Water conservation and protein metabolism in northern elephant seal pups during the postweaning fast

Urine production and N output were monitored in northern elephant seal (Mirounga angustirostris) pups progressing through 10 weeks of a natural postweaning fast. Urine output declind by 84% (to 69±12 ml·day^–1) at 10 weeks (P<0.05). Glomerular filtration rate at 10 weeks was 51% of the 67±3 ml serum·min^–1 observed during week 1 (P<0.05). Urine N excretion fell by 69% to 1.2±0.17 g·day^–1, while urinary concentration increased (P<0.05). Serum urea declined from an initial 11 mmol·1^–1 to 5–7 mmol·1^–1 by 5 weeks. The fall in urinary N loss (and thus amino acid oxidation) was concomitant with depressed metabolic rate. Therefore, protein contributed little toward meeting energy demands (i.e., <4% of average metabolic rate) throughout fasting. These data indicate that fasting pups improve water conservation and minimize protein catabolism during prolonged natural fasts without an exogenous source of water.Abbreviations AA amino acid(s) - AMR average metabolic rate - ANOVA one-way analysis of variance - BMR basal metabolic rate - BUN blood urea nitrogen - EP end product - EWL evaporative water loss - [Gr]_s serum creatinine concentration - GFR glomerular filtration rate - LBM lean body mass - LML Long Marine Laboratory - MR metabolic rate - NEFA non-esterified fatty acids - RMR resting metabolic rate - TCA tricarboxylic acid - U:C ulinary urea: creatinine concentration ratio     

Protein catabolism in suckling and fasting northern elephant seal pups (Mirounga angustirostris)

Nursing elephant seal pups are hypothesized to be preadapted to the postweaning fast, yet no comparison of lipid or protein use for meeting metabolic costs has been made between these contrasting nutritional periods. To address this, protein catabolism was estimated in five elephant seal pups from measurements of urea turnover made twice during nursing and twice during the postweaning fast. Changes in body composition were measured in ten separate weaned pups via tritiated water dilution and matched to fasting urea turnover measurements in order to assess errors in protein catabolism derived from urea turnover rates. Estimates of lean mass loss based upon urea turnover and tritiated water dilution were in general agreement, supporting estimates of protein catabolism derived from urea turnover measurements. Protein catabolism was estimated to contribute less than 4% to the average metabolic rate of suckling and fasting pups implying strict protein conservation during both periods and supporting the shypothesis that suckling pups are pre-adapted to fasting. It is proposed that strict protein conservation across suckling and fasting compensates for relative reductions in maternal investment associated with the abbreviated lactation period of the elephant seal.     

Blood viscosity in phocid seals: possible adaptations to diving

1. Mean corpuscular volume (MCV) and mean corpuscular hemoglobin concentration (MCHC) of phocid seal red blood cells (RBC) are elevated compared to those of most terrestrial mammalian species. The influence of these characteristics on blood flow was revealed by viscosity (VIS) measurements. 2. RBC morphology and VIS of whole blood from 7 harbor seals and 5 northern elephant seals were compared with blood of the domestic pig. Samples were analysed for RBC count, white blood cell (WBC) count, total plasma proteins, hematocrit (HCT), MCV and MCHC. Viscosity measurements were made at shear rates from 11.5 to 230.4 s-1 on a Wells-Brookfield cone-plate viscometer at 37 degrees C. 3. Mean values for HCT (%), MCV (micron 3) and MCHC (%) were, respectively: elephant seal: 57, 176, 44; harbour seal: 53, 105, 38; domestic pig: 28, 54, 34. Pig blood was reconstituted to match seal blood HCTs. VIS determinations showed that seal and pig blood conform to the general mammalian dependence of VIS upon shear rate and HCT. 4. Seal blood VIS was 28% (harbour seal) and 16% (elephant seal) less than pig blood VIS at low shear (P less than 0.05). Seal blood carried more hemoglobin per unit volume than did pig blood reconstituted to the same HCT. Fewer, larger RBC with higher MCHC, and hence elevated oxygen storage, accompanied by reduced VIS and reduced flow resistance near stasis suggests that this feature of phocid seal blood is an adaptation to circulatory redistribution during long dives.     

