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In long‐distance migrants, preparation for migration is typically associated with increases in fat and body mass, and with an enlargement of pectoralis muscle mass that likely improves flight performance. Although changes in muscle mass or size have been well described in migratory birds, potential changes in muscle ultrastructure during this transition still deserves scrutiny. Using outdoor captive snow buntings (Plectrophenax nivalis n = 15) measured during their transition into a spring migratory phenotype as a model system, we studied changes in pectoralis muscle ultrastructure and predicted that muscle fiber diameter could increase in parallel with the gain in body mass. We also expected that larger fibers could either recruit satellite cells to support cellular maintenance and protein turnover, increase myonuclear domain (cytoplasm per nuclei) with a potential increase in protein turnover load per myonucleus, or existing myonuclei could undergo endoreduplication. Buntings increased body mass by 46% within a month, largely due to a > 6‐fold increase in body fat. However, this increase in body mass was also associated with a 36% increase in muscle fiber diameter. Both pectoralis muscle mass (r2 = 0.57–0.77) and fiber diameter (r2 = 0.32) correlated with total body mass, without any change in the number of nuclei per fiber. Consequently, variation in myonuclear domain (i.e. the amount of cytoplasm per nucleus), was also positively associated with body mass (r2 = 0.51). Therefore, buntings preparing for migration may experience an increase in muscle contraction force due to larger muscle fibers, but this is also coupled with increases in myonuclear domain, which may force these cells to increase protein production to safeguard satellite cells. [ABSTRACT FROM AUTHOR]

Habitat selection theory suggests that environmental features selected at coarse scales reveal fundamental factors affecting animal fitness. When these factors vary across seasons, they may lead to large-scale movements, including long-distance seasonal migrations. We analyzed the seasonal habitat selection of 25 satellite-tracked Arctic hares from a population on Ellesmere Island (Nunavut, Canada) that relocated over 100 km in the fall. Since no other lagomorph is known to perform such extensive movements, this population offered an ideal setting to test animal movement and habitat selection theory. On summer grounds hares selected low elevation areas, while on winter grounds they selected high vegetation biomass, high elevation, and steep slopes. During fall relocation, they alternated between stopover and traveling behavioral states (ratio 2:1). Stopover locations were characterized by higher vegetation heterogeneity and lower rugosity than traveling locations, while vegetation biomass and elevation interacted to explain stopover locations in a more complex way. The selected combination of environmental features thus varied across seasons and behavioral states, in a way broadly consistent with predictions based on the changing food and safety needs of hares. Although causality was not demonstrated, our results improve our understanding of long-distance movements and habitat selection in Arctic hares, as well as herbivore ecology in the polar desert. Results also provide strong support to animal movement and habitat selection theory, by showing how some important hypotheses hold when tested in a species phylogenetically distinct from most animal models used in this research field. [ABSTRACT FROM AUTHOR]

Phenotypic flexibility has received considerable attention in the last decade; however, whereas many studies have reported amplitude of variation in phenotypic traits, much less attention has focused on the rate at which traits can adjust in response to sudden changes in the environment. We investigated whole animal and muscle phenotypic changes occurring in black-capped chickadees (Poecile atricapillus) acclimated to cold (−5°C) and warm (20°C) temperatures in the first 3 h following a 15°C temperature drop (over 3 h). Before the temperature change, cold-acclimated birds were consuming 95% more food, were carrying twice as much body fat, and had 23% larger pectoralis muscle fiber diameters than individuals kept at 20°C. In the 3 h following the temperature drop, these same birds altered their pectoralis muscle ultrastructure by increasing the number of capillaries per fiber area and the number of nuclei per millimeter of fiber by 22%, consequently leading to a 22% decrease in myonuclear domain (amount of cytoplasm serviced per nucleus), whereas no such changes were observed in the warm-acclimated birds. To our knowledge, this is the first demonstration of such a rapid adjustment in muscle fiber ultrastructure in vertebrates. These results support the hypothesis that chickadees maintaining a cold phenotype are better prepared than warm-phenotype individuals to respond to a sudden decline in temperature, such as what may be experienced in their natural wintering environment. [ABSTRACT FROM AUTHOR]

Current life history theory suggests that during times of extreme energetic demand, animals may need to trade off physiological capacity of certain functions to maintain other more important biological functions. This might be especially true during reproduction, as parents try to balance their current reproductive demands with the maintenance of themselves and their offspring, while also considering their future fitness. Here, we experimentally manipulated the brood size of black-capped (Poecile atricapillus) and boreal (Poecile hudsonicus) chickadees, giving them ± 2 nestlings (increased/decreased group) or leaving their nest at the same size (without manipulation; natural comparator group). Parents did not adjust their field metabolic rate to reflect the new brood size; however, nest visitation rate was higher in the decreased group. We found little difference between groups in terms of parental body composition or inflammatory response, with the exception that parents in the decreased group had a higher mass. Nestlings in the increased brood group had lower masses over the course of growth, including right before fledging. The manipulation also appeared to have an impact on tarsus length, but was best explained by a combination of group, species, hatch order and interactions. Nestlings in the decreased group had lower circulating IgY antibodies , but the manipulation had a minimal impact on inflammatory response. We found that, when parents were challenged to work harder, instead of altering aspects of their own physiology, the parents pushed the costs onto their nestlings, resulting in lower quality offspring. [ABSTRACT FROM AUTHOR]

Food availability might sometimes be unpredictable for wild birds. To alleviate this possible food limitation, millions of households in North America provide food supplementation to bird populations. However, the ecoimmunological impacts of this supplementation on free-living birds are largely unclear. Therefore, we compared immune function and body composition of three groups of free-living black-capped chickadees (Poecile atricapillus) that were provided either constant food supplementation (“supplemented”), interrupted food supplementation (“interrupted”), or no food supplementation (“unsupplemented”). At capture, all three groups had similar body mass and fat scores. All three groups also had similar levels of circulating immunoglobulin Y antibodies and complement lysis ability, two measures of constitutive immune function. Supplemented and interrupted groups mounted a somewhat similar body mass and temperature response to injection with lipopolysaccharide; however, the supplemented group had a higher haptoglobin (acute-phase protein) response to lipopolysaccharide injection compared to the interrupted group. This study demonstrates that birds maintained similar levels of fat despite their level of food supplementation; however, sudden removal of supplemental food might elicit a short-term decline in aspects of immunity. Future studies should investigate how food supplementation might impact induced or adaptive aspects of immune function to add to our understanding of immunology in free-living animals. [ABSTRACT FROM AUTHOR]

Using red knots (Calidris canutus) as a model, we determined how changes in mass and metabolic activity of organs relate to temperature-induced variation in metabolic performance. In cold-acclimated birds, we expected large muscles and heart as well as improved oxidative capacity and lipid transport, and we predicted that this would explain variation in maximal thermogenic capacity (Msum). We also expected larger digestive and excretory organs in these same birds and predicted that this would explain most of the variation in basal metabolic rate (BMR). Knots kept at 5°C were 20% heavier and maintained 1.5 times more body fat than individuals kept in thermoneutral conditions (25°C). The birds in the cold also had a BMR up to 32% higher and a Msum 16% higher than birds at 25°C. Organs were larger in the cold, with muscles and heart being 9-20% heavier and digestive and excretory organs being 21-36% larger than at thermoneutrality. Rather than the predicted digestive and excretory organs, the cold-induced increase in BMR correlated with changes in mass of the heart, pectoralis, and carcass. Msum varied positively with the mass of the pectoralis, supracoracoideus, and heart, highlighting the importance of muscles and cardiac function in cold endurance. Cold-acclimated knots also expressed upregulated capacity for lipid transport across mitochondrial membranes [carnitine palmitoyl transferase (CPT)] in their pectoralis and leg muscles, higher lipid catabolism capacity in their pectoralis muscles [β-hydroxyacyl CoA-dehydrogenase (HOAD)], and elevated oxidative capacity in their liver and kidney (citrate synthase). These adjustments may have contributed to BMR through changes in metabolic intensity. Positive relationships among Msum, CPT, and HOAD in the heart also suggest indirect constraints on thermogenic capacity through limited cardiac capacity. [ABSTRACT FROM AUTHOR]

