Population genetic structure of parthenogenetic flatworm populations with occasional sex

1. Sexual populations are expected to perform better in fluctuating environments than asexuals because recombination provides the potential to adapt to changing environments due to increased genetic variation. Nevertheless, some asexual species show comparably high levels of genotypic diversity. Such diversity might be achieved through gene flow between coexisting sexual and asexual populations or through sexual events within asexual populations. 2. Evidence for occasional sex in the flatworm Schmidtea polychroa was previously found at one specific site that is inhabited by parthenogenetic forms. There, varying rates of sex between subpopulations, reaching up to 12%, were observed. Past recurrent sexual processes left a significant genetic signature in the population genetic structure of this population. In the present study, we examined the population genetic structure of six independent metapopulations (lakes) of the freshwater planarian flatworm S. polychroa, to confirm the presence of occasional sex and that its population genetic consequences can be generalised. 3. Using microsatellites, we found varying rates of occasional sex among subpopulations. Metapopulations showed medium to high levels of genotypic diversity that correlated with the rate of sex. 4. We conclude that occasional sex has considerable consequences for population genetic structure of parthenogenetic species and promotes diversity that might allow response to the particular type of selection that is usually predicted to favour sexual reproduction. This reproductive strategy provides genetic characteristics required for selection to act on, and might, therefore, explain the success of this parthenogenetic species.     

Reproductive strategy and population variability in the facultative apomict Hieracium pilosella (Asteraceae)

Molecular studies of apomictic plant species often detect more genotypic variation than predicted from their assumed reproductive mode. The two most commonly invoked mechanisms to explain these high levels of variation are recombination, via facultative sexuality, and mutation. The potential for sexual reproduction in the facultative apomict Hieracium pilosella (Asteraceae) was determined at three field sites by artificially pollinating with the closely related, but morphologically distinct, H. aurantiacum. The level of genotypic variation at the three sites was recorded using inter-simple sequence repeats (ISSRs). There was a significant, positive relationship between the measured potential for sexual reproduction and population genotypic variability, indicating that sex has played a role in the structuring of these populations; however, a causal relationship cannot be stated because of the use of regression. We also applied the recently developed method of compatibility analysis. Compatibility analysis can determine, using the occurrence of "character incompatibilities," whether patterns of variation observed in populations are most parsimoniously explained by mutation or recombination. Compatibility analysis also indicated that sexual reproduction had played a role in generating genotypic diversity in these populations. Combining these different types of data may give a greater understanding of the potential for the generation of genotypic diversity in facultative apomictic populations.     

The effects of sex and mutation rate on adaptation in test tubes and to mouse hosts by Saccharomyces cerevisiae

Some hypotheses for the evolution of sex focus on adaptation to changing or heterogeneous environments, but these hypotheses have rarely been tested. We tested for advantages of sex and of increased mutation rates in yeast strains in two contrasting environments: a standard and relatively homogeneous laboratory environment of minimal medium in test tubes, and the variable environment of a mouse brain experienced by pathogenic strains. Evolving populations were founded as equal mixtures of sexual and obligately asexual genotypes. In the sexuals, cycles of sporulation, meiosis, and mating were induced approximately every 50 mitotic generations, with the asexuals undergoing sporulation but not ploidy cycles or recombination. In both environments, replicate negative control populations established with the same pair of strains were propagated with neither mating nor meiosis. In test tubes with no sex induced, sexuals were fixed in all five replicates within 250 mitotic generations, whereas in mice with no sex induced, asexuals were fixed in all four replicates by 170 generations. Inducing sex altered these outcomes in opposite directions in test tubes and mice, decreasing the fixation frequencies of sexuals in test tubes but increasing them in mice. These contrasts with asexual controls suggest an advantage for sex in mice but not in test tubes, although there was no difference between test tubes and mice in the numbers of populations fixed-for sexuals. In analogous experiments testing for an advantage of increased mutation rates, wild-type genotypes became fixed at the expense of mutators in every replicate of both test tube and mouse populations, indicating a disadvantage for mutators in both environments. Increased rates of point mutation do not appear to accelerate adaptation.     

