Milk composition in the eastern quoll, Dasyurus viverrinus (Marsupialia:Dasyuridae)

The milk constituents of Dasyurus viverrinus, a carnivorous marsupial, exhibited major quantitative and qualitative changes during the course of lactation. The milk produced in the early stages of lactation was dilute, about 13-16% (w/w) solids before 3 weeks with carbohydrate representing the major fraction. In the latter stages of lactation the milk was concentrated, around 30% solids, and lipid was the predominant fraction. Palmitic acid was the major fatty acid present in early-stage milk but oleic acid became predominant in milk after 10 weeks post-partum. The changes in milk composition in D. viverrinus were similar to those described for the milks of herbivorous marsupials which therefore suggests that this pattern may be uniform throughout the Marsupialia.     

Changes in milk carbohydrates during lactation in the eastern quoll, Dasyurus viverrinus (Marsupialia)

1. Milk samples were obtained at various stages of lactation from 16 captive eastern quolls. 2. The mean milk carbohydrate concentration was 7.4% (w/v; total hexose) at 8 weeks post partum, decreasing to 5.2% at 17 weeks and to less than 2% at the end of lactation. 3. The predominant monosaccharide constituent of the carbohydrates was galactose, followed by glucosamine, glucose and sialic acid. 4. Thin layer and gel chromatography showed that the milk carbohydrates consisted of lactose and a variety of higher oligosaccharides, some of which appeared to be identical to oligosaccharides of known structure previously isolated from milk of the tammar wallaby, Macropus eugenii. 5. In general, both the quantitative and qualitative changes observed in the milk carbohydrates of the eastern quoll were similar to those documented for macropodids.     

Milk carbohydrates of marsupials. II. Quantitative and qualitative changes in milk carbohydrates during lactation in the tammar wallaby (Macropus eugenii)

Milk was collected at various stages of lactation from a group of tammar wallabies, M. eugenii, in which parturition had been synchronized. The milk carbohydrate was determined by a phenol-sulfuric acid method which had been modified to give equal colour yields for galactose and glucose. The mean carbohydrate content increased gradually during the first 6 months of lactation to a peak of 13 g hexose/100 ml of milk, but then fell rapidly to much lower values, over the following 2 months. Throughouth lactation, galactose was the predominant monosaccharide constituent of acid hydrolysates of the milk carbohydrate. Glucose, glucosamine, galactosamine and sialic acid were the only other monosaccharides present. Qualitative changes were investigated by gel filtration and thin-layer chromatography. During the first 6 months post partum the milk carbohydrate was composed of a variety of oligosaccharides including lactose, but from 8 months onwards it consisted mainly of free monosaccharides. Between 6 and 8 months an intermediate pattern was observed, i.e. a mixture of lower oligosaccharides and free monosaccharides. In two animals which suckled both a new-born pouch young and a young at foot, the mammary gland supplying the new-born secreted milk which was rich in oligosaccharides, whereas that supplying the young at foot produced milk in which the carbohydrates were mainly free monosaccharides, and which had a much lower carbohydrate content.     

Milk composition of three free-ranging African elephant (Loxodonta africana africana) cows during mid lactation

Data are presented that indicate the dynamic changes of nutrients in milk from three free ranging African elephant (Loxodonta africana africana) cows during lactation. At the respective collection times of 12, 14 and 18 months of lactation the nutrient content was 47.3, 52.0 and 68.6 g protein; 60.7, 87.4 and 170.8 g fat; 1.6, 2.1 0.5 g lactose and 20.9, 21.5 and 8.6 g oligosaccharides per kg milk. The protein fraction respectively consisted of 18.0, 31.7 and 45.9 g caseins/kg milk and of 29.3, 20.3 and 22.7 g whey proteins/kg milk. Electrophoresis and identification of protein bands showed that polymorphs of one whey protein may be present in elephant's milk similar to polymorphs of alpha-lactalbumin found in cow's milk. From the middle of the lactation time lactose was replaced by oligosaccharides as major carbohydrate, and the major compound of these was identified as isoglobotriose by 1H NMR spectroscopy. The lipid fraction contains a high content, of capric and lauric acids, approximately 70% of the total fatty acids, and low content of myristic, palmitic and oleic acids. During these lactation times the content of short chain fatty acids, capric and caprylic acids increased, while fatty acids lauric acid and longer decreased.     

