Effects of litter size, age of young, and parity on foster mother behaviour in Rattus norvegicus

Maternal behaviour in the presence of newborn pups is determined by the age and size of the litter independent of parity of the mother, previous experience with different sized litters, and the number of days after delivery the measurements are taken. After fostering 10-day-old young, maternal behaviour is influenced by the age and size of the litter and also by the mother's previous experience with different sized litters.     

Effects of age and parity on litter size and offspring sex ratio in golden hamsters (Mesocricetus auratus)

Golden hamsters that were mated repeatedly from 55 days of age produced 6-12 litters. Litter size at birth rose between the 1st and 2nd litters, peaked on the 3rd, and declined steadily after the 5th litter. Offspring sex ratio (% male) at birth followed a similar pattern: increasing between the 1st and 2nd litters, remaining high through the 3rd, and becoming increasingly female-biased thereafter. Weaning success decreased sharply after the 6th litter and most dams failed to raise any young to weaning after the 9th litter. These sequential effects on litter size, offspring sex ratio and weaning success were also observed in females mated once at different ages, but they occurred considerably later in life, i.e. increasing parity hastened the effects of advanced age. These age- and parity-related changes in litter composition are consistent with the Trivers-Willard hypothesis that physiologically-stressed females would skew offspring sex ratios to favour daughters. However, since the observed changes in sex ratio were probably due to differential prenatal mortality, their adaptive significance is unclear.     

Rearing a second generation of cotton-top tamarins (Saguinus oedipus oedipus) in captivity

The average age at first parturition in captive-born female cotton-top tamarins was 31 months. Only 18% of full-term young born were successfully reared by primiparous females, but success increased with parity and 71% of babies born at 4th and 5th pregnancies were raised. 15% of litters were single births, 61% were twins and 24% were triplets. There was a seasonal distribution of births, with a clear peak in the spring months. The average interbirth interval was 294 days.     

Behaviour of does and leverets of the European hare (Lepus europaem) whilst nursing

After birth in one particular place, the leverets of a litter dispersed after a few days. In the evening, the leverets were observed gathering at, or near, their birth place to await nursing. There the doe would arrive later, about one hour after sunset, to nurse the young gathered there. Individual recognition between leverets and their doe seemed of no importance, and meeting was dependent on time and place. The young were nursed for only a few minutes and their urine, excreted during nursing, was licked up by the doe, which finished nursing by jumping away from the young. The leverets dispersed again and kept apart from each other until they gathered again the next evening. Usually leverets are nursed for four weeks, but longer nursing periods were found in the last litters of the breeding season. The nursing behaviour is probably adapted to prevent discovery of leverets by predators.     

Differential Growth of Own and Alien Pups in Mixed Litters of Mice: A Role for Genomic Imprinting?

Female mice are said to be unable to distinguish own from alien offspring and will indiscriminately nurse each other's young in communal nests. Here, we present results of a split‐litter experiment that tested whether offspring growth was affected depending on whether they were nursed by their own or unrelated foster mothers. Pups of reciprocal crosses between C57/B6 and CBA/Ca strains were fostered in mixed litters that consisted half of their natural siblings and half of unrelated littermates of the reciprocal genotype. Analysing the relative growth of the two pup types showed that offspring gained proportionally more weight when nursed by their own mothers than their cross‐fostered litter mates during the period from day 15 until weaning, during which maternal provisioning effort contributes to pup weight gain. Before day 15 of the pups’ life, however, we found no advantage of being nursed by biological mothers, and we suggest that this may be due to the effects of paternally expressed genes in young that mask their maternal identity, thus favouring indiscriminate nursing of all young in a communal nest.     

Dominance Rank and Parental Investment in Swine (Sus scrofa domesticus)

Using domestic swine, we tested the general prediction from life history theory that females increase their investment in offspring with increasing age and parity. Because increased investment may have a greater beneficial impact on the lifetime reproduction of sons than daughters, we also tested the prediction that older females would invest more in sons than in daughters compared to younger females. Finally, we examined whether age- or parity-related patterns of change in reproductive effort were associated with differences in the social dominance ranks of females. Female swine from a large number of domestic breeds were assigned to social groups, and their dominance ranks were determined based on the outcome of agonistic encounters. The prediction that older females produce larger litters was supported, but the increase was related only to age, not to parity. Across all ages, high-ranking females produced a greater proportion of sons than low-ranking females. Contrary to our prediction, there was no rank-related change in the proportion of sons born with increasing age or parity. However, the mean body masses offspring born to high-ranking females increased with increasing maternal age and parity, but this was not the case for offspring of low-ranking females. Studies of free-ranging groups of swine are needed to determine whether an increase in body mass at birth would have different effects on the reproduction of sons or daughters.     

