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- Maternal thyroid hormones are essential for neural development in ZebrafishPublication . Campinho, Marco António; Saraiva, João; Florindo, Claudia; Power, Deborah M.Teleost eggs contain an abundant store of maternal thyroid hormones (THs), and early in zebrafish embryonic development, all the genes necessary for TH signaling are expressed. Nonetheless the function of THs in embryonic development remains elusive. To test the hypothesis that THs are fundamental for zebrafish embryonic development, an monocarboxilic transporter 8 (Mct8) knockdown strategy was deployed to prevent maternal TH uptake. Absence of maternal THs did not affect early specification of the neural epithelia but profoundly modified later dorsal specification of the brain and spinal cord as well as specific neuron differentiation. Maternal THs acted upstream of pax2a, pax7, and pax8 genes but downstream of shha and fgf8a signaling. The lack of inhibitory spinal cord interneurons and increased motoneurons in the mct8 morphants is consistent with their stiff axial body and impaired mobility. The mct8 mutations are associated with X-linked mental retardation in humans, and the cellular and molecular consequences of MCT8 knockdown during embryonic development in zebrafish provides new insight into the potential role of THs in this condition.
- Chemical diplomacy in male tilapia: urinary signal increases sex hormone and decreases aggressionPublication . Saraiva, João; Keller-Costa, Tina; Hubbard, Peter; Rato, Ana; Canario, AdelinoAndrogens, namely 11-ketotestosterone (11KT), have a central role in male fish reproductive physiology and are thought to be involved in both aggression and social signalling. Aggressive encounters occur frequently in social species, and fights may cause energy depletion, injury and loss of social status. Signalling for social dominance and fighting ability in an agonistic context can minimize these costs. Here, we test the hypothesis of a 'chemical diplomacy' mechanism through urinary signals that avoids aggression and evokes an androgen response in receiver males of Mozambique tilapia (Oreochromis mossambicus). We show a decoupling between aggression and the androgen response; males fighting their mirror image experience an unresolved interaction and a severe drop in urinary 11KT. However, if concurrently exposed to dominant male urine, aggression drops but urinary 11KT levels remain high. Furthermore, 11KT increases in males exposed to dominant male urine in the absence of a visual stimulus. The use of a urinary signal to lower aggression may be an adaptive mechanism to resolve disputes and avoid the costs of fighting. As dominance is linked to nest building and mating with females, the 11KT response of subordinate males suggests chemical eavesdropping, possibly in preparation for parasitic fertilizations.
- Social organization and endocrine profiles of Australoheros facetus, an exotic freshwater fish in southern PortugalPublication . Baduy, F.; Silva, FBV; Guerreiro, Pedro M; Canario, A.V.M.; L. Saraiva, JoãoAustraloheros facetus is a neotropical cichlid and an exotic species in the Guadiana and Odelouca basins (Southern Portugal). In this research, we aimed to characterize the main behavioural patterns and circulating hormones, 17 beta-estradiol (E-2) for females, and testosterone, 11-ketotestosterone (11KT) and cortisol for both sexes, during the formation of social groups, a crucial step in the life-history of A. facetus. A pair-breeding strategy with territorial behaviour and aggressive interactions was found, with a positive correlation between dominance and size for both sexes. There were no significant differences between non-territorial and territorial individuals in the baseline levels of hormones, but 11KT was higher in males when they became territorial, as E-2 for territorial females, while there was no clear pattern for testosterone. In contrast, cortisol was higher in non-territorial males and correlated negatively with social dominance. These results suggest that size is the main driver for social dominance and that formation of stable hierarchies result in higher circulating 11KT and lowers stress in territorial males. Related to A. facetus' behaviour, aggressiveness and biparental care may be an advantage during the invasion process of A. facetus, facilitating colonization of new habitats.
- Birth date predicts alternative life-history pathways in a fish with sequential reproductive tacticsPublication . Fagundes, Teresa; Simões, Mariana G.; L. Saraiva, João; Ros, Albert F. H.; Gonçalves, David; Oliveira, Rui F.1. In species with plastic expression of alternative reproductive tactics (ARTs), individuals of the same sex, usually males, can adopt different reproductive tactics depending on factors such as body size. 2. The 'birth date hypothesis' proposes that condition-dependent expression of ARTs may ultimately depend on birth date, because individuals born at different times of the year may achieve different sizes and express different reproductive tactics accordingly. However, this has rarely been tested. 3. Here, we tested this hypothesis in a fish with ARTs, the peacock blenny (Salaria pavo). A long-term (6 years) mark-recapture study demonstrated that ARTs in the peacock blenny were sequential and that males may follow at least two alternative life-history pathways: a nestholder pathway, in which males express the nest-holder tactic from their first breeding season onwards, and a parasitic pathway, where males reproduce on their first breeding season as sneaker males and subsequently as nest-holders. 4. We have found evidence of a birth date effect on the expression of ARTs in the peacock blenny. Males following the nest-holder pathway are born earlier and are larger at the first breeding season than males following the parasitic pathway, but they have similar growth curves. 5. The mechanisms underlying a birth date effect are far from clear and might be diverse. We have not found support for a mechanism of body size threshold triggering sexual maturation and subsequent ARTs. A mechanism of tactic determination that is strictly based on timing of first maturation is also unlikely. 6. A proxy of lifetime reproductive success shows crossing (body size associated) fitness curves for the two main life-history pathways.
