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Physiological Adaptations for Breeding in Birdsis the most current and comprehensive account of research on avian reproduction. It develops two unique themes: the consideration of female avian reproductive physiology and ecology, and an emphasis on individual variation in life-history traits. Tony Williams investigates the physiological, metabolic, energetic, and hormonal mechanisms that underpin individual variation in the key female-specific reproductive traits and the trade-offs between these traits that determine variation in fitness. The core of the book deals with the avian reproductive cycle, from seasonal gonadal development, through egg laying and incubation, to chick rearing. Reproduction is considered in the context of the annual cycle and through an individual's entire life history. The book focuses on timing of breeding, clutch size, egg size and egg quality, and parental care. It also provides a primer on female reproductive physiology and considers trade-offs and carryover effects between reproduction and other life-history stages. In each chapter, Williams describes individual variation in the trait of interest and the evolutionary context for trait variation. He argues that there is only a rudimentary, and in some cases nonexistent, understanding of the physiological mechanisms that underpin individual variation in the major reproductive life-history traits, and that research efforts should refocus on these key unresolved problems by incorporating detailed physiological studies into existing long-term population studies, generating a new synthesis of physiology, ecology, and evolutionary biology.

Twenty years ago, Albert Bennett published a paper in the influential book New directions in ecological physiology arguing that individual variation was an 'underutilized resource'. In this paper, I review our state of knowledge of the magnitude, mechanisms and functional significance of phenotypic variation, plasticity and flexibility in endocrine systems, and argue for a renewed focus on inter-individual variability. This will provide challenges to conventional wisdom in endocrinology itself, e.g. re-evaluation of relatively simple, but unresolved questions such as structure-function relationships among hormones, binding globulins and receptors, and the functional significance of absolute versus relative hormone titres. However, there are also abundant opportunities for endocrinologists to contribute solid mechanistic understanding to key questions in evolutionary biology, e.g. how endocrine regulation is involved in evolution of complex suites of traits, or how hormone pleiotropy regulates trade-offs among life-history traits. This will require endocrinologists to embrace the raw material of adaptation (heritable, individual variation and phenotypic plasticity) and to take advantage of conceptual approaches widely used in evolutionary biology (selection studies, reaction norms, concepts of evolutionary design) as well as a more explicit focus on the endocrine basis of life-history traits that are of primary interest to evolutionary biologists (cf. behavioural endocrinology).

1. Glucocorticoids hormones (GCs) are intuitively important for mediation of age-dependent vertebrate life-history transitions through their effects on ontogeny alongside underpinning variation in life-history traits and trade-offs in vertebrates. These concepts largely derive from the ability of GCs to alter energy allocation, physiology and behaviour that influences key life-history traits involving age-specific life-history transitions, reproduction and survival. 2. Studies across vertebrates have shown that the neuroendocrine stress axis plays a role in the developmental processes that lead up to age-specific early life-history transitions. While environmental sensitivity of the stress axis allows for it to modulate the timing of these transitions within species, little is known as to how variation in stress axis function has been adapted to produce interspecific variation in the timing of life-history transitions. 3. Our assessment of the literature confirms that of previous reviews that there is only equivocal evidence for correlative or direct functional relationships between GCs and variation in reproduction and survival. We conclude that the relationships between GCs and life-history traits are complex and general patterns cannot be easily discerned with current research approaches and experimental designs. 4. We identify several future research directions including: (i) integration of proximate and ultimate measures, including longitudinal studies that measure effects of GCs on more than one life-history trait or in multiple environmental contexts, to test explicit hypotheses about how GCs and life-history variation are related and (ii) the measurement of additional factors that modulate the effects of GCs on life-history traits (e.g. GC receptors and binding protein levels) to better infer neurendocrine stress axis actions. 5. Conceptual models of HPA/I axis actions, such as allostatic load and reactive scope, to some extent explicitly predict the role of GCs in a life-history context, but are descriptive in nature. We propose that GC effects on life-history transitions, survival probabilities and fecundity can be modelled in existing quantitative demographic frameworks to improve our understanding of how GC variation influences life-history evolution and GC-mediated effects on population dynamics