Age at first mating affects parental effort and fecundity of female Mongolian gerbils

We examined effects of age at first mating on both parental effort and fecundity of female Mongolian gerbils, Meriones unguiculatus. We found that, with increasing age at first mating and resulting age at first parturition, female gerbils: (1) were more likely to retrieve young removed from the nest, (2) spent more time both in contact with and nursing young, and (3) provided an environment in which pups grew more rapidly. Older mothers were also less likely to become pregnant than were younger mothers and, if successful in delivering a second litter, showed longer interlitter intervals and delivered smaller second litters. Between delivery and weaning of first litters, older mothers lost more weight than did younger mothers. We discuss these findings as consistent with the prediction from life-history theory that parental effort should increase with age-related decreases in residual reproductive value. Furthermore, and as predicted by parental investment theory, older mothers delivered reliably more male-biased second litters than did younger mothers. Because of the different sex ratios of litters born to older and younger dams, we anticipate discovery of differences in reproductive and parental behaviours of offspring of dams of varying ages as a result of differences in the intrauterine exposure of their young to testosterone. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

The growth of the Mongolian gerbil, Meriones unguiculatus, from birth to maturity

Birthweights and sex of young were recorded in 357 litters of Mongolian gerbils, Meriones unguiculatus. The liveweights of individual young in 47 litters were recorded every 5 days up to 40 days of age and related to the number of young suckling. Growth curves are presented for both sexes from birth to 150 days of age.     

Optimal offspring size in a small mammal: an exception to the tradeoff invariant life-history rule

Offspring size and number were examined in a captive population of wild guinea pigs ( Cavia aperea ), and findings were compared with models of optimal offspring size for small litters. Median and modal litter size was two, regardless of maternal size or parity. Females producing their second litter tended to have litters that were larger than average. In contrast, young females that were still growing never had litters that were larger than average. Mean offspring size decreased and variation in offspring size tended to decrease with increasing litter size. Optimal offspring size models, in which offspring survival depended on the amount of resources invested, as well as litter size, predict such a trend. Little support was found for Charnov and Downhower's (1995) tradeoff invariant life-history rule that the range in offspring sizes between litters is inversely proportional to the size of the litter. Cavia aperea may be an exception to this rule because pup mass at birth did not reflect total reproductive investment, because conversion of resources into litter mass may not be linearly related to litter size and because resources were not equally partitioned among offspring within large litters. Experimental data are needed to determine the relevance of these results among mammals in general.     

Explaining the seasonal decline in litter size in European ground squirrels

In European ground squirrels Spermophilus citellus as in many ground squirrel species. late born litters are composed of fewer young than early born litters. Two alternative though not mutually exclusive hypotheses may explain this seasonal pattern of change in litter size. On the one hand. the production of few large young late in the season may be an adaptation to time limitations on the offspring. that have to complete growth and fattening prior to hibernation. Then one would expect a trade-off between offspring number and size as the breeding season progresses. At its extreme. this hypothesis would predict that total maternal effort should be equal independent of litter size. Alternatively. litter size may be determined by physiological limitations on the mother. in that highly constrained mothers breed later and produce smaller litters. Then one would expect reduced overall maternal effort in highly constrained mothers of smaller litters. In this case. a trade-off between litter size and offspring size would not be expected. We found that total maternal effort in terms of gestation length and the duration of lactation increased with increasing litter size. thus supporting the second hypothesis. Lactation was not terminated at natal emergence. It extended a relatively long period of time beyond the time of first litter emergence depending on litter size. During prolonged lactation. individual young of large litters made up body mass to young of small litters. As a consequence. juvenile weaning body mass was unaffected by litter size although offspring body mass at natal emergence was inversely related to litter size. This additional weight gain in young of large litters compensated for initial survival disadvantages and presumably affected fecundity at yearling age.     

