Milk intake,growth and energy consumption in pups of ice-breeding grey seals (Halichoerus grypus) from the Gulf of St. Lawrence,Canada

In this study we document growth, milk intake and energy consumption in nursing pups of icebreeding grey seals (Halichoerus grypus). Change in body composition of the pups, change in milk composition as lactation progresses, and mass transfer efficiency between nursing mothers and pups are also measured. Mass transfer efficiency between mother-pup pairs (n=8) was 42.5±8.4%. Pups were gaining a daily average of 2.0±0.7 kg (n=12), of which 75% was fat, 3% protein and 22% water. The total water influx was measured to be 43.23±8.07 ml·kg^-1·day^-1. Average CO₂ production was 0.85±0.20 ml·g^-1·h^-1, which corresponds to a field metabolic rate of 0.55±0.13 MJ·kg^-1·day^-1, or 4.5±0.9 times the predicted basal metabolic rate based on body size (Kleiber 1975). Water and fat content in the milk changed dramatically as lacation progressed. At day 2 of nursing, fat and water content were 39.5±1.9% and 47.3±1.5%, respectively, while the corresponding figures for day 15 were 59.6±3.6% fat and 28.4±2.6% water. Protein content of the milk remained relatively stable during the lactation period with a value of 11.0±0.8% at day 2 and 10.4±0.3% at day 15. Pups drank an average of 3.5±0.9 kg of milk daily, corresponding to a milk intake of 1.75 kg per kg body mass gained. The average daily energy intake of pups was 82.58±19.80 MJ, while the energy built up daily in the tissue averaged 61.72±22.22 MJ. Thus, pups assimilated 74.7% of the energy they received via milk into body tissue. The lactation energetics of ice-breeding grey seals is very similar to that of their land-breeding counterparts.Abbreviations bm body mass - BMR basal metabolic rate - FMR field metabolic rate - IU international unit - RQ respiration quotient - HTO tritiated water - HT¹⁸O doubly labeled water - TBW total body water - VHF very high frequency     

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     

Energy intake and utilisation by nursing bearded seal (Erignathus barbatus) pups from Svalbard,Norway

In this study we measure energy intake via milk in nursing bearded seal (Erignathus barbatus) pups and determine how this energy is allocated into metabolism and storage of new tissues. This was accomplished using longitudinal mass gain records and the doubly labelled water technique on nursing pups in combination with cross-sectional data on changes in milk composition from bearded seal mothers. The pups (n=3) were all less than a week old at the start of the experiments. Pups gained 3.3±0.4 kg·day^-1 of which 50% was fat, 14% protein and 36% water. Average daily water influx for the pups was 69.5±9.0 ml · kg^-1· day^-1. Average CO₂ production during the study period was 0.99±0.10 ml·g^-1·h^-1, which corresponds to a field metabolic rate of 642±67 kJ·kg^-1· day^-1, or 6.0±0.5 times the predicted basal metabolic rate according to Kleiber (1975). The pups drank an average of 7.6±0.5 kg of milk daily. This corresponds to a daily energy intake of 154±8 MJ, 47±14% of which was stored as new body tissue. Despite this high energy intake bearded seal pups do not get as fat as do other nursing phocids. This is in part due to their larger body size but also due to their very active aquatic lifestyle and the lower and more consistent fat content of the milk compared to other phocid species. Bearded seal mothers forage during lactation and may also be involved in teaching their pups to feed independently. All these data suggest that the lactation strategy of bearded seals differs from the phocid norm.     

