Male Antarctic fur seals: neglected food competitors of bioindicator species in the context of an increasing Antarctic krill fishery.

The fishery for Antarctic krill is currently managed using a precautionary, ecosystem-based approach to limiting catch, with performance indices from a long-term monitoring program focused on several krill-dependent predators that are used to track ecosystem health. Concerns over increased fishing in concentrated areas and ongoing efforts to establish a Marine Protected Area along the Peninsula, a key fishing region, is driving the development of an adaptive management system for the fishery. The cumulative effects of fishing effort and interactions among krill-dependent predators and their performance is at present neglected in the CCAMLR Ecosystem Monitoring Program. However, we show considerable overlap between male Antarctic fur seals and the krill fishery in a complex mosaic, suggesting potential for cumulative impacts on other krill dependent predators. A holistic view is required as part of future efforts to manage the krill fishery that incorporates various sources of potential impacts on the performance of bioindicator species, including the fishery and its interactions with various krill dependent predators.

A global cline in a colour polymorphism suggests a limited contribution of gene flow towards the recovery of a heavily exploited marine mammal.

Evaluating how populations are connected by migration is important for understanding species resilience because gene flow can facilitate recovery from demographic declines. We therefore investigated the extent to which migration may have contributed to the global recovery of the Antarctic fur seal (Arctocephalus gazella), a circumpolar distributed marine mammal that was brought to the brink of extinction by the sealing industry in the eighteenth and nineteenth centuries. It is widely believed that animals emigrating from South Georgia, where a relict population escaped sealing, contributed to the re-establishment of formerly occupied breeding colonies across the geographical range of the species. To investigate this, we interrogated a genetic polymorphism (S291F) in the melanocortin 1 receptor gene, which is responsible for a cream-coloured phenotype that is relatively abundant at South Georgia and which appears to have recently spread to localities as far afield as Marion Island in the sub-Antarctic Indian Ocean. By sequencing a short region of this gene in 1492 pups from eight breeding colonies, we showed that S291F frequency rapidly declines with increasing geographical distance from South Georgia, consistent with locally restricted gene flow from South Georgia mainly to the South Shetland Islands and Bouvetøya. The S291F allele was not detected farther afield, suggesting that although emigrants from South Georgia may have been locally important, they are unlikely to have played a major role in the recovery of geographically more distant populations.

Extreme diversity in the songs of Spitsbergen's bowhead whales.

Almost all mammals communicate using sound, but few species produce complex songs. Two baleen whales sing complex songs that change annually, though only the humpback whale (Megaptera novaeangliae) has received much research attention. This study focuses on the other baleen whale singer, the bowhead whale (Balaena mysticetus). Members of the Spitsbergen bowhead whale population produced 184 different song types over a 3-year period, based on duty-cycled recordings from a site in Fram Strait in the northeast Atlantic. Distinct song types were recorded over short periods, lasting at most some months. This song diversity could be the result of population expansion, or immigration of animals from other populations that are no longer isolated from each other by heavy sea ice. However, this explanation does not account for the within season and annual shifting of song types. Other possible explanations for the extraordinary diversity in songs could be that it results either from weak selection pressure for interspecific identification or for maintenance of song characteristics or, alternatively, from strong pressure for novelty in a small population.

Biology of the Greenland shark Somniosus microcephalus.

Greenland shark Somniosus microcephalus is a potentially important yet poorly studied cold-water species inhabiting the North Atlantic and Arctic Oceans. Broad-scale changes in the Arctic ecosystem as a consequence of climate change have led to increased attention on trophic dynamics and the role of potential apex predators such as S. microcephalus in the structure of Arctic marine food webs. Although Nordic and Inuit populations have caught S. microcephalus for centuries, the species is of limited commercial interest among modern industrial fisheries. Here, the limited historical information available on S. microcephalus occurrence and ecology is reviewed and new catch, biological and life-history information from the Arctic and North Atlantic Ocean region is provided. Given the considerable by-catch rates in high North Atlantic Ocean latitudes it is suggested that S. microcephalus is an abundant predator that plays an important, yet unrecognized, role in Arctic marine ecosystems. Slow growth and large pup sizes, however, may make S. microcephalus vulnerable to increased fishing pressure in a warming Arctic environment.

