Spatio-temporal variation in territory quality and oxidative status: a natural experiment in the Seychelles warbler (Acrocephalus sechellensis)

1. Fluctuations in the quality of the habitat in which an animal lives can have major consequences for its behaviour and physiological state. In poor-quality habitat with low food availability, metabolically intensive foraging activity is likely to result in increased generation of reactive oxygen species, while scarcity of food can lead to a weakening of exogenously derived antioxidant defences. The consequent oxidant/antioxidant imbalance may lead to elevated oxidative stress. 2. Although the link between food availability and oxidative stress has been studied in the laboratory, very little is known about this relationship in the wild. Here, we investigate the association between territory quality (measured through food availability) and oxidative stress in the Seychelles warbler (Acrocephalus sechellensis). 3. Seychelles warblers are insectivorous birds that inhabit a fixed feeding territory year round. Individuals experience profound and rapid local fluctuations in territory quality within these territories, owing to changing patterns of vegetation defoliation resulting from seasonal changes in prevailing wind direction and wind-borne salt spray. 4. As expected, oxidant generation (measured as reactive oxygen metabolites; ROMs) was higher when territory quality was low, but there was no correlation between territory quality and antioxidant capacity (OXY). The negative correlation between territory quality and ROMs was significant between individuals and approached significance within individuals, indicating that the pattern resulted from individual responses to environmental variation. 5. ROMs and OXY levels within individuals were positively correlated, but the relationship between territory quality and ROMs persisted after including OXY as a covariate, implying that oxidative stress occurs in low territory quality conditions. 6. Our results indicate that the oxidative stress balance of an individual is sensitive to relatively short-term changes in territory quality, which may have consequences for the birds' fitness.     

Plasma reactive oxygen metabolites and non-enzymatic antioxidant capacity are not affected by an acute increase of metabolic rate in zebra finches

Understanding the sources of variation in oxidative stress level is a challenging issue due to the implications of oxidative stress for late age diseases, longevity and life-history trade-offs. Reactive oxygen species that cause oxidative stress are mostly a by-product of energy metabolism and it is therefore often assumed that oxidative stress is proportional to energy consumption. In mammals, an increased metabolic rate induced by cold exposure generally increases oxidative stress. However, compared to mammals, birds generate fewer free radicals per ATP produced and hence it is not obvious that, in birds, a cold-induced increase of metabolic rate increase oxidative stress. We tested whether cold-induced increase in metabolic rate increased oxidative stress in zebra finches by exposing individuals to cold and warm overnight temperatures. We registered metabolic rate and plasma levels of non-enzymatic antioxidants and reactive oxygen metabolites (ROMs), a measure of oxidative damage. Metabolic rate was on average 88 % higher in cold compared to warm temperature, with females being stronger affected than males. However, temperature had no effect on plasma antioxidants or our measure of oxidative damage. Middle-age birds had higher levels of plasma antioxidants than younger and older birds, but age was unrelated to ROMs. Birds showed repeatability of plasma ROMs across temperatures but not of non-enzymatic antioxidants. In contrast to similar studies in mammals, our results do not show evidence of increased oxidative stress in plasma after an acute cold-induced increase of metabolic rate but research in more bird species is needed to assess the generality of this pattern.     

Aggressive and non-aggressive personalities differ in oxidative status in selected lines of mice (Mus musculus)

Mice selected for aggression and coping (long attack latency (LAL), reactive coping strategy; short attack latency (SAL), pro-active coping strategy) are a useful model for studying the physiological background of animal personalities. These mice also show a differential stress responsiveness, especially in terms of hypothalamic-pituitary-adrenal axis reactivity, to various challenges. Since the stress response can increase the production of reactive oxygen species, we predicted that the basic oxidative status of the lines could differ. We found that LAL showed higher serum antioxidant capacity (OXY) than SAL, while no differences emerged for reactive oxygen metabolites (ROMs) or the balance between ROMs and OXY, reflecting oxidative stress. Moreover, the lines showed inverse relationships between ROMs or OXY and body mass corrected for age. The results indicate that variation in oxidative status is heritable and linked to personality. This suggests that different animal personalities may be accompanied by differences in oxidative status, which may predict differences in longevity.     

