Rates of oxygen consumption in four species of seabird at Palmer Station, Antarctic peninsula

1. We determined standard metabolic rates (SMR) of wild-caught adults of the Adelie penguin, southern giant fulmar, blue-eyed shag, and South Polar skua at Palmer Station, Antarctica, during January and February 1981. Oxygen consumption was measured volumetrically in a closed system at temperatures between 2 and 12 degrees C. 2. Mean SMR varied between 0.82 l O2/kg per hr for male fulmars and 1.30 l O2/kg per hr for unsexed adult skuas. Values were 174-198% of those predicted by the Lasiewski-Dawson equation for nonpasserines. 3. The SMR of the Adelie penguin was considerably higher than that reported in other studies and higher than most values for other species of penguins. 4. Our measurements of oxygen consumption agree with some estimates of metabolism based upon loss of mass by fasting birds during incubation.     

Regional heterothermy and conservation of core temperature in emperor penguins diving under sea ice

Temperatures were recorded at several body sites in emperor penguins (Aptenodytes forsteri) diving at an isolated dive hole in order to document temperature profiles during diving and to evaluate the role of hypothermia in this well-studied model of penguin diving physiology. Grand mean temperatures (+/-S.E.) in central body sites during dives were: stomach: 37.1+/-0.2 degrees C (n=101 dives in five birds), pectoral muscle: 37.8+/-0.1 degrees C (n=71 dives in three birds) and axillary/brachial veins: 37.9+/-0.1 degrees C (n=97 dives in three birds). Mean diving temperature and duration correlated negatively at only one site in one bird (femoral vein, r=-0.59, P<0.05; range <1 degrees C). In contrast, grand mean temperatures in the wing vein, foot vein and lumbar subcutaneous tissue during dives were 7.6+/-0.7 degrees C (n=157 dives in three birds), 20.2+/-1.2 degrees C (n=69 in three birds) and 35.2+/-0.2 degrees C (n=261 in six birds), respectively. Mean limb temperature during dives negatively correlated with diving duration in all six birds (r=-0.29 to -0.60, P<0.05). In two of six birds, mean diving subcutaneous temperature negatively correlated with diving duration (r=-0.49 and -0.78, P<0.05). Sub-feather temperatures decreased from 31 to 35 degrees C during rest periods to a grand mean of 15.0+/-0.7 degrees C during 68 dives of three birds; mean diving temperature and duration correlated negatively in one bird (r=-0.42, P<0.05). In general, pectoral, deep venous and even stomach temperatures during diving reflected previously measured vena caval temperatures of 37-39 degrees C more closely than the anterior abdominal temperatures (19-30 degrees C) recently recorded in diving emperors. Although prey ingestion can result in cooling in the stomach, these findings and the lack of negative correlations between internal temperatures and diving duration do not support a role for hypothermia-induced metabolic suppression of the abdominal organs as a mechanism of extension of aerobic dive time in emperor penguins diving at the isolated dive hole. Such high temperatures within the body and the observed decreases in limb, anterior abdomen, subcutaneous and sub-feather temperatures are consistent with preservation of core temperature and cooling of an outer body shell secondary to peripheral vasoconstriction, decreased insulation of the feather layer, and conductive/convective heat loss to the water environment during the diving of these emperor penguins.     

The winter migration of Adelie penguins breeding in the Ross Sea sector of Antarctica

Satellite telemetry was used to monitor the migratory movements of a single Adelie penguin (Pygoscelis adeliae) from Cape Hallett (72.31°S, 170.21°E) following the 1997/1998 breeding season. Locations were obtained using the ARGOS satellite system and compared with the migratory paths taken by two penguins from the Northern Colony at Cape Bird, Ross Island (77.22°S, 166.48°E) following the 1990/1991 breeding season. Although the sample sizes are small, if representative they would indicate that: (1) Adelie penguins breeding in the Ross Sea follow a common migratory path, (2) Adelie penguins breeding in the Ross Sea may travel to a common over-winter feeding ground west and north of the Balleny Islands, and (3) Adelie penguins breeding at 77°S on Ross Island travel nearly twice the distance during their over-winter migration as do those penguins breeding at Cape Hallett and colonies further north. While the Cape Hallett penguin was tracked successfully for 172 days, a record for Adelie penguins, the problem of long-term attachment of transmitters to penguins remains.     