Lifespan ontogeny of breeding and reproductive success in Mongolian gerbils

An analysis was made of demographic data collected from a colony of Mongolian gerbils (Meriones unguiculatus). The data on births of over 3800 gerbils provide evidence on several aspects of development and fecundity. Approximately 4% more females than males survived to weaning. Mean number of pups per dam per year was estimated to be 48.3, greater reproductive success than reported previously. The shortest period of gestation resulting in viable pups was 25 days. Implantation was delayed by the presence of suckling pups. When gestation was prolonged, there was a direct relationship between the number of pups being suckled concurrently and the length of the delay. Length of gestation did not affect the size of the subsequent litter. Female gerbils gave birth as early as 72 days of age. Gerbils continued to reproduce until after 2 years of age in some cases. There were fewer pups per litter born to older females, fewer pups per litter with increasing parity, and longer intervals between litters with increasing age.     

The influence of prenatal and postnatal fraternity size on reproduction in mice

Effects of prenatal and postnatal fraternity size (size of litter in which an individual is reared) on age at vaginal opening, growth, and subsequent litter production were examined by rearing mice in standardized prenatal and postnatal fraternities in a 3 X 3 factorial design. Prenatal fraternity size was standardized by reducing litters of 14 or more to either 14, 10, or 6 fetuses on Day 9 of gestation. Postnatal fraternity size was standardized by assigning pups randomly to nurse in litters of 5, 10, or 15 pups within 24 h of birth. Both prenatal and postnatal fraternity size affected growth of the mice (p = 0.02 and p less than 0.01, respectively) with mice reared in small fraternities attaining greater weights throughout the study. Prenatal fraternity size had a negative linear effect on litter size at second parity and tended to have a positive linear effect on age at vaginal opening. Postnatal fraternity size affected age at vaginal opening (p less than 0.01), litter size at first parity (p less than 0.01), and litter size at second parity (p = 0.03). These effects were positive and linear for age at vaginal opening (p less than 0.01; small fraternity size associated with younger age at vaginal opening) and negative and linear for litter size at first and second parity (p less than 0.01 and p = 0.10, respectively; smaller fraternity size associated with larger litter size). There was no interaction between prenatal and postnatal fraternity size effects on age at vaginal opening or litter size (p greater than 0.20).     

Mating relationships and breeding suppression in the dwarf mongoose

Dwarf mongooses live in packs containing a dominant breeding pair. The alpha female produces litters at regular intervals, usually three times per year. Other sexually mature females come into oestrus in synchrony with the alpha female and occasionally become pregnant but may not raise their offspring. Some females which had not been visibly pregnant nursed the young of other females. Early in the alpha female's oestrous cycle the alpha male maintains proximity and copulates with her exclusively, attacking any other adult males which approach. Later he also copulates with other adult females and the beta male mates with the alpha female. The alpha pair are likely to be the parents of the great majority of young born in the pack.     

Reproduction in a laboratory colony of the pouched mouse, Saccostomus campestris

Pouched mice ovulate spontaneously and have a 4-day cycle (3-5 days). The variation was caused by prolonged oestrus. The vagina opened at about 34 days of age and the first oestrus was experienced at 44 days of age. Females experienced several sterile cycles before their first conception, which occurred at an age of about 56 days. The gestation period in most cases was 21 days and implantation occurred about 4-5 days after mating. The females were not receptive post partum. Litter size varied from 3 to 13 with a mean of 7.1 in young primiparous females and 7.9 in adult multiparous females, indicating that fecundity did not increase with age or parity in this species. Mortality of young was highest during the first 2 days post partum. The young were not attached to the nipples and were weaned at 25 days of age. Females did not cycle during lactation. After lactation most females exhibited one or two oestrous cycles without mating or became oestrous and mated without conceiving, resulting in a litter interval of about 53 days.     

Co-operative Rearing by Slender-tailed Meerkats (Suricata suricatta) in the Southern Kalahari

Slender-tailed meerkats or suricates, Suricata suricatta, are small, gregarious and largely insectivorous mongooses which inhabit the arid and semi-arid areas of southern Africa. We describe four elements of co-operative rearing of the young: 1. babysitting at the den; 2. creching on foraging trips; 3. provisioning with prey items away from the den; and 4. allonursing , including spontaneous lactation. Individual band members differed markedly in contributions to co-operative rearing. All adults and yearlings guarded the den and helpers actively defended young against predators. Breeders babysat significantly less than did non-breeding adults. Differences in age–sex class contributions were also evident in creching behaviour. The feeding of kittens by provisioners extended from soon after their first emergence from the den at 2–3 wk of age to effective foraging independence at 10–12 wk. Breeding females initially contributed little care other than nursing. Allonursing occurred in six of 25 closely observed litters, including three incidents of spontaneous lactation when subordinates nursed the young of higher-ranking females. Such co-operation is likely to be critical to the survival of these highly sociable mongooses in a semi-arid environment such as the Kalahari where food availability can fluctuate enormously.     