n-acetyl-beta-hexosaminidase activity in human breast milk

1. The lysosomal enzyme, N-acetyl-beta-hexosaminidase (HEX) is present in human breast milk. It is composed predominantly of "A" (heat-labile) and "B" (heat-stable) isozymes which coelute with the corresponding major serum isozymes on DE-52 ion-exchange chromatography. 2. Total HEX activity in "early" milk obtained at 2.8 +/- 1.4 weeks post partum, is approx. 2.5-fold higher (87 +/- 29 nmol/60'/mg protein. n = 10) than that of pregnancy serum (35.7 nmol/60'/mg protein) prior to delivery. 3. These levels increase to greater than 3-fold (110 +/- 20 nmol/60'/mg protein, n = 13) as the milk matures (10.3 +/- 4.2 weeks). 4. The specific activity of HEX A in the milk changes little with time post partum, because absolute 5. In contrast, HEX B specific activity is increased, as absolute levels (per volume) remain constant in the face of decreasing milk protein content, 5. These changes result in a high degree of correlation (r = 0.81) between time of lactation and % HEX A observed.     

Milk Composition during Lactation Suggests a Mechanism for Male Biased Allocation of Maternal Resources in the Tammar Wallaby (Macropus eugenii)

Recent research has found empirical evidence in support of the Trivers-Willard Hypothesis that offspring sex allocation is correlated with maternal investment. Tammar wallabies birthing sons have higher investment ability; however a mechanism for sex specific differential allocation of maternal resources in wallabies remains elusive. In metatherians the majority of maternal investment is during lactation. To examine if differential allocation occurs during lactation, we measured total milk protein, lipid and carbohydrates, from mothers with male and female pouch young, during phase 2B (100–215 days post partum) and phase 3 (215–360 days post partum) of lactation. Mothers of sons allocated significantly higher levels of protein than mothers of daughters during phase 2B of lactation, however no sex specific difference in maternal allocation was found for lipids, carbohydrates, or any milk component during phase 3 of lactation. We were unable to measure milk production to establish any differences in the amount of milk allocated. However, with the production of more milk comes a dilution effect on milk components. Given that we find no apparent dilution of milk components may suggest equality in milk production. Offspring body weight at 14 months of age was related to protein allocation during phase 2B of lactation, providing a maternal mechanism for differential allocation with fitness consequences. We believe collection of earlier phase 2A (0–100 days post partum) milk may yield important results given that differential investment in metatherians may be most apparent early in lactation, prior to any significant maternal investment, when a decision on termination of investment can be made with very little energetic loss to the mother. Interestingly, small mothers did not birth sons and better maternal condition was associated with raising sons. These data are in support of TWH and demonstrate a potential mechanism through which condition dependent and sex specific maternal investment may occur.     

Replacement of the aliphatic chains of Clostridium acetobutylicum by exogenous fatty acids: regulation of phospholipid and glycolipid composition.

