Audience effect is context dependent in Siamese fighting fish, Betta splendens

Investigations of communication networks in animals have focusedprimarily on determining whether animals extract informationfrom peripheral contests (eavesdropping) or respond to the presenceof bystanders (audience effect). The possibility that an animal'sresponse to being watched might be context dependent, however,has been explored in far less detail. This study investigatedthe influence of two contexts, exposure to audiences of differentsexes and presence or absence of a nest, on the aggressive behaviorof interacting male Siamese fighting fish, Betta splendens.Males interacted in the presence (male, female) or absence ofan audience in three different nest conditions (0, 1, or 2 nests).Audience sex and territorial status influenced aggressive behaviorin the interacting males, but a strong audience x nest interactionalso was uncovered. Males were more aggressive when neithermale had a nest and a male audience was present than when afemale or no audience was present. Males also were more aggressivewhen only one male had a nest and a male audience was presentthan when a female or no audience was present. When both maleshad nests and a male audience was present, however, males wereless aggressive than when only one male or neither male hada nest. In sum, aggressive behavior was influenced by the interactionbetween audience and nest; neither nest nor audience alone wassufficient to explain the results. Male Siamese fighting fishalter their behavior based on both external cues, the sex ofthe audience, and internal cues, reproductive state and resourcepossession. Our results emphasize the importance of consideringaspects of an animal's environment when examining audience effectsand communication networks in general.     

Sexual size dimorphism and male contest in wild Siamese fighting fish

Larger males of the wild Siamese fighting fish Betta splendens were more successful in male contests. There were no differences in fighting duration among treatments. Comparing agonistic behaviour between large and small males in ±1 and ±2 S.D treatments, larger males attacked, chased and performed total agonistic behaviour more than smaller males. There were no differences between larger and smaller males concerning other agonistic behaviour during fighting. Females presented with two potential mates of different sizes did not prefer larger males.     

Bubble nest habitat characteristics of wild Siamese fighting fish

Siamese fighting fish Betta splendens in Thailand inhabit shallow water amongst dense emergent vegetation near the margin of rice paddy fields. Nesting water was low in dissolved oxygen, pH, salinity but high in free carbon dioxide and temperature. This fish aggregated with a mean population density of 1·7 fish m^-2 and an equal adult sex ratio. Males were heavier, longer and wider than females. The size of their bubble nest is in proportion to their weight and length.     

Timing of presentation of an audience: aggressive priming and audience effects in male displays of Siamese fighting fish (Betta splendens)

Studies of animal communication often underestimate the presence of individuals other than the signaller-receiver dyad. Signalling interactions often occur in the presence of non-participating individuals (audiences); the effect of these individuals upon the dynamics of interactions has been called the audience effect. Recent studies of fighting fish Betta splendens have shown that the presence of a male audience can increase aggression during interactions. However, in many of these studies males were allowed to see the audience prior to the interaction, thus such pre-exposure may have facilitated aggressive behaviour (aggressive priming). Here we present results of two experiments designed to examine the relative importance of priming and audience effects on the dynamics of aggressive interactions. Males that were pre-exposed showed higher levels of aggression during subsequent interactions regardless of the presence or absence of an audience. When only one of the interactants had been pre-exposed to the audience, the non-exposed male showed similar increases in aggressive behaviour, i.e. matching the level of aggression showed by his opponent. Taken together these results suggest that aggressive priming may have resulted in an over-estimation of the audience effect in previous studies. The results still highlight the importance of social environment in determining the dynamics and outcomes of aggressive contests.     

Instrumental conditioning of choice behavior in male Siamese fighting fish (Betta splendens)

Despite the differences in the response of male Betta splendens toward various stimuli, no research has attempted to determine the preference for a live conspecific versus a mirror presentation. A submerged T-maze was used to present both stimuli to healthy male B. splendens (N=16). The results indicated that subjects' start box and swimway latencies decreased significantly over the 30 trials. Moreover, the analysis of choices demonstrated a modest, but statistically significant, preference for the live conspecific over the mirror presentation. The results are discussed in terms of the stimuli qualities that elicit an aggressive response in B. splendens and the implications for common experimental procedures.     