Disruptive effects of persistent organohalogen contaminants on thyroid function in white whales (Delphinapterus leucas) from Svalbard.

We analysed levels of 56 organohalogen contaminants (OHCs) including brominated flame retardants, polychlorinated biphenyls (PCBs), and organochlorine pesticides in the blubber of white (beluga) whales (Delphinapterus leucas) from Svalbard, Norway (N=12; 6 adults [5 males and 1 female] and 6 subadults [4 males and 2 females]) collected in 1996-2001. We also measured circulating levels of thyroid hormones (THs) and thyroid stimulating hormone (TSH) in the whales. The results confirm that OHC levels in these white whales are among the highest levels recorded in wildlife from Svalbard, and at the high end of the range when compared to white whales from the North American Arctic. A projection to latent structure (PLS) model (subadults and adult males grouped together) revealed that known or suspected thyroid disruptive contaminants (polybrominated diphenylether [PBDE]-28, -47, -99, -100, and -154, hexachlorobenzene [HCB], and PCB-105) were negatively correlated with circulating levels of total thyroxin (TT4), free T4 (FT4) and free triiodothyronine (FT3). Most of these negative relationships were also confirmed using partial correlations controlling for length (and thus age) of the whales. The positive correlations of TT4, FT4 and FT3 with hexabromocyclododecane (HBCD), α-hexachlorocyclohexane (α-HCH), chlorinated bornanes CHB-40 and CHB-62 revealed by the PLS model were not confirmed by partial correlations. TH levels in the present study appeared to be somewhat lower than levels measured in beluga whales from the Canadian Arctic. However, we were not able to determine if this was caused by different levels of OHCs, or differences in biological factors (e.g. age, sex, moulting status, and season) and analytical methods between the studies. Although the sample sizes were low and statistical models cannot depict the biological cause-effect relationships, this study suggests negative influences of specific OHCs, particularly PBDEs, on thyroid hormone levels in white whales. The impact this might have on individual and population health is unknown.

The derivation of radionuclide transfer parameters for and dose-rates to an adult ringed seal (Phoca hispida) in an Arctic environment.

Radionuclide transfer parameters and dose-rates for an adult ringed seal from Svalbard have been determined based on empirical and estimated tissue activity concentrations and detailed dietary and habitat information. Whole-body equivalent concentration factors determined for anthropogenic radionuclides ranged from 10(1) ((90)Sr) to 10(2) ((137)Cs, (238)Pu and (239,240)Pu), while natural radionuclides ranged from 10(2) ((210)Pb) to 10(4) ((210)Po). Employing a dietary composition of 40% fish, 40% zooplankton and 20% benthic invertebrates, a whole-body biological half-life of 29 days was derived for (137)Cs. A total dose-rate of approximately 0.19microGyh(-1) (1.7mGya(-1)) was derived for an adult ringed seal; this dose-rate is virtually entirely attributable to the internal components of (210)Po and (40)K. The dose-rates associated with the presence of anthropogenically derived radionuclides in the present assessment fall many orders of magnitude below the dose-rates at which any biological effects would be expected.

Probabilistic biokinetic modelling of radiocaesium uptake in Arctic seal species: verification of modelled data with empirical observations.

The necessity to provide information about radionuclide concentrations in Arctic marine species has been heightened in recent years due to a number of accidents in Arctic regions involving nuclear vessels and the presence of a large number of potential radioactive contamination sources. The provision for such information is largely dependent on the use of radionuclide uptake and transfer models. The uptake of radionuclides in Arctic seal species in this study has been modelled using a probabilistic biokinetic approach. In this paper, model results are compared with empirical data from relevant samples taken within the Arctic region. Results indicate that the model performs well when estimating concentrations of (137)Cs in two seal species for both median values and reproduction of the distribution of data values, but not as well for a third seal species. Likely factors affecting the results are the probability density functions used for the input parameters.