Reduced recombination in gynodioecious populations of a facultative apomictic orchid

Many plants combine sexual reproduction with some form of asexual reproduction to different degrees, and lower genetic diversity is expected with asexuality. Moreover, the ratios of sexual morphs in species with gender dimorphism are expected to vary in proportion to the reproductive success of the sexual process. Hence, sex ratios can directly influence the genetic structure and diversity of a population. We investigated genotypic diversity in 23 populations of a facultative, apomictic gynodioecious orchid, Satyrium ciliatum, to examine the effect on genotypic diversity of variation in the frequency of females and in the amount of sexual reproduction. The study involved one pure female, seven gynodioecious (both females and hermaphrodites present) and 15 hermaphroditic populations. Pollinia receipt was higher in hermaphroditic than in gynodioecious populations. Analyses of variation in ISSRs demonstrated that genotypic diversity was high in all populations and was not significantly different between hermaphroditic and gynodioecious populations. We used character compatibility analysis to determine the extent to which recombination by sexual reproduction contributed to genotypic diversity. The results indicate that the contribution of recombination to genotypic diversity is higher in hermaphroditic than in gynodioecious populations, consistent with the finding that hermaphroditic populations received higher amounts of pollinia. Our finding of reduced recombination in gynodioecious populations suggests that maintenance of sex in hermaphrodites plays an important role in generating genotypic diversity in this apomictic orchid.     

Life cycle strategies and genotypic variability in populations of aphids

Populations of the rose aphid (Macrosiphum rosae) from various latitudes show differences in their life cycles. In warm climates they are exclusively parthenogenetic, whereas in cold climates sexual reproduction allows the aphid to overwinter in the egg stage. In temperate zones both holocyclic and anholocyclic clones occur within the same population. It is assumed that the reproductive mode has consequences for the genotypic structure of the populations. Continuous parthenogenesis results in low genogypic variability caused by selection and/or random genetic drift. The greater genotypic variability of holocyclic populations is due to recombination during the sexual phase. This yiew is supported by the study of populations which revealed high genotypic variability and homogeneity between populations in areas with harsh winters (e.g. Norway). In temperate countries (e.g. England) the genotypic variability was lower but the uniformity between populations still high, and in warm climates (e.g. the Canary Isles) the variability was lowest and heterogeneity between populations largest.     

The ecology and genetics of fitness in Chlamydomonas. VIII. The dynamics of adaptation to novel environments after a single episode of sex

According to classical evolutionary theory, sexual recombination can generate the variation necessary to adapt to changing environments and thereby confer an evolutionary advantage of sexual over asexual reproduction. Using the green alga, Chlamydomonas reinhardtii, we investigated the effect of a single sexual episode on adaptation of heterotrophic growth on different carbon sources. In an initial mixture of isolates, sex was induced and the resulting offspring constituted the sexual populations, along with any unmated vegetative cells; the unmated mixture of isolates represented the asexual populations. Mean and variance in division rates (i.e., fitness) were measured four times during approximately 50 generations of vegetative growth in the dark on all possible combinations of four carbon sources. Consistent with effects of recombination of epistatic genes in linkage disequilibrium, sexual populations initially had a higher variance in fitness, but their mean fitness was lower than that of asexual populations, possibly due to recombinational load. Subsequently, fitness of sexual populations exceeded that of asexual ones, but finally they regained parity in both mean and variance of fitness. Although recombination was not more effective on more complex substrates, these results generally support the idea that sex can accelerate adaptation to novel environments.     