The membrane lipid aliphatic chains of Clostridium acetobutylicum ATCC 4259 have been extensively modified by growth in biotin-free medium containing vitamin-free casein hydrolysate supplemented with either elaidic acid, oleic acid, or mixtures of palmitic and oleic acids. Growth with elaidic acid resulted in polar lipids containing 88.6% 18:1 acyl chains and 94.5% 18:1 ether-linked chains. Growth with oleic acid resulted in comparable levels of enrichment of the lipids with 18:1 chains and C19 chains containing cyclopropane rings. When cells were grown with mixtures of palmitic and oleic acids, the ether-linked chains of the plasmalogens were greater than or equal to 64% 18:1 plus C19 chains containing cyclopropane rings at all ratios of oleic to palmitic acid in the medium. The acyl chains reflected the palmitic acid content of the medium more closely. Marked changes were observed in both phospholipid and glycosyldiglyceride compositions as the lipid acyl and ether-linked chains became more enriched with unsaturated and cyclopropane chains. The ratio of the glycerol acetal of plasmenylethanolamine to phosphatidylethanolamine increased, the ratio of cardiolipin to phosphatidylglycerol decreased, and the ratio of diglycosyldiglyceride to monoglycosyldiglyceride increased. However, the monoglycosyldiglyceride/diglycosyldiglyceride ratio was lower for cells grown on 100% oleic acid than for cells grown on 60 or 80% oleic acid. In the membranes of cells grown on 100% oleic acid, the ratio of glycolipids to phospholipids was lower than that found in cells grown on 60% oleic acid. These results indicate that C. acetobutylicum regulates its polar lipid composition in a complex manner involving phospholipids and glycosyldiglycerides. These changes can affect the equilibria between those lipids that form bilayers and those lipids that tend to form nonlamellar phases when enriched with unsaturated aliphatic chains. Phosphoglycolipids of unknown structure were also observed in cells grown either with biotin or with fatty acids. The content of the most abundant phosphoglycolipid also varied with the degree of unsaturation of the cellular lipids.     

Lipids and fatty acid composition of developing winged bean seeds

The moisture, lipids and fatty acid composition of developing winged bean (Psophocarpus tetragonolobus) seeds were studied. The moisture content decreased steadily as the seeds matured. The lipid content increased gradually and reached a maximum ca 6 weeks after flowering (WAF). In the early stage (2 WAF) of the developing seeds there were more polar lipids (glycolipids and phospholipids) than neutral lipids but, as the seeds developed, neutral lipids gradually accumulated while the polar lipids decreased until 6 WAF. Thereafter, both the neutral lipid and polar lipid levels remained little changed. The amounts of palmitic and stearic acids decreased, but the level of behenic acid increased as the seeds matured. On the other hand, the oleic acid content increased while that of linolenic acid decreased rapidly as the seeds matured. The concentration of linoleic acid, however, fluctuated during the development of the seeds.     

Late gestational nutrient restriction: effects on ewes' metabolic and homeorhetic adaptation, consequences for lamb birth weight and lactation performance

We investigated the effect of 50% nutrient restriction during the last 6 weeks of gestation on twin-pregnant ewes' plasma glucose, non-esterified fatty acid, beta-hydroxybutyrate, insulin, IGF-1 and leptin concentrations and the effects on lamb birth weight and ewes' lactation performance. Plasma metabolite and hormone concentrations in restricted ewes suggest that maternal tissues were being mobilised. Despite the ewes' adaptations their lambs weighed significantly less at birth. Furthermore, colostrum and milk yields were markedly reduced up until the latest measurement at 3 weeks post partum despite ad libitum access to feed. Reduced milk yields coincided with reduced plasma IGF-1 concentration pre partum in nutrient restricted ewes indicating, that mammary gland development may have been compromised. The present data suggest that leptin is not involved in the regulation of early lactation changes in feed intake and energy balance. It is concluded that severely reduced nutrient availability in late gestation affects fetal growth in utero and has a prolonged negative effect on lactation performance.     

Gangliosides in bovine milk. Changes in content and distribution of individual ganglioside levels during lactation.

Bovine milk undergoes changes in its ganglioside contents during the different stages of lactation. These contents are higher in colostrum (7.5 mg of lipid-bound NeuAc/kg) than in transitional (2.3 mg) or mature (1.4 mg) milk. The sialic acid content of milk follows a similar profile to that of gangliosides with the highest content during the first few days post partum followed by a gradual decrease towards the end of the period studied. When the individual distribution of gangliosides was examined throughout the course of lactation, several changes were also found. GD3 is the major ganglioside (about 60-70%) found; its content decreases from the first to the fifth day, increasing towards the end of the period considered. GM3, GD3 and GT3, sialyllactosylceramide-containing gangliosides account for 80-90% of the total lipid-bound NeuAc content. The most striking change in the ganglioside pattern was the gradual increase in G3.     

