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In this review, we consider the definitions and experimental approaches to emotional contagion and prosocial behaviour in mammals and explore their evolutionary conceptualisation for studying their occurrence in the evolutionarily divergent vertebrate group of ray-finned fish. We present evidence for a diverse set of fish phenotypes that meet definitional criteria for prosocial behaviour and emotional contagion and discuss conserved mechanisms that may account for some preserved social capacities in fish. Finally, we provide some considerations on how to address the question of interdependency between emotional contagion and prosocial response, highlighting the importance of recognition processes, decision-making systems, and ecological context for providing evolutionary explanations. • Emotional contagion can explain ancestral forms of prosocial behaviour in fish. • The testable basis is state transfer triggering appropriate benefit from others' acts. • Recognition processes are a paramount component to elucidating this link. • Comparative approaches can reveal evolutionary substrates for higher-order functions. [ABSTRACT FROM AUTHOR]

In pretty much any species, an individual's survival and reproduction depends crucially on the outcome of interactions with other individuals. Key interactions may take place between individuals of the same species but also between individuals belonging to different species. However, the most accepted definition of social behavior only considers interactions between conspecifics. Here, we argue that the distinction between intra‐ and interspecific interactions is largely artificial and hinders the integration of the historically separately developed concepts. At the ultimate level, given that the ecological landscape of organisms is composed both by interactions with conspecifics and with heterospecifics, and both types of interactions may have evolutionary consequences. Although intraspecific interactions usually have a higher impact in fitness because in most species interactions relevant for reproduction (mating, parenting) exclusively involve conspecifics, and interactions relevant for survival are more probable between conspecifics because they share the same ecological niche, hence competing for the same resources (e.g., food, shelter), there are notable exceptions in both fitness components (e.g., heterospecific mating in parthenogenic all‐female species; heterospecific brood parasitism; heterospecific aggression in sympatric species that compete for shared resources). At the proximate level, behaviors and cognitive decision‐making rules used to interact with other organisms may be shared between intra‐ and interspecific interactions, and the mechanistic differences between conspecific social behaviors used in distinct functional domains, such as mating, aggression, or parenting, can be more expressive than those found within the same functional domain between conspecific and heterospecific behavior. Therefore, there are neither fundamental conceptual (ultimate) reasons, nor key differences in mechanisms underlying behaviors involved in conspecific vs. heterospecific interactions that support the exclusion of interspecific interactions from the conceptual framework of social behavior. [ABSTRACT FROM AUTHOR]

The alarm substance in fish is a pheromone released by injured individuals after a predator attack. When detected by other fish, it triggers fear/defensive responses, such as freezing and erratic movement behaviours. Such responses can also help other fish in the shoal to modulate their own behaviours: decreasing a fear response if conspecifics have not detected the alarm substance (social buffering) or triggering a fear response if conspecifics detected the alarm substance (social contagion). Response variation to these social phenomena is likely to depend on sex. Because males have higher-risk life-history strategies than females, they may respond more to social buffering where they risk not responding to a real predator attack, while females should respond more to social contagion because they only risk responding to a false alarm. Using zebrafish, we explored how the response of males and females to the presence/absence of the alarm substance is modified by the alarmed/unalarmed behaviour of an adjacent shoal of conspecifics. We found that, in social buffering, males decreased freezing more than females as expected, but in social contagion males also responded more than females by freezing at a higher intensity. Males were, therefore, more sensitive to visual information, while females responded more to the alarm substance itself. Because visual information updates faster than chemical information, males took more risks but potentially more benefits as well, because a quicker adjustment of a fear response allows to save energy to other activities. These sex differences provide insight into the modifying effect of life-history strategies on the use of social information. [ABSTRACT FROM AUTHOR]

Group living animals must be able to express different behavior profiles depending on their social status. Therefore, the same genotype may translate into different behavioral phenotypes through socially driven differential gene expression. However, how social information is translated into a neurogenomic response and what are the specific cues in a social interaction that signal a change in social status are questions that have remained unanswered. Here, we show for the first time, to our knowledge, that the switch between statusspecific neurogenomic states relies on the assessment of fight outcome rather than just on self- or opponent-only assessment of fighting ability. For this purpose, we manipulated the perception of fight outcome in male zebrafish and measured its impact on the brain transcriptome using a zebrafish whole genome gene chip. Males fought either a real opponent, and a winner and a loser were identified, or their own image on a mirror, in which case, despite expressing aggressive behavior, males did not experience either a victory or a defeat. Massive changes in the brain transcriptome were observed in real opponent fighters, with losers displaying both a higher number of differentially expressed genes and of coexpressed gene modules than winners. In contrast, mirror fighters expressed a neurogenomic state similar to that of noninteracting fish. The genes that responded to fight outcome included immediate early genes and genes involved in neuroplasticity and epigenetic modifications. These results indicate that, even in cognitively simple organisms such as zebrafish, neurogenomic responses underlying changes in social status rely on mutual assessment of fighting ability. [ABSTRACT FROM AUTHOR]

Androgens respond to social challenges and this response has been interpreted as a way for males to adjust androgen-dependent behavior to social context. However, the androgen responsiveness to social challenges varies across species and a conceptual framework has been developed to explain this variation according to differences in the mating system and parental care type, which determines the regimen of challenges males are exposed to, and concomitantly the scope (defined as the difference between the physiological maximum and the baseline levels) of response to a social challenge. However, this framework has been focused on territorial species and no clear predictions have been made to gregarious species (e.g. shoaling fish), which although tolerating same-sex individuals may also exhibit intra-sexual competition. In this paper we extend the scope of this conceptual framework to shoaling fish by studying the endocrine response of zebrafish ( Danio rerio ) to social challenges. Male zebrafish exposed to real opponent agonistic interactions exhibited an increase in androgen levels (11-ketotestosterone both in Winners and Losers and testosterone in Losers). This response was absent in Mirror-fighters, that expressed similar levels of aggressive behavior to those of winners, suggesting that this response is not a mere reflex of heightened aggressive motivation. Cortisol levels were also measured and indicated an activation of the hypothalamic–pituitary–interrenal axis in Winners of real opponent fighters, but not Losers or in Mirror-fighters. These results confirm that gregarious species also exhibit an endocrine response to an acute social challenge. [ABSTRACT FROM AUTHOR]