Reproduction and life history strategies of the common jellyfish, Aurelia aurita, in relation to its ambient environment

The scyphozoan Aurelia aurita (Linnaeus) is a cosmopolitan species, having been reported from a variety of coastal and shelf sea environments around the world. It has been extensively studied over the last 100 years or so, and examination of the literature reveals three striking features: (1) the presence of populations in a wide range of environmental conditions; (2) large inter-population differences in abundance and life history patterns over large and small spatial scales; and (3) inter-annual variability in various aspects of its population dynamics. A. aurita is clearly a highly flexible species that can adapt to a wide range of environmental conditions. While various physiological and behavioural characteristics explain how A. aurita populations can take advantage of their surrounding environment, they do not explain what governs the observed temporal and spatial patterns of abundance, and the longevity or lifespan of populations. Understanding these features is necessary to predict how bloom populations might form. In a given habitat, the distribution and abundance of benthic marine invertebrates have been found to be maintained by four factors: larval recruitment (sexual reproduction), migration, mortality and asexual reproduction. The aims of this review are to determine the role of reproduction and life history strategies of the benthic and pelagic phases of A. aurita in governing populations of medusae, with special attention given to the dynamic interaction between A. aurita and its surrounding physical and biological environment.     

Clonality and recombination in the arctic plant Saxifraga cernua

The circumarctic clonal plant Saxifraga cernua reproduces efficiently via bulbils, largely depends on insects for pollination and appears to set seed very rarely. However, high levels of genotypic variation observed at small spatial scales in the arctic archipelago of Svalbard have been taken as evidence of occasional sexual reproduction. Here we assess the relative contributions of mutation and recombination to random amplified polymorphic DNA variation in four populations in East Greenland and re-analyse the Svalbard data. Greater variation due to recombination was predicted in Greenland than in Svalbard, because the higher summer temperatures and longer growing season likely increase the chances for sexual reproduction. Although we observed higher levels of genotypic diversity in Greenland than in Svalbard, matrix incompatibility and linkage disequilibrium measures provided no evidence of more sexuality, suggesting differences in glacial/postglacial history. The genetic structure and spatial distribution of clones suggest that clonal migration may increase variability in local populations, which is consistent with frequent large-scale migration in this species inferred from a circumarctic analysis of chloroplast DNA haplotypes. We conclude that a combination of somatic mutations and sexual reproduction has contributed to the observed patterns of genotypic diversity in the Greenland and Svalbard populations of S. cernua , and that sexual reproduction is important in maintaining genotypic diversity, despite the rarity of observations of seed setting.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 152 , 209–217.     

The population genetics of clonal and partially clonal diploids

The consequences of variable rates of clonal reproduction on the population genetics of neutral markers are explored in diploid organisms within a subdivided population (island model). We use both analytical and stochastic simulation approaches. High rates of clonal reproduction will positively affect heterozygosity. As a consequence, nearly twice as many alleles per locus can be maintained and population differentiation estimated as F(ST) value is strongly decreased in purely clonal populations as compared to purely sexual ones. With increasing clonal reproduction, effective population size first slowly increases and then points toward extreme values when the reproductive system tends toward strict clonality. This reflects the fact that polymorphism is protected within individuals due to fixed heterozygosity. Contrarily, genotypic diversity smoothly decreases with increasing rates of clonal reproduction. Asexual populations thus maintain higher genetic diversity at each single locus but a lower number of different genotypes. Mixed clonal/sexual reproduction is nearly indistinguishable from strict sexual reproduction as long as the proportion of clonal reproduction is not strongly predominant for all quantities investigated, except for genotypic diversities (both at individual loci and over multiple loci).     

The effect of newly induced mutations on the fitness of genotypes and populations of yeast (Saccharomyces cerevisiae)

This paper analyses the fate of artificially induced mutations and their importance to the fitness of populations of the yeast, Saccharomyces cerevisiae, an increasingly important model organism in population genetics.

Diploid strains, treated with UV and EMS, were cultured asexually for approximately 540 generations and under conditions where the asexual growth was interrupted by a sexual phase.

