Sharing is living: The role of habitat heterogeneity in the coexistence of closely related species.

In biologically diverse ecosystems, an essential process to support competing species to coexist is ecological differentiation. Habitat heterogeneity is, hence, important in establishing species abundance and richness, favoring the coexistence of species due to habitat partition. In this context, shading and species thermal tolerance can be good factors to elucidate the role of habitat heterogeneity in the habitat partition among closely related species. Herein, we study shading effects in microhabitat selection, behavior, and physiological limitation on two species of fiddler crabs (Leptuca leptodactyla and Leptuca uruguayensis). Indeed, shading conditions influenced fiddler crabs species proportion over time, with L. leptodactyla more associated with nonshaded/warmer areas while the L. uruguayensis to shaded/cooler ones. They also adjusted their behavior differently from each other to deal with thermal stress. Finally, we have demonstrated that these effects are related to species' physiological limitations. We conclude that biologically diverse ecosystems, such as intertidal regions from estuaries (e.g., mudflats and mangroves), support the coexistence between closely related species by reducing competition due to habitat partition.

Innate response based on visual cues of sympatric and allopatric predators in Nile tilapia.

Predators are an important selective pressure for prey, and responses to visual exposure to non-predators and to allopatric and sympatric predators may assist the understanding of how prey animals recognize and distinguish potential threats. Here, we visually exposed predator-naïve Nile tilapia (Oreochromis niloticus) to aquaria with only water (handling-negative control), a non-predator fish (Cyprinus carpio), a South American catfish (Pseudoplatystoma coruscans, allopatric predator), and an African catfish (Clarias gariepinus; sympatric predator). We found that Nile tilapia displayed antipredator responses to both predator catfish but not to non-predator fish or negative control conditions. These antipredator responses, however, were stronger to the sympatric catfish in comparison to the allopatric animal. We concluded that Nile tilapia is able to innately distinguish between a sympatric and an allopatric predator based on visual cues. We assume that the innate recognition of the allopatric predator could be attributed to similarities in the body morphology and movement (key stimuli) of the South American and African catfish. Although this is plausible, the mechanism of the innate allopatric recognition remains unknown and deserves future investigation.

Feeding regimen modulates zebrafish behavior.

Here we show that the feeding regimen modulates zebrafish (Danio rerio) behavior. With regard to the time elapsed between feeding and behavioral evaluation, fish fed 3 h before behavioral evaluation in the novel tank test (NTT) showed decreased activity and a trend toward an anxiolytic reaction (increased use of the upper section of the aquarium) in comparison to fish fed 0.5, 6, 12, 24 or 48 h before testing, although differences were not statistically significant for all comparisons. Activity and use of the upper section of the aquarium did not differ significantly among the other treatments. Regarding feeding frequency, fish fed once a day showed higher anxiety-like behavior (decreased use of the upper section of the aquarium) in comparison to fish fed twice a day, but feeding four or six times per day or only every second day did not result in differences from feeding twice a day. Feeding frequency had no effect on activity level. Metabolically, fish fed once a day presented decreased levels of glucose and glycogen and increased lactate when compared to the regular feeding (fish fed twice a day), suggesting that feeding regimen may modulate carbohydrate metabolism. Mechanistically, we suggest that the metabolic changes caused by the feeding regimen may induce behavioral changes. Our results suggest that the high variability of the results among different laboratories might be related to different feeding protocols. Therefore, if issues pertaining to the feeding regimen are not considered during experiments with zebrafish, erroneous interpretations of datasets may occur.

Modulation of Cortisol Responses to an Acute Stressor in Zebrafish Visually Exposed to Heterospecific Fish During Development.

Studies regarding predator-prey relationships have often focused on prey assessment and the responses to predation risk, but few have verified the relationship in the presence or absence of visual contact with a predator (e.g., tiger oscar, Astronotus ocellatus) or a nonpredator (e.g., goldfish, Carassius auratus) during the developmental phase, which could alter several physiological and neuroendocrine mechanisms in adulthood. Herein, we determined responses to physical (chasing with a net) and biological stressors (visualization to predator) in adult zebrafish raised in visual contact with a predator or nonpredator fish. We demonstrated that adult naive zebrafish show a more intense cortisol stress response than fish housed in visual contact with the stimulus fish (predator or nonpredator) when larvae, and that this alteration is related with movement specificity of the stimulus fish.