In winter, temperate resident birds are often faced with periodic low natural food availability. This reduction or unpredictability in resource availability might then have a negative impact on immune function, given that immune system support is highly resource dependent. We investigated the balance between energetic and immune management in captive black-capped chickadees (Poecile atricapilus) by manipulating the predictability of resources. The control group received food ad lib. every day, while the experimental group received a reduced amount of food on random days and food ad lib. on all other days. We measured two key metrics of energetic management (body and fat mass) as well as a suite of immune system components. Compared with control birds, experimental birds maintained significantly higher total body and fat mass, had lower acute phase protein concentrations, and had decreased body temperature and lost more body mass during the fever response following injection with lipopolysaccharides. Interestingly, birds in both groups had similar levels of complement lysis, delayed-type hypersensitivity response (phytohemagglutinin), and primary antibody production (keyhole limpet hemocyanin). This experiment demonstrates that black-capped chickadees strategically increase their fat mass in response to decreased food availability and that this might allow the birds to maintain most of the immune system unaltered, except some of the most costly immune components. [ABSTRACT FROM AUTHOR]

Small birds inhabiting regions with cold winter climates show seasonally flexible metabolic phenotypes, with the winter phenotype characterized by increments of summit metabolic rate (M) and cold tolerance. In the study reported here, we focused on variations in M as a metric of metabolic performance because it is positively correlated with cold tolerance in birds and positively related to overwinter survival in small mammals, although the latter has yet to be demonstrated in birds. Temperature appears to be a prominent driver of seasonal metabolic phenotypes in birds, as evidenced by the correlation between inter- and intra-seasonal variation in M and temperature variation, by recent temperature variables serving as better predictors of M variation than long-term climate variables, and by the induction of M variation by experimental cold exposure. In contrast, photoperiod and social status do not appear to be prominent drivers of metabolic flexibility in birds studied to date. Because skeletal muscle is the primary thermogenic tissue in birds, studies of the mechanistic underpinnings of metabolic flexibility have focused on skeletal muscles, particularly flight muscles. At the level of the skeletal muscle, two potential mechanisms exist for increasing thermogenic capacity, namely, muscle hypertrophy and elevated cellular metabolic intensity. Correlative studies suggest consistent winter increments in flight muscle size, with a potential regulatory role for the muscle growth inhibitor myostatin. Recent experimental studies in small birds, including modification of flight costs, cold acclimation, and exercise training, also suggest that muscle size is an important driver of metabolic flexibility in birds. Therefore, the focus of our study was on the seasonal regulation of muscle size and its contribution to metabolic flexibility. Future studies should address fitness consequences of M variation, the relative roles of muscle hypertrophy, and other factors (e.g., oxygen and substrate transport, cellular metabolic intensity) promoting M variation, as well as the molecular mechanisms underlying seasonal phenotypes. [ABSTRACT FROM AUTHOR]

Reversible phenotypic flexibility allows organisms to better match phenotypes to prevailing environmental conditions and may produce fitness benefits. Costs and constraints of phenotypic flexibility may limit the capacity for flexible responses but are not well understood nor documented. Costs could include expenses associated with maintaining the flexible system or with generating the flexible response. One potential cost of maintaining a flexible system is an energetic cost reflected in the basal metabolic rate (BMR), with elevated BMR in individuals with more flexible metabolic responses. We accessed data from thermal acclimation studies of birds where BMR and/or Msum (maximum cold-induced metabolic rate) were measured before and after acclimation, as a measure of metabolic flexibility, to test the hypothesis that flexibility in BMR (ΔBMR), Msum (ΔMsum), or metabolic scope (Msum − BMR; ΔScope) is positively correlated with BMR. When temperature treatments lasted at least three weeks, three of six species showed significant positive correlations between ΔBMR and BMR, one species showed a significant negative correlation, and two species showed no significant correlation. ΔMsum and BMR were not significantly correlated for any species and ΔScope and BMR were significantly positively correlated for only one species. These data suggest that support costs exist for maintaining high BMR flexibility for some bird species, but high flexibility in Msum or metabolic scope does not generally incur elevated maintenance costs. [ABSTRACT FROM AUTHOR]

Reproductive success of seabirds is tightly associated with availability of their prey for which the spatiotemporal distribution may be influenced by sea surface temperature (SST) fluctuations. The objective of this study was to investigate whether Northern gannets ( Morus bassanus ) from the largest colony in North America (Bonaventure Island, Quebec, Canada) were in negative nutritional state during the unprecedented low reproductive success year of 2012, and whether this was associated with changes in SST anomalies and diet. The incubation period of gannets in 2012 was characterized by a significant decline, from early to late incubation, in plasma triglyceride levels that was associated with an increase in plasma corticosterone levels. However, no changes in plasma glycerol and β-hydroxybutyrate levels were noted. SST anomalies recorded in this area (south of the Gulf of St. Lawrence) during the breeding period were consistently higher in 2012 compared to the previous year (a better reproductive success year). Based on signatures of stable carbon (δ 13 C) and nitrogen (δ 15 N) isotopes in gannet red blood cells and in whole fish homogenates of three major preys (mackerel, herring, and capelin), a minor dietary shift was noted between those years and incubation periods. In light of these findings, it is suggested that the extreme warm-water perturbation event that prevailed in the Gulf of St. Lawrence during summer 2012 was associated with a rapid deterioration of nutritional condition of Bonaventure Island gannets during the incubation. These suboptimal physiological changes likely contributed to the dramatic decline in reproductive success reported in this colony. [ABSTRACT FROM AUTHOR]

Reaction norms reflect an organisms' capacity to adjust its phenotype to the environment and allows for identifying trait values associated with physiological limits. However, reaction norms of physiological parameters are mostly unknown for endotherms living in natural conditions. Black-capped chickadees (Poecile atricapillus) increase their metabolic performance during winter acclimatization and are thus good model to measure reaction norms in the wild. We repeatedly measured basal (BMR) and summit (Msum) metabolism in chickadees to characterize, for the first time in a free-living endotherm, reaction norms of these parameters across the natural range of weather variation. BMR varied between individuals and was weakly and negatively related to minimal temperature. Msum varied with minimal temperature following a Z-shape curve, increasing linearly between 24°C and −10°C, and changed with absolute humidity following a U-shape relationship. These results suggest that thermal exchanges with the environment have minimal effects on maintenance costs, which may be individual-dependent, while thermogenic capacity is responding to body heat loss. Our results suggest also that BMR and Msum respond to different and likely independent constraints. [ABSTRACT FROM AUTHOR]

In small resident bird species living at northern latitudes, winter cold acclimatization is associated with an increase in pectoral muscle size and haematocrit level, and this is thought to drive the seasonal increase in summit metabolic rate (...Msum, a measure of maximal shivering thermogenic capacity). However, evidence suggesting that pectoral muscle size influences ...Msum is correlational and the link between haematrocrit level and ...Msum remains to be demonstrated. We experimentally tested the relationship between pectoral muscle size and ...Msum by manipulating muscle size using a feather clipping protocol in free-living wintering black-capped chickadees (Poecile atricapillus). This also allowed us to investigate the link between haematocrit and thermogenic capacity. After a first series of measures on all birds, we cut half of the flight feathers of experimental individuals (N=14) and compared their fat and pectoral muscle scores, ...Msum and haematocrit level at recapture with their previous measures and with those of control birds (N=17) that were captured and recaptured at comparable times. Results showed that: (1) experimental birds developed larger pectoral muscles than control individuals and (2) mass-independent ...Msum was up to 16% higher in birds expressing large pectoral muscles. ...Msum was also positively correlated with haematocrit, which was not affected by the experimental manipulation. These findings demonstrate that, for a given body mass, large pectoral muscles are associated with a higher ...Msum in black-capped chickadees and that oxygen carrying capacity likely supports thermogenesis in this species. [ABSTRACT FROM AUTHOR]

In small resident bird species living at northern latitudes, winter cold acclimatization is associated with an increase in pectoral muscle size and haematocrit level, and this is thought to drive the seasonal increase in summit metabolic rate (Ṁsum, a measure of maximal shivering thermogenic capacity). However, evidence suggesting that pectoral muscle size influences Ṁsum is correlational and the link between haematrocrit level and Ṁsum remains to be demonstrated. We experimentally tested the relationship between pectoral muscle size and Ṁsum by manipulating muscle size using a feather clipping protocol in free-living wintering black-capped chickadees (Poecile atricapillus). This also allowed us to investigate the link between haematocrit and thermogenic capacity. After a first series of measures on all birds, we cut half of the flight feathers of experimental individuals (N=14) and compared their fat and pectoral muscle scores, Ṁsum and haematocrit level at recapture with their previous measures and with those of control birds (N=17) that were captured and recaptured at comparable times. Results showed that: (1) experimental birds developed larger pectoral muscles than control individuals and (2) mass-independent Ṁsum was up to 16% higher in birds expressing large pectoral muscles. Ṁsum was also positively correlated with haematocrit, which was not affected by the experimental manipulation. These findings demonstrate that, for a given body mass, large pectoral muscles are associated with a higher Ṁsum in black-capped chickadees and that oxygen carrying capacity likely supports thermogenesis in this species. [ABSTRACT FROM AUTHOR]