Fatty acid metabolism in fasting elephant seal pups

Summary The turnover of two plasma free fatty acids (FFA) were measured in 5 northern elephant seal pups (Mirounga angustirostris) after approximately 2 months of post-weaning fasting. Turnover was determined using simultaneous bolus injections of¹⁴C-palmitate,³H-oleate and Evans blue (EB) administered via an indwelling extradural intravertebral catheter. At this time in their natural fast, the seals exhibited plasma FFA levels and turnover values higher than reported for other marine mammals and most terrestrial carnivores. There was no consistent difference between plasma FFA turnover measured by palmitate or oleate tracers. The results imply that FFA metabolism is the primary source of energy during fasting. This is interesting in light of previous observations of minimal ketoacid accumulation and low levels of glucose and protein energy production during fasting in these juvenile seals.     

Renal function in suckling and fasting pups of the northern elephant seal.

Elephant seals fast for prolonged periods without access to water. This is made possible, in part, by reductions in urine production. However, the mechanisms involved in reducing urine production are not understood. In this study, glomerular filtration rate (GFR) was measured in five northern elephant seal pups (Mirounga angustirostris) via the inulin clearance technique. Measurements were made during day 9 and day 18-22 of nursing and the second and eighth week of the postweaning fast. Plasma aldosterone and cortisol concentrations, quantified by radioimmunoassay, were measured in eight other weanlings during the second and eighth week of the fast. Mean GFR was 79.3+/-29.3 ml/min during the early suckling period and 78.2+/-17.1, 89.8+/-52.7, and 80.4+/-12.2 ml/min during the late suckling, early fasting and late fasting periods, respectively. Differences between nursing and fasting were insignificant, possibly because reduced protein oxidation during suckling and rapid recruitment of protein for tissue synthesis obviated the need for postprandial hyperfiltration. Alternatively, maintenance of GFR during fasting may facilitate urea concentration by compensating for reductions in the fractional excretion of urea. It is further hypothesized that aldosterone is primarily responsible for mediating renal water reabsorption in this system.     

Energy gain and loss during lactation and postweaning in southern elephant seal pups (Mirounga leonina) at King George Island

Deuterium-labeled water was used to measure changes in the proximate body composition during the lactation period and after weaning in southern elephant seal pups at King George Island, South Shetland Islands, Antarctica. During the lactation period (23.0 ± 1.4 days) pups gained a mean of 4.9 ± 0.5 kg/day (n=7). Of the total mass gain (112 ± 8 kg), 38% was water, 48% was fat, and 11% was protein. This represented an increase in total body gross energy of 2437 ± 145 MJ. The proportion of body mass represented by fat was less than 2% at birth, increasing to 35 ± 2% at weaning. We followed the pups during a mean period of 36 ± 3 days after weaning. During this period, pups had a mean loss of 1.21 ± 0.10 kg/day (n=7) comprising 39% water, 48% fat, and 12% protein. The energy cost over this period was 952 ± 168 MJ, which represented, on average, 39% of the total energy gained during the suckling period. Accepted: 3 January 2000     

Fasting and lactation effect fat-soluble vitamin A and E levels in blood and their distribution in tissue of grey seals (Halichoerus grypus)

Grey seals among other phacoids represent a good model to study the mobilisation, transfer and deposition of fat-soluble components such as vitamins in lactating females and suckling pups because during the lactation period mothers may fast completely while secreting large quantities of high fat milks, and pups deposit large amounts of fat as blubber. The level of vitamins A and E in different tissues (liver, adipose tissue, kidney, heart, skeletal muscle, testis) and blood plasma of adult grey seal females and males changed as a result of fasting and lactation; changes were also observed in pups. The most obvious effects were a significant increase of retinol and a decrease of vitamin E levels in plasma of females with the onset of lactation as well as a substantial decrease in liver vitamin E. In suckling pups both retinol and vitamin E levels in plasma increased with the onset of suckling; after weaning no changes in retinol but a significant decrease in plasma vitamin E was observed. While liver vitamin A levels tended to be unaffected by suckling or post-weaning fast, liver vitamin E levels increased with the uptake of milk substantially (P<0.01) and returned at weaning to low levels similar to that in fetuses. Adipose tissue levels of vitamin A and E in both females and pups were only marginally affected by lactation, suckling or post-weaning fast. Results indicate that both plasma and liver levels of vitamin A and E are affected by the mobilisation, absorption and deposition of these components during lactation in seals to a much greater extent than adipose tissue, from which fat-soluble vitamins are mobilized at rates similar to that of lipids.     

