Probing aggressive motivation in a cichlid fish

The duration of startles provides an inverse measure of motivation to resume the previous activity. Here, we use a novel method in which one convict cichlid fish (Amatitlania nigrofasciata) of a competing pair was startled independently of the opponent. Fish were given various opponents and the mean startle duration determined. This mean was negatively correlated with the mean use of highly escalated ‘frontal activities’ such as biting and frontal display, but not the less escalated lateral displays or tail beating. Thus the startle duration was a reliable surrogate measure of the most escalated components of aggressive interactions. That is, it provided a motivational probe for aggressiveness of individual fish. Fight motivation is often determined in terms of fight duration or physiological costs for losers, who reveal the costs they are prepared to pay. We discuss various potential advantages of the motivational probe over previous measures, particularly with respect to winners and losers and different times during the interactions.     

Male courtship preferences demonstrate discrimination against allopatric colour morphs in a cichlid fish

Whether premating isolation is achieved by male‐specific, female‐specific or sex‐independent assortative preferences often depends on the underlying evolutionary processes. Here we test mate preferences of males presented with females of different allopatric colour variants of the cichlid fish Tropheus sp., a Lake Tanganyika endemic with rich geographical colour pattern variation, in which the strength of sexual isolation varies between populations. We conducted two‐way mate choice experiments to compare behaviour of males of a red‐bodied morph (population Moliro) towards females from their own population with behaviour towards females from four allopatric populations at different stages of phylogenetic and phenotypic divergence. Males courted same‐population females significantly more intensely than females of other populations, and reduced their heteromorphic courtship efforts both with increasing genetic and increasing phenotypic distinctness of the females. In particular, females of a closely related red‐bodied population received significantly more courtship than either genetically distinct, similarly coloured females (‘Kirschfleck’ morph) or genetically related, differently coloured females (‘yellow‐blotch’ morph), both of which were courted similarly. Genetically and phenotypically distinct females (Tropheus polli) were not courted at all. Consistent with previous female‐choice experiments, female courtship activity also decreased with increasing genetic distance from the males’ population. Given successful experimental and natural introgression between colour morphs and the pervasive allopatry of related variants, we consider it unlikely that assortative preferences of both sexes were driven by direct selection during periods of secondary contact or, in turn, drove colour pattern differentiation in allopatry. Rather, we suggest that sexual isolation evolved as by‐product of allopatric divergence.     

A complex mode of aggressive mimicry in a scale-eating cichlid fish

Aggressive mimicry is an adaptive tactic of parasitic or predatory species that closely resemble inoffensive models in order to increase fitness via predatory gains. Although similarity of distantly related species is often intuitively implicated with mimicry, the exact mechanisms and evolutionary causes remain elusive in many cases. Here, we report a complex aggressive mimicry strategy in Plecodus straeleni, a scale-eating cichlid fish from Lake Tanganyika, which imitates two other cichlid species. Employing targeted sequencing on ingested scales, we show that P. straeleni does not preferentially parasitize its models but—contrary to prevailing assumptions—targets a variety of co-occurring dissimilar looking fish species. Combined with tests for visual resemblance and visual modelling from a prey perspective, our results suggest that complex interactions among different cichlid species are involved in this mimicry system.     

Social familiarity modulates personality trait in a cichlid fish

Personality traits, such as exploration–avoidance, are expected to be adaptive in a given context (e.g. low-risk environment) but to be maladaptive in others (e.g. high-risk environment). Therefore, it is expected that personality traits are flexible and respond to environmental fluctuations, given that consistency across different contexts is maintained, so that the relative individual responses in relation to others remains the same (i.e. although the magnitude of the response varies the differences between high and low responders are kept). Here, we tested the response of male cichlid fish (Oreochromis mossambicus) to a novel object (NO) in three different social contexts: (i) social isolation, (ii) in the presence of an unfamiliar conspecific, and (iii) in the presence of a familiar conspecific. Males in the familiar treatment exhibited more exploratory behaviour and less neophobia than males in either the unfamiliar or the social isolation treatments. However, there were no overall correlations in individual behaviour across the three treatments, suggesting a lack of consistency in exploration–avoidance as measured by the NO test in this species. Moreover, there were no differences in cortisol responsiveness to an acute stressor between the three treatments. Together, these results illustrate how behavioural traits usually taken as measures of personality may exhibit significant flexibility and lack the expected consistency across different social contexts.     