Animals communicate through the exchange of signals. However, third-party individuals can detect and intercept signals not directly sent to them, a phenomenon known as eavesdropping, and the presence of bystanders can influence the signalling behaviour of interacting conspecifics, a phenomenon named the audience effect. So far, research done on audience effects and eavesdropping has been mainly focused on their function, rather than on their proximate mechanisms. For this reason, we were interested in testing the occurrence of audience effects on male zebrafish, a genetically tractable model organism that is emerging as a major candidate for the study of the neural basis of social behaviour. Here, pairs of males were exposed to a mixed-sex shoal, which was used as an audience, at two different times: (1) during a contest between them, to test for an audience effect and (2) before the contest, to test whether this prior exposure influences subsequent agonistic behaviour (i.e. aggressive priming). We analysed the pairs' aggressive signalling during the contest by measuring variables that characterize both the individuals' behaviour and the interaction, and found that pre-exposure to an audience induced a shorter latency to display, an increase in the time dominants spent chasing subordinates and a shorter time to resolve the agonistic interaction. Also, exposure to the audience during the interaction led to a higher number of interactions in which displays occurred, a higher number of resolved interactions with displays and a decrease in the escalation of aggression for resolved interactions. These results add zebrafish to the literature on the audience effect and, most importantly, open the way for the study of the neural mechanisms involved in the processing of social information in a model organism. [ABSTRACT FROM AUTHOR]

The nonapeptide arginine vasotocin (AVT) and its mammalian homologue arginine vasopressin play a key role in the regulation of social behaviour across vertebrates. In teleost fishes, three AVT neuronal populations have been described in the preoptic area (POA): the parvocellular (pPOA), the magnocellular (mPOA) and the gigantocellular (gPOA). Neurons from each of these areas project both to the pituitary and to other brain regions, where AVT is supposed to regulate neural circuits underlying social behaviour. However, in the fish species studied so far, there is considerable variation in which AVT neuronal populations are involved in behavioural modulation and in the direction of the effect. In this study, the association between AVT neuronal phenotypes and social status was investigated in the Mozambique tilapia (Oreochromis mossambicus). This species is an African female mouth-brooding cichlid fish in which males form breeding aggregations in which dominant males establish territories and subordinate males to act as floaters. With respect to sex differences in AVT neuronal phenotypes, females have a larger number of AVT neurons in the pPOA and mPOA. Within males, AVT appeared associated with social subordination, as indicated by the larger cell body areas of AVT neurons in mPOA and gPOA nuclei of non-territorial males. There were also positive correlations between submissive behaviour and the soma size of AVT cells in all three nuclei and AVT cell number in the mPOA. In summary, the results provide evidence for an involvement of AVT in the modulation of social behaviour in tilapia, but it was not possible to identify specific roles for specific AVT neuronal populations. The results presented here also contrast with those previously published for another cichlid species with a similar mating system, which highlights the species-specific nature of the pattern of association between AVT and social behaviour even within the same taxonomic family. © 2015 S. Karger AG, Basel [ABSTRACT FROM AUTHOR]

Vasotocin (VT) has been associated with the regulation of different aspects of social behavior (e.g., mating and aggression). Given the fact that androgens are also known to regulate reproductive behavior, we hypothesized that VT and androgens could be interacting, rather than acting independently, in the regulation of reproductive behavior. In the present study, we aimed to understand the effect of VT and its interaction with gonadal hormones (putatively androgens) on different aspects of reproductive behavior of a polygynous and territorial cichlid fish, the Mozambique tilapia (Oreochromis mossambicus). Using a within-subject design, we treated territorial males, that were previously castrated or sham-operated, with different dosages of VT as well as with a V1A receptor antagonist (Manning compound) and subsequently analyzed their behavior towards females and towards an intruder male. Our results showed that VT affected the behavior of territorial males towards females but not towards males. Specifically, VT-treated males interacted less with females (i.e., spent less time touching the transparent partition that allowed visual contact with females) and were less aggressive towards females than saline-treated males. Moreover, in sham-operated males, blocking V1A receptors increased the frequency of bites towards females in comparison to saline-treated males, but not in castrated males. This result suggests that VT down-regulates aggressiveness towards females through the action of V1A receptors in the gonads (putatively decreasing androgen secretion), and that androgens up-regulate this behavior. In summary, our results suggest that VT may modulate social behavior, through an interaction with gonadal hormones. [ABSTRACT FROM AUTHOR]

Highlights: [•] We examined female mate choice mechanisms in a captive neotropical passerine. [•] Male attributes were less important to females than novelty in preference trials. [•] Females behaved aggressively towards familiar males when novel ones were available. [•] Males were unresponsive behaviourally to female contact with novel male. [•] They also did not respond physiologically to threat of novel males. [ABSTRACT FROM AUTHOR]

‘Social competence’ refers to the ability of an individual to optimise its social behaviour depending on available social information. Although such ability will enhance social interactions and thus raise Darwinian fitness, its evolutionary and ecological significance has been largely ignored. Social competence is based on behavioural flexibility. We propose that the study of social competence requires an integrative approach that aims to understand how the brain translates social information into flexible behavioural responses, how flexibility might be constrained by the developmental history of an individual or by trade-offs with other (ecological) competences, and how social plasticity feeds back on fitness. Finally we propose a hypothesis of how social competence can become a driver of social evolution. [Copyright &y& Elsevier]

