Perinatal variation and covariation of oxidative status and telomere length in yellow-legged gull chicks.

The perinatal period is critical to survival and performance of many organisms. In birds, rapid postnatal growth and sudden exposure to aerial oxygen around hatching markedly affect the chick redox status, with potentially negative consequences on physiology mediated by oxidative stress. In addition, telomere length (TL) undergoes reduction during birds' early life, partly depending on oxidative status. However, relatively few studies have focused specifically on the changes in oxidative status and TL that occur immediately after hatching. In this study of the yellow-legged gull Larus michahellis, we found that chicks undergo a marked increase in plasma total antioxidant capacity and a marked decrease in the concentration of pro-oxidant molecules during the first days after hatching. In addition, TL in erythrocytes decreased by 1 standard deviation over the 4 days post-hatching. Body mass and tarsus length covaried with total antioxidant capacity and concentration of pro-oxidants in a complex way, that partly depended on sex and laying order, suggesting that oxidative status can affect growth. Moreover, TL positively covaried with the concentration of pro-oxidant molecules, possibly because retention of high concentrations of pro-oxidant molecules results from mechanisms of prevention of their negative effects, including reduction in TL. Thus, this study shows that chicks undergo marked variation in oxidative status, which predicts growth and subsequent TL, prompting for more studies of the perinatal changes in the critical post-hatching stages

Physiological increase of yolk testosterone level does not affect oxidative status and telomere length in gull hatchlings.

Conditions experienced during early-life can cause the onset of oxidative stress, resulting in pervasive effects on diverse life-history traits, including lifespan. In birds, maternally-transferred egg substances may exert positive or negative influence over the offspring phenotype. Among these, testosterone can upregulate the bioavailability of certain antioxidants but simultaneously promotes the production of pro-oxidants, leading to an oxidative stress situation, which is one of the main forces causing telomere attrition However, no study has investigated the role of this androgen on telomere dynamics in birds and little is known about the effects of yolk testosterone on oxidative status in early-life of these species. We physiologically increased the levels of yolk testosterone by in ovo injections in yellow-legged gull (Larus michahellis) to evaluate the effects induced by this androgen on hatchlings plasma total antioxidant capacity, amount of pro-oxidant molecules and telomere length at hatching. Testosterone supplementation did not increase hatchling body growth, did not result in the overproduction of pro-oxidant molecules nor a reduction of antioxidant capacity. Accordingly, telomere length at hatching was not affected by testosterone treatment, although hatchlings from the third-laid eggs showed shorter telomeres than their siblings from first- and second-laid eggs, independently of testosterone treatment. Our results suggest that injection of physiological levels of testosterone does not induce oxidative stress to hatchlings and, consequently do not affect telomere dynamics during early post-natal periods

Independent and combined effects of egg pro- and anti-oxidants on gull chick phenotype

Oviparous mothers transfer to their eggs components that have both independent and combined effects on offspring phenotype. The functional interaction between egg components, such as antioxidants and hormones, suggests that a change in the concentration of one component will have effects on offspring traits that depend on the concentration of other interacting components. However, the combined effects of variation in different egg components are virtually unknown. Bird eggs contain vitamin E, a major antioxidant, and also maternal corticosterone. The independent consequences of variation in the egg concentrations of these compounds for offspring phenotype are largely unknown and no study has investigated their combined effects. We manipulated the concentration of vitamin E and corticosterone in the eggs of the yellow-legged gull (Larus michahellis) by administering a physiological (2 s.d.) dose both independently and in combination. We tested for an effect on chick post-natal growth, plasma antioxidant capacity (TAC) and oxidative compounds (TOS). Separate administration of vitamin E or corticosterone caused a reduction in body mass relative to controls, whereas the combined administration of the two compounds reversed their negative effects. These results suggest that maternal egg components, such as antioxidants and steroid hormones, interact and mothers must balance their concentrations in order to achieve optimal offspring phenotype. The functional relationship between vitamin E and corticosterone is corroborated by the observation of positive covariation between these compounds

