Suchergebnisse

To understand how traits used in animal communication evolved and are maintained as honest signals, we need to understand the mechanisms that prevent cheating. It has been proposed that honest signaling is guaranteed by the costs associated with the signal expression. However, the nature of these costs is still under debate. Melanin-based signals are intriguing because their expression seems to be tightly controlled by genes and the resource involved (i.e. melanin) seems to be not limited. However, in vertebrates, low levels of a key intracellular antioxidant (i.e. glutathione) are needed to promote melanogenesis. We propose that melanin-based ornaments can signal the ability to cope with oxidative stress because those individuals with low enough levels of glutathione, such as those required for melanin production, should manage well the whole of the antioxidant machinery in order to maintain a certain oxidative status. We analysed the expression of a melanin-based signal: the well-known black stripe of the great tit (Parus major). Great tit nestlings were injected with a specific inhibitor of glutathione production (DL-buthionine-S,R-sulfoximine; BSO) throughout their development. BSO effectively decreased intracellular glutathione levels without apparent side effects on growth or body condition. Instead, treated nestlings developed black breast stripes 70–100% larger than controls. Moreover, treated nestlings also compensated the decrease in glutathione levels by increasing the levels of circulating antioxidants. Results indicate that melanin-based signals can be at least partially permeable to environmental influences such as those associated to oxidative stress. They also reveal a potential handicap associated to the expression of this kind of signals. Finally, although other contributing factors could have been present, our findings emphasize the role of oxidative stress in shaping the evolution of animal signals in general and, in particular, those produced by pigments. ; Financial support was obtained from the CGL2006-10357-C02-02 and CGL2007-61395BOS projects from Ministerio de Educación y Ciencia (Spanish Government). I.G. benefited from a FPI grant, whereas C.A.-A. from a Ramón y Cajal fellowship, both from Ministerio de Educación y Ciencia. ; Peer reviewed

Interest in the imbalance between the production of reactive oxygen species and the state of the antioxidant machinery—that is, oxidative stress—has recently grown among comparative physiologists and evolutionary/behavioral ecologists. The number and types of markers used to estimate oxidative stress is, however, under debate. The study of covariation among these markers is necessary to better interpret the information content of each independent variable. Here, the covariation in levels of 10 blood parameters in a group of zebra finches (Taeniopygia guttata) as nestlings and adults was analyzed across a large data set. Total glutathione levels in erythrocytes were negatively correlated with plasma carotenoid values in nestlings only, supporting the implication of carotenoids in the antioxidant machinery during a particularly stressful period of life. Plasma lipid levels (triglycerides [TRGs]) as well as plasma antioxidant capacity—the latter tested with and without control for uric acid levels—showed individual consistency with age. Plasma TRG and uric acid levels were strongly correlated with plasma lipid peroxidation and antioxidant capacity, respectively, suggesting an influence of recent intake or mobilization of energy stores on these variables. The meaning of oxidative stress markers, whether corrected or uncorrected for levels of nutritional metabolites, remains to be explored. Experiments manipulating diet composition and oxidative stress are necessary to confirm or reject the hypothesized causalities. ; A.A.R.-H. was funded by a Formación de Personal de Investigación grant (BES-2010-035013; Ministerio de Economía y Competitividad [MINECO], Spanish government). Financial support was obtained from the projects CGL-2009-10883-C02-02 and CGL2012-40229-C02-01 (MINECO). ; Peer Reviewed

This studywas funded by Consejería de Educación y Ciencia, Junta de Comunidades de Castilla la Mancha (project ref.: PII1I09-0271-5037) and Ministerio de Economía y Competitividad (CGL2009-10883-C02-02 and CGL2012-40229-C02-01) from the Spanish Government. I.G. benefited from a postdoctoral contract of the CSIC JAE-Doc programme. ; Peer reviewed