The influence of sex, line, and fight experience on aggressiveness of the Siamese fighting fish in intrasexual competition

We examined the influence of sex, line, i.e., broods from different parents, and previous fight experience on the aggressiveness of the Siamese fighting fish Betta splendens in intrasexual competition. The innate aggressiveness of the fish against their mirror images was measured on the day prior to the direct fight with other individuals, and it was found to be influenced by the line type but not by the sex. In the direct fight with other individuals, the males invested more effort in the fight than the females. In addition, the individuals of a particular line that exhibited a lower innate aggressiveness spent less time in the direct fight and were often losers when compared with those of other lines. After the direct fight with other individuals, the aggressiveness of the fish against their mirror images was remarkably influenced by the outcome of the direct fight, i.e., the winners exhibited more aggressive behavior, whereas the losers exhibited a lesser degree of aggressive behavior. This influence of the previous fight experience on subsequent aggressiveness was the greatest in the individuals of the line that have exhibited the lowest innate aggressiveness. However, the positive effect of the winning experience or the negative effect of the losing experience on subsequent aggressiveness decreased following several days after the previous fight increased.     

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.     

The pharyngeal organ in the buccal cavity of the male Siamese fighting fish, Betta splendens, supplies mucus for building bubble nests

The male Siamese fighting fish, Betta splendens, builds a bubble nest on the water surface to care for offspring during the reproductive period. To our knowledge, this study is the first to determine the composition of the bubble nest and to compare the pharyngeal organs of male and female Siamese fighting fish to determine the relationship between the pharyngeal organ and the ability to make bubble nests. Dot blots of the bubble nest probed with periodic acid-Schiff's (PAS) staining and Ponceau S solution revealed that the contents of the nest are glycoprotein rich. Dissection of the heads of Siamese fighting fish showed that the pharyngeal organ is located in the position through which inhaled air passes. The epithelial structure of the pharyngeal organ of the Siamese fighting fish, like that of other teleosts, has numerous wrinkles and papillae. Mucous goblet cells were observed on the epithelium of pharyngeal organs in male and female fish. The pharyngeal organ was found to be larger in male than in female fish. In addition, the epithelium of the pharyngeal organ in male fish has a greater number of mucous goblet cells than that in female fish. In Siamese fighting fish, this sexual dimorphism of the pharyngeal organ suggests that the male fish secretes more glycoprotein-rich mucus to build the bubble nest. Future work will focus on the type of mucous cells found in the epithelium of the pharyngeal organ that contributes to bubble formation and will determine the components of the mucus in the bubble nest.     

The novel application of an immunological technique reveals the immunosuppressive effect of phytoestrogens in Betta splendens

The novel application of a standard technique to assess cell‐mediated immune response to phytohaemagglutinin (PHA) injected in the caudal peduncle in a small fish (<2 g) to test the immunosuppressive effect of three phytoestrogens: genistein, equol and β‐sitosterol is described. Individual Betta splendens exposed to these oestrogenic chemicals produced weaker inflammatory responses to PHA than did control individuals, suggesting that phytoestrogens are immunosuppressive. This technique should enable immune function in fish species, too small to provide sufficient blood for conventional immunological measures, to be assessed.     

A non-social and isolate rearing condition induces an irreversible shift toward continued fights in the male fighting fish (Betta splendens)

Effects of rearing conditions were examined in the development of agonistic behaviors in the male fighting fish. In group-I (highly social), fish were communally reared. In group-II (highly social and isolate), fish were individually housed and exposed to the group-I fish through transparent walls until the sexual maturity (from 6 to 12 weeks post-hatch). In group-III (social and isolate), individually housed fish were similarly exposed to other fish within the group. In group-IV (non-social and isolate), individually housed fish were further visually isolated. Agonisitc behaviors were compared among males of the groups-II, -III, and -IV in their fights against the group-I male. The group-IV males showed significantly higher rate of wins than the groups-II and -III males, without differences in the incidence of agonistic behaviors (butt-or-bite, chase, and gill-cover erect) before the termination of the mutual fights. Increased incidence of agonistic behaviors was found after the termination (particularly in the unilateral chase), suggesting that the group-IV males continued to fight even after the opponent male displayed a submission. The aggression was also enhanced in the group-II, when they were thereafter reared in a social isolation after the sexual maturation; a critical period was thus not found. The enhanced aggression was not reversed in the group-IV, when they were thereafter exposed to social stimuli; shift to the continued fights was irreversible. Possible fitness gain of the enhanced aggression was discussed in terms of the adjustability to altered biological resources.     

