Sex-specific responses to GnRH challenge, but not food supply, in kittiwakes: Evidence for the "sensitivity to information" hypothesis.

Seasonal timing of breeding is usually considered to be triggered by endogenous responses linked to predictive cues (e.g., photoperiod) and supplementary cues that vary annually (e.g., food supply), but social cues are also important. Females may be more sensitive to supplementary cues because of their greater role in reproductive timing decisions, while males may only require predictive cues. We tested this hypothesis by food-supplementing female and male colonial seabirds (black-legged kittiwakes, Rissa tridactyla) during the pre-breeding season. We measured colony attendance via GPS devices, quantified pituitary and gonadal responses to gonadotropin releasing hormone (GnRH) challenge, and observed subsequent laying phenology. Food supplementation advanced laying phenology and increased colony attendance. While female pituitary responses to GnRH were consistent across the pre-breeding season, males showed a peak in pituitary sensitivity at approximately the same time that most females were initiating follicle development. The late peak in male pituitary response to GnRH questions a common assumption that males primarily rely on predictive cues (e.g., photoperiod) while females also rely on supplementary cues (e.g., food availability). Instead, male kittiwakes may integrate synchronising cues from their social environment to adjust their reproductive timing to coincide with female timing.

The effects of food supply on reproductive hormones and timing of reproduction in an income-breeding seabird.

Current food supply is a major driver of timing of breeding in income-breeding animals, likely because increased net energy balance directly increases reproductive hormones and advances breeding. In capital breeders, increased net energy balance increases energy reserves, which eventually leads to improved reproductive readiness and earlier breeding. To test the hypothesis that phenology of income-breeding birds is independent of energy reserves, we conducted an experiment on food-supplemented ("fed") and control female black-legged kittiwakes (Rissa tridactyla). We temporarily increased energy costs (via weight handicap) in a 2 × 2 design (fed/unfed; handicapped/unhandicapped) during the pre-laying period and observed movement via GPS-accelerometry. We measured body mass, baseline hormones (corticosterone; luteinising hormone) before and after handicap manipulation, and conducted a gonadotropin-releasing hormone challenge. Females from all treatment groups foraged in similar areas, implying that individuals could adjust time spent foraging, but had low flexibility to adjust foraging distance. Consistent with the idea that income breeders do not accumulate reserves in response to increased food supply, fed birds remained within an energy ceiling by reducing time foraging instead of increasing energy reserves. Moreover, body mass remained constant until the onset of follicle development 20 days prior to laying regardless of feeding or handicap, implying that females were using a 'lean and fit' approach to body mass rather than accumulating lipid reserves for breeding. Increased food supply advanced endocrine and laying phenology and altered interactions between the hypothalamic-pituitary-adrenal axis and the hypothalamic-pituitary-gonadal axis, but higher energy costs (handicap) had little effect. Consistent with our hypothesis, increased food supply (but not net energy balance) advanced endocrine and laying phenology in income-breeding birds without any impact on energy reserves.

Post-natal glucocorticoid elevation affects GnRH-induced luteinizing hormone concentration in female house sparrows.

Most non-mammalian studies investigating the long-term effects of early-life stressor exposure on endocrine regulation have focused on the hypothalamic-pituitaryadrenal (HPA) axis. However, the hypothalamic-pituitary-gonadal (HPG) axis may more directly affect fitness by regulating reproduction. Changes in HPG axis regulation could allow vertebrates to adaptively mitigate negative effects of early-life stressor exposure. However, only a few studies have examined long-term effects of early-life stressor experience on the HPG axis, and these have found mixed results. Here, we evaluate long-term effects of post-natal corticosterone exposure on the HPG axis in adult female house sparrows (Passer domesticus). We elevated circulating corticosterone non-invasively in wild nestling house sparrows between 8 and 11 days post-hatching, and then brought birds into captivity at fledging. Early in their first breeding season (ages 285-353d post-hatching), females were given a gonadotropin releasing hormone (GnRH) challenge. We found that early-life corticosterone exposure interacted with current condition such that females exposed to elevated post-natal corticosterone had higher baseline and GnRH-induced luteinizing hormone (LH) concentration than control females, but only if they had a high mass. Our results suggest that female house sparrows may mitigate negative impacts of early-life corticosterone exposure by investing in early reproduction, but only when current energetic condition allows.

