Modelling sub-binomial variation in the frequency of sex combinations in litters of pigs

In some samples of pig litters, the number of males per litter exhibits significant sub-binomial dispersion: there are fewer unisexual and more "sex-balanced" litters than expected under a binomial model. It is argued here that the explanation is that the sex of zygotes may be subject to hormonal control with the consequence that, within a litter, the zygotes do not all have the same probability of being male. This variation in probability is approximated by a quadratic function of the time during oestrus that a zygote is formed so as to parallel the variation in maternal hormone levels. Other explanations are examined briefly, but each seems to have serious weaknesses.     

Ecological genetics of the rotifer Brachionus plicatilis in Soda Lake,Nevada, USA

In an earlier study we found that the rotifer Brachionus plicatilis in Soda Lake, Nevada (USA) maintains extensive genetic variation for three isozymes. This variation was present both within and between different collections. No evidence was found that there was a seasonal succession of distinct electromorphs. In this paper we ask: is the genetic variation maintained by selective neutrality among electromorphs? Parthenogenetic females from ten clones representing seven different composite electromorphs were selected for life table analysis. Substantial main effects of genotype or interactions of genotype with experimental food quantities and temperatures, were found for the intrinsic rates of increase, net reproductive rates, and mean life spans of these clones. We conclude that these fitness differences permit us to reject the hypothesis that genetic variation is maintained because of selective neutrality among clones. Instead, it seems likely that directional selectional may be either too weak or of too brief a duration in Soda Lake for clonal exclusion to occur.     

Sexual Antagonism,Temporally Fluctuating Selection,and Variable Dominance Affect a Regulatory Polymorphism in Drosophila melanogaster

Understanding how genetic variation is maintained within species is a major goal of evolutionary genetics that can shed light on the preservation of biodiversity. Here, we examined the maintenance of a regulatory single-nucleotide polymorphism (SNP) of the X-linked Drosophila melanogaster gene fezzik. The derived variant at this site is at intermediate frequency in many worldwide populations but absent in populations from the ancestral species range in sub-Saharan Africa. We collected and genotyped wild-caught individuals from a single European population biannually over a period of 5 years, which revealed an overall difference in allele frequency between the sexes and a consistent change in allele frequency across seasons in females but not in males. Modeling based on the observed allele and genotype frequencies suggested that both sexually antagonistic and temporally fluctuating selection may help maintain variation at this site. The derived variant is predicted to be female-beneficial and mostly recessive; however, there was uncertainty surrounding our dominance estimates and long-term modeling projections suggest that it is more likely to be dominant. By examining gene expression phenotypes, we found that phenotypic dominance was variable and dependent upon developmental stage and genetic background, suggesting that dominance may be variable at this locus. We further determined that fezzik expression and genotype are associated with starvation resistance in a sex-dependent manner, suggesting a potential phenotypic target of selection. By characterizing the mechanisms of selection acting on this SNP, our results improve our understanding of how selection maintains genetic and phenotypic variation in natural populations.     

Does bird predation influence the spot-number variation in Maniola jurtina (Lepidoptera)?

For six years samples of the satyrine butterfly Maniolajurtina L. were collected on small islands in southern Sweden and scored for beak marks, i.e. damage on wings presumably resulting from attacks by birds; in the present study area mainly Red-backed Shrike Lanius collurio L. An overall mean of 8% (5–1496 in different years) of the females and 1396 (10–2296) of the males had beak marks. Both sexes of M. jurtina show variation in spot-number on the hindwings and there is some evidence that in some years the frequencies of different spot-number phenotypes with beak marks differ significantly from randomness. Although it is not known how beak mark frequencies are related to predation pressure (reflecting rates of predation or escapes?) it is suggested that, due to behavioural differences of spot-number phenotypes, birds act as a selective factor influencing the spot-number variation. The results are discussed in relation to a recent model proposed by Brakefield attempting to account for variation in spot-number in M. jurtina.     