Abstract: Studies on birds and mammals indicate that sexual traits may signal superior health because active immunity, like inflammatory responses to infections, is suppressive to the production of androgens that facilitate the expression of these traits. Here we test this possible pathway for honest signaling in a teleost species, Sarotherodon galilaeus, by activating the immune system with sheep red blood cells (SRBC), which is a non-pathogenic T- and B-cell stimulating antigen. Two weeks after the start of treatment adult males injected with SRBC showed a significant increase in antibody production in comparison with control males. The variation in specific antibody production was negatively related with variation in both testosterone and 11-ketotestosterone levels. This suggests that investment in immune protection is incompatible with increased activity of the hypothalamic-pituitary-gonadal axis. However, opposite to our expectation no difference in androgen levels was found between placebo and SRBC treatment suggesting that immune activation did not cause androgen suppression in our studied species. [Copyright &y& Elsevier]

There is a wealth of game theoretical approaches to the evolution and maintenance of cooperation between unrelated individuals and accumulating empirical tests of these models. This contrasts strongly with our lack of knowledge on proximate causes of cooperative behaviour. Marine cleaning mutualism has been used as a model system to address functional aspects of conflict resolution: client reef fish benefit from cleaning interactions through parasite removal, but cleaner fish Labroides dimidiatus prefer client mucus. Hence, feeding against their preference represents cooperative behaviour in cleaners. Cleaners regularly cheat non-predatory clients while they rarely cheat predatory clients. Here, we asked how precisely cleaners can adjust service quality from one interaction to the next. We found that non-predatory clients receive a better service if the previous client was a predator than if the previous client was a non-predator. In a related laboratory experiment, a hand-net used as a stressor resulted in cleaners feeding more against their preference in subsequent interactions. The combination of the cleaners' behaviour in the two studies shows that the cleaners' service quality for a given client species is not fixed, but it can be manipulated. The results suggest that short-term stress is one factor that causes cleaners to increase their levels of cooperation, a hypothesis that is amenable to further experiments manipulating the endocrine system. [ABSTRACT FROM AUTHOR]

The ability to respond to stress is vital to the survival of any living organism, though sustained reactions can become detrimental to the health and welfare of animals. Stress responses of vertebrates are known through several studies in their physiological, behavioural and psychological components, under acute and chronic contexts. In fish, the physiological and behavioural aspects of stress are considerably well known phenomena and show striking similarities to those of other vertebrates. However, the psychological component is not well known. Some authors deny mental experiences to fish on the basis of their lack of neocortex. Nevertheless, recent studies have shown neuroendocrine, cognitive and emotional processes in fish that are not only equivalent to other vertebrates, but also allow inferring some forms of mental representation. The integration of psychological elements in fish stress physiology is insufficiently studied, but, as discussed in this article, there is already indirect evidence to admit that some form of stimuli appraisal can take place in fish. This fact has profound implications on the regulation of the stress response, as well as on fish welfare and its management. [ABSTRACT FROM AUTHOR]

In social species animals should fine-tune the expression of their social behavior to social environments in order to avoid the costs of engaging in costly social interactions. Therefore, social competence, defined as the ability of an animal to optimize the expression of its social behavior as a function of the available social information, should be considered as a performance trait that impacts on the Darwinian fitness of the animal. Social competence is based on behavioral plasticity which, in turn, can be achieved by different neural mechanisms of plasticity, namely by rewiring or by biochemically switching nodes of a putative neural network underlying social behavior. Since steroid hormones respond to social interactions and have receptors extensively expressed in the social behavioral neural network, it is proposed that steroids play a key role in the hormonal modulation of social plasticity. Here, we propose a reciprocal model for the action of androgens on short-term behavioral plasticity and review a set of studies conducted in our laboratory using an African cichlid fish (Oreochromis mossambicus) that provide support for it. Androgens are shown to be implicated as physiological mediators in a wide range of social phenomena that promote social competence, namely by adjusting the behavioral response to the nature of the intruder and the presence of third parties (dear enemy and audience effects), by anticipating territorial intrusions (bystander effect and conditioning of the territorial response), and by modifying future behavior according to prior experience of winning (winner effect). The rapid behavioral actions of socially induced short-term transient changes in androgens indicate that these effects are most likely mediated by nongenomic mechanisms. The fact that the modulation of rapid changes in behavior is open to the influence of circulating levels of androgens, and is not exclusively achieved by changes in central neuromodulators, suggests functional relevance of integrating body parameters in the behavioral response. Thus, the traditional view of seeing neural circuits as unique causal agents of behavior should be updated to a brain–body–environment perspective, in which these neural circuits are embodied and the behavioral performance (and outcomes as fitness) depends on a dynamic relationship between the different levels. In this view hormones play a major role as behavioral modulators. [ABSTRACT FROM PUBLISHER]

In many territorial species androgens respond to social interactions. This response has been interpreted as a mechanism for adjusting aggressive motivation to a changing social environment. Therefore, it would be adaptive to anticipate social challenges and reacting to their clues with an anticipatory androgen response to adjust agonistic motivation to an imminent social challenge. Here we test the hypothesis of an anticipatory androgen response to territorial intrusions using classical conditioning to establish an association between a conditioned stimulus (CS = light) and an unconditioned stimulus (US = intruder male) in male cichlid fish (Oreochromis mossambicus). During the training phase conditioned males (CS-Us paired presentations) showed a higher decrease in latency for agonistic response toward the intruder than unconditioned males (Cs-Us unpaired presentations). In the test trial, conditioned males showed an increase in androgen levels (i.e., testosterone and illketotestosterone) relative to baseline, in response to the Cs alone. This increase was similar to that of control males exposed to real intruders after Cs, whereas unconditioned males showed a decrease in androgen levels in response to the Cs. Furthermore, conditioned males were significantly more aggressive than unconditioned males during the post-CS period on test trial, even though the intruder male was not present during this period. These results reveal the occurrence of a conditioned androgen response that may give territorial males an advantage in mounting a defense to upcoming territorial intrusions, if the ability to readily elevate androgens does not co-vary with other traits that bear costs. [ABSTRACT FROM AUTHOR]