Levels of toxaphene congeners in white whales (Delphinapterus leucas) from Svalbard, Norway.

This study reports concentrations of three pesticide toxaphene congeners (CHBs; CHB-26, -50 and -62) from the blubber of ten adult, male white whales (Delphinapterus leucas) from Svalbard, Norway. The CHB congeners that occurred at the highest levels in the blubber of the white whales were, as expected, CHB-26 (4636+/-1992 (SD) ng/g l.w.) and CHB-50 (6579+/-2214 ng/g l.w.); CHB-62 (232+/-231 ng/g l.w.) was also present, but at much lower concentrations. The mean level of the sum of the three CHBs (SigmaCHBs = 11,447+/-4208 ng/g l.w.) in this study is more than twice the mean concentrations of the well-known organochlorine (OC) pollutants SigmaDDTs (sum of pp'-DDT, pp'-DDE, pp'-DDD) and SigmaPCBs (sum of 27 PCB congeners) previously reported from the same individual white whales. The concentrations of CHBs in white whales from Svalbard are at the high end of the range for concentrations of these compounds compared to other Arctic white whale populations. Additionally, the contribution of CHBs to the overall OC burden is larger in white whales from Svalbard compared with their counterparts from other areas in the Arctic. Male white whales from Svalbard have several orders of magnitude higher concentrations of SigmaCHBs compared to seals and polar bears (Ursus maritimus) from the same area. The high levels of CHBs in these whales, and their dominance in the OC pattern, suggests that white whales in Svalbard are exposed to high levels of this group of contaminants. Further studies are needed to investigate possible effects of CHBs and other OC contaminants on the white whale population in Svalbard.

Accumulation, metabolism, and food-chain transfer of chlorinated and brominated contaminants in subadult white whales (Delphinapterus leucas) and narwhals (Monodon monoceros) from Svalbard, Norway.

The concentrations and patterns of polychlorinated biphenyls (PCBs), chlorinated pesticides, and polybrominated diphenyl ethers (PBDEs) were studied in white whales (Delphinapterus leucas) and narwhals (Monodon monoceros) from Svalbard, Norway. In addition, their main food items were included in the study. In the whales, a broad range of pollutants was found in relatively high concentrations. PCBs and pesticides were approximately 3000 and 8000 ng/g lipid, respectively, for white whales and three times higher for narwhals. PBDEs 47 were approximately 70 ng/g lipid for white whales and 170 ng/g lipid for narwhals. Compared with other marine mammals from the same area, contaminant levels are among the highest levels ever measured. These high levels are likely in part because of a decreased capacity to metabolize contaminants. Metabolic indices indicated that most compounds accumulate to the same degree in white whales and narwhals, but for some toxaphenes and chlordanes, narwhals might have a decreased metabolism and consequently a higher accumulation. The three-times-higher contaminant levels in blubber of narwhals was further explained by substantially higher contaminant levels in their more benthic diet. The high levels and broad pattern of accumulating pollutants make white whales and narwhals excellent indicators for a wide range of contaminants in the Arctic.

Partitioning of persistent organic pollutants in grey seal (Halichoerus grypus) mother-pup pairs.

