. Twenty fish from each of the six treatment groups were then subjected to three individual-based behavioural tests: basal locomotor activity, boldness, and escape response. Following behavioural characterization, the fish were released into a near-natural experimental stream. Results showed differences in escape behaviour between emergence groups in a net restraining test, but the social rearing environment did not affect individual behavioural expression. Emergence time and social environment had no significant effects on survival, growth, and migration status in the stream, although migration propensity was 1.4 to 1.9 times higher for early emerging individuals that were reared separately. In addition, despite individuals showing considerable variation in behaviour across treatment groups, this was not translated into differences in growth, survival, and migration status. Hence, our study adds to the view that fitness (i.e., growth and survival) and life-history predictions from laboratory measures of behaviour should be made with caution and ideally tested in nature.     

The Ontogeny of Antipredator Behaviour in Paradise Fish Larvae (Macropodus opercularis): The Recognition of Eyespots

The ontogeny of antipredator behaviour of paradise fish was examined using a simple testing method. The results showed that the ability to recognize eyespots appears in larvae between 15 and 20 days old. At the same time, two horizontally placed eyespots produced the most pronounced effect on the behaviour of the larvae. Testing different inbred strains revealed that the ontogeny of antipredator behaviour is also under broad genetic control, as the response to a model predator was strong in one strain but weak in another. The results suggest that the recognition of eyespots coincides with larvae becoming more conspicuous to predators by the appearance of pigmentation and increasing level of swimming activity (exploration).     

Parasite-induced change in host behaviour and susceptibility to predation in an eye fluke-fish interaction

Trophically transmitted parasites may increase their transmission efficiency by altering the behaviour of infected hosts to increase their susceptibility to predation by target hosts (the next host in the life cycle). The parasite Diplostomum spathaceum (Trematoda) reduces the vision of its fish intermediate hosts: its metacercariae lodge themselves in the eyes of fish and induce cataract formation, which gives them the opportunity to affect fish behaviour. We examined whether D. spathaceum eye flukes change the preference of fish for the surface layers of the water column or their escape behaviour, which could make the fish more vulnerable to predation by bird hosts. We also studied the influence of parasites on the susceptibility of fish to artificial aerial predators that were able to catch fish from the water surface. Infected and control fish did not differ in their preference for the surface layers but infected fish showed less escape behaviour when a black plate was drawn over the water surface. They were also more easily caught by human ‘predators’ dipping a net into the tank. Thus, infected fish should be easier prey for gulls and terns, implying that the ability of D. spathaceum eye flukes to alter fish behaviour may be a parasite strategy evolved to enhance transmission.     

Benthivorous fish reduce stream invertebrate drift in a large-scale field experiment

Drift as a low-energy cost means of migration may enable stream invertebrates to leave risky habitats or to escape after encountering a predator. While the control of the diurnal patterns of invertebrate drift activity by fish predators has received considerable interest, it remains unclear whether benthivorous fish reduce or increase drift activity. We performed a large-scale field experiment in a second-order stream to test if invertebrate drift was controlled by two benthivorous fish species (gudgeon Gobio gobio and stone loach Barbatula barbatula). An almost fishless reference reach was compared with a reach stocked with gudgeon and loach, and density and structure of the invertebrate communities in the benthos and in the drift were quantified in both reaches. The presence of gudgeon and stone loach reduced the nocturnal drift of larvae of the mayfly Baetis rhodani significantly, in contrast to the findings of most previous studies that fish predators induced higher night-time drift. Both drift density and relative drift activity of B. rhodani were lower at the fish reach during the study period that spanned 3 years. Total invertebrate drift was not reduced, by contrast, possibly due to differences in vulnerability to predation or mobility between the common invertebrate taxa. For instance, Chironomidae only showed a slight reduction in drift activity at the fish reach, and Oligochaeta showed no reduction at all. Although benthic community composition was similar at both reaches, drift composition differed significantly between reaches, implying that these differences were caused by behavioural changes of the invertebrates rather than by preferential fish consumption. The direction and intensity of changes in the drift activity of stream invertebrates in response to the presence of benthivorous fish may depend on the extent to which invertebrate taxa can control their drifting behaviour (i.e. active versus passive drift). We conclude that invertebrate drift is not always a mechanism of active escape from fish predators in natural streams, especially when benthos-feeding fish are present.     

Can Paradise Fish (Macropodus opercularis,Anabantidae) Recognize a Natural Predator? An Ethological Analysis

Predator recognition and avoidance by paradise fish have been studied with allopatric species and model experiments. The effect of sympatric predators has not been investigated. Here I report that reactions of paradise fish towards a sympatric predator (Channa micropeltes) are quantitatively different from those shown towards an allopatric predator or different harmless species of fishes. I investigate the possible cues eliciting this differential response and show that visual as well as olfactory stimuli play roles. Olfactory stimuli from the sympatric predator alone elicit an elevated level of activity from paradise fish; the appearance of the sympatric predator (with or without olfactory stimuli) results in an exploratory and display reaction. I speculate what visual stimuli may play roles in predator recognition in paradise fish and I suggest that previously asserted key stimuli such as the eyes of the encountered heterospecific fish may not differentiate the harmful species from innocuous. I conclude that the antipredatory behavior of paradise fish may be affected by both genetic factors and learning and that the relative importance of the former or latter factor may vary depending on the situation.     

Exploratory and fear behaviour in the Macropodus opercularis subspecies, M. o. opercularis and M. o. concolor

The exploratory and fear behaviours of the black paradise fish M. o. concolor, a subspecies of Macropodus opercularis, were studied and compared with those of an outbred and two inbred populations of M. o. opercularis. Adult females of both subspecies were used in the behaviour tests (open-field, novel-object approach, restricted space), and their behaviour in novel and startling situations was described by recording the sequence of ethologically defined units. It was found that M. o. concolor more often responded passively to novelty and fear-evoking stimuli. In 11 out of 14 test measures they were less active (p less than 0.01 and 0.001) than the outbred M. o. opercularis, but were very similar to our extremely timid strain P. Subtle differences of behaviour units were found between the subspecies, e.g. the absence of interrupted movements and unoriented panic behaviour in M. o. concolor. It is supposed that alternative passive and active strategies represent genetic polymorphism of defensive behaviour.