Chemical communication of predation risk in zebrafish does not depend on cortisol increase.

We investigated chemical cues among groups of zebrafish (Danio rerio) when communicating information about the risk of predation. We found that visual cues of the predator (tiger Oscar, Astronotus ocellatus) did not increase whole-body cortisol levels in groups of zebrafish but that water conditioned by these (donor) zebrafish stressed (target) conspecifics, thereby increasing whole-body cortisol. This finding was confirmed when these zebrafish groups were in different aquaria and communicated exclusively via water transfer. This result indicates that the stress induced in the target zebrafish does not depend on an increase in whole-body cortisol levels in the donor zebrafish. Because cortisol participation is rejected in this predation-risk communication, other chemicals from the stress systems should be investigated.

Stomach fullness modulates prey size choice in the frillfin goby, Bathygobius soporator.

Behaviours related to foraging and feeding in predator-prey systems are fundamental to our understanding of food webs. From the perspective of a predator, the selection of prey size depends upon a number of factors including prey vulnerability, prey size, and the predator's motivation to eat. Thus, feeding motivation and prey visual cues are supposed to influence predator decisions and it is predicted that prey selection by visual cues is modulated by the predator's stomach fullness prior to attacking a prey. This study was conducted using an animal model from the rocky shores ecosystem, a predatory fish, the frillfin goby Bathygobius soporator, and a benthic prey, the mottled shore crab Pachygrapsus transversus. Our results demonstrate that frillfin gobies are capable of visually evaluating prey size and that the size evaluation process is modulated by the level of stomach fullness. Predators with an empty stomach (0% fullness) attacked prey that was larger than the predicted optimal size. Partially satiated predators (50% stomach fullness) selected prey close to the optimal size, while fully satiated predators (100% stomach fullness) showed no preference for size. This finding indicates an integrative response of the predator that depends on the input of both internal and external sensory information when choosing prey. Predator perceptions of visual cues (prey size) and stomach fullness modulate foraging decisions. As a result, a flexible feeding behaviour emerges, evidencing a clearly adaptive response in line with optimal foraging theory predictions.

Ventilation rates indicate stress-coping styles in Nile tilapia.

Behavioural responses to stress can form distinct profiles in a wide range of animals: proactive and reactive profiles or coping styles. Stress responsiveness can also differentiate between the behavioural profiles. The tendency to regain feed intake following transfer to a novel social-isolation tank (the speed of acclimation) can discriminate between proactive or reactive profiles. Consequently, differential stress responsiveness can be linked to this feeding behaviour trait. This study shows that ventilation rates of Nile tilapia, Oreochromis niloticus (L.), correlate with the rate of feeding resumption, following transfer to a novel social-isolation aquarium. Therefore, ventilation rate (VR) indicates coping styles; consequently, VR is a proxy for the way fish will deal with environmental challenges.

The correlation between subordinate fish eye colour and received attacks: a negative social feedback mechanism for the reduction of aggression during the formation of dominance hierarchies.

Eye darkening has been linked to social status in fish. The subordinate's eyes darken, while the eyes of the dominant fish become pale. Although this phenomenon has been described in salmonid fishes and in the African cichlid Nile tilapia Oreochromis niloticus, it is unclear whether eye darkening correlates with a reduction in aggressive behaviour. Thus, we evaluated the link between social status and eye darkening. We evaluated whether the eye colours of subordinate fish correlate with the frequency of received attacks in a neotropical fish, the pearl cichlid Geophagus brasiliensis. We paired pearl cichlids and quantified both the aggressive behaviour and the eye darkening of each fish. As has been described for Nile tilapia and Atlantic salmon, a clear-cut hierarchical relationship formed, where dominance and subordination were associated with pale and dark eye colours, respectively. Initially, eye colour darkening was positively correlated with the frequency of received attacks; however, a negative association occurred following eye darkening, in which the intensity of aggressive interactions decreased. Thus, fish that initially received a high number of attacks signalled subordination more rapidly and intensely (rapid and dramatic eye darkening), thereby inducing a negative social feedback mechanism that led to reduced aggression.