Social context affects behavior,preoptic area gene expression,and response to D2 receptor manipulation during territorial defense in a cichlid fish

Social context often has profound effects on behavior, yet the neural and molecular mechanisms which mediate flexible behavioral responses to different social environments are not well understood. We used the African cichlid fish, Astatotilapia burtoni, to examine aggressive defense behavior across three social contexts representing different motivational states: a reproductive opportunity, a familiar male and a neutral context. To elucidate how differences in behavior across contexts may be mediated by neural gene expression, we examined gene expression in the preoptic area, a brain region known to control male aggressive and sexual behavior. We show that social context has broad effects on preoptic gene expression. Specifically, we found that the expression of genes encoding nonapeptides and sex steroid receptors are upregulated in the familiar male context. Furthermore, circulating levels of testosterone and cortisol varied markedly depending on social context. We also manipulated the D2 receptor (D2R) in each social context, given that it has been implicated in mediating context‐dependent behavior. We found that a D2R agonist reduced intruder‐directed aggression in the reproductive opportunity and familiar male contexts, while a D2R antagonist inhibited intruder‐directed aggression in the reproductive opportunity context and increased aggression in the neutral context. Our results demonstrate a critical role for preoptic gene expression, as well as circulating steroid hormone levels, in encoding information from the social environment and in shaping adaptive behavior. In addition, they provide further evidence for a role of D2R in context‐dependent behavior.     

Removal of both antennae influences the courtship and aggressive behaviors in male crickets

To test whether insect antennae are necessary for eliciting courtship and aggression toward appropriate partners, we antennectomized adult male crickets (Gryllus bimaculatus) and observed their behavior toward other antennectomized males and intact females. At 7 days after removal of both antennae, pairs of antennectomized males were placed together; 80% displayed courtship behavior, generating courtship song by rubbing their forewings together, toward other antennectomized males, and 20% displayed aggressive behavior. Only 45% courted intact females. No intact males courted antennectomized males, and 80% displayed aggressive behavior. All intact males courted females. The results for males with one antenna removed were essentially the same as for intact males. These findings indicate that a high proportion of male crickets with both antennae removed court other males and fail to display male-male aggression, demonstrating that removal of antennae from male crickets induces male-male courtship and that an antenna is necessary for the expression of male-male aggression. Moreover, brain serotonin (5-hydroxytryptamine; 5-HT) levels in male crickets were significantly reduced at 7 days after removal of antennae. The reduction of 5-HT was detected primarily in the central body of the brain. Thus, 5-HT in the central body of the male cricket brain may be involved in the behavioral changes.     

Wasteful carotenoid coloration and its effects on territorial behavior in a cichlid fish

The famously diverse body coloration of cichlid fish serves communicative functions in mating and social interactions including competition for resources. Here, we examined the effects of a color pattern trait—the width of a yellow bar on a black body—on territorial competition in males and females of a color variant (“Ikola”) of the Lake Tanganyika cichlid Tropheus. First, measuring integumentary carotenoid concentrations in the yellow and black body regions, we established that wider yellow bars require more carotenoids allocated to body coloration. However, we also detected high carotenoid concentrations in the black body regions (>?100 µg/g fresh skin), raising questions about the function of non-displayed integumentary carotenoids. Behavioral experiments showed that fish with wider bars were quicker to explore an unfamiliar area of the tank. In experiments including presentations of fish dummies, the bar width of ‘territorial’ dummies had no effect on the latency time which test fish took to intrude into the dummies’ territories. However, male test fish performed fewer aggressive acts against wide-barred than against narrow-barred dummy competitors. Our results suggest that intimidation by wide bars as well as correlations between bar width and explorative behavior may contribute to mediating success in territorial Tropheus “Ikola”.

     

Lateralized kinematics of predation behavior in a Lake Tanganyika scale-eating cichlid fish

Behavioral lateralization has been documented in many vertebrates. The scale-eating cichlid fish Perissodus microlepis is well known for exhibiting lateral dimorphism in its mouth morphology and lateralized behavior in robbing scales from prey fish. A previous field study indicated that this mouth asymmetry closely correlates with the side on which prey is attacked, but details of this species' predation behavior have not been previously analyzed because of the rapidity of the movements. Here, we studied scale-eating behavior in cichlids in a tank through high-speed video monitoring and quantitative assessment of behavioral laterality and kinematics. The fish observed showed a clear bias toward striking on one side, which closely correlated with their asymmetric mouth morphologies. Furthermore, the maximum angular velocity and amplitude of body flexion were significantly larger during attacks on the preferred side compared to those on the nonpreferred side, permitting increased predation success. In contrast, no such lateral difference in movement elements was observed in acoustically evoked flexion during the escape response, which is similar to flexion during scale eating and suggests that they share a common motor control pathway. Thus the neuronal circuits controlling body flexion during scale eating may be functionally lateralized upstream of this common motor pathway.     