Migrating songbirds on stopover prepare for,and recover from,oxidative challenges posed by long‐distance flight

Managing oxidative stress is an important physiological function for all aerobic organisms, particularly during periods of prolonged high metabolic activity, such as long‐distance migration across ecological barriers. However, no previous study has investigated the oxidative status of birds at different stages of migration and whether that oxidative status depends on the condition of the birds. In this study, we compared (1) energy stores and circulating oxidative status measures in (a) two species of Neotropical migrants with differing migration strategies that were sampled at an autumn stopover site before an ecological barrier; and (b) a species of trans‐Saharan migrant sampled at a spring stopover site after crossing an ecological barrier; and (2) circulating oxidative measures and indicators of fat metabolism in a trans‐Saharan migrant after stopovers of varying duration (0–8 nights), based on recapture records. We found fat stores to be positively correlated with circulating antioxidant capacity in Blackpoll Warblers and Red‐eyed Vireos preparing for fall migration on Block Island, USA, but uncorrelated in Garden Warblers on the island of Ponza, Italy, after a spring crossing of the Sahara Desert and Mediterranean Sea. In all circumstances, fat stores were positively correlated with circulating lipid oxidation levels. Among Garden Warblers on the island of Ponza, fat anabolism increased with stopover duration while oxidative damage levels decreased. Our study provides evidence that birds build antioxidant capacity as they build fat stores at stopover sites before long flights, but does not support the idea that antioxidant stores remain elevated in birds with high fuel levels after an ecological barrier. Our results further suggest that lipid oxidation may be an inescapable hazard of using fats as the primary fuel for flight. Yet, we also show that birds on stopover are capable of recovering from the oxidative damage they have accrued during migration, as lipid oxidation levels decrease with time on stopover. Thus, the physiological strategy of migrating songbirds may be to build prophylactic antioxidant capacity in concert with fuel stores at stopover sites before a long‐distance flight, and then repair oxidative damage while refueling at stopover sites after long‐distance flight.     

Seasonal variation in the regulation of redox state and some biotransformation enzyme activities in the barn swallow (Hirundo rustica L.)

Little is known of the normal seasonal variation in redox state and biotransformation activities in birds. In long-distance migratory birds, in particular, seasonal changes could be expected to occur because of the demands of migration and reproduction. In this study, we measured several redox parameters in the barn swallow (Hirundo rustica L.) during the annual cycle. We captured the wintering barn swallows before spring migration in South Africa, and we captured the barn swallows that arrived in spring, bred in summer, and migrated in autumn in Finland. The redox status and biotransformation activities of barn swallows varied seasonally. Wintering birds in South Africa had high biotransformation activities and appeared to experience oxidative stress, whereas in spring and summer, they showed relatively low redox (superoxide dismutase [SOD], catalase [CAT], and glutathione reductase [GR]) and biotransformation enzyme activities. Autumn birds had very low biotransformation enzyme activities and low indication of oxidative stress but high activity of some redox enzymes (GR and glucose 6-phosphate dehydrogenase [G6PDH]). High activities of some redox enzymes (SOD, GR, and G6PDH) seem to be related to migration, whereas low activities of some redox enzymes (SOD, CAT, and GR) may be associated with breeding. Barn swallows in South Africa may experience pollution-related oxidative stress, which may hamper interpretation of normal seasonal variation in redox parameters.     

Effects of carotenoid supplementation on colour expression,oxidative stress and body mass in rehabilitated captive adult kestrels (<Emphasis Type="Italic">Falco tinnunculus</Emphasis>)