Summary 1. Seasonal carry-over effects may be important structuring components of avian life-history cycles. However, little is known on physiological transitions between stages and on phenotypic compromises that may be made at such time to improve fitness. 2. We studied postmigratory body remodelling in red knots ( Calidris canutus islandica) arriving on the Arctic breeding grounds. Our objectives were to determine the effects of body reconstruction and preparation for breeding on maintenance energy costs and to determine whether weather conditions can force compromises between functions required for postmigration recovery of body composition, thermoregulation and breeding. 3. During two consecutive springs at the northernmost land on Earth (Alert, Ellesmere Island, Canada, 82°30′N), we monitored changes in knots post-arrival body stores. Using ultrasonography, we also tracked changes in gizzard size, an indicator of gut size, and pectoral muscle thickness, not only an endogenous protein source but also a thermogenic organ. We measured basal metabolic rate (BMR) throughout reconstruction and compared it with BMR of nonbreeding red knots wintering in the Dutch Wadden Sea. 4. Arriving knots faced temperatures up to 13 °C lower than during midwinter. Birds arrived with large body stores and pectoral muscles, which declined in size while they grew their gizzard and prepared for breeding. BMR at arrival was indistinguishable from winter BMR and increased linearly throughout reconstruction. BMR increased up to 69% faster in females than males, likely due to the development of their reproductive organs. 5. Birds had lower body stores but larger muscles in the colder year, and muscle loss was correlated with the warming of spring temperatures. Therefore, muscles would not only serve as a nutrient source, but their thermogenic function could also provide the flexibility to cope with high thermostatic costs in the spring. However, retaining muscles for shivering may limit protein recirculation and delay the onset of breeding. 6. Postmigratory recovery therefore involves significant energy costs and arriving birds likely have to make physiological compromises, depending on spring conditions, which may impact on fitness. Although this period is clearly critical in the life cycle of red knots, it is one of the least understood life-history stages in Arctic-breeding shorebirds. [ABSTRACT FROM AUTHOR]

Phenotypic flexibility in shorebirds has been studied mainly in the context of adjustments to migration and to quality of food; little is known on how birds adjust their phenotype to harsh winter conditions. We showed earlier that red knot (Calidris canutus islandica) can acclimate to cold by elevating body mass. This goes together with larger pectoral muscles, i.e., greater shivering machinery, and thus, better thermogenic capacity. Here, we present results of a yearlong experiment with indoor captive knots to determine whether this strategy is part of their natural seasonal phenotypic cycle. We maintained birds under three thermal regimes: constant cold (5°C), constant thermoneutrality (25°C) and natural seasonal variation between these extremes (9–22°C). Each month we measured variables related to the birds’ endurance to cold and physiological maintenance [body mass, thickness of pectoral muscles, summit metabolic rate (Msum), food intake, gizzard size, basal metabolic rate (BMR)]. Birds from all treatments expressed synchronized and comparable variation in body mass in spite of thermal treatments, with a 17–18% increase between the warmest and coldest months of the year; which appeared regulated by an endogenous driver. In addition, birds living in the cold exhibited a 10% higher average body mass than did those maintained at thermoneutrality. Thickness of the pectoral muscle tracked changes in body mass in all treatments and likely contributed to greater capacity for shivering in heavier birds. Consequently, Msum was 13% higher in cold-acclimated birds compared to those experiencing no thermoregulation costs. However, our data also suggest that part of maximal heat production comes from nonshivering processes. Birds facing cold conditions ate up to 25% more food than did birds under thermoneutral conditions, yet did not develop larger gizzards. Seasonal variation in BMR followed changes in body mass, probably reflecting changes in mass of metabolically active tissues. Just as cold-exposed birds, red knots in the variable treatment increased body mass in winter, thereby improving cold endurance. During summer, however, they maintained a lower body mass and thermogenic capacity compared to cold-exposed birds, similar to individuals kept at thermoneutrality. We conclude that red knots acclimate to seasonal variations in ambient temperature by modulating body mass, combining a preprogrammed increase in mass during winter with a capacity for fine-tuning body mass and thermogenic capacity to temperature variations. [ABSTRACT FROM AUTHOR]

Interest in phenotypic flexibility has increased dramatically over the last decade, but flexibility during reproduction has received relatively little attention from avian scientists, despite its possible impact on fitness. Because most avian species maintain atrophied reproductive organs when not active, reproduction in birds requires major tissue remodeling in preparation for breeding. Females undergo rapid (days) recrudescence and regression of their reproductive organs at each breeding attempt, while males grow their organs ahead of time at a much slower rate (weeks) and may maintain them at maximal size throughout the breeding season. Reproduction is associated with significant metabolic costs. Egg production leads to a 22%-27% increase in resting metabolic rate (RMR) over non-reproductive values. This is partly due to the activity of the oviduct, an organ that may allow females to adjust reproductive investment by modulating egg size and quality. In males, gonadal recrudescence may lead to a 30% increase in RMR, but the data are inconsistent and general conclusions regarding energetic costs of reproduction in males will require more research. Recent studies on captive female zebra finches describe the impacts of these costs on daily energy budgets and highlight the strategies used by birds to maintain their investment in reproduction when energy is limited. Whenever possible, birds use behavioral flexibility as a first means of saving energy. Decreasing locomotor activity saves energy during challenges such as egg production or exposure to cold temperatures and is an efficient way to buffer variation in individual daily energy budgets. However, when behavioral flexibility is not possible, birds must rely on flexibility at the physiological level to meet energy demands. In zebra finches breeding in the cold, this results in a reduced pace of laying, likely due to down-regulation of both reproductive and non-reproductive function, allowing females to defend minimal egg size and maintain reproductive success. More research involving a range of species in captive and free-living conditions is needed to determine how phenotypic flexibility during tissue remodeling and early reproductive investment translates to natural conditions and affects fitness. [ABSTRACT FROM AUTHOR]

Pre-flight fuelling rates in free-living red knots Calidris canutus, a specialized long-distance migrating shorebird species, are positively correlated with latitude and negatively with temperature. The single published hypothesis to explain these relationships is the heat load hypothesis that states that in warm climates red knots may overheat during fuelling. To limit endogenous heat production (measurable as basal metabolic rate BMR), birds would minimize the growth of digestive organs at a time they need. This hypothesis makes the implicit assumption that BMR is mainly driven by digestive organ size variation during pre-flight fuelling. To test the validity of this assumption, we fed captive knots with trout pellet food, a diet previously shown to quickly lead to atrophied digestive organs, during a fuelling episode. Birds were exposed to two thermal treatments (6 and 24°C) previously shown to generate different fuelling rates in knots. We made two predictions. First, easily digested trout pellet food rather than hard-shelled prey removes the heat contribution of the gut and would therefore eliminate an ambient temperature effect on fuelling rate. Second, if digestive organs were the main contributors to variations in BMR but did not change in size during fuelling, we would expect no or little change in BMR in birds fed ad libitum with trout pellets. We show that cold-acclimated birds maintained higher body mass and food intake (8 and 51%) than warm-acclimated birds. Air temperature had no effect on fuelling rate, timing of fuelling, timing of peak body mass or BMR. During fuelling, average body mass increased by 32% while average BMR increased by 15% at peak of mass and 26% by the end of the experiment. Our results show that the small digestive organs characteristic of a trout pellet diet did not prevent BMR from increasing during premigratory fuelling. Our results are not consistent with the heat load hypothesis as currently formulated. [ABSTRACT FROM AUTHOR]