Environmental conditions during early life may shape phenotype in adulthood. Early adverse conditions may increase the oxidative stress in adults, which could affect their reproductive output and survival. It has also been hypothesized that the larger the reproductive investment, the higher the oxidative stress. We tested this and the potential influence of early oxidative stress on how individuals respond to a reproductive stimulation. The synthesis of the antioxidant glutathione was inhibited in captive zebra finches (Taeniopygia guttata) during growth. In adulthood, the expression of a carotenoid-based sexual signal, bill redness, increased in both sexes, with females also being heavier than controls. The social context of control and glutathione-inhibited males was then manipulated to stimulate precopulatory reproductive investments. Males were individually caged in front of a female or another male. We predicted that males enduring lower early antioxidant levels and placed close to a female should pay the highest cost of a hypothetical increase in bill redness in terms of oxidative damage. However, early conditions only influenced the male's phenotype via their partners. Males caged with females showed increases in circulating pigment (carotenoid) levels, but only when females endured early low antioxidant values. This was probably related to the higher attractiveness of these females. Nevertheless, the bill redness of males did not differ during the social manipulation. Moreover, males facing females from any early condition group showed lower oxidative damage levels in plasma lipids. This result agrees with some findings in rodents, also in captivity. However, the effect may be due to increased triglyceride levels and body mass in males not facing females, as variation in these traits explained oxidative damage variability. The importance of considering housing conditions and life history when interpreting oxidative stress-related trade-offs is highlighted. ; AR was funded by a Formación de Personal de Investigación grant (BES-2010-035013; Ministerio de Economía y Competitividad, MINECO, Spanish Government). Financial support was obtained from the projects CGL-2009-10883-C02-02 and CGL2012-40229-C02-01 (MINECO, Spain). ; Peer Reviewed

The synthesis of melanins, which are the most common animal pigments, is influenced by glutathione (GSH), a key intracellular antioxidant. At high GSH levels, pheomelanin (the lightest melanin form) is produced, whereas production of eumelanin (the darkest melanin form) does not require GSH. Oxidative damage typically increases with age, and age-related decreases in GSH have accordingly been found in diverse organisms. Therefore, there should be positive associations between the capacity to produce eumelanic traits, GSH levels, and senescence, whereas there should be negative associations between the capacity to produce pheomelanic traits, GSH levels, and senescence. We explored this hypothesis in a free-ranging population of wild boars Sus scrofa of different ages. As expected from the fact that pheomelanogenesis consumes GSH, levels of this antioxidant in muscle tended to be negatively related to pheomelanization and positively related to eumelanization in pelage, and the degree of pelage pheomelanization was positively related to oxidative damage as reflected by levels of thiobarbituric- acid-reactive substances (TBARS), which is consistent with the hypothesis that pheomelanin synthesis has physiological costs. In our cross-sectional sample, GSH levels did not show senescence effects, and we did not detect senescence effects in pelage melanization. Prime body condition and low TBARS levels were also associated with hair graying, which is attributable to a loss of melanin produced by oxidative stress, thus raising the possibility that hair graying constitutes a signal of resistance to oxidative stress in wild boars. Our results suggest that the degree of melanization is linked to GSH levels in wild boars and that their antioxidant damage shows senescence effects. © 2012 by The University of Chicago. All rights reserved. ; During writing, I.G. was supported by a Marie Curie Intra European Fellowship (PIEF-GA-2009-252145) within the Seventh European Community Framework Programme. Laboratory analyses were funded by the Ministerio de Ciencia e Innovación, Spanish Government (project nos. CGL2009-10883-C02-02 and CGL2009-10652). ; Peer Reviewed

Colorful ornaments have been the focus of sexual selection studies since the work of Darwin. Yellow to red coloration is often produced by carotenoid pigments. Different hypotheses have been formulated to explain the evolution of these traits as signals of individual quality. Many of these hypotheses involve the existence of a signal production cost. The carotenoids necessary for signaling can only be obtained from food. In this line, carotenoid-based signals could reveal an individual's capacity to find sufficient dietary pigments. However, the ingested carotenoids are often yellow and became transformed by the organism to produce pigments of more intense color (red ketocarotenoids). Biotransformation should involve oxidation reactions, although the exact mechanism is poorly known. We tested the hypothesis that carotenoid biotransformation could be costly because a certain level of oxidative stress is required to correctly perform the conversion. The carotenoid-based signals could thus reveal the efficiency of the owner in successfully managing this challenge. In a bird with ketocarotenoid-based ornaments (the red-legged partridge Alectoris rufa), the availability of different carotenoids in the diet (i.e. astaxanthin, zeaxanthin and lutein) and oxidative stress were manipulated. The carotenoid composition was analyzed and quantified in the ornaments, blood, liver and fat. A number of oxidative stress biomarkers were also measured in the same tissues. First, we found that color and pigment levels in the ornaments depended on food levels of those carotenoids used as substrates in biotransformation. Second, we found that birds exposed to mild levels of a free radical generator (diquat) developed redder bills and deposited higher amounts of ketocarotenoids (astaxanthin) in ornaments. Moreover, the same diquat-exposed birds also showed a weaker resistance to hemolysis when their erythrocytes were exposed to free radicals, with females also enduring higher oxidative damage in plasma lipids. Thus, higher color production would be linked to higher oxidative stress, supporting the biotransformation hypothesis. The recent discovery of an avian oxygenase enzyme involved in converting yellow to red carotenoids may support our results. Nonetheless, the effect could also depend on the abundance of specific substrate carotenoids in the diet. Birds fed with proportionally higher levels of zeaxanthin showed the reddest ornaments with the highest astaxanthin concentrations. Moreover, these birds tended to show the strongest diquat-mediated effect. Therefore, in the evolution of carotenoid-based sexual signals, a biotransformation cost derived from maintaining a well-adjusted redox machinery could coexist with a cost linked to carotenoid acquisition and allocation (i.e. a resource allocation trade-off). ; Esther García-de Blas was supported by a predoctoral grant (JAE-PRE) from the Consejo Superior de Investigaciones Científicas (CSIC) co-financed by Fondo Social Europeo (EU). This study was funded by Consejería de Educación y Ciencia, Junta de Comunidades de Castilla la Mancha (project ref.: PII1I09-0271-5037) and Ministerio de Economía y Competitividad (CGL2009-10883-C02-02 and CGL2015-69338-C2-2-P) from the Spanish Government. ; Peer Reviewed

