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Prenatal resource allocation to offspring can be influenced by maternal environment and offspring value, and affect offspring survival. An important pathway for flexible maternal allocation is via egg components such as nutrients and hormones. In cooperative breeders, females with helpers may increase resource allocation to eggs—‘differential allocation’—or reduce it—‘load-lightening’. Yet, helper effects on egg composition have been poorly studied. Moreover, it is unknown how helpers’ presence modulates laying order effects on egg content and survival. Here, we investigated how maternal allocation varied with group size and laying order in the cooperatively breeding sociable weaver (Philetairus socius). We estimated interactive effects of helpers and laying order on allocation to egg mass, yolk nutrients—yolk mass, proteins, lipids, carotenoids, vitamin A and vitamin E—and hormones—testosterone, androstenedione, and corticosterone. Results concurred with the ‘differential allocation’ predictions. Females with more helpers produced later-laid eggs with heavier yolks and more lipids, and laid eggs overall richer in lipids. Proteins, antioxidants, and hormones were not found to vary with helper number. We then analyzed how helper number modulated laying order effects on survival. Females with more helpers did not specifically produce later-laid eggs with higher survival, but eggs laid by females with more helpers were overall more likely to fledge. These findings show that some egg components (yolk mass, lipids) can positively vary according to females’ breeding group size, which may improve offspring fitness.

Maternal effects are thought to be essential tools for females to modulate offspring development. The selective deposition of avian maternal hormones could therefore allow females to strategically adjust the phenotype of their offspring to the environmental situation encountered. However, at the time of egg formation, several contaminants are also transferred to the egg, including per- and polyfluoroalkyl substances (PFAS) which are ubiquitous organic contaminants with endocrine disrupting properties. It is, however, unknown if they can disrupt maternal hormone deposition. In this study we explored relationships between female PFAS burden and maternal deposition in the eggs of steroids (dihydrotestosterone, androstenedione and testosterone), glucocorticoids (corticosterone) and thyroid hormones (triiodothyronine and thyroxine) in a population of the Arctic-breeding black-legged kittiwake (Rissa tridactyla). Egg yolk hormone levels were unrelated to female hormone plasma levels. Second-laid eggs had significantly lower concentrations of androstenedione than first-laid eggs. Triiodothyronine yolk levels were decreasing with increasing egg mass but increasing with increasing females' body condition. Testosterone was the only transferred yolk hormone correlated to maternal PFAS burden: specifically, we found a positive correlation between testosterone in yolks and circulating maternal perfluorononanoic acid (PFNA), perfluorodecanoic acid (PFDcA) and perfluoroundecanoic acid (PFUnA) in first-laid eggs. This correlative study provides a first insight into the potential of some long-chain perfluoroalkyl acids to disrupt maternal hormones deposition in eggs and raises the question about the consequences of increased testosterone deposition on the developing embryo.

Adaptations resulting from co-evolutionary interactions between avian brood parasites and their hosts have been extensively studied, yet the physiological mechanisms underlying antiparasitic host defences remain little known. Prolactin, one of the main hormones involved in the regulation of avian parental behaviour, might play a key role in the orchestration of the host responses to avian brood parasitism. Given the positive association between prolactin and parental behaviour during incubation, decreasing prolactin levels are expected to facilitate egg-rejection decisions. We tested this prediction by implanting Eurasian blackbird (Turdus merula) females with an inhibitor of prolactin secretion, bromocriptine mesylate, to experimentally decrease their plasma prolactin levels. Bromocriptine mesylate-implanted individuals ejected mimetic model eggs at higher rates, and showed shorter latency to egg ejection, than placebo-treated birds. To our knowledge, this is the first experimental evidence that behavioural host defences against avian brood parasitism are mediated by prolactin.

