In birds, the use of corticosterone (Cort) implants is a frequent tool aimed at simulating systemic elevations of this hormone and studying effects on biological traits (e.g. physiology, morphology, behavior). This manipulation may alter adrenocortical function, potentially changing both baseline (Cort) and stress-induced (Cort) plasma Cort levels. However, implant effects on the latter trait are rarely measured, disregarding downstream consequences of potentially altered stress responses. Here, we analyzed the effects of Cort implants on both Cort and Cort in nestling and adult European white storks, In addition, we performed a review of 50 studies using Cort implants in birds during the last two decades to contextualize stork results, assess researchers' patterns of use and infer current study biases. High and low doses of Cort implants resulted in a decrease of both Cort (31-71% below controls) and Cort (63-79% below controls) in storks. Our literature review revealed that Cort generally increases (72% of experiments) whereas Cort decreases (78% of experiments) following implant treatment in birds. Our results challenge and expand the prevailing assumption that Cort implants increase circulating Cort levels because: (i) Cort levels show a quadratic association with implant dose across bird species, and decreased levels may occur at both high and low implant doses, and (ii) Cort implants also decrease Cort levels, thus producing stress-hyporesponsive phenotypes. It is time to work towards a better understanding of the effects of Cort implants on adrenocortical function, before addressing downstream links to variation in other biological traits.
Soaring landbirds typically exploit atmospheric uplift as they fly overland, displaying a highly effective energy-saving locomotion. However, large water bodies lack thermal updrafts, potentially becoming ecological barriers that hamper migration. Here we assessed the effects of a sea surface on the migratory performance of GPS-tagged white storks (Ciconia ciconia) before, during and after they crossed the straits of Gibraltar. Oversea movements involved only flapping and gliding and were faster, traversed in straighter, descending trajectories and resulted in higher movement-related energy expenditure levels than overland, supporting the water barrier hypothesis. Overland movements at both sides of the sea straits resulted in tortuous routes and ascending trajectories with pre-crossing flights showing higher elevations and more tortuous routes than post-crossing, thus supporting the barrier negotiation hypothesis. Individual positions at both ends of the sea narrow were predicted by zonal winds and storks´ location at entry in the European hinterland, and birds did not show compensational movements overland in anticipation to subsequent wind displacements oversea. The length of the water narrow at departure shore, the elevation therein and the winds on route affected major components of sea crossing performance (such as distances and times overwater, minimum elevations, climb angles, speeds and energy expenditure), supporting the departure position and oversea winds hypotheses. In summary, our study provides a prime example at high temporal resolution of how birds adjust their behavior and physiology as they interact with the changing conditions of the travelling medium, reallocating resources and modifying their movement to overcome an ecological barrier.
In birds, the magnitude of the adrenocortical stress response can be down‐regulated during specific life‐history stages. Such modulation likely occurs when the effects of mounting robust corticosterone (Cort) elevations interfere with the normal progression of critical lifecycle activities (e.g. development, molt, migration, reproduction). The developmental hypothesis posits that altricial birds should display a ‘stress hyporesponsive period’ during the early post‐natal life stages, characterized by reduced adrenocortical stress responses compared to adult birds and a gradual age‐related increase. Such modulation would allow avoiding the potential deleterious effects that long‐term elevations of circulating Cort might exert on growth and development, when the physiological and behavioral abilities to cope with disturbance are limited. Two proximate hypotheses have been proposed to explain this age‐dependent pattern of Cort secretion. The ‘maturation hypothesis’ proposes a progressive age‐related growth, maturation and enhanced sensitivity to sensory input of the Hypothalamic‐Pituitary‐Adrenal (HPA) axis tissues, whereas the ‘negative feedback attenuation hypothesis’ proposes a gradual attenuation in the intensity of the negative feedback in the HPA axis. Here we tested these hypotheses by experimentally inducing negative feedback on the HPA axis via dexamethasone (DEX) treatment in nestling white storks Ciconia ciconia. Nestling age positively affected stress‐induced plasma Cort (STRESS‐Cort) levels during experimental handling and restraint, thus supporting the developmental hypothesis. DEX treatment significantly reduced STRESS‐Cort levels compared to saline (SAL) treatment, thus eliciting the expected negative feedback on the HPA axis. However, inter‐ and intra‐individual comparisons indicated no age effects on the intensity of the negative feedback exerted by DEX. Our results do not support the negative feedback attenuation hypothesis and suggest that progressive maturation of the HPA axis tissues is the proximate mechanism responsible for age‐related changes in the stress response during avian post‐natal development. We encourage further tests of the proposed proximate mechanisms during migration, breeding and molt.
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