Species with alternative reproductive tactics offer the opportunity to analyse how behavioural and morphological traits are tuned to produce successfully competing phenotypes within one sex. In the teleost fish Salaria pavo, alternative reproductive tactics are sequential. The older ornamented males compete for access to females by guarding a cavity to which they attract females to spawn. Ornamented males that are found without a nest are called ‘floaters’. Younger mature males which are too small to compete with nest-holders attempt to ‘sneak’ as female mimics into successful nests and release their sperm to fertilize freshly spawned eggs. The alternative tactics in S. pavo are associated with different levels of the androgens testosterone and 11-ketotestosterone, which have been found to suppress immune function in several teleost fish. A field study was carried out to analyse the relationship between these reproductive tactics, androgens and blood levels of lymphocytes as a monitoring method of immune function. We expected highest investment in the immune system in sneakers because these have the lowest androgen levels and functionally because investing in self-maintenance increases their future prospect to switch tactic and to reproduce as a nest-holder. Indeed, the relative count of lymphocytes correlated negatively with body size and thus was highest in sneakers and lowest in nest-holders. In concordance, 11-ketotestosterone and testosterone levels were found to be higher in floaters and nest-holders than in sneakers. However, no correlation was found between individual levels of testosterone or 11-ketotestosterone and lymphocytes. Thus, a trade-off between reproductive traits associated with high androgen levels and immunocompetence might exist at the level of alternative tactics but this might not be explained by acute immunosuppressive effects of circulating androgens. [ABSTRACT FROM AUTHOR]

Background: Marine cleaning interactions in which cleaner fish or shrimps remove parasites from visiting 'client' reef fish are a textbook example of mutualism. However, there is yet no conclusive evidence that cleaning organisms significantly improve the health of their clients. We tested the stress response of wild caught individuals of two client species, Chromis dimidiata and Pseudanthias squamipinnis, that had either access to a cleaner wrasse Labroides dimidiatus, or to cleaner shrimps Stenopus hispidus and Periclimenes longicarpus, or no access to cleaning organisms. Results: For both client species, we found an association between the presence of cleaner organisms and a reduction in the short term stress response of client fish to capture, transport and one hour confinement in small aquaria, as measured with cortisol levels. Conclusion: It is conceivable that individuals who are more easily stressed than others pay a fitness cost in the long run. Thus, our data suggest that marine cleaning mutualisms are indeed mutualistic. More generally, measures of stress responses or basal levels may provide a useful tool to assess the impact of interspecific interactions on the partner species. [ABSTRACT FROM AUTHOR]

The challenge hypothesis (Wingfield et al. 1990, American Naturalist, 136, 829–846) predicts varying androgen responses to mating, breeding or territorial behaviour in avian males. At the interspecific level, the highest androgen responsiveness has been observed in males from monogamous species with paternal incubation, and the lowest in males from promiscuous, nonpaternal species. Studies of a number of vertebrate species have discussed the extension of the challenge hypothesis predictions to nonavian vertebrates, but a general ‘vertebrate consensus’ has not been achieved. For this quantitative review we included data from all vertebrate species available in the literature into several meta-analyses. We distinguished between the effects of androgens on sexual, aggressive and paternal behaviour and the effects of behaviour on androgen levels and compared these effects between taxa, mating systems and types of parental care. We observed large variations between taxa in all data sets. Nevertheless, at the vertebrate level the challenge hypothesis predictions originating from the avian literature were confirmed for the modulation of androgen responsiveness (1) to sexual behaviour by paternal care but not by mating system, and (2) to paternal behaviour by mating system but not the degree of paternal care. In contrast, our results provide (3) no support for the predicted modulation of androgen levels in response to agonistic interactions by mating and parental care system at the vertebrate level. Furthermore, our meta-analyses suggest that the effects of exogenous testosterone on sexual and agonistic behaviour, as a rule of thumb, may be expected to be larger in nonpaternal than in paternal systems. [Copyright &y& Elsevier]

Abstract: The occurrence of intra-sexual variation in reproduction is a widespread phenomenon in teleosts. One such form of variation consists in the occurrence of alternative male types: males that invest resources in mate attraction and males that exploit the investment of the former males, by trying to sneak fertilizations during spawning. These alternative reproductive tactics can be classified according to their plasticity during the life span of the individuals (i.e., fixed vs. sequential vs. reversible). Furthermore, the differences between morphs within a given species may involve a set of different traits, including reproductive behavior, the differentiation of male morphological traits, and the patterns of gonad tissue allocation and the differentiation of gonadal accessory glands. In this paper, we review the available data on four species exhibiting different types of intra-sexual plasticity in reproduction that have been studied in our lab. The data on the proximate mechanisms, androgens and forebrain arginine-vasotocin (AVT), underlying these alternative tactics suggest that between-morph differences in androgen levels, especially in 11-ketotestosterone, are especially present in species where the alternative male types have evolved morphological traits that are tactic-specific (i.e., sexual polymorphisms) and that differences in AVT appear to be related to between-morph differences in the expression of courtship behavior. Therefore, this comparative approach leads us to propose that the different endocrine systems are involved in the differentiation of different sets of traits that make up alternative phenotypes, and that the differentiation of alternative tactics is not controlled by a single endocrine system (e.g., androgens). [Copyright &y& Elsevier]

In most species females do not exhibit conspicuous sexual behaviours and female mate preferences are often measured by means of the time spent close to males. In spite of its widespread use, in only a few studies has this measure been validated as a reliable indicator of female mate preference. Sex-role-reversed species offer a better opportunity to test female preferences, as females usually court males. We tested in a sex-role-reversed population of the blenny Salaria pavo (Risso) if the time spent by females close to males related to the number of courtship displays directed towards males, and whether these measures of female preference reliably predicted matings. Females were simultaneously presented with two males behind a glass partition and the time spent close to each male was measured. We then allowed females to have access to the males' compartments and measured the courtship behaviours performed by females towards each male and recorded with which of the males females spawned. Females spent more time close to the male that subsequently received more courtship displays, and the preferred male had a more developed head crest. However, both measures of female preference failed to predict matings. Females were often attacked by males and probably had, in some occasions, to spawn with the less preferred male. Females that spawned with the previously preferred male had more swollen bellies than females that spawned with the less preferred male, suggesting a male preference towards more ripe females. These results validate the use of "time spent close to a sexual partner" as a measure of female preference in S. pavo. We argue that matings may not always reliably indicate mate preferences, particularly in the less choosy sex. [ABSTRACT FROM AUTHOR]