International audience; In altricial species, parents brood their chicks constantly before leaving them unattended sometimes for extendedperiods when they become thermally independent. During this second phase, there is sometimes important interindividualdifferences in parental attendance and the fitness costs and benefits of parental strategies have previouslybeen extensively investigated. However, the impact of parental presence on offspring behaviors andstress physiology has been overlooked. Here, we examined the influence of parental presence on offspringhormonal and behavioral stress sensitivities in snow petrel chicks. We demonstrated for the first time in a wildbird species that attended chicks had lower stress-induced corticosterone levels and a lower probability to showdefensive behavior compared to the alone chicks. This reduced stress sensitivity is certainly explained by thewell-known link between corticosterone and nutritional status, and by the recent delivery of meals to theattended chicks and the improvement of their nutritional status. It may also be explained by the parental protectionagainst predators or inclement weather, or/and by the psychosocial comfort of parental presence for theoffspring. Overall, these results suggest that the presence of a parent in the nest reduces offspring stress sensitivityin wild birds. Further studies would now be required to disentangle the impact of nutritional status andparental presence on stress sensitivity and to better understand the potential impact of parental presence andcirculating corticosterone levels on growth and cognitive development in wild birds.

International audience; 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/un-handicapped) 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.

Environmental contaminants affect ecosystems worldwide and have deleterious effects on biota. Non-essential mercury (Hg) and lead (Pb) concentrations are well documented in some taxa and are described to cause multiple detrimental effects on human and wildlife. Additionally, essential selenium (Se) is known to be toxic at high concentrations but, at lower concentrations, Se can protect organisms against Hg toxicity. Crocodilians are known to bioaccumulate contaminants. However, the effects of these contaminants on physiological processes remain poorly studied. In the present study, we quantified Hg, Pb and Se concentrations in spectacled caimans (Caiman crocodilus) and investigated the effects of these contaminants on several physiological processes linked to osmoregulatory, hepatic, endocrine and renal functions measured through blood parameters in 23 individuals. Mercury was related to disruption of osmoregulation (sodium levels), hepatic function (alkaline phosphatase levels) and endocrine processes (corticosterone levels). Lead was related to disruption of hepatic functions (glucose and alanine aminotransferase levels). Selenium was not related to any parameters, but the Se:Hg molar ratio was positively related to the Na+ and corticosterone concentrations, suggesting a potential protective effect against Hg toxicity. Overall, our results suggest that Hg and Pb alter physiological mechanisms in wild caimans and highlight the need to thoroughly investigate the consequences of trace element contamination in crocodilians.

International audience; Glucocorticoids, androgens, and prolactin regulate metabolism and reproduction, but they also play critical roles in immunomodulation. Since the introduction of avian malaria to Hawaii a century ago, low elevation populations of the Hawaii Amakihi (Chlorodrepanis virens) that have experienced strong selection by avian malaria have evolved increased resilience (the ability to recover from infection), while high elevation populations that have undergone weak selection remain less resilient. We investigated how variation in malaria selection has affected corticosterone, testosterone, and prolactin hormone levels in Amakihi during the breeding season. We predicted that baseline corticosterone and testosterone (which have immunosuppressive functions) would be reduced in low elevation and malaria-infected birds, while stress-induced corticosterone and prolactin (which have immunostimulatory functions) would be greater in low elevation and malaria-infected birds. As predicted, prolactin was significantly higher in malaria-infected than uninfected females (although more robust sample sizes would help to confirm this relationship), while testosterone trended higher in malaria-infected than uninfected males and, surprisingly, neither baseline nor stress-induced CORT varied with malaria infection. Contrary to our predictions, stress-induced corticosterone was significantly lower in low than high elevation birds while testosterone in males and prolactin in females did not vary by elevation, suggesting that Amakihi hormone modulation across elevation is determined by variables other than disease selection (e.g., timing of breeding, energetic challenges). Our results shed new light on relationships between introduced disease and hormone modulation, and they raise new questions that could be explored in experimental settings.

Even though adaptations resulting from co-evolutionary interactions between avian brood parasites and their hosts have been well studied, the hormonal mechanisms underlying behavioural host defences remain largely unexplored. Prolactin, the main hormone mediating avian parental behaviour, has been hypothesized to play a key role in the orchestration of host responses to brood parasitic eggs. Based on the positive association between plasma prolactin and parental attachment to eggs, decreasing levels of this hormone are expected to facilitate egg-rejection decisions in parasitized clutches. We tested this prediction by implanting Eurasian blackbirds (Turdus merula) females with an inhibitor of prolactin secretion, bromocriptine mesylate, to experimentally low their prolactin levels. We found that bromocriptine-treated females rejected mimetic model eggs at higher rates than placebo-treated individuals. To our knowledge, this is the first experimental evidence that host responses to brood parasitism are mediated by the primary endocrine pathway that orchestrates the expression of avian parental care.