The aggressive behavior of Nile tilapia introduced into novel environments with variation in enrichment.

Many studies show environmental enrichment is correlated with benefits to captive animals; however, one should not always assume this positive relationship given that enrichment increases the amount of resources that a territorial animal must defend and possibly affects its aggressive dynamics. In this study, we tested if environmental enrichment affects aggressive interactions in the aggressive fish Nile tilapia (Oreochromis niloticus). We compared fights staged between pairs of male tilapia of similar size (= matched in resource holding potential) in a novel arena that was either barren or enriched, to examine whether enrichment enhances territory value in line with theoretical predictions, with the potential for compromised welfare. We evaluated time elapsed until the first attack (latency), frequency of aggressive interactions and fight duration. We detected fight dynamic differences at the pair level. Higher resource value generated increased aggression but had no effect on fight duration or latency. This conclusion is in line with game theory predictions concerning resource value and contradicts the theory that enrichment of the environment will serve welfare purposes.

Oxidative stress biomarkers and aggressive behavior in fish exposed to aquatic cadmium contamination

The objective of this study was to investigate the possible link between cadmium exposure, hepatic markers of oxidative stress and aggressive behavior in Nile tilapia (Oreochromis niloticus). Fish were first exposed to 0.75 mg/L CdCl2 for 15 days (12 isolated fish for each group) and afterward a behavioral test was performed. Fish from the control and cadmium-exposed groups were paired for 1 h (6 pairs of fish per group) for determination of aggressiveness parameters. Immediately after the behavioral test, the animals were sacrificed and the liver was used to determine biochemical parameters. Cadmium decreased aggression in Nile tilapia. Subordinate animals exposed to cadmium showed decreased glutathione peroxidase (GSH-Px) activity compared to dominant ones. No alterations were observed in selenium-dependent glutathione peroxidase Se-GSH-P and Cu-Zn superoxide dismutase activities, but total superoxide dismutase activity was increased in subordinate animals exposed to cadmium compared to subordinate control. Catalase activity was increased in cadmium-exposed fish. Lipoperoxide concentrations also increased in cadmium exposed fish indicating that cadmium toxicity may affect oxidative stress biomarkers in Nile tilapia. Social stress induced lipoperoxidation in Nile tilapia, and subordinate animals exposed to cadmium responded with lower activities of liver antioxidant enzymes compared to dominant fish. The present study shows that cadmium exposure is capable of inducing changes in the social status and oxidative stress parameters in this species.(AU)

O objetivo deste estudo foi investigar a possível relação entre a exposição ao cádmio, os marcadores hepáticos de estresse oxidativo e o comportamento agressivo em tilápia do Nilo (Oreochromis niloticus). Os peixes foram primeiramente expostos à concentração de 0,75 mg/L de CdCl2 durante 15 dias (12 peixes isolados em cada grupo), após o qual o teste de comportamento foi aplicado. Peixes do grupo controle e do grupo exposto ao cádmio foram pareados durante 1 hora (6 pares de peixes por grupo) para determinação de parâmetros de agressividade. Imediatamente após o teste comportamental, os animais foram sacrificados e o fígado foi coletado para as determinações bioquímicas. O cádmio diminuiu a agressão em tilápias do Nilo. Animais subordinados e expostos ao cádmio mostraram uma redução na atividade da enzima glutationa peroxidase (GSH-Px), quando comparados aos dominantes. Nenhuma alteração foi observada na atividade da glutationa peroxidase dependente de selênio (Se-GSH-P) e na superóxido dismutase Cu Zn, mas a atividade total da superóxido dismutase aumentou em animais subordinados e expostos ao cádmio, quando comparado ao controle subordinado. A atividade da catalase aumentou em peixes expostos ao cádmio. As concentrações de lipoperóxido aumentaram em peixes expostos ao cádmio, indicando que a toxicidade deste pode afetar os biomarcadores de estresse oxidativo em tilápias do Nilo. O estresse social induziu lipoperoxidação em tilápia do Nilo e animais subordinados e expostos ao cádmio responderam com baixas atividades de enzimas antioxidantes hepáticas do que em peixes dominantes. O presente estudo mostra que a exposição ao cádmio é capaz de induzir alterações no status social e nos parâmetros de estresse oxidativo nesta epécie.(AU)