Abdominal odours of young,low-ranking European rabbit mothers are less attractive to pups: an experiment with animals living under natural breeding conditions

Lactating rabbit females (Oryctolagus cuniculus) emit abdominal, ventral odour cues that are involved in guiding the pups to the nipples. We tested the impact of the females’ social rank and age on the emission of this odour signal; both factors are highly collinear in wild rabbits and we did not aim to disentangle them in our study. We predicted that the abdominal odours of young, low-ranking females, which are usually more stressed and have a comparatively lower body condition, should be less attractive to pups than those of older, high-ranking females. For the experiment, we caught females from a population of wild-type European rabbits living in a large field enclosure. Pairs of adult females were placed in boxes with wire-mesh floor fixed over an arena with three rabbit pups aged 4–7 days old. First, we evaluated the effectiveness of our two-choice design by testing lactating females (LF) against non-lactating (NLF) controls (n = 17 pairs). In accordance with recent studies, the pups showed a clear preference for lactating females by means of a higher exploration time and more local searching under the ventral region of these females. In a second experiment, the pups chose between lactating 1-year-old, low-ranking females (Y/LRF) and older, high-ranking females (O/HRF) (n = 10 pairs). These latter assays suggested a comparatively lower attractiveness of Y/LRF females to pups searching for milk. Although the pups showed no significant preference in exploration behaviour, they preferably directed local searching events to the abdominal region of O/HRF females. In conclusion, our study suggests that mother–offspring chemosensory communication is modified by the mother’s age and social rank, i.e., by a complex of factors that are strongly linked in rabbit females in the wild.     

Influenza Infection Neutralizes the Attractiveness of Male Odour to Female Mice (Mus musculus)

This study aimed to determine if female house mice, Mus musculus domesticus, are able to assess a male's infection status from odour cues. We collected urine from male mice before, during, and after they were experimentally infected with influenza, a respiratory virus. Females spent more time investigating urine collected from males while they were uninfected than when they were infected. Also 70 % of females released into a large enclosure preferred to nest in boxes containing urine collected from uninfected rather than infected males. This is the first evidence that mice can discriminate virally infected individuals through chemical signals and the first evidence that infection causes odour changes in the urine. To determine if the odour of infected males is repulsive, we presented females with urine samples and neutral water blanks. Normal urine collected from uninfected males was more attractive, whereas urine collected during infection was as attractive as water. This indicates that rather than being aversive, influenza infection abolishes the attractiveness of a male's odour. A similar effect also occurs when male mice are infected with coccidian gut parasites (Kavaliers & Colwell 1995, Proc. R. Soc. Lond. 261B, 31–35). One proximate reason for the neutralization of the attractiveness of a male's odour may be a decrease in serum androgen concentrations during infection.     

Foraging and mate-finding in the silver Y moth, Autographa gamma (Lepidoptera: Noctuidae) under the risk of predation

Animal foraging and reproductive behaviour is influenced by other simultaneous demands such as predator avoidance. The trade-offs between these demands may depend on sex or mating experience. This study demonstrates that the olfactory-mediated foraging and mate-seeking behaviours in the silver Y moths, Autographa gamma , are affected by auditory cues mimicking their bat predators. Both males and females changed their foraging behaviour under simulated predation risk. Fewer moths reached the odour source following sound stimulation and the time to find the odour source increased by up to 250%. However, there were no significant differences between male and female ability to reach the plant odour source or the duration of the flight towards the source when stimulated with ultrasound. Hence females are not more cautious than males when observed in the same behavioural context. Risk-taking in males was independent of whether they were flying toward a flower odour or sex pheromones having equal attractive value. This indicates that the trade-off between olfactory and acoustic cues is independent the type of odour. Mated females were not as strongly affected by sound as non-mated, indicating that flower odours have a higher adaptive value for mated females, suggesting that some processes following mating experience influence the trade-off between flower odours and simulated bat sounds.     