International audience; To maximize fitness, parents may trade-off time and energy between parental care and self-maintenance. In vertebrates, prolactin and corticosterone are two important hormones that regulate parental investment because they stimulate parental care and mobilize energy, respectively. Further, concentrations of both hormones change in response to disturbances. One of the most important parental care behaviors in birds is incubation, since small changes in egg temperature have large effects on offspring. We investigated how prolactin and corticosterone may mediate parental incubation constancy (i.e., the daily amount of time spent incubating eggs) and regulation of egg temperature. We collected blood samples from female wood ducks (Aix sponsa) near the start and end of the incubation period to measure baseline and stress-induced (30 min after capture and restraint) hormone concentrations. We also quantified incubation constancy and egg temperature using artificial egg temperature loggers. As expected, prolactin decreased and corticosterone increased after 30 min of capture and restraint. Corticosterone concentrations (baseline and stress-induced) were negatively related to body mass, but were not related to incubation constancy. In contrast, prolactin concentrations (baseline and stress-induced) were higher at the end than the start of the incubation period, and stress-induced prolactin concentrations were positively related to incubation constancy following a nest disturbance (i.e., capture). Further, prolactin (baseline and stress-induced) concentrations were positively related to egg temperatures, but only after the disturbance. These results suggest that prolactin may be associated with the regulation of parental incubation constancy and resulting heat-transfer after a disturbance, which may ultimately affect offspring development.

International audience; In the last decade, house sparrow populations have shown a general decline, especially in cities. Avian malaria has been recently suggested as one of the potential causes of this decline, and its detrimental effects could be exacerbated in urban habitats. It was initially thought that avian malaria parasites would not have large negative effects on wild birds because of their long co-evolution with their hosts. However, it is now well-documented that they can have detrimental effects at both the primo- and chronical infection stages. In this study, we examined avian malaria infection and its physiological and morphological consequences in four populations of wild house sparrows (2 urban and 2 rural). We did not find any relationship between the proportions of infected individuals and the urbanisation score calculated for our populations. However, we observed that the proportion of infected individuals increased during the course of the season, and that juveniles were less infected than adults. We did not detect a strong effect of malaria infection on physiological, morphological and condition indexes. Complex parasite dynamics and the presence of confounding factors could have masked the potential effects of infection. Thus, longitudinal and experimental studies are needed to understand the evolutionary ecology of this very common, but still poorly understood, wild bird parasite.

Future environmental variations linked to climate change are expected to influence precipitation regimes, and thus drinking water availability. Dehydration can be a particularly challenging physiological state for most organisms, yet no study has examined the effect of dehydration on the functioning of the Hypothalamus-Pituitary-Adrenal axis in wild endothermic animals, despite its central role in maintaining homeostasis. In this study, we experimentally imposed a temporary water shortage (∼20 hours) to captive House sparrows in order to investigate the consequences of short-term dehydration on baseline and stress-induced corticosterone levels. As expected, water-deprived birds displayed higher plasma osmolality and haematocrit. Additionally, water-deprived birds had lower defecation rates, suggesting that the mechanisms allowing cecal water absorption may be triggered very rapidly during water-deprivation. Baseline, but not stress-induced corticosterone levels were higher in water-deprived birds. Taken together, these results suggest that water-restriction may have critical consequences on several corticosterone-related traits such as energy budget (protein catabolism and possibly feeding reduction), enhanced mobility (to promote water acquisition) and potential responses to predators (thirst threshold overriding the acute stress response). Owing to the possible fitness consequences of such components of the day-to-day life of birds, future studies should aim at investigating the influence of future changes in precipitation regimes and drinking water availability on bird populations.