Phenotypic flexibility is a phenomenon where physiological functions in animals are reversibly adjusted in response to ecological constraints. Research usually focuses on effects of single constraints, but under natural conditions animals face a multitude of restrictions acting simultaneously, and potentially generating conflicting demands on the phenotype. We investigated the conflicting demands of low temperatures and a low quality diet on the phenotype of a shorebird, the red knot Calidris canutus. We tested the effects of switching diet from a high quality trout food to low quality hard-shelled bivalves in captive birds acclimated to temperatures reflecting natural winter conditions. Feeding on bivalves generated a digestive constraint forcing the birds to increase the height and width of their gizzard by 66% and 71%, respectively, over 30 days. The change in gizzard size was associated with an initial 15% loss of body mass and a reduction in size of the pectoral muscles by 11%. Because pectoral muscle size determines summit metabolic rate (Msum, an indicator of cold endurance), measured Msum declined by 9%. Therefore, although the birds were acclimated to cold, gizzard growth led to a loss of cold endurance. We propose that cold-acclimated knots facing a digestive constraint made a phenotypic compromise by giving-up cold hardiness for digestive capacity. Field studies suggest that phenotypic compromises occur in free-living red knots as well and help improve survival. [ABSTRACT FROM AUTHOR]

Previous experiments showed reduction of basal metabolic rate (BMR) in birds facing energetic challenges. We alternately exposed two groups of red knots (Calidris canutus) to either 6 h or 22 h of food availability for periods of 22 d. Six h of access to food led to a 6%-10% loss of body mass over the first 8 d, with nearly all of the birds' daily energy expenditures supported by body nutrient stores during the first 2 d. Birds responded by increasing feeding behavior and food intake, but the response was slow. There were no gains of mass before day 15, which suggests a digestive bottleneck and a period of physiological adjustment. Food-restricted birds exhibited decreases in pectoral-muscle thickness and BMR in association with a loss of body mass. Although a decrease in BMR saves energy, savings represented only 2%-7% of the daily energy spent in excess of that acquired during the deficit period. Red knots did not downregulate mass-independent BMR. On the bases of recent independent findings and the pattern of mass gain observed when food access was switched from 6 h to 22 h, we suggest that these birds routinely maintain nutrient stores as a buffer against periods of energy shortages, thereby precluding the need for downregulation of mass-independent BMR. [ABSTRACT FROM AUTHOR]

Basal metabolic rate (BMR) in animals is interpreted as reflecting the size and metabolic intensity of energy-consuming tissues. However, studies investigating relationships between the mass of specific organs and interindividual variation in BMR have produced inconsistent patterns with regard to which organs have the largest impact on BMR variation. Because of the known flexibility in organ mass and metabolic intensity within individual organs, relationships between BMR and body-composition variables are bound to be context specific. Altricial nestlings are excellent models to illustrate this phenomenon because of the extreme variation in body composition occurring during growth. Using European starlings at three age classes, we studied changes in body composition together with its effect on variation in resting metabolic rate (RMR) in order to highlight the context-specific nature of these relationships. Our data suggest a transition in metabolic costs during growth in starling nestlings. During the linear phase of growth, energy is mainly consumed by tissue-synthesis processes, with fast-growing organs having a large influence on RMR. In the plateau phase of growth, the energy expenditure is transferred to functional costs, with high-intensity organs having a predominant effect on RMR variation. Our data illustrates the context-specific nature of organ mass-metabolic rate correlations, which complicates inter- and intraspecific comparisons of BMR. In the future, such comparisons must be done while taking the physiological state of the study animal into account. [ABSTRACT FROM AUTHOR]

It is axiomatic that whole-animal metabolism, measured for example as daily energy expenditure (DEE), plays a central role in determining reproductive success and survival (fitness) in all organisms. Nevertheless, strong evidence for consistent systematic relationships between DEE and either individual traits (age, sex, body size), environmental factors (e.g. food availability, temperature) or 'fitness' traits (e.g. number of offspring, survival) remains far from compelling in birds and mammals. Recently, we suggested that female birds might utilise complex, individually variable energy management strategies to meet the metabolic demands of reproduction, generating a wide spectrum of effects on reproductive DEE, from overcompensation (net decrease in DEE) to additive effects (net increase in DEE). Here we show that this individually variable adjustment or 'plasticity' in energy expenditure associated with egg production is repeatable among individuals between successive breeding attempts in female zebra finches (Taeniopygia guttata). Our study highlights the importance (a) of measuring 'plasticity' or change associated with transitions of physiological state (e.g. non-breeding to breeding) based on multiple measurements of the same individual, and (b) of extending consideration of how selection might drive the evolution of phenotypic plasticity per se to include physiological and metabolic traits. [ABSTRACT FROM AUTHOR]

Even though molt involves both endocrine and energetic changes in bird bodies, this study is among the first to combine assessments of energy costs together with thyroid hormone variations in molting birds. Individual shorebirds (red knots Calidris canutus islandica) were measured while in full summer and winter plumage as well as during peak of molt. Molt was associated with a 9.8% increase in average mass-independent basal metabolic rate (BMR) above nonmolting levels. Individual plasma levels of thyroxine (T4) were correlated with individual rate of body feather renewal, confirming that T4 is related to body molt but also showing that it is potentially regulating its rate. Across seasons, mass-independent average heat loss measured as conductance gradually declined with conductance during molt falling between measured values for summer and winter. During the molting period, however, body molting rate was positively correlated with thermal conductance, indicating that for a given ambient temperature below thermoneutrality, the fastest molting birds were losing more body heat. Across seasons, triiodothyronine (T3), a hormone typically upregulated in response to a cold stimulus, was correlated with individual thermal conductance and BMR. We suggest that the increased heat loss of fast-molting birds leads to a cold-acclimatization response that may be partly responsible for the elevated BMR measured during molt. This could be mediated through a stimulatory effect of T3 on BMR in response to increased heat loss. Our interpretation is supported by a positive relationship between the individual changes in conductance and the change in BMR from summer to the molting period. [ABSTRACT FROM AUTHOR]

In the calidrine sandpiper red knot (Calidris canutus), the weeks preceding takeoff for long-distance migration are characterized by a rapid increase in body mass, largely made up of fat but also including a significant proportion of lean tissue. Before takeoff, the pectoral muscles are known to hypertrophy in preparation for endurance flight without any specific training. Because birds facing cold environments counterbalance heat loss through shivering thermogenesis, and since pectoral muscles represent a large proportion of avian body mass, we asked the question whether muscle hypertrophy in preparation for long-distance endurance flight would induce improvements in thermogenic capacity. We acclimated red knots to different controlled thermal environments: 26°C, 5°C, and variable conditions tracking outdoor temperatures. We then studied within-individual variations in body mass, pectoral muscle size (measured by ultrasound), and metabolic parameters [basal metabolic rate (BMR) and summit metabolic rate (Msum)] throughout a 3-mo period enclosing the migratory gain and loss of mass. The gain in body mass during the fattening period was associated with increases in pectoral muscle thickness and thermogenic capacity independent of thermal acclimation. Regardless of their thermal treatment, birds showing the largest increases in body mass also exhibited the largest increases in Msum. We conclude that migratory fattening is accompanied by thermoregulatory side effects. The gain of body mass and muscle hypertrophy improve thermogenic capacity independent of thermal acclimation in this species. Whether this represents an ecological advantage depends on the ambient temperature at the time of fattening. [ABSTRACT FROM AUTHOR]

Seasonal acclimatization and experimental acclimation to cold in birds typically results from increased shivering endurance and elevated thermogenic capacity leading to improved resistance to cold. A wide array of physiological adjustments, ranging from biochemical transformations to organ mass variations, are involved in this process. Several studies have shown that improved cold endurance is accompanied by increases in summit metabolic rate (Msum), a measure of maximal heat production and an indicator of the level of sustainable thermogenic capacity. However, improved endurance to cold can also be achieved without significant changes in Msum. The same is true for basal metabolic rate (BMR), which is known to increase in association with cold acclimatization or acclimation in some species but not in others. We investigated cold acclimation in a migrant shorebird known for extreme physiological flexibility, the red knot (Calidris canutus, the northerly wintering subspecies islandica). We measured BMR and Msum over two months in birds caught in the wild and transferred to experimentally controlled conditions representative of aspects of their seasonal thermal environment (two groups at constant 25°C, one group at constant 4°C and two groups experiencing variable outdoor temperatures). Birds maintained in both cold and variable ambient temperatures showed a 14-15% higher body mass, 33-45% higher food intake, and 26% and 13% elevations in BMR and Msum, respectively, compared with birds kept at thermoneutrality. These results, together with data on alimentary tract size and pectoral muscle thickness measured by ultrasonography, suggest that red knots acclimate to cold primarily through modulation of (lean) body mass components. Heavier individuals have larger muscles, which allow higher maximal heat production and better thermal compensation. Cold acclimation effects on BMR are most probably due to changes in the size of visceral organs, although not the alimentary tract in this specific case. The liver, known for its thermogenic capacity, is a probable candidate. Overall, our results indicate that relatively small changes in body mass and muscle size allow enough reserve capacity in terms of heat production to cope with typical wintering ambient temperature variations as measured on the red knot's wintering grounds. [ABSTRACT FROM AUTHOR]