The question of why maternal stress influences offspring phenotype is of significant interest to evolutionary physiologists. Although embryonic exposure to maternally derived glucocorticoids (i.e., corticosterone) generally reduces offspring quality, effects may adaptively match maternal quality with offspring demand. We present results from an interannual field experiment in European starlings (Sturnus vulgaris) designed explicitly to examine the fitness consequences of exposing offspring to maternally derived stress hormones. We combined a manipulation of yolk corticosterone (yolk injections) with a manipulation of maternal chick‐rearing ability (feather clipping of mothers) to quantify the adaptive value of corticosterone‐induced offspring phenotypes in relation to maternal quality. We then examined how corticosterone‐induced “matching” within this current reproductive attempt affected future fecundity and maternal survival. First, our results provide support that low‐quality mothers transferring elevated corticosterone to eggs invest in daughters as predicted by sex allocation theory. Second, corticosterone‐mediated sex‐biased investment resulted in rapid male‐biased mortality resulting in brood reduction, which provided a better match between maternal quality and brood demand. Third, corticosterone‐mediated matching reduced investment in current reproduction for low‐quality mothers, resulting in fitness gains through increased survival and future fecundity. Results indicate that the transfer of stress hormones to eggs by low‐quality mothers can be adaptive since corticosterone‐mediated sex‐biased investment matches the quality of a mother to offspring demand, ultimately increasing maternal fitness. Our results also indicate that the branding of the proximate effects of maternal glucocorticoids on offspring as negative ignores the possibility that short‐term phenotypic changes may actually increase maternal fitness.

Female birds incur costs associated with increased egg production, including reductions in chick provisioning ability, in future fecundity, in survival, and in egg and chick viability. It should be possible to identify the components of the physiological system underlying reproduction, or the specific reproductive traits themselves, that explain these costs, but this has proved to be difficult, in part because of marked, but unexplained, individual variation in these traits. Resolving the physiological and evolutionary consequences of this individual variation represents an exciting challenge for the future. Several mechanisms have been proposed for the cost of egg production (e.g., protein depletion and impaired flight muscle function; immunosuppression), which assume relatively simple resource-allocation trade-offs. I argue that such mechanisms provide an unsatisfactory explanation for costs that can occur over months or even years. A more productive approach for future research will be to focus on hormonally mediated, non-resource-based costs of egg production caused by pleiotropic effects of reproductive hormones that can operate over the longer time scales at which costs of reproduction are expressed.

Glucocorticoid (GC) hormones are important phenotypic mediators across vertebrates, but their circulating concentrations can vary markedly. Here we investigate macroevolutionary patterning in GC levels across tetrapods by testing seven specific hypotheses about GC variation and evaluating whether the supported hypotheses reveal consistent patterns in GC evolution. If selection generally favors the \"supportive\" role of GCs in responding effectively to challenges, then baseline and/or stress-induced GCs may be higher in challenging contexts. Alternatively, if selection generally favors \"protection\" from GC-induced costs, GCs may be lower in environments where challenges are more common or severe. The predictors of baseline GCs were all consistent with supportive effects: levels were higher in smaller organisms and in those inhabiting more energetically demanding environments. During breeding, baseline GCs were also higher in populations and species with fewer lifetime opportunities to reproduce. The predictors of stress-induced GCs were instead more consistent with the protection hypothesis: during breeding, levels were lower in organisms with fewer lifetime reproductive opportunities. Overall, these patterns indicate a surprising degree of consistency in how some selective pressures shape GCs across broad taxonomic scales; at the same time, in challenging environments selection appears to operate on baseline and stress-induced GCs in distinct ways.

It has long been recognized that mass loss during breeding could be adaptive (e.g., by ameliorating the costs of increased parental activity). However, many studies still commonly interpret mass loss as evidence of “stress” or a cost of reproduction (i.e., a negative effect of high workload during chick provisioning). Despite several studies reporting evidence in support of both hypotheses, the ecological and/or life‐history contexts under which mass loss may be viewed as a “cost” or an adaptive strategy are still unclear. Here, we used a long‐term dataset from a breeding population of European starlings (Sturnus vulgaris) to investigate natural annual and individual variation in body mass and mass loss and to test whether mass loss during chick rearing represents a phenotypically plastic trait that varies predictably in relation to ecological context and individual quality. While there was significant annual variation in incubation mass, chick‐rearing mass, and mass change, there were no systematic relationships between mass loss and current breeding success or future fecundity and survival. In addition, we found no evidence of intra‐annual repeatability of mass loss between first and second broods ( = .00) but moderate interannual repeatability of mass loss (R = .61) during first broods, suggesting differences in mass loss under different selective pressures. However, we found no covariation between residual intra‐individual variation in mass loss for first broods and other reproductive or life‐history traits. We therefore found no support for the idea that mass loss reflects “reproductive stress” in our system: there were no negative relationships between mass loss and either current or future reproduction and survival (local return rate). Our results are consistent with mass loss being an individually plastic trait, with individuals using mass loss to “level the playing field” and individually optimize reproductive effort and fitness within their specific ecological context and relative to their individual quality for a given breeding attempt. It has long been recognized that avian mass loss during breeding could be adaptive, though many studies still commonly interpret mass loss as evidence of “stress” or a cost of reproduction. Here, we used a long‐term dataset from a breeding population of European starlings to investigate the ecological and/or life‐history contexts under which mass loss may be viewed as a “cost” or an adaptive strategy and, specifically, test whether mass loss during chick rearing represents a phenotypically plastic trait that varies predictably in relation to ecological context and individual quality. Overall, we found no support for the idea that mass loss reflects “reproductive stress” in our system and instead suggest that mass loss may represent an individually plastic trait that individuals use to “level the playing field” and individually optimize reproductive effort and fitness.