Life-history theory predicts that, to optimize their fitness, individuals should increase their reproductive effort as their residual reproductive value decreases. Accordingly, several studies have shown that individuals downregulate their glucocorticoid stress response (a proxy of reproductive investment in vertebrates) as they age, and as the subsequent reproductive value decreases. However, and surprisingly, results appear inconsistent, suggesting that the environmental context or the individual state may affect the relationship between age and reproductive effort. Here, we tested for the first time this hypothesis, and more specifically, whether this attenuation of the corticosterone stress response with advancing age depends on the energetic status of individuals. We compared the influence of age on the corticosterone stress response between fasting and non-fasting breeding snow petrels ( Pagodroma nivea ), an extremely long-lived bird. As expected, we found that the corticosterone stress response was attenuated in old petrels, but only when they were not fasting. Interestingly, this pattern was not apparent in fasting petrels, suggesting that old birds downregulate their corticosterone stress response and increase their parental investment only when they are in good body condition. At the ultimate level, old individuals may maintain a strong corticosterone stress response when fasting because the survival costs of increased stress resistance and parental effort might then outweigh their reproductive benefits.

International audience; In birds and other vertebrates, there is good evidence that females adjust the allocation of hormones in their eggs in response to prenatal environmental conditions, such as food availability or male phenotype, with profound consequences for life history traits of offspring. In insects, there is also evidence that females deposit juvenile hormones (JH) and ecdysteroids (ESH) in their eggs, hormones that play a key role in regulating offspring growth and metamorphosis. However, it is unclear whether females adjust their hormonal deposition in eggs in response to prenatal environmental conditions. Here we address this gap by conducting an experiment on the burying beetle Nicrophorus vespilloides, in which we manipulated the presence of the male parent and the size of the carcass used for breeding at the time of laying. We also tested for effects of the condition (i.e., body mass) of the parents. We then recorded subsequent effects on JH and ESH concentrations in the eggs. We found no evidence for an effect of these prenatal environmental conditions (male presence and carcass size) on hormonal concentration in the eggs. However, we found that females reduced their deposition of JH when mated with heavier males. This finding is consistent with negative differential allocation of maternal hormones in response to variation in the body mass of the male parent. We encourage further work to investigate the role of maternally derived hormones in insect eggs.

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.

Future environmental variations linked to climate change are expected to influence precipitation regimes and thus drinking water availability. Dehydration can be a particularly challenging physiological state for most organisms, yet no study has examined the effect of dehydration on the functioning of the hypothalamic-pituitary-adrenal axis in wild endothermic animals, despite its central role in maintaining homeostasis. In this study, we experimentally imposed a temporary water shortage (∼20 h) on captive house sparrows in order to investigate the consequences of short-term dehydration on baseline and stress-induced corticosterone levels. As expected, water-deprived birds displayed higher plasma osmolality and haematocrit. Additionally, water-deprived birds had lower defecation rates, suggesting that the mechanisms allowing caecal water absorption may be triggered very rapidly during water deprivation. Baseline but not stress-induced corticosterone levels were higher in water-deprived birds. Taken together, these results suggest that water restriction may have critical consequences on several corticosterone-related traits such as energy budget (protein catabolism and possibly feeding reduction), enhanced mobility (to promote water acquisition) and potential responses to predators (thirst threshold overriding the acute stress response). Owing to the possible fitness consequences of such components of the day-to-day life of birds, further studies should aim at investigating the influence of future changes in precipitation regimes and drinking water availability on bird populations.

International audience; Urban landscapes are associated with abiotic and biotic environmental changes thatmay result in potential stressors for wild vertebrates. Urban exploiters have physiological,morphological, and behavioral adaptations to live in cities. However, there isincreasing evidence that urban exploiters themselves can suffer from urban conditions,especially during specific life‐history stages. We looked for a link between thedegree of urbanization and the level of developmental stress in an urban exploiter(the house sparrow, Passer domesticus), which has recently been declining in multipleEuropean cities (e.g., London, UK). Specifically, we conducted a large‐scale study andsampled juvenile sparrows in 11 urban and rural sites to evaluate their feather corticosterone(CORT) levels. We found that juvenile feather CORT levels were positivelycorrelated with the degree of urbanization, supporting the idea that developinghouse sparrows may suffer from urban environmental conditions. However, we didnot find any correlation between juvenile feather CORT levels and body size, mass,or body condition. This suggests either that the growth and condition of urban sparrowsare not impacted by elevated developmental CORT levels, or that urban sparrowsmay compensate for developmental constraints once they have left the nest.Although feather CORT levels were not correlated with baseline CORT levels, wefound that feather CORT levels were slightly and positively correlated with the CORTstress response in juveniles. This suggests that urban developmental conditions maypotentially have long‐lasting effects on stress physiology and stress sensitivity in thisurban exploiter.

