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1.
Several hymenopteran parasitoids are infected with parthenogenesis‐inducing (PI) Wolbachia. Infected wasps produce daughters instead of sons from unfertilized eggs. Thus far, little is known about the direct effects of PI Wolbachia on their host's fitness. Here, we report reduced competitive ability due to Wolbachia infection in a minute parasitoid wasp, Trichogramma kaykai Pinto and Stouthamer (Hymenoptera: Trichogrammatidae). Immature survival of infected individuals in a host parasitized by a single infected female, laying a normal clutch of eggs, was lower than those parasitized by a single uninfected individual. When the offspring of infected and uninfected females shared the same host, the infected immatures had significantly lower survival rates than their uninfected counterparts. The survival rate of infected immatures was higher when they competed with other infected immatures from a different infected parent than in competition with uninfected immatures of conspecific wasps. Thus, the host Trichogramma can suffer a substantial reduction in fitness when it is infected with the PI Wolbachia. We discuss why such a reduction is to be expected when populations of infected and uninfected individuals co‐occur, and how the reduced competitive ability of PI Wolbachia influences the spread of the bacteria in the field.  相似文献   

2.
Kremer N  Huigens ME 《Molecular ecology》2011,20(17):3496-3498
A huge variety of Arthropod species is infected with endosymbiotic Wolbachia bacteria that manipulate their host’s reproduction to invade populations. In addition to vertical transmission from mother to offspring through the egg cytoplasm, it has been demonstrated through phylogenetic analyses and natural transfer experiments that horizontal transmission of Wolbachia (i.e. contagion) can occur between Arthropod hosts. More recently, factors influencing horizontal transfer have also been explored. While it is clear that horizontal transmission between species plays a major role in the evolutionary history of Wolbachia infections among insects, its role in the spread of a new infection through a host population, notably through within‐species transfers, remained unknown. In this issue of Molecular Ecology, Kraaijeveld et al. (2011) present the first evidence that horizontal transmission played a key role in the early spread of parthenogenesis‐inducing Wolbachia through the parasitoid wasp Leptopilina clavipes. To support their finding, the authors studied genetic variation in three types of markers, including host nuclear DNA, mitochondrial DNA and Wolbachia DNA. Specifically, they examined potential associations between their diversity patterns. No diversity was detected in Wolbachia genes, indicating that a single Wolbachia strain must have infected and spread through L. clavipes. In addition, a correlation between substantial variation in mitochondrial and nuclear genotypes suggested that horizontal transmission played an important role in the current clonal genetic variation in this wasp. Such horizontal transmission could be facilitated by a specific host ecology (e.g. parasitoid wasps sharing the same host resource) and potentially impact co‐evolution between host and symbiont.  相似文献   

3.
Wolbachia are endocellular bacteria known for manipulating the reproductive systems of many of their invertebrate hosts. Wolbachia are transmitted vertically from mother to offspring. In addition, new infections result from horizontal transmission between different host species. However, to what extent horizontal transmission plays a role in the spread of a new infection through the host population is unknown. Here, we investigate whether horizontal transmission of Wolbachia can explain clonal genetic variation in natural populations of Leptopilina clavipes, a parasitoid wasp infected with a parthenogenesis‐inducing Wolbachia. We assessed variance of markers on the nuclear, mitochondrial and Wolbachia genomes. The nuclear and mitochondrial markers displayed significant and congruent variation among thelytokous wasp lineages, showing that multiple lineages have become infected with Wolbachia. The alternative hypothesis in which a single female became infected, the daughters of which mated with males (thus introducing nuclear genetic variance) cannot account for the presence of concordant variance in mtDNA. All Wolbachia markers, including the hypervariable wsp gene, were invariant, suggesting that only a single strain of Wolbachia is involved. These results show that Wolbachia has transferred horizontally to infect multiple female lineages during the early spread through L. clavipes. Remarkably, multiple thelytokous lineages have persisted side by side in the field for tens of thousands of generations.  相似文献   

