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Reproduction is a demanding activity, since organisms must produce and, in some cases, protect and provision their progeny. Hence, a central tenet of life-history theory predicts that parents have to trade parental care against body maintenance. One physiological cost thought to be particularly important as a modulator of such trade-offs is oxidative stress. However, evidence in favour of the hypothesis of an oxidative cost of reproduction is contradictory. In this study, we manipulated the brood size of wild barn swallows Hirundo rustica soon after hatching of their nestlings to test whether an increase in nestling rearing effort translates into an increased oxidative damage and a decreased antioxidant protection at the end of the nestling rearing period. We found that, while plasma oxidative damage was unaffected by brood size enlargement, females rearing enlarged broods showed a decrease in plasma non-enzymatic antioxidants during the nestling rearing period. This was not the case among females rearing reduced broods and among males assigned to either treatment. Moreover, individuals with higher plasma oxidative damage soon after the brood size manipulation had lower plasma non-enzymatic antioxidants at the end of the nestling rearing period, suggesting that non-enzymatic antioxidants were depleted to buffer the negative effects of high oxidative damage. Our findings point to antioxidant depletion as a potential mechanism mediating the cost of reproduction among female birds.

Variation in baseline glucocorticoid (cort) levels can be attributed, at least in part, to differences in energetic demands confronting individuals. Elevated baseline cort levels are routinely interpreted as indicating individuals in poor condition, with low relative fitness. However, when greater reproductive effort increases energetic demands, individuals with high cort might paradoxically be those with the highest fitness. Here, we experimentally test the hypothesis that increased reproductive demand causes increases in baseline cort (the Cort-Adaptation hypothesis). We measured maternal baseline cort before and after experimentally enlarging and reducing brood sizes in tree swallows (Tachycineta bicolor). Females with experimentally enlarged broods had greater increases in baseline cort and fledged more offspring than females with reduced broods. Additionally, females with greater increases in baseline cort had higher offspring-provisioning rates than females with lower changes in cort. These findings demonstrate that increased reproductive demand can cause increased baseline cort. As yet, we do not know if these increases in cort cause increased allocation of resources towards reproduction, but the positive relationship between parental behaviour and cort suggests that increased cort does not always interfere with reproductive investment, and might instead facilitate it.

Glucocorticoid (cort) hormones are increasingly applied in studies of free-ranging animals, with elevated baseline cort levels generally assumed to indicate individuals or populations in worse condition and with lower fitness (the Cort-Fitness Hypothesis). The relationship between cort and fitness is rarely validated and studies investigating the cort-fitness relationship often find results inconsistent with the Cort-Fitness Hypothesis. The inconsistency of these studies may result in part from variation in the cort-fitness relationship across life history stages. Here we address the following questions in a two-year study in free-ranging tree swallows (Tachycineta bicolor): (1) Do baseline cort levels correlate with fitness within a life history stage? (2) Does the cort-fitness relationship vary across different life history stages? (3) Does the cort-fitness relationship vary across life history stages within an individual? (4) Does reproductive effort influence cort levels, and do cort levels influence reproductive effort? We measured baseline cort and fitness components in female birds of known breeding stages. We find correlations between baseline cort levels and fitness within some life history stages, but the relationship shifts from negative during early breeding to positive during late breeding, even within the same individuals. A positive relationship between baseline cort and fitness components during the nestling period suggests that reproductive investment may elicit higher cort levels that feedback to reallocate more effort to reproduction during critical periods of nestling provisioning. Our findings provide reason to question the Cort-Fitness Hypothesis, and have implications for the application of cort measures in monitoring the condition of populations of conservation concern.

The heterophil to lymphocyte (HL) ratio in the peripheral blood is increasingly being recognized as a reliable indicator of stress in birds. In this study we examined whether HL ratio, as well as the proportion of heterophils and lymphocytes, varied systematically with sampling time, date, age, climate conditions or with measures of maternal investment in female tree swallows (Tachycineta bicolor) over three breeding seasons. We found that HL ratios showed significant annual variation, which appeared to be driven by annual changes in the proportion of heterophils. HL ratios were higher among those females laying larger clutches, suggesting a potential cost of reproduction. Variation in body condition also appeared to affect stress levels of females, as decreases in body condition were associated with elevated HL ratios. Among females that we sampled over multiple breeding seasons, we were unable to detect significant repeatability for both HL ratios and proportion of heterophils, although proportion of lymphocytes showed low but significant repeatability within individuals. We therefore suggest that caution should be exercised in using these measures for illustrating the inherent quality or health of individuals over time frames beyond the current breeding attempt, or as predictors of future reproductive potential.

Parents feeding altricial nestlings have to trade-off the competing demands of self-maintenance and reproductive investment over their lifetime. Corticosterone, a glucocorticoid hormone released by birds in response to stressors, might play a key role in regulating parental investment when conditions unexpectedly deteriorate. However, birds breeding in unpredictable environmental conditions have been shown not to increase circulating levels of corticosterone as a response to bad weather to avoid nest abandonment when investment in offspring is high or when the probability of re-nesting is low. We investigated whether parent barn swallows Hirundo rustica, a passerine bird whose aerial insect food varies greatly in abundance depending on weather, also belongs to those species or whether it responds with an activation of the hypothalamo-pituitary-adrenal axis to natural variation in insect availability. We correlated plasma corticosterone levels of parents with weather conditions, the availability of aerial insects and parental body condition. Plasma corticosterone concentrations increased when mean daytime temperature declined, and consequently insect availability decreased and body condition of the parents deteriorated. Low temperatures also had a negative effect on body mass of the nestlings and there was a negative relationship between circulating corticosterone of parents and body mass of nestlings. We conclude that corticosterone is probably involved in the regulation of parental investment.