Analysis of variance of first-year growth in captive garter snakes (Thamnophis elegans) by family and sex

Although body size in reptiles has important consequences for their ecology, it is quite variable in most species, presumably because growth in these animals is influenced by many factors. In this study, we analyse the effects of family membership and sex on first-year growth of garter snakes (Thamnophis elegans) held under identical conditions in the laboratory. Litters, but not the two sexes, differed significantly in mean size at birth. Larger snakes ate more food and had a higher absolute (but not relative) growth rate in length than smaller snakes. Thus, there was significant variation in length increase among litters, but not between the sexes. All else being equal, variance in size at birth could contribute significantly to variance in adult body size.     

From breeding pairs to fox towns: the social organisation of arctic fox populations with stable and fluctuating availability of food

Food availability can impact group formation in Carnivora. Specifically, it has been suggested that temporal variation in food availability may allow a breeding pair to tolerate additional adults in their territory at times when food abundance is high. We investigate group occurrence and intraspecific tolerance during breeding in a socially flexible canid, the arctic fox (Vulpes lagopus). We compare Iceland and Sweden where resource conditions differ considerably. A breeding pair was the most common social unit in both populations, but as predicted, groups were more frequent where food abundance varied substantially between years (Sweden: 6 %) than where food availability was stable (Iceland: ≤2 %). Within Sweden, supplemental feeding increased group occurrence from 6 to 21 %, but there was no effect of natural variation in lemming (Lemmus lemmus) availability since group formation was rare also at lemming highs. Thus, additional factors appeared to influence the trade-off between intraspecific territoriality and tolerance. We report two cases where related females showed enduring social relationships with good-neighbour strategies. Related females also engaged in alloparental behaviour in a ‘fox town’ with 31 foxes (4 adults, 3 litters). In contrast, when unrelated foxes bred close to each other, they moved or split their litters during summer, presumably because of territorial conflict. We suggest that fluctuating food availability is linked to group formation in this Arctic carnivore, but also when food availability increases, additional factors such as relatedness, alloparental benefits, competition and predator defence appear necessary to explain group formation.     

FINE-GRAINED SPATIAL AND TEMPORAL VARIATION IN SELECTION DOES NOT MAINTAIN GENETIC VARIATION IN ERIGERON ANNUUS

Because interactions among plants are spatially local, the scale of environmental heterogeneity can have large effects on evolutionary dynamics. However, very little is known about the spatial patterns of variation in fitness and the relative magnitude of spatial and temporal variation in selection. Replicates of 12 genotypes of Erigeron annuus (Asteraceae) were planted in 288 locations within a field, separated by distances of 0.1 to 30.0 m, and replicated in two years. In a given year, most spatial variation in relative fitness (genotype-environment [G × E] interactions for fitness) occurred over distances of only 50 cm. Year effects were as large or larger than the spatial variation in fitness; in particular there was a large, three-way, genotype-year-environment interaction at the smallest spatial scale. The genetic correlation of fitness across years at a given location was near zero, 0.03. Thus, the relative fitness of genotypes is spatially unpredictable and a map of the selective environment has constantly shifting locations of peaks and valleys. Including measurements of soil nutrients as covariates in the analysis removed most of the spatial G × E interaction. Vegetation and microtopography had no effect on the G × E terms, suggesting that differential response to soil nutrients is the cause of spatial variation in fitness. However, the slope of response to NH₄ and P0₄ was negative; therefore the soil nutrients are probably just indicators of other, unknown, environmental factors. We explored via simulation the evolutionary consequences of spatial and temporal variation in fitness and showed that, for this system, the spatial scale of variation was too fine grained (by a factor of 3 to 5) to be a powerful force maintaining genetic variation in the population. The inclusion of both spatial and temporal variation in fitness actually reduced the coexistence of genotypes compared to pure spatial models. Thus the presence of spatial or temporal variation in selection does not guarantee that it is an effective evolutionary force maintaining diversity. Instead the pattern of selection favors generalist genotypes.     