In the common Azorean rock-pool blenny, Parablennius parvicornis, males exhibit alternative reproductive morphologies: (1) larger males defend nest sites, provide parental care, have anal glands (involved in pheromone release), testicular glands, and low gonad:body weight ratio (GSI) and (2) smaller, younger, males do not defend nests, have reduced glands and high GSI. These smaller non-nesting males behave as satellites (associated with nests) or sneakers (moving among nests), attempting to achieve parasitic fertilizations via sperm competition. In non-mammals, arginine vasotocin (AVT) is a key hypothalamic peptide involved in the control of reproductive behavior and physiology, and several fish species that exhibit alternative male reproductive morphs show polymorphism in AVT brain chemistry. We conducted an immunocytochemical study to generate comparative data on this intertidal blenny. Our analysis showed no difference in AVT-immunoreactive cell number or size between the male morphs, which is consistent with studies on other fish, including blennies. The number of AVT cells was positively correlated to fish body mass, while cell size showed no such relation. If corrected for body mass, the smaller non-nesting males have significantly more cells than the large nesting males. Our data suggest that the size and number of forebrain AVT cells develops initially to allow for reproduction in the young non-nesting males and this pattern does not appear to change when males take on the nesting morphotype later in life. This result appears to be consistent in many fishes with alternative male morphotypes. [Copyright &y& Elsevier]

In general, animals prefer to mate with individuals they have observed being courted or in close association with sexual partners. This phenomenon of mate copying has been demonstrated in several species, but so far no study has provided substantial evidence that it is adaptive. Furthermore, mate copying has been viewed only in the context of females copying other females or males copying other males. In the peacock blenny, Salaria pavo, parasitic males (sneakers) may gain an advantage by copying the association patterns of females with bourgeois males (large males that defend nests). We tested the sneaker''s preference for one of two males and subsequently presented the nonpreferred male to the sneaker in the company of females, while the preferred male was presented alone. If the association of females with bourgeois males influences the sneaker''s preference, we predicted the sneaker would spend more time close to the nonpreferred male, when females were no longer present. We found that (1) sneakers preferred to associate with the larger of the two males and (2) when the previously nonpreferred male was presented in the company of females the sneakers tended to approach that male sooner, although not significantly so, and to spend more time close to it. Thus, parasitic males seem to choose host males both by independent mechanisms (larger males were preferred) and by nonindependent mechanisms (males observed with females were preferred). We discuss the adaptive value of sneakers choosing males by each of these mechanisms. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved. [Copyright &y& Elsevier]

Androgens are classically thought of as the sex steroids controlling male reproduction. However, in recent years evidence has accumulated showing that androgens can also be affected by the interactions between conspecifics, suggesting reciprocal interactions between androgens and behaviour. These results have been interpreted as an adaptation for individuals to adjust their agonistic motivation and to cope with changes in their social environment. Thus, male–male interactions would stimulate the production of androgens, and the levels of androgens would be a function of the stability of its social environment [‘challenge hypothesis’, Gen. Comp. Endocrinol. 56 (1984) 417]. Here the available data on social modulation of androgen levels in male teleosts are reviewed and some predictions of the challenge hypothesis are addressed using teleosts as a study model. We investigate the causal link between social status, territoriality and elevated androgen levels and the available evidence suggests that the social environment indeed modulates the endocrine axis of teleosts. The association between higher androgen levels and social rank emerges mainly in periods of social instability. As reported in the avian literature, in teleosts the trade-off between androgens and parental care is indicated by the fact that during the parental phase breeding males decreased their androgen levels. A comparison of androgen responsiveness between teleost species with different mating and parenting systems also reveals that parenting explains the variation observed in androgen responsiveness to a higher degree than the mating strategy. Finally, the adaptive value of social modulation of androgens and some of its evolutionary consequences are discussed. [Copyright &y& Elsevier]

In the Azorean rock-pool blenny, sexually active males may adopt alternative reproductive tactics. In the present paper the relationship between the presence of satellite males and the reproductive success of nest-holders was investigated by comparing nests with and without an associated satellite male. Males with an associated satellite male suffered more conspecific intrusions but they did not display a higher frequency of attacks towards conspecifics. Nest-holder males were more aggressive towards other conspecifics than towards satellites and the tolerance of nest-holders towards satellites was inversely correlated with the time spent by the satellites in the breeding territory, which suggests control by the nest-holder male of the satellite investment in shared territorial defence. Nest-holders with an associated satellite male had higher condition factors and received more female visits and more spawnings. These results bear two possible interpretations. (1) Nest-holders benefit from the presence of a satellite male by increased attractiveness of their nests to females; satellite males are mutualists helping to defend the nest-owner's territory and to attract females, which is why they are tolerated. (2) Satellite males associate preferentially with more successful nest-holder males which have higher condition factors, and by doing so have more opportunities to achieve parasitic fertilizations. Only experiments will allow these two hypotheses to be distinguished. [ABSTRACT FROM AUTHOR]

