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Genetic dimension of the coevolution of virulence-resistance in Drosophila -- parasitoid wasp relationships.

Variations observed in parasite virulence and host resistance may be the outcome of coevolutionary processes. Recent theoretical developments have led to a 'geographic mosaic theory' of coevolution according to which there are some localities where reciprocal selection occurs (hot spots) and others where it is strongly reduced (cold spots). Studies of host-parasitoid systems back this up, revealing a geographical variation of traits subjected to antagonistic selection governed by variations in the strength of the ecological interactions. A more detailed analysis of the genetic basis of these geographic variations in a model system -- the interaction between Drosophila melanogaster and its specific parasitoid Leptopilina boulardi -- suggests that cold spots and hot spots are also driven by the amount of genetic variation available for the trait considered. Our approach, based on isolating reference strains, has been found to predict the result of sympatric interactions and it will be helpful in identifying the selective forces responsible for the coevolution. In this model, host resistance to a standardised reference strain is a weak predictor of the outcome of interactions in the field, and the main parameter accounting for the geographic variations is the number of host species available, with less parasitoid virulence towards D. melanogaster being found in areas displaying a more diversified host community.

Animals↗

Genetics of female functional virginity in the parthenogenesis-Wolbachia infected parasitoid wasp Telenomus nawai (Hymenoptera: Scelionidae).

A lepidopteran egg parasitoid species Telenomus nawai consists of two distinct populations with different reproductive modes. One is a completely thelytokous population consisting of females only, whereas the other displays arrhenotokous reproduction where fertilized eggs develop into diploid females and unfertilized eggs into haploid males. Thelytoky in T. nawai is caused by a bacterial symbiont, the parthenogenesis-inducing (PI) Wolbachia. Recent theoretical studies have shown that when a PI-Wolbachia is spreading in a population, mutations that allow uninfected females to produce more male offspring will spread rapidly eventually becoming fixed. The consequence of such a mutation is that sexual reproduction is no longer successful in infected females. Here we determine the genetic basis of the females' inability to reproduce sexually by introgressing the genome of a thelytokous line into an arrhenotokous line. The results suggest that the mutations are recessive and inherited either as a single-locus major gene with some modifiers, or as two partially linked loci.

Animals↗

Single-locus sex determination in the parasitoid wasp Cotesia glomerata (Hymenoptera: Braconidae).

The parasitoid Cotesia glomerata usually produces female-biased sex ratios in the field, which are presumably caused by inbreeding and local mate competition (LMC); yet, sibling mating increases the production of males, leading to the male-biased sex ratio of broods in the laboratory. Previous studies have suggested that the sex allocation strategy of C. glomerata is based on both partial LMC in males and inbreeding avoidance in females. The current study investigated the presence of single-locus complementary sex determination (sl-CSD) as a sex-determining mechanism in this species through inbreeding experiment, cytological examination and microsatellite analysis. Cytological examination detected diploid males in nine of 17 single pairs of sibling mating, thus in agreement with the proportion of matched matings predicted by the sl-CSD model. Sex ratio shifts in these matched sibling matings were consistent with the sl-CSD model with less viable diploid males. The haploid males have a single set of maternal chromosomes (n = 10), whereas diploid males possess a double set of chromosomes (2n = 20). Microsatellite analyses confirmed that diploid males produced from the matched matings inherited segregating genetic materials from both parents. Thus, this study provides the first solid evidence for the presence of sl-CSD as a sex-determining mechanism in the braconid genus Cotesia.

Animals↗

A new specific gene for wasp cellular immune resistance in Drosophila.

Larvae of Drosophila melanogaster produce a haemocytic reaction against eggs of the parasitoid, Asobara tabida, which leads to the formation of a multicellular capsule surrounding the foreign object. The same phenomenon was observed with the parasitoid, Leptopilina boulardi. Concerning the resistance of D. melanogaster to L. boulardi, a single major segregating locus with the resistant allele dominant to the susceptible one was found. The host strain susceptible to this parasitoid species was found to be highly immune reactive against the eggs of A. tabida. The inheritance of the capacity to encapsulate A. tabida was analysed by comparing reciprocal crosses made using inbred resistant and susceptible parental strains. We conclude that differences in the encapsulation capacity are inherited autosomally, with the reactive phenotype showing complete dominance over the non-reactive one. These data suggest the existence of two independent gene systems, each being concerned with the recognition of one species only.

Animals↗

Absence of single-locus complementary sex determination in the braconid wasps Asobara tabida and Alysia manducator.

In species with single-locus complementary sex determination (sl-CSD), sex is determined by multiple alleles at a single locus. In the haplodiploid Hymenoptera, sl-CSD results in females, if individuals are heterozygous at the sex locus, and in males, if individuals are hemizygous (haploid males) or homozygous (diploid males). Several hymenopteran species have been shown to have sl-CSD, but in several others sl-CSD is absent and the phylogenetic distribution remains unclear. In the family Braconidae, all four species tested so far were shown to possess sl-CSD. In this study, inbreeding experiments were used to test for the presence of sl-CSD in two species belonging to a subfamily of the Braconidae, Asobara tabida and Alysia manducator (Alysiinae). In both species inbreeding experiments showed no difference in brood size or sex ratio compared to the (outbred) control group. Furthermore, the sex ratios found in the inbreeding treatment differed significantly from the sex ratios expected under sl-CSD. Therefore, we conclude that sl-CSD is absent in these species. This study is the first to show the lack of sl-CSD in species of the Braconidae family and that hymenopteran sex-determining mechanisms can vary, even within a family.