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.     

Energetic cost of foraging in free-diving emperor penguins.

Hypothesizing that emperor penguins (Aptenodytes forsteri) would have higher daily energy expenditures when foraging for their food than when being hand-fed and that the increased expenditure could represent their foraging cost, we measured field metabolic rates (FMR; using doubly labeled water) over 4-d periods when 10 penguins either foraged under sea ice or were not allowed to dive but were fed fish by hand. Surprisingly, penguins did not have higher rates of energy expenditure when they dove and captured their own food than when they did not forage but were given food. Analysis of time-activity and energy budgets indicated that FMR was about 1.7 x BMR (basal metabolic rate) during the 12 h d(-1) that penguins were lying on sea ice. During the remaining 12 h d(-1), which we termed their "foraging period" of the day, the birds were alert and active (standing, preening, walking, and either free diving or being hand-fed), and their FMR was about 4.1 x BMR. This is the lowest cost of foraging estimated to date among the eight penguin species studied. The calculated aerobic diving limit (ADL(C)), determined with the foraging period metabolic rate of 4.1 x BMR and known O(2) stores, was only 2.6 min, which is far less than the 6-min ADL previously measured with postdive lactate analyses in emperors diving under similar conditions. This indicates that calculating ADL(C) from an at-sea or foraging-period metabolic rate in penguins is not appropriate. The relatively low foraging cost for emperor penguins contributes to their relatively low total daily FMR (2.9 x BMR). The allometric relationship for FMR in eight penguin species, including the smallest and largest living representatives, is kJ d(-1)=1,185 kg(0.705).     

An emperor penguin population estimate: the first global, synoptic survey of a species from space

Our aim was to estimate the population of emperor penguins (Aptenodytes fosteri) using a single synoptic survey. We examined the whole continental coastline of Antarctica using a combination of medium resolution and Very High Resolution (VHR) satellite imagery to identify emperor penguin colony locations. Where colonies were identified, VHR imagery was obtained in the 2009 breeding season. The remotely-sensed images were then analysed using a supervised classification method to separate penguins from snow, shadow and guano. Actual counts of penguins from eleven ground truthing sites were used to convert these classified areas into numbers of penguins using a robust regression algorithm.We found four new colonies and confirmed the location of three previously suspected sites giving a total number of emperor penguin breeding colonies of 46. We estimated the breeding population of emperor penguins at each colony during 2009 and provide a population estimate of ~238,000 breeding pairs (compared with the last previously published count of 135,000-175,000 pairs). Based on published values of the relationship between breeders and non-breeders, this translates to a total population of ~595,000 adult birds.There is a growing consensus in the literature that global and regional emperor penguin populations will be affected by changing climate, a driver thought to be critical to their future survival. However, a complete understanding is severely limited by the lack of detailed knowledge about much of their ecology, and importantly a poor understanding of their total breeding population. To address the second of these issues, our work now provides a comprehensive estimate of the total breeding population that can be used in future population models and will provide a baseline for long-term research.     

The cost of a hot meal: facultative specific dynamic action may ensure temperature homeostasis in post-ingestive endotherms.

1. Resting metabolic rate was determined in Adelie penguins (Pygoscelis adeliae) that had been fed warm and cold ingesta. 2. Post-ingestion metabolic rate was found to be related to food temperature rather than to specific dynamic action. 3. Calorimetric calculations indicate that up to 13% of Adelie penguin daily energy expenditure may be used heating ingested food to body temperature. 4. Heating food costs are predicted to be higher in endotherms living in cold regions and in species with low assimilation efficiencies, but may be minimized by appropriate foraging behaviour.     

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.     

The long engagement of the emperor penguin

In birds, courtship is generally short relative to the whole breeding cycle. Emperor penguins (Aptenodytes forsteri), however, are an exception as their courtship period is much longer (ca. 6 weeks) than the courtship of other penguin species. This strategy may appear surprising, as it is especially costly to fast and endure drastic climatic conditions for long periods at the colony (1.5 and up to 4 months for females and males, respectively). We examined here the reasons of this extended courtship period and found that emperor penguins returned earlier to the colony when primary oceanic production before breeding was high. This suggests that emperor penguins return to the colony as soon as primary oceanic production in summer allows them to replenish their body reserves. The extended period of time spent at the colony during courtship may therefore result from an evolutionary process that confers advantages to emperor penguins that arrive earlier at the colony by reducing predation risks and offering better chances of securing a partner.     

