Sveinn Are Hanssen

Cost of reproduction in a long-lived bird: incubation effort reduces immune function and future reproduction

Life-history theory predicts that increased current reproductive effort should lead to a fitness cost. This cost of reproduction may be observed as reduced survival or future reproduction, and may be caused by temporal suppression of immune function in stressed or hard-working individuals. In birds, consideration of the costs of incubating eggs has largely been neglected in favour of the costs of brood rearing. We manipulated incubation demand in two breeding seasons (2000 and 2001) in female common eiders (Somateria mollissima) by creating clutches of three and six eggs (natural range 3–6 eggs). The common eider is a long-lived sea-duck where females do not eat during the incubation period. Mass loss increased and immune function (lymphocyte levels and specific antibody response to the non-pathogenic antigens diphtheria and tetanus toxoid) was reduced in females incubating large clutches. The increased incubation effort among females assigned to large incubation demand did not lead to adverse effects on current reproduction or return rate in the next breeding season. However, large incubation demand resulted in long-term fitness costs through reduced fecundity the year after manipulation. Our data show that in eiders, a long-lived species, the cost of high incubation demand is paid in the currency of reduced future fecundity, possibly mediated by reduced immune function.

Costs of immunity: immune responsiveness reduces survival in a vertebrate

Immune defences are undoubtedly of great benefit to the host, reducing the impact of infectious organisms. However, mounting immune responses also entails costs, which may be measured by inducing immune responses against artificial infections. We injected common eider (Somateria mollissima) females with three different non–pathogenic antigens, sheep red blood cells (SRBC), diphtheria toxoid and tetanus toxoid, early in their incubation period. In the group of females that mounted a humoral immune response against SRBC, the return rate was only 27%, whereas the group of females that did not mount a response against SRBC had a return rate of 72–. Moreover, responding against diphtheria toxoid when also responding against SRBC led to a further reduction in return rate. These results are repeatable, as the same effect occurred independently in two study years. The severely reduced return rate of females producing antibodies against SRBC and diphtheria toxoid implies that these birds experienced considerably impaired long–term survival. This study thus documents severe costs of mounting humoral immune responses in a vertebrate. Such costs may explain why many organisms suppress immunity when under stress or when malnourished, and why infections may sometimes be tolerated without eliciting immune responses.

Reduced immunocompetence and cost of reproduction in common eiders

Immunocompetence may be especially important in long-lived species where infectious organisms may have detrimental effects upon future reproductive value of hosts. The resource demand for immunocompetence may reduce resource availability for reproduction and a trade-off between these traits has therefore been proposed. Capital breeders, such as the common eider (Somateria mollissima), rely upon accumulated body reserves during reproduction. Eiders lose more than 40% of pre-breeding body mass during egglaying and incubation and many females abandon their ducklings to other females after hatching. Results from our observational study show that levels of leukocytes (i.e., lymphocytes, heterophils and heterophil/lymphocyte ratio) are not related to body mass early in the incubation period. However, eider females with low initial body mass showed signs of immunosuppression (i.e., decreased late levels of lymphocytes) and increased response towards stressors (i.e., increased heterophil/lymphocyte ratio) later in the incubation period. Moreover, females with low lymphocyte levels more frequently abandoned their brood, and females abandoning young had an increased return rate to the next breeding season. However, among brood abandoning females return rate was lower for the females with low lymphocyte levels. These results suggest that immunosuppression, as indicated by low lymphocyte levels, is a reproductive cost that may be partly compensated for by abandoning young.

Incubation start and egg size in relation to body reserves in the common eider

Abstract. Avian incubation is often initiated before all eggs are laid. In altricial birds this has been proposed to facilitate brood reduction through asynchronous hatching. However, in precocial birds eggs normally hatch synchronously even if incubation has started before all eggs are laid. Patterns of incubation start may be the adaptive trait selected for both in altricial and precocial species. Several hypotheses have been proposed to explain the timing of incubation start in birds. Decreasing egg-size after incubation start may be adaptively related to an early incubation start, either to ensure synchronous hatching or to decrease fitness cost of late hatched eggs. We have measured individual body condition, egg size and start of incubation in common eider Somateria mollissima, a precocial sea-duck which does not feed during the incubation period. Females in poor body condition start to incubate earlier in the laying sequence than those in good body condition. Furthermore females in poor body condition lay smaller final eggs than females in good body condition. The laying of smaller eggs late in the sequence is therefore probably related to energetic or nutritional state. We propose that females in poor body condition start to incubate early to shorten the nest period in order to reduce their mass loss, but at the cost of reduced size and growth of the ducklings from the eggs laid after incubation start. Females in good body condition on the other hand postpone incubation start at the cost of a longer incubation period and a higher mass loss to the benefit of synchronized hatching and a higher survival of ducklings.