Fat transfer and energetics during lactation in the hooded seal: the roles of tissue lipoprotein lipase in milk fat secretion and pup blubber deposition

Hooded seals (Cystophora cristata) lactate for 3.6 days during which females simultaneously fast and transfer large amounts of energy to their pups through fat-rich milk. Pups grow rapidly, principally due to blubber deposition. Lipoprotein lipase (LPL), the primary enzyme responsible for tissue uptake of triglyceride fatty acids, may strongly influence both maternal milk fat secretion and pup blubber deposition. We measured the energetic costs of lactation (using hydrogen isotope dilution, ³H₂0), milk composition, prolactin, and LPL activity (post-heparin plasma LPL [PH LPL], blubber, mammary gland and milk; U) in six females. PH LPL and blubber LPL were measured in their pups. Females depleted 216.3 MJ · day⁻¹ of body energy and fat accounted for 59% of maternal mass loss and 90% of postpartum body energy loss, but maternal body composition changed little. Maternal blubber LPL was negligible (0.0–0.2 U), while mammary LPL was elevated (1.8–2.5 U) and was paralleled by changes in prolactin. Estimated total mammary LPL activity was high (up to 20,000 U · animal⁻¹) effectively favoring the mammary gland for lipid uptake. Levels of total blubber LPL in pups increased seven-fold over lactation. Pups with higher PH LPL at birth had greater relative growth rates (P = 0.025). Pups with greater blubber stores and total blubber LPL activity had elevated rates of fat deposition (P = 0.035). Accepted: 4 May 1999     

Mass transfer from mothers to pups and mass recovery by mothers during the post-breeding foraging period in southern elephant seals (Mirounga leonina) at King George Island

Mass transfer from mother to pup during the lactation period, and mass recovery for the same females during the foraging period were measured in the southern elephant seal at King George Island, Antarctica. During the 19.2 ± 0.9-day lactation period measured (which represented 87% of the entire nursing), females lost a mean mass of 10.56 ± 1.76 kg/day (n = 27), while their pups gained a mean mass of 5.27 ± 1.1 kg/day. There was a correlation between daily body weight gain in pups and daily weight loss by their mothers. Pup weaning mass was positively related to maternal post-partum mass. Serial samples showed that weight losses by females and gains by their pups were not linear over lactation, but showed lower values at the beginning and at the end of lactation. During the 60.5 ± 6.2-day foraging phase between the end of lactation and molt, females gained 2.21 ± 0.65 kg/day (n = 12), or 54% of the mass lost during nursing. Growth rates reported here are higher than those reported in other breeding sites. However, the ratio of body mass loss by females to gain by their pups was similar, suggesting that higher growth rates and greater weaning mass at South Shetland are due to a higher mean weight of females on arrival at this breeding site. The foraging period was shorter and the mass gained greater than those measured at South Georgia; this could be related to relatively shorter distances to foraging areas. Received: 20 September 1996 / Accepted: 28 April 1997     

Lipoprotein lipase activity and its relationship to high milk fat transfer during lactation in grey seals

Lipoprotein lipase regulates the hydrolysis of circulating triglyceride and the uptake of fatty acids by most tissues, including the mammary gland and adipose tissue. Thus, lipoprotein lipase is critical for the uptake and secretion of the long-chain fatty acids in milk and for the assimilation of a high-fat milk diet by suckling young. In the lactating female, lipoprotein lipase appears to be regulated such that levels in adipose tissue are almost completely depressed while those in the mammary gland are high. Thus, circulating fatty acids are directed to the mammary gland for milk fat production. Phocid seals serve as excellent models in the study of lipoprotein lipase and fat transfer during lactation because mothers may fast completely while secreting large quantities of high fat milks and pups deposit large amounts of fat as blubber. We measured pup body composition and milk fat intake by isotope (deuterium oxide) dilution and plasma post-heparin lipoprotein lipase activity in six grey seal (Halichoerus grypus) mother-pup pairs at birth and again late in the 16-day laction period. Maternal post-heparin lipoprotein lipase activity increased by an average of four-fold by late lactation (P=0.027), which paralleled an increase in milk fat concentration (from 38 to 56%; P=0.043). Increasing lipoprotein lipase activity was correlated with increasing milk fat output (1.3–2.1 kg fat per day) over lactation (P=0.019). Maternal plasma triglyceride (during fasting) was inversely correlated to lipoprotein lipase activity (P=0.027) and may be associated with the direct incorporation of longchain fatty acids from blubber into milk. In pups, post-heparin lipoprotein lipase activity was already high at birth and increased as total body fat content (P=0.028) and the ratio of body fat: protein incrased (P=0.036) during lactation. Although pup plasma triglyceride increased with increasing daily milk fat intake (P=0.023), pups effectively cleared lipid from the circulation and deposited 70% of milk fat consumed throughout lactation. Lipoprotein lipase may play an important role in the mechanisms involved with the extraordinary rates of fat transfer in phocid seals.Abbreviations FFA free fatty acid - HL hepatic lipase - LPL lipoprotein lipase - PH-HL post-heparin hepatic lipase - PH-LPL post-heparin lipoprotein lipase - VLDL very low density lipoprotein     