Phocid seals have lipid rich milk, which is known to serve as a transfer medium through which persistent organic pollutants (POPs) move from mother to offspring during lactation. However, knowledge on this generational transfer of different POPs and the partitioning of these compounds in maternal and offspring tissues over the course of the lactation are limited. In this study we examined the qualitative and quantitative partitioning of a range of chlorinated POPs in maternal blubber, blood and milk as well as in pup blubber, collected early in the lactation period and late in the lactation period. In the lactating female, the high-chlorinated and hydrophobic compounds were passed less efficiently into the milk than the low-chlorinated compounds and more water-soluble compounds. Significantly, lower maternal blood concentrations than in maternal blubber biopsies suggest a stratification of POP concentrations in the blubber column of lactating female and lower concentrations in the metabolic active inner layers. Over the course of lactation, there was a significant increase in maternal blood and milk concentrations of POPs as opposed to no change in maternal blubber biopsy concentrations. This was most apparent for the hydrophobic and high-chlorinated compounds. The most likely explanation for this is that the metabolic active inner blubber layer, from which the milk lipids are derived from, is in steady state with the circulatory system, while the outer layers are more static and only slowly respond to changes in concentrations elsewhere in the body. The concentrations of the high-chlorinated and hydrophobic compounds were substantially lower in pup blubber than in maternal blubber. This probably relates the combined effect of these compounds stratification in maternal blubber and their slow transfer into the milk. The present study shows that the more hydrophobic and high-chlorinated compounds come to steady state less quickly in the different tissues than the more water-soluble and low-chlorinated compounds in the lactating female and her offspring. This has implications for which matrices to choose when sampling for assessing the toxicological risk of POPs in seals.

Precocial diving and patent foramen ovale in bearded seal (Erignathus barbatus) pups.

In this study we used a Doppler ultrasonic device, in combination with a sonographic contrast medium, to test whether free-living bearded seal (Erignathus barbatus) pups have a closed (anatomically or functionally) foramen ovale. A total of 17 examinations were performed on 12 individual pups with a body mass range of 29-103 kg (0-21 days old). These examinations showed that young bearded seal pups dive with a patent foramen ovale (PFO), and that this structure starts to close, at least functionally, during the 2nd week of life. The wide range in the timing of closure (one animal 21 days old still had a PFO) indicates that a closed foramen ovale is not crucial for the diving that these seals perform at this age. The primary function of diving during the 1st week of life is to avoid surface predation and only moderate diving ability is sufficient to achieve this goal. However, some of the diving performed by bearded seal pups with a PFO would likely be sufficient to create intravenous bubble formation during breath-hold diving in humans. Special adaptations in the seals, such as collapsible lungs and diving with minimal lung air volume, probably prevent this from happening.

Global distribution of halogenated dimethyl bipyrroles in marine mammal blubber.

Four halogenated dimethyl bipyrroles (HDBPs), hypothesized to be naturally produced, were quantitated in marine mammal blubber from a number of species obtained from various locations worldwide. HDBPs were found in samples from all locations studied. Concentrations of total HDBPs (SigmaHDBPs) ranged from 0.4 ng/g lipid weight in ringed seals (Phoca hispida) from the White Sea to 2,540 ng/g lipid weight in Dall's porpoise (Phocoenoides dalli) from the northwestern North Pacific Ocean. At their highest levels, SigmaHDBPs made up 11% of the total quantitated organohalogen body burden of adult male Dall's porpoises. In two beluga (Delphinapterus leucas) data subsets, it was found that males contained significantly higher concentrations of SigmaHDBPs than females. No significant effects of age or sex on SigmaHDBPs were observed in harbor seal (Phoca vitulina) and bowhead whale (Balaena mysticetus) data subsets. The geographical distribution of concentrations did not resemble that of the ubiquitous anthropogenic organohalogen, polychlorinated biphenyl congener CB-153. Higher concentrations of HDBPs and different patterns of congeners were observed in samples from Pacific as opposed to non-Pacific Ocean influenced environments. Concentrations of HDBPs in beluga from the Arctic and St. Lawrence River were similar. Their high abundance in north Pacific Ocean biota and widespread occurrence suggest that HDBPs undergo extensive transport from a source located primarily in the Pacific Ocean. Evidence from HDBP congener patterns indicates that both ocean currents and atmospheric transport likely play a role in the movement of HDBPs. These results imply that HDBPs and anthropogenic organohalogens have different sources and support the natural production hypothesis.

