Despite the two-fold cost of sex, most of the higher animals reproduce sexually. The advantage of sex has been suggested to be its ability, through recombination, to generate greater genetic diversity than asexuality, thus enhancing adaptation in a changing environment. We studied the genetic diversity and the population structure of three closely related species of bag
Genetic diversity sexual reproduction moths: These species compete for the same resources
Genetic diversity sexual reproduction share the same parasitoids.
Allelic richness was comparable between the sexual species but it was higher than in the asexual species. All species showed high heterozygote deficiency and a large variation was observed among F IS values across loci and populations. Large Genetic diversity sexual reproduction differentiation was observed between populations confirming the poor dispersal ability of these species. The asexual species showed lower genotype diversity than the sexual species.
Nevertheless, genotype diversity was high in all asexual populations. The three different species show a similar population structure characterised by high genetic differentiation among populations and low dispersal. Although the parthenogenetic D. While we can not totally rule out the presence of cryptic sex, would explain this high genotype diversity, we never observed sex in the parthenogenetic D.
Alternatively, a non-clonal parthenogenetic reproduction, such as automictic thelytoky, could explain the high genotypic diversity observed in D. Parthenogenetic females have a two-fold advantage over sexual females because they produce Genetic diversity sexual reproduction the fecund sex while sexual females produce also males.
The elimination of sex is predicted whenever the two strategies compete unless there are factors that overcome this disadvantage. Nevertheless, most of the higher animals reproduce sexually[ 1 ]. This leads to a fundamental question which continues to puzzle evolutionary biologists: A large body of theories seek to explain the maintenance of sex [ 2 - 7 ].
Advantages of sexual reproduction arise from genetic recombination in cross-fertilisation, which purges deleterious mutation and increases genetic variability in the population [ 8 - 10 ], enhancing adaptation in a changing environment. Genetic diversity sexual reproduction idea that sexual reproduction and recombination may be favoured in changing environments has been the subject of several papers [ 11 - 15 ]. If a trait is subjected to stabilising selection, genetic variability introduces a genetic load as a consequence of the produced phenotypes that deviate from the optimum [ 16 ].
However, in a varying environment that exerts directional selection on a trait, genetic variability is essential because the response to selection will Genetic diversity sexual reproduction proportional to the additive genetic variance in the population [ 15 ]. Under the mutation accumulation theory, the persistence of asexual lineages is more problematic unless asexuals are able to minimise Genetic diversity sexual reproduction competition with sexuals through high dispersal rates [ 17 ].
Under the Red Queen hypothesis for sex [ 18 ], we should expect that heavy directional selection exerted by parasites can favour greater genetic in host populations. Parasites are more likely to infect Genetic diversity sexual reproduction common genotypes while rare genotypes, produced by sexual females, may escape infection [ 19 - 23 ].
Genetic diversity sexual reproduction reproduction is expected to be an unstable long term strategy since asexual females can only generate offspring with new genotypes through mutation. The parasite hypothesis relies on several critical assumptions: The all-else-equal assumption e.
The difference in population genetic structure between competing sexuals and asexuals may determine difference in the parasite infection load. Asexual hosts can persist in the long term, even in the presence of parasites, if they out-disperse their parasites [ 2627 ]. The parasite hypothesis also assumes that sexual
Genetic diversity sexual reproduction harbour higher levels of genetic diversity than asexual populations.
The parasite hypothesis does not select for sex per se, but for diversity [ 28 ]. Thus, high clonal diversity could erode any advantage sex.
Howard Genetic diversity sexual reproduction Lively [ 29 ] theoretically showed that host-parasite coevolution could lead to the accumulation of clones with different resistance genotypes and, in turn, to the elimination of sexual populations. Few with coexisting sexual and asexual competitors are known.
So, comparisons of genetic Genetic diversity sexual reproduction between coexisting sexual and asexual populations are scarce: Additional comparisons are needed to further evaluate the parasite hypothesis for sex.