Genetic determinism for within-litter birth weight variation and its relationship with litter weight and litter size in the Ripollesa ewe breed

Birth weight plays a central role in lamb survival and growth, and the knowledge of its genetic determinism has become essential in worldwide selection programmes. Within this context, within-litter birth weight variation (BWV) has been suggested as an attractive trait to homogenise litters in prolific species, although it has not been analysed in sheep. The objective of this study was to ascertain whether maternal additive genetic variance exists for BWV in Ripollesa ewes, and to study its genetic, permanent environmental and residual relationships with litter weight (LW) and litter size (LS) at birth. Data were recorded in the Ripollesa experimental flock of the Universitat Autònoma of Barcelona, between 1986 and 2005, and included 1 662 litters from 380 ewes, with 712 records of BWV and 1 530 records of LW. Traits were analysed with a multivariate animal model solved through Bayesian methodologies, and with a threshold characterisation of LS. Additionally, the effect of BWV on lamb survival was studied. Additive genetic variance was observed for BWV (h² = 0.061), as well as for LW (h² = 0.200) and LS (h² = 0.141). Nevertheless, genetic correlations among those traits were not substantial (BWV and LW = 0.151; BWV and LS = − 0.219; LW and LS = − 0.320) and suffered from a high degree of uncertainly, with the null correlation included within the highest posterior interval at 95%. Within-litter birth weight variation and LS showed a negative and large permanent environmental correlation ( − 0.872), and LW and LS were negatively correlated due to residual ( − 0.762) and permanent environmental ( − 0.449) random sources of variation. Within-litter birth weight variation influenced lamb mortality during the first 7 days of life (P < 0.05), increasing and decreasing survivability in heavier and lighter littermates, respectively. Nevertheless, stillbirths and lambs died after the 1st week of life were not affected by BWV (P>0.05). The low heritability found indicates that slow genetic progress may be expected from selecting for BWV. Close to zero genetic correlations suggest that this selection will probably not affect LS and LW, although some significant permanent and residual correlations must be taken into account. Further studies are needed to understand better the genetic architecture among these three reproductive traits.     

Litter decomposition and nitrogen and phosphorus dynamics in peatlands and uplands over 12 years in central Canada

The large accumulation of organic matter in peatlands has been partially attributed to litter decomposition rates, which are slowed by a high water table. To test this, we examined whether there were significant differences in the decomposition and N and P dynamics of ten foliar litters and wood blocks at three pairs of upland forest and peatland sites in the transitional grassland, high boreal and low subarctic regions of central Canada, using litterbags collected over a 12-year period. At two of the three pairs, the decomposition rate, as determined by proportion of the original mass remaining after 12 years and by the exponential decay coefficient (k), was faster overall at the upland than at the peatland. In the third pair, there was no significant difference, despite the water table being close to the peat surface; warmer soil temperatures in the peatland than the upland may be the cause. In general, there were small losses or gains of N in the litters after 12 years, compared to the original litter, though there were some differences among litter types and sites, net gains in N likely reflecting the higher exogenous N availability. P was lost from most litters at the two northern pairs of sites, but at the transitional grassland pair, there were large net gains in P and greater variation among litters. The N:P ratio in the original litters ranged from 5 to 26 and after 12 years the ratio narrowed, with the site average of the ten litters ranging from 13 to 22, varying with the soil ratio. Decomposition rates and N and P dynamics after 12 years are different between upland and peatland sites: although the water table is a primary control on these differences, other factors such as temperature and soil nutrient status are also important.     

Multiple paternity in wild house mice (Mus musculus musculus): effects on offspring genetic diversity and body mass

Multiple mating is common in many species, but it is unclear whether multiple paternity enhances offspring genetic diversity or fitness. We conducted a survey on wild house mice (Mus musculus musculus), and we found that in 73 pregnant females, 29% of litters had multiple sires, which is remarkably similar to the 23–26% found in feral populations of Mus musculus domesticus in the USA and Australia, respectively. The question is: How has selection maintained multiple mating in these subspecies since the evolutionary divergence, ca. 2800–6000 years ago? We found no evidence that multiple paternity enhanced females’ litter size, contrary to the fertility assurance or genetic benefits hypotheses. Multiple paternity was associated with reduced mean and variance in offspring body mass, which suggests that females allocate fewer resources or that there is increased intrauterine conflict in multiple-versus single-sired litters. We found increased allelic diversity (though not heterozygosity) in multiple-sired litters, as predicted by the genetic diversity hypothesis. Finally, we found that the dams’ heterozygosity was correlated with the mean heterozygosity of their offspring in single-and multiple-sired litters, suggesting that outbred, heterozygous females were more likely to avoid inbreeding than inbred, homozygous females. Future studies are needed to examine how increased genetic diversity of litters and smaller mean (and variance) offspring body mass associated with multiple paternity affect offspring fitness.     