4.
In some parasitoid wasps, infection by the micro‐organism Wolbachia leads to asexual reproduction. Within the Hymenoptera, the limits of distribution of parthenogenesis inducing Wolbachia have not yet been established. To address this issue, we screened all known thelytokous social hymenopteran species using a PCR assay. None was infected, and therefore we conclude that worker thelytoky evolves independently of Wolbachia in ants and bees. This supports the previously proposed hypothesis that a sex determining system based on heterozygosity may form a proximate limitation to Wolbachia‐induced parthenogenesis.  相似文献   

5.
Trichogramma wasps (Hymenoptera: Trichogrammatidae) are egg parasitoids commonly employed in augmentative biological control releases against a variety of mainly lepidopteran pests. By exploiting the mechanism by which the endosymbiotic bacterium Wolbachia induces parthenogenesis in this genus, we created a set of completely homozygous Wolbachia‐infected recombinant isofemale lines (RILs), each consisting of a different combination of the genome of two well‐characterized lines of Trichogramma pretiosum Riley. We subsequently use 16 of these RILs to investigate the effect of genetic variation on various measures associated with offspring production under laboratory conditions. Unsurprisingly, substantial differences were found between the RILs in their propensity to parasitize hosts, the number of hosts they parasitize, and the levels of mortality in their offspring. Such measures can be used to choose an optimal line for biological control purposes. A method was also developed to characterize the 16 RILs using their allelic state at five loci. Essentially, this binary system uses high‐resolution melt analysis to resolve identity at each locus, with alleles originating from either the grandmaternal or grandpaternal line, and is such that each RIL can be distinguished from each other RIL by their allelic state at one or more loci. The method facilitates the easy diagnosis of line origin when two or more lines are competing with each other in competition assays, allowing for the design of more complicated tests of parasitoid quality. Future field experiments will determine which genetic line performs best under more realistic biological control conditions. The fact that these lines are infected with parthenogenesis‐inducing Wolbachia will allow for prolonged rearing without appreciable change in their genetic makeup, which should lead to a predictable biological control performance.  相似文献   

6.
Wolbachia bacteria are obligatory intracellular parasites of arthropods and have been detected in about 70 species of parasitic wasps and three parasitoid flies. Wolbachia are transmitted cytoplasmically (maternally) and modify host reproduction in different ways to enhance their own transmission: parthenogenesis induction (PI), cytoplasmic incompatibility (CI), or feminization (F) of genetic males. Only PI and CI are known in parasitoids. PI-Wolbachia cause thelytoky in otherwise arrhenotokous parasitoids by generating diploid (rather than haploid) unfertilized wasp eggs. CI-Wolbachia cause incompatibility of crosses between infected males and uninfected females because the paternally derived chromosomes fail to decondense and are destroyed after syngamy. More complex situations arise when hosts harbor multiple infections, which can lead to bidirectional incompatibility and may be involved in parasitoid speciation. The relative fitness of infected and uninfected hosts is important to the population dynamics of Wolbachia, and more data are needed. Evolutionary conflict should be common between host genes, Wolbachia genes, and other "selfish" genetic elements. Wolbachia-specific PCR primers are now available for several genes with different rates of evolution. These primers will permit rapid screening in future studies of spatial and temporal patterns of single and multiple infection. Molecular phylogenies show that CI- and PI-Wolbachia do not form discrete clades. In combination with experimental transfection data, this result suggests that host reproductive alterations depend on the interaction between attributes of both Wolbachia and host. Moreover, Wolbachia isolates from closely related hosts do not usually cluster together, and phylogenies suggest that Wolbachia may have radiated after their arthropod hosts. Both results support considerable horizontal transmission of Wolbachia between host species over evolutionary time. Natural horizontal transmisson between parasitoids and their hosts, or with entomoparasitic nematodes or ectoparasitic mites, remains a tantalizing but equivocal possibility. Received: November 27, 1998 / Accepted: January 15, 1999  相似文献   