International audience; Per- and poly-fluoroalkyl substances (PFAS) raised increasing concerns over the past years due to their persistence and global distribution. Understanding their occurrence in the environment and their disruptive effect on the physiology of humans and wildlife remains a major challenge in ecotoxicological studies. Here, we investigate the occurrence of several carboxylic and sulfonic PFAS in 105 individuals of three seabird species (27 great black-backed gull Larus marinus; 44 lesser black-backed gull Larus fuscus graellsii; and 34 European herring gull Larus argentatus) from South western France. We further estimated the relationship between plasma concentrations of PFAS and i) the body condition of the birds and ii) plasma concentrations of thyroid hormone triiodothyronine (TT3). We found that great and lesser black-backed gulls from South Western France are exposed to PFAS levels comparable to highly contaminated species from other geographical areas, although major emission sources (i.e. related to industrial activities) are absent in the region. We additionally found that PFAS are negatively associated with the body condition of the birds in two of the studied species, and that these results are sex-dependent. Finally, we found positive associations between exposure to PFAS and TT3 in the great black-backed gull, suggesting a potential disrupting mechanism of PFAS exposure. Although only three years of data have been collected, we investigated PFAS trend over the study period, and found that great black-backed gulls document an increasing trend of plasma PFAS concentration from 2016 to 2018. Because PFAS might have detrimental effects on birds, French seabird populations should be monitored since an increase of PFAS exposure may impact on population viability both in the short- and long-term.

International audience; Artificial light at night (ALAN) can disrupt adaptive patterns of physiology and behavior that promote high fitness, resulting in physiological stress and elevation of steroid glucocorticoids (corticosterone, CORT in birds). Elevated CORT may have particularly profound effects early in life, with the potential for enduring effects that persist into adulthood. Research on the consequences of early-life exposure to ALAN remains limited, especially outside of the laboratory, and the effects of early-life light exposure on CORT concentrations in wild nestling birds remain to be elucidated. We used an experimental setup to test the hypothesis that ALAN elevates CORT concentrations in developing free-living birds, by exposing nestling great tits (Parus major) to ALAN inside nest boxes. We measured CORT in feathers grown over the timeframe of the experiment (7 nights), such that CORT concentrations represent an integrative metric of hormone release over the period of nocturnal light exposure, and of development. We also assessed the relationships between feather CORT concentrations, body condition, nestling size rank and fledging success. In addition, we evaluated the relationship between feather CORT concentrations and telomere length. Nestlings exposed to ALAN had higher feather CORT concentrations than control nestlings, and nestlings in poorer body condition and smaller brood members also had higher CORT. On the other hand, telomere length, fledging success, and recruitment rate were not significantly associated with light exposure or feather CORT concentrations. Results indicate that exposure to ALAN elevates CORT concentrations in nestlings, which may reflect physiological stress. In addition, the organizational effects of CORT are known to be substantial. Thus, despite the lack of effect on telomere length and survivorship, elevated CORT concentrations in nestlings exposed to ALAN may have subsequent impacts on later-life fitness and stress sensitivity.

Parasites often prompt sub-lethal costs to their hosts by eliciting immune responses. These costs can be hard to quantify but are crucial to our understanding of the host's ecology. Energy is a fundamental currency to quantify these costs, as energetic trade-offs often exist between key fitness-related processes. Daily energy expenditure (DEE) comprises of resting metabolic rate (RMR) and energy available for activity, which are linked via the energy management strategy of an organism. Parasitism may play a role in the balance between self-maintenance and activity, as immune costs can be expressed in elevated RMR. Therefore, understanding energy use in the presence of parasitism enables mechanistic elucidation of potential parasite costs. Using a gradient of natural parasite load and proxies for RMR and DEE in a wild population of breeding European shags (Phalacrocorax aristotelis), we tested the effect of parasitism on maintenance costs as well as the relationship between proxies for RMR and DEE. We found a positive relationship between parasite load and our RMR proxy in females but not males, and no relationship between proxies for RMR and DEE. This provides evidence for increased maintenance costs in individuals with higher parasite loads and suggests the use of an allocation energy management strategy, whereby an increase to RMR creates restrictions on energy allocation to other activities. This is likely to have fitness consequences as energy allocated to immunity is traded off against reproduction. Our findings demonstrate that understanding energy management strategies alongside fitness drivers is central to understanding the mechanisms by which these drivers influence individual fitness., Summary: Demonstration of the sub-lethal costs of endo-parasitism for a wild population of seabirds via increased maintenance metabolism.

