Consistent behavioral phenotype differences between inbred mouse strains in the IntelliCage.

The between-laboratory effects on behavioral phenotypes and spatial learning performance of three strains of laboratory mice known for divergent behavioral phenotypes were evaluated in a fully balanced and synchronized study using a completely automated behavioral phenotyping device (IntelliCage). Activity pattern and spatial conditioning performance differed consistently between strains, i.e. exhibited no interaction with the between-laboratory factor, whereas the gross laboratory effect showed up significantly in the majority of measures. It is argued that overall differences between laboratories may not realistically be preventable, as subtle differences in animal housing and treatment will not be controllable, in practice. However, consistency of strain (or treatment) effects appears to be far more important in behavioral and brain sciences than the absolute overall level of such measures. In this respect, basic behavioral and learning measures proved to be highly consistent in the IntelliCage, therefore providing a valid basis for meaningful research hypothesis testing. Also, potential heterogeneity of behavioral status because of environmental and social enrichment has no detectable negative effect on the consistency of strain effects. We suggest that the absence of human interference during behavioral testing is the most prominent advantage of the IntelliCage and suspect that this is likely responsible for the between-laboratory consistency of findings, although we are aware that this ultimately needs direct testing.

Sexual growth dimorphism affects birth sex ratio in house mice.

We present the first empirical evidence that mammalian sex-ratio deviations result from variation in adult-weight sexual dimorphism via correlated effects on blastocyst development. Two selection lines of mice exhibiting high and low sexual dimorphism in adult weight showed correlated sexual weight differences at birth and at weaning, caused by relatively decelerated growth of males in the low line from before birth. The sex ratio at birth was significantly female-biased in the low line, and significantly lower than in the highly dimorphic line. Concomitantly, blastomere numbers were at significantly higher variance in the low than in the highly dimorphic line, owing to an increased frequency of slowly growing blastocysts. Since low-dimorphism mice produced more corpora lutea and more female pups than the high-dimorphism mice, but not more males, birth sex-ratio bias most parsimoniously resulted from the loss of slowly growing male blastocysts. This is in agreement with the observation that sex-ratio skews in mammals arise when timing of uterine responsiveness (i.e. its temporally limited capacity for implantation) varies in relation to sex-specific embryonic growth rates. Hence, natural mammalian sex-ratio variation that stems from developmental asynchrony might be a by-product of natural selection for sexual dimorphism in adult weight.

Timing of mating, developmental asynchrony and the sex ratio in mice.

According to the developmental asynchrony hypothesis, changing the time of mating within the estrous cycle could alter the interval between completion of blastocyst development and uterine responsiveness for implantation. This may then lead to sex ratio skews in animals that exhibit sex-differential blastocyst development, because uterine stage may now benefit either slow (female) or fast (male) developing blastocysts. To test this hypothesis, the responses of two strains of mice to altered mating dynamics were compared. In a strain that exhibits higher male than female blastocyst developmental rates, sex ratios became significantly female-biased when mated late during the estrous cycle as opposed to early mating. However, timing of mating did not affect sex ratios in a strain with synchronous development of male and female preimplantation embryos. Hence, it is concluded that developmental asynchrony between male and female blastocysts on the one hand, and blastocysts and uterus on the other, are indeed responsible for the effect of timing of mating on litter sex ratios in mice.

Design variability in web geometry of an orb-weaving spider.

We studied the effect of several variables (environmental and physiological) on web geometry in the garden cross spider Araneus diadematus. Variables were: web support, wind, temperature, humidity, and silk supply. All had an effect. The spiders generally attempted to fit their webs to the shape of the supporting frame (standard, small, vertical, or horizontal). Windy conditions (0.5 m s-1) during web construction caused spiders to build smaller and rounder webs, laying down fewer capture spirals while increasing the distances between capture-spiral meshes. Decreasing temperature from 24 degrees to 12 degrees C caused the capture spiral to have fewer and wider spaced meshes, which did not change overall capture area but reduced the length of capture-spiral threads laid down. Subsequent increase of temperature to 24 degrees C restored the number of meshes laid down, but the wider mesh was retained, causing the capture area to be increased over initial control values. Decreased humidity (from 70 to 20% rH) had the effect of reducing web and capture-spiral size, the latter by reducing mesh number while keeping mesh spacing constant. Subsequent increase of humidity to control level (70%) restored web and capture area. However, this was achieved by laying down capture meshes at larger distances, rather than returning to initial mesh numbers. Silk supply also had a strong effect. Webs built in unnaturally rapid succession by the same spider (4 in 24 h when 1 is the norm) became sequentially smaller, had fewer radii, shorter capture spirals, and were wider meshed.

Effects of mating dynamics and crowding on sex ratio variance in mice.

Mating units of six virgin females and one adult stud male were established to test for the effects of timing of mating and crowding of pregnant females on litter sex ratios in mice. Females either copulated during periods when no other female of the mating unit copulated simultaneously (single mating condition) or when more than one female copulated (multiple matings condition). Two crowding conditions were imposed on the animals: the females of 14 mating units were placed into individual cages after mating (isolated condition), while females of the other 13 mating units remained in the original group until shortly before littering (crowded condition). Sex ratio variance did not deviate from random expectation in litters arising from the multiple matings periods. However, in litters arising from single mating periods, extreme sex ratios were found significantly less frequently than expected by chance. Higher sex ratio variance in litters arising from multiple matings periods is attributed to the timing of mating being at higher variance under this condition, which is known to affect sex ratios in other rodents. Crowding significantly reduced sex ratio variance further. Reduced sex ratio variance under single mating and crowded conditions is speculated to follow from competition for resources between preimplantation embryos, which may be further increased by stressful effects of crowding. Loss of embryos after implantation appeared not to be responsible for the above effects.