7.
Wolbachia are endosymbiotic bacteria known to manipulate the reproduction of their hosts. These manipulations are expected to have consequences on the population genetics of the host, such as heterozygosity levels, genetic diversity and gene flow. The parasitoid wasp Tetrastichus coeruleus has populations that are infected with parthenogenesis‐inducing Wolbachia and populations that are not infected. We studied the population genetics of T. coeruleus between and within Wolbachia‐infected and uninfected populations, using nuclear microsatellites and mitochondrial DNA. We expected reduced genetic diversity in both DNA types in infected populations. However, migration and gene flow could introduce new DNA variants into populations. We therefore paid special attention to individuals with unexpected (genetic) characteristics. Based on nuclear and mitochondrial DNA, two genetic clusters were evident: a thelytokous cluster containing all Wolbachia‐infected, parthenogenetic populations and an arrhenotokous cluster containing all uninfected, sexual populations. Nuclear and mitochondrial DNA did not exhibit concordant patterns of variation, although there was reduced genetic diversity in infected populations for both DNA types. Within the thelytokous cluster, there was nuclear DNA variation, but no mitochondrial DNA variation. This nuclear DNA variation may be explained by occasional sex between infected females and males, by horizontal transmission of Wolbachia, and/or by novel mutations. Several females from thelytokous populations were uninfected and/or heterozygous for microsatellite loci. These unexpected characteristics may be explained by migration, by inefficient transmission of Wolbachia, by horizontal transmission of Wolbachia, and/or by novel mutations. However, migration has not prevented the build‐up of considerable genetic differentiation between thelytokous and arrhenotokous populations.  相似文献   

8.

Background  

The maternally inherited, bacterial symbiont, parthenogenesis inducing (PI) Wolbachia, causes females in some haplodiploid insects to produce daughters from both fertilized and unfertilized eggs. The symbionts, with their maternal inheritance, benefit from inducing the production of exclusively daughters, however the optimal sex ratio for the nuclear genome is more male-biased. Here we examine through models how an infection with PI-Wolbachia in a previously uninfected population leads to a genomic conflict between PI-Wolbachia and the nuclear genome. In most natural populations infected with PI-Wolbachia the infection has gone to fixation and sexual reproduction is impossible, specifically because the females have lost their ability to fertilize eggs, even when mated with functional males.  相似文献   

9.
Population divergence in sexual traits is affected by different selection pressures, depending on the mode of reproduction. In allopatric sexual populations, aspects of sexual behavior may diverge due to sexual selection. In parthenogenetic populations, loss‐of‐function mutations in genes involved in sexual functionality may be selectively neutral or favored by selection. We assess to what extent these processes have contributed to divergence in female sexual traits in the parasitoid wasp Leptopilina clavipes in which some populations are infected with parthenogenesis‐inducing Wolbachia bacteria. We find evidence consistent with both hypotheses. Both arrhenotokous males and males derived from thelytokous strains preferred to court females from their own population. This suggests that these populations had already evolved population‐specific mating preferences when the latter became parthenogenetic. Thelytokous females did not store sperm efficiently and fertilized very few of their eggs. The nonfertility of thelytokous females was due to mutations in the wasp genome, which must be an effect of mutation accumulation under thelytoky. Divergence in female sexual traits of these two allopatric populations has thus been molded by different forces: independent male/female coevolution while both populations were still sexual, followed by female‐only evolution after one population switched to parthenogenesis.  相似文献   

10.
Wolbachia are endosymbiotic bacteria known to manipulate the reproduction of their hosts by, for example, inducing parthenogenesis. In most cases of Wolbachia‐induced parthenogenesis, the infection is fixed and the entire host population consists of females. In the absence of males and sexual reproduction, genes involved in sexual reproduction are not actively maintained by selection. Accumulation of neutral mutations or selection against maintenance of sexual traits may lead to their loss or deterioration. In addition, females may lose the ability to reproduce sexually due to ‘functional virginity mutations’ that may spread concomitantly with the Wolbachia infection through a population. The parasitoid wasp Tetrastichus coeruleus (Nees) (Hymenoptera: Eulophidae) forms an ideal model to study the decay of sexual functionality, because it has both Wolbachia‐infected, parthenogenetic populations and uninfected, sexual populations. We compared several components of sexual functionality of arrhenotokous (sexual) and thelytokous (parthenogenetic) T. coeruleus females. First, we tested whether arrhenotokous and thelytokous females were equally attractive and receptive to males. Second, we examined whether mating is costly to females by measuring the life span of mated and virgin females. Last, we studied the morphology of the spermathecae of arrhenotokous and thelytokous females. Mated females had shorter life spans than virgin females, showing that mating carried a fitness cost. Two sexual traits of thelytokous females have degraded compared to arrhenotokous females. Arrhenotokous and thelytokous females were equally attractive to males, but thelytokous females were unreceptive to males. Furthermore, there was a clear difference in spermathecal morphology between arrhenotokous and thelytokous females. Our data do not allow distinction between the various potential causes of such degradation. Although the longevity cost of mating may indicate selection against the maintenance of costly sexual traits, accumulation of neutral mutations, functional virginity mutations, manipulation by Wolbachia, and/or the genetic distance between the two populations may all have contributed to the decay of sexual traits in thelytokous females.  相似文献   