Body temperature changes induced by huddling in breeding male emperor penguins

Huddling is the key energy-saving mechanism for emperor penguins to endure their 4-mo incubation fast during the Antarctic winter, but the underlying physiological mechanisms of this energy saving have remained elusive. The question is whether their deep body (core) temperature may drop in association with energy sparing, taking into account that successful egg incubation requires a temperature of about 36 degrees C and that ambient temperatures of up to 37.5 degrees C may be reached within tight huddles. Using data loggers implanted into five unrestrained breeding males, we present here the first data on body temperature changes throughout the breeding cycle of emperor penguins, with particular emphasis on huddling bouts. During the pairing period, core temperature decreased progressively from 37.5 +/- 0.4 degrees C to 36.5 +/- 0.3 degrees C, associated with a significant temperature drop of 0.5 +/- 0.3 degrees C during huddling. In case of egg loss, body temperature continued to decrease to 35.5 +/- 0.4 degrees C, with a further 0.9 degrees C decrease during huddling. By contrast, a constant core temperature of 36.9 +/- 0.2 degrees C was maintained during successful incubation, even during huddling, suggesting a trade-off between the demands for successful egg incubation and energy saving. However, such a limited drop in body temperature cannot explain the observed energy savings of breeding emperor penguins. Furthermore, we never observed any signs of hyperthermia in huddling birds that were exposed to ambient temperatures as high as above 35 degrees C. We suggest that the energy savings of huddling birds is due to a metabolic depression, the extent of which depends on a reduction of body surface areas exposed to cold.     

Effects of corticosterone treatment on growth, development, and the corticosterone response to handling in young Japanese quail (Coturnix coturnix japonica)

Corticosterone, a glucocorticoid secreted during stress responses, has a range of actions that help birds respond to stressors. Although effects of corticosterone treatment have been described in several avian species, the impacts of defined increases in plasma corticosterone on early development and on corticosterone stress responses are little known. These issues were addressed by providing quail with different doses of corticosterone in drinking water from days 8 to 38 post-hatch. The corticosterone dose consumed by each bird during treatment days 15-30 was calculated by measuring water intake. The corticosterone dose was inversely, but weakly, correlated with weights of the bursa, thymus, spleen, liver, testes, oviduct, muscle, and body, and positively correlated with peritoneal fat deposition. When birds were divided into groups based on their corticosterone intake, weights of the spleen, thymus, bursa, muscle, testes, and oviduct were significantly reduced in birds receiving the highest doses; with the exception of muscle, similar reductions were also observed in birds receiving medium doses, and thymic growth was inhibited in birds receiving low doses. The acute corticosterone stress response was measured by handling birds for 15 min. Plasma corticosterone was transiently increased at 15 min in control birds in response to the handling stressor. Some birds consuming low doses of corticosterone had corticosterone responses similar to control birds. Initial corticosterone concentrations were elevated in birds consuming higher doses of corticosterone. Plasma corticosterone in these birds decreased from 0 to 15 min, then increased from 15 to 30 min. The initial decrease could be due to corticosterone clearance, whilst the increase could indicate that the birds had a greater response than control birds to isolation as a stressor. Corticosterone treatment may have reduced the strength of corticosterone negative feedback within the hypothalamo-pituitary-adrenal axis. The results indicate that individuals and organs differ in their sensitivity to corticosterone. Moreover, elevated plasma corticosterone may disrupt the acute corticosterone stress response, and may thus reduce the ability of birds to cope with stressors.     

Morphological aspects of the heart of the northern rockhopper penguin (Eudyptes chrysocome moseleyi ): possible implication in diving behavior and ecology?