Differences in fertility and suckling patterns between primiparous and multiparous rhesus mothers (Macaca mulatta)

Primiparous females gave birth around the same time as multiparous nonlactating females, and earlier than did multiparous lactating females. No differences in birth sex ratio were found between primiparous and multiparous females. During the breeding season following birth, primiparous mothers returned to oestrus later than did multiparous mothers, and while few primiparous mothers conceived successfully during that season, most multiparous mothers did. Primiparous females suckled their infants more frequently than did multiparous females at all ages; infants of primiparous females also made more nipple contacts per bout, and had shorter sucking bouts. When mothers came into oestrus, suckling frequency drastically increased for primiparous females, but not for multiparous females, magnifying the differences between the two groups. After the first oestrus, suckling frequency declined for all mothers, but multiparous mothers had consistently lower suckling frequencies than did primiparous mothers. The high suckling frequency, and numerous nipple contacts per bout, found among primiparous mothers are likely to be related to the low reproductive chances that these females faced during the breeding season. Multiparous mothers seemed to compensate for their low suckling frequency by lengthening the suckling bouts, and this suckling pattern did not hinder their reproduction. It is argued that primiparous mothers might have to suckle their infants more frequently because they can only produce milk at slow rates, being in this way forced into a reproductively inhibiting suckling pattern. However, the delay in subsequent reproduction could be ultimately advantageous for primiparous mothers if it enhanced infant survival, and allowed the mothers to regain physical condition before reproducing again.     

Brief communication: Self‐suckling in Barbary macaque (Macaca sylvanus) mothers before and after the death of their infant

We report here self‐suckling in four wild female Barbary macaques (Macaca sylvanus), living in two troops (i.e. “Flat face” and “Large” troop) in the middle‐Atlas Mountains, Morocco. The four females lost their infants due to predation or for unknown causes. Self‐suckling was observed before and after the infants died in the four females living in the “Flat face” troop. When the infants were still alive, self‐suckling was of short duration and it was probably a method to improve milk flow when the infant switched from one nipple to the other. After the infants died, self‐suckling lasted significantly longer and the females were apparently drinking their own milk. Self‐suckling was never observed among the four lactating females in the “Large” troop (including one monkey who lost her infant) and it could thus represent a cultural difference. Moreover, self‐suckling after the death of an infant may be explained by the energetic and immunological benefits that a monkey may gain from drinking their own milk. Finally, self‐suckling may have a stress‐releasing effect on the mothers who have lost their infants. Am J Phys Anthropol, 2009. © 2009 Wiley‐Liss, Inc.     

Bioassay approaches to observing behavioural responses of adult female Cydia molesta to host plant odour

Abstract:  Three different olfactometers were evaluated in order to develop a bioassay procedure testing for the olfactory responses of Cydia molesta . Females were tested individually using a linear and a Y-tube olfactometer, and in groups using a dual-choice arena. Room temperature, relative humidity, light intensity, airflow, age of the moths and odour source were kept constant during experiments. The odour source tested was a green shoot of peach excised 10 min before experiments started. Cydia molesta females showed a significant response to this plant odour in all olfactometers. A number of qualitative aspects were found to be in favour of the dual choice arena as a tool for screening potentially attractive odour sources. It allowed for a differentiation of the response of mated and virgin females. Experimental conditions allowed the circadian rhythm of insects to be mimicked. Manipulation of individuals is reduced and flight is not precluded.     

The attraction of virgin female hide beetles (Dermestes maculatus) to cadavers by a combination of decomposition odour and male sex pheromones

ABSTRACT: INTRODUCTION: The hide beetle Dermestes maculatus (Coleoptera: Dermestidae) feeds as an adult and larva on decomposing animal remains and can also be found on human corpses. Therefore, forensic entomological questions with regard to when and how the first receptive females appear on carcasses are important, as the developmental stages of their larvae can be used to calculate the post-mortem interval. To date, we know that freshly emerged males respond to the cadaver odour of post-bloated carcasses (approximately 9 days after death at Tmean = 27degrees), being attracted by benzyl butyrate. This component occurs at its highest concentration at this stage of decay. The aim of our study was to determine the principle of attraction of virgin females to the feeding and breeding substrate. For this purpose, we tested the response of these females to headspace samples of piglet cadavers and male sex pheromones [(Z9)-unsaturated fatty acid isopropyl esters] in a Y-olfactometer. Because we expected that such an odour combination is of importance for virgin female attraction, we tested the following two questions: 1) Are virgin female hide beetles attracted by a combination of cadaver odour and male sex pheromones? 2) During which decomposition stage do the first virgin females respond to cadaver odour when combined with male sex pheromones? RESULTS: We found that young virgin females were attracted to the cadaver by a combination of cadaver odour and male sex pheromones. Neither cadaver odour alone nor male sex pheromones alone was significantly more attractive than a solvent control. Our results also gave a weak indication that the first young virgin females respond as early as the post-bloating stage to its associated decomposition odour when combined with male sex pheromones. CONCLUSIONS: Our results indicate that freshly emerged males possibly respond to cadaver odour and visit carcasses before virgin females. Being attracted to cadavers when male sex pheromone is perceived as well, virgin females can optimise their reproductive possibilities.     