Frequency distributions for chloroplast genes in Chlamydomonas zygote clones: Evidence for random drift

Mitochondria and chloroplasts of eucaryotic cells contain populations of DNA molecules. In certain cases, e.g., the chloroplasts of Chlamydomonas reinhardtii and the mitochondria of Saccharomyces cerevisiae, organelles contributed by the two parents are known to fuse in the zygote, creating a single population of DNA molecules. In a cross, this population will include molecules of both parental genotypes. There is reason to suspect that organelle DNA molecules in this population are selected randomly for replication and recombination. This would result in random changes in the frequency of a particular allele or genotype within the organelle gene pool of a single zygote and also within its clone of progeny cells. A given gene frequency would increase in some zygote clones and decrease in others, analogous to random drift of gene frequencies in small Mendelian populations. To test this, we have examined the distribution of chloroplast gene frequencies among the zygote clones produced in each of a number of crosses of Chlamydomonas. These distributions are typically U or L shaped as predicted by the random drift hypothesis. They include uniparental zygote clones, in which a chloroplast allele from one parent has been fixed (frequency 100%) and the alternative allele from the other parent has been lost (frequency 0%). Among the remaining (biparental) zygote clones, there is a linear distribution of allele frequencies, showing a great increase in variance over the input frequencies. In these experiments both biparental and uniparental zygotes show a bias favoring chloroplast alleles from the mt⁺ (maternal) parent, and there is no statistically significant mode at the allele frequency of 0.5 corresponding to the equal input of alleles from the maternal and paternal (mt^?) parents. The observed distributions support the hypothesis that both uniparental inheritance and the high variance of allele frequencies among zygote clones are due to random drift of allele frequencies, coupled with a directional force which favors fixation of the maternal allele. In addition, statistical analysis of the data shows a strong but incomplete tendency for linked chloroplast markers to be fixed or lost together in uniparental zygotes. Possible cellular and molecular mechanisms for these observations are discussed.     

Seasonal variation in fertility, fecundity and litter sex ratio in laboratory and wild stocks of house mice (Mus domesticus)

Data compilations were made for three parameters pertaining to reproduction in a domestic strain (14 years) and a wild stock (4 years) of commensal house mice: (a) the percentage of females mated that produced litters; (b) the average number of pups per litter; and (c) the sex ratio of the pups in the litters. Fecundity and fertility varied seasonally in both domestic and wild mice. More females become pregnant and litter sizes were larger in the spring, summer, and fall months than during the winter season. Sex ratios also varied seasonally with more male biased litters produced during the 4 winter months. There appear to be seasonal shifts in productivity for both stocks of mice and these seasonal trends have not been altered by domestication under laboratory conditions. In spite of the fact that house mice are generally opportunistic, it is possible that there has been selection in the mice for shifting rates of production in relation to the best seasons of the year in terms of climate and resource availability. The higher production of males during summer months may be geared toward greater success in dispersal at that season and a higher probability that the males can find a territory and mate successfully in summer rather than winter. These results have potential implications for animal breeders and for those who maintain mouse colonies to produce animals for scientific investigations.     