Sexual segregation in vertebrate foraging niche is often associated with sexual size dimorphism (SSD), i.e., ecological sexual dimorphism. Although foraging behavior of male and female seabirds can vary markedly, differences in isotopic (carbon, δ13C and nitrogen, δ15N) foraging niche are generally more pronounced within sexually dimorphic species and during phases when competition for food is greater. We examined ecological sexual dimorphism among sympatric nesting Pygoscelis penguins asking whether environmental variability is associated with differences in male and female pre-breeding foraging niche. We predicted that all Pygoscelis species would forage sex-specifically, and that higher quality winter habitat, i.e., higher or lower sea ice coverage for a given species, would be associated with a more similar foraging niche among the sexes. P2/P8 primers reliably amplified DNA of all species. On average, male Pygoscelis penguins are structurally larger than female conspecifics. However, chinstrap penguins were more sexually dimorphic in culmen and flipper features than Adélie and gentoo penguins. Adélies and gentoos were more sexually dimorphic in body mass than chinstraps. Only male and female chinstraps and gentoos occupied separate δ15N foraging niches. Strong year effects in δ15N signatures were documented for all three species, however, only for Adélies, did yearly variation in δ15N signatures tightly correlate with winter sea ice conditions. There was no evidence that variation in sex-specific foraging niche interacted with yearly winter habitat quality. Chinstraps were most sexually size dimorphic followed by gentoos and Adélies. Pre-breeding sex-specific foraging niche was associated with overall SSD indices across species; male chinstrap and gentoo penguins were enriched in δ15N relative to females. Our results highlight previously unknown trophic pathways that link Pygoscelis penguins with variation in Southern Ocean sea ice suggesting that each sex within a species should respond similarly in pre-breeding trophic foraging to changes in future winter habitat.

Events on the non‐breeding grounds and on migration can influence the timing of reproduction and the productivity of migratory songbirds. We show, using data from a 12‐year study in Revelstoke, BC, Canada, that the onset of breeding in yellow warblers (Setophaga petechia) is linked to weather conditions on migration, specifically the speed of crosswinds experienced over the western flyway during a 2‐week period from May 18 to 31. During 2015–2017, we investigated whether this carryover effect was due to crosswind effects on the timing of arrival, the reproductive state and mass of females on arrival, or a combination of other effects that delayed egg laying. In these years, female arrival dates varied with age but were independent of year, growing degree days, or crosswinds on migration. Though crosswinds experienced by females during the 14‐day period before their arrival on the breeding ground had no significant effect on their reproductive state (plasma triglyceride levels) or mass (controlling for tarsus length) on arrival, crosswinds had an effect on the time interval between arrival and egg laying. The time interval between arrival and egg laying was also shorter if females arrived with elevated plasma triglyceride levels and longer if females arrived early in the season. Carryover effects from crosswinds experienced during migration on the timing of breeding and reproductive success of yellow warblers are likely to arise due to their effect on arrival date and how rapidly birds can transition from a migratory to a reproductive physiological state. Both wind‐speed experienced on migration (14 days before arrival) and reproductive state on arrival (plasma triglyceride concentration) independently influenced the time interval between arrival and egg laying. Crosswinds that delay breeding have a significant effect on reproductive success; female yellow warblers that initiate reproduction late decrease both their chance to raise at least one nestling and the number of nestlings fledged. Carryover effects from crosswinds experienced during migration on the timing of breeding and reproductive success of yellow warbler (Setophaga petechia) are likely to arise due to their effect on arrival date and how rapidly birds can transition from a migratory to a reproductive physiological state. Both wind‐speed experienced on migration (14 days before arrival) and reproductive state on arrival (plasma triglyceride concentration) independently influenced the time interval between arrival and egg laying.