Differential investment and costs during avian incubation determined by individual quality: an experimental study of the common eider (Somateria mollissima)

Individuals of different quality may have different investment strategies, shaping responses to experimental manipulations, thereby rendering the detection of such patterns difficult. However, previous clutch–size manipulation studies have infrequently incorporated individual differences in quality. To examine costs of incubation and reproductive investment in relation to changes in clutch size, we enlarged and reduced natural clutch sizes of four and five eggs by one egg early in the incubation period in female common eiders (Somateria mollissima), a sea duck with an anorectic incubation period. Females that had produced four eggs (lower quality) responded to clutch reductions by deserting the nest more frequently but did not increase incubation effort in response to clutch enlargement, at the cost of reduced hatch success of eggs. Among birds with an original clutch size of five (higher quality), reducing and enlarging clutch size reduced and increased relative body mass loss respectively without affecting hatch success. In common eiders many females abandon their own ducklings to the care of other females. Enlarging five–egg clutches led to increased brood care rate despite the higher effort spent incubating these clutches, indicating that the higher fitness value of a large brood is increasing adult brood investment. This study shows that the ability to respond to clutch–size manipulations depends on original clutch size, reflecting differences in female quality. Females of low quality were reluctant to increase investment at the cost of lower hatch success, whereas females of higher quality apparently have a larger capacity both to increase incubation effort and brood care investment.

Costs of reproduction in common eiders ( Somateria mollissima ): An assessment of relationships between reproductive effort and future survival and reproduction based on observational and experimental studies

The two traditional approaches to the study of costs of reproduction, correlational and experimental, have been used in parallel in a breeding colony of common eiders ( Somateria mollissima ) and were compared in this paper. The analysis of the observational data was based on a two-strata capture-recapture model, the strata being defined on the basis of the clutch size laid by individual females in a given year. The best model according to AIC C indicated substantial variation in survival, recapture and transition rates, but overall a pattern emerged: females laying large clutches have a somewhat higher survival and much higher capture rate than females laying small clutches, and transition from large to small clutch size occurs much more frequently than the reverse transition. The analysis of the experimental data (adding/removing one egg) showed that no clear effect was found on either survival or transition rates. We conclude by suggesting (1) that condition should be included in multi-strata models in addition to reproductive effort; (2) that a specific study design for estimating the proportion of non-breeding females should be implemented, and (3) that non-breeding (a non-observable state in this study) may be influenced by previous reproduction events.

A first evaluation of the usefulness of feathers of nestling predatory birds for non-destructive biomonitoring of persistent organic pollutants

In previous studies, feathers of adult predatory birds have been evaluated as valid non-destructive biomonitor matrices for persistent organic pollutants (POPs). In this study, we assessed for the first time the usefulness of nestling raptor feathers for non-destructive biomonitoring of POPs. For this purpose, we collected body feathers and blood of nestlings from three avian top predators from northern Norway: northern goshawks (Accipiter gentilis), white-tailed eagles (Haliaeetus albicilla) and golden eagles (Aquila chrysaetos). We were able to detect a broad spectrum of legacy POPs in the nestling feathers of all three species (Σ PCBs: 6.78-140ng g(-1); DDE: 3.15-145ng g(-1); Σ PBDEs: 0.538-7.56ng g(-1)). However, these concentrations were lower compared to other studies on raptor species, probably due to the aspect of monitoring of nestlings instead of adults. Besides their analytical suitability, nestling feathers also appear to be biologically informative: concentrations of most POPs in nestling feathers showed strong and significant correlations with blood plasma concentrations in all species (p<0.050; 0.775<r<0.994). In addition, the reported correlations between feathers and blood plasma were much higher than those previously reported for adult individuals. Accumulation profiles and species-specific differences were in accordance with other toxicological studies on avian species and generally in agreement with the specific ecology of the studied species. In summary, our results indicate that the use of nestling feathers of northern raptors may be a valid and promising non-destructive biomonitoring strategy for POPs in their ecosystems.