We compared the heart morphology of the small, deep-diving northern rockhopper penguin to the hearts of small, shallow-diving and large, deep-diving penguin species. The rockhopper penguin had a heart larger than expected for its body mass, and its heart weight/body weight was significantly greater than in the larger Adélie penguin. We found the rockhopper's right ventricle weight/heart weight to be significantly greater than this relationship in both the larger chinstrap and Adélie penguins. The relationship of the right to left ventricular weights in the rockhopper heart is not different to that of the large, deepest-diving emperor penguin. A larger heart in the rockhopper penguin might be related to its diving behavior and ecology if it contributes to diving efficiency during foraging by increasing lung perfusion during surface recovery. This would lead to decreased surface time. Accepted: 20 May 2000     

Subcutaneous clostridial infection in Adelie penguins in Hope Bay,Antarctica

During the 2000–2001 breeding season in Hope Bay, Antarctica, two adult Adelie penguins were found dead with lesions compatible with subcutaneous clostridial infection. Clostridium cadaveris was isolated from the musculature and the subcutaneous tissue of one of these two penguins, whereas Clostridium sporogenes was isolated from the subcutaneous tissue of the other penguin. Escherichia coli and Staphylococcus spp. were isolated from both animals. This is the first report of subcutaneous clostridial infection in Antarctic Adelie penguins.     

Refeeding signal in fasting-incubating king penguins: changes in behavior and egg temperature

This study is directed toward understanding the process of feeding stimulation ("refeeding signal") that has been suggested to operate below a body mass threshold or critical metabolic status in spontaneously fasting birds. Behavior and egg temperature (T(egg)) were continuously monitored by video monitoring and biotelemetry, respectively, in fasting-incubating king penguins kept in a pen to prevent relief by the partner until spontaneous egg abandonment. Penned birds fasted 10 days more and lost 1.2 kg more than birds relieved normally by their partner, abandoning the egg about 1 wk after reaching a critical body mass. Definitive egg abandonment was preceded by transitory abandonments of progressively increasing duration during which time the birds went further and further away from their egg. There were marked interindividual differences but on average transitory abandonments began 36 +/- 5 h before the definitive abandonment and were paralleled by resumption of display songs signaling the readiness of the bird to depart for feeding. T(egg) was maintained at around 35.7 degrees C during normal incubation but significantly decreased the last 2 days before egg abandonment. These changes are interpreted as reflecting a stimulation to refeed at a threshold body mass corresponding to a critical fat store depletion. Thus the fasting-incubating king penguin appears to be an interesting animal model for understanding the long-term metabolic control of feeding behavior in relation to energy status.     

Lack of consistent hormonal responses to capture during the breeding season of the bearded dragon, Pogona barbata

The bearded dragon (Pogona barbata: Agamidae) is a diurnal, oviparous, multi-clutching lizard from Australia. We examined plasma hormonal responses to capture in males and females during the spring breeding season. Corticosterone concentrations at capture (0 h; < or =3 min after capture) were low (males: 1.81+/-0.63 ng/ml; females 2. 23+/-0.47 ng/ml) and within sexes were unrelated to the time of the day, snout-vent length or, in females, reproductive condition (vitellogenic, gravid, assumed spent). Corticosterone concentrations at capture were significantly and inversely correlated with body condition in males, but not in females. Unexpectedly, neither sex showed significant changes in mean concentrations of corticosterone at 3.5 or 24 h after capture compared with 0 h values. Corticosterone concentrations at 3.5 h after capture did not differ between dragons bled at capture or not. Concentrations of progesterone in both the sexes did not change between 0 h and 3.5 or 24 h after capture. Testosterone concentrations in males at capture were moderate (10.1+/-2.2 ng/ml), and unchanged at 3.5 h after capture. The adrenocortical axis of adult bearded dragons in the breeding season seems remarkably unresponsive to capture compared with many other reptiles. Low adrenocortical responses to capture may be a feature of reptiles known to adjust well to captivity.     

The effects of prey demography on humpback whale (Megaptera novaeangliae) abundance around Anvers Island, Antarctica

Baleen whales and Adelie penguins in the near-shore waters around the Antarctic Peninsula forage principally on Antarctic krill. Given the spatial overlap in the distribution of these krill predators (particularly humpback whales) and their dependence on krill, the goals of this paper are to determine if the inter-annual community structure and relative abundance of baleen whales around Anvers Island is related to krill demography and abundance, and if the potential exists for inter-specific interactions between Adelie penguins and baleen. We use whale sightings and prey data from both net tows and Adelie penguin stomach samples to correlate the abundance of humpback whales with krill demography and abundance from 1993 to 2001. We find significant relationships between whale abundance and the size–frequency distribution of krill targeted by Adelie penguins, as well as the foraging success of Adelie penguins. These findings suggest both krill predators share common prey preferences in the upper portions of the water column around Anvers Island. These findings highlight the need for better knowledge of baleen whale foraging ecology and inter-specific interactions with penguins, as sea ice and krill populations around the Antarctic Peninsula are affected by rapid changes in climate.     