Contaminants and energy expenditure in an Arctic seabird: Organochlorine pesticides and perfluoroalkyl substances are associated with metabolic rate in a contrasted manner.

Basal metabolic rate (BMR), the minimal energetic cost of living in endotherms, is known to be influenced by thyroid hormones (THs) which are known to stimulate in vitro oxygen consumption of tissues in birds and mammals. Several environmental contaminants may act on energy expenditure through their thyroid hormone-disrupting properties. However, the effect of contaminants on BMR is still poorly documented for wildlife. Here, we investigated the relationships between three groups of contaminants (organochlorines (OCs), perfluoroalkyl substances (PFASs), and mercury) with metabolic rate (MR), considered here as a proxy of BMR and also with circulating total THs (thyroxine (TT4) and triiodothyronine (TT3)) in Arctic breeding adult black-legged kittiwakes (Rissa tridactyla) from Svalbard, during the chick rearing period. Our results indicate a negative relationship between the sum of all detected chlordanes (∑CHLs) and MR in both sexes whereas perfluorotridecanoate (PFTrA) and MR were positively related in females only. MR was not associated with mercury. Additionally, levels of TT3 were negatively related to ∑CHLs but not to PFTrA. The findings from the present study indicate that some OCs (in both sexes) and some PFASs (only in females) could disrupt fine adjustment of BMR during reproduction in adult kittiwakes. Importantly, highly lipophilic OCs and highly proteinophilic PFASs appear, at least in females, to have the ability to disrupt the metabolic rate in an opposite way. Therefore, our study highlights the need for ecotoxicological studies to include a large variety of contaminants which can act in an antagonistic manner.

Intra-clutch ratio of yolk progesterone level changes with laying date in rockhopper penguins: a strategy to influence brood reduction?

Hatching asynchrony in avian species generally leads to a size hierarchy among siblings, favouring the first-hatched chicks. Maternally deposited hormones affect the embryo and chick's physiology and behaviour. It has been observed that progesterone, a hormone present at higher levels than other steroid hormones in egg yolks, is negatively related to body mass in embryos, chicks and adults. A differential within-clutch progesterone deposition could therefore be linked to the size hierarchy between siblings and to the resulting brood reduction. We tested whether yolk progesterone levels differed between eggs according to future parental ability to feed the entire clutch in wild rockhopper penguins Eudyptes chrysocome. This species presents a unique reversed egg-size dimorphism and hatching asynchrony, with the larger second-laid egg (B-egg) hatching before the smaller first-laid egg (A-egg). Yolk progesterone levels increased only slightly with female body mass at laying. However, intra-clutch ratios were not related to female body mass. On the other hand, yolk progesterone levels increased significantly with the date of laying onset for A-eggs while they decreased for B-eggs. Early clutches therefore had proportionally more progesterone in the B-egg compared to the A-egg while late clutches had proportionally less progesterone in the B-egg. We propose that females could strategically regulate yolk progesterone deposition within clutches according to the expected food availability during chick growth, an adaptive strategy to adjust brood reduction to conditions. We also discuss these results, relating to yolk progesterone, in the broader context of other yolk steroids.

Measuring corticosterone in seabird egg yolk and the presence of high yolk gestagen concentrations.

Large inter-species differences have been found in yolk corticosterone amounts in avian eggs. While some studies have failed to detect significant amounts of corticosterone, in other species high amounts have been recorded, such as in a recent study of southern rockhopper penguins Eudyptes chrysocome chrysocome. However, attention has been drawn recently to the fact that many antibodies for corticosterone measurement cross-react with other steroids present in the yolk. In particular, progesterone and related substances can occur in yolk in high concentrations, such that also low cross-reactions of corticosterone assays may lead to measurement errors. We thus performed high-performance liquid chromatographic (HPLC) analyses of yolk extracts and determined the concentration of immunoreactive corticosterone, as well as cross-reacting progesterone and cortisol in egg yolks of southern rockhopper penguins and imperial shags Phalacrocorax atriceps albiventer. We found that high gestagen concentrations in the yolk result in large measurement errors for yolk corticosterone, even when the cross-reactivity seems small. This was observed for both species. We further found species-specific differences in the actual corticosterone amounts present in the egg yolks.