Yolk vitamin E positively affects prenatal growth but not oxidative status in yellow-legged gull embryos

Parental effects occur whenever the phenotype of parents or the environment that they experience influences the phenotype and fitness of their offspring. In birds, parental effects are often mediated by the size and biochemical quality of the eggs in terms of maternally transferred components. Exogenous antioxidants are key egg components that accomplish crucial physiological functions during early life. Among these, vitamin E plays a vital role during prenatal development when the intense metabolism accompanying rapid embryo growth results in overproduction of pro-oxidant molecules. Studies of captive birds have demonstrated the positive effect of vitamin E supplementation on diverse phenotypic traits of hatchling and adult individuals, but its effects on embryo phenotype has never been investigated neither in captivity nor under a natural selection regime. In the present study, we experimentally tested the effect of the in ovo supplementation of vitamin E on morphological traits and oxidative status of yellow-legged gull (Larus michahellis) embryos. The supplementation of vitamin E promoted somatic growth in embryos soon before hatching, but did not affect their oxidative status. Our results suggest that maternally transferred vitamin E concentrations are optimized to prevent imbalances of oxidative status and the consequent raise of oxidative damage in yellow-legged gull embryos during prenatal development

Effect of yolk corticosterone on begging in the yellow-legged gull

Behavioral lateralization is widespread across vertebrates. The development of lateralization is affected by both genetic and environmental factors. In birds, maternal substances in the egg can affect offspring lateralization via activational and/or organizational effects. Corticosterone affects the development of brain asymmetry, suggesting that variation in yolk corticosterone concentration may also influence post-natal behavioral lateralization, a hypothesis that has never been tested so far. In the yellow-legged gull (Larus michahellis), we increased yolk corticosterone concentration within physiological limits and analyzed the direction of lateralization of hatchlings in reverting from supine to prone position ('RTP' response) and in pecking at dummy parental bills to solicit food provisioning ('begging' response). We found that corticosterone treatment negatively affected the frequency of begging and it may cause a slight leftward lateralization. However, the direction of lateralization of the RTP response was not affected by corticosterone administration. Thus, our study shows a maternal effect mediated by corticosterone on a behavioral trait involved in parent-offspring communication during food provisioning events. The findings on lateralization are not conclusive due to the weak effect size but provide information for further ecological and evolutionary studies, investigating mechanisms underlying the development of lateralization

Carotenoid-based skin coloration signals antioxidant defenses in the brown trout (Salmo trutta)

Carotenoid-based signals may function as indicators of individual quality because, being exclusively obtained from the diet, they indicate the ability of individuals to intake high-quality food. Moreover, carotenoids are involved in several important physiological functions, including antioxidant defense, so that carotenoid-based colorations have been suggested to reflect the antioxidant status of their bearers. The present correlative, cross-sectional study aimed at investigating if the skin carotenoid-based coloration is a signal of antioxidant defenses in the brown trout (Salmo trutta Linnaeus, 1758). We investigated the relationships between carotenoid-based coloration traits (including the number, density and redness of red spots, as well as the ventral yellowness), and both non-enzymatic (plasma and liver total antioxidant capacity) and enzymatic antioxidant defenses (activity of hepatic superoxide dismutase\ua0\u2014SOD, catalase\ua0\u2014CAT and glutathione peroxidase\ua0\u2014GPx). We found significant positive covariations between antioxidant defenses and carotenoid-based skin coloration, in terms of ventral yellowness. Brown trout individuals displaying intense carotenoid-based coloration (i.e., ventral yellowness) had a high non-enzymatic antioxidant capacity both in plasma and in liver and, interestingly, an elevated activity of hepatic SOD and CAT. Our data suggest that carotenoid-based skin colorations may be considered a signal of individual quality in terms of antioxidant defenses in the brown trout

Contrasting effects of increased yolk testosterone content on development and oxidative status in gull embryos