Sows with high milk production had both a high feed intake and high body mobilization

Selection for increased litter size have generated hyper-prolific sows that nurses large litters, however limited knowledge is available regarding the connection between milk production, feed intake and body mobilization of these modern sows. The aim of the current study was to determine what characterized sows with high milk production and nursing large litters, differences between sows of different parities and effects of lactational performance on next reproductive cycle. In total 565 sows (parity 1 to 4) were studied from 7 days before farrowing until weaning. On day 2 postpartum litters were standardized to 14 piglets. Weight and back fat thickness of sows were measured at day 7 prepartum, day 2 postpartum and at weaning. Litters were weighed at day 2 and at weaning. Pearson correlation coefficients between variables were calculated and regression models were developed. The average daily feed intake (ADFI) of the sows was 6.1±1.1 kg/day, average daily gain (ADG) of the litter was 2.92±0.53 kg/day and sows weaned 13.0±1.1 piglets. First parity sows generally had a lower ADFI and milk production and a decrease in total born piglets in next litter compared with parity 2 to 4 sows, which could be explained by a relatively higher proportion of their body reserves being mobilized compared with multiparous sows. The ADG of the litter was positively related by ADFI of the sows, litter size and BW loss and increasing the ADFI with 1 kg/day throughout lactation likely increased the ADG of the litter with 220 to 440 g/day in parity 1 to 4, respectively. Increasing the ADFI by 1 kg/day reduced the BW loss with 6.6 to 13.9 kg of parity 1 to 4 sows, respectively, during lactation, whereas increasing the average milk yield with 1 kg/day raised the BW loss with 4.3 to 21.0 kg of the four parities during lactation. The number of total born piglets in the next litter was positively related to the number of piglets born in the previous litter. In conclusion, both a high feed intake and a high mobilization of body reserves was a prerequisite for a high milk production. The sows might be very close to the physical limit of what they can ingest and future research should therefore, focus on optimizing the dietary energy and nutrient concentrations of diets for lactating hyper-prolific sows and herein distinguish between primiparous and multiparous sows.     

Energy intake and milk production in mink (MUSTELA vison)‐effect of litter size

Energy intake and milk production were measured in 12 mink dams raising litters of 3, 6 and 9 kits one to four weeks post partum by means of balance experiments and measurements of milk intake of the kits by the water isotope dilution technique. The dams were fed ad libitum on a conventional wet mink diet (DM: 323 g/kg; CP: 173 g/kg; ME: 4.4 MJ/kg). Milk samples collected from dams with corresponding litter sizes and lactation weeks, and body composition of kits nursed by these dams, were analysed for content of DM, ash, N and fat. The ME and drinking water consumption were higher in dams nursing 9 kits than in dams nursing 3 kits. The N and water balances as well as the live weight of dams were not affected by litter size. Daily milk production was higher in dams nursing 9 kits than in dams nursing 3 kits. The DM, N and fat content of the milk increased during lactation, but were not affected by litter size. Individual kit live weight was higher in litters of 3 than in litters of 6 and 9 kits four weeks post partum. The DM and fat content of the kits were lowest in kits from litters of 9 kits, whereas these kits had the highest protein content. Daily ME for maintenance of kits and the efficiency of utilisation of ME in milk for body gain were estimated to 356 kJ/kg^0.75, k_p ≈0.53 and k_f ≈0.71, respectively. In conclusion, daily milk production increased with increasing litter size, but not in proportion to the number of kits, indicating that milk production limits the growth rate of the young. In the fourth week of lactation, milk production was not different between dams nursing 6 or 9 kits, indicating a maximum capacity.     