Energy reserve utilization in northern elephant seal (<Emphasis Type="Italic">Mirounga angustirostris</Emphasis>) pups during the postweaning fast: size does matter

During fasting most mammals preferentially utilize lipid reserves for energy while sparing protein reserves. This presents a potential problem for marine mammals that also depend on lipids as a major component of blubber, the primary thermoregulatory structure. Because of this dual function for lipid, rates of lipid and protein utilization should be closely regulated during the postweaning fast in northern elephant seals (Mirounga angustirostris). To quantify energy expenditure during the fast, we measured body mass and composition of 60 pups at 2.3+/-0.2 days and 55.9+/-0.3 days postweaning in 1999 and in 2000. Body condition differed significantly between years. At weaning, body mass (125.9+/-3.8 kg) and percentage lipid content (39.3+/-0.6% of body mass) in 2000 were significantly greater than body mass (115.2+/-3.1 kg) and percentage lipid content (35.8+/-0.6%) in 1999. In general, percentage lipid content increased with body mass, and fatter pups utilized lipid at relatively higher rates during the fast. Lipid fueled 85-95% and 88-98% of energy expended by pups in 1999 and 2000, respectively. Postweaning fast duration (32-78 days) was positively correlated with body mass and hence lipid content at weaning. This suggests that body composition at weaning influences lipid utilization patterns and ultimately the duration of the postweaning fast in northern elephant seal pups.     

Characterization of blubber fatty acid signatures in northern elephant seals (Mirounga angustirostris) over the postweaning fast

Phocids routinely fast for extended periods. During these fasts, energetic requirements are met primarily through the catabolism of blubber lipid. To assess whether fatty acid (FA) composition changes during the postweaning fast in northern elephant seals, blubber biopsies were acquired longitudinally from 43 pups at 2.3 ± 1.5 and 55.2 ± 3.7 days postweaning in 1999 and 2000. At weaning, short-chain monounsaturated FA (SC-MUFA, ≤18 carbons) dominated the blubber while saturated FA (SFA) were found in the next highest proportion. The major FA (all ≥1 % by mass) comprised approximately 91 % of total blubber FA. In both years, 18:1n-9 and 16:0 were the most prevalent FA. Major FA mobilized during the fast consisted of polyunsaturated FA (PUFA), SFA, and SC-MUFA. Long-chain MUFA (>18 carbons) tended to be conserved. The fractional mobilization value of 20:5n-3 was the highest, resulting in significant reductions of this PUFA. Although concentrations of some blubber FA changed significantly during the postweaning fast, the general FA signature of blubber was similar at weaning and near the end of the fast. Changes in some FA differed across years. For example, the concentration of 20:4n-6, a minor PUFA, was significantly reduced in 1999 but not in 2000. FA mobilization patterns in northern elephant seal pups are somewhat similar to those reported previously for other fasting phocids and terrestrial mammals, though there are some notable differences. Differences in FA mobilization patterns across mammalian species may be related to differences in diets, geographical distribution, environmental factors, physiological adaptations, and life history stage.     

Microbiome maturation during a unique developmental window

Shortly after birth, mammals are colonized by a multitude of microbes derived from the mother and the environment. Studies in model organisms have demonstrated that the structure and composition of the gut microbiome of offspring steadily mature with increasing diversity during nursing and weaning (Sommer & Bäckhed, 2013). This period of microbiome assembly is critical for young mammals because the gut microbes they acquire will help train their immune system (Lathrop et al., 2011) with potential long‐lasting effects on their health (Cox et al., 2014). In an article in this issue of Molecular Ecology, Stoffel et al. (2020) investigated the gut microbiota of northern elephant seals (Mirounga angustirostris) during a key developmental window. A month after giving birth, elephant seal mothers stop nursing their pups and return to the sea. As a consequence, their pups go from a diet of milk rich in fat to abruptly enter a post weaning fasting period which lasts for about two months while they remain with the colony. This particular life‐history trait therefore offered the authors a unique and exciting opportunity to evaluate intrinsic factors contributing to gut microbiota development in a wild marine mammal.     