Urban landscapes are associated with abiotic and biotic environmental changes that may result in potential stressors for wild vertebrates. Urban exploiters have physiological, morphological, and behavioral adaptations to live in cities. However, there is increasing evidence that urban exploiters themselves can suffer from urban conditions, especially during specific life‐history stages. We looked for a link between the degree of urbanization and the level of developmental stress in an urban exploiter (the house sparrow, Passer domesticus), which has recently been declining in multiple European cities (e.g., London, UK). Specifically, we conducted a large‐scale study and sampled juvenile sparrows in 11 urban and rural sites to evaluate their feather corticosterone (CORT) levels. We found that juvenile feather CORT levels were positively correlated with the degree of urbanization, supporting the idea that developing house sparrows may suffer from urban environmental conditions. However, we did not find any correlation between juvenile feather CORT levels and body size, mass, or body condition. This suggests either that the growth and condition of urban sparrows are not impacted by elevated developmental CORT levels, or that urban sparrows may compensate for developmental constraints once they have left the nest. Although feather CORT levels were not correlated with baseline CORT levels, we found that feather CORT levels were slightly and positively correlated with the CORT stress response in juveniles. This suggests that urban developmental conditions may potentially have long‐lasting effects on stress physiology and stress sensitivity in this urban exploiter.

International audience; In wild vertebrates, conspicuousness often signals a high phenotypic quality and is therefore associated with fitness benefits (“the handicap principle hypothesis”). However, conspicuous individuals usually face a higher risk of predation because they are easier to detect. Therefore, conspicuous individuals are expected to modify their behavioural and their physio logical stress responses to limit the negative effect of their conspicuousness on survival. We examined the link between plumage coloration and the stress response in a bird species (the rock pigeon, Columbia livia) by comparing two groups of pigeons with different plumage characteristics: ‘blue-bar’ birds (lighter birds) and ‘checker’ birds (darker birds). Specifically, we measured several behavioural and physiological components of the stress response (breath rate, corticosterone, and tonic immobility). Although breath rate did not differ between ‘checker’ and ‘blue bar’ birds, the corticosterone stress response of ‘blue-bar’ birds was greater relative to that of ‘checker’ birds. Moreover, ‘checker’ birds were more likely to initiate tonic immobility relative to ‘blue bar’ birds. Therefore, this study suggests that the behavioural and physiological stress responses of wild birds may differ according to their plumage coloration. To conclude, our results suggest that plumage colorations may be functionally linked to specific behavioral and physiological sensitivities to stressors.

The poorly known winter foraging ecology of the king penguin, a major Southern Ocean consumer, was investigated at the subantarctic Crozet Islands where the largest global population breeds. Blood δ13C and δ15N values were used as proxies of the birds’ foraging habitat and diet, respectively, and circulating prolactin levels helped in determining the birds’ reproductive status. Plasma prolactin concentrations showed that king penguin adults of unknown breeding status (n = 52) that were present at the colony in winter were in fact breeders and failed breeders, but were not non ‐breeders. Circulating prolactin was neither related to δ13C nor δ15N values, thus suggesting that both breeders and failed breeders used the same foraging habitats and fed on the same prey. Plasma and blood cell isotopic values depicted four new relevant biological features on the feeding strategies of king penguins during the critical winter period: (1) 42% of the birds foraged in the distant Antarctic Zone, but 58% fed primarily in subantarctic waters (δ13C), (2) they preyed upon myctophids in both zones (δ15N), (3) individuals were consistent in their foraging strategies over the winter months (δ13C and δ15N), and (4) a higher proportion of females (77%–80%) than males (27%–31%) favored feeding in distant Antarctic waters (δ13C). This study highlights trophic connectivity between subantarctic and Antarctic ecosystems and hence the key role of energy export from Antarctic waters to sustain breeding populations of subantarctic predators, including during the Austral winter.

