Corticosterone in migrating songbirds during endurance flight

The specific role of the glucocorticoid hormone corticosterone in regulating the migratory stages of flight and refueling in free-living migrants is as yet poorly studied, because these stages are difficult to identify in the field. Night-migrating songbirds provide an excellent model to investigate how corticosterone correlates with behavior and physiology because they fly during the night and rest and forage during the day. We measured baseline corticosterone and the adrenocortical response to restraint in 9 free-ranging songbird species: 3 night-migrating species, 3 day-migrating species, and 3 day-migrating irruptive species. Baseline corticosterone of night migrants was higher in birds caught out of nocturnal migration than in birds resting and foraging, and on the same level as in day migrants, suggesting that a rise in circulating corticosterone may facilitate the heightened metabolic processes of active flight, in particular protein breakdown. Stress-induced corticosterone levels increased in both actively flying birds and birds resting and foraging. The increase was highest in landing birds, which are possibly most sensitive to stress when arriving at an unfamiliar place. Migratory endurance flight is thus characterized by corticosterone concentrations that are lower than those associated with acute stressful and life-threatening episodes. In addition, the responsiveness to stress increased with decreasing fat score in a night-migrating species. Corticosterone approaches therefore stressful concentrations only when fat depots are nearly depleted, possibly to promote protein catabolism and to trigger a change in behavior, i.e., a switch to landing and searching for food.     

Seasonal and diel transitions in physiology and behavior in the migratory dark-eyed junco

Body mass, fat stores, activities of lipogenic and lipolytic enzymes, and plasma corticosterone were measured throughout seasonal and diel transitions from fall through spring encompassing the non-migratory stages of early and mid winter, the prealternate molt, and the spring migratory stage in captive dark-eyed juncos to determine the physiological mechanisms underlying adaptations for migration. On a seasonal basis, lipid enzymes and corticosterone varied little throughout the stages even though the birds underwent dramatic alterations in mass, fat deposition, behavior, and activation of the reproductive axis. By contrast, diel changes were found in lipogenesis, lipolysis, muscle lipoprotein lipase, and plasma corticosterone when comparing birds in the two phases of spring migration--active flight and resting, as during times of stopover. In these two phases of migration, coordination of the lipogenic and lipolytic systems appear to maximize storage of fatty acids during rest and delivery/utilization during flight. Diel patterns of corticosterone revealed fairly consistent peaks during the night time (23:00) throughout the nonmigratory period. The profile of this pattern altered during the migratory period with variation between the flight and resting phases. In sum, the results from these captive studies offer a new approach for studying the regulation of migratory physiology in free-living birds.     

The daily pattern of autumn bird migration in the northern Sahara

The temporal pattern of migration by passerine birds during the night, and their arrival during the day at the Egyptian coast and in the northern Sahara Desert was investigated. The mean direction of nocturnal migration at the coast was south-southeast, while at all desert sites it was south-southwest.
Birds arrived at the Egyptian coast only during the second half of the night which is explained by the fact that no birds could have taken off from the Mediterranean Sea. At least some of the birds landed at the coast where they spent the day before taking off shortly after sunset. These birds passed the desert sites at the expected time of day assuming a ground speed of 18 m per second. However, the origin of the birds passing the desert sites early at night is unclear. They must either have spent the day in the desert north of the study sites or they had overflown the Egyptian coast in the afternoon without landing.
The landing of birds during the day at the desert sites was bimodal. This pattern of arrival is explained either by some birds having landed at the Egyptian coast in the early morning before continuing, or by deteriorating conditions later in the day during flight or when resting in the desert, that obliged them to seek shelter at the desert sites.
A correlation between the number of migrants observed during the night and the number of resting birds in the desert on the following day suggests that an unknown proportion of birds might regularly use an intermittent migratory strategy that includes rest periods by day when crossing the desert, whereas others might adapt a non-stop migratory strategy.     