Localized oxygen use of healthy and low back pain individuals during controlled trunk movements.

Individuals who have low back pain (LBP) have significantly different motion characteristics than healthy individuals. However, the cause of these differences is unknown. Oxygen use of the erector spinae muscle was examined while simultaneously monitoring motion characteristics to determine whether oxygen use differed between healthy and LBP individuals. Thirty volunteers were classified as healthy, structural, or muscular-based LBP. A near-infrared spectrometer monitored oxygen use and blood volume in the lumbar region. Results showed significant differences in oxygen use but not blood volume between healthy and LBP subjects with muscular-based disorders. Inability of the muscular group to use oxygen in a manner similar to the healthy group indicates different processes at the tissue level, indicating that differences in oxygen use may provide insight into why motion patterns differ between healthy and LBP groups.

Chemical characterisation of the oligosaccharides in hooded seal (Cystophora cristata) and Australian fur seal (Arctocephalus pusillus doriferus) milk.

Carbohydrates were extracted from hooded seal milk, Crystophora cristata (family Phocidae). Free oligosaccharides were separated by gel filtration and then purified by ion exchange chromatography, gel filtration and preparative thin layer or paper chromatography and their structures determined by 1H-NMR. The hooded seal milk was found to contain inositol and at least nine oligosaccharides, most of which had lacto-N-neotetraose or lacto-N-neohexaose as core units, similar to those in milk of other species of Carnivora such as bears (Ursidae). Their structures were as follows: Gal(beta1-4)Glc (lactose); Fuc(alpha1-2)Gal(beta1-4)Glc (2'-fucosyllactose); Gal(beta1-4)GlcNAc(beta1-3)Gal(beta1-4)Glc (lacto-N-neotetraose); Fuc(alpha1-2)Gal(beta1-4)GlcNAc(beta1-3)Gal(beta1-4)Glc (lacto-N-fucopentaose IV); Gal(beta1-4)GlcNAc(beta1-3)[Gal(beta1-4)GlcNAc(beta1-6)]Gal(1-4)Glc (lacto-N-neohexaose); Fuc(alpha1-2)Gal(beta1-4)GlcNAc(beta1-3)[Gal(beta1-4)GlcNAc(beta1-6)]Gal(beta1-4)Glc (monofucosyl lacto-N-neohexaose a); Gal(beta1-4)GlcNAc(beta1-3)[Fuc(alpha1-2)Gal(beta1-4)GlcNAc(beta1-6)]Gal(beta1-4)Glc (monofucosyl lacto-N-neohexaose b); Fuc(alpha1-2)Gal(beta1-4)GlcNAc(beta1-3)[Fuc(alpha1-2)Gal(beta1-4)GlcNAc(beta1-6)]Gal(beta1-4)Glc (difucosyl lacto-N-neohexaose); Gal(beta1-4)GlcNAc(beta1-3)Gal(beta1-4)GlcNAc(beta1-3)Gal(beta1-4)Glc (para lacto-N-neohexaose); Fuc(alpha1-2)Gal(beta1-4)GlcNAc(beta1-3)Gal(beta1-4)GlcNAc(beta1-3)Gal(beta1-4)Glc (monofucosyl para lacto-N-neohexaose). Milk of the Australian fur seal, Arctophalus pusillus doriferus (family Otariidae) contained inositol but no lactose or free oligosaccharides. These results, therefore, support the hypothesis that the milk of otariids, unlike that of phocids, contains no free reducing saccharides.

Concentrations and patterns of organochlorine contaminants in white whales (Delphinapterus leucas) from Svalbard, Norway.