The metabolic cost of egg production in birds (passerines) has been measured as a 16-27% increase in basal or resting metabolic rate (BMR and RMR, respectively) when comparing non-breeding values with those in egg-producing individuals. However, available data to date have been obtained in free-living birds and may thus be confounded by the effect of variable ecological conditions on non-reproductive physiological machinery (organ mass or metabolic activity) which might contribute to measured variation in RMR. Here, we show that in captive, controlled conditions, the process of egg formation induces a 22% increase in RMR in female zebra finches Taeniopygia guttata. Among individuals, variation in laying RMR is independent of egg mass, clutch size or total clutch mass. Importantly, we show that individual variation in both non-breeding and laying RMR is repeatable over periods of at least 8-10 months, i.e. individual variation in RMR remained constant over time for any given physiological state. This suggests that the metabolic cost of egg formation should respond to selection. However, we also show that in males, but not females, repeatability of RMR declines over time even when birds are kept in constant controlled conditions. [ABSTRACT FROM AUTHOR]

Little is known about the energy costs of egg production in birds. We showed in previous papers that, during egg production, European starlings (Sturnus vulgaris) undergo a 22% increase in resting metabolic rate (RMR) and that the maintenance and activity costs of the oviduct are responsible for 18% of the variation in elevated laying RMR. Therefore, other energy-consuming physiological mechanisms must be responsible for the remaining unexplained variation in elevated laying RMR. Yolk precursor [vitellogenin (VTG) and very-low-density lipoprotein (VLDL)] production is likely to be costly because it signifies a marked increase in the biosynthetic activity of the liver. We documented the pattern of yolk precursor production in response to daily injections of 17β-estradiol (E[SUB2]) in zebra finches (Taeniopygia guttata). Based on this pattern we carried out an experiment in order to evaluate the metabolic costs of producing VTG and VLDL. Our E[SUB2] treatment resulted in a significant increase in plasma VTG and VLDL levels within the natural breeding range for the species. Although RMR was measured during the period of active hepatic yolk precursor production, it did not differ significantly within individuals in response to the treatment or when comparing E[SUB2]-treated birds with sham-injected birds. This could mean that yolk precursor production represents low energy investment. However, we discuss these results in light of possible adjustments between organs that could result in energy compensation. [ABSTRACT FROM AUTHOR]

Body composition in vertebrates is known to show phenotypic plasticity, and changes in organ masses are usually rapid and reversible. One of the most rapid and reversible changes is the transformation of the female avian reproductive organs before breeding. This provides an excellent system to investigate the effects of plasticity in organ size on basal metabolic rate (BMR) through relationships between organ masses and BMR. We compared body composition of female European starlings (Sturnus vulgaris) during various reproductive stages over 3 yr and investigated the pattern of changes in reproductive and nonreproductive organ mass during follicular development and ovulation. Furthermore, we analyzed the relationship between organ mass and resting metabolic rate (RMR) in nonbreeding, laying, and chick-rearing females. Our analysis revealed marked variation in organ masses between breeding stages but no consistent pattern among years except for kidney and pectoralis muscle. Furthermore, changes in nonreproductive organs did not parallel the cycle of growth and regression of the reproductive organs. The oviduct gained 62% of its 22-fold increase in mass in only 3 d, and oviduct regression was just as rapid and began even before the final egg of the clutch was laid, with 42% of the oviduct mass lost before laying of the final egg. In laying females, 18% of variation in mass-corrected RMR was explained by the mass of the oviduct (r⊃2 = 0.18, n = 80, P < 0.0005), while pectoralis muscle mass in nonbreeding individuals and liver and gizzard mass in chick-rearing females were the only organs significantly related to RMR (r⊃2 = 0.31-0.44). We suggest that the nonreproductive organs are affected more by changes in local ecological conditions than the reproductive state itself and that the activity and maintenance cost of the oviduct is high enough that selection has led to a very tight size-function relationship for this organ. [ABSTRACT FROM AUTHOR]

Animal migration contributes largely to the seasonal dynamics of High Arctic ecosystems, linking distant habitats and impacting ecosystem structure and function. In polar deserts, Arctic hares are abundant herbivores and important components of food webs. Their annual migrations have long been suspected, but never confirmed. We tracked 25 individuals with Argos satellite telemetry to investigate the existence of migration in a population living at Alert (Ellesmere Island, Nunavut, Canada). During fall, 21 hares undertook directional, long-distance movements in a southwestern direction towards Lake Hazen. Daily movement rates averaged 1.3 ± 0.5 km, 4.3 ± 1.6 km, and 1.7 ± 0.9 km before, during, and after relocation, respectively. Straight-line and minimum cumulative distances traveled averaged 98 ± 18 km (range: 72–148 km) and 198 ± 62 km (range: 113–388 km), respectively. This is the first report of large-scale seasonal movements in Arctic hares and, surprisingly, in any lagomorph species. These movements may be part of an annual migratory pattern. Our results redefine our understanding of the spatial ecology of Arctic hares, demonstrate unsuspected mobility capacities in lagomorphs, and open new perspectives regarding the ecological dynamics of the northern polar deserts. [ABSTRACT FROM AUTHOR]

Several species of passerines leave their nest with unfinished feather growth, resulting in lower feather insulation and increased thermoregulatory demands compared to adults. However, feather insulation is essential for avian species breeding at northern latitudes, where cold conditions or even snowstorms can occur during the breeding season. In altricial arctic species, increased heat loss caused by poor feather insulation during growth could be counter-adaptative as it creates additional energy demands for thermoregulation. Using flow-through respirometry, we compared resting metabolic rate at thermoneutrality (RMRt), summit metabolic rate (Msum) and heat loss (conductance) in adult and juvenile snow buntings on their summer and winter grounds. In summer, when buntings are in the Arctic, juveniles had a 12% higher RMRt, likely due to unfinished growth, and lost 14% more heat to the environment than adults. This pattern may result from juveniles fledging early to avoid predation at the cost of lower feather insulation. Surprisingly, an opposite pattern was observed at lower latitudes on their wintering grounds. Although they showed no difference in RMRt and Msum, adults were losing 12% more heat than juveniles. We suggest that this difference is due to poorer insulative property of plumage in adults stemming from energetic and time constraints encountered during their post-breeding molt. High plumage insulation in first-winter juvenile buntings could be adaptive to reduce thermoregulatory demands and maximize survival in the first winter of life, while adults could use behavioral strategies to compensate for their greater rate of heat loss. [ABSTRACT FROM AUTHOR]

It is generally assumed that small birds improve their shivering heat production capacity by developing the size of their pectoralis muscles. However, some studies have reported an enhancement of thermogenic capacity in the absence of muscle mass variation between seasons or thermal treatments. We tested the hypothesis that an increase in muscle mass is not a prerequisite for improving avian thermogenic capacity. We measured basal (BMR) and summit (Msum) metabolic rates of black capped chickadees (Poecile atricapillus) acclimated to thermoneutral (27 °C) and cold (−10 °C) temperatures and obtained body composition data from dissections. Cold acclimated birds consumed 44% more food, and had 5% and 20% higher BMR and Msum, respectively, compared to individuals kept at thermoneutrality. However, lean dry pectoralis and total muscle mass did not differ between treatments, confirming that the improvement of thermogenic capacity did not require an increase in skeletal muscle mass. Nevertheless, within temperature treatments, Msum was positively correlated with the mass of all measured muscles, including the pectoralis. Therefore, for a given acclimation temperature individuals with large muscles do benefit from muscle size in term of heat production but improving thermogenic capacity during cold acclimation likely requires an upregulation of cell functions. [ABSTRACT FROM AUTHOR]