The developmental asynchrony hypothesis for sex ratio manipulation.

This paper presents the hypothesis that developmental asynchrony between embryos of both sexes, on the one hand, and offspring development and responsiveness of maternal reproductive organs, on the other, may result in significant sex ratio biases at birth in mammals, and in birds. It is argued that the developmental asynchrony hypothesis may account for a significant proportion of findings on sex ratio skews, including the inconsistency of evidence and the lack of successful selection attempts. This mechanism could be used by the maternal organism adaptively to alter the sex ratio by control of the relative time of insemination within the oestrous cycle or by alteration of reproductive hormone levels around the time of ovulation and early embryonic development. However, the developmental asynchrony hypothesis may also imply that many of the sex ratio biases can be viewed as by-products of constraining selection on control of fertility and fecundity and on sexual dimorphic growth rates.

Potential mechanisms for sex ratio adjustment in mammals and birds.

Sex ratio skews in relation to a variety of environmental or parental conditions have frequently been reported among mammals and, though less commonly, among birds. However, the adaptive significance of such sex ratio variation remains unclear. This has, in part, been attributed to the absence of a low-cost physiological mechanism for sex ratio manipulation by the parent. It is shown here that several recent findings in reproductive biology are suggestive of many potential pathways by which gonadotropins and steroid hormones could interfere with the sex ratio at birth. And these hormone levels are well-known to be influenced by many parameters which have been invoked in correlating with offspring sex ratios. Hence, it is argued that the significant, but inconsistent sex ratio biases reported in mammalian and avian populations are coherent with current knowledge on reproductive physiology in those species. However, whether such variations can be viewed at as a consequence of physiological constraint or as adaptive sex ratio adjustment, has still to be determined.

Sex ratio manipulation in wild house mice: the effect of fetal resorption in relation to the mode of reproduction.

First generation laboratory-born descendants of wild-caught house mice (Mus musculus domesticus Rutty) were bred to produce litters of primipares and of dams that had conceived a second litter either after lactational anestrus or within the postpartum estrus. At the day of birth, pups were sexed and the number of implanted and resorbed embryos was determined to evaluate the influence of mode of reproduction on litter gender composition and its relation to fetal resorption. No significant deviations from an even sex ratio occurred in the sample. The results indicate that primipares produced litters with subnormal dispersion of the gender distribution, but this could not unequivocally be attributed to fetal resorption. No significant bias in the litter gender composition was detectable within litters conceived after lactational anestrus. In contrast, the dispersion of the gender distribution was significantly supernormal in the litters of dams inseminated at postpartum estrus. Within this group, fetal resorption had a significant effect upon the sex ratio, and this relationship was significantly affected by the number of implanted embryos: resorbing dams produced male-biased litters at small and intermediate numbers of implantation sites and female-biased litters when the number of implanted embryos was large. It is argued that this is most likely attributable to sex-selective fetal resorption.

Sex ratio and litter size in relation to parity and mode of conception in three inbred strains of mice.

Breeding records from three inbred strains of mice (BALB/c ABom, C57BL/10ScSn, C3H/He/Kon) were examined with respect to the effects of parity and mode of conception upon litter size and sex ratio at birth. Litters from 3 modes of conception were considered: litters of primipares, litters of multipares conceived during postpartum oestrus and litters conceived after lactational anoestrus. Litters of multipares were assigned to one of these latter groups according to the inter-litter intervals. Parity had no significant effect upon the sex ratio but had a significant one upon the litter size, which did not vary between the strains when first litters were excluded from analysis. The expected variations in response to the mode of conception were found in BALB/c ABom mice but both the effects on the litter size as well as on the sex ratio varied significantly between the strains. Litter size reduction per se could be ruled out to be the cause of the sex ratio variations found. Rather, it is suggested that sex-specificity of embryonic loss depends upon the mode of conception.

Sex-specific embryonic mortality during concurrent pregnancy and lactation in house mice.

To evaluate the importance of lactational stress for sex ratio manipulation in postpartum inseminated house mice, 163 sexually inexperienced male and female BALB/c ABom inbred mice were mated and stud males were removed the day following the initial parturition. Randomly chosen dams were allowed to suckle their young for either one or 14 or 21 days. The results showed that two weeks of lactation enhance the incidence of total abortion of the litter conceived postpartum. If lactation continued another week, dams which gave birth to a second litter lost relatively more embryos during uterine development and the percentage of lost embryos was increased by the number of pups suckled. Among these dams, the number of embryos lost in utero correlated positively with viable litter sex ratios (percentage male pups) at second parturition, indicating that lactational stress enhances embryonic mortality and that this additional mortality predominantly affects female embryos.

Effect of food restriction on reproduction and lactation in house mice mated post partum.

Primiparous, post-partum mated BALB/c bom inbred mice were allowed to raise litters of 6 young until Day 22 of lactation: 11 of 25 females were restricted to 60% of food consumption of ad-libitum fed dams after stud male removal at Day 2 of lactation. Since weight gain of restricted females during lactation was not inhibited and infanticidal behaviour was not enhanced, food deprivation can be considered to have been relatively mild. However, none of the food-restricted dams gave birth to a second litter whereas 12 of the 14 ad-libitum fed mice littered. This pregnancy failure is suggested to be due to implantation failure or abortion shortly after implantation, which is attributed to maternal manipulation rather than to immediate consequences of energetic demands of lactated young. The dynamics of the weight changes of dams and young suggest that milk production in suckling house mice drops most markedly between Days 17 and 18 of lactation, irrespective of whether the dams are non-pregnant, pregnant, or food-restricted.