Growth rates of 100 randomly sampled diploid clones were estimated at the beginning and at the end of the experiment. After the induction of sporulation the growth rates of 100 randomly sampled spores were measured. UV and EMS treatment decreases the average growth rate of the clones significantly but increases the variability in comparison to the untreated control. After selection over approximately 540 generations, variability in growth rates was reduced to that of the untreated control. No increase in mean population fitness was observed. However, the results show that after selection there still exists a large amount of hidden genetic variability in the populations which is revealed when the clones are cultivated in environments other than those in which selection took place. A sexual phase increased the reduction of the induced variability.     

Population structure,seasonal genotypic differentiation,and clonal diversity of weedy dandelions in three Boston area populations (Taraxacum sp.)

Weedy dandelions have a worldwide distribution and thrive in urban environments despite a lack of sexual reproduction throughout most of its range. North American dandelions, introduced from Eurasia, are believed to be primarily, if not exclusively, apomictic triploids. In some European populations, apomicts co‐occur with diploid sexual individuals and hybridizations can create genetically unique apomicts, which may subsequently disperse and establish new populations globally. Using six nuclear microsatellite markers and a cpDNA intergenic spacer, we investigate the impact of this unusual natural history on population structure and diversity in three urban Boston area dandelion populations. Our results show high levels of genetic diversity within populations, spatial population structure, and seasonal genotypic differentiation in flowering times. We find evidence that sexual reproduction and recombination, presumably in Europe, and extensive gene flow drive these patterns of diversity and create the appearance of panmixia despite the lack of evidence for local sexual reproduction.     

Patterns, sources and ecological implications of clonal diversity in apomictic Ranunculus carpaticola (Ranunculus auricomus complex, Ranunculaceae)

Sources and implications of genetic diversity in agamic complexes are still under debate. Population studies (amplified fragment length polymorphisms, microsatellites) and karyological methods (Feulgen DNA image densitometry and flow cytometry) were employed for characterization of genetic diversity and ploidy levels of 10 populations of Ranunculus carpaticola in central Slovakia. Whereas two diploid populations showed high levels of genetic diversity, as expected for sexual reproduction, eight populations are hexaploid and harbour lower degrees of genotypic variation, but maintain high levels of heterozygosity at many loci, as is typical for apomicts. Polyploid populations consist either of a single AFLP genotype or of one dominant and a few deviating genotypes. genotype/genodive and character incompatibility analyses suggest that genotypic variation within apomictic populations is caused by mutations, but in one population probably also by recombination. This local facultative sexuality may have a great impact on regional genotypic diversity. Two microsatellite loci discriminated genotypes separated by the accumulation of few mutations ('clone mates') within each AFLP clone. Genetic diversity is partitioned mainly among apomictic populations and is not geographically structured, which may be due to facultative sexuality and/or multiple colonizations of sites by different clones. Habitat differentiation and a tendency to inhabit artificial meadows is more pronounced in apomictic than in sexual populations. We hypothesize that maintenance of genetic diversity and superior colonizing abilities of apomicts in temporally and spatially heterogeneous environments are important for their distributional success.     

Elevated temperature changes female costs and benefits of reproduction

Despite obvious benefits, reproduction also imposes severe costs on females. Such costs and benefits are highly sensitive to environmental factors. Rapidly changing conditions may thus disturb a finely poised balance between the two and pose a challenge to natural populations. A more complete understanding of reproduction and population fitness across different environments is, hence, crucial. In particular, sexual selection could either be beneficial or detrimental when conditions change abruptly. Here Tribolium castaneum females were subjected to mating treatments with or without sexual selection (virginity, monogamy, polyandry) replicated at standard versus elevated temperatures. We found a substantial survival cost of reproduction at the standard, but not at the elevated temperature. Reproductive success was similar across mating treatments at the standard temperature, but at elevated temperature we detected a significant benefit of polyandry compared to monogamy. These findings indicate that environmental heterogeneity can strongly influence the balance between costs and benefits when sexual selection is allowed to act. Furthermore, reproduction may be critically affected by changes in temperature with potentially profound consequences for population fitness.     