- The effect of tank cover on welfare of farmed Nile tilapiaPublication . Saraiva, João L.; Nogueirinha, Margarida; Teodósio, Rita; Aragão, Cláudia; Engrola, Sofia; Arechavala Lopez, PabloWelfare in aquaculture is a pressing topic. One of the main measures to improve the life of farmed fish is implementing environmental enrichment, which can include the addition of structural covers to rearing tanks. Here we test the effects of tank covers (fully covered, 50 % covered and uncovered) in an all-male population (N = 75 tagged individuals: 5 individuals per tank, 5 tanks per treatment) of farmed Nile tilapia using behavioural and physiological welfare indicators. Video recordings were performed over 21 days to evaluate overall swimming (swim bursts, freezing on the bottom or hovering in the water column) and aggression events (bites and chases). At the end of the experiment, blood was collected to assess circulating cortisol. Results showed a clearly lower welfare in the partially covered tanks regarding behavioural and physiological variables. The present study highlights the need to correctly study 1) the ethology of the farmed species and 2) the enrichment strategy to be implemented, before any measure is taken. While environmental enrichment is undoubtedly a much needed area of research and a promising solution to improve the welfare of farmed fish, a proper ethological assessment must be made a priori to avoid detrimental effects of poorly devised approaches.
- Environmental modulation of androgen levels and secondary sex characters in two populations of the peacock blenny Salaria pavoPublication . Saraiva, João L.; Gonçalves, David M.; Oliveira, Rui F.Morphology and endocrinology were studied in two populations of the peacock blenny Salaria pavo, with different regimes of sexual selection imposed by differences in nest site availability. The peacock blenny is a small, sexually dimorphic benthic fish that presents exclusive paternal care of the clutch and inhabits rocky shores of the Mediterranean and adjacent Atlantic areas. In a population from the Gulf of Trieste (Northern Adriatic sea) inhabiting rocky shores where nest sites are abundant, male-male competition for nests is low, males court females and a low frequency of alternative reproductive tactics (small, parasitic female-mimicking sneaker males that change tactic into nest holders in subsequent breeding seasons) occurs. Conversely at Ria Formosa, a coastal lagoon in Southern Portugal, where nest sites are scarce and highly aggregated, male-male competition for nests is very high, there is sex-role reversal with female courtship and a high frequency of alternative reproductive tactics is observed. Concomitantly, at Ria Formosa nest holder males are larger and present more developed secondary sex characters and higher levels of 11 KT than at the Gulf of Trieste. However, the gonads of nest holders and parasitic males were larger in the Gulf of Trieste population. Competition for nests at Ria Formosa seems to promote more developed secondary sex characters in nest site scarcity conditions, while competition for females at the Gulf of Trieste seems to be spurring sperm competition among males in populations where nest sites are more abundant. 11 KT was thus associated with the development and expression of secondary sex characters in contrasting environments. These results exemplify how the modulation of behavioral plasticity and secondary sex characters by the social environment can be mediated by androgens. (c) 2009 Elsevier Inc. All rights reserved.
- Taxonomic re-evaluation of the non-native cichlid in Portuguese drainagesPublication . Carecho, João; Baduy, F.; Silva, FBV; Guerreiro, Pedro; Saraiva, João L.; Ribeiro, Filipe; Veríssimo, AnaA non-native cichlid fish firstly reported in Portugal in 1940 was originally identified as Cichlasoma facetum (Jenyns 1842) based on specimens reported from “Praia de Mira” (Vouga drainage, northwestern Portugal). Currently, the species is known only from three southern Portuguese river drainages, namely Sado, Arade and Guadiana, and no other record has been made from Praia de Mira or the Vouga drainage since the original record. The genus Cichlasoma has since suffered major taxonomic revisions: C. facetum has been considered a species-complex and proposed as the new genus Australoheros, including many species. Given the current taxonomic re-arrangement of the C. facetum species group, we performed a taxonomic re-evaluation of species identity of this non-native cichlid in Portuguese drainages using morphological and molecular analyses. Morphological data collected on specimens sampled in the Sado river drainages confirmed the identification as Australoheros facetus. Moreover, nucleotide sequences of the cytochrome b gene obtained from specimens from Sado, Arade and Guadiana showed the existence of a single haplotype across drainages, which was 100% identical to A. facetus specimens collected in native Argentinean waters (i.e. Uruguay River). The current non-native distribution range of the species in Portugal results from human-mediated introductions across the southern drainages.