The use of remote sensing for developing land cover maps in the Arctic has grown considerably in the last two decades, especially for monitoring the effects of climate change. The main challenge is to link information extracted from satellite imagery to ground covers due to the fine-scale spatial heterogeneity of Arctic ecosystems. There is currently no commonly accepted methodological scheme for high-latitude land cover mapping, but the use of remote sensing in Arctic ecosystem mapping would benefit from a coordinated sharing of lessons learned and best practices. Here, we aimed to produce a highly accurate land cover map of the surroundings of the Canadian Forces Station Alert, a polar desert on the northeastern tip of Ellesmere Island (Nunavut, Canada) by testing different predictors and classifiers. To account for the effect of the bare soil background and water limitations that are omnipresent at these latitudes, we included as predictors soil-adjusted vegetation indices and several hydrological predictors related to waterbodies and snowbanks. We compared the results obtained from an ensemble classifier based on a majority voting algorithm to eight commonly used classifiers. The distance to the nearest snowbank and soil-adjusted indices were the top predictors allowing the discrimination of land cover classes in our study area. The overall accuracy of the classifiers ranged between 75 and 88%, with the ensemble classifier also yielding a high accuracy (85%) and producing less bias than the individual classifiers. Some challenges remained, such as shadows created by boulders and snow covered by soil material. We provide recommendations for further improving classification methodology in the High Arctic, which is important for the monitoring of Arctic ecosystems exposed to ongoing polar amplification. [ABSTRACT FROM AUTHOR]

Adaptive explanations for both high and low body mass-independent basal metabolic rate (BMR) in endotherms are pervasive in evolutionary physiology, but arguments implying a direct adaptive benefit of high BMR are troublesome from an energetic standpoint. Here, we argue that conclusions about the adaptive benefit of BMR need to be interpreted, first and foremost, in terms of energetics, with particular attention to physiological traits on which natural selection is directly acting. We further argue from an energetic perspective that selection should always act to reduce BMR (i.e., maintenance costs) to the lowest level possible under prevailing environmental or ecological demands, so that high BMR per se is not directly adaptive. We emphasize the argument that high BMR arises as a correlated response to direct selection on other physiological traits associated with high ecological or environmental costs, such as daily energy expenditure (DEE) or capacities for activity or thermogenesis. High BMR thus represents elevated maintenance costs required to support energetically demanding lifestyles, including living in harsh environments. BMR is generally low under conditions of relaxed selection on energy demands for high metabolic capacities (e.g., thermoregulation, activity) or conditions promoting energy conservation. Under these conditions, we argue that selection can act directly to reduce BMR. We contend that, as a general rule, BMR should always be as low as environmental or ecological conditions permit, allowing energy to be allocated for other functions. Studies addressing relative reaction norms and response times to fluctuating environmental or ecological demands for BMR, DEE, and metabolic capacities and the fitness consequences of variation in BMR and other metabolic traits are needed to better delineate organismal metabolic responses to environmental or ecological selective forces. [ABSTRACT FROM AUTHOR]

Small resident bird species living at northern latitudes increase their metabolism in winter, and this is widely assumed to improve their chances of survival. However, the relationship between winter metabolic performance and survival has yet to be demonstrated. Using capture-mark-recapture,we followed a population of free-living black-capped chickadees (Poecile atricapillus) over 3 yr and evaluated their survival probability within and among winters. We also measured the size-independent body mass (Ms), hematocrit (Hct), basal metabolic rate (BMR), and maximal thermogenic capacity (Msum) and investigated how these parameters influenced survival within and among winters. Results showed that survival probability was high and constant both within (0.92) and among (0.96) winters. They also showed that while Ms, Hct, and BMR had no significant influence, survival was positively related to Msum—following a sigmoid relationship— within but not among winter. Birds expressing an Msum below 1.26W(i.e., similar to summer levels) had a !50% chance of survival, while birds with anMsumabove 1.35Whad at least a 90% chance of surviving through the winter. Our data therefore suggest that black-capped chickadees that are either too slow or unable to adjust their phenotype from summer to winter have little chances of survival and thus that seasonal upregulation of metabolic performance is highly beneficial. This study is the first to document in an avian system the relationship between thermogenic capacity and winter survival, a proxy of fitness. [ABSTRACT FROM AUTHOR]

Avian basal metabolic rate (BMR) and summit metabolic rate (Msum) vary in parallel during cold acclimation and acclimatization, which implies a functional link between these variables. However, evidence suggests that these parameters may reflect different physiological systems acting independently. We tested this hypothesis in white-throated sparrows (Zonotrichia albicollis) acclimated to two temperatures (287 and 287C) and two diets (0% and 30% cellulose). We expected to find an uncoupling of Msum and BMR whereMsum, a measure of maximal shivering heat production, would reflect muscle and heart mass variation and would respond only to temperature, while BMR would reflect changes indigestive and excretory organs in response to daily food intake, responding to both temperature and diet. We found that the gizzard, liver, kidneys, and intestines responded to treatments through a positive relationship with food intake. BMR was 15% higher in cold-acclimated birds and, as expected, varied with food intake and the mass of digestive and excretory organs. In contrast, although Msum was 19% higher in cold-acclimated birds, only heartmassresponded totemperature (118%in the cold). Pectoral muscles did not change in mass with temperature but were 8.2% lighter on the cellulose diet. Nevertheless, Msum varied positively with the mass of heart and skeletal muscles but only in coldacclimated birds. Our results therefore suggest that an upregulation of muscle metabolic intensity is required for cold acclimation. This study increases support for the hypothesis that BMRandMsum reflect different physiological systems responding in parallel to constraints associated with cold environments. [ABSTRACT FROM AUTHOR]

In birds, acclimation and acclimatization to temperature are associated with changes in basal (BMR), summit (M) and maximal (MMR) metabolic rates but little is known about the rate at which species adjust their phenotype to short-term temperature variations. Our aims were (1) to determine the pattern of metabolic adjustments following a rapid temperature change, (2) to determine whether performance varies at similar rates during exposure to warm or cold environments, and (3) to determine if BMR, M and MMR change at comparable rates during thermal acclimation. We measured these parameters in white-throated sparrows ( Zonotrichia albicollis), black-capped chickadees ( Poecile atricapillus), and snow buntings ( Plectrophenax nivalis) after acclimation to 10 °C (day 0) and on the 4th and 8th days of acclimation to either −5 or 28 °C. Birds changed their metabolic phenotype within 8 days with patterns differing among species. Sparrows expressed the expected metabolic increases in the cold and decreases at thermoneutrality while performance in chickadees and buntings was not influenced by temperature but changed over time with inverse patterns. Our results suggest that BMR varies at comparable rates in warm and cold environments but changes faster than M and MMR, likely due to limitations in the rate of change in organ size and function. They also suggest that maximal metabolic capacity is lost faster in a warm environment than it is gained in a cold environment. With the expected increase in temperature stochasticity at northern latitudes, a loss of thermogenic capacity during warm winter days could, therefore, be detrimental if birds are slow to readjust their phenotype with the return of cold days. [ABSTRACT FROM AUTHOR]

Rising global temperatures are expected to increase reproductive costs for wildlife as greater thermoregulatory demands interfere with reproductive activities. However, predicting the temperatures at which reproductive performance is negatively impacted remains a significant hurdle. Using a thermoregulatory polygon approach, we derived a reproductive threshold temperature for an Arctic songbird—the snow bunting (Plectrophenax nivalis). We defined this threshold as the temperature at which individuals must reduce activity to suboptimal levels (i.e. less than four-time basal metabolic rate) to sustain nestling provisioning and avoid overheating. We then compared this threshold to operative temperatures recorded at high (82° N) and low (64° N) Arctic sites to estimate how heat constraints translate into site-specific impacts on sustained activity level. We predict buntings would become behaviourally constrained at operative temperatures above 11.7°C, whereupon they must reduce provisioning rates to avoid overheating. Low-Arctic sites had larger fluctuations in solar radiation, consistently producing daily periods when operative temperatures exceeded 11.7°C. However, high-latitude birds faced entire, consecutive days when parents would be unable to sustain required provisioning rates. These data indicate that Arctic warming is probably already disrupting the breeding performance of cold-specialist birds and suggests counterintuitive and severe negative impacts of warming at higher latitude breeding locations. [ABSTRACT FROM AUTHOR]

Small avian species wintering at northern latitudes typically show increases in basal metabolic rate (BMR) and maximal thermogenic capacity (Msum). Those are widely assumed to reflect changes in body composition, with enlargement of digestive and excretory organs resulting in elevated winter BMR and larger body muscles driving the increase in Msum. Using free-living black-capped chickadees (Poecile atricapillus) as our model species, we investigated seasonal changes in body composition and tested for relationships between mass variations of body organs and variability of both BMR and Msum. Our results confirmed the expected winter increase in mass of body muscles and cardiopulmonary organs (heart + lungs) and showed that 64% of the observed Msum variations throughout the year were explained by changes in these organs. In contrast, we found little support for an effect of the digestive organs (gizzard + intestines) on BMR seasonal changes. Instead, this variable was mainly influenced by variations in mass of body muscles and excretory organs (liver + kidney), explaining up to 35% of its variability. [ABSTRACT FROM AUTHOR]