Changes in alpha-lactalbumin, total lactose, UDP-galactose hydrolase and other factors in tammar wallaby (Macropus eugenii) milk during lactation

alpha-Lactalbumin was isolated from milk of M. eugenii and its concentration in milk samples taken at various times during lactation (0-40 weeks post partum) was determined by single radial immunodiffusion using rabbit antiserum to the purified protein. The alpha-lactalbumin concentration remained almost constant throughout lactation even though the concentration of total lactose (free lactose plus lactose contained in oligosaccharides) fell to zero after 34 weeks post partum. This fall in lactose was accompanied by a rise in the free galactose and glucose concentrations and marked increases in UDP-galactose hydrolase, nucleotide pyrophosphatase, alkaline phosphatase and acid beta-galactosidase activities. It is suggested that the in vitro hydrolysis of UDP-galactose was due to nucleotide pyrophosphatase and that this enzyme may also play a role in vivo late in lactation by making UDP-galactose unavailable for the synthesis of lactose. Alternatively, lactose and lactose-containing oligosaccharides might be degraded by the acid beta-galactosidase during or after secretion.     

Stereospecific biosynthesis of triacylglycerols in mammary glands from lactating rats.

Microsomal plus cytosol preparations from the mammary gland of lactating rats are capable of incorporating palmitic acid and oleic acid into triacylglycerols. These triacylglycerols are similar in structure to those found in rat milk, where palmitic acid tends to be confined to the sn-2-position of the glycerol. Both glycerol 3-phosphate and dihydroxyacetone phosphate function as acyl acceptors. The enzymic synthesis of triacylglycerols appears in late pregnancy, increases rapidly during early lactation, but disappears within 3 days of weaning.     

Milk secretion in the rat: progressive changes in milk composition during lactation and weaning and the effect of diet.

1. Progressive changes in the composition of milk from rats has been studied from day 0 to 20 of lactation and for 3 days following separation of the dams and pups at day 20 post partum. 2. The changes in concentration of Na, K and lactose suggested that secretion both prepartum and following weaning occurred by a paracellular mechanism whereas a transcellular pathway existed during established lactation. 3. The concentration of total protein and casein increased gradually throughout lactation. In contrast, the concentration of serum albumin increased and transferrin decreased markedly during early lactation. The fat content of milk declined 3-fold within 5 days of birth but the concentration of Ca, Mg and inorganic P increased. The concentration of each of these milk constituents remained constant during established lactation. 4. Following weaning the pronounced decline in lactose, K and inorganic P was negatively correlated with an increase in all other milk constituents except fat. 5. Rats fed a low energy diet produced milk with a lower fat content but with an unaltered concentration of protein and carbohydrate. The growth rate of these litters was similar for the first 5 days of lactation when compared to litters from dams fed a high energy diet. The growth rate of litters thereafter and following weaning was greater for rats fed a high energy diet.     

Glucose metabolism in crossbred Holstein cattle feeding on two types of roughage at different stages of lactation

An experiment was performed to study the glucose kinetics of crossbred Holstein cattle feeding on either hay or 5% urea-treated rice straw during early lactation (30 days post partum), mid-lactation (120 days post partum) and late lactation (210 days post partum). Two breeds: Holstein FriesianxRed Sindhi (50:50 = 50% HF) and Holstein FriesianxRed Sindhi (87.5:12.5 = 87.5% HF) were used. In early lactating 87.5% HF animals feeding on either hay or urea-treated rice straw, the high milk yields and lactose secretion were related to glucose uptake by the udder and udder blood flow as compared with those of 50% HF animals. Marked decreases in udder blood flow, glucose uptake, lactose secretion and milk yield were apparent in mid- and late lactation of both groups of 87.5% HF animals. In contrast, both groups of 50% HF animals showed no significant changes in udder blood flow, udder glucose uptake, lactose secretion and milk yields throughout the course of lactation. Total glucose entry rate using 3-[3H] glucose infusion, recycling of glucose carbon and plasma glucose clearance significantly increased during late lactation for 50 and 87.5% HF animals feeding on urea-treated rice straw. The utilization rates of glucose using [U-(14)C] glucose infusion were not significantly different among groups of animals and periods of lactation. It can be concluded that 87.5% HF animals have the genetic potential for a higher milk yield, but a shorter peak yield and poorer persistence in comparison with 50% HF animals. Changes in the utilization of glucose by the mammary gland for milk production in both groups of crossbred animals during feeding on either hay or urea-treated rice straw would be dependent on intramammary changes.     