Field energetics of free-living, lactating and non-lactating echidnas (Tachyglossus aculeatus)

We measured daily energy expenditure (DEE) and water turnover rates in lactating and non-lactating short beaked echidnas (Tachyglossus aculeatus) using the doubly labelled water technique during the lactation period in spring. Reproductively inactive echidnas were on average significantly heavier (median: 3354 g; range: 2929-3780 g; N=4) than lactating females (median: 2695 g; range: 2690-2715 g; N=3) during the equivalent time period. The median water flux rate of lactating echidnas (152 ml day(-1); range: 120-198 ml day(-1)) did not differ significantly from that of non-lactating females (170 ml day(-1); range: 128-227 ml day(-1)). The median DEE of echidnas that were lactating was 645 kJ day(-1) (range: 581-850 kJ day(-1)), which was not different from the median DEE of non-reproductive control females (763 kJ day(-1); range: 720-766 kJ day(-1)). Lactating females somehow compensate for the energy costs of milk production, resulting in a daily energy budget that is not different from that of non-reproductive females. At least part of their energy minimising strategy could involve the use of moderate heterothermy, allowing a greater proportion of daily energy expenditure to diverted to milk production.     

Lactation in the Japanese monkey (Macaca fuscata): Yield and composition of milk and nipple preference of young

Changes in the yield and chemical composition of milk, together with those in the nipple preferences and suckling behavior of infants, were followed in three lactating Japanese monkeys rearing single infants under laboratory conditions at monthly intervals for six months after their paturition. Milk accumulating in the mammary glands during a 4-hr separation of the infant was collected by milking under anesthesia with the aid of a physiological dose of oxytocin. The stage of full lactation appeared to last for about two or three months after parturition. The milk at this stage contained 14.0% of total solids, 4.2% of lipids, 1.6% of proteins, and 6.2% of lactose, and the concentrations of Na, Cl, and K in the milk water were 7.4, 15.1, and 5.9 mM, respectively. From the 9th or 13th week of lactation onwards, according to the individual, the milk composition changed appreciably and the time during which the young left hold of the mother's nipple also increased. The weaning of young in the Japanese monkey appears to begin during the 3rd or 4th month of age. The milk production then wanes and almost ceases by the end of the 6th month. The nipple preference of the young was generally established completely by the 3rd, or the 8th at latest, week of age. However, no difference in either the yield or composition of the milk secreted was found between the mammary glands of the preferred and non-preferred sides. Without regard to the apparent nipple preference, infants seem to ingest milk from the mother's breasts of both sides. Accurate estimation of the rate of milk production could not be achieved, but the rate was expected to be 150–200 g/day at the height of lactation based on the relation between the mother's body weight and the milk production rates of other primate species. A disparity was noted between the observed and estimated values for the milk yield, and a diurnal fluctuation in the milk secretion of the Japanese monkey was inferred. This study was conducted through the official system of the Primate Research Institute, Kyoto University, for cooperative research work with other institutions.     

Human toxoplasmosis acquired by ingestion of goat's milk

Acute toxoplasmosis in three members of the same family was related to drinking unpasteurized goat's milk. The goats were reared, for milk production in a peri-domestic area. Based on indirect immunofluorescent reactions antibody titers greater than 1: 1024 were recorded in eight out of fifteen animals examined and were highest in the five lactating adult females. Toxoplasma was isolated, by inoculation of mice, from the milk of one of these females. Dogs reared in the same house showed no symptoms of acute toxoplasmosis and low antibody titers were detected in these animals. It was also found that the human infections could not have been due to the ingestion of food contaminated with oocysts.     