Alleles↗

Meiotic and mitotic instability of two EMS-produced centric fragments in the haplodiploid wasp Nasonia vitripennis.

Terminal deletions that result in chromosomal fragments with centromeres (centric fragments) are relatively easy to generate and study in the haplodiploid insect Nasonia. We investigated the transmission stability of two chromosomal fragments generated by chemical mutagenesis. Visible mutations at the R locus (peach-233 and St-DR) and a linked body-colour mutant (purple) were used to track transmission of the centric fragments (which lack the purple locus and are wild-type at the R locus). Transmission rates in meiotic oogenesis were low (medians 0.15-0.18) and comparable to previous data on centric fragments in this species. The homologous chromosome genetic background strongly affected meiotic stability of one centric fragment (CF2) but not the other (CF1). Specifically, in peach/scarlet R locus heterozygous females, CF2 showed a normal segregation proportion with the chromosome bearing the scarlet allele (0.16), but near complete failure to segregate with peach (0.0002). Data show that this is due to loss of CF2 in eggs receiving peach, rather than to preferential segregation of CF2 with scarlet or mortality of CF2-bearing males. CF1 shows typical segregation ratios with both chromosomes. We hypothesize that deletions (or rearrangements) associated with the peach-233 mutant inhibit proper pairing and segregation of CF2. Consistent with the model, CF2 did segregate with chromosomes that had undergone recombination between peach and purple (a body-colour mutation 10 cM from peach), indicating that the domains inhibiting segregation are closely linked to peach. Mitotic instability also differed between the two fragments; reduced mitotic stability may relate to absence of telomeres on these centric fragments. Given the relative ease of generating and tracking terminal deletions in Nasonia, we propose this as a good system for studying mitotic and meiotic stability of centric fragments. Finally, results are discussed in relation to the evolution of B chromosomes from centric fragments.

Animals↗

Genetic variation and founder effects in the parasitoid wasp, Diaeretiella rapae (M'intosh) (Hymenoptera: Braconidae: Aphidiidae), affecting its potential as a biological control agent.

The effects of recent colonization on the aphid parasitoid, Diaeretiella rapae (M'Intosh) (Hymenoptera: Braconidae: Aphidiidae), in Western Australia were investigated. When compared with populations from the Old World, the results of a microsatellite analysis show that the insects have low allelic length and low allele frequency variation, revealing that these individuals experienced a significant founder effect. Marked genetic differentiation between populations was also revealed, which has potentially important implications for host utilization in this species when introduced to a new geographical area(s). Low genetic variation and gene flow in a founder population could limit evolutionary potential in Australia, including the ability of a population to mount a response to newly introduced hosts, such as the Russian wheat aphid, Diuraphis noxia (Mordvilko). Although the actual importance of genetic diversity in the success of biological control agents is unclear, current theory concerning the potential impact of genetic bottlenecks on additive genetic variance is discussed.

Animals↗

The population genetics of a biological control introduction: mitochondrial DNA and microsatellie variation in native and introduced populations of Aphidus ervi, a parisitoid wasp.

Introductions of biological control agents may cause bottlenecks in population size despite efforts to avoid them. We examined the population genetics of Aphidius ervi (Hymenoptera: Braconidae), a parasitoid that was introduced to North America from Western Europe in 1959 to control pea aphids. To explore the phylogeographical relationships of A. ervi we sequenced 1249 bp of mitochondrial DNA (mtDNA) from 27 individuals from the native range and 51 individuals from the introduced range. Most individuals from Western Europe, the Middle East and North America shared one of two common haplotypes, consistent with the known history of the introduction. However, some A. ervi from the Pacific Northwest have a haplotype that is most similar to haplotypes found in Japan, raising the possibility of a second accidental introduction. To examine population structure and assess whether a bottleneck occurred upon introduction to North America, we assayed variation at 5 microsatellite loci in 62 individuals from 2 native populations and 230 individuals from 6 introduced populations. Introduced samples had fewer rare alleles than native samples (F1,34 = 13.5, P = 0.0008), but heterozygosity did not differ significantly. These results suggest that a mild bottleneck occurred in spite of the introduction of over 1000 individuals. Using a hierarchical Bayesian approach, the founding population size was estimated to be 245 individuals. amova showed significant genetic differentiation between the European and North American samples, and a Bayesian assignment approach clustered individuals into four groups, with most European individuals in one group and most North American individuals in the other three. These results highlight that genetic changes are associated with founder events in rapidly growing natural populations, even when the founding population size is relatively large.

Analysis of Variance↗

Worker interests and male production in Polistes gallicus, a Mediterranean social wasp.

The resolution of social conflict in colonies may accord with the interests of the most numerous party. In social insect colonies with single once-mated queens, workers are more closely related to the workers' sons than they are to the queens' sons. Therefore, they should prefer workers to produce males, against the queen's interests. Workers are capable of producing males as they arise from unfertilized eggs. We found Polistes gallicus to have colonies of single, once-mated queens, as determined by microsatellite genotyping of the workers, so worker interests predict worker male production. In colonies lacking queens, workers produced the males, but not in colonies with original queens. Thus worker interests were expressed only when the queen was gone. The high fraction of missing queens and early end to the colony cycle relative to climate so early in the season is surprising and may indicate a forceful elimination of the queen.

Animals↗