The mechanisms that make that the costs of producing high-quality signals are unaffordable to low-quality signalers are a current issue in animal communication. The size of the melanin-based bib of male house sparrows Passer domesticus honestly signals quality. We induced the development of new bibs while treating males with buthionine-sulfoximine (BSO), a substance that depletes the levels of the antioxidant glutathione (GSH) and the amino acid cysteine, two elements that switch melanogenesis from eumelanin to pheomelanin. Final bib size is negatively related to pheomelanin levels in the bib feathers. BSO reduced cysteine and GSH levels in all birds, but improved phenotypes (bibs larger than controls) were only expressed by high-quality birds (BSO birds with largest bibs initially). Negative associations between final bib size and cysteine levels in erythrocytes, and between pheomelanin and cysteine levels, were observed in high-quality birds only. These findings suggest that a mechanism uncoupling pheomelanin and cysteine levels may have evolved in low-quality birds to avoid producing bibs of size not corresponding to their quality and greater relative costs. Indeed, greater oxidative stress in cells was not observed in low-quality birds. This may represent the first mechanism maintaining signal honesty without producing greater relative costs on low-quality signalers. ; I.G. benefited from a postdoctoral contract of the CSIC JAE-Doc program and, during writing, from a Ramón y Cajal Fellowship. his study was funded by Consejería de Educación y Ciencia, Junta de Comunidades de Castilla-La Mancha (project ref.: PII1I09-0271-5037), and Ministerio de Economía y Competitividad (CGL2009-10883-C02-02 and CGL2012-40229-C02-01) from the Spanish Government. ; Peer Reviewed

Many animal species show ornaments with yellow-orange-red colors produced by carotenoid pigments. Such traits have evolved as reliable signals of individual quality because of the costs inherent to their production or maintenance. In animal tissues, carotenoids are often found combined with free fatty acids, as carotenoid esters, which may confer more stability to coloration than free carotenoids. Surprisingly, the potential relevance of carotenoid esterification in the expression of animal sexual signals has been virtually ignored. Moreover, the sources of variability of esterified carotenoid levels are barely known, because most studies have not quantified their concentrations. Here, carotenoids in the ornaments (bill, eye rings, and legs) of red-legged partridges Alectoris rufa were quantified in their free and esterified forms. Carotenoid ester levels were the best predictors of leg color, whereas the redness of the other traits was better explained by free carotenoids. Nonetheless, total carotenoid levels (the sum of free and esterified forms) were always significantly correlated to redness. Young partridges had lower levels of free and esterified carotenids in the legs than did older individuals. Also, wild animals had higher ester levels and a higher proportion of carotenoids in esterified forms in all traits than did captive partridges. Probable physiological mechanisms explaining these patterns are discussed. ; Esther García-de Blas was supported by a predoctoral grant (JAE-PRE program) from the Consejo Superior de Investigaciones Científicas cofinanced by Fondo Socia Europeo. Lorenzo Pérez-Rodríguez was supported by a "Juan de la Cierva" postdoctoral contract from the Spanish Ministerio de Ciencia e Innovación (JCI-2008-2059). This study was funded by Consejería de Educación y Ciencia, Junta de Comunidades de Castilla la Mancha (PII1I09-0271-5037), and Ministerio de Economía y Competitividad (CGL2009-10883-C02-02) from the Spanish government. ; Peer reviewed