Shifts in specific serum protein concentrations: apolipoprotein-B and albumin in the southern elephant seal during the breeding and molting periods in Antarctica

Serum protein values in Southern elephant seals (Mirounga leonina) were analyzed during the breeding and molting periods at the 25 de Mayo Island (King George Island), Antarctica, during the 1999/2000 and 2000/2001 field seasons, in order to study the reaction of adults to fasting, and of pups to nursing and fasting. The following analyses were carried out: Total Proteins (TP) and Albumin (Alb) were analyzed by a colorimetric technique, and Apolipoprotein B (Apo-B) was determined by the immunodiffusion technique on agarose gel plates. The general ranges of average values for total proteins and specific fractions during the breeding period were: TP (g/dl) = 5.12–9.83; Alb (g/dl) = 1.72–5.71 and Apo-B (mg/dl) = 10–266, and during the post-breeding period: TP = 4.85–9.45; Alb = 2.06–4.20 and Apo-B = 13–232. The essential obtained data were: (a) fasting does not impact adult males, except for a significant decrease of Apo-B during breeding; (b) fasting does not impact adult females, except for a significant decrease in TP during molting; (c) suckling increases significantly TP and Apo-B in pups; (d) post-weaning fast decreases significantly all measured serum components in pups. We can conclude that the adults are adapted to long term fasting, without any metabolic downside they maintain homeostasis during this period, as shown by the serum data. The pups clearly react to both suckling and the post-weaning fast, which are periods driven almost exclusively by lipid chemistry. These impacts may be seen in the chemistry of serum proteins.     

Diving behaviour in relation to water temperature in the southern elephant seal: foraging implications

Summary A time-depth-temperature recorder provided a continuous record of diving by a female southern elephant seal in relation to water temperature for 27 days (1939 dives) after completion of moult. Mean maximum dive depth was 391±2.6 m and the overall maximum was 775 m. Dives lasted on average 17.5±0.09 min. Most dives showed a rapid descent to the discontinuity between the cold surface water and warmer deep water. Consequently the seal spent 57% of its time while diving at a depth of 200–400 m when it may have been foraging. This strongly suggests that the seal was exploiting a food source at the discontinuity between vertically stratified water masses. The water temperature data also indicated that the seal was diving in waters south of the Antarctic Polar Front and at some distance from the northern edge of the pack ice. The seal spent 88% of its time under water. Normal surface intervals between dives lasted an average of 2.1 ± 0.1 min whereas 16 extended surface intervals (>10 min duration) lasted 32.7±4.6 min. Dives were deeper during the day than at night and all but one extended surface interval occurred at night. The pattern of dives was similar to records from northern elephant seals but this is the first study to show how diving behaviour relates to water temperature.     

Prolonged fasting activates hypoxia inducible factors-1α, -2α and -3α in a tissue-specific manner in northern elephant seal pups

Hypoxia inducible factors (HIFs) are important regulators of energy homeostasis and cellular adaptation to low oxygen conditions. Northern elephant seals are naturally adapted to prolonged periods (1–2 months) of food deprivation (fasting) which result in metabolic changes that may activate HIF-1. However, the effects of prolonged fasting on HIFs are not well defined. We obtained the full-length cDNAs of HIF-1α and HIF-2α, and partial cDNA of HIF-3α in northern elephant seal pups. We also measured mRNA and nuclear protein content of HIF-1α, -2α, -3α in muscle and adipose during prolonged fasting (1, 3, 5 & 7 weeks), along with mRNA expression of HIF-mediated genes, LDH and VEGF. HIF-1α, -2α and -3α are 2595, 2852 and 1842 bp and encode proteins of 823, 864 and 586 amino acid residues with conserved domains needed for their function (bHLH and PAS) and regulation (ODD and TAD). HIF-1α and -2α mRNA expression increased 3- to 5-fold after 7 weeks of fasting in adipose and muscle, whereas HIF-3α increased 5-fold after 7 weeks of fasting in adipose. HIF-2α protein expression was detected in nuclear fractions from adipose and muscle, increasing approximately 2-fold, respectively with fasting. Expression of VEGF increased 3-fold after 7 weeks in adipose and muscle, whereas LDH mRNA expression increased 12-fold after 7 weeks in adipose. While the 3 HIFα genes are expressed in muscle and adipose, only HIF-2α protein was detectable in the nucleus suggesting that HIF-2α may contribute more significantly in the up-regulation of genes involved in the metabolic adaptation during fasting in the elephant seal.     