Red fish, blue fish: trade-offs between pigmentation and immunity in Betta splendens

Carotenoid pigments are responsible for many examples of sexuallyattractive red, orange, and yellow coloration in animals andplay an important role in antioxidant and immune defenses. Becausevertebrates cannot synthesize carotenoids, limited dietary availabilitymay impose a trade-off between maintaining ornamental colorationand health. We used an experimental approach to test the carotenoidtrade-off hypothesis in the fighting fish Betta splendens, byexamining whether carotenoid allocation strategies differ amongconspecifics that exhibit a gradient of body coloration fromblue to red. We found that male redness is underlain by carotenoidsand that females preferred to associate with red males overblue males, suggesting a sexually-selected advantage to beingred. Moreover, we found strong experimental support for thecarotenoid trade-off hypothesis, as individuals that variedin color did not appear to allocate carotenoids equally to bothimmune response and coloration. Redder fish given supplementalcarotenoids increased in both immune response (to a phytohemagglutinationchallenge) and redness compared with controls. In contrast,bluer fish given supplemental carotenoids did not become morered but instead benefited immunologically more so than eithercontrol or redder supplemented fish. These results enhance ourunderstanding of the evolution and plasticity of carotenoidmobilization and utilization pathways in animals.     

Effect of a Dummy Audience on Male–Male Interactions in Siamese Fighting Fish, Betta splendens

Recent research on Siamese fighting fish, Betta splendens, and other taxa has demonstrated that an audience can cause males to alter their behavior in aggressive interactions. One factor not taken into account in these studies is how exactly the audience influences these interactions. It is possible that a live audience may interact with the subjects, creating an active communication network rather than a signaler–receiver dyad with a passive audience. Here, we used a dummy audience to control for information exchange between the audience and the interactants that might cause them to modify their behavior. Audience treatments included dummies of male and female B. splendens, a dummy cichlid, and a control condition with no audience present. The presence of a dummy audience did not influence male–male interactions. However, males spent the most time near the audience tank when the audience was a dummy of a B. splendens. This suggests that some factor other than the physical presence of the audience is responsible for the modification of behavior found in previous audience effect studies in Siamese fighting fish. However, we cannot rule out definitively that our dummy audience is viewed as unimportant by the opponents and, thus, ignored. Further research is necessary to determine which component of the audience is important for producing audience effects.     

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.     

The energetic costs of fighting in the house cricket, Acheta domesticus L

The cost of performing an agonistic behavior, or tactic, willhave consequences for an individual's rate of cost accrual,the tactic's evolutionary stability if used as an assessmentsignal, and its pattern of use in the behavioral choreographyof a contest. Few studies have attempted to quantify the costsof fighting, particularly with regard to energy expenditure.Flow-through respirometry revealed that house cricket malescan expend energy at relatively high rates when fighting (morethan eight times resting levels) depending on the particulartactic performed. Acoustic signalling (stridulation) constitutedthe least costly of seven measured agonistic tactics, whilewrestling with an opponent, the most energetically costly tactic,consumed oxygen at a net rate more than 40 times that of stridulation.Low-cost tactics are used by opponents more frequently thanmore costly tactics, providing evidence for an escalating tacticalconvention based on energetic costs. The moderate energeticcost of some tactics suggests they may function as reliablesignals in the assessment of wrestling ability. Net energy costsper contest increased linearly with contest duration for bothcontest winners and losers, but winners tended to expend moreenergy per contest than losers. The likely fitness effects ofenergy expended while fighting are discussed. The results ofthis study indicate that energy expenditure is an importantcost shaping contest strategies in Acheta domesticus     

The effect of body coloration and group size on social partner preferences in female fighting fish (Betta splendens)

Females of the fighting fish Betta splendens have been shown to associate with other B. splendens females in a manner reminiscent of shoaling behavior. Since body coloration varies dramatically in this species, and since body coloration has been shown to affect shoalmate choice in other species of fish, we examined the influence of body coloration on association preferences in female B. splendens. In dichotomous choice tests, B. splendens females spent more time swimming near groups of females (regardless of coloration) than swimming near an empty chamber, and chose to swim near fish of similar coloration to their own when choosing between two distinctly colored groups of females. When examining the interplay between body coloration and group size, focal fish spent more time swimming near larger groups (N = 5) of similarly colored fish than swimming near an individual female of similar coloration. However, focal fish showed no preference when presented with an individual female of similar coloration and a larger group of females of dissimilar coloration. These results suggest that association choices in B. splendens females are strongly affected by both body coloration and by group size.