11.
Whereas sexual reproduction may facilitate adaptation to complex environments with many biotic interactions, simplified environments are expected to favour asexual reproduction. In agreement with this, recent studies on invertebrates have shown a prevalence of asexual species in agricultural (simplified) but not in natural (complex) environments. We investigated whether the same correlation between reproductive mode and habitat can be found in different populations within one species. The parasitoid wasp Tetrastichus coeruleus forms an ideal model to test this question, since it occurs both in natural and agricultural environments. Further, we investigated whether Wolbachia infection caused parthenogenesis in female‐biased populations. In contrast to the general pattern, in Dutch and French natural areas, we found Wolbachia‐infected, highly female‐biased populations that reproduce parthenogenetically. In contrast, populations on Dutch agricultural fields were not infected with Wolbachia, showed higher frequencies of males and reproduced sexually. However, we also found a female‐only, Wolbachia‐infected population on agricultural fields in north‐eastern United States. All Wolbachia‐infected populations were infected with the same Wolbachia strain. At this moment, we do not have a convincing explanation for this deviation from the general pattern of ecology and reproductive mode. It may be that asparagus agricultural fields differ from other crop fields in ways that favour sexual reproduction. Alternatively, Wolbachia may manipulate life history traits in its host, resulting in different fitness pay‐offs in different habitats. The fixation of Wolbachia in the United States populations (where the species was introduced) may be due to founder effect and lack of uninfected, sexual source populations.  相似文献   

12.
Among eukaryotes, sexual reproduction is by far the most predominant mode of reproduction. However, some systems maintaining sexuality appear particularly labile and raise intriguing questions on the evolutionary routes to asexuality. Thelytokous parthenogenesis is a form of spontaneous loss of sexuality leading to strong distortion of sex ratio towards females and resulting from mutation, hybridization or infection by bacterial endosymbionts. We investigated whether ecological specialization is a likely mechanism of spread of thelytoky within insect communities. Focusing on the highly specialized genus Megastigmus (Hymenoptera: Torymidae), we first performed a large literature survey to examine the distribution of thelytoky in these wasps across their respective obligate host plant families. Second, we tested for thelytoky caused by endosymbionts by screening in 15 arrhenotokous and 10 thelytokous species for Wolbachia, Cardinium, Arsenophonus and Rickettsia endosymbionts and by performing antibiotic treatments. Finally, we performed phylogenetic reconstructions using multilocus sequence typing (MLST) to examine the evolution of endosymbiont‐mediated thelytoky in Megastigmus and its possible connections to host plant specialization. We demonstrate that thelytoky evolved from ancestral arrhenotoky through the horizontal transmission and the fixation of the parthenogenesis‐inducing Wolbachia. We find that ecological specialization in Wolbachia's hosts was probably a critical driving force for Wolbachia infection and spread of thelytoky, but also a constraint. Our work further reinforces the hypothesis that community structure of insects is a major driver of the epidemiology of endosymbionts and that competitive interactions among closely related species may facilitate their horizontal transmission.  相似文献   