International audience; In a rapidly urbanizing world, trace element pollution may represent a threat to human health and wildlife, and it is therefore crucial to assess both exposition levels and associated effects of trace element contamination on urban vertebrates. In this study, we investigated the impact of urbanization on trace element contamination and stress physiology in a wild bird species, the common blackbird (Turdus merula), along an urbanization gradient (from rural to moderately urbanized areas). Specifically, we described the contamination levels of blackbirds by 4 non-essential (Ag, Cd, Hg, Pb) and 9 essential trace elements (As, Co, Cr, Cu, Fe, Mn, Ni, Se, Zn), and explored the putative disrupting effects of the non-essential element contamination on corticosterone levels (a hormonal proxy for environmental challenges). We found that non-essential trace element burden (Cd and Pb specifically) increased with increasing urbanization, indicating a significant trace element contamination even in medium sized cities and suburban areas. Interestingly, the increased feather non-essential trace element concentrations were also associated with elevated feather corticosterone levels, suggesting that urbanization probably constrains birds and that this effect may be mediated by trace element contamination. Future experimental studies are now required to disentangle the influence of multiple urban-related constraints on corticosterone levels and to specifically test the influence of each of these trace elements on corticosterone secretion.

International audience; In the context of global change, the physiological and hormonal stress responses have received much attentionbecause of their implications in terms of allostasis. However, most studies have focused on glucocorticoidsonly as the “common” response to stressorswhile neglecting other endocrine axes and hormones (e.g. prolactin,thyroid hormones) that play a crucial role in metabolic adjustments. Interestingly, the responsiveness of all theseendocrine axes to stress may depend on the energetic context and this context-dependent stress response hasbeen overlooked so far. In the wild, temperature can vary to a large extent within a short time window and ambienttemperature may affect these metabolic-related endocrine axes, and potentially, their responsiveness to anacute stressor. Here,we explicitly tested this hypothesis by examining the effect of a standardized stress protocolon multiple hormonal responses in the rock pigeon (Columbia livia). We tested the effect of an acute restraintstress on (1) corticosterone levels, (2) prolactin levels, and (3) thyroid hormone levels (triiodothyronine,thyroxine) in pigeons that were held either at cool temperature (experimental birds) or at room temperature(control birds) during the stress protocol. Although we found a significant influence of restraint stress on mosthormone levels (corticosterone, prolactin, and thyroxine), triiodothyronine levels were not affected by the restraintstress. This demonstrates that stressors can have significant impact on multiple endocrine mechanisms.Importantly, all of these hormonal responses to stress were not affected by temperature, demonstrating thatthe exposure to cold temperature does not affect the way these hormone levels change in response to handlingstress. This suggests that some endocrine responses to temperature decreases may be overridden by the endocrineresponses to an acute restraint stress.

International audience; Understanding sources of reproductive variation can inform management and conservation decisions, population ecology and life-history theory. Annual reproductive variation can drive population growth rate and can be influenced by factors from across the annual cycle (known as carry-over effects). The majority of studies, however, focus solely on the role of current environmental events. Past events often influence future reproductive decisions and success but can be logistically difficult to collect and quantify, especially in migratory species. Recent work indicates that glucocorticoids may prove good indicators to evaluate carry-over effects across life-history transitions. Here, we evaluated three different measures of glucocorticoid physiology (feathers, faeces and plasma) to evaluate the predictability of future breeding decision in the harlequin duck (Histrionicus histrionicus). We collected tail and back feathers, plasma and faeces for glucocorticoid analysis, and fitted female harlequin ducks with very high-frequency transmitters to track their breeding decisions. Both back feathers (moulted immediately before the current season) and faecal glucocorticoid metabolites were identified as important predictive factors of reproductive decisions; high concentrations of glucocorticoid metabolites in back feathers and faeces predicted a higher likelihood of reproductive deferral for the year. Although back and tail feather corticosterone concentrations were correlated, tail feathers (moulted at the end of the previous breeding season) did not predict breeding decisions. Plasma corticosterone concentrations were collected over too broad a time range after capture to be useful in this study. This study demonstrates the utility of non-invasive corticosterone metrics in predicting breeding decisions and supports the use of feathers to measure carry-over effects in migratory birds. With this technique, we identified the prenuptial moult as an important life-history phase that contributes to reproductive decisions. Identification of critical life-history phases is paramount to efficient management of species.