Regulation of Lipid Droplet Size in Mammary Epithelial Cells by Remodeling of Membrane Lipid Composition—A Potential Mechanism

Milk fat globule size is determined by the size of its precursors—intracellular lipid droplets—and is tightly associated with its composition. We examined the relationship between phospholipid composition of mammary epithelial cells and the size of both intracellular and secreted milk fat globules. Primary culture of mammary epithelial cells was cultured in medium without free fatty acids (control) or with 0.1 mM free capric, palmitic or oleic acid for 24 h. The amount and composition of the cellular lipids and the size of the lipid droplets were determined in the cells and medium. Mitochondrial quantity and expression levels of genes associated with mitochondrial biogenesis and polar lipid composition were determined. Cells cultured with oleic and palmitic acids contained similar quantities of triglycerides, 3.1- and 3.8-fold higher than in controls, respectively (P < 0.0001). When cultured with oleic acid, 22% of the cells contained large lipid droplets (>3 μm) and phosphatidylethanolamine concentration was higher by 23 and 63% compared with that in the control and palmitic acid treatments, respectively (P < 0.0001). In the presence of palmitic acid, only 4% of the cells contained large lipid droplets and the membrane phosphatidylcholine concentration was 22% and 16% higher than that in the control and oleic acid treatments, respectively (P < 0.0001). In the oleic acid treatment, approximately 40% of the lipid droplets were larger than 5 μm whereas in that of the palmitic acid treatment, only 16% of the droplets were in this size range. Triglyceride secretion in the oleic acid treatment was 2- and 12-fold higher compared with that in the palmitic acid and control treatments, respectively. Results imply that membrane composition of bovine mammary epithelial cells plays a role in controlling intracellular and secreted lipid droplets size, and that this process is not associated with cellular triglyceride content.     

Changes in the yield, and carbohydrate, lipid and protein content of milk during lactation in the rat

The milk yield and composition was studied during the first three lactations of a group of rats. Milk yield increased steadily throughout the three lactations, but was somewhat lower during the first than subsequent lactations. Protein concentration was similar during all three lactations and varied little with stage of lactation. In contrast the lactose concentration, which was reasonably constant for the first 8 days post partum, increased thereafter two-fold by the end of the period studied in all three lactations. However, the N-acetyl-neuraminyl lactose concentration showed somewhat reciprocal changes. Considerable variations in the triacylglycerol concentration was found during the first lactation but few changes were observed during subsequent lactations. The free fatty acid concentration was at all times low and showed no significant changes during or between lactations. At most stages of lactation in raw milk, the major fatty acids are palmitate, oleate and linoleate. However, as lactation progresses there is an increase in the proportion of medium-chain saturated fatty acids and a corresponding decrease in the proportion of long chain unsaturated fatty acids in milk fat. Clearly the composition of milk is not invariable but changes both during and between lactations. Such changes may be expected to have some influence on the metabolism of the offspring.     