Flight by night or day? Optimal daily timing of bird migration

Many migratory bird species fly mainly during the night (nocturnal migrants), others during daytime (diurnal migrants) and still others during both night and day. Need to forage during the day, atmospheric structure, predator avoidance and orientation conditions have been proposed as explanations for the widespread occurrence of nocturnal migration. However, the general principles that determine the basic nocturnal-diurnal variation in flight habits are poorly known. In the present study optimal timing of migratory flights, giving the minimum total duration of the migratory journey, is evaluated in a schematic way in relation to ecological conditions for energy gain in foraging and for energy costs in flight. There exists a strong and fundamental advantage of flying by night because foraging time is maximized and energy deposition can take place on days immediately after and prior to the nocturnal flights. The increase in migration speed by nocturnal compared with diurnal migration will be largest for birds with low flight costs and high energy deposition rates. Diurnal migration will be optimal if it is associated with efficient energy gain immediately after a migratory flight because suitable stopover/foraging places have been located during the flight or if energy losses during flight are substantially reduced by thermal soaring and/or by fly-and-forage migration. A strategy of combined diurnal and nocturnal migration may be optimal when birds migrate across regions with relatively poor conditions for energy deposition (not only severe but also soft barriers). Predictions about variable timing of migratory flights depending on changing foraging and environmental conditions along the migration route may be tested for individual birds by analysing satellite tracking results with respect to daily travel routines in different regions. Documenting and understanding the adaptive variability in daily travel schedules among migrating animals constitute a fascinating challenge for future research.     

Corticosterone and time-activity budget: an experiment with Black-legged kittiwakes

In vertebrates, the well established increase in plasma corticosterone in response to food shortage is thought to mediate adjustments of foraging behavior and energy allocation to environmental conditions. However, investigating the functional role of corticosterone is often constrained by the difficulty to track time-activity budget of free-ranging animals. To examine how an experimental increase in corticosterone affects the activity budget of male Black-legged kittiwakes (Rissa tridactyla), we used miniaturized activity loggers to record flying/foraging, presence on the sea surface and nest attendance. To investigate how corticosterone affects allocation processes between self-foraging and foraging devoted to the brood, we monitored body mass change of males from capture (day 0) to recapture (day 3). Among control birds, males in poor condition at day 0 spent significantly more time flying/foraging and less time attending the nest site than did males in good condition. Corticosterone treatment affected time spent flying/foraging in interaction with body condition at day 0: corticosterone-implanted males in good condition spent more time flying/foraging than control ones; this was not observed in poor condition males. In control birds, change in body mass was negatively correlated with body condition at day 0. This was reinforced by corticosterone treatment and, on average, corticosterone-implanted males gained much more mass than controls. These results suggest that in Black-legged kittiwakes, body condition and corticosterone levels can interact to mediate foraging decisions and possibly energy allocation: when facing stressful environmental conditions, birds in good body condition may afford to increase the time spent foraging probably to maintain brood provisioning, whereas poor body condition birds seemed rather to redirect available energy from reproduction to self-maintenance.     

Plasma corticosterone increases during migratory restlessness in the captive white-crowned sparrow Zonotrichia leucophrys gambelli

Plasma corticosterone increases during the period of spring migration in a variety of bird species. Long-distance migrants show elevations in corticosterone specifically in association with the stage of flight, suggesting that corticosterone may support flight-related processes, for example, locomotor activity and/or energy mobilization. The pattern of corticosterone secretion as it relates to migratory flight has hitherto not been clearly described in migrants that frequently interrupt flight to refuel, for example, the Gambel's white-crowned sparrow (Zonotrichia leucophrys gambelii). The Gambel's white-crowned sparrow fuels by day and expresses peak migratory activity during the first few hours of night. To determine if plasma corticosterone increases in association with the stage of migratory flight also in this short-bout migrant, we induced captive white-crowned sparrows to enter into the migratory condition by placing photosensitive birds on long days (16L:8D) and then evaluated birds for plasma corticosterone and locomotor activity during four time points of the day. Patterns found in long-day birds were compared to those observed in short-day controls (8L:16D). Differences in energy metabolism as determined from plasma metabolites were also evaluated. We found that locomotor activity and corticosterone were significantly elevated at the onset of the dark period, but only in long-day birds. Plasma beta-hydroxybutyrate (a ketone body) was also elevated. Thus, findings suggest that plasma corticosterone and ketogenesis increase in association with migratory restlessness in a short-bout migrant. In fact, corticosterone may play a regulatory role, because it shows a trend to increase already before night-time activity.     

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.     