Carotenoids are considered a limited resource for animals because they are not synthesised by the body. Birds use carotenoids, mainly xanthophylls, for physiological functions, such as anti-oxidant activity, and for colour expression; hence, they need to shunt carotenoids between competitive demands. Recent studies suggest that the anti-oxidant role of xanthophylls might not be as important as previously thought and that at high concentrations they may, in fact, acquire pro-oxidant properties. In this work, we studied the effects of a moderate xanthophyll supplementation (115 mg of carotenoids/kg diet/day; 4 weeks) on serum carotenoids, serum concentration of reactive oxygen metabolites (ROMs), serum anti-oxidant capacity (OXY), the degree of oxidative stress (OS; ROMs/OXY × 1,000), body mass, and skin colour, in rehabilitated captive adult Eurasian kestrels (Falco tinnunculus). The supplementation caused increased levels of serum carotenoids (∼90%), ROMs (∼82%), OS (∼115%) and an immediate loss of body mass (∼6.2%), but it did not affect OXY and tarsi skin hue. The red (∼16%) and yellow (∼15%) colorimetric components were increased after the first week of supplementation and the effect persisted during the rest of the experiment. Two months after the end of supplementation, serum carotenoids, OS and ROMs returned to baseline levels, however the body mass did not. Our findings suggest that, above a certain physiological threshold, carotenoids can cause detrimental effects. This is relevant for the trade-off between expression of sexual signals and the costs of maintaining/producing them.     

Effects of T-cell-mediated immune response on avian oxidative stress

We evaluated the oxidative cost paid by birds when coping with an immune challenge. We used the phytohaemagglutinin skin test (PHA) to assess the effects of the T-cell-mediated immune response on the concentration of reactive oxygen metabolites (ROMs), total antioxidant barrier (OXY) and total serum carotenoid concentration in wild nestlings of the Eurasian kestrel (Falco tinnunculus). Immunostimulation caused increased levels of ROMs, decreased OXY and increased circulating levels of carotenoids. These results suggest that an immune challenge can increase avian oxidative stress, and that carotenoids were remobilised from other tissues likely because their circulating levels were not sufficiently high to sustain an effective immune response.     

Interaction between sexual steroids and immune response in affecting oxidative status of birds

One hypothesis explaining the honesty of secondary sexual traits regulated by testosterone (T) is that T can impair the balance between pro-oxidant compounds and antioxidant defences, favouring a status of oxidative stress that only good quality individuals can sustain (oxidative handicap hypothesis). In the present study, we evaluated for the first time the effects of sexual steroids, T and its metabolites 5-α-dihydrotestosterone (DHT) and estradiol (E2) on oxidative damage and plasma non-enzymatic antioxidant capacity, while birds are faced by an oxidative challenge induced by an immune stimulation with sheep red blood cells. We used male and female diamond doves Geopelia cuneata, a species that shows an orange-red periorbital ring, whose size and color are strongly affected by androgens, but not by estrogens. Immunization increased oxidative damage in all groups, regardless of hormone treatment. It also decreased anti-oxidant capacity in all groups, except for testosterone treated birds. The ratio of oxidative damage over anti-oxidant capacity (oxidative stress) was increased in both immunological challenged controls and E2 birds, while challenged birds treated with androgens did not differ from non-challenged birds. The response of males and females to our treatments never differed. Our results undermine the idea that T can induce honest signalling through a pro-oxidant activity.     

Response to oxidative stress as a welfare parameter in swine

In pigs, the genetic selection for lean, large muscle blocks and fast growth has been linked to an increased prevalence of metabolic diseases such as porcine stress syndrome and mulberry heart disease. These diseases are associated with cardiovascular inadequacy, which may lead to oxidative stress. In the present study, reactive oxygen metabolites (ROMs) and the anti-oxidant power (OXY) in sera of different swine groups were investigated. The following groups were selected (each around 80 kg body weight): wild boars (WB), Cinta Senese (CS), and Landrace x Large White (LxLW), the latter as both specific pathogen-free (SPF) and intensively farmed animals. In addition, a group of LxLW agonic sows (AS) was also investigated; this group is known to be under oxidative stress. Two colorimetric micro-methods were used to measure ROMs and OXY; ROMs were expressed as mM H(2)O(2) and OXY as microM HOCl neutralised. Between groups, average ROM and OXY values were found to be significantly different by one-way ANOVA (P < 0.001). ROM levels were lower in WB (13.41 +/- 1.85) and CS (19.27 +/- 1.68), and highest in LxLW (42.00 +/- 1.36). OXY values ranged from 260.10 +/- 22.13 (WB) to 396.90 +/- 9.83 (LxLW). Only one swine group (the CS group) showed a significant, positive correlation between ROM and OXY values. The AS group even showed a negative correlation between ROM and OXY values. These results imply satisfactory environmental coping occurred only within the CS group. Results are discussed in the light of animal welfare legislation, food safety and consumers' protection.     

