Molecular evolution and phylogenetic utility of Wolbachia ftsZ and wsp gene sequences with special reference to the origin of male-killing

A detailed assessment of the evolution and phylogenetic utility of two genes, ftsZ and wsp, was used to investigate the origin of male-killing Wolbachia, previously isolated from the ladybird Adalia bipunctata and the butterfly Acraea encedon. The analysis included almost all available sequences of B-group Wolbachia and two outgroup taxa and showed that (1) the two gene regions differ in phylogenetic utility, (2) sequence variation is here correlated with phylogenetic information content, (3) both genes show significant rate heterogeneity between lineages, (4) increased substitution rates are associated with homoplasy in the data, (5) wsp sequences of some taxa appear to be subject to positive selection, and (6) only a limited number of clades can be inferred with confidence due to either lack of phylogenetic information or the presence of homoplasy. With respect to the evolution of male-killing, the two genes nevertheless seemed to provide unbiased information. However, they consistently produce contradictory results. Current data therefore do not permit clarification of the origin of this behavior. In addition, A. bipunctata was found to be a host to two recently diverged strains of male-killing Wolbachia that showed increased substitution rates for both genes. Moreover, the wsp gene, which codes for an outer membrane protein, was found to be subject to positive selection in these taxa. These findings were postulated to be the product of high selection pressures due to antagonistic host-symbiont interactions in this ladybird species. In conclusion, our study demonstrates that the results of a detailed phylogenetic analysis, including characterization of the limitations of such an approach, can serve as a valuable basis for an understanding of the evolution of Wolbachia bacteria. Moreover, particular features of gene evolution, such as elevated substitution rates or the presence of positive selection, may provide information about the dynamics of Wolbachia-host associations

Studies of biased sex-ratios in Adalia bipunctata L

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Inherited microorganisms, sex-specific virulence and reproductive parasitism

Parasites show an amazing repertoire of adaptations, highlighted by complex life cycles that allow both survival in the host and transmission among hosts. However, there is one heterogeneous group of microorganisms whose adaptations are perhaps even more surprising: parthenogenesis induction, feminization of genetic males, killing of male hosts and sperm-mediated sterilization of uninfected eggs. The common feature of these microorganisms is their mode of transmission: inheritance from mother to offspring. Here, we present an introduction to hereditary symbiosis, focusing on microsporidia and bacteria that manipulate host reproduction in arthropods (reproductive parasites). We also discuss the implications of one of these microorganisms, Wolbachia, for the control of arthropod pests and vectors and for the therapy of filarial diseases. Finally, we discuss whether some parasites of vertebrates might show sex-specific virulence. Copyrigh

Symbiont-mediated protection

Despite the fact that all vertically transmitted symbionts sequester resources from their hosts and are therefore costly to maintain, there is an extraordinary diversity of them in invertebrates. Some spread through host populations by providing their hosts with fitness benefits or by manipulating host sex ratio, but some do not: their maintenance in host lineages remains an enigma. In this review, I explore the evolutionary ecology of vertically transmitted symbionts and their impact on host resistance, and provide an overview of the evidence for the three-way interactions between these symbionts, natural enemies and invertebrate hosts. A number of recent empirical and theoretical studies suggest that vertically transmitted symbionts may protect their hosts from pathogens. If this ‘symbiont-mediated protection’ is widespread, it is likely that vertically transmitted symbionts contribute significantly to variation in measures of invertebrate resistance to natural enemies

Rapidly Shifting Sex Ratio across a Species Range

SummarySex ratios are subject to distortion by a range of inherited parasites [1]. Although it has been predicted that the presence of these elements will result in spatial and temporal variation in host sex ratio [2–4], testing of this hypothesis has been constrained by availability of historical data. We here determine spatial and temporal variation in sex ratio in a interaction between a butterfly and male-killing Wolbachia bacteria [5] by assaying infection presence in museum specimens, and