长距离迁徙鸟类对应于能量积累状态的取食行为调整

迁徙鸟类能够预计到迁徙过程中对能量需求的增加和迁徙途中获得能量的不确定性。最佳迁徙理论指出：迁徙停留期的一系列决策受到体内能量状态、取食机会和迁徙时间的影响。利用刚完成春季跨越墨西哥湾迁徙的鸫类,我们研究了取食行为、体内能量状态和能量积累速度的相互关系。我们用雾网捕获了停留的鸫,然后给每只鸟进行了环志和称重,并估测了表皮下积累的脂肪。为了检测体内能量积累大小对取食行为的影响,我们把在野外观察到的鸟分成肥、瘦两组。当一天里被捕获鸟的平均体重低于相关种的瘦体重时,这一天被观察到的取食鸟被归到瘦组,反之就属于肥组。我们同时对一部分鸟用有色环进行了标记,以便能在野外观察到它们时能准确地知道每一只鸟的初始能量积累状态。在春天完成跨越墨西哥湾迁徙以后的鸫大约有50％在到达停留地时就已经消耗了所有的表皮下脂肪。与肥组鸟相比,瘦组的个体在停留期扩展了取食方法和取食基底,增加了取食速度。与此相关的是,瘦鸟表现出体重积累更多、速度更快。我们的数据表明迁徙鸟在到达迁徙中途停留地时的能量状态会影响到它们停留期间的取食行为和继续迁徙时的生理机能,从而影响停留期的长短和在迁徙途中停留与否的决定[动物学报51(1)：12—23,2005]。     

Within night correlations between radar and ground counts of migrating songbirds

ABSTRACT.   Studies comparing numbers of nocturnal migrants in flight with numbers of migrants at stopover sites have produced equivocal results. In 2003, we compared numbers of nocturnal migrants detected by radar to numbers of passerines observed at the Atlantic Bird Observatory in southwestern Nova Scotia, Canada. Numbers of nocturnal migrants detected by radar were positively correlated with numbers of migrants as determined by mist-netting, censuses, and daily estimated totals (daily estimates of birds present based netting and census results and casual observations) the following day. On nights with winds favorable for migration (tailwinds), the peak correlation between ground counts and radar counts the night before occurred just after sunset. On nights with unfavorable winds (headwinds), the correlation increased through the night, with a peak just before sunrise. The patterns of correlation are consistent with a scenario where birds accumulate at the coastline during periods of unfavorable wind, likely because they are not willing to cross a major ecological barrier, the Gulf of Maine. On nights with favorable winds, many birds departed, but some, possibly after testing wind conditions, apparently decided not to cross the Gulf of Maine and returned. Our results suggest that combining data collected using different methods to generate a daily estimated total provides the best estimate of the number of migrants present at a stopover site. Simultaneous studies at multiple locations where different census methods are used, making more effective use of temporal data (both from radar and diurnal counts), will more clearly elucidate patterns of flight behavior by migratory songbirds and the relationship between ground counts and counts of birds aloft.     

Monitoring flight calls of migrating birds from an oil platform in the northern Gulf of Mexico

ABSTRACT.   Millions of birds migrate across the Gulf of Mexico each year. However, most studies of migration in this region involve sampling onshore locations during the day, potentially underrepresenting the diversity and abundance of migrants passing the region. We evaluated a potential solution to this problem by recording the flight calls of passing migrants from an oil platform located southeast of the Alabama coast in the Gulf of Mexico. We detected 2762 calls during 30 nights from 9 September to 2 November 1999, and were able to identify 2329 calls to species. Flight calls by nine species of birds represented 23% of all identified calls. The greatest number of calls during one night (1017 calls) and during a 1-h period (257 calls) were recorded on 10 September. The greatest number of calls was recorded 8 h after sunset, with a secondary peak 2 h after sunset. The peak prior to sunrise may indicate the formation of flocks at dawn, and the peak after sunset may have been caused by the first wave of migrants reaching the platform. However, call counts varied extensively, with 98% of all calls recorded during 13 nights and 40% on a single night, possibly resulting from hourly and nightly differences in bird numbers aloft, atmospheric conditions, and artificial lighting conditions. Although recording on oil platforms can be difficult because of mechanical, wind, and wave noise, our results suggest great potential for describing the species composition of passing vocal migrants and the temporal patterns of flight-calling behavior if quiet recording locations can be found. Moreover, flight call monitoring could be a critically important tool for bird conservation in this region, given recent proposals to develop wind power and the potential bird mortality associated with such developments.     

