Ontogeny of thermoregulatory mechanisms in king penguin chicks (Aptenodytes patagonicus)

The rapid maturation of thermoregulatory mechanisms may be of critical importance for optimising chick growth and survival and parental energy investment under harsh climatic conditions. The ontogeny of thermoregulatory mechanisms was studied in growing king penguin chicks from hatching to the full emancipation observed at 1 month of age in the sub-Antarctic area (Crozet Archipelago). Newly hatched chicks showed small, but significant regulatory thermogenesis (21% rise in heat production assessed by indirect calorimetry), but rapidly became hypothermic. Within a few days, both resting (+32%) and peak (+52%) metabolic rates increased. The first week of life was characterised by a two-fold rise in thermogenic capacity in the cold, while thermal insulation was not improved. During the second and third weeks of age, thermal insulation markedly rose (two-fold drop in thermal conductance) in relation to down growth, while resting heat production was slightly reduced (-13%). Shivering (assessed by electromyography) was visible right after hatching, although its efficiency was limited. Thermogenic efficiency of shivering increased five-fold with age during the first weeks of life, but there was no sign of non-shivering thermogenesis. We conclude that thermal emancipation of king penguin chicks may be primarily determined by improvement of thermal insulation after thermogenic processes have become sufficiently matured. Both insulative and metabolic adaptations are required for the rapid ontogeny of thermoregulation and thermal emancipation in growing king penguin chicks.     

Early changes in plasma hormones and metabolites during fasting in king penguin chicks

Summary Chicks of the king penguin (Aptenodytes patagonica) can tolerate a fast of 4–6 months during the subantarctic winter. The aim of this work was to study their initial response to food deprivation. Nine chicks were starved for 18 days. Two phases of starvation were defined according to changes in the specific daily loss in body mass: it decreased by 92% in phase I (6.6±0.3 days) and remained steady and low in phase II. Phase I was marked by a large decline in protein utilization, indicated by decreases in plasma levels of alanine (58%), uric acid (89%) and urea (76%) together with a decrease in circulating corticosterone (60%) and thyroxine (75%). In phase I, plasma insulin concentration decreased (61%) in some birds, but did not change in others; plasma pancreatic glucagon was stable whereas gut-glucagon decreased by 75%. Free fatty acids and -hydroxybutyrate concentrations gradually rode during the fast to 5 to 6 times pre-fast levels. Glycemia remained unchanged. Phase II was characterized by no change in plasma concentrations of protein-derived metabolites and by no or little change in circulating hormone levels. From comparison with previous data, we conclude that there are similar early adjustments to food deprivation in king penguin chick, rat and man: (1) a decrease in resting metabolic rate, (2) a decrease in protein utilization, and (3) mobilization of fat stores. The key, adaptations to long-term fasting in these species are therefore effectiveness in protein sparing and ability to prolong this situation.Abbreviations GLI glucagon-like immunoreactivity - PG pancreatic glucagon - -OHB -hydroxybutyrate - FFA free fatty acids - T ₃ triiodothyronine - T ₄ thyroxine     

Effects of food availability on fledging condition and post-fledging survival in king penguin chicks

Effects of summer food shortage on king penguin Aptenodytes patagonicus chicks were studied at South Georgia. Two cohorts were compared, fledging in the austral summers of 1992 (n = 32) and 1994 (n = 33) when availability of food was judged good and poor, respectively. The former cohort had a higher pre-fledging mean mass (12.78 kg vs ≤ 10.03 kg), fledged earlier (median 5 January vs 21 January), and a higher proportion was re-sighted within 2 years of fledging (28% vs 0%). Within 4 years, 47% of the former cohort had been re-sighted (i.e. post-fledge survival); in addition, one was observed at the Falkland Islands, and 22% had bred (i.e. recruitment) in their colony of origin. The re-sighted chicks of the 1992 cohort fledged earlier than those not re-sighted (median 24 December vs 10 January), but it remain unclear if they were heavier at fledging. All chicks in this study (n = 65) were marked with both transponders (subcutaneously implanted) and flipper bands (on one flipper), and no losses of any markings were found (controlled up to 4 years afterwards). Therefore, data on chick post-fledging survival and recruitment were not adjusted for losses of markings, as has been done in other studies. Received: 21 October 1996 / Accepted: 2 February 1997     