Assessing the cost of helping: the roles of body condition and oxidative balance in the Seychelles warbler (Acrocephalus sechellensis)

In cooperatively breeding species, helping close relatives may provide important fitness benefits. However, helping can be energetically expensive and may result in increased generation of reactive oxygen species. Consequently, an oxidant/antioxidant imbalance can lead to higher oxidative stress susceptibility. Given the potential costs of helping, it may be that only individuals with a sufficiently good body condition and/or stable oxidative balance can afford to help. Knowledge about relationships between social status and oxidative balance in cooperatively breeding systems is still limited. Studying these relationships is important for understanding the costs of helping and physiological pressures of reproduction. Here we evaluate the relationship between helping behaviour, body condition and oxidative balance in a wild population of the cooperatively breeding Seychelles warbler (Acrocephalus sechellensis). In this species, some subordinate individuals help dominant birds with the rearing of young, while others refrain from any assistance. We assessed body condition and oxidative parameters of birds of different social status caught during different breeding stages. We found that, prior to breeding, female subordinates that did not subsequently help (non-helpers) had significantly lower body condition and higher ROMs (reactive oxygen metabolites) than helpers and dominants. During the later stages of breeding, body condition was low in dominants and helpers, but high in non-helpers. Differences in oxidative balance between individuals of different social status were found only during nest care: Dominant males occupied with guarding behaviours tended to have relatively high oxidative stress susceptibility. Furthermore, dominant and helper females showed elevated antioxidant capacity (measured as OXY) in the weeks just prior to egg-laying, possibly representing a change in their reproductive physiology. The results imply that an individuals' oxidative balance may be influenced by factors related to reproduction, which can differ with sex and--within cooperative breeding systems--social status.     

Environmental and genetic components of oxidative stress in wild kestrel nestlings (Falco tinnunculus)

In this study, we estimated the environmental and genetic components of two variables related to avian oxidative stress using wild nestlings of the Eurasian kestrel (Falco tinnunculus). The study was carried out during two breeding seasons. In the first season, we assessed the between- and within-nest resemblance in serum reactive oxygen metabolites (ROMs) and total serum antioxidant barrier (OXY). In the second season, we carried out a cross-fostering experiment to determine the importance of environmental and genetic factors on ROMs and OXY. The 23.5% of ROMs variance was explained by the nest of origin, indicating a main genetic component. In contrast, the 52.8% of OXY variance was explained by the nest of rearing, indicating that this variable was more influenced by environmental components. These findings suggest that variations in ROMs and OXY could reflect, respectively, the expression of different genetic polymorphisms and differences in dietary uptake of antioxidants.     

Oxidative stress,circulating antioxidants,and dietary preferences in songbirds

Oxidative stress is an unavoidable consequence of metabolism and increases during intensive exercise. This is especially problematic for migratory birds that metabolize fat to fuel long-distance flight. Birds can mitigate damage by increasing endogenous antioxidants (e.g. uric acid) or by consuming dietary antioxidants (e.g. tocopherol). During flight, birds may increase protein catabolism of lean tissue which may increase circulating uric acid and many birds also consume an antioxidant-rich frugivorous diet during autumn migration. We evaluated three related hypotheses in a migratory passerine: (1) protein consumption is positively related to circulating antioxidants, (2) a dietary oxidative stressor [i.e. polyunsaturated fatty acid (PUFA)] influences antioxidant capacity and oxidative damage, and (3) oxidative stress influences dietary antioxidant preferences. White-throated Sparrows (Zonotrichia albicollis) consuming a high protein diet increased circulating uric acid; however, uric acid, antioxidant capacity, and oxidative stress did not differ between birds consuming a high PUFA versus a low PUFA diet, despite increased oxidative damage in high PUFA birds. Birds did not prefer antioxidant-rich diets even when fed high PUFA, low protein. We conclude that White-throated Sparrows successfully mitigated oxidative damage associated with a high PUFA diet and mounted an endogenous antioxidant response independent of uric acid, other circulating antioxidants, and dietary antioxidants.     

