Baseline and stress-induced plasma corticosterone during long-distance migration in the bar-tailed godwit, Limosa lapponica

The specific roles of corticosterone in promotion of avian migration remain unclear even though this glucocorticosteroid is elevated in many migrating bird species. In general, glucocorticosteroids promote metabolic homeostasis and may elicit effects on feeding and locomotion. Because the migratory stages of refueling and flight are characterized by distinct behaviors and physiology, the determination of corticosterone levels during each stage should help identify potential processes in which corticosterone is involved. We measured baseline levels of corticosterone in bar-tailed godwits (Limosa lapponica) during two distinct stages of migration: (1) immediately after arrival at a false stopover site just short of the Wadden Sea and (2) throughout the subsequent 4-wk refueling period on the Wadden Sea. Plasma corticosterone was higher in arriving than in refueling birds. In addition, corticosterone increased with size-corrected body mass during the refueling phase, suggesting that corticosterone rises as birds prepare to reinitiate flight. Therefore, elevated corticosterone appears associated with migratory flight and may participate in processes characterizing this stage. We also performed a capture stress protocol in all birds and found that corticosterone increased in both arriving and refueling godwits. Therefore, the normal course of migration may be typified by corticosterone concentrations that are lower than those associated with stressful and life-threatening episodes.     

Baseline and stress-induced plasma corticosterone concentrations of mice selectively bred for high voluntary wheel running

The hypothalamic-pituitary-adrenal (HPA) axis is important in regulating energy metabolism and in mediating responses to stressors, including increasing energy availability during physical exercise. In addition, glucocorticoids act directly on the central nervous system and influence behavior, including locomotor activity. To explore potential changes in the HPA axis as animals evolve higher voluntary activity levels, we characterized plasma corticosterone (CORT) concentrations and adrenal mass in four replicate lines of house mice that had been selectively bred for high voluntary wheel running (HR lines) for 34 generations and in four nonselected control (C) lines. We determined CORT concentrations under baseline conditions and immediately after exposure to a novel stressor (40 min of physical restraint) in mice that were housed without access to wheels. Resting daytime CORT concentrations were approximately twice as high in HR as in C mice for both sexes. Physical restraint increased CORT to similar concentrations in HR and C mice; consequently, the proportional response to restraint was smaller in HR than in C animals. Adrenal mass did not significantly differ between HR and C mice. Females had significantly higher baseline and postrestraint CORT concentrations and significantly larger adrenal glands than males in both HR and C lines. Replicate lines showed significant variation in body mass, length, baseline CORT concentrations, and postrestraint CORT concentrations in one or both sexes. Among lines, both body mass and length were significantly negatively correlated with baseline CORT concentrations, suggesting that CORT suppresses growth. Our results suggest that selection for increased locomotor activity has caused correlated changes in the HPA axis, resulting in higher baseline CORT concentrations and, possibly, reduced stress responsiveness and a lower growth rate.     

Resource use for reproduction depends on spring arrival time and wintering area in an arctic breeding shorebird

Resources for egg production may come from body reserves stored before breeding (“capital breeders”) or from food acquired at the breeding site (“income breeders”). Arctic migrants were long thought to be capital breeders, because they often arrive at a time when local food availability is still limited. However, later evidence suggested that arctic breeding shorebirds are primarily income breeders, or that they use a mixed strategy depending on laying date. We explored the relationship between laying date and resource use for reproduction in the pectoral sandpiper Calidris melanotos breeding in the Alaskan arctic by contrasting carbon isotope (δ¹³C) values of the local diet and of maternal plasma, cellular blood, feather and claw with those of the eggs produced. Our results revealed that early breeding females utilize resources for egg production that were acquired recently at staging areas, whereas later breeding females mostly relied on nutrients derived from local food sources. These findings suggest that the resource allocation strategy used for reproduction differs among females, and varies depending on the timing of arrival and the start of reproduction. The arrival date at the breeding ground and laying date may critically depend on non‐breeding season events such as winter habitat choice, staging areas or migration routes. By comparing maternal feather δ¹³C, claw δ¹³C and feather δD, we examined whether non‐breeding season events influenced the use of resources for egg production through variation in capture date or clutch initiation date. Female pectoral sandpipers originating from moulting areas characterized by higher (more positive) δD signatures were caught earlier and started laying earlier, and they used stored resources for reproduction. Using regional maps of δD values for precipitation in the wintering sites in South America, we compared the spatial variation in the observed feather δD signatures. This analysis indicated that female pectoral sandpipers with higher δD signatures, presumably coming from more north‐easterly wintering sites in southern America, started laying earlier and used mostly stored resources for egg production, compared to females that wintered (or at least moulted) further south. Our results thus show that winter moulting habitat is linked to breeding resource allocation strategy in this high‐arctic breeding shorebird.     