Changes in plasma fatty acids indicate change in nutritional status in developing Weddell seal pups

The concentrations and the fatty acid compositions (weight percent) of three plasma lipid fractions (phospholipids, cholesteryl esters, and triacylglycerols) were examined in five Weddell seal pups through suckling, the post-weaning fast and periods of independent foraging. The principal plasma fatty acids in Weddell seal pups during suckling were the same as those previously reported for milk samples collected from lactating Weddell seals. In contrast to the stable milk fatty acid composition suggested by the previous study, significant changes in the fatty acid composition of plasma lipids occurred during suckling. These included an increase in the weight% of 20:4(n-6) and 18:0 in phospholipids, an increase of 20:4(n-6) in cholesteryl esters, and an increase of 20:1(n-9) and a decrease of 20:5(n-3) in triacylglycerols. Weaning and the subsequent fasting period were accompanied by a dramatic drop in the plasma concentration and 14:0 content of triacylglycerols and by marked decrease and increase in the weight% of 20:1(n-9) and 20:5(n-3), respectively, in this lipid fraction. Weight% of 14:0 in triacylglycerols and/or plasma concentration of triacylglycerols clearly distinguish post-weaning from suckling. However, fasting versus foraging pups could not be clearly distinguished. Received: 7 October 1996 / Accepted: 20 May 1997     

Age, sex, and reproductive state influence free amino acid concentrations in the fasting elephant seal

Long-term fasting is a component of northern elephant seal (Mirounga angustirostris) life history requiring physiological adaptations to nitrogen conservation. Plasma free amino acids (FAAs) were determined for five elephant seal pups during the second and eighth weeks of the postweaning fast, six lactating female seals at 4-6 and 25 d postpartum, and seven sexually competitive adult male seals taken midway through the breeding season. Total FAAs declined in lactating females (11%) and pups (30%) with time fasting, but cystine concentration more than doubled in pups while decreasing by approximately 43% in lactating females. Methionine concentration significantly increased (approximately 68%) across lactation in adult females but was low for all classes of seal. Alanine was the most abundant FAA in adult males, and glycine became the dominant FAA in adult females late in lactation. Glutamine dominated the FAAs of weaned pups across the fast. Reductions in total FAAs of weanlings mirrored reductions in protein catabolism, but reductions in total FAAs also occurred in lactating females concomitant with an increase in protein catabolism. Observed variation in FAA concentrations may reflect ontogenetic requirements for certain amino acids in fasting weanlings. Similarly, increases in specific FAA concentrations across lactation may reflect variations in FAA flux resulting from the nutrient demands of lactogenesis.     

Early sexual dimorphism in the developing gut microbiome of northern elephant seals

The gut microbiome is an integral part of a species' ecology, but we know little about how host characteristics impact its development in wild populations. Here, we explored the role of such intrinsic factors in shaping the gut microbiome of northern elephant seals (Mirounga angustirostris) during a critical developmental window of 6 weeks after weaning, when the pups stay ashore without feeding. We found substantial sex differences in the early‐life gut microbiome, even though males and females could not yet be distinguished morphologically. Sex and age both explained around 15% of the variation in gut microbial beta diversity, while microbial communities sampled from the same individual showed high levels of similarity across time, explaining another 40% of the variation. Only a small proportion of the variation in beta diversity was explained by health status, assessed by full blood counts, but clinically healthy individuals had a greater microbial alpha diversity than their clinically abnormal peers. Across the post‐weaning period, the northern elephant seal gut microbiome was highly dynamic. We found evidence for several colonization and extinction events as well as a decline in Bacteroides and an increase in Prevotella, a pattern that has previously been associated with the transition from nursing to solid food. Lastly, we show that genetic relatedness was correlated with gut microbiome similarity in males but not females, again reflecting early sex differences. Our study represents a naturally diet‐controlled and longitudinal investigation of how intrinsic factors shape the early gut microbiome in a species with extreme sex differences in morphology and life history.     

Enhancement of antibody synthesis in rats by feeding cis-9,trans-11 conjugated linoleic acid during early life

Previous studies have demonstrated that the intake of a 1% conjugated linoleic acid (CLA) diet in an 80:20 mixture of cis-9,trans-11 and trans-10,cis-12 exerts age-specific effects on the immune system: immunoglobulin enhancement and proliferative down-modulation in neonatal and adult rats, respectively. The present study evaluates the influence of the same diet on antibody synthesis of early infant Wistar rats during suckling and/or after weaning. Dietary supplementation was performed during suckling and early infancy (4 weeks), only during suckling (3 weeks), or only in early infancy (1 week). CLA content in plasma and serum immunoglobulin (Ig) G, IgM and IgA concentration were determined. Proliferation, cytokines and Ig production were evaluated on isolated splenocytes. Cis-9,trans-11- and trans-10,cis-12-CLA isomers were detected in the plasma of all CLA-supplemented animals, and the highest content was quantified in those rats supplemented over the longest period. These rats also exhibited higher concentrations of serum IgG, IgM and IgA. Moreover, splenocytes from CLA-supplemented rats showed the highest IgM and IgG synthesis and interleukin (IL)-6 production, whereas their proliferative ability was lower. In summary, in infant rats, we observed both the enhance antibody synthesis previously reported in neonates, and the reduced lymphoproliferation previously reported in adults.     