Coccidian infection causes oxidative damage in greenfinches

The main tenet of immunoecology is that individual variation in immune responsiveness is caused by the costs of immune responses to the hosts. Oxidative damage resulting from the excessive production of reactive oxygen species during immune response is hypothesized to form one of such costs. We tested this hypothesis in experimental coccidian infection model in greenfinches Carduelis chloris. Administration of isosporan coccidians to experimental birds did not affect indices of antioxidant protection (TAC and OXY), plasma triglyceride and carotenoid levels or body mass, indicating that pathological consequences of infection were generally mild. Infected birds had on average 8% higher levels of plasma malondialdehyde (MDA, a toxic end-product of lipid peroxidation) than un-infected birds. The birds that had highest MDA levels subsequent to experimental infection experienced the highest decrease in infection intensity. This observation is consistent with the idea that oxidative stress is a causative agent in the control of coccidiosis and supports the concept of oxidative costs of immune responses and parasite resistance. The finding that oxidative damage accompanies even the mild infection with a common parasite highlights the relevance of oxidative stress biology for the immunoecological research.     

Individual consistency and covariation of measures of oxidative status in greenfinches

Oxidative stress results from a mismatch between production of reactive oxygen species (ROS) and the organism's capacity to mitigate their damaging effects by building up sufficient antioxidant protection and/or repair mechanisms. Because ROS production is a universal consequence of cellular metabolism and immune responses, evolutionary animal ecologists have become increasingly interested in involvement of oxidative stress as a proximate mechanism responsible for the emergence of trade-offs related to the evolution of life-history and signal traits. Among the most practical problems pertinent to ecological research on oxidative stress is finding a combination of biomarkers of oxidative status that can be applied to typical wild animal models such as small birds, mammals, and reptiles. This study describes covariation and individual consistency of eight parameters of oxidative status in a small passerine bird, wild-caught captive greenfinch (Carduelis chloris). We measured two markers of plasma antioxidant potential--total antioxidant capacity (TAC) and oxygen radical absorbance (OXY)--and concentrations of one lipophilic (carotenoids) and two hydrophilic (uric acid and ascorbate) antioxidants in plasma. We also measured total glutathione (GSH) concentration and superoxide dismutase (SOD) activity in erythrocytes. Oxidative damage was assessed on the basis of plasma malondialdehyde (MDA) levels, measured by high-performance liquid chromatography. Plasma carotenoids, TAC, and erythrocyte GSH showed significant individual consistency over an 8-d period, indicating that those variables reflected more persistent differences between individuals than plasma OXY, MDA, and uric acid. We did not detect any strong or moderate correlations between the studied parameters, which suggests that all of these biomarkers contain potentially unique information. Injection of a synthetic mimetic of SOD and catalase--EUK-134--did not affect any of the parameters of oxidative status. Capability of phagocytes to produce oxidative burst was not associated with MDA, indicating that under our experimental conditions, ROS production by phagocytes was not a strong determinant of oxidative damage. Altogether these findings suggest that attempts to characterize oxidative balance should use a wide range of biomarkers, and further studies of oxidative status in wild animals may benefit from the experimental induction of oxidative stress.     