Selective retention of n-3 and n-6 fatty acids in human milk lipids in the face of increasing proportions of medium chain-length (C10-14) fatty acids

This paper reports the results of our analysis of the impact high levels of de novo fatty acids have on the proportions of essential and non-essential fatty acids in human milk lipids. The data for seven fatty acids (linoleic, alpha-linolenic, arachidonic (AA), docosahexaenoic (DHA), palmitic, stearic and oleic) were derived from several studies conducted in Nigeria. The proportion by weight of each of these fatty acids was plotted versus the proportion of C10-14 fatty acids. As the proportion of C10-14 fatty acids increased from 15 to 65%, there was not a proportional decrease in the percentages of all seven fatty acids, but, instead, preferential incorporation of the essential fatty acids, AA and DHA into the triacylglycerol component of the milk. At the same time, the proportions of stearic and oleic acid declined by 69% and 86%, respectively. However, the proportions of linoleic acid, palmitic acid, DHA, AA and alpha-linolenic acid, in milk lipids decreased by only 44%, 40%, 39%, 28% and 2.3%, respectively. These observations indicate that as the contribution of C10-14 fatty acids increases, essential fatty acids are preferentially incorporated into milk triacylglycerols at the expense of oleic acid and stearic acid.     

Milk composition and growth in wild and captive Tasmanian bettongs,<Emphasis Type="Italic"> Bettongia gaimardi</Emphasis> (Marsupialia)

Changes in milk composition (total solids, carbohydrate, protein, lipid and calculated gross energy content) during lactation in three groups of Tasmanian bettongs ( Bettongia gaimardi): free-living animals (wild group), captive animals offered a diet of dry dog food and apples ad libitum (ad lib group), and captive animals fed restricted amounts of the same diet (maintenance group) were related to growth rates (measured as body mass and head length) of their young. There were no significant differences in the concentration of milk solids among the three groups, but the wild group had higher lipid concentrations and the maintenance group had higher carbohydrate but lower protein concentrations. For all three groups, milk total solids increased through lactation from ca. 25% to ca. 45% and carbohydrate concentrations decreased from ca. 18% to about 3%. Protein concentrations increased from ca. 5% to ca. 10% in the wild and ad lib groups, but only from 4% to 8% in the maintenance group. Lipid concentrations increased in the wild and ad lib groups from ca. 4% to ca. 18%, but in the maintenance group only to ca. 7%. Calculated gross energy content of milk increased through lactation in the wild and ad lib groups (from ca. 500 kJ.100 ml(-1) to ca. 1,000 kJ.100 ml(-1)), but there was no significant increase in the maintenance group. The volume of milk produced increased to a peak just prior to permanent pouch vacation by the young, when the gross energy output in milk was 120-150 kJ.3 h(-1) in the wild and ad lib groups. On a daily basis this is equivalent to the milk energy output of larger wallabies, and helps to explain the relatively high growth rates of young Tasmanian bettongs. There were significant differences in growth rates among the groups, with the heaviest young always in the ad lib group. Thus differences in milk composition resulting from different planes of nutrition can lead to differences in growth rates of marsupial young.     

Comparison of the fatty acid profile of Spanish infant formulas and Galician women breast milk

The importance of dietary lipids during childhood is evident, as they are necessary for correct growth and development of the newborn. When breastfeeding is not possible, infant formulas are designed to mimic human milk as much as possible to fulfill infant’s requirements. However, the composition of these dairy products is relatively constant, while human milk is not a uniform bio-fluid and changes according to the requirements of the baby. In this study, breast milk samples were donated by 24 Spanish mothers in different lactation stages and different infant formulas were purchased in supermarkets and pharmacies. Gas chromatography coupled to flame ionization detection was used for the fatty acid determination. Compared to breast milk, first-stage formulas are apparently very similar in composition; however, no major differences were observed in the fatty acid profiles between formulas of different lactation stages. The Galician women breast milk has a fatty acid profile rich in oleic acid, linoleic acid, arachidonic acid, and docosahexaenoic acid. When comparing human milk with formulas, it becomes evident that the manufacturers tend to enrich the formulas with essential fatty acids (especially with α-linolenic acid), but arachidonic and docosahexaenoic acid levels are lower than in breast milk. Additionally, the obtained results demonstrated that after 1 year of lactation, human milk is still a good source of energy, essential fatty acids, and long-chain polyunsaturated fatty acids for the baby.     

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.