Basal and stress-induced corticosterone levels of garden warblers,Sylvia borin,during migration

Summary Plasma levels of the metabolically and behaviorally active corticosteroid hormone, corticosterone, were studied in garden warblers in the laboratory and in the field during the autumnal migratory phase. Garden warblers showing nocturnal migratory activity in the laboratory had elevated levels of corticosterone at the end of the dark phase and low levels during daytime. When nocturnal migratory activity was experimentally disrupted by food deprivation and subsequent refeeding or after spontaneous termination of migratory activity this rhythm was absent. Garden warblers stopping over in the Sahara desert during autumnal migration had low levels of corticosterone. Levels were negatively correlated with fat stores and body mass in birds sampled throughout the day. These levels were generally lower than those associated with stress in response to repeated handling and blood sampling. The results suggest (1) the existence of diel changes in adrenocortical hormonal activity that could be involved in regulation of migration, and (2) that garden warblers carrying large fat depots are not stressed by prolonged flight or lack of appropiate feeding areas during migration over the desert.Abbreviations ACTH adrenocorticotrope hormone - cpm counts per minute - EtOH ethanol - RIA radioimmunoassay     

Serum prolactin levels and crop-sac development in ring doves during a breeding cycle

Using a turkey prolactin radioimmunoassay, the serum prolactin levels of male and female ring doves (Streptopelia risoria) during the breeding cycle were measured and their circulating prolactin levels were compared to crop-sac weight on a within-bird basis. During the early phase of the incubation period, crop weight showed a delayed response to prolactin stimulation and there was no correlation; during the midincubation period when prolactin and crop-sac weight were increasing, there was a strong positive correlation; around hatching, when prolactin was at its peak, there was no correlation. There were again strong positive correlations at later samples, when squabs were developing and both prolactin and crop-sac weight were declining. Thus, it appears that the correlation between the circulating prolactin level and crop-sac development depends on the stage of the breeding cycle. While males and females showed similar pattern of circulating prolactin during the period of incubation and parental care, only females consistently showed a postovulatory rise of prolactin. These results were discussed in the context of the role of prolactin in the breeding cycle.     

Repeatability of baseline and stress-induced corticosterone levels across early life stages in the Florida scrub-jay (Aphelocoma coerulescens)

Recent studies have posited that the pattern of glucocorticoid secretion within an individual represents a stable, fixed physiological trait. To test this hypothesis, we assessed the repeatability of baseline and stress-induced corticosterone (CORT) secretion across developmental stages and years in Florida scrub-jays. We sampled individuals from multiple cohorts repeatedly from the age of 11 days post-hatch up to 4 years of age. We found a significant degree of repeatability within individuals in stress-induced corticosterone levels, i.e., the amount of hormone secreted during a standardized stress protocol (corrected integrated corticosterone). However, baseline corticosterone levels were not statistically repeatable, although there was some indication that nestling levels predicted levels at 1 year of age. The results of this study indicate that stress-induced CORT levels are consistent within individual scrub-jays, and the degree to which a young jay mounts an acute stress response appears to be somewhat “set” by the age of nutritional independence. Thus stress-induced corticosterone secretion appears to be a stable, repeatable trait within individuals and as such may be subject to natural selection.     

An experimental test of the relationship between temporal variability of feeding opportunities and baseline levels of corticosterone in a shorebird

In this study, we tested the hypothesis that baseline corticosterone levels increase with a change from constant to variable feeding schedules. Captive red knots, Calidris canutus, were presented with food that was either available during the same time each day (constant) or starting at variable times during the day. Food intake rates, frequency of aggressive interactions, and baseline levels of corticosterone were measured. In the majority of cases, red knots showed higher plasma corticosterone concentrations during feeding schedules that were irregular than when food was available at consistent times. These findings are supported by a previous study that showed that red knots take a long time to adjust to the newly offered, predictable conditions of their aviary environment. The frequency of conflicts in the different groups and (size-corrected) body mass were not correlated with average corticosterone level. The results are examined in the light of literature showing that increases in corticosterone in response to acute, unpredictable events mediate behavioral responses such as increased explorative behavior and memory. For red knots that have to find their food on the temperate-zone mudflats in Western Europe, an increased circulating corticosterone level may be adaptive during periods when the patchily distribution of buried bivalves and the burying behavior of such prey presents them with a variable and unpredictable food supply.     