Differential Regulation of Adipokines May Influence Migratory Behavior in the White-Throated Sparrow (Zonotrichia albicollis)

White-throated sparrows increase fat deposits during pre-migratory periods and rely on these fat stores to fuel migration. Adipose tissue produces hormones and signaling factors in a rhythmic fashion and may be controlled by a clock in adipose tissue or driven by a master clock in the brain. The master clock may convey photoperiodic information from the environment to adipose tissue to facilitate pre-migratory fattening, and adipose tissue may, in turn, release adipokines to indicate the extent of fat energy stores. Here, we present evidence that a change in signal from the adipokines adiponectin and visfatin may act to indicate body condition, thereby influencing an individual''s decision to commence migratory flight, or to delay until adequate fat stores are acquired. We quantified plasma adiponectin and visfatin levels across the day in captive birds held under constant photoperiod. The circadian profiles of plasma adiponectin in non-migrating birds were approximately inverse the profiles from migrating birds. Adiponectin levels were positively correlated to body fat, and body fat was inversely related to the appearance of nocturnal migratory restlessness. Visfatin levels were constant across the day and did not correlate with fat deposits; however, a reduction in plasma visfatin concentration occurred during the migratory period. The data suggest that a significant change in the biological control of adipokine expression exists between the two migratory conditions and we propose a role for adiponectin, visfatin and adipose clocks in the regulation of migratory behaviors.     

Heritability of arrival date in a migratory bird

The behaviour of long-distance migratory birds is assumed to partly be under the influence of genes, as demonstrated by selection experiments. Furthermore, competition for early arrival among males may lead to condition-dependent migration associated with fitness benefits of early arrival achieved by individuals in prime condition. Here I present field data on the repeatability and the heritability of arrival date in a trans-equatorial migratory bird, the barn swallow Hirundo rustica, and I test for a genetic correlation between arrival date and the expression of a condition-dependent secondary sexual character. The repeatability was statistically significant and the heritability of arrival date was estimated to be 0.54 (s.e. = 0.15). There was no significant evidence of this estimate being inflated by environmental or maternal condition during rearing. Arrival date and migration are condition dependent in the barn swallow, with males with the most exaggerated secondary sexual characters also arriving the earliest. There was a significant genetic correlation between arrival date and tail length in male barn swallows, providing indirect evidence for a genetic basis of this condition dependence. Given the high level of heritability, arrival date could readily respond to selection caused by environmental change.     

A Continent-Wide Migratory Divide in North American Breeding Barn Swallows (Hirundo rustica)

Populations of most North American aerial insectivores have undergone steep population declines over the past 40 years but the relative importance of factors operating on breeding, wintering, or stopover sites remains unknown. We used archival light-level geolocators to track the phenology, movements and winter locations of barn swallows (Hirdundo rustica; n = 27) from populations across North America to determine their migratory connectivity. We identified an east-west continental migratory divide for barn swallows with birds from western regions (Washington State, USA (n = 8) and Saskatchewan, Canada (n = 5)) traveling shorter distances to wintering areas ranging from Oregon to northern Colombia than eastern populations (Ontario (n = 3) and New Brunswick (n = 10), Canada) which wintered in South America south of the Amazon basin. A single swallow from a stable population in Alabama shared a similar migration route to eastern barn swallows but wintered farther north in northeast Brazil indicating a potential leap frog pattern migratory among eastern birds. Six of 9 (67%) birds from the two eastern populations and Alabama underwent a loop migration west of fall migration routes including around the Gulf of Mexico travelling a mean of 2,224 km and 722 km longer on spring migration, respectively. Longer migration distances, including the requirement to cross the Caribbean Sea and Gulf of Mexico and subsequent shorter sedentary wintering periods, may exacerbate declines for populations breeding in northeastern North America.     

The effects of radiation on sperm swimming behavior depend on plasma oxidative status in the barn swallow (Hirundo rustica)