Milk composition and lactational output in the greater spear-nosed bat, Phyllostomus hastatus

Growth rates of mammalian young are closely linked to the ability of the mother to provide nutrients; thus, milk composition and yield provide a direct measure of maternal investment during lactation in many mammals. We studied changes in milk composition and output throughout lactation in a free-ranging population of the omnivorous bat, Phyllostomus hastatus. Fat and dry matter of milk increased from 9 to 21% and from 21 to 35% of wet mass, respectively, throughout lactation. Energy increased from 6 to 9 kJ · g⁻¹ wet mass, primarily due to the increase in fat concentration. Total sugar levels decreased slightly but non-significantly. Mean sugar level was 4.0% of wet mass. Protein concentration increased from 6 to 11% of wet mass at peak lactation and then decreased as pups approached weaning age. Total milk energy output until pups began to forage was 3609 kJ. Milk levels of Mg, Fe, Ca, K, and Na averaged 0.55 ± 0.26, 0.23 ± 0.2, 8.75 ± 4.17, 5.42 ± 2.11, and 9.87 ± 4.3 mg · g⁻¹ dry matter, respectively. Of the minerals studied, calcium appears to be most limiting in this species. The high degree of variability in foraging time, milk composition and milk yield between individuals at the same stage of lactation could potentially yield high variance in reproductive success among females of this polygynous species. Accepted: 23 January 1997     

Postweaning duration and body composition changes in Southern elephant seal (Mirounga leonina) pups at King George Island.

Weaning mass in southern elephant seals is highly variable, the heaviest pups being three times as heavy as the lightest ones. After weaning, pups undergo an extensive postweaning period in which they draw on their reserves. To quantify the energy expenditure during the postweaning period, changes in mass, body composition, and postweaning duration were measured in southern elephant seals at King George Island, South Shetland Islands, Antarctica. Overall, mean pup weaning mass was 154 +/- 26 kg (n=117) and did not differ between sexes. Mean minimum postweaning duration was 42.5 +/- 7.5 d. Heavier animals at weaning had lower mass-specific mass loss rates than lighter ones, and a faster depletion of body reserves was associated with a shorter postweaning period. The proportion of body mass represented by fat at weaning was 37% +/- 4% (n=47) and did not differ between sexes. Of these pups, 36 were recaptured after a mean period of 36 d after weaning. On average, total mass loss measured in these animals (39 kg) was composed of 39% water, 47% fat, and 12% protein. The composition of mass loss was not significantly different between sexes and was not related to weaning mass or total body energy reserves. However, fatter animals at weaning lost more fat per kilogram lost than thinner ones. Late in the fast, males and females appeared to be in a similar body condition. Nevertheless, the overall proportion of body mass represented by fat at this time was lower than that presented by the same animals at weaning. We estimated that during the postweaning period pups lost, on average, 30% of their mass at weaning. This comprised approximately 35% of the energy and 32% of the fat in the pup's body.     

Lactation costs in southern elephant seals at King George Island,South Shetland Islands

Labelled-water methodology was used to quantify energy costs and energy transfer efficiency in 18 mother-pup pairs of southern elephant seals (Mirounga leonina) during lactation. During the lactation period, mothers lost a mean mass of 227±47 kg. Mass loss included 22% of the protein, 60% of the fat, and 51% of the energy in the mothers body upon arrival. Total body-energy reserves at parturition explained 69% of the variation in the total lactation costs and 50% of the variation in the pups body energy at weaning. On average, pups retained 48% of the mass, 49% of protein, 53% of fat and 51% of energy lost by their mothers. Greater, fatter females showed a decrease in the efficiency of energy and fat transfer and, at the same time, an increase in the efficiency of protein transfer. This may be due to an increased use of protein as metabolic fuel, as fat demands for milk production increase. There was no evidence that greater total lactation costs influence the ability of mothers to produce a pup in the next breeding season.     