Sow and piglet factors determining variation of colostrum intake between and within litters

Colostrum intake has a short- and long-term beneficial impact on piglet performance and mortality. Sows’ colostrum production and piglets’ colostrum intake are limited and highly variable. The present study investigated sow and piglet factors explaining the variation of colostrum intake between and within litters. The CV for colostrum intake and birth weight (BW_b) of all piglets within a litter was calculated to evaluate the variation of colostrum intake and BW_b within a litter (colostrum and litter BW_b heterogeneity, respectively). A total of 1937 live-born piglets from 135 litters from 10 commercial herds were included. Colostrum intake per piglet averaged 371±144 g and was affected by breed (P=0.02). It was lower when oxytocin was administered to the sow during parturition (P=0.001) and with increased litter size (P<0.001). It was higher when the interval between birth and first suckling decreased (t_FS, P<0.001). Colostrum intake was positively influenced by BW_b (P<0.001) and this association was more pronounced in piglets from Topigs (P=0.03) and Hypor (P=0.03) sows compared with piglets from Danbred sow breeds. The positive relationship between colostrum intake and BW_b was more pronounced when t_FS lasted longer (P=0.009). Heterogeneity in colostrum intake averaged 31±11%, it increased when oxytocin was applied during farrowing (P=0.004) and when stillbirth occurred (P=0.006). Colostrum heterogeneity was positively associated with litter size (P<0.001) and litter BW_b heterogeneity (P=0.01). The positive relationship between colostrum and litter BW_b heterogeneity was more pronounced when oxytocin was applied during farrowing (P=0.04). The present study demonstrated that oxytocin should be used cautiously in sows during farrowing. Farrowing and colostrum management should prevent or counteract the adverse influences of stillbirth, large and heterogeneous litters on colostrum intake and colostrum heterogeneity. The study also confirmed the expected association between BW_b and colostrum intake and indicated that the impact of BW_b on colostrum intake was different among breeds (Hypor v. Danbred) and dependent on piglets’ latency to first suckling. Hence, colostrum management should focus on low birth weight piglets, especially in some breeds, and low colostrum intake in low birth weight piglets can be counteracted by shortening the t_FS.     

Evolution of color variation in dragon lizards: quantitative tests of the role of crypsis and local adaptation

Abstract Many animal species display striking color differences with respect to geographic location, sex, and body region. Traditional adaptive explanations for such complex patterns invoke an interaction between selection for conspicuous signals and natural selection for crypsis. Although there is now a substantial body of evidence supporting the role of sexual selection for signaling functions, quantitative studies of crypsis remain comparatively rare. Here, we combine objective measures of coloration with information on predator visual sensitivities to study the role of crypsis in the evolution of color variation in an Australian lizard species complex (Ctenophorus decresii). We apply a model that allows us to quantify crypsis in terms of the visual contrast of the lizards against their natural backgrounds, as perceived by potential avian predators. We then use these quantitative estimates of crypsis to answer the following questions. Are there significant differences in crypsis conspicuousness among populations? Are there significant differences in crypsis conspicuousness between the sexes? Are body regions “exposed” to visual predators more cryptic than “hidden” body regions? Is there evidence for local adaptation with respect to crypsis against different substrates? In general, our results confirmed that there are real differences in crypsis conspicuousness both between populations and between sexes; that exposed body regions were significantly more cryptic than hidden ones, particularly in females; and that females, but not males, are more cryptic against their own local background than against the background of other populations. Body regions that varied most in contrast between the sexes and between populations were also most conspicuous and are emphasized by males during social and sexual signaling. However, results varied with respect to the aspect of coloration studied. Results based on chromatic contrast (“hue’ of color) provided better support for the crypsis hypothesis than did results based on achromatic contrast (“brightness’ of color). Taken together, these results support the view that crypsis plays a substantial role in the evolution of color variation and that color patterns represent a balance between the need for conspicuousness for signaling and the need for crypsis to avoid predation.     

A naturally occurring variant of Hsp90 that is associated with decanalization

The heat shock protein Hsp90 has been the focus of many studies since it was suggested that it acts to mediate the buffering of phenotypic variation. Hsp90-mediated buffering may result in the accumulation of cryptic genetic variation that, when released either as a consequence of environmental or genetic stress, increases the evolvability of a population. Recent studies using laboratory-induced mutations of Hsp90 and/or chemical inhibition to disrupt Hsp90 function confirm that Hsp90 can buffer cryptic genetic variation. We have previously identified a naturally occurring variant in the charged linker region of the Hsp90 gene, and now examine whether this variant is associated with altered levels of trait variability. The variant is associated with the release of cryptic genetic variation for canalized morphological (bristle) traits, but not for uncanalized morphological (wing and bristle) traits, and the effect on canalized traits depends on culture temperature. This suggests that natural genetic variation in Hsp90 may mediate the evolution of canalized morphological traits even if it does not influence the expression of variation for uncanalized traits.     