GLUCOSE KINETICS IN NEONATAL ELEPHANT SEALS DURING POSTWEANING APHAGIA

Northern elephant seal ( Mirounga angustirostris )pups undergo extended periods of terrestrial aphagia after weaning and exhibit a paradoxical fasting hyperglycemia. To investigate the details of glucose metabolism during this period, reversible and irreversible radiotracers were used to determine the body mass of glucose, and rates of glucose turnover, recycling, and oxidation in fasting seal pups. A typical 75 kg pup has a glucose mass of about 4.5 g (60 mg/kg), and a blood glucose concentration of about 174 mg/dl. Blood glucose removal rate was about 30 grams per day (17 mg/kg · h^-1), but less than 2.5% of this glucose was oxidized, contributing less than 1% of the total metabolic rate. About 20% of the glucose pool was removed from the blood per hour, yielding a turnover time in the vascular space of about five hours. Most glucose removed from the blood was returned to the blood by recycling. Such recycling may contribute to mechanisms which prolong survival during fasting, such as high rates of triacyclglycerol turnover, synthesis of new protein pools, low ketone levels, and the Cori cycle which is important during diving.     

A profile of carbohydrate metabolites in the fasting northern elephant seal

Northern elephant seals endure prolonged periods of food deprivation at multiple life-history stages and simultaneous with energetically costly activities—including reproduction and development. Most mammals decrease their energy expenditure while fasting, with simultaneous reductions in gluconeogenesis and circulating glucose concentration. Paradoxically, elephant seals maintain high rates of both energy expenditure and gluconeogenesis, and high blood glucose concentrations throughout fasting. We therefore characterized the suite of changes that occur in carbohydrate metabolites during fasting in northern elephant seals. Using a broad-based metabolomics platform we investigated fasting during two states—lactation in adult females and the post-weaning developmental period in pups. A total of 227 metabolites were detected in seal plasma; 31 associated with carbohydrate metabolism were analyzed in the present study. Several compounds showed similar responses during lactation and the post-weaning fast (e.g. glycerol and mesaconate) whereas other compounds displayed quite different abundances between groups (e.g. citrate and pyruvate). This work found that, while the changes that occur with fasting were frequently similar in lactating females and developing pups, the relative abundance of compounds often varied markedly. These differences suggest that the metabolic strategies used to endure prolonged fasts are influenced by life-history or nutrient constraints.     

Body condition at weaning affects the duration of the postweaning fast in gray seal pups (Halichoerus grypus)

Gray seals (Halichoerus grypus) undergo a terrestrial postweaning fast (PWF) that depletes energy reserves acquired during the suckling interval. Plasticity in PWF duration may ensure that pups of variable body condition depart for sea with adequate energy reserves. To test this hypothesis, we examined body condition of 30 gray seal pups at weaning and monitored their PWF duration. On average, fat accounted for 47.3% +/- 0.7% of their 53.2 +/- 1.3-kg weaning mass. Although fasting duration averaged 21 +/- 1.1 d (n = p28), there was considerable variation in fasting duration (9 to > 31 d) and the resulting age when pups departed to sea (26 to > 49 d). Percent fat at weaning(38.6%-54.6%) was positively correlated with fasting duration(n = 28, r = 0.376, P = 0.0489). In contrast, total body gross energy (735.3-1,447.4 MJ) and body mass (39.0-66.0 kg) were not correlated with fasting duration. Thus, body composition,not overall body reserves, predicted fasting duration, but the effect was weak, indicating that other factors also account for the observed variation in fasting duration. We speculate that pups with greater percent fat more effectively utilized lipid and conserved protein while meeting metabolic costs throughout the PWF. As a result, fatter pups extended the PWF duration,which may be critical for development of diving physiology and may have facilitated their survivorship to age 1.