Cocktail-party effect in king penguin colonies

The king penguin, Aptenodytes patagonicus, breeds without a nest in colonies of several thousands of birds. To be fed, the chick must recognize the parents in a particularly noisy environment using only vocal cues. The call an adult makes when seeking the chick is emitted at a high amplitude level. Nevertheless, it is transmitted in a colonial context involving the noise generated by the colony and the screening effect of the bodies, both factors reducing the signal-to-noise ratio. In addition, the adult call is masked by a background noise with similar amplitude and spectral and temporal characteristics, enhancing the difficulty for the chick in finding its parents. We calculate that the maximum distance from the caller at which its signal can be differentiated from the background noise (signal-to-noise ratio equal to 1) should not exceed 8 to 9 m in a feeding area. But our tests show that, in fact, chicks can discriminate between the parental call and calls from other adults at a greater distance, even when call intensity is well below that of the noise of simultaneous calls produced by other adults. This capacity to perceive and extract the call of the parent from the ambient noise and particularly from the calls of other adults, termed the ''cocktail-party effect'' in speech intelligibility tests, enhances the chick''s ability to find its parents.     

Sexual harassment of a king penguin by an Antarctic fur seal

Males of gregarious pinnipeds are often aggressive to conspecifics and sexual coercion of females is commonplace. Males of some pinniped species have been known to attempt interspecific copulation, occasionally being successful in producing hybrid offspring. The most extreme case of interspecific sexual coercion reported concerned species from different families. We report a case of interspecific sexual harassment bridging the rank of vertebrate class.     

Ornamental colors reveal age in the king penguin

We investigated whether delayed plumage maturation occurred in king penguins (Aptenodytes patagonicus). Therefore we examined the relationships between age and sex on spectral properties and size of two colored plumage patches and a UV-reflective beak spot, using known-age cohorts. Unlike the colored patch on the breast, we found age differences in ear and beak coloration. These results suggest that head ornaments in king penguins could signal sexual maturity or social status. No sex differences were found in the intensity of colored ornaments, which can result from mutual mate choice or genetic correlation between sexes. Size of colored patches did not relate to age or sex.     

Maternal telomere length inheritance in the king penguin

Telomeres are emerging as a biomarker for ageing and survival, and are likely important in shaping life-history trade-offs. In particular, telomere length with which one starts in life has been linked to lifelong survival, suggesting that early telomere dynamics are somehow related to life-history trajectories. This result highlights the importance of determining the extent to which telomere length is inherited, as a crucial factor determining early life telomere length. Given the scarcity of species for which telomere length inheritance has been studied, it is pressing to assess the generality of telomere length inheritance patterns. Further, information on how this pattern changes over the course of growth in individuals living under natural conditions should provide some insight on the extent to which environmental constraints also shape telomere dynamics. To fill this gap partly, we followed telomere inheritance in a population of king penguins (Aptenodytes patagonicus). We tested for paternal and maternal influence on chick initial telomere length (10 days old after hatching), and how these relationships changed with chick age (at 70, 200 and 300 days old). Based on a correlative approach, offspring telomere length was positively associated with maternal telomere length early in life (at 10 days old). However, this relationship was not significant at older ages. These data suggest that telomere length in birds is maternally inherited. Nonetheless, the influence of environmental conditions during growth remained an important factor shaping telomere length, as the maternal link disappeared with chicks'' age.     

Impact of predation on king penguin Aptenodytes patagonicus in Crozet Archipelago

Predation can have major effects on population dynamics, but predator–prey interactions in marine ecosystems have rarely been studied. While the king penguin is one of the most studied seabirds, little is known about the impact of predation on its population dynamics. Here, we determine the impact of the main predators (giant petrels and skuas) on king penguin breeding success taking into account the nocturnal predation of petrels. We found that predation is the most important source of breeding failure for king penguins. The smallest chicks within crèches are the most hunted. The periphery of the colony suffers the highest risk of predation during summer. Our study shows the unequal quality of some areas inside the colony in terms of predation risk and breeding success, and points out the importance of timing in successful breeding.     