Hormone-mediated maternal effects generate variation in offspring phenotype. In birds, maternal egg testosterone (T) exerts differential effects on offspring traits after hatching, suggesting that mothers experience a trade-off between contrasting T effects. However, there is very little information on T pre-natal effects. In the yellow-legged gull (Larus michahellis), we increased yolk T concentration within physiological limits and measured the effects on development and oxidative status of late-stage embryos. T-treated embryos had a larger body size but a smaller brain than controls. Males had a larger brain than females, controlling for overall size. T treatment differentially affected brain mass and total amount of pro-oxidants in the brain depending on laying order. T-treatment effects were not sex dependent. For the first time in the wild, we show contrasting T prenatal effects on body mass and brain size. Hence, T may enforce trade-offs between different embryonic traits, but also within the same trait during different developmental periods

Yolk testosterone affects growth and promotes individual-level consistency in behavioral lateralization of yellow-legged gull chicks

Behavioral lateralization is common in animals and may be expressed at the individual- and at the population-level. The ontogenetic processes that control lateralization, however, are largely unknown. Well-established sex-dependence in androgen physiology and sex-dependent variation in lateralization have led to the hypothesis that testosterone (T) has organizational effects on lateralization. The effects of T exposure in early life on lateralization can be efficiently investigated by manipulating T levels in the cleidoic eggs of birds, because the embryo is isolated from maternal and sibling physiological interference, but this approach has been adopted very rarely. In the yellow-legged gull (Larus michahellis) we increased yolk T concentration within the physiological limits and tested the effects on the direction of lateralization in two functionally fundamental behaviors (begging for parental care and escape to cover) of molecularly sexed hatchlings. We also speculated that T may intervene in regulating consistency, rather than direction of lateralization, and therefore tested if T affected the 'repeatability' of lateral preference in consecutive behavioral trials. T treatment had no effect on the direction of lateralization, but enhanced the consistency of lateral preference in escape responses. Sex did not predict lateralization. Neither behavior was lateralized at the population-level. We therefore showed for the first time in any species an effect of egg T on consistency in lateralization. The implications of the effect of T for the evolution of trade-offs in maternal allocation of egg hormones, and the evolutionary interpretations of findings from our studies on lateralization among unmanipulated birds are discussed

Within- and Among-Clutch Variation of Yolk Perfluoroalkyl Acids in a Seabird from the Northern Adriatic Sea

Perfluoroalkyl substances (PFAS) are surface-active agents used in diverse industrial and commercial applications. They contaminate both freshwater and marine ecosystems, are highly persistent, and accumulate through trophic transfer. Seabirds are exposed to environmental contaminants due to their high trophic position in food webs and relatively long lifespan. We measured levels of 10 perfluoroalkyl acids (PFAAs) in egg yolks of yellow-legged gulls (Larus michahellis) breeding in the northern Adriatic Sea (Northeast Italy). We examined variations in PFAAs within clutches (between eggs of different laying order) and among clutches. Perfluorooctane sulfonate (PFOS) was the most abundant yolk PFAA (mean = 42.0 ng/g wet wt), followed by perfluorooctanoic acid (PFOA; 3.8 ng/g wet wt) and perfluorododecanoic acid (PFDoDa; 2.8 ng/g wet wt). The 11PFAAs averaged 57.4 ng/g wet weight, ranging between 26.5 and 115.0 ng/g wet weight. The PFAA levels varied substantially among clutches (0.29\u20130.79 of the total variation), whereas the effects of laying order were considerably weaker (0.01\u20130.13). Egg-laying order effects were detected for 11PFAAs, PFOS, perfluorononanoic acid (PFNA), perfluorodecanoic acid (PFDA), perfluoroundecanoic acid (PFUnA), and PFDoDa, whereby the last-laid eggs exhibited lower PFAA concentrations than early-laid eggs. Our results indicate that seagulls from the northern Adriatic basin deposit measurable amounts of PFAAs in their eggs. The large among-clutches differences in PFAAs suggest that exposure of yellow-legged gull females to these compounds is highly variable