Keywords: Arctic; Lepus arcticus; migration; movement ecology; satellite telemetry EN Arctic Lepus arcticus migration movement ecology satellite telemetry 1 5 5 03/02/22 20220301 NES 220301 Long-distance movements undertaken by animals are among the most fascinating biological phenomena. " Demography and Ecology of the Arctic Hare (Lepus arcticus) in Southwestern Newfoundland." Describing the exceptional fall movement of BBYY is nevertheless a significant step forward, opening new research avenues about the locomotion capacities, seasonal ecology and even social behavior of Arctic hares, as they may be migrating in groups. [Extracted from the article]

Winter requires physiological adjustments in northern resident passerines. Cold acclimatization is generally associated with an increase in physiological maintenance costs, measured as basal metabolic rate (BMR), and cold endurance, reflected by summit metabolic rate ( M). However, several northern species also form social groups in winter and a bird's hierarchical position may influence the size of its metabolically active organs as well as its BMR. Winter metabolic performance in these species may therefore reflect a complex set of adjustments to both seasonal climatic variations and social environment. We studied the effect of social status on parameters of cold acclimatization (body mass, size of fat reserves and pectoral muscles, BMR and M) in free-living black-capped chickadees ( Poecile atricapillus). Birds that were structurally large and heavy for their body size, mostly dominant individuals, carried more fat reserves and had larger pectoral muscles. However, social status had little effect on metabolic performance in the cold. Indeed, M was independent of social rank while mass-corrected BMR was slightly lower in dominant individuals, likely due to a statistical dilution effect caused by large metabolically inactive fat reserves. BMR and M, whether considered in terms of whole-animal values, corrected for body mass or body size were nevertheless correlated, suggesting a functional link between these metabolic components. Our results therefore indicate that the energy cost of social dominance is not a generalized phenomenon in small wintering birds. [ABSTRACT FROM AUTHOR]

Current research in birds suggests that a conflict should exist during reproduction for the role of the glucocorticoid corticosterone (CORT). While elevated levels have been correlated with the increased energetic demand of raising offspring, elevated CORT levels have traditionally been implicated in reproductive abandonment. We examined the relationship between CORT and nest desertion in breeding wild female European starlings (Sturnus vulgaris) incorporating analyses of both total circulating levels and ‘free’, unbound CORT through analysis of corticosteroid-binding globulin (CBG). Free baseline CORT levels of nest-abandoning birds were significantly higher than nonabandoning birds within each stage, with chick-rearing birds exhibiting the highest free baseline CORT levels, while concurrently remaining the most resistant stage to nest desertion. Elevated free baseline CORT levels in chick-rearing birds were not due to increased total CORT secretion, but rather to a decrease in CBG levels. Overall, our results suggest that CORT and CBG interact to play a role in mediating the increased energetic demand of offspring, while minimizing the chances of nest desertion, thereby alleviating any potential behavioral conflict for CORT during reproduction. Furthermore, these results demonstrate that the traditional view of the role of CORT during reproduction is much more complex than previously appreciated. Together with mounting evidence, we suggest that elevated corticosteroid levels are an inherent and necessary part of reproduction in nonmammalian tetrapods. [Copyright &y& Elsevier]

Neonicotinoids are insecticides widely used as seed treatments that appear to have multiple negative effects on birds at a diversity of biological scales. Adult birds exposed to a low dose of imidacloprid, one of the most commonly used neonicotinoids, presented reduced fat stores, delayed migration and potentially altered orientation. However, little is known on the effect of imidacloprid on birds growth rate despite studies that have documented disruptive effects of low imidacloprid doses on thyroid gland communication. We performed a 2 × 2 factorial design experiment in Zebra finches, in which nestling birds were exposed to a very low dose (0.205 mg kg body mass - 1 ) of imidacloprid combined with food restriction during posthatch development. During the early developmental period, imidacloprid exposure resulted in an improvement of body condition index in treated nestlings relative to controls. Imidacloprid also led to compensatory growth in food restricted nestlings. This early life neonicotinoid exposure also carried over to adult age, with exposed birds showing higher lean mass and basal metabolic rate than controls at ages of 90–800 days. This study presents the first evidence that very low-dose neonicotinoid exposure during early life can permanently alter adult phenotype in birds. [ABSTRACT FROM AUTHOR]

Cold acclimatization (phenotypic adjustments to cope with cold conditions) is an imperative requirement for birds living at high latitudes during the cold depths of winter. Despite the significant remodelling of key phenotypic traits and energetic costs associated with elevating cold endurance, winter cold acclimatization can also provide further carryover benefits to subsequent stages in species wintering, migrating and breeding in cold environments (e.g. the Arctic).We tested this beneficial carryover hypothesis using outdoor captive Arctic‐breeding snow buntings Plectrophenax nivalis, a cold specialist known for its impressive wintering thermogenic capabilities. We compared changes in phenotypic traits supporting cold acclimatization—body composition (body, fat, lean mass, pectoral muscle thickness), oxygen carrying capacity (haematocrit), thermogenic capacity and endurance (Msum, time to Msum), cold tolerance (Ta at Msum) and maintenance energy expenditure (BMR)—between the wintering, migratory and arrival/summer stages.Body mass (+31%), fat mass (+226%) and BMR (+13%) increased relative to the winter phenotype, likely to support the added costs of migration—that is the migratory upregulation hypothesis. In contrast, lean mass, pectoral muscle thickness, haematocrit and thermogenic capacity remained high and stable at winter level across stages in support of the thermal carryover hypothesis. The maintenance of these traits likely offers spare capacity for unpredictable cold environments expected during migration and breeding in the Arctic.Our results thus suggest that birds can extend the long‐term advantages of winter phenotypic adjustments through additional benefits to thermogenic capacity during subsequent life‐history stages. These benefits likely make it possible for Arctic‐breeding birds to maximize success across diverse life‐history stages in the face of extreme cold conditions. [ABSTRACT FROM AUTHOR]

In altricial avian species, nutrition can significantly impact nestling fitness by increasing their survival and recruitment chances after fledging. Therefore, the effort invested by parents towards provisioning nestlings is crucial and represents a critical link between habitat resources and reproductive success. Recent studies suggest that the provisioning rate has little or no effect on the nestling growth rate. However, these studies do not consider prey quality, which may force breeding pairs to adjust provisioning rates to account for variation in prey nutritional value. In this 8-year study using black-capped (Poecile atricapillus) and boreal (Poecile hudsonicus) chickadees, we hypothesized that provisioning rates would negatively correlate with prey quality (i.e., energy content) across years if parents adjust their effort to maintain nestling growth rates. The mean daily growth rate was consistent across years in both species. However, prey energy content differed among years, and our results showed that parents brought more food to the nest and fed at a higher rate in years of low prey quality. This compensatory effect likely explains the lack of relationship between provisioning rate and growth rate reported in this and other studies. Therefore, our data support the hypothesis that parents increase provisioning efforts to compensate for poor prey quality and maintain offspring growth rates. [ABSTRACT FROM AUTHOR]

Arctic animals inhabit some of the coldest environments on the planet and have evolved physiological mechanisms for minimizing heat loss under extreme cold. However, the Arctic is warming faster than the global average and how well Arctic animals tolerate even moderately high air temperatures (Ta) is unknown.Using flow‐through respirometry, we investigated the heat tolerance and evaporative cooling capacity of snow buntings (Plectrophenax nivalis; ≈31 g, N = 42), a cold specialist, Arctic songbird. We exposed buntings to increasing Ta and measured body temperature (Tb), resting metabolic rate (RMR), rates of evaporative water loss (EWL), and evaporative cooling efficiency (the ratio of evaporative heat loss to metabolic heat production).Buntings had an average (±SD) Tb of 41.3 ± 0.2°C at thermoneutral Ta and increased Tb to a maximum of 43.5 ± 0.3°C. Buntings started panting at Ta of 33.2 ± 1.7°C, with rapid increases in EWL starting at Ta = 34.6°C, meaning they experienced heat stress when air temperatures were well below their body temperature. Maximum rates of EWL were only 2.9× baseline rates at thermoneutral Ta, a markedly lower increase than seen in more heat‐tolerant arid‐zone species (e.g., ≥4.7× baseline rates). Heat‐stressed buntings also had low evaporative cooling efficiencies, with 95% of individuals unable to evaporatively dissipate an amount of heat equivalent to their own metabolic heat production.Our results suggest that buntings' well‐developed cold tolerance may come at the cost of reduced heat tolerance. As the Arctic warms, and this and other species experience increased periods of heat stress, a limited capacity for evaporative cooling may force birds to increasingly rely on behavioral thermoregulation, such as minimizing activity, at the expense of diminished performance or reproductive investment. [ABSTRACT FROM AUTHOR]