Moulting fast and time constraint for reproduction in the king penguin

King penguins (Aptenodytes patagonicus) replace their complete plumage during a 3- to 4-week moulting fast on shore. After moulting, birds forage at sea for 2-3 weeks to store energy reserves for reproduction. Using an automatic identification and weighing system coupled with visual observations, we investigated the trade-off between the moulting fast and the following breeding attempt, in terms of body condition and in relation to time constraints, in free-living king penguins. King penguins reached their lowest body mass of the yearly cycle at the end of the moulting fast; this was lower than that at the end of the incubation fasts and not different from the body mass at egg desertion. Later in the season, the duration of the moulting fast became shorter and old feather loss occurred earlier after arrival ashore. Postmoulting foraging trips were shorter, but at the expense of body condition at the beginning of courtship. These results are discussed in relation to the synchronisation of the breeding cycle with food availability and mate choice.     

Ontogeny of thermoregulation and energy metabolism in pygoscelid penguin chicks

The ontogeny of thermoregulation and energy metabolism of chinstrap (Pygoscelis antarctica) and gentoo (P. papua) penguins was studied on King George Island, South Shetland Island, Antarctica. The major findings of this study are: Chinstrap and gentoo penguin chicks hatched completely poikilothermic, due to their poor heat-production ability at low ambient temperatures. They were able to maintain high body temperatures and metabolic rates only by being brooded by adults. Newly hatched chinstrap penguin chicks had, at a specified ambient temperature, significantly higher metabolic rates than newly hatched gentoos. Moreover, chinstrap chicks maintained a significantly higher body temperature. It is suggested that this is a non-acclimatory metabolic adaptation of chinstrap penguin chicks to the lower mean temperatures of their breeding areas. On the 15th day after hatching, chinstrap chicks were completely, and gentoo chicks almost completely, homeothermic. In spite of their high thermogenic capacity from about day 10, chicks were not at that time capable of controlling heat dissipation, and were still dependent on their parents. In older downy chicks and fledglings, heat loss at low temperatures, expressed as heat conductance (CA), was similar to that found for the adults of other penguin species. Just before moulting the CA of chicks was lower than after moulting. Moulting alone did not cause a clear increase in CA. Towards the end of their stay on land the CA of pre-fledged gentoos decreased by 31%. This decrease was not connected with the development of feathers or growth in the chicks' weight. The combination of the low CA and high SMR of chicks gave very low lower critical temperatures, near -15 degrees C. The wide thermoneutral zones of the chicks covered the whole range of air temperature variations in the breeding colonies of both species studied on King George Island. The CA values of homeothermic chinstrap chicks were not lower than those of gentoos, despite the more southern breeding range of the former species. The older chicks of both species are well protected against cold. Any further increase in insulation in chinstrap chicks would be of no adaptative importance.     

Metabolic fate of yolk fatty acids in the developing king penguin embryo

This study examines the metabolic fate of total and individual yolk fatty acids (FA) during the embryonic development of the king penguin, a seabird characterized by prolonged incubation (53 days) and hatching (3 days) periods, and a high n-3/n-6 polyunsaturated FA ratio in the egg. Of the approximately 15 g of total FA initially present in the egg lipid, 87% was transferred to the embryo by the time of hatching, the remaining 13% being present in the internalized yolk sac of the chick. During the whole incubation, 83% of the transferred FA was oxidized for energy, with only 17% incorporated into embryo lipids. Prehatching (days 0-49), the fat stores (triacylglycerol) accounted for 58% of the total FA incorporated into embryo lipid. During hatching (days 49-53), 40% of the FA of the fat stores was mobilized, the mobilization of individual FA being nonselective. At hatch, 53% of the arachidonic acid (20:4n-6) of the initial yolk had been incorporated into embryo lipid compared with only 15% of the total FA and 17-24% of the various n-3 polyunsaturated FA. Similarly, only 32% of the yolk's initial content of 20:4n-6 was oxidized for energy during development compared with 72% of the total FA and 58-66% of the n-3 polyunsaturated FA. The high partitioning of yolk FA toward oxidization and the intense mobilization of fat store FA during hatching most likely reflect the high energy cost of the long incubation and hatching periods of the king penguin. The preferential partitioning of 20:4n-6 into the structural lipid of the embryo in the face of its low content in the yolk may reflect the important roles of this FA in tissue function.