Blubber was collected from live-captured, adult male white whales (Delphinapterus leucas) from Svalbard, Norway, and analysed for levels and patterns of organochlorine (OC) contaminants. The OC compounds analysed were HCB, dieldrin, sigmaHCH (alpha-HCH, beta-HCH and gamma-HCH), sigmaChl (heptachlor epoxide, oxychlordane, cis-chlordane, trans-nonachlor, and cis-nonachlor), sigmaDDT (pp'-DDT, pp'-DDE and pp'-DDD) and sigmaPCB (27 PCB congeners). The major OC compounds detected in the blubber were sigmaPCB (5103+/-1874 ng/g l.w.) and sigmaDDT (5108+/-1089 ng/g l.w.), which made up 70% of the sigmaOC. These compounds were followed in prevalence by sigmaChl (2872+/-1177 ng/g l.w.), which contributed 20% of the sigmaOC burden. SigmaHCH, HCB and dieldrin were present, but at low concentrations. This OC pattern is typical of top predators in Arctic marine food chains. OC levels in white whales from Svalbard are lower than white whales from the St Lawrence River in Canada and are generally similar to values reported previously for other Arctic white whale stocks. Some geographic patterns in relative prevalence of various OC compounds appear to be quite consistent among various marine mammal species in the Arctic. PCB and DDT concentrations in Svalbard's white whales are below the levels that are thought to have negative effects on reproduction or the immune system.

Diving development in nursing harbour seal pups.

This study investigated physiological and behavioural aspects of diving development in pups of the harbour seal Phoca vitulina. Behavioural data (4280 h, 6027 dives) from time/depth recorders (N=13) deployed on pups aged 0-19 days are presented concomitantly with physiological measurements (N=8, sampled both early and late in the nursing period) of blood oxygen stores and body composition. Pups grew from 12.6+/-1.8 kg (mean age 2 days, total body fat 16+/-4 %) to 22.2+/-2.5 kg (mean age 16 days, total body fat 35+/-5 %; means +/- S.D.) over the duration of the experiment. Pups less than 5 days of age had an elevated haematocrit and reduced plasma volume compared with older pups. Although plasma volume and blood volume increased, mass-specific blood oxygen stores (total haemoglobin) fell during the study period. Simultaneously, the following behavioural indicators of diving ability increased: the proportion of time spent in the water, dive depth, dive duration, bottom time and maximum daily swimming velocity. In addition, the proportion of dives that were identified by cluster analyses as being U-shaped increased significantly with age. On the basis of the measured blood oxygen stores, less than 1 % of the recorded dives exceeded the calculated aerobic dive limit. Thus, development in blood oxygen stores or rates of oxygen consumption did not seem to restrain the rate of neonatal dive development in harbour seals. It appears that behavioural modifications (experience and learning) may be the primary rate-limiting factors for ontogeny of diving skills in neonates of this species.

Levels and patterns of PCBs and OC pesticides in harbour and grey seals from the St Lawrence Estuary, Canada.

Blubber samples from harbour (Phoca vitulina) and grey seals (Halichoerus grypus) captured in the St Lawrence Estuary were analysed for PCBs and OC pesticides. Concentrations of sigma PCB, sigma DDT, sigma CHLOR and mirex were higher in harbour than in grey seals, while sigma HCH and HCB were similar in the two species. Age vs. concentration plots showed that sigma PCB, sigma DDT and sigma CHLOR concentrations increased with age in males, but plateaued at sexual maturity in females. Concentrations of sigma HCH decreased and mirex increased with age for post-weaning animals, regardless of gender. HCB did not show age-related trends. PCB congener and OC pesticide patterns varied within harbour seals (based on gender and maturity) and between harbour and grey seals. PCB and OC concentrations in harbour seals were lower than those seen in a sample of significantly older beluga whales. Both species remain in the Estuary year-round. PCB and OC pesticide concentrations were higher in resident harbour seals than in either grey or harp seals that visit the Estuary seasonally. PCB and OC pesticide concentrations in harbour seals from the Estuary were generally higher than in harbour seals from other regions of North America. They were comparable to, or higher than those from the NE Atlantic Ocean and the North Sea, and lower than those from the Wadden and Baltic Seas.