Evolutionary traction: the cost of adaptation and the evolution of sex

The advantage of sexual reproduction remains a puzzle for evolutionary biologists. Everything else being equal, asexual populations are expected to have twice the number of offspring produced by similar sexual populations. Yet, asexual species are uncommon among higher eukaryotes. In models assuming small populations, high mutation rates, or frequent environmental changes, sexual reproduction seems to have at least a two-fold advantage over asexuality. But the advantage of sex for large populations, low mutation rates, and rare or mild environmental changes remains a conundrum. Here we show that without recombination, rare advantageous mutations can result in increased accumulation of deleterious mutations ('evolutionary traction'), which explains the long-term advantage of sex under a wide parameter range.     

Influence of parthenogenetic reproduction on the genotypic constitution and evolutionary success of populations and species

The life cycle of Daphnia commonly includes parthenogenesis, sexual reproduction, and diapause. Although there are no genotypic transformations during diapause, it separates the clonal and pseudopanmictic states of the population. During parthenogenetic reproduction primarily polymorphic natural populations gradually degenerate into a mixture of a few clones. As resting eggs are usually produced sexually (= by means of sexual reproduction), after the diapause the vast diversity of individual genotypes normal for panmictic populations is observed. It is the competitive interactions between parthenogenetic clones which again eventually decrease the genotypic polymorphism. Forms in which the sexual process and diapause are rare (or species which are represented by parthenogenetic populations in one part of their area and by bisexual ones in another) demonstrate the most significant differences between clonal and panmictic populations.Parthenogenesis, apomixis and other kinds of reproduction without genetic recombination are widely spread in branchiopod crustaceans (Notostraca, Anostraca, Conchostraca, Cladocera) and in many other animals and plants. Moreover, reproduction without recombination plays an important role in evolution of faunas and floras. It is emphasized that non-recombinating races and species having high heterozygosity but a low level of genetic variation, enjoy short term advantages, but die out after change in the environment.     

Sexual reproduction and the evolution of microbial pathogens

Three common systemic human fungal pathogens--Cryptococcus neoformans, Candida albicans and Aspergillus fumigatus--have retained all the machinery to engage in sexual reproduction, and yet their populations are often clonal with limited evidence for recombination. Striking parallels have emerged with four protozoan parasites that infect humans: Toxoplasma gondii, Trypanosoma brucei, Trypanosoma cruzi and Plasmodium falciparum. Limiting sexual reproduction appears to be a common virulence strategy, enabling generation of clonal populations well adapted to host and environmental niches, yet retaining the ability to engage in sexual or parasexual reproduction and respond to selective pressure. Continued investigation of the sexual nature of microbial pathogens should facilitate both laboratory investigation and an understanding of the complex interplay between pathogens, hosts, vectors, and their environments.     

ALLOZYMIC VARIATION IN LOCAL APOMICTIC POPULATIONS OF LEMNA MINOR (LEMNACEAE)

Flowering occurrence and allozymic variation were studied in eight local populations of Lemna minor in eastern Ontario, Canada. After 2 years of survey, not a single flower was observed. This absence of flowering suggests the possibility of loss of sexual reproduction. This may have had no net adverse effect on fitness given the simple life history and prolific vegetative propagation in L. minor. However, the absence of sexual reproduction may limit genotypic diversity. The allozymic analysis detected 18 loci from 13 enzyme systems. Large deviations from Hardy-Weinberg equilibrium were common because of an excess of homozygotes for several enzyme systems. The genotypic diversity within these eight populations had a mean D value of 0.973 with an average number of genotypes per population of 19.6. These results suggest that genotypic diversity within these populations is not severely limited by the rarity of sexual reproduction. The mean genotypic distance index (D₁₄ = 0.801) suggests a high degree of differentiation between populations. The mean number of populations per genotype was 1.78. Using a Mantel test, the genotypic distance matrix was not significantly related to the population-to-population distance matrix (t = -0.161, P = 0.413). Although rare events of sexual reproduction may help maintain genetic variation, somatic mutations and multiple origins of clones may be important factors maintaining genetic diversity both within and between populations of L. minor.     