Non-lactating versus lactating females: a comparison of sex steroids,sexual coloration,and sexual behavior in Japanese macaques

Female Japanese macaques are seasonal breeders distinguished by their red-colored hindquarters, face, and nipple skin areas. Intensity of coloration seems to be associated with sexual attractiveness, behavior, and fluctuating sex steroids. Our aim was to investigate whether the color intensity of these regions differed between lactating (LA) and non-lactating (NLA) females during sexually inactive (SI) and active (SA) phases. Coloration scores of 19 adult females were classified using color tables. Estrogen and progesterone metabolites were determined in fecal samples. Weekly comparison between both groups revealed significantly increased coloration of the hindquarters area from week 13 (SI) until the end of the observation period, and for the nipple skin throughout the SI and SA periods. Face coloration differed marginally. Hormonally, NLA females showed significantly increased excretion rates of sex steroids at the end of the SI phase and throughout the whole SA period. Logistic regression analyses between elevated fecal steroids and nipple coloration disclosed a significant relationship for NLA females during the SI period. This connection persisted and included hindquarter coloration during the SA period. NLA females showed increased intromission with ejaculation, but no difference was found for intromission without ejaculation. In conclusion, results demonstrate increased endocrine excretion rates for NLA females during the whole observation period, paralleled by an enhanced, fertility-signaling sexual attractiveness.     

The energetics of lactation in cooperatively breeding meerkats Suricata suricatta

Species may become obligate cooperative breeders when parents are unable to raise their offspring unassisted. We measured the daily energy expenditure of mothers, helpers and offspring during peak lactation in cooperatively breeding meerkats Suricata suricatta using the doubly labelled water technique. Lactating mothers expended more energy per day than allo-lactating subordinate females, non-lactating females or suckling offspring. Metabolizable energy intakes of lactating mothers were calculated from isotope-based estimates of offspring milk energy intake, and were not significantly different from the previously suggested maximal limit for mammals. Allo-lactating females were the only category of animals that lost weight during the period of study, probably because they spent more time babysitting than non-lactating females. Daily energy expenditure (DEE) of lactating mothers increased with litter size but decreased with the number of helpers. Calculations show that for every 10 helpers, even in the absence of allo-lactators, mothers are able to reduce their DEE during peak lactation by an amount equivalent to the energy cost of one pup. These results indicate that helpers have beneficial energetic consequences for lactating mothers in an obligate cooperatively breeding mammal.     

The effect of dietary fat content on lactation energetics in the European hare (Lepus europaeus)

European hares selectively feed on plants with high fat and hence energy content. We hypothesized that these dietary requirements limit the ability of hares to adjust daily food intake during periods of high energy requirements, namely lactation. Our measurements in captive lactating females show that does kept on a low-fat diet increased food intake compared to does on a high-fat diet but assimilated significantly lower amounts of energy. Further, does fed a low-fat diet showed a prolonged rise of food intake during lactation, reduced milk energy content and lower milk mass production at large litter sizes. We hypothesize that impaired milk production under suboptimal fat supply is due to the inability of females to increase the capacity of nutrient-processing organs rapidly enough to meet the high energy demands of precocial juveniles with high metabolic costs. Thus, in hares, the production of precocial young may be viewed as a constraint, caused by their inability to dig thermally buffered burrows, rather than as an adaptive reproductive strategy. We suggest that the interaction of lactation energetics, dietary requirements, and reduced plant diversity in modern agricultural landscapes has facilitated the decline of hare populations across Europe over the last decades.     

Olfactory memory in the mosquito Culex quinquefasciatus

The cosmotropical urban mosquito Culex quinquefasciatus Say (Diptera: Culicidae) uses chemical cues to locate suitable water pools for oviposition. Although gravid females are innately attracted to or repelled by certain compounds, this study found that an individual mosquito's preferences for these odours could be altered greatly by prior experience. Mosquitoes reared in water containing skatole, at a level normally repellent to ovipositing females, preferred to oviposit in water containing that compound rather than in water with an otherwise attractive odour compound (P-cresol). This behaviour occurred regardless of whether mosquitoes were tested individually or in groups of up to 50 per cage. The F1 progeny of conditioned mosquitoes did not exhibit the parental preference, but were as susceptible to conditioning as their parents. Moreover, rearing mosquitoes in infusions of hay or animal (guinea-pig) faeces produced a similar although less dramatic change, such that the innate propensity for hay infusion could be cancelled by rearing in guinea-pig faeces infusion. The results demonstrated a change in odour preference by Cx. quinquefasciatus following exposure to the odour during development or pupal eclosion, suggesting that some form of larval conditioning or early adult imprinting occurred. Precisely when that conditioning occurred remains to be determined.     