Milk production and composition in reindeer (Rangifer tarandus): effect of lactational stage

Milk yield and composition of major milk constituents were measured in captive, nursing reindeer. Registration of milk production was performed during two successive lactations (2001 and 2002). The milk yield was significantly affected by week of lactation (P<0.001) and by individual (P<0.001). The lactation curve had an asymmetrical peak 3 weeks postpartum and the milk yield at peak lactation was 983 g/day (range 595-1239). The length of lactation varied from 24 to 26 weeks and average total milk production was 99.5 kg. From peak lactation the milk production decreased linearly (P<0.001) until milk production was terminated. Mean values for content of major milk constituents were 15.5% fat, 9.9% protein and 2.5% lactose. The content of fat and protein increased markedly with the lactation stage (P<0.001), while lactose showed a slight decrease (P<0.001). The milk composition was significantly affected by stage of lactation (P<0.001). There was a marginally significant decrease in protein:fat ratio (P=0.06) as protein was substituted by fat with stage of lactation. The caloric value of the milk averaged 8.7 kJ/g and increased significantly with the stage of lactation (P<0.001). The overall increase in milk gross energy content during lactation was 67.6%. The energy output averaged 7996 kJ/day at peak lactation and decreased significantly during the course of lactation (P=0.002).     

Variation in milk production and lactation performance in grey seals and consequences for pup growth and weaning characteristics

Phocid seals are one of the few groups of mammals capable of sustaining the energetic demands of lactation entirely through body nutrient stores while fasting. Lactation performance of the female in turn influences the rate and pattern of pup growth. We examined variation in and patterns of milk composition and production, maternal energy output, and pup growth and energy deposition over the entire lactation period in 18 grey seal mother-pup pairs using hydrogen isotope (3H2O and D2O) dilution. Milk composition was independent of maternal mass and nutrient stores, indicating dependence on other physiological and genetic factors. Heavier females lactated longer (r2=0.653, P<0.001), had higher total milk outputs (r2=0.652, P<0.001), and produced larger pups at weaning (r2=0.417, P=0.005). While fatter females lactated for longer periods of time (r2=0.595, P<0.001), females with a larger lean body mass at parturition produced more milk (r2=0.579, P<0.001). Total milk energy output was the strongest predictor of pup weaning mass, which, along with the pup's efficiency of energy storage, accounted for 91% of the variation in weaning mass. Nevertheless, there was sufficient plasticity in milk composition and energy output that some smaller females produced relatively large pups. Few females appeared to deplete body nutrients to the point where it might limit the duration of lactation.     

Southern elephant seals: IgM concentration in milk of cows and serum of their pups

Serum and milk Immunoglobulin M (IgM) concentrations in 11 mother-pup pairs were measured in southern elephant seals (Mirounga leonina) throughout lactation during 2 breeding seasons at King George Island. Samples were obtained sequentially throughout the suckling period (approximately 23 days). The IgM concentration was measured by single radial immunodiffusion on agarose plates. Milk IgM concentrations showed significant differences throughout lactation, with the highest concentrations on the 1st day (x=989.7 mg/dL skimmed milk; SD=433.2) followed by a sharp fall during the next 3–6 days of the suckling period. The ratio of milk IgM/serum IgM concentrations from mothers ranged from 0.21 to 21.92, with highest values in the 1st day of lactation (x=8.25, SD=5.4) and a decrease in concentration as lactation progressed. This was due to the fact that, throughout lactation, milk IgM concentrations fell while serum IgM values showed an increasing trend. Pups showed the lowest serum IgM values in the 1st day of the suckling period (x=13.0 mg/dL, SD=4.3) with an increasing trend as lactation progressed. Despite the high IgM concentrations of milk at day 1 of lactation, by 1 week of age pups had serum IgM concentrations only slightly greater than at birth. This suggests that much of this Ig was newly formed and little or no milk IgM was absorbed during the 1st week. Possibly, the function of ingested IgM is to provide local immunity in the pup's gut, during the first few days of postnatal life. Accepted: 26 March 2000     

Milking strategy in subantarctic fur seals Arctocephalus tropicalis breeding on Amsterdam Island: evidence from changes in milk composition.