Effect of reproductive modes and environmental heterogeneity in the population dynamics of a geographically widespread clonal desert cactus

The dynamics of plant populations in arid environments are largely affected by the unpredictable environmental conditions and are fine-tuned by biotic factors, such as modes of recruitment. A single species must cope with both spatial and temporal heterogeneity that trigger pulses of sexual and clonal establishment throughout its distributional range. We studied two populations of the clonal, purple prickly pear cactus, Opuntia macrocentra, in order to contrast the factors responsible for the population dynamics of a common, widely distributed species. The study sites were located in protected areas that correspond to extreme latitudinal locations for this species within the Chihuahuan Desert. We studied both populations for four consecutive years and determined the demographic consequences of environmental variability and the mode of reproduction using matrix population models, life table response experiments (LTREs), and loop and perturbation analyses. Although both populations seemed fairly stable (population growth rate, λ∼1), different demographic parameters and different life cycle routes were responsible for this stability in each population. In the southernmost population (MBR) LTRE and loop and elasticity analyses showed that stasis is the demographic process with the highest contributions to λ, followed by sexual reproduction, and clonal propagation contributed the least. The northern population (CR) had both higher elasticities and larger contributions of stasis, followed by clonal propagation and sexual recruitment. Loop analysis also showed that individuals in CR have more paths to complete a life cycle than those in MBR. As a consequence, each population differed in life history traits (e.g., size class structure, size at sexual maturity, and reproductive value). Numerical perturbation analyses showed a small effect of the seed bank on the λ of both populations, while the transition from seeds to seedlings had an important effect mainly in the northern population. Clonal propagation (higher survival and higher contributions to vital rates) seems to be more important for maintaining populations over long time periods than sexual reproduction.     

Variation and covariation among life-history traits in Rumex acetosella from a successional old-field gradient

In this study morphological variation and the potential for competition to affect biomass and seedling selection of the families of five populations of Rumex acetosella L. sampled along a successional old-field gradient have been investigated. Seeds from 25 families were submitted to four competitive regimes: no competition (one plant per pot), medium competition (two plants/ pot taking plants from the same population), high within-population competition (four individuals from the same population in a pot) and high between-population competition (four individuals from two different populations in a pot). Eight traits were analysed after 3 months of growth for variation among families within populations. A significant difference among families within the two older populations was recorded for sexual biomass and related components. High sensitivity of these traits to density was observed in all populations except the youngest, suggesting specialization to particular environmental conditions in late successional populations, and a good adaptive capacity to buffer environmental variation in the pioneer population. Little significant interaction between competitive regimes and families within populations was found, i.e. genotypes within each population showed little variation in their response to environmental variation. Genotypic variance decreased with increasing competitive conditions for the majority of the traits. However, the percentage of variance in sexual reproduction explained by family was stable among treatments. Tradeoffs between vegetative reproduction and sexual reproduction were recorded at the population level along the successional gradient, with increasing competitive conditions. As succession proceeds, we observed a decrease in sexual reproduction and an increase in vegetative reproduction. At the family level, correlation among traits were similar when plants were grown in the absence of competition and at high density, with a significant negative correlation between sexual reproduction and vegetative reproduction. For both sprout number and sexual biomass, the performance of families grown under all the treatments was positively correlated. Together these results indicate allocational constraints on the reproductive biology of R. acetosella that may be favoured by natural selection and have influenced population differentiation along the successional gradient. However, they also revealed that the potential exists for evolutionary specialization through plasticity, in response to variation in environmental conditions.     

Clonal diversity and turnover in an overwintering Daphnia pulex population,and the effect of fish predation

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