Body condition and feather molt of a migratory shorebird during the non‐breeding season

Migratory shorebirds have some of the highest fat loads among birds, especially species which migrate long distances. The upland sandpiper Bartramia longicauda makes long‐distance migrations twice a year, but variation in body condition or timing of feather molt during the non‐breeding season has not been studied. Molt is an important part of the annual cycle of migratory birds because feather condition determines flight performance during migration, and long‐distance movements are energetically costly. However, variation in body condition during molt has been poorly studied. The objective of our field study was to examine the timing and patterns of feather molt of a long distance migratory shorebird during the non‐breeding season and test for relationships with body size, fat depots, mass, and sex. Field work was conducted at four ranches in the Northern Campos of Uruguay (Paysandú and Salto Departments). We captured and marked 62 sandpipers in a 2‐month period (Nov–Jan) during four non‐breeding seasons (2008–2012). Sex was determined by genetic analyses of blood samples taken at capture. Molt was measured in captured birds using rank scores based on published standards. Body mass and tarsus length measurements showed female‐biased sexual size dimorphism with males smaller than females. Size‐corrected body mass (body condition) showed a U‐shaped relationship with the day of the season, indicating that birds arrived at non‐breeding grounds in relatively good condition. Arriving in good body condition at non‐breeding grounds is probably important because of the energetic demands due to physiological adjustments after migration and the costs of feather molt.     

Effects of switches in nutrient levels during the life cycle of spring wheat

Accumulation of cations in roots and shoots and influx of K⁺(⁸⁶Rb⁺) and Ca²⁺(⁴⁵Ca²⁺) into the roots were investigated in spring wheat (Triticum aestivum L. cv. Svenno). Plants were sampled at four main developmental stages: tillering, shooting, heading and grain filling. The effects of switches between a high and a low supply of nutrients were characterized. Growth of the shoots and roots was affected by the switches. A high supply of nutrients at the seedling stage and towards tillering supported a high growth rate, while a further high supply of nutrients increased vegetative growth and delayed grain filling. An early high supply of nutrients followed by a low supply at shooting, heading and grain filling accelerated root growth and growth of the main culm. Switches of the mineral supply gave only small changes in concentration of Mg in the plants. Generally, the K⁺(⁸⁶Rb⁺) influx into the roots decreased during ontogenesis, while Ca²⁺(⁴⁵Ca²⁺) influx increased more or less independent of the switches between nutrients levels.     

Effects of pair migratory behavior on breeding phenology and success in a partially migratory shorebird population

In migratory systems, variation in individual phenology can arise through differences in individual migratory behaviors, and this may be particularly apparent in partial migrant systems, where migrant and resident individuals are present within the same population. Links between breeding phenology and migratory behavior or success are generally investigated at the individual level. However, for breeding phenology in particular, the migratory behaviors of each member of the pair may need to be considered simultaneously, as breeding phenology will likely be constrained by timing of the pair member that arrives last, and carryover effects on breeding success may vary depending on whether pair members share the same migratory behavior or not. We used tracking of marked individuals and monitoring of breeding success from a partially migrant population of Eurasian oystercatchers (Haematopus ostralegus) breeding in Iceland to test whether (a) breeding phenology varied with pair migratory behavior; (b) within‐pair consistency in timing of laying differed among pair migratory behaviors; and (c) reproductive performance varied with pair migratory behavior, timing of laying, and year. We found that annual variation in timing of laying differed among pair migratory behaviors, with resident pairs being more consistent than migrant and mixed pairs, and migrant/mixed pairs breeding earlier than residents in most years but later in one (unusually cold) year. Pairs that laid early were more likely to replace their clutch after nest loss, had higher productivity and higher fledging success, independent of pair migratory behavior. Our study suggests that the links between individual migratory behavior and reproductive success can vary over time and, to a much lesser extent, with mate migratory behavior and can be mediated by differences in laying dates. Understanding these cascading effects of pair phenology on breeding success is likely to be key to predicting the impact of changing environmental conditions on migratory species.     

Diurnal variability in the detection of waders on their breeding grounds

When censusing breeding waders in large upland areas, the time of day has a major influence on the numbers of birds detected.     