13.
Wolbachia pipientis is an obligately intracellular bacterium infecting a number of arthropod and nematode species. At the body level, Wolbachia infection may cause parthenogenesis, feminization of genetic males, male killing, or cytoplasmic incompatibility; it may also be asymptomatic. Of special interest is DNA transfer from Wolbachia to the host insect genome, which was discovered recently. At the cellular level, the effects caused by Wolbachia have been studied more poorly. Only one of the known insect cell lines has been obtained from an insect species (the mosquito Aedes albopictus) infected by Wolbachia. In this study, a continuous cell line Dm2008Wb1 has been obtained from embryos of Drosophila melanogaster infected under natural conditions. Wolbachia both persists in a primary cell culture and is retained upon its transformation into a continuous culture. The presence of this bacterium in cells in a free form is evidenced by the fact that tetracycline treatment can cure the cells of Wolbachia and by successful transfer of Wolbachia to another cell line (S2), where it has not been detected before.  相似文献   

14.
All Hymenoptera have a haplodiploid mode of sex determination. Although most species reproduce by arrhenotokous parthenogenesis, there are many thelytokous species, in which unfertilized eggs develop into diploid females. Thelytoky can be genetic or due to microbial infection. In the large Chalcidoidea superfamily, thelytokous parthenogenesis is almost always associated with infection of endosymbionts of the genera Wolbachia, Cardinium, and Rickettsia. Thripoctenus javae (Girault) (Hymenoptera: Eulophidae) is a larval parasitoid of the greenhouse thrips Heliothrips haemorrhoidalis (Bouché) (Thysanoptera: Thripidae), an important worldwide pest. Both the host and its parasitoid reproduce by thelytokous parthenogenesis. The main goal of this study was to test whether endosymbiotic bacteria, either those known to induce thelytokous parthenogenesis or other sex‐manipulators, are responsible for thelytoky of two geographically distinct populations of T. javae. We used sequencing of ribosomal ITS2 and 28S‐D2 and mitochondrial COI genes to molecularly characterize the two populations, antibiotic and heat treatments, and FISH of ovaries, for thelytoky studies. It was impossible to revert thelytokous individuals back to sexual reproduction and no evidence of bacterial infection was found in parthenogenetic T. javae females. This makes T. javae the second chalcidoid in which thelytokous reproduction appears not to be associated with the presence of bacterial endosymbionts.  相似文献   

15.
Parthenogenesis-inducing (PI) Wolbachia bacteria are reproductive parasites that cause infected (W +) female haplodiploid parasitoids to produce daughters without fertilization by males. Theoretically, PI Wolbachia infection should spread to fixation within Trichogramma populations as males are no longer required to produce female offspring. Infections in some naturally occurring Trichogramma populations are, however, maintained at frequencies ranging from 4 to 26%. Here we describe discrete equation models to examine if the PI Wolbachia infection in Trichogramma populations can be maintained at relatively low frequencies by mating regularity. Model outcomes suggest the probability of W + females mating could stabilize Wolbachia infection frequency at low levels in Trichogramma populations. The primary mechanism maintaining low-level PI Wolbachia infection in Trichogramma populations is reducing the survivorship from egg to adult in infected relative to uninfected females. The model successfully demonstrates that the relatively low PI Wolbachia infection frequency in host populations can be maintained by fertilization, or male rescue, of infected eggs, which avoids potentially hazardous gamete duplication that occurs during Wolbachia-induced parthenogenesis.  相似文献   

16.
The alpha‐proteobacteria of the genus Wolbachia is a widespread group of maternally inherited endosymbionts of arthropod and nematode hosts. Wolbachia infection induces a range of host phenotypes, including cytoplasmic incompatibility, male killing, feminization, and induction of thelytokous parthenogenesis. Heterogony (cyclical parthenogenesis) is a remarkable characteristic of oak gallwasps, Cynipini, the largest tribe of the Cynipidae. A few species of Cynipini are exceptional in that they are univoltine and exhibit thelytokous parthenogenesis, probably because they lost the arrhenotokous generation of their heterogonic ancestor species due to Wolbachia infection. In this study, the presence of Wolbachia was detected using polymerase chain reaction primers for the wsp genes in a thelytokous parthenogenetic species [Dryocosmus kuriphilus (Yasumatsu)] (Hymenoptera: Cynipidae: Cynipini). Approximately 29.8 and 87.1% of adults of the Zhuzhou and Fuzhou strains, respectively, were infected with Wolbachia while all females of the remaining four strains collected from other localities in China were Wolbachia free. The length of the wsp fragment of Zhuzhou and Fuzhou strains was found to be 573 and 561 bp, respectively. The nucleotide sequence of the bacterial wsp fragment indicated that the endosymbiotic bacteria of the Zhuzhou and Fuzhou strains are members of supergroup A, but belong to different clades; they probably originated from two independent infection events. In conclusion, thelytokous parthenogenesis of D. kuriphilus is not caused by Wolbachia infection and the deletion of the arrhenotokous generation is thus not associated with such an infection.  相似文献   