- A global assessment of welfare in farmed fishes: the FishEthoBasePublication . Saraiva, João; Arechavala Lopez, Pablo; Castanheira, Maria Filipa; Volstorf, Jenny; Heinzpeter Studer, BilloFish welfare is an essential issue that needs to be tackled by the aquaculture industry. In order to address it, studies have been limited to a small number of species and the information is generally scattered. In order to have a consistent overview of the welfare of farmed fishes, we present the FishEthoBase, an open-access database that ultimately aims to provide information on the welfare of all fish species currently farmed worldwide. Presently with 41 species, this database is directed to all stakeholders in the field and targets not only to bridge the gaps between them but also to provide scientific information to improve the welfare of fish. The current text explains the database and presents an analysis of the welfare scores of 41 species, suggesting that (i) the general welfare state of farmed fishes is poor, (ii) there is some potential for improvement and (iii) this potential is related to research on species’ needs, but (iv) there are many remaining knowledge gaps and (v) current fish farming technologies do not seem to fully address welfare issues. The existence of a framework, such as the FishEthoBase, is proposed as fundamental to the design of strategies that improve the welfare of farmed fish.
- A multi-component pheromone in the urine of dominant male tilapia (Oreochromis mossambicus) reduces aggression in rivalsPublication . Keller-Costa, Tina; Saraiva, João L.; Hubbard, Peter; Barata, Eduardo Nuno; Canário, Adelino V. M.Males often use scent to communicate their dominance, and to mediate aggressive and breeding behaviors. In teleost fish, however, the chemical composition of male pheromones is poorly understood. Male Mozambique tilapia, Oreochromis mossambicus, use urine that signals social status and primes females to spawn. The urinary sex pheromone directed at females consists of 5 beta-pregnane-3 alpha,17 alpha,20 beta-triol 3-glucuronate and its 20 alpha-epimer. The concentration of these is positively correlated with male social rank. This study tested whether dominant male urine reduces aggression in receiver males, and whether the pregnanetriol 3-glucuronates also reduce male-male aggression. Males were allowed to fight their mirror image when exposed to either: i) water control or a chemical stimulus; ii) dominant male urine (DMU); iii) C18-solid phase (C18-SPE) DMU eluate; iv) C18-SPE DMU eluate plus filtrate; v) the two pregnanetriol 3-glucuronates (P3Gs); or vi) P3Gs plus DMU filtrate. Control males mounted an increasingly aggressive fight against their image over time. However, DMU significantly reduced this aggressive response. The two urinary P3Gs did not replicate the effect of whole DMU. Neither did the C18-SPE DMU eluate, containing the P3Gs, alone, nor the C18-SPE DMU filtrate to which the two P3Gs were added. Only exposure to reconstituted DMU (C18-SPE eluate plus filtrate) restored the aggression-reducing effect of whole DMU. Olfactory activity was present in the eluate and the polar filtrate in electro-olfactogram studies. We conclude that P3Gs alone have no reducing effect on aggression and that the urinary signal driving off male competition is likely to be a multi-component pheromone, with components present in both the polar and non-polar urine fractions.
- Changes in the gene expression profiles of the brains of male European eels (Anguilla anguilla) during sexual maturationPublication . Churcher, Allison; Pujolar, Jose M.; Milan, Massimo; Hubbard, Peter; Martins, Rute S. T.; L. Saraiva, João; Huertas, Mar; Bargelloni, Luca; Patarnello, T.; Marino, Ilaria A. M.; Zane, Lorenzo; Canario, Adelino V. M.Background: The vertebrate brain plays a critical role in the regulation of sexual maturation and reproduction by integrating environmental information with developmental and endocrine status. The European eel Anguilla anguilla is an important species in which to better understand the neuroendocrine factors that control reproduction because it is an endangered species, has a complex life cycle that includes two extreme long distance migrations with both freshwater and seawater stages and because it occupies a key position within the teleost phylogeny. At present, mature eels have never been caught in the wild and little is known about most aspects of reproduction in A. anguilla. The goal of this study was to identify genes that may be involved in sexual maturation in experimentally matured eels. For this, we used microarrays to compare the gene expression profiles of sexually mature to immature males. Results: Using a false discovery rate of 0.05, a total of 1,497 differentially expressed genes were identified. Of this set, 991 were expressed at higher levels in brains (forebrain and midbrain) of mature males while 506 were expressed at lower levels relative to brains of immature males. The set of up-regulated genes includes genes involved in neuroendocrine processes, cell-cell signaling, neurogenesis and development. Interestingly, while genes involved in immune system function were down-regulated in the brains of mature males, changes in the expression levels of several receptors and channels were observed suggesting that some rewiring is occurring in the brain at sexual maturity. Conclusions: This study shows that the brains of eels undergo major changes at the molecular level at sexual maturity that may include re-organization at the cellular level. Here, we have defined a set of genes that help to understand the molecular mechanisms controlling reproduction in eels. Some of these genes have previously described functions while many others have roles that have yet to be characterized in a reproductive context. Since most of the genes examined here have orthologs in other vertebrates, the results of this study will contribute to the body of knowledge concerning reproduction in vertebrates as well as to an improved understanding of eel biology.