Foraging by deep-diving birds is not constrained by an aerobic diving limit: a model of avian depth-dependent diving metabolic rate

The theoretical aerobic diving limit (tADL) specifies the duration of a dive after which oxygen reserves available for diving are depleted. The tADL has been calculated by dividing the available oxygen stores by the diving metabolic rate (DMR). Contrary to diving mammals, most diving birds examined to date exceed the tADL by a large margin. This discrepancy between observation and theory has engendered two alternative explanations suggesting that dive duration is extended either anaerobically or by depressing aerobic metabolism. Current formulations of tADL uncritically assume that DMR is independent of depth. However, diving birds differ from other vertebrate divers by having a larger respiratory system volume and by retaining air in their plumage while diving, thereby elevating buoyancy. Because air compresses with depth, diving power requirement decreases with depth. Following this principle, we modeled DMR to depth for Adelie and little penguins and reformulated the tADL accordingly. The model's results suggest that < approximately 5% of natural dives by Adelie penguins exceed the reformulated tADL(d), or maximal aerobic depth, and none in the more buoyant little penguin. These data suggest that, for both small and large species, deep diving birds rarely if ever exceed tADL(d).     

Genetic diversity in populations of Gentoo penguins (Pygoscelis papua)

RAPD analysis was used to examine the extent of genetic polymorphism in two populations of Gentoo penguin (Pygoscelis papua) from Antarctic Islands (Petermann and Livingston). The chosen two of three 10 mer oligonucleotide primers accordingly to preliminary results showed different levels of polymorphism in Gentoo penguins at Petermann Island (from 23.53 to 42.86%) and Livingston Island (from 52.94 to 57.14%). Nei's similarity coefficients were in range from 0.5606 (when Gentoo genome profiles were compared with RAPD profiles of two related penguin species: Pygoscelis adeliae (Adelie) and Pygoscelis antarctica (Chinstrep)) to 0.9281 among observed Gentoo penguin populations. Nei's distances values ranged from 0.0746 to 0.5787 among the populations and species. The obtained results will be used for further estimation of genetic diversity of Gentoo penguins and determination of their taxonomic status.     

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.     

Trapping initiates stress response in breeding and non-breeding house sparrows Passer domesticus: implications for using unmonitored traps in field studies

In response to a variety of unpredictable conditions, birds secrete the steroid hormone corticosterone, which has numerous effects on physiology and behavior. A standardized protocol of handling and restraint has been demonstrated to elicit a robust corticosterone response in many species of birds. In contrast, comparatively little is known about the effects of capture technique on corticosterone secretion in wild birds. Setting up multiple live traps checked at regular intervals allows field researchers to capture many birds in a short period of time. However, one potential drawback of this technique is that birds may spend unknown lengths of time in traps prior to sampling. Many birds appear to remain calm and/or feed during this period, potentially leading researchers to assume that corticosterone secretion is unaffected by trapping. We assessed the impact of being left in traps for up to 30 minutes on baseline corticosterone and subsequent corticosterone responses to restraint in non‐breeding and breeding house sparrows Passer domesticus. Traps were baited with seed, and birds were either removed immediately after entry (controls), or left in the trap undisturbed for 15 or 30 min. Upon removal, birds were subjected to a standardized handling/restraint protocol in which blood samples were collected within 3 min, and again at regular intervals for 60 min. Analysis of blood samples revealed that both non‐breeding and breeding sparrows that were held in the traps had significantly higher baseline corticosterone than controls, and showed no further increase in corticosterone secretion in response to handling. However, corticosterone responses to trapping differed seasonally. Our study indicates that although birds did not exhibit prolonged escape behavior while trapped, entry into a walk‐in trap initiated a robust stress response. Taken together with data from a previous study, our data suggest that ornithologists should consider species‐specific and stage‐specific effects of trapping on physiology when designing field studies.