Resident temperate passerines adjust their phenotypes to cope with winter constraints, with peak performance in metabolic traits typically occurring during the coldest months. However, it is sparsely known whether cold-adapted northern species make similar adjustments when faced with variable seasonal environments. Life in near-constant cold could be associated with limited flexibility in traits underlying cold endurance. We investigated this by tracking individual physiological changes over five consecutive winters in snow buntings (Plectrophenax nivalis), an Arctic-breeding migratory passerine typically confronted with nearly constant cold. Buntings were held in an outdoor aviary and exposed to seasonal temperature variation typical of temperate zone climates. We measured phenotypic changes in body composition (body, fat, and lean mass, pectoralis muscle thickness), oxygen transport capacity (hematocrit), metabolic performance (basal metabolic rate [BMR] and summit metabolic rate [ M sum]), thermogenic endurance (time to reach M sum), and cold tolerance (temperature at M sum). Snow buntings showed flexibility in functions underlying thermogenic capacity and cold endurance comparable to that observed in temperate resident passerines wintering at similar latitudes. Specifically, they increased body mass (13%), fat mass (246%), hematocrit (23%), pectoralis muscle thickness (8%), and M sum (27%). We also found remarkable cold tolerance in these birds, with individuals reaching M sum in helox at temperatures equivalent to less than −90°C in air. However, in contrast with resident temperate passerines, lean mass decreased by 12%, and there was no clear increase in maintenance costs (BMR). Our results show that the flexibility of traits underlying thermal acclimatization in a cold-adapted northern species is comparable to that of temperate resident species living at lower latitudes and is therefore not limited by life in near-constant cold. [ABSTRACT FROM AUTHOR]

Body temperature variation in response to acute stress is typically characterized by peripheral vasoconstriction and a concomitant increase in core body temperature (stress-induced hyperthermia). It is poorly understood how this response differs between species and within individuals of the same species, and how it is affected by the environment. We therefore investigated stress-induced body temperature changes in a non-model species, the Black-capped Chickadee, in two environmental conditions: outdoors in low ambient temperature (mean: − 6.6 °C), and indoors, in milder ambient temperature close to thermoneutrality (mean: 18.7 °C). Our results show that the change in body temperature in response to the same handling stressor differs in these conditions. In cold environments, we noted a significant decrease in core body temperature (− 2.9 °C), whereas the response in mild indoor conditions was weak and non-significant (− 0.6 °C). Heat loss in outdoor birds was exacerbated when birds were handled for longer time. This may highlight the role of behavioral thermoregulation and heat substitution from activity to body temperature maintenance in harsh condition. Importantly, our work also indicates that changes in the physical properties of the bird during handling (conductive cooling from cold hands, decreased insulation from compression of plumage and prevention of ptiloerection) may have large consequences for thermoregulation. This might explain why females, the smaller sex, lost more heat than males in the experiment. Because physiological and physical changes during handling may carry over to affect predation risk and maintenance of energy balance during short winter days, we advice caution when designing experimental protocols entailing prolonged handling of small birds in cold conditions. [ABSTRACT FROM AUTHOR]

Background Meniscus lesions are common musculoskeletal knee injuries which often lead to pain, limitation and compensations during functional tasks, such as descending stairs. This study investigated the effect of patellar taping with tension and without tension on three-dimensional (3D) kinematics of the knee during a slow step descent task in patients with meniscal lesions. Methods Ten patients diagnosed with a meniscal lesion, confirmed by magnetic resonance imaging, underwent five, step descent movements at slow speed under three different conditions: 1) no taping; 2) tension-free taping; and 3) patellar taping with medial tension. 3D kinematic data were recorded from the injured knee using an eight-camera infrared Vicon motion analysis system. Maximum and minimum angle values and total range of motion (maximum/minimum value) in three movement planes during single-limb stance were compared using a repeated measure ANOVA. Findings Results showed a significant increase in the maximum and minimum angle value in the sagittal plane (mean differences = 2.4° and 4.2°, respectively) and a decrease in the transverse plane (− 6.3° and − 2.2°, respectively) for the patellar taping condition compared to the no taping condition. A decreased rotational angle range when comparing the patellar taping to the no taping (− 4.1°) and tension-free taping (− 3.1°) conditions was also observed. These changes remained significant when pain was considered as a covariate in the analysis. The tension applied to the patellar tape played a role in controlling the sagittal and transverse plane step-down movement among patients in our study. Interpretation These results support the use of patellar taping with a medially oriented tension to help to reduce the transversal plane movement of the knee in this population and they bring new light to the taping effect. [ABSTRACT FROM AUTHOR]

Ultrasonography has proven to be a valuable noninvasive method of measure of muscle size in birds, but validation of its use in birds as small as black-capped chickadees (Poecile atricapillus; 11 g) is scarce. The effect of observers and life state (dead or alive) of models used for calibration on measurement quality is also poorly documented. Using 31 dead and 22 live chickadees, linear regressions between ultrasound and dissection measurements of pectoral and thigh muscles were fitted and compared between five different observers. R² values varied greatly between observers and were generally weaker in live birds, ranging between 0.02 and 0.59, despite high repeatability of measurement. Using equations calculated from dead birds to estimate muscle mass of live birds yielded much higher measurement errors (9%-18%) than when using equations calculated from live birds (5%-8%). Our results suggest that with careful training and using only calibration from live birds, ultrasonography can be a useful but limited tool to estimate muscle size of birds as small as the black-capped chickadee. [ABSTRACT FROM AUTHOR]

Plasma lipid metabolites triglycerides (TRIG) and glycerol (GLY) are used as indicators of fattening rate and nutritional condition in migratory birds. Requiring only one blood sample, they could also be used for studying daily and seasonal fattening rates in relation with habitat quality or weather variations in species wintering in cold climates. Using black-capped chickadees exposed to three experimental temperatures (0 °C, 15 °C, and 30 °C), the goal of this experiment was to determine the relationship between plasma levels of TRIG and GLY and fattening rate measured over periods from a few hours to the previous two days. Results showed that birds maintained in the cold had metabolite levels 39–81% higher than those at thermoneutrality, likely reflecting the size of their fat reserves, and that TRIG and total GLY were highly correlated across treatments. Fattening rate was also higher at 0 °C (+ 35%) and 30 °C (+ 24%) relative to that measured at 15 °C and, as expected, was positively correlated with metabolite levels across treatments. However, despite fattening rates similar to that observed at the other temperatures, the relationships were uncoupled at 30 °C, implying that the technique may not be easily applicable at temperatures within or close to thermoneutrality. We also found a strong individual effect in the relationships between fattening rate and TRIG levels, suggesting high individual consistency in these parameters in conditions of unrestricted food access such as in captivity. Our study confirms that plasma TRIG and GLY levels can be used as relative indexes of condition and fattening rates in wintering passerines. [ABSTRACT FROM AUTHOR]

Stochastic winter weather events are predicted to increase in occurrence and amplitude at northern latitudes and organisms are expected to cope through phenotypic flexibility. Small avian species wintering in these environments show acclimatization where basal metabolic rate (BMR) and maximal thermogenic capacity (MSUM) are typically elevated. However, little is known on intra-seasonal variation in metabolic performance and on how population trends truly reflect individual flexibility. Here we report intra-seasonal variation in metabolic parameters measured at the population and individual levels in black-capped chickadees (Poecile atricapillus). Results confirmed that population patterns indeed reflect flexibility at the individual level. They showed the expected increase in BMR (6%) and MSUM (34%) in winter relative to summer but also, and most importantly, that these parameters changed differently through time. BMR began its seasonal increase in November, while MSUM had already achieved more than 20% of its inter-seasonal increase by October, and declined to its starting level by March, while MSUM remained high. Although both parameters co-vary on a yearly scale, this mismatch in the timing of variation in winter BMR and MSUM likely reflects different constraints acting on different physiological components and therefore suggests a lack of functional link between these parameters. [ABSTRACT FROM AUTHOR]