Bag worm moths Lepidoptera: Psychidae provide an attractive case for investigating the coexistence of sexual and asexual reproduction in the same locations. In Lepidoptera, parthenogenetic reproduction is very rare. However, in the family Psychidae and especially among Genetic diversity sexual reproduction species, parthenogenesis seems to have evolved several times [ 3132 ].
A parthenogenetic Dahlica fennicellaSuomalainen and two sexual species Siederia rupicolellaSauter and D. In these small insects 3—6 mmadult females are always wingless, sessile and incapable of dispersing. Males are always winged but their Genetic diversity sexual reproduction ability is very limited and they can only fly short distances between 10 and m. Life cycle from egg to adult takes from
Genetic diversity sexual reproduction to two years, but the adults only live 3—6 days [ 33 ].
In central Finland, these bag worm moth species occur patchily in wooded habitats. The proportion of sexually and parthenogenetically reproducing species varies between locales Genetic diversity sexual reproduction the total absence of the sexual species to only their presence.
Psychid larvae are often infected by at least two common species of Hymenopteran parasitoids, e. This result could argue in favour of the parasite hypothesis for the maintenance of sex. In light of this previous result, we investigated the genetic variability and the population structure of three closely related species of bag worm moths, two strictly sexual Siederia rupicolella and Dahlica charlottae and one parthenogenetic
Genetic diversity sexual reproduction. All were polymorphic in the two sexual species whereas twelve were polymorphic in the asexual species, with only fumaric acid FUM being monomorphic.
No more than two bands were observed at Genetic diversity sexual reproduction loci in the asexual D. Ewens-Watterson [ 36 ] and Chakraborty's [ 37 ] test of neutrality indicated that the polymorphism observed, at the scale investigated, can confidently be assumed to be neutral.
Isoenzymatic loci scored for Siederia rupicolellaDahlica charlottae and D. Recipes for buffers used are found at http: Sufficient sampling of all three species in each population for population genetic analyses was not possible.
There were two reasons for this; 1 although all three species were present to some extent in each location, they were not all abundant,
Genetic diversity sexual reproduction D. While sexual females secrete pheromones to attract males and do not lay eggs before mating, parthenogenetic females lay eggs in their larval case immediately after hatching
Genetic diversity sexual reproduction pupa. Species determination for sexual females was performed by experimental mating with a male.
Because the adults are very short lived, females can mated with males hatching only few days Genetic diversity sexual reproduction restricting the sample size. Sample sizes, average number of alleles per locus, allelic richness, proportion different genotypes kSimpson's index D and Evenness E D and E calculated for asexual D. In the last column are presented the averaged values per species and the F ST values among populations.
Allelic richness ranged from 1. The expected heterozygosity Hs ranged from 0. Allele richness and gene diversity Hs were similar Genetic diversity sexual reproduction the two sexual species permutation: Both sexual species
Genetic diversity sexual reproduction significantly higher allele richness and Hs than the asexual species permutation tests: The proportion of different clones k was also high in the asexual species, ranging from 0.
Evenness was very similar among asexual
Genetic diversity sexual reproduction and it was very close to 1 because of the high genotype diversity. Significant deviation from the Hardy-Weinberg equilibrium was observed in most the loci in Genetic diversity sexual reproduction three species.
In the sexual species, this deviation was due to
Genetic diversity sexual reproduction deficiency. The F IS values, over all loci indicated a significant deficiency of heterozygotes
Genetic diversity sexual reproduction all populations of both sexual species with the exception Genetic diversity sexual reproduction Isosaari Isa and Sippulanniemi 1 Sip 1 populations of D.
The exclusion of this locus, however, did not change any of our results. F IS values and frequency of null alleles for each population of the three species at each locus. Null alleles might cause deviation from H-W proportion.
The overall differentiation was significantly different in the three species. While the mean F ST value of D. Sixty Genetic diversity sexual reproduction different genotypes were detected among the 86 samples of D.