Metabolic correlates of leg length in breeding arctic shorebirds: the cost of getting high

Aim We test the hypothesis that tarsus length in all shorebirds breeding in the Canadian arctic shows an evolutionary response to average metabolic stress encountered across the breeding range, such that birds nesting in metabolically stressful environments have relatively shorter legs. Longer‐legged birds living in colder environments will experience greater metabolic costs because their torsos are elevated farther away from the ground's wind‐dampening boundary layer. Methods We use weather data (temperature, wind speed, global solar radiation) from 27 arctic weather stations measured over 37 years, and a previously published model of heat transfer, to characterize the metabolic harshness over the breeding season of the ranges of each of the 17 shorebirds of the family Charadriidae nesting in the Canadian arctic. Results After controlling for the lengths of two other body extremities (wing and bill), there was a significant negative relationship between tarsus length and mean metabolic harshness. This result was obtained whether species were treated as independent data points, or in a comparative analysis using standardized independent contrasts. Main conclusions We support a unique extension of Allen's rule: body‐supporting appendages of homeotherms may be shorter in colder environments so as to take advantage of a boundary layer effect, thereby reducing metabolic costs.     

Testosterone and corticosterone during the breeding cycle of equatorial and European stonechats (Saxicola torquata axillaris and S. t. rubicola)

Northern-temperate male birds show seasonal changes in testosterone concentrations with a peak during the breeding season. Many tropical birds express much lower concentrations of testosterone with slight elevations during breeding. Here we describe testosterone and corticosterone concentrations of male stonechats from equatorial Kenya during different substages of breeding and molt. This tropical species has a short breeding season of approximately 3 months. We compare their hormone concentrations to previously published data of males of a northern-temperate relative, the European stonechat, also a seasonal breeder but with a breeding season of approximately 5 months. Equatorial stonechats show a pronounced peak of testosterone during the nest-building and laying stage. During all other stages, testosterone concentrations are low, similar to other year-round territorial tropical bird species. Corticosterone concentrations peak also during the nest-building and laying stage suggesting that this period of maximum female fecundity is a demanding period for the male. Equatorial stonechats have significantly lower concentrations of testosterone than European stonechats during all stages, except during the nest-building and laying stage. During this stage of maximum female fertility, testosterone levels tend to be higher in equatorial than in European stonechats. Our results suggest that equatorial stonechats belong to a group of tropical bird species that are characterized by a short breeding season and a brief high peak of testosterone during the female's fertile period. Such brief, but substantial peaks of testosterone may be common in tropical birds, but they may easily be missed if the exact breeding stage of individual birds is not known.     

Resource allocation in a social wasp: effects of breeding system and life cycle on reproductive decisions

Organisms must make important decisions on how to allocate resources to reproduction. We investigated allocation decisions in the social wasp Vespula maculifrons to understand how social insects make reproductive choices. We first determined how annual colonies apportioned resources to growth and reproduction by analysing developing brood. In contrast to expectations, colonies invested in both growth (workers) and reproduction (males) simultaneously. In addition, colonies showed evidence of producing males in pulses and reversing their reproductive choices by decreasing investment in males late in the season. This reversal is consistent with theory suggesting that colonies decrease production in males if fitness of late emerging males is low. To further investigate reproductive decisions within colonies, we determined if the male mates of multiply-mated queens varied in their reproductive success over time. Sperm use by queens did vary over time suggesting that male success may depend on sperm clumping within the female reproductive tract. Finally, we tested if colony sex ratio conformed to expectations under kin selection theory that nestmate relatedness would positively correlate with investment in new queens if workers controlled sex allocation. Surprisingly, the proportion of queens produced by colonies was negatively correlated with nestmate relatedness, suggesting that allocation may be shaped by advantages arising from increased genetic diversity resulting from multiple mating by queens. Overall, our study suggests that the reproductive decisions of colonies are flexible and may depend both on environmental cues arising from energetic needs of the colony and genetic cues arising from mating behaviours of queens.     

Changes in polyamine levels in various organs of Bombyx mori during its life cycle

Polyamines in various organs of larval, pupal, and moth stages of Bombyx mori, were assayed by high-performance ion-exchange chromatography and paper and thin-layer chromatography. Putrescine and spermidine were especially abundant in the silk gland, gonads, mucous gland, and sucking stomach; spermine was also present in them, but at much lower concentrations. Both norspermidine and norspermine were detected in almost all organs examined, while their precursor 1,3-diaminopropane was found only in a limited number of organs. Low concentrations of sym-homospermidine were observed in the silk gland and ovary. Cadaverine content was particularly high in the mucous gland which contained diapause eggs and the sucking stomach. Diapause eggs contained much higher levels of cadaverine than non-diapause eggs. The concentrations of most polyamines in the silk glands remained rather constant during the larval stage, and decreased markedly at the pupal stage. Polyamines in gonads, in contrast, did not decrease at the pupal stage, but putrescine, diaminopropane, and norspermidine rather increased during the pupal and moth stages.     

Habituation and habitat changes can moderate the impacts of human disturbance on shorebird breeding performance

Disturbance by humans is widely expected to reduce the reproductive fitness of nesting birds if disturbance reduces nest attentiveness, and unattended eggs experience increased risk of predation or exposure to potentially lethal temperature extremes. Yet, relatively few studies have examined the physiological or behavioural mechanisms whereby disturbance influences reproductive fitness, or the extent to which the costs of disturbance may be reduced through habituation. We compared the behavioural responses, egg temperatures and reproductive success of shore-nesting white-fronted plovers Charadrius marginatus to disturbance at two breeding sites experiencing low versus high human recreational activity, respectively. Daytime nest attentiveness decreased with increasing experimental disturbance at both sites, but this relationship differed between sites; for any given level of disturbance, incubating birds at the more disturbed site had greater nest attentiveness. They achieved this through habituation, allowing a closer human approach before leaving the nest, and returning to the nest faster after a disturbance event. Despite lower average daytime nest attentiveness at the more disturbed site, incubation temperatures did not differ significantly between sites. Nest mortality, mostly by natural mammalian and corvid predators, was significantly lower at the site experiencing high recreational activity. However, chick mortality was significantly greater at the more disturbed site, most likely because of predation by domestic dogs. Chick mortality may have been increased by the habitation of chicks, whose escape responses were much reduced at the more disturbed site. Nonetheless, annual fecundity was substantially higher at the more disturbed site, showing that the overall reproductive fitness of wild birds is not always compromised by human disturbance and urbanization.     

Rapid changes in the size of different functional organ and muscle groups during refueling in a long-distance migrating shorebird.

The adaptive value of size changes in different organ and muscle groups was studied in red knots (Calidris canutus islandica) in relation to their migration. Birds were sampled on five occasions: at arrival in Iceland in May 1994, two times during subsequent refueling, at departure toward, and on return from, the high arctic breeding grounds. During their 24-d stopover in May, body mass increased from 144.3 to 214.5 g. Mass gains were lowest over the first week (0.85 g/d, only fat-free tissue deposited). Over the subsequent 10 d, average mass increased by 5.0 g/d (fat contributing 78%), and over the last week before takeoff, it increased by 2.0 g/d (fat contributing over 100% because of loss of lean components). There were no sex differences in body and fat mass gains. Over the first interval, lean masses of heart, stomach, and liver increased. During the middle 10 d, sizes of leg muscle, intestine, liver, and kidneys increased. Stomach mass decreased over the same interval. In the last interval before takeoff, the stomach atrophied further and the intestine, leg muscles, and liver became smaller too, but pectoral muscles and heart increased in size. Sizes of "exercise organs" such as pectoral muscle and heart were best correlated with body mass, whereas sizes of organs used during foraging (leg muscles) and nutrient extraction (intestine, liver) were best correlated with rate of mass gain. Kidneys changed little before takeoff, which suggests that they are needed as much during flight as during refueling.     

Tropical winter habitat limits reproductive success on the temperate breeding grounds in a migratory bird

Identifying the factors that control population dynamics in migratory animals has been constrained by our inability to track individuals throughout the annual cycle. Using stable carbon isotopes, we show that the reproductive success of a long-distance migratory bird is influenced by the quality of habitat located thousands of kilometres away on tropical wintering grounds. For male American redstarts (Setophaga ruticilla), winter habitat quality influenced arrival date on the breeding grounds, which in turn affected key variables associated with reproduction, including the number of young fledged. Based on a winter-habitat model, females occupying high-quality winter habitat were predicted to produce more than two additional young and to fledge offspring up to a month earlier compared with females wintering in poor-quality habitat. Differences of this magnitude are highly important considering redstarts are single brooded, lay clutches of only three to five eggs and spend only two-and-a-half months on the breeding grounds. Results from this study indicate the importance of understanding how periods of the annual cycle interact for migratory animals. Continued loss of tropical wintering habitat could have negative effects on migratory populations in the following breeding season, minimizing density-dependent effects on the breeding grounds and leading to further population declines. If conservation efforts are to be successful, strategies must incorporate measures to protect all the habitats used during the entire annual cycle of migratory animals.     

Changes in blood histamine and bradykininogen levels during different periods of the oestrous cycle in goats.

Blood histamine and plasma bradykininogen levels were estimated in eighteen normally cyclic goats at different periods of the oestrous cycle. The histamine level increased significantly (P smaller than 0.05) on Day 18 of the oestrous cycle. The bradykininogen level decreased significantly (P smaller than 0.05) towards the end of oestrus.