Olfactometer Responses of Plum Curculio <Emphasis Type="Italic">Conotrachelus nenuphar</Emphasis> (Herbst) (Coleoptera: Curculionidae) to Host Plant Volatiles,Synthetic Grandisoic Acid,and Live Conspecifics

The plum curculio Conotrachelus nenuphar (Herbst) (Coleoptera: Curculionidae) is a major pest of pome and stone fruit, but will also attack other fruits. Males produce the aggregation pheromone grandisoic acid; emitting only the (+)-enantiomer which is attractive to both sexes of the univoltine and multivoltine strains, while the synthetic racemic mixture contains optical isomers with equal amounts of (+)- and (?)-enantiomers. Synergy between odours can increase trap captures and improve monitoring techniques, therefore tests were performed in a dual-choice olfactometer with odours attractive to plum curculios according to literature to determine 1) under what physiological conditions (mating status, age, starvation period) these odours are attractive or repulsive, 2) if the (+)-enantiomer or the odour of live males synergizes with host plant volatiles, and 3) if there is a difference in response between plum curculio strains. Females were exposed to: benzaldehyde; trans-2-hexenal; apples; extracts of: plums, apples, blueberries; grandisoic acid; and live males. Plum essence was found to be the most attractive host-plant odour for both immature and mature virgin females, and immature whole apples were attractive to starved females, while trans-2-hexenal, McIntosh apple essence, benzaldehyde along with the combination of benzaldehyde and plume essence was found to be repulsive. Starvation, age, and mated status all influence response to odours. No synergistic or additive affects were observed between any of the odour combinations tested, including the combination of both the natural and synthetic pheromone and plum essence or apples.     

Accurate spatial and temporal transgene expression driven by a 3.8-kilobase promoter of the bovine β-casein gene in the lactating mouse mammary gland

The spatial, temporal, and hormonal pattern of expression of the β-casein gene is highly regulated and confined to the epithelial cells of the lactating mammary gland. Previous studies have shown that 1.7 kb of the bovine β-casein promoter were able to drive cell-specific and hormone-dependent expression to a mouse mammary cell line but failed to induce accurate expression to the mammary gland of transgenic mice. We investigated here the ability of 3.8 kb of the bovine β-casein gene promoter to drive the expression of the human growth hormone (hGH) gene in transgenic mice. A Northern blot analysis using total RNA obtained from different tissues of lactating and nonlactating females revealed the presence of hGH mRNA only in the mammary gland of lactating females. hGH mRNA was not detectable in the mammary gland of virgin females or males. A developmental analysis showed that hGH mRNA only peaked on parturition, resembling more closely the bovine β-casein temporal expression pattern rather than the murine. In situ hibridization studies performed on mammary gland sections showed that the cellular pattern of hGH expression was homogeneous in all lobules from heterozygous and homozygous transgenic mice. Silver grain counts on the tissue sections highly correlated with the hGH contents in the milk determined by radioimmunoassay (r = 0.996). Thus 3.8 kb of the bovine β-casein promoter direct a high-level expression of a reporter gene to the lactating mammary gland of transgenic mice in a tissue-specific and developmentally regulated manner. Mol. Reprod. Dev. 49:236–245, 1998. © 1998 Wiley-Liss, Inc.     

Suckling behaviour and fertility in rhesus macaques (Macaca multatta)

Lactation has been proposed as the main proximate mechanism regulating female fertility among non–seasonally breeding mammals. Less is known about the effects of lactation upon fertility among seasonal species, where photoperiodic control of reproduction is widely believed to have an overriding influence. In neither case is it clear which of the different components of lactation influence fertility and in which ways. In this paper, the links between suckling patterns and the likelihood of conception during the following breeding season are examined. Data were collected at the Madingley rhesus colony, where reproduction is strictly seasonal. It is shown that fertility was depressed in lactating females when compared to non–lactating ones. Within the lactating group, mothers who suckled their infants frequently, and with a great number of independent nipple contacts in each bout, failed to conceive during the following breeding season. It is suggested that early differences in suckling patterns might reflect differences in maternal condition; and that differences during the breeding season may affect the quality of the ovulations. A considerable decrease in both the frequency of suckling and the number of nipple contacts per bout seemed to be required for conception to occur. On the other hand, neither the duration of the suckling bouts, nor the duration of the intervals between bouts, were found to be associated with the likelihood of conception.     