In the peacock blenny, Salaria pavo, a species with courtship sex-role reversal, smaller, younger males mimic the courtship behavior and the nuptial coloration of females in order to get access to nests during spawning and to parasitize egg fertilization from nest-holder males. Later in their life, sneakers transform both morphologically and behaviorally into nest-holder males. In the present paper we investigate the activational role of 11-ketotestosterone (KT), the most potent androgen in most teleost species, to promote the switch between tactics in sneaker males of S. pavo. Sneakers were implanted either with KT or with control (i.e. castor oil) silastic implants. A week after implantation they were subjected to a set of behavioral tests and morphometric measurements. KT treatment promoted the differentiation of secondary sex characters, such as the anal glands, and inhibited the expression of female courtship behavior. KT-treated sneakers also showed a trend toward less frequent display of female nuptial coloration. There was no effect of KT treatment on the expression of typical nest-holder male behavior. Finally, there was no effect of KT treatment on the number or soma size of arginine vasotocin neurons in the preoptic area, which are often associated with the expression of vertebrate sexual behavior. Thus, KT seems to play a key role in mating tactic switching by inhibiting the expression of female courtship behavior and by promoting the development of male displaying traits (e.g. anal glands). The lack of a KT effect on behavior typical of nest-holding males and vasotocinergic preoptic neurons suggests that a longer time frame or other endocrine/social signals are needed for the initiation of these traits in males that are switching tactics.Copyright © 2001 S. Karger AG, Basel [ABSTRACT FROM AUTHOR]

Abstract This paper presents an analysis of the distributional patterns of blenniids (Pisces: Blenniidae) in the north-eastern Atlantic. Two peaks of species diversity were found, both in terms of number of species and number of endemics: one in the tropical African coast and another in the Mediterranean Sea. A cluster analysis of similarity values (Jaccard coefficient) among the eastern Atlantic zoogeographical areas, revealed the following groups: a north temperate group, a tropical group formed by the tropical African coast and Mauritania, another group formed by the islands of Cape Verde, a south temperate group (South Africa), and a southern Atlantic group formed by the islands of Ascension and St Helena. Within the north temperate group, the subgroups with higher similarities were: Azores and Madeira, Canary Islands and Morocco, and the Mediterranean and the Atlantic coast of the Iberian Peninsula. Based on affinity indices, the probable directions of faunal flows were inferred. The tropical coast of Africa and the Mediterranean emerged from this analysis as probable speciation centres of the north-eastern Atlantic blenniid fauna. The Mediterranean may have also acted as a refuge during glacial periods. [ABSTRACT FROM AUTHOR]

Presents a study on visual communication in a marine fish, Salaria pavo, using video recordings. Design of the video monitors; Controversy in using video playbacks in the study of animal visual communication; Details on the reactions to the stimulus of the Salaria pavo species; Discussion on reaction differences of species.

Discusses the relationship between social status and male secondary sex traits in cichlid fish at the onset of sexual maturation. Differential expression of morphology and behavioral traits according to social status; Effect of social status on the development of the genital papillae; Predictor of androgen levels in cichlid fishes.

Presents a study that examined the maternal aggression in cichlid mossambicus during the fish mouthbrooding cycle. Methodology used in the study; Comparison between the behaviour of brooding female with non-incubating females; Changes in the agressive behaviors and feeding activities during the brooding cycle of the female cichlid fish; Reasons for the increased maternal agression of female fish during the cycle.

Dominance hierarchies were studied during 8 weeks in eight heterosexual captive groups of Oreochromis mossambicus at the onset of sexual maturity. Linearity was assessed for each group on a weekly basis based on the Landau's index (b) and on the procedure described by Appleby [Appleby MC (1983): Animal Behaviour 31:600-608]. Week-to- week stability of the hierarchical structures was measured using the Burk's stability index (b) and Spearman rank correlation coefficients for the ranks of fishes in consecutive weeks. The two measures of stability showed a high level of concordance but the Spearman rank correlations were more conservative in detecting cases of stability. Although most dominance structures were linear they were unstable from week to week. Dominant individuals were less likely to experience rank reversals than subordinates. Rank reversals were especially likely among hierarchical neighbors. Dominance hierarchies did not stabilize in the time span of this study. [ABSTRACT FROM AUTHOR]

Focuses on a study which investigated the making of mudballs by the European fiddler crab. Differences in mudballing behavior between the male and female of the species; Behavior of the males when the mudball were removed; Behavior of the females in relation to the number of mudballs outside the male burrows.

Describes a method for recording the coordinates of a fish in an aquarium in a three-dimensional space using a single video camera and a mirror. Review of related literature; Assumptions of the lens formula.

Knowing which of correlated traits are more strongly targeted by selection is crucial to understand the evolutionary process. For example, it could help in understanding how behavioural and cognitive adaptations to social living have evolved.Social competence is the ability of animals to optimize their social behaviours according to the demands of their social environment. It is a behavioural performance trait that expresses how well a whole organism performs complex social tasks, such as choosing mates, raising offspring, participating in dominance hierarchies, solving conflicts or forming social bonds. Non‐social competence, on the other hand, is the ability of animals to optimize their non‐social behaviours according to the demands of their non‐social environment, such as finding food or avoiding predators.Social and non‐social cognition are correlated lower‐level traits of social and non‐social competence, respectively, encompassing the underlying psychological and neural mechanisms of behaviour that allow animals to acquire, encode, store and recall information about their social and non‐social environments.Here, we employ the theoretical framework that selection acts on performance traits first and on lower‐level traits only secondarily, to propose a new approach to the study of the evolution of social cognition.We hypothesize that when selection favours social competence, the cognitive system becomes more adapted to the social domain, making species biased for social information, and increasing their degree of sociality. The opposite can happen when selection favours non‐social competence.The level of specialization that the cognitive system can attain depends on whether social and non‐social competence are correlated with the same cognitive lower‐level traits. This in turn will determine whether species will evolve a type of social cognition that is general—that contributes with cognitive abilities that can be used in both social and non‐social environments—or modular—that contributes with cognitive abilities that are specific to the social environment. A free Plain Language Summary can be found within the Supporting Information of this article. [ABSTRACT FROM AUTHOR]

The fitness benefits of social life depend on the ability of animals to affiliate with others and form groups, on dominance hierarchies within groups that determine resource distribution, and on cognitive capacities for recognition, learning and information transfer. The evolution of these phenotypes is coupled with that of neuroendocrine mechanisms, but the causal link between the two remains underexplored. Growing evidence from our research group and others demonstrates that the tools available in zebrafish, Danio rerio, can markedly facilitate progress in this field. Here, we review this evidence and provide a synthesis of the state‐of‐the‐art in this model system. We discuss the involvement of generalized motivation and cognitive components, neuroplasticity and functional connectivity across social decision‐making brain areas, and how these are modulated chiefly by the oxytocin‐vasopressin neuroendocrine system, but also by reward‐pathway monoamine signaling and the effects of sex‐hormones and stress physiology. [ABSTRACT FROM AUTHOR]