Climate change and the increasing impact of polar bears on bird populations

The Arctic is becoming warmer at a high rate, and contractions in the extent of sea ice are currently changing the habitats of marine top-predators dependent on ice. Polar bears (Ursus maritimus) depend on sea ice for hunting seals. For these top-predators, longer ice-free seasons are hypothesized to force the bears to hunt for alternative terrestrial food, such as eggs from colonial breeding birds. We analyzed time-series of polar bear observations at four locations on Spitsbergen (Svalbard) and one in east Greenland. Summer occurrence of polar bears, measured as the probability of encountering bears and the number of days with bear presence, has increased significantly from the 1970/80s to the present. The shifts in polar bear occurrence coincided with trends for shorter sea ice seasons and less sea ice during the spring in the study area. This resulted in a strong inverse relationship between the probability of bear encounters on land and the length of the sea ice season. Within, 10 years after their first appearance on land, polar bears had advanced their arrival dates by almost 30 days. Direct observations of nest predation showed that polar bears may severely affect reproductive success of the barnacle goose (Branta leucopsis), common eider (Somateria mollissima) and glaucous gull (Larus hyperboreus). Nest predation was strongest in years when the polar bears arrived well before hatch, with more than 90% of all nests being predated. The results are similar to findings from Canada, and large-scale processes, such as climate and subsequent habitat changes, are pinpointed as the most likely drivers in various parts of the Arctic. We suggest that the increasing, earlier appearance of bears on land in summer reflects behavioral adaptations by a small segment of the population to cope with a reduced hunting range on sea ice. This exemplifies how behavioral adaptations may contribute to the cascading effects of climate change.

Anti-parasite treatment removes negative effects of environmental pollutants on reproduction in an Arctic seabird

Recent studies have shown that the detrimental effects of anthropogenic pollutants may be worse if organisms are exposed to natural stress. In this study, we examined whether negative effects of organochlorines (OCs) could be influenced by parasites. In two breeding seasons, we administered an anti-helminthic drug to groups of breeding glaucous gulls (Larus hyperboreus), whereas control groups were placebo treated. In all birds, blood residues of the most important OCs in the study population (hexachlorobenzene, oxychlordane, p,p′-dichlorodiphenyldichloroethylene and polychlorinated biphenyl), were measured. The relationships between OCs and fitness components (i.e. nesting success and return rate between breeding seasons) were then compared between the birds receiving anti-parasite treatment and the controls. Among untreated males, higher blood residues of OCs were associated with lowered nesting success, while in males receiving anti-parasite treatment, there was no detrimental effect of OCs on fitness. Return rate was not affected by treatment or OCs. Our findings suggest that parasites may be an important factor in triggering reproductive effects of such pollutants, and that relatively low levels of OCs may have serious reproductive consequences in natural populations when stress from other sources is high.

Relationships between organohalogen contaminants and blood plasma clinical–chemical parameters in chicks of three raptor species from Northern Norway

Organohalogen contaminants (OHCs) may affect various physiological parameters in birds including blood chemistry. We therefore examined blood plasma clinical-chemical parameters and OHCs in golden eagle, white-tailed eagle and goshawk chicks from Northern Norway. Correlation analyses on pooled data showed that alkaline phosphatase (ALKP), glucose and creatinine were significantly negatively correlated to various OHCs (all: p<0.05; r: -0.43 to -0.55; n=23), while alanine aminotransferase (ALAT), total protein, cholesterol, uric acid, total bilirubin, ratios protein:creatinine and uric acid:creatinine were significantly positively correlated to various OHCs (all: p<0.05; r: 0.43-0.96). Based on these relationships, we suggest that the OHC concentrations found in certain raptor chicks of Northern Scandinavia may impact blood plasma biochemistry in a way that indicates impacts on liver, kidney, bone, endocrinology and metabolism. In order to elaborate further on these relationships and mechanisms, we recommend that a larger study should take place in the near future.