Genetic diversity sexual reproduction two of them were shared among different populations. both cases the two populations are geographically distant, and three populations in the same area Sip1, 2 and 3 and Hn, Pih and Isa, respectively did not share any genotypes. Analysis of isozyme variation in three species of psychid moths revealed that the genetic diversity of the sexual species D. Allele richness, gene and genotypic diversity were also higher in the sexual species than in the asexual species.
Higher genotype diversity in sexual than in asexual populations is most likely the logical result of recombination and was an expected result.
The sexual populations also showed higher allele diversity, which is a more intriguing outcome [ 30 ]. One possible explanation for this difference is Genetic diversity sexual reproduction the asexual lineage retained only a portion of the diversity of its sexual ancestor.
Alternatively, a lower Genetic diversity sexual reproduction locus diversity in the asexual species could reflect lower population sizes compared
Genetic diversity sexual reproduction the
Genetic diversity sexual reproduction species. A lower population size is also suggested by the higher differentiation F ST among
Genetic diversity sexual reproduction. This was in contrast with a previous analysis with allozyme markers which found limited diversity among samples of D.
This amount of genotype diversity higher than that recently observed in Potamopyrgus snails [ 20 ] and that reported in animals reviewed in[ 40 ] and in plants reviewed in [ 41 ] in the previous allozymes literature. Interestingly, clonal lineages of D. Only two genotypes were shared among distant populations.
The lack of a common broadly adapted haplotype Genetic diversity sexual reproduction over different populations is in conflict with the hypothesis of the general-purpose-genotype [ 42 ]. Instead, adaptation to different microclimates or other specific environmental conditions of these locales could the presence of many different genotypes, as suggested by Vrijenhoek's [ 43 ] frozen niche variation hypothesis.
However, we found no significant differences in morphology, size and life-history characters between two different D. Although several studies have reported allozymes as not neutral reviewed in [ Genetic diversity sexual reproduction ]in our study there were no indications that they deviate from neutrality, thus these markers are expected to be subjected more to drift than to selection.
High genotypic diversity could indicate the presence of cryptic sex in the parthenogenetic species. Although we cannot completely rule out this hypothesis, we never observed sex in the species. All parthenogenetic females lay eggs immediately after hatching from pupa and never show the characteristic behaviour of sexual females when they secrete pheromones to attract potential mates Kumpulainen et al.
Moreover, mitochondrial sequences from sexual and asexual females clearly indicate these are two different species Grapputo et al. This high genotypic diversity could also be explained by alternative types of parthenogenesis involving recombination, such as the automictic thelytoky [ 46 ].
Sexual reproduction can increase genetic variation but reduce species diversity
Genetic diversity in populations...
So, comparisons of genetic diversity between coexisting sexual and asexual populations are scarce: Table 2 Sample sizes, average number of alleles per locus, allelic richness, proportion of different genotypes k , Simpson's index D and Evenness E D and E calculated for asexual D. Mutikainen and four anonymous reviewers for their valuable comments on previous versions of the manuscript. Ecological implications of clonal diversity in parthenogenetic morphospecies. To explore the impact of sex on biodiversity, the researchers developed new theoretical models, based on ecological and evolutionary dynamics, to connect the mode of reproduction - whether sexual or asexual - to the origin of species and resulting biodiversity.
However, we found no significant differences in morphology, size and life-history characters between two different D.
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In spite of the two-fold cost of sex, greater of the higher animals reproduce sexually. The improvement of gender has antiquated suggested to be its ability, auspices of recombination, to generate greater genetic multiplicity than asexuality, thus enhancing adaptation in a changing environment. We studied the genetic variety and the population construct of three closely concomitant species of bag worm moths: These species joust for the same resources and divide up the coequal parasitoids.
Allelic richness was comparable interpolated the fleshly species but it was higher than in the asexual species. All species showed far up heterozygote non-existence and a large modulation was observed among F IS values across loci and populations. Large genetic differentiation was observed interpolated populations confirming the snuff dispersal adeptness of these species. The asexual species showed diminish genotype diverseness than the sexual species.
Nevertheless, genotype diversity was high in all asexual populations.
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