Group living animals can use the behavior of others as cues for the presence of threat in the environment and adjust their behavior accordingly. Therefore, different social phenomena that modulate the response to threat, such as social buffering, social transmission (contagion), and facilitation of alarm responses can be seen as different manifestations of social information use in threat detection. Thus, social phenomena that are functionally antagonistic, such as social buffering and social transmission of fear, may rely on shared neurobehavioral mechanisms related to the use of social information in decision-making about the presence of threat in the environment. Here, we propose a unifying conceptual framework for the study of social information use in threat perception based on signal detection theory. [ABSTRACT FROM PUBLISHER]

For male vertebrates, androgens are considered physiological mediators of the trade-off between mating and parenting effort. About 30 years ago, the challenge hypothesis provided a conceptual framework to explain the variation in androgen levels among individuals and species, primarily as a function of male competition and parental care. Initially developed in—and applied to—birds, the challenge hypothesis was rapidly adopted for other vertebrate groups and even insects. Experimental evidence on birds, however, offers limited support for the challenge hypothesis in terms of explaining androgen responses to social challenges from other males. Rather than throwing the baby out with the bathwater, we advance a modified challenge hypothesis 2.0, in which male–female interactions are more important than male–male interactions in mediating rapid changes in plasma androgen concentrations. The predictions we generate are supported by current evidence from birds and can be tested in other animal taxa. [ABSTRACT FROM AUTHOR]

Background: Animal models enable targeting autism-associated genes, such as the shank3 gene, to assess their impact on behavioural phenotypes. However, this is often limited to simple behaviours relevant for social interaction. Social contagion is a complex phenotype forming the basis of human empathic behaviour and involves attention to the behaviour of others for recognizing and sharing their emotional or affective state. Thus, it is a form of social communication, which constitutes the most common developmental impairment across autism spectrum disorders (ASD). Methods: Here we describe the development of a zebrafish model that identifies the neurocognitive mechanisms by which shank3 mutation drives deficits in social contagion. We used a CRISPR-Cas9 technique to generate mutations to the shank3a gene, a zebrafish paralogue found to present greater orthology and functional conservation relative to the human gene. Mutants were first compared to wild types during a two-phase protocol that involves the observation of two conflicting states, distress and neutral, and the later recall and discrimination of others when no longer presenting such differences. Then, the whole-brain expression of different neuroplasticity markers was compared between genotypes and their contribution to cluster-specific phenotypic variation was assessed. Results: The shank3 mutation markedly reduced social contagion via deficits in attention contributing to difficulties in recognising affective states. Also, the mutation changed the expression of neuronal plasticity genes. However, only downregulated neuroligins clustered with shank3a expression under a combined synaptogenesis component that contributed specifically to variation in attention. Limitations: While zebrafish are extremely useful in identifying the role of shank3 mutations to composite social behaviour, they are unlikely to represent the full complexity of socio-cognitive and communication deficits presented by human ASD pathology. Moreover, zebrafish cannot represent the scaling up of these deficits to higher-order empathic and prosocial phenotypes seen in humans. Conclusions: We demonstrate a causal link between the zebrafish orthologue of an ASD-associated gene and the attentional control of affect recognition and consequent social contagion. This models autistic affect-communication pathology in zebrafish and reveals a genetic attention-deficit mechanism, addressing the ongoing debate for such mechanisms accounting for emotion recognition difficulties in autistic individuals. [ABSTRACT FROM AUTHOR]

Group living animals use social and asocial cues to predict the presence of reward or punishment in the environment through associative learning. The degree to which social and asocial learning share the same mechanisms is still a matter of debate. We have used a classical conditioning paradigm in zebrafish, in which a social (fish image) or an asocial (circle image) conditioned stimulus (CS) have been paired with an unconditioned stimulus (US=food), and we have used the expression of the immediate early gene c-fos to map the neural circuits associated with each learning type. Our results show that the learning performance is similar to social and asocial CSs. However, the brain regions activated in each learning type are distinct and a community analysis of brain network data reveals segregated functional submodules, which seem to be associated with different cognitive functions involved in the learning tasks. These results suggest that, despite localized differences in brain activity between social and asocial learning, they share a common learning module and social learning also recruits a specific social stimulus integration module. Therefore, our results support the occurrence of a common general-purpose learning module, that is differentially modulated by localized activation in social and asocial learning. Zebrafish in a classical conditioning paradigm display similar learning performance, yet distinct brain region activation, in the context of social- or asocial-conditioned stimuli, suggesting the occurrence of a general-purpose learning module that is differentially modulated by localized activation. [ABSTRACT FROM AUTHOR]

Androgens are the principal sex steroids controlling reproduction and aggression in male fish, but their production can also be affected by social interactions. Here we show that androgen concentrations are not significantly increased in cichlid fish (Oreochromis mossambicus) that are fighting their own image in a mirror, despite their aggressive behaviour towards the virtual intruder. Our results indicate that the hormonal response normally triggered in male contests is not induced under these circumstances by the act of fighting itself, and that it may therefore depend on some indicator of relative fighting ability that cannot be delivered by a mirror-image challenger. [ABSTRACT FROM AUTHOR]

Emotional contagion is the most ancestral form of empathy. We tested to what extent the proximate mechanisms of emotional contagion are evolutionarily conserved by assessing the role of oxytocin, known to regulate empathic behaviors in mammals, in social fear contagion in zebrafish. Using oxytocin and oxytocin receptor mutants, we show that oxytocin is both necessary and sufficient for observer zebrafish to imitate the distressed behavior of conspecific demonstrators. The brain regions associated with emotional contagion in zebrafish are homologous to those involved in the same process in rodents (e.g., striatum, lateral septum), receiving direct projections from oxytocinergic neurons located in the pre-optic area. Together, our results support an evolutionary conserved role for oxytocin as a key regulator of basic empathic behaviors across vertebrates. [ABSTRACT FROM AUTHOR]

Comments on the relationship between aggression, dominance and androgens. Potential role of testosterone as a physiological mediator between social status and the expression of male character; Dominance of testosterone in humans.