Role of the low-affinity glucocorticoid receptor in the regulation of behavior and energy metabolism in the migratory red knot Calidris canutus islandica

Plasma corticosterone increases in association with migratory flight in the red knot Calidris canutus islandica, suggesting that corticosterone may promote migratory activity and/or energy mobilization in this species. This hypothesis is supported by general effects of glucocorticoids, which include stimulation of locomotion and the mobilization of energy depots. We experimentally examined the role of elevated corticosterone levels in the migratory red knot by comparing foraging behavior, flight frequency, and plasma metabolites between vehicle-injected controls and birds treated with RU486, an antagonist to the genomic low-affinity glucocorticoid receptor (GR). We predicted that RU486 treatment would interfere with energy mobilization. However, we expected no effects on flight activity because recent studies suggest that glucocorticoids affect locomotion through a nongenomic receptor. Finally, because glucocorticoids exert permissive effects on food intake, we postulated that RU486 treatment in the red knot would interfere with feeding. Results were consistent with the latter prediction, suggesting that the GR participates in the promotion of hyperphagia, the intense feeding state that is characteristic of the migratory condition. RU486 treatment did not affect flight frequency, suggesting that corticosterone may support migratory activity through a receptor other than the GR. Energy metabolism (as determined through plasma metabolites) was also unaffected by RU486, possibly because energetic demands experienced by captive birds were low.     

Corticosterone and growth hormone levels in shorebirds during spring and fall migration stopover.

Large numbers of shorebirds stop over at Delaware Bay during spring migration and undergo major mass increases within a two- to three-week period. We studied plasma levels of corticosterone and growth hormone in three species of migrants that use this site, sanderlings, Calidris alba, semipalmated plovers, Charadrius semipalmatus, and semipalmated sandpipers, Calidris pusilla. Semipalmated sandpipers were also studied at a fall migration stopover in Manomet, Massachusetts. These two hormones were chosen because they modulate the physiological processes of lipogenesis/lipolysis and promote increased feeding in birds. The stress response was not suppressed in the shorebirds studied, and plasma levels of corticosterone were elevated compared to other studies. We believe that the high levels of corticosterone relate to the rapid fat deposition that takes place at this stop-over site. There was a significant negative correlation between plasma growth hormone and body mass, indicating the lipolytic effects of the growth hormone. Because the lighter birds are recent arrivals to Delaware Bay they may have elevated plasma growth because of fat breakdown during flight to this stop-over site. High levels of growth hormone may also result in protein synthesis, replenishing tissues broken down during the previous migratory bout. J. Exp. Zool. 284:645-651, 1999.     

Modulation of prolactin but not corticosterone responses to stress in relation to parental effort in a long-lived bird

We tested the hypothesis that parental effort modulates the magnitude of corticosterone and prolactin responses to stress in a long-lived bird, the Black-legged kittiwake (Rissa tridactyla). To do so, we compared corticosterone and prolactin responses to capture/restraint stress between chick-rearing birds and failed breeders (no parental effort). We predicted that (1) the increase in plasma corticosterone levels in response to stress should be lower in chick-rearing birds, (2) the decrease in plasma prolactin levels in response to stress should be lower in chick-rearing birds, and (3) as both sexes care for the chick, there should be no sex difference in the hormonal response to stress. Baseline plasma corticosterone and prolactin levels were higher in chick-rearing birds and were not influenced by body condition. Failed breeders were in better condition than chick-rearing individuals. Corticosterone response to stress was unaffected by parental effort as both chick-rearing and failed birds exhibited a robust corticosterone increase. Prolactin response to stress was however clearly influenced by parental effort: chick-rearing birds showed a modest 9% prolactin decrease whereas in failed birds prolactin concentrations fell by 41%. Body condition did not influence hormonal responses to stress. When facing stressful condition, breeding kittiwakes attenuate their prolactin response to stress while enhancing their secretion of corticosterone. Increasing corticosterone secretion triggers foraging efforts and diminishes nest attendance whereas an attenuation of prolactin response to stress maintains parental behavior. We suggest that this hormonal mechanism facilitates a flexible time-budget that has been interpreted as a buffer against environmental variability.     