Early learning influences species assortative mating preferences in Lake Victoria cichlid fish

The Lake Victoria 'species flock' of cichlids is puzzling because reproductive isolation often occurs in the absence of substantial ecological differences among species. Theory predicts that this cannot evolve with most genetic mechanisms for mate choice. We provide the first evidence that learning, in the form of sexual imprinting, helps maintain reproductive isolation among closely related cichlid species. Using a cross-fostering experiment, we show that young females develop a sexual preference for males of their foster mothers' species, even reversing species assortative mating preferences. We suggest that learning creates favourable conditions for reproductive isolation to evolve.     

Tidal cycling and parental behavior of the cichlid fish,Biotodoma cupido

Synopsis Parental behavior of the substrate-brooding cichlid, Biotodoma cupido, was studied in a small creek entering the lower Essequibo River, Guyana, where the freshwaters are affected by semi-diurnal tides. Physico-chemical variables of the tidal cycle were associated with the parental behavior of B. cupido. During late ebb and early flood tides, while off-spring were nest dependent, parents displayed intense aggression toward brood predators, mainly characins. At low tide, when the concentration of dissolved oxygen decreased to about 4 mg 1^–1 and that of free carbon dioxide increased to 28 mg 1^–1, parents entered a state of somnolence and brood predators vanished. Early flood tide brought an immediate and dramatic reversal of hypoxic and hypercarbic conditions and an associated renewal of aggressive and predatory activity. At very low tide, parents orally transferred the brood to a secondary nest depression in deeper water. The significance of water-level fluctuation to the evolution of this behavior, as well as that of parent-brood itineracy and the related phenomena of oral incubation and movable nests, is discussed.     

Mating tactics and male-male courtship in the lek-breeding cichlid Oreochromis mossambicus

Data are presented on the breeding behaviour of Oreochromis mossambicus under captive conditions. Males tended to synchronize their occupation of territories and breeding activities. Different male mating tactics were observed, namely establishing a breeding territory, acting as a floater, or behaving as a sneaker. The majority of spawnings observed involved dominant males and were subjected to interference from other males. Males were found to court other males that frequently responded to these attempts by adopting a female-like behaviour. Results are discussed in terms of a probable time constraint in territoriality, which promotes male-male competition and a low level of sex discrimination by territorial fish.     

Depth equilibration of a shallow-water cichlid fish

The cichlid Melanochromis exasperatus has a natural distribution which is restricted to a single island in Lake Malawi, Africa. Diving observations indicated that it does not occur below 4.5 m depth. Measurements of depth equilibration in a pressure tank showed that males can equilibrate to a depth of 25 m and females to 18 m. Mean maximum sustained rate of depth equilibration was 2 m day⁻¹. M. exasperatus survived a pressure decrease of 40% every 12 h from the maximum equilibration depth to the surface. The fish swim actively during the day but rest on the bottom at night when they are equilibrated to a depth 1 m deeper than ambient. Reduced activity at night is reflected in a drop in oxygen consumption to nearly half of the daytime active value.     

Differences in aggressive behavior between convict cichlid color morphs: amelanistic convicts lose even with a size advantage

Convict cichlids (Archocentrus nigrofasciatus) are a territorial, monogamous, and biparental Central American cichlid fish. Convicts exist in two common color morphs: the wild-type (WT) black-barred form and an amelanistic (AM) barless morph. Color morphs affect aggressive interactions in other species of fish. We staged fights between males of each color morph with varying size asymmetries and found that WT males were able to overcome a size disadvantage by increasing their rate of aggressive behavior. AM males lost more often when smaller than their opponent, apparently because they did not increase their rate of aggressive behavior when at a size disadvantage. We discuss two possible hypotheses to explain these findings: (1) that there are genetic differences in aggressive behavior between the morphs and (2) that AM fish are disadvantaged in staged contests because they are unable to signal via changes in bar coloration.