Milk composition was investigated throughout the 10-mo pup-rearing period in subantarctic fur seals (Arctocephalus tropicalis) breeding on Amsterdam Island. The mean milk composition was 42.8% +/- 5.7% lipid, 12.1% +/- 1.5% protein, and 42.6% +/- 7.3% water. Subantarctic fur seals breeding on Amsterdam Island produced one of the richest milks ever reported in otariids (20.4 +/- 2.9 kJ/g), with lipid content contributing 85% of total gross energy. The high lipid levels measured in the milk of subantarctic fur seals breeding on Amsterdam Island is consistent (i) with the relatively long time lactating females spend at sea, due to the relatively poor local trophic conditions near the colony that necessitate that they travel long distances to reach the foraging grounds, and (ii) with the consequently short time mothers spend with their pups ashore. Milk composition changed according to the time mothers were fasting ashore: milk produced during the first 2 d spent ashore, when more than 80% of milk transfer occurred, had higher levels of lipids, proteins, and gross energy than milk produced later during the visit ashore, suggesting that the pups were fed with two types of milk during a suckling period. Throughout the year, mothers in good condition produced milk of higher lipid content than others, suggesting that individual foraging skills contribute to enhance milk quality. Milk lipid and gross energy content varied with pup age, according to quadratic relationships, increasing during the earlier stages of lactation before reaching asymptotic values when pups were 180 d old. The stage of lactation appears to be a better predictor of milk lipid content than the duration of the preceding foraging trip, suggesting that either changes in the nutritional requirements of the pup and/or seasonal changes in trophic conditions act on milk composition. These changes in milk quality may also be related to changes in maternal care; lactating subantarctic fur seals apparently reallocate their body reserves toward gestation rather than lactation at the end of the pup-rearing period.     

Energy transfer and female condition in nursing harp seals Phoca groenlandica

Lactating female harp seals, most with their pups, were collected on the "whelping" ice in the Gulf of St. Lawrence in 1976 and from 1978 to 1980. During lactation females lost weight at an average rate of 3.17±0.52 (SE_b) kg d⁻¹, for a total energy loss of approximately 250000 kcal. Pups grew at a mean rate of 2.78±0.19 kg d⁻¹ for a gain in production energy of about 194000 kcal. Compared to 1976, adult females sampled in 1978 to 1980 had lower energy reserves at the onset of lactation. Coincidentally there has been a decrease in newborn lengths and girths. Although compensatory growth in weight appears to occur, between-year differences in newborn lengths persist to weaning. Various reproductive strategies of female harp seals faced with reduced energy stores are discussed.     

Production and composition of Iberian sow's milk and use of milk nutrients by the suckling Iberian piglet