Sperm are highly susceptible to reactive oxygen species (ROS) that can damage sperm DNA and structure, resulting in reduced fertilizing capacity. Exposure to radioactive contamination can also impair sperm swimming behavior and fertilizing ability, both through a reduction of sperm DNA integrity and via an increased generation of reactive oxygen species (ROS). However, the relationship between individual oxidative status and sperm swimming behavior has never been investigated in any wild population of animals exposed to radioactive contamination. We studied the motility of sperm collected from barn swallows, Hirundo rustica, breeding under different levels of radioactive contamination following the Chernobyl accident in 1986, in relation to individual oxidative status. We tested the hypothesis that the degree of impairment of sperm swimming behavior by radioactive contamination depended on plasma antioxidant capacity, the level of reactive oxygen metabolites (ROMs) and oxidative stress (sensu Costantini et al. 2006), a better oxidative status being associated with higher sperm motility. Sperm behavior parameters were subjected to principal component (PC) analysis, which extracted four PCs explaining 86% of the variance in sperm motility. PC2, representing sperm with high track velocity and ample lateral head displacement, was significantly predicted by the interaction between radiation level and either oxidative damage or oxidative stress. Contrary to our predictions, the highest values of PC2 were associated with relatively high radiation levels, particularly for high levels of either ROMs or oxidative stress. In addition, there was a tendency for values of PC3 (representing the percent of motile sperm) and PC4 (representing slow sperm with high beat cross frequency) to depend on the interaction between radiation level and total plasma antioxidant protection. Our results confirm the importance of oxidative status in determining the genetic and physiological outcome of exposure to radioactive contamination, complementing previous studies relating sperm abnormality to circulating levels of specific antioxidants. Our results also complement previous evidence that oxidative damage of sperm was negatively related to sperm motility, thus indicating a possible trade-off in quenching pro-oxidant compounds in the plasma and the seminal fluid.     

Corticosterone secretion patterns prior to spring and autumn migration differ in free-living barn swallows (Hirundo rustica L.)

Recent studies of long-distance migratory birds show that behavioural and physiological changes associated with predictable or unpredictable challenges during the annual cycle are distinctively regulated by hormones. Corticosterone is the primary energy regulating hormone in birds. Corticosterone levels are elevated during stresses but they are also modulated seasonally according to environmental conditions and life-history demands. We measured the baseline and stress-induced levels of corticosterone in the barn swallow (Hirundo rustica L.) just before spring and autumn migrations in South Africa and Finland, respectively. Barn swallows completing their pre-breeding moult had low body condition (residual body mass) and high baseline corticosterone levels in the wintering grounds. In contrast, baseline corticosterone levels in Finland were low and not related to residual mass. These data contradict the first prediction of the migration modulation hypothesis (MMH) by showing no association with baseline corticosterone levels and pre-migratory fuelling. Yet, the adrenocortical response to the capture and handling stress was notably blunted in South Africa compared to a strong response in Finland. Further, individuals that had started fuelling in Finland showed a reduced response to the handling stress. Taken together, elevated baseline corticosterone levels and high residual mass may blunt the adrenocortical response in long-distance migrants and aerial feeders such as the barn swallow. This observation lends support to the second prediction of the MMH.     

Long-term trends in first arrival and first egg laying dates of some migrant and resident bird species in northern Italy

Climate change is affecting the phenology of seasonal events in Europe and the Northern Hemisphere, as shown by several studies of birds’ timing of migration and reproduction. Here, we analyse the long-term (1982–2006) trends of first arrival dates of four long-distance migratory birds [swift (Apus apus), nightingale (Luscinia megarhynchos), barn swallow (Hirundo rustica), and house martin (Delichon urbicum)] and first egg laying dates of two migrant (swift, barn swallow) and two resident species [starling (Sturnus vulgaris), Italian sparrow (Passer italiae)] at a study site in northern Italy. We also addressed the effects of local weather (temperature and precipitation) and a climate index (the North Atlantic Oscillation, NAO) on the interannual variability of phenological events. We found that the swift and the barn swallow significantly advanced both arrival and laying dates, whereas all other species did not show any significant temporal trend in either arrival or laying date. The earlier arrival of swifts was explained by increasing local temperatures in April, whereas this was not the case for arrival dates of swallows and first egg laying dates of both species. In addition, arrival dates of house martins were earlier following high NAO winters, while nightingale arrival was earlier when local spring rainfall was greater. Finally, Italian sparrow onset of reproduction was anticipated by greater spring rainfall, but delayed by high spring NAO anomalies, and swift’s onset of reproduction was anticipated by abundant rainfall prior to reproduction. There were no significant temporal trends in the interval between onset of laying and arrival in either the swift or the barn swallow. Our findings therefore indicate that birds may show idiosyncratic responses to climate variability at different spatial scales, though some species may be adjusting their calendar to rapidly changing climatic conditions.