17.
Wolbachia is an endocytoplasmic bacterium responsible for various reproductive modifications in arthropods. In several species, Wolbachia induces a phenomenon called cytoplasmic incompatibility (CI), whereby crosses between a Wolbachia-infected male and a healthy female are incompatible. In haplodiploid species reproducing with arrhenotokous parthenogenesis, CI crosses produce only parthenogenetic males, inducing a male-biased sex ratio in the population. Here, we used two modeling approaches to evaluate the respective influences of demographic and biological parameters on Wolbachia fixation probability and on the sex ratio peak occurring during a Wolbachia invasion, and compared these parameters to values reported in the literature. Results suggest that the impact of Wolbachia invasion on population dynamics remains relatively limited, especially for parasitoids with high rates of sib-mating. The consequences for introduction of the parasitoids for biological control are discussed.  相似文献   

18.
Wolbachia is a widespread endosymbiont that induces dramatic manipulations of its host's reproduction. Although there has been substantial progress in the developing theory for Wolbachia–host interactions and in measuring the effects of Wolbachia on host fitness in the laboratory, there is a widely recognized need to quantify the effects of Wolbachia on the host fitness in the field. The wasp Anagrus sophiae, an egg parasitoid of planthoppers, carries a Wolbachia strain that induces parthenogenesis, but its effects on the fitness of its Anagrus host are unknown. We developed a method to estimate the realized lifetime reproductive success of female wasps by collecting them soon after they die naturally in the field, counting the number of eggs remaining in their ovaries and quantifying Wolbachia density in their body. We sampled from a highly infected A. sophiae population and found no evidence for Wolbachia virulence and possible evidence for positive effects of Wolbachia on realized reproductive success.  相似文献   

19.
Aims: The Aedes albopictus C7‐10 cell line was infected with Wolbachia strains wRi and wAlbB to create C7‐10R and C7‐10B cell lines, respectively. We compared two different methods, fluorescence in situ hybridization staining and SYTO11 staining, to describe these new Wolbachia infections in C7‐10. Methods and Results: Both staining methods were as efficient to stain Wolbachia. A formula was developed to quantify Wolbachia infection. The infection levels in C7‐10B and C7‐10R differed. The live stain SYTO11 was found to be useful to visualize Wolbachia in replicating host cells. Its potential cytotoxic effect at high concentration was investigated. Conclusions: C7‐10 supported two Wolbachia infections, constituting new tools to study Wolbachia–host interactions. The different infection levels suggest that wRi and wAlbB have different requirements for their survival in C7‐10 host cell line. Observation of SYTO11‐stained live cells gave new insights on Wolbachia segregation pattern during host cell mitosis. Significance and Impact of the Study: Wolbachia‐induced phenotypes in their arthropod and worm hosts could potentially be used to control pest populations. However, the mechanisms underlying these phenotypes are difficult to study because of Wolbachia’s intracellular lifestyle. The Wolbachia infections in C7‐10 described here could be used as in vitro models to investigate Wolbachia biology.  相似文献   

20.
Abstract Wolbachia are endosymbiotic bacteria that infect numerous arthropod species. Previous studies in Panama and Australia revealed that the majority of fig wasp species harbor Wolbachia infections, but that similar patterns of incidence have evolved independently with different wasp species and Wolbachia strains on the two continents. We found Wolbachia infections in 25/47 species (53%) of fig wasp associated with 25 species of Chinese figs. Phylogenetic analyses of Wolbachia wsp sequences indicated that very similar strains are not obviously found in either closely related or ecologically linked fig wasps species. The extremely high prevalence of Wolbachia in fig wasps (over 50% of species infected) is not constrained by geographical origin and is a recurrent theme of fig wasp/Wolbachia interactions.  相似文献   

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