Sixteen purebred Iberian (IB) sows were used in two consecutive trials to determine the efficiency of conversion of sow's milk into piglet body weight (BW) gain and the relationship between milk protein and body protein retention and between milk energy yield and body energy retention in the nursing IB piglet. In each trial, four sows were selected in order to evaluate their milk production, litter growth and nutrient balance measurements, together with four additional sows for milk sampling. Litter size was equalized to six piglets. Daily milk yield (MY) was determined weekly by the weigh-suckle-weigh technique over a 34-day lactation period. Piglets were weighed individually at birth and then weekly from day 5 of lactation. Milk samples were collected on days 5, 12, 19, 26 and 34 post partum. The comparative slaughter procedure was used to determine piglet nutrient and energy retention. One piglet from each litter was slaughtered at birth and four on the morning of day 35. Total MY was on average 5.175 ± 0.157 kg/day. The average chemical composition (g/kg) of the milk was 179 ± 4 dry matter, 53.4 ± 1.0 CP, 58.5 ± 3.8 fat, 10.4 ± 0.3 ash and 56.9 ± 2.3 lactose. Milk gross energy (GE) was 4.626 ± 0.145 MJ/kg. Milk intake per piglet tended to increase in trial 2 (832 v. 893 g/day; P = 0.066). Piglet BW gain contained (g/kg) 172.1 ± 1.3 protein, 151.5 ± 3.5 fat, 41.4 ± 0.6 ash and 635 ± 3 water and 10.127 ± 0.126 MJ GE/kg. Throughout the 34-day nursing period, the piglets grew at an average rate of 168 ± 3 g/day. The ratio of daily piglet BW gain to daily MY was 0.195 ± 0.002 g/g and the gain per MJ milk GE intake was 41.9 ± 0.5 g/MJ. The overall efficiency of protein accretion (g CP gain/g CP milk intake) was low and declined in trial 2 (0.619 v. 0.571; P = 0.016). Nutrient and energy deposition between birth and weaning were 27.4 ± 0.5 g/day protein, 24.2 ± 0.8 g/day fat and 1615 ± 40 kJ/day energy. Piglet energy requirements for maintenance were 404 kJ metabolizable energy (ME)/kg BW0.75. ME was used for growth with a net efficiency of 0.584. These results suggest that poor efficiency in the use of sow's milk nutrients rather than a shortage in milk nutrient supply might explain the low growth rate of the suckling IB piglet.     

Experimental evidence of seawater drinking in juvenile hooded (Cystophora cristata) and harp seals (Phoca groenlandica)

This study was undertaken to measure whether young harp seals (Phoca groenlandica) and hooded seals (Cystophora cristata) drink seawater and, if so, to investigate how the excess salt load is handled. Blood and urine samples were collected from hooded seal pups (n=3) and harp seal pups (n=3) after 2 weeks of freshwater exposure, at intervals during 3 weeks of seawater exposure and, finally, after 2 weeks of re-exposure to fresh water. Total water turnover, as measured by injection of tritiated water, was 2200 ml · day⁻¹ and 3300 ml · day⁻¹ in hooded seals and harp seals, respectively. The extent of mariposia was taken as the difference between total water turnover and influx of water through food (free and metabolic water) and respiratory water exchange. Seawater drinking amounted to 14% and 27% of total water turnover (r_H2O) for the hooded seals and harp seals, respectively. Further evidence of mariposia was obtained from an increase in the excretion rate of the urine osmolytes Na⁺, Cl⁻ and Mg²⁺, during the period of seawater exposure. It is concluded that water influx due to seawater drinking can not be excluded as a source of error when estimating food consumption of free-ranging harp seals and hooded seals, by use of labeled water techniques. Accepted: 11 May 2000     

Seasonal variation in energy intake and condition of harp seals: Is there a harp seal morph? Problems for bioenergetic modelling

Seasonal changes in mass, fat depth, condition index and energy intake were measured in eight captive harp seals (Phoca groenlandica) over a 15-month period. Two of the five adult females were pregnant but lost their pups before term. There were large shifts in all variables measured. The range of these changes in the individual adult animals were for mass, 15–89%, blubber depth 56–240%, and condition index 13–23% (girth x 100)/length. Daily energy consumption varied from as little as 1.16 MJ to as much as 70.97 MJ. The two young males showed larger seasonal changes than the adults. Consistent with our earlier studies of harbour seals (Phoca vitulina) , there was either a significantly negative or no correlation between energy intake and changes in mass or fat. Mass and blubber thickness were positively related in the male seals, however, this was true of the females only during spring and summer. Water and air temperature varied indirectly with changes in blubber thickness in all but two seals. It is proposed that metabolic rate varies seasonally in seals. The implications of these findings are discussed in the light of their impact on bioenergetic modelling.     