Comparison of the "nursing" and other parental behaviors of nulliparous and lactating female rats

Virgin female rats display maternal behaviors after continuous exposure to pups (sensitization) that are in some respects similar to those of postpartum females. We herein provide a detailed comparison of the "nursing" and other parental behaviors of maternally sensitized virgin females and postpartum lactating dams. Ovariectomized and intact virgin females were exposed to pups until displaying maternal behavior. On the females' fourth day of maternal responsiveness, the pups were removed for 3 h and then returned, and subject-litter interactions were observed for 45 min. Behavior of maternal virgins was compared with that of lactating dams observed on day 4 postpartum interacting with either suckling pups or pups unable to suckle due to perioral anesthesia. Ovariectomy had no effect on behavior of virgins. Retrieval and licking of pups were deficient in virgins compared with lactating dams. Suckled dams showed prolonged kyphosis (upright crouched nursing), whereas nonsuckled dams displayed little kyphosis but rather were often in a hunched position over pups. Some aspects of quiescent "nursing" behaviors of virgins were surprisingly similar to those of suckled dams, including the latency to and duration of quiescence. Nonsuckling pup stimulation elicited more kyphosis in virgins than in lactating dams, which was still much less than in suckled dams. Virgins also "nursed" pups in hunched and prone postures. Differences between sensitized and postpartum females in their maternal behaviors likely reflect differences in motivation as well as sensory inputs they receive from pups. In particular, sensory regulation of "nursing" behaviors is influenced by reproductive state because nonsuckling pups elicit different postural responses in sensitized and lactating mothers.     

Effect of milkflow rate and presence of a floating nipple on abnormal sucking between dairy calves

The aim of this study was to investigate if access to an artificial teat compared to an open bucket would decrease abnormal sucking in calves held in pairs, and if the calves would perform less abnormal sucking if they spent more time drinking or sucking the milk. In total 16 calves of Swedish Red and White cattle housed in pairs were used. The eight groups were given 2.5l of whole milk per calf, twice a day in one of the following treatments; bucket with fast flow (control), bucket with slow flow, floating nipple with fast flow and floating nipple with slow flow. All groups were subjected to the four different treatments in a random balanced order. Two buckets were connected with a rubber pipe in order to control the milk flow. In the fast treatments, the milk poured from one bucket to the other in 40s, and in the slow treatments the milk poured from one bucket to the other in 10min. Each group had the treatment for 6 days before behavioural observations were done on day 7. These were made by instantaneous recordings at 20s intervals for 30min, starting when the calves received their milk in the morning and afternoon. The number of recordings of abnormal sucking was lower than the control in all treatments (P<0.001). The lowest recording of abnormal sucking was found, when the calves had both a floating nipple and a slow flow. Sucking on other objects in the pen had the highest number of recordings, when they had a nipple and a fast flow (P<0.01), but the sucking on the empty nipple represented 95% of this category. The calves tended to lick and bite more on the bucket, the nipple and pen fittings in the two treatments with fast flow (P<0.1) than in the slow treatment with nipple. It is concluded that both the performance of sucking and the time taken to ingest milk are important in order to decrease abnormal sucking between calves.     

Maternal Effort is State Dependent: Energetic Limitation or Regulation?

Many small altricial rodents have a postpartum oestrus and are often simultaneously pregnant and lactating. Negative influences of concurrent pregnancy and lactation on both lactational performance and the litter in utero are commonly observed and have been interpreted as resulting from high simultaneous energetic demands of gestation and lactation. We studied these effects in the precocial guinea‐pig (Cavia aperea f. porcellus) that, like many altricial rodents, has a postpartum oestrus, but in which the peaks of energy expenditure on lactation and gestation are widely separated. This life history allowed to investigate whether physiological regulation other than by energetic limitations may be responsible for allocation conflicts arising when lactation and gestation overlap. By comparing simultaneously pregnant and lactating females with lactating non‐pregnant females, we show that females in the former group nurse less and wean earlier than females of the latter group. In a comparison of litter size, litter mass, and pup mortality of females that had not been lactating during pregnancy with females that had been simultaneously pregnant and lactating, we show that the latter do not reduce investment in the following litter. In our study, energetic constraints on ad libitum fed females are unlikely and we therefore suggest that the results must be explained by regulatory constraints on lactational effort. We point out that this explanation has not been excluded for the effects observed in altricial small mammals.