Anionic binding site and 2,3-DPG effect in bovine hemoglobin.

It is generally believed that bovine hemoglobin (BvHb) interacts weakly with 2,3-diphosphoglycerate (2,3-DPG) in a chloride-free media and not at all in the presence of physiological concentrations of chloride (100 mM). This lack of interaction has raised several questions at both structural and evolutionary levels. Results obtained in this study via 31P nuclear magnetic resonance (NMR) show that, even in the presence of 100 mM chloride ions, 2,3-DPG does, in fact, interact with bovine deoxy-Hb. This spectroscopic observation has been confirmed by oxygen binding experiments, which have also shown that, under certain conditions, chloride and 2,3-DPG may display a synergistic effect in modifying the oxygen affinity of bovine hemoglobin. It could be that this synergistic effect has its structural basis in a conformational modification induced by 2,3-DPG, possibly causing extra chloride anions to approach the positive charges which constitute the anion binding site. Another possibility, not necessarily an alternative, is the additional chloride binding site recently identified [Fronticelli, C., Sanna, M. T., Perez-Alvarado, G. C., Karavitis, M., Lu, A.-L., and Brinnigar, W. S. (1995) J. Biol. Chem 270, 30588-30592] involving lysine beta76 that in bovine Hb substitutes for the alanine residue present in human hemoglobin. All of these findings are in agreement with the very low enthalpy of oxygenation that characterizes bovine Hb when both chloride and 2,3-DPG are present in concomitance. The results reported here clearly show that bovine hemoglobin does react with 2, 3-DPG and is functionally affected by this organic phosphate. Hence, the very low intraerythrocytic concentration of 2,3-DPG (0.5 mM) in adult bovine red blood cells is the result of metabolic adaptation which cannot be explained solely by the different amino acid sequence at the level of the 2,3-DPG binding site.

Energetics during nursing and early postweaning fasting in hooded seal (Cystophora cristata) pups from the Gulf of St Lawrence, Canada.

In this study we measured growth and milk intake and calculated energy intake and its allocation into metabolism and stored tissue for hooded seal (Cystophora cristata) pups. In addition, we measured mass loss, change in body composition and metabolic rate during the first days of the postweaning fast. The mean body mass of the hooded seal pups (n = 5) at the start of the experiments, when they were new-born, was 24.3 +/- 1.3 kg (SD). They gained an average of 5.9 +/- 1.1. kg.day-1 of which 19% was water, 76% fat and 5% protein. This corresponds to an average daily energy deposition of 179.8 +/- 16.0 MJ. The pups were weaned at an average body mass of 42.5 +/- 1.0 kg 3.1 days after the experiment was initiated. During the first days of the postweaning fast the pups lost an average of 1.3 +/- 0.5 kg of body mass daily, of which 56% was water, 16% fat and 28% protein. During the nursing period the average daily water influx for the pups was 124.6 +/- 25.8 ml.kg-1. The average CO2 .production during this period was 1.10 +/- 0.20 ml.g-1.h-1, which corresponds to a field metabolic rate of 714 +/- 130 kJ. kg-1. day-1, or 5.8 +/- 1.1 times the predicted basal metabolic rate according to Kleiber (1975). During the postweaning fast the average daily water influx was reduced to 16.1 +/- 6.6 ml.kg-1. The average CO2 production in this period was 0.58 +/- 0.17 ml.g-1.h-1 which corresponds to a field metabolic rate of 375 +/- 108 kJ.kg-1.day-1 or 3.2 +/- 0.9 times the predicted basal metabolic rate. Average values for milk composition were 33.5% water, 58.6% fat and 6.2% protein. The pups drank an average of 10.4 +/- 1.8 kg of milk daily, which represents an energy intake of 248.9 +/- 39.1 MJ.day-1. The pups were able to store 73.2 +/- 7.7% of this energy as body tissue.