Mass transfer efficiency between harp seal (Phoca groenlandica) mothers and their pups during lactation

We investigated the efficiency of mass transfer in lactating harp seals through serial measurements on individual mother-pup pairs during the whelping seasons of 1988 and 1989. We also compared the influence of longitudinal versus cross-sectional sampling on estimates of the efficiency of mass transfer. Among longitudinally sampled pairs, pups grew at an average rate of 2·3 ± 0·5 (mean ± S.D.) kg/d (N = 20). The concomitant mass loss by females averaged 3·1 ± 0·8 kg/d (N = 19). The mean efficiency of mass transfer was 77·0 ± 13·6% (N= 19 pairs).
Estimates of pup growth and female mass loss from regressions of cross-sectional data were 2·0 kg/d and 3·1 kg/d, respectively. These values produce an estimate of 65% for the efficiency of mass transfer.
Consistent with the high efficiency of mass transfer, harp seal females contribute less of their total body mass to nursing ( c. 28%) than most other phocids examined. The resulting energy savings may be important for females of an ice-breeding species, which migrate a long distance shortly after weaning their pups     

Feast or famine: evidence for mixed capital–income breeding strategies in Weddell seals

Evolved patterns of resource expenditure for reproduction have resulted in a life history continuum across species. A strictly capital-breeding strategy relies extensively on stored energy for reproduction, whereas income breeding uses energy acquired throughout the reproductive period. However, facultative income breeding has been shown in some classically capital-breeding animals, and was originally thought to provide a nutritional refuge for smaller females incapable of securing sufficient reserves during pre-partum foraging. We examined milk composition and milk output for the Weddell seal to determine to what degree lactation was aided by food intake, and what factors contributed to its manifestation. Milk composition was independent of maternal post-partum mass and condition, but did change over lactation. Changes were most likely in response to energetic and nutritional demands of the pup at different stages of development. During early lactation, females fasted and devoted 54.9% of total energy loss to milk production. Later in lactation 30.5% more energy was devoted to milk production and evidence suggested that larger females fed more during lactation than smaller females. It appears that Weddell seals may exhibit a flexible strategy to adjust reproductive investment to local resource levels by taking advantage of periods when prey are occasionally abundant, although it is restricted to larger females possessing the physiological capacity to dive for longer and exploit different resources during lactation. This supports the assumption that although body mass and phylogenetic history explain most of the variation in lactation patterns (20–69%), the remaining variation has likely resulted from physiological adaptations to local environmental conditions. Our study confirms that Weddell seals use a mixed capital–income breeding strategy, and that considerable intraspecific variation exists. Questions remain as to the amount of energy gain derived from the income strategy, and the consequences for pup condition and survival. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Breeding biology of weddell seals (Leptonychotes weddellii) at Drescher Inlet,riiser larsen ice shelf,Antarctica

Summary The breeding biology of Weddell seals (Leptonychotes weddellii) was studied at Drescher Inlet (Antarctica) during the austral late winter and spring 1986. The whelping season was estimated to last about 40 days, with a peak in pupping at late October. No post natal mortality was observed during the whole study. The attachment between parous females and their pups was strong during the first five weeks after parturition. From the third week on mothers spent more time in the water than the pups. Data on daily peak haul-out time of pups, post natal presence of pups on the ice and distribution of births during the whelping season, indicate that censuses should be carried out between 12.00 hours and 16.00 hours local time and during three weeks after the peak of the pupping season. Pups weaned between 6 and 7 weeks of age. Hormone concentrations (oestradiol-17 and progesterone) indicated that females did not come into oestrus during lactation. Throughout the whole lactation period the mother dominates the relationship with her pup. Growth of the pups was linear during the first 5 weeks post partum, during which period birth weight (x=29.1kg) increased more than 3 1/2 times. Until the 6th week of age the pups gained on average 2.6 kg/day, the respective weight loss in cows was 5.8 kg/day. The latter represents an average loss of nearly 38% of the initial weight at parturition. During the course of lactation, suckling frequency decreased, whereas the length of each bout increased resulting in an approximately constant total suckling duration per week. Of the energy used by females during the first 5 weeks, 52% was consumed by pups. Pups only took milk and the cows did not feed at all during the whole period of observation. Therefore the realized growth in pups, 10 g/min of suckling, was totally derived from energy stored by the cows.