Epidemiological Studies of Piglet Diarrhoea in Intensively Managed Danish Sow Herds

The study comprised 70,796 litters in 104 sow herds, observed from 1976 through 1982. Weaning age decreased from approx. 42 days to approx. 30 days during the observation period. Diseases and symptoms were recorded together with production parameters (feeding, barn construction, economic returns etc.). The mean incidence rate of pre-weaning diarrhoea was 6.8 % of litters, with considerable inter-herd differences (incidence rates from 0 to approx. 50%). There was a slight increase in incidence during the autumn (August through October). Incidence rates increased with litter size, with a peak incidence in litters of 11–13 piglets, and decreased with increasing parity of the sow. There was a positive association between occurrence of arthritis and pre-weaning diarrhoea in the litters, and litters from sows with post parturient disease (MMA complex) had 1.8 times higher risk of getting diarrhoea than litters from healthy sows. No important differences among breeds were found. Small herds (< 200 farrowings per year) had higher incidence rates than large herds (400–499 farrowings per year). Herds with a gilt proportion above 30 % had an incidence rate of 12.3 %, i. e. nearly twice as high as the overall mean (6.8 %). There was a trend towards a higher incidence rate in litters kept in traditional pens (i. e. large pens with solid floor and loose sows) than in intensive pens (i. e. small pens with slatted flooring and tethered sows). The overall pre-weaning mortality rate was 16.2 % of pigs born, half of which was due to stillbirths (6.3 %) and overlaid piglets (2.2 %). In litters with pre-weaning diarrhoea, the mortality rate was 19 %, compared to 13 % in litters without occurrence of diarrhoea. This difference accounts for an excess loss of 0.6 piglets from birth to weaning in diarrhoeic vs. non-diarrhoeic litters. Piglets from litters with pre-weaning diarrhoea had reduced weight gain. Thus, on the average, they were 2.2 days older at 25 kg bodyweight and weighed 0.4 kg less at 30 days than piglets from non-diarrhoeic litters. Also, litters with pre-weaning diarrhoea had a significantly increased risk of post-weaning diarrhoea. The present information forms a basis for defining acceptable disease thresholds in suckling litters in intensively managed herds.     

Influence of some sow characteristics on within-litter variation of piglet birth weight

Within-litter variation of piglet birth weight (BW0) is associated with an increased piglet mortality and a high variability in pig weight at weaning and weight or age at slaughter. Data collected in two experimental herds were used to quantify within-litter variability in BW0 and to assess the influence of factors mainly related to the sow. Within 24 h after birth, piglets born alive were individually weighed and stillborn piglets were collectively (first data set) or individually (second data set) weighed. The first data set was restricted to litters with no or only one stillborn piglet (3338 litters). It was used to assess the influence of genetic selection on BW0 variation by comparing litter characteristics before (1994 to 1996) and after (2001 to 2004) the development of hyperprolific sows in this herd. The second data set included all litters (n = 1596) from sows born between 2000 and 2004. For each litter, mean BW0 (mBW0) and its coefficient of variation (CVBW0) were calculated. Then, variance analyses were performed to test the influence of litter size, parity, year of sow birth and season at conception. Prolificacy improvement was associated with an increased CVBW0 in litters from pure Large White (LW) and Landrace × Large White (LR × LW) crossbred sows. The CVBW0 averaged 21% and was significantly influenced by litter size and parity. It increased from 15% to 24% when litter size varied from less than 10 piglets to more than 15 piglets. The proportion of small piglets (i.e. weighing less than 1 kg) increased concomitantly. The CVBW0 was not repeatable from a parity to the following. It was lowest for first and second parities (20%) and thereafter increased progressively. The CVBW0 was positively related to sow's backfat thickness gain during gestation. Taking into account litter size, parity, year of sow birth and season at conception explained 20% of BW0 variation. Thus, major part of heterogeneity is due to other factors, presumably including embryo genotype, on the one hand, and factors that influence embryo and foetus development, such as epigenetic factors, on the other hand.     

Reproductive biology of captive female Eurasian lynx, Lynx lynx

Studies on wild Eurasian lynx (Lynx lynx) have revealed variation in reproduction between areas, years and individuals. In order to explore potential causes for this variation other than food supply, we analysed data from captive lynx, which provide conditions with minimal environmental variation as all were fed ad libitum. Data from 37 individual female lynx were available from 20 zoos in Norway, Sweden, Finland, Switzerland and the Czech Republic. Data on 177 reproductive events (where a male was available to the female at mating time) are presented. Of these events, 85% resulted in litters being born. Average litter size was 1.95, with a variation from 1 to 4. The mean birth date was 26th May, and sex ratio was not significantly different from parity. The probability of reproduction was related to age, with fewer litters produced by the very young (2–3-year old), and no sign of a senescence effect. However, a clear effect of senescence on litter size was evident. The captive lynx did not have higher reproductive rates than wild lynx, indicating that either factors other than food supply are driving the variation in wild lynx reproduction, or that a factor such as stress may be causing additional variation in the captive population.