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.     

Phenology and synchrony of roe deer breeding in northern Italy

The timing and synchrony of births of roe deerCapreolus capreolus Linnaeus, 1758 population were examined for 7 consecutive years in northern Italy. From 1997 to 2003, 136 fawns, 65 females and 71 males, from 117 litters were captured. The average date of fawning (30th of May) was constant among years and between sexes. The onset of parturition was 19th of May while the total length of the parturition season was 25 days (from 19th of May to 13th of June). Births were highly synchronous, with 80% of births occurring in 17 days. Synchrony of birth did not exhibit any significant interannual or intersexual variation. The stability of the birth pattern was confirmed by the parturition date of 13 radiocollared females who all gave birth within 4 days in consecutive fawning seasons. In agreement with previous findings, our data support the hypothesis that the roe deer is constrained at both ends of the fawning season by the short time between the fawning season and rut and by the need to match optimal climate conditions and forage quality to face late gestation and lactation. All these factors result in a relatively stable and synchronous birth season.     

Letter: Notes on "heat loss from a Newtonian animal"

Parker, Baker &; Smith (1972) have demonstrated mathematically that given the evolution of sexual reproduction, disruptive selection for the production of either many small gametes or a few large gametes may occur, resulting in a stable polymorphism of “sperm” and “egg” producers. Their model for the evolution of anisogamy requires only that zygote fitness (F) increase steeply with increases in zygote volume (V) (for Foc∝ V^x, x must be greater than 1·5) and that a sufficiently broad range of zygote productivity-size variants exist in the population (the higher the value of x, the broader the range needed). They suggest that anisogamy is almost universal in multicellular organisms but relatively rare in unicellular organisms because only for the former is an investment in extra gametic reserves at the expense of the number of gametes produced likely to be worthwhile in terms of increasing the survival probability of the zygote. In this note a graphical analysis and evidence from the anisogamous Protista will be presented concerning this hypothesis.     

An overview of natural variation studies in the Arabidopsis thaliana circadian clock

Circadian clocks are ubiquitous mechanisms that provide an adaptive advantage by predicting subsequent environmental changes. In the model plant Arabidopsis thaliana (Arabidopsis), our understanding of the complex genetic network among clock components has considerably increased during these past years. Modeling has predicted the possibility of additional component to systematically and functionally complete the clock system. Mutagenesis screens have in the past been successfully employed to detect such novel components. With the advancement in sequencing technologies and improvements in statistical approaches, the extensive natural variation present in Arabidopsis accessions has emerged as a powerful alternative in functional gene discovery. In this review article, we review the previous efforts in mapping natural alleles affecting various clock parameters and will discuss further potentials of such natural-variation studies in physiological and ecological contexts.     

Metric variation and species recognition in the fossil record

One of the first and most important tasks of the paleontologist is classifying specimens into species. Species recognition commonly involves sorting specimens on the basis of qualitative and quantitative similarities and differences. Often, however, variation in simple metric characters like tooth size or jaw length plays an important role in debates about whether a sample comprises a single species or more than one morphologically similar species. For example, Simpson, Roe, and Lewontin 1 suggested that a fossil sample showing a coefficient of variation greater than 10.0 was likely to comprise more than one species. Well‐known controversies over species recognition in which metric sample variation has been important have simmered for years, focusing on hominids, hominoids, and other extinct primates. Some of these have been resolved; others have not. For example, Pilbeam and Zwell 2 convincingly demonstrated multiple species among South African hominids by showing that metric tooth size variation was too great to be reasonably interpreted as sexual dimorphism. But metric variation continues to play a role in debates about whether Australopithecus afarensis 3 , 4 and Homo habilis 5 - 9 each comprise a single species or two or more separate species. Similarly, there has been steady debate about the number of species present in African Proconsul. Some favor an interpretation of a single extremely dimorphic species, 10 - 12 while others favor an interpretation of two or more species. 13