International audience; The hormonal stress response is flexible and can be modulated by individuals according to its costs and benefits. Therefore, it is predicted that parents in poor body condition should modify their hormonal stress response, and thus, redirect energy allocation processes from parental care to self-maintenance when stressors occur. To test this prediction, most studies on free-living vertebrates have only focused on the stress response while the stress recovery – how quickly hormonal levels return to baseline values – has been neglected. Moreover, most studies have only focused on corticosterone – the primary mediator of allostasis – without paying attention to prolactin despite its major role in mediating parental behaviors. Here, we examined the effect of a short-termfasting event on the corticosterone and prolactin stress responses and recoveries, andwe subsequently explored their relationshipswith parental decision in the snowpetrel (Pagodroma nivea). By comparing the hormonal profiles of fasting and non-fasting snow petrels, we showed that parents modulate their corticosterone (but not prolactin) stress response according to their energetic status. We also described for the first time the hormonal stress recoveries in wild birds and found that they did not differ between fasting and non-fasting birds. Importantly, egg neglect was negatively correlated with circulating prolactin but not corticosterone levels in this species, demonstrating therefore a complex link between body condition, parental behavior and circulating corticosterone and prolactin levels. Wesuggest that both corticosterone and prolactin play a major role in theway parents adjust to stressors. This multiple signaling may allowparents to fine-tune their response to stressors, and especially, to activate specific allostasis-related mechanisms in a timely manner.

PubMedID: 22504343 One of the fundamental principles of the life-history theory is that parents need to balance their resources between current and future offspring. Deserting the dependent young is a radical life-history decision that saves resources for future reproduction but that may cause the current brood to fail. Despite the importance of desertion for reproductive success, and thus fitness, the neuroendocrine mechanisms of brood desertion are largely unknown. We investigated two candidate hormones that may influence brood desertion in the Kentish plover Charadrius alexandrinus: prolactin ('parental hormone') and corticosterone ('stress hormone'). Kentish plovers exhibit an unusually diverse mating and parental care system: brood desertion occurs naturally since either parent (the male or the female) may desert the brood after the chicks hatch and mate with a new partner shortly after. We measured the hormone levels of parents at hatching using the standard capture and restraint protocol. We subsequently followed the broods to determine whether a parent deserted the chicks. We found no evidence that either baseline or stress-induced prolactin levels of male or female parents predicted brood desertion. Although stress-induced corticosterone levels were generally higher in females compared with males, individual corticosterone levels did not explain the probability of brood desertion. We suggest that, in this species, low prolactin levels do not trigger brood desertion. In general, we propose that the prolactin stress response does not reflect overall parental investment in a species where different parts of the breeding cycle are characterized by contrasting individual investment strategies. © 2012 Elsevier Inc.. Deutscher Akademischer Austauschdienst OMFB-00365/2010, 22895WE Hungarian Scientific Research Fund Hungarian Scientific Research Fund: K81953, PD76862 TUBITAK-108Y329, OMFB-00581/2009 The study was supported by the Hungarian Scientific Research Fund (OTKA, K81953 , PD76862 ) and the Hungarian-French Intergovernmental S&T Cooperation Programme (OMFB-00365/2010, Egide 22895WE). Fieldwork was supported by a grant from the Hungarian–Turkish Intergovernmental S&T Cooperation Programme ( OMFB-00581/2009 , TUBITAK-108Y329 ). CK was supported by a DAAD fellowship. At the CEBC, we thank Colette Trouvé for her technical assistance in performing the prolactin and corticosterone assays. We are grateful to Peter Sharp and Alexander Badyaev for discussion and useful advice on the methods. We are grateful for two anonymous referees for their excellent comments that improved the manuscript.

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.

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.