• We tested how the social environment and cortisol exposure (acute/chronic) affect adult neurogenesis in zebrafish. • Cell proliferation is dependent on the valence of the social environment (positive > negative). • Cell proliferation is independent of either acute or chronic cortisol exposure. • Thus, there is a key role of the social environment in adult neurogenesis in zebrafish independently of cortisol modulation. In Mammals adult neurogenesis is influenced by environmental conditions, and the glucocorticoid hormones (GC) play a major role in this regulation. In contrast in fish, the study of the effects of cortisol on the regulation of environmental driven adult neurogenesis has produced conflicting results. While in some species elevated cortisol levels impair cell proliferation, in others, it promotes cell proliferation and differentiation. This lack of consistency may be explained by methodological differences across studies, namely in the stimuli and/or cortisol treatments used. Here, we tested the effects of the social environment on adult neurogenesis, considering a positive and a negative social context, and different durations of cortisol exposure. We hypothesise that there is an interaction between the valence of the social environment and cortisol, such that elevated acute cortisol experienced during social interactions only have a detrimental effect on neurogenesis in negative social contexts. Therefore, fish were exposed to a positive (conspecific shoal) or negative (predator) social experience, and the interaction between the valence of the social context and cortisol exposure (acute and chronic) was tested. Our results indicate that adult neurogenesis is modulated by the social environment, with the number of newly generated cells being dependent on the valence of the social information (positive > negative). These effects were independent of cortisol, either for acute or chronic exposure, highlighting the social environment as a key factor in the modulation of cell proliferation in the adult zebrafish brain, and rejecting a role for cortisol in this modulation. [ABSTRACT FROM AUTHOR]

Climate change is a growing threat to marine organisms and ecosystems, and it is already modifying ocean properties by, for example, increasing temperature and decreasing pH. Increasing water temperature may also lead to an impairment of primary productivity and an overall depletion of available zooplankton. Understanding how the crossover between warming and zooplankton availability impacts fish populations has paramount implications for conservation and mitigation strategies. Through a cross factorial design to test the effects of ocean temperature and food availability in a temperate marine teleost, Pomatochistus flavescens, we showed that hindered feeding impacted sheltering and avoidance behaviour. Also, low food availability impaired fish reproduction, particularly male reproduction, as the expression of cyp11b1, a gene with a pivotal role in the synthesis of the most important fish androgen, 11-ketotestosterone, was significantly reduced under a low food regime. In contrast, temperature alone did not affect reproductive success, but offspring showed increased saturated fatty acid content (embryos) and increased lipid peroxidation (larvae). Altogether, food availability had a stronger effect on fitness, showing that coping with elevated temperatures, an ability that may be expected in shallow-water fish, can be indirectly impacted, or even overwhelmed, by the effects of ocean warming on primary productivity and downstream ecological processes. [ABSTRACT FROM AUTHOR]

Reports on research to determine whether the so-called challenge hypothesis can be extended to bystanders. Statement that an individual's androgen production responds to their social interactions with other group members; Measurement of the concentrations of testosterone and 11-ketotestosterone in adult male cichlid fish; Production of an androgen surge in spectators; Conclusion that androgens may eventually be viewed not only as sex steroids but also as competition hormones.

Public information is widely available at low cost to animals living in social groups. For instance, bystanders may eavesdrop on signaling interactions between conspecifics and use it to adapt their subsequent behavior towards the observed individuals. This social eavesdropping ability is expected to require specialized mechanisms such as social attention, which selects social information available for learning. To begin exploring the genetic basis of social eavesdropping, we used a previously established attention paradigm in the lab to study the brain gene expression profile of male zebrafish (Danio rerio) in relation to the attention they paid towards conspecifics involved or not involved in agonistic interactions. Microarray gene chips were used to characterize their brain transcriptomes based on differential expression of single genes and gene sets. These analyses were complemented by promoter region-based techniques. Using data from both approaches, we further drafted protein interaction networks. Our results suggest that attentiveness towards conspecifics, whether interacting or not, activates pathways linked to neuronal plasticity and memory formation. The network analyses suggested that fos and jun are key players on this response, and that npas4a, nr4a1 and egr4 may also play an important role. Furthermore, specifically observing fighting interactions further triggered pathways associated to a change in the alertness status (dnajb5) and to other genes related to memory formation (btg2, npas4b), which suggests that the acquisition of eavesdropped information about social relationships activates specific processes on top of those already activated just by observing conspecifics. [ABSTRACT FROM AUTHOR]

Sociality relies on motivational and cognitive components that may have evolved independently, or may have been linked by phenotypic correlations driven by a shared selective pressure for increased social competence. Furthermore, these components may be domain‐specific or of general‐domain across social and non‐social contexts. Here, we used zebrafish to test if the motivational and cognitive components of social behavior are phenotypically linked and if they are domain specific or of general domain. The behavioral phenotyping of zebrafish in social and equivalent non‐social tests shows that the motivational (preference) and cognitive (memory) components of sociality: (1) are independent from each other, hence not supporting the occurrence of a sociality syndrome; and (2) are phenotypically linked to non‐social traits, forming two general behavioral modules, suggesting that sociality traits have been co‐opted from general‐domain motivational and cognitive traits. Moreover, the study of the association between single nucleotide polymorphisms (SNPs) and each behavioral module further supports this view, since several SNPs from a list of candidate "social" genes, are statistically associated with the motivational, but not with the cognitive, behavioral module. Together, these results support the occurrence of general‐domain motivational and cognitive behavioral modules in zebrafish, which have been co‐opted for the social domain. [ABSTRACT FROM AUTHOR]