Postexercise ketosis in night-migrating passerine birds

This study investigated the postexercise metabolism of six species of free-living, night-migrating passerine birds (European robin, pied flycatcher, wheatear, redstart, blackcap, and garden warbler). The birds were caught during autumn migration out of their nocturnal flight, and their metabolism changed from a fasting, highly active state to a fasting, resting state. Concentrations of six plasma metabolites of the fat, carbohydrate, and protein metabolism were measured during up to 10 h of recovery time. The metabolic changes indicated a biphasic pattern: (a) a quick first response to the reduced energy demands during the first 20 min of recovery, suggested by an increase and subsequent decrease of free fatty acid levels, and (b) subsequently, a postexercise ketosis and a reduction of lipolysis and proteolysis, suggested by high beta-hydroxy-butyrate and low free fatty acid, glycerol, triglyceride, and uric acid levels. This metabolic pattern differs from that of humans and rats, in which ketosis starts immediately postexercise or is absent in trained subjects. Since migrating birds are naturally adapted to endurance exercise, it is hypothesized that the high and long-lasting postexercise ketosis does not evoke physiological problems (such as hypoglycemia) but, by contrast, increases the ability of birds to rely on lipids, to a very high extent, during and after flight and decreases the dependence on glucose and glucogenic amino acids. Differences between species in fat stores and metabolic pattern support this hypothesis.     

Comparative activity pattern during foraging of four albatross species

The activity patterns of foraging Yellow‐nosed Diomedea chlororhynchos, Sooty Phoebetria fusca, Black‐browed D. melanophris impavida and Grey‐headed Albatross D. chrysostoma were compared using loggers recording the timing of landing and take‐offs, as well as the duration of bouts in flight or on the water, and the overall time spent in flight. The four species spent a similar proportion of their foraging time in flight (56–65%). During the day they were mostly flying (77–85% of the daylight period) whereas at night they were mainly (61–71%) sitting on the water. The amount of time spent in flight during the daytime foraging period was related to the amount of time spent sitting on the water at night. Differences between species occurred in the duration of bouts in flight and on the water as well as in the frequency of landings and in the time elapsed between successive landings. Yellow‐nosed Albatrosses were more active than the other species, with more frequent short bouts in flight and more frequent successive landings at short intervals. Sooty Albatrosses landed or took‐off less often than the other species and were more active just before dusk. Black‐browed and Grey‐headed Albatrosses were more active at night, especially the first part of the night and far from the colonies. Their trips consisted of a commuting part and a foraging part. Black‐browed Albatrosses landed more often during the foraging than the commuting part, suggesting that they were not searching when travelling. The study suggests that there is no fundamental difference between the overall activity budgets of the four species although they show distinctive diet, morphology and life history traits. The differences observed between the four species were related mainly to differences in foraging technique. Comparison with the Wandering Albatross, the only species for which data were available previously, suggest that this larger species might differ completely in foraging technique from the smaller albatrosses.     

The three-dimensional flight of red-footed boobies: adaptations to foraging in a tropical environment?

In seabirds a broad variety of morphologies, flight styles and feeding methods exist as an adaptation to optimal foraging in contrasted marine environments for a wide variety of prey types. Because of the low productivity of tropical waters it is expected that specific flight and foraging techniques have been selected there, but very few data are available. By using five different types of high-precision miniaturized logger (global positioning systems, accelerometers, time depth recorders, activity recorders, altimeters) we studied the way a seabird is foraging over tropical waters. Red-footed boobies are foraging in the day, never foraging at night, probably as a result of predation risks. They make extensive use of wind conditions, flying preferentially with crosswinds at median speed of 38 km h(-1), reaching highest speeds with tail winds. They spent 66% of the foraging trip in flight, using a flap-glide flight, and gliding 68% of the flight. Travelling at low costs was regularly interrupted by extremely active foraging periods where birds are very frequently touching water for landing, plunge diving or surface diving (30 landings h(-1)). Dives were shallow (maximum 2.4 m) but frequent (4.5 dives h(-1)), most being plunge dives. While chasing for very mobile prey like flying fishes, boobies have adopted a very active and specific hunting behaviour, but the use of wind allows them to reduce travelling cost by their extensive use of gliding. During the foraging and travelling phases birds climb regularly to altitudes of 20-50 m to spot prey or congeners. During the final phase of the flight, they climb to high altitudes, up to 500 m, probably to avoid attacks by frigatebirds along the coasts. This study demonstrates the use by boobies of a series of very specific flight and activity patterns that have probably been selected as adaptations to the conditions of tropical waters.     

Oxidative Stress in Endurance Flight: An Unconsidered Factor in Bird Migration

Migrating birds perform extraordinary endurance flights, up to 200 h non-stop, at a very high metabolic rate and while fasting. Such an intense and prolonged physical activity is normally associated with an increased production of reactive oxygen and nitrogen species (RONS) and thus increased risk of oxidative stress. However, up to now it was unknown whether endurance flight evokes oxidative stress. We measured a marker of oxidative damage (protein carbonyls, PCs) and a marker of enzymatic antioxidant capacity (glutathione peroxidase, GPx) in the European robin (Erithacus rubecula), a nocturnal migrant, on its way to the non-breeding grounds. Both markers were significantly higher in European robins caught out of their nocturnal flight than in conspecifics caught during the day while resting. Independently of time of day, both markers showed higher concentrations in individuals with reduced flight muscles. Adults had higher GPx concentrations than first-year birds on their first migration. These results show for the first time that free-flying migrants experience oxidative stress during endurance flight and up-regulate one component of antioxidant capacity. We discuss that avoiding oxidative stress may be an overlooked factor shaping bird migration strategies, e.g. by disfavouring long non-stop flights and an extensive catabolism of the flight muscles.     

Regulation of protein breakdown and adrenocortical response to stress in birds during migratory flight

During long-term fasting at rest, protein utilization is maintained at low levels until it increases at a threshold adiposity. This study examines 1) whether such a shift in energy substrate use also occurs during endurance exercise while fasting, 2) the role of corticosterone, and 3) the adrenocortical response to an acute stressor. Ten species of migrating birds caught after an endurance flight over at least 500 km were examined. Plasma uric acid and corticosterone levels were low in birds with fat stores >5% of body mass and high in birds with smaller fat stores. Corticosterone levels were very high in birds with no visible fat stores and emaciated breast muscles. Corticosterone levels increased with handling time only in birds with large fat stores. These findings suggest that 1) migrating birds with appreciable fat stores are not stressed by endurance flight, 2) a metabolic shift (increased protein breakdown), regulated by an endocrine shift (medium corticosterone levels), occurs at a threshold adiposity, as observed in birds at rest, 3) adrenocortical response to an acute stressor is inhibited after this shift, and 4) an adrenocortical response typical for an emergency situation (high corticosterone levels) is only reached when muscle protein is dangerously low.     

Baseline and stress-induced levels of corticosterone during different life cycle substages in a shorebird on the high arctic breeding grounds

After a migratory flight of several thousand kilometers to their high arctic breeding grounds, red knots (Calidris canutus islandica, Scolopacidae) showed high baseline concentrations of plasma corticosterone (58 ng/mL). Such high baseline corticosterone levels may be conditional for the right behavioral and metabolic adjustments to environmental and social stresses that shorebirds experience on arrival in an unpredictable tundra breeding environment. Despite the high baseline levels of corticosterone, red knots still showed a marked stress response during the postarrival period, with corticosterone concentrations increasing significantly during a 60-min period of confinement. Baseline levels of corticosterone declined as the breeding season progressed. Red knots with brood patches, that is, birds that had completed egg laying and commenced incubation, had a reduced adrenocortical response to the stress of confinement compared with red knots with no, or with half-developed, brood patches. This is consistent with the idea that birds breeding in extreme environments with short breeding seasons may exhibit a decreased adrenocortical response to stressful events to prevent high corticosterone concentrations from inducing interruptions of reproductive behavior.     

Energetics and metabolite profiles during early flight in American robins (Turdus Migratorius)

Although birds use fat as the primary fuel for migratory flights, carbohydrate and protein catabolism could be significant in the early stages of flight while pathways of fatty acid transport and oxidation are induced. The fuel mixture of long distance migrant birds can also be affected by the rate of water loss, where birds catabolize more protein to increase endogenous water production under dehydrating flight conditions. Despite many studies investigating flight metabolism, few have focused on the metabolic response to flight during the switchover to fat catabolism in migrants, and none have examined the effect of ambient conditions on fuel selection during early flight. We investigated the effect of water loss on the metabolic response to short duration flight in the American robin (Turdus migratorius). Birds were flown in a climatic wind tunnel and changes in body composition and plasma metabolites were measured. As flight duration increased, there was a gradual switchover from carbohydrate and protein catabolism to fat catabolism. Plasma metabolite profiles indicate that the mobilization of fat occurred within 20 min of initiating flight. Plasma glucose decreased and uric acid increased with flight duration. Ambient humidity did not affect fuel mixture. Thus, it seems that the utilization of fat may be delayed as migrants initiate flight. Short-hop migrants may exploit high rates of endogenous water production resulting from carbohydrate and protein catabolism early in flight to offset high water loss associated with low humidity. Rapid catabolism of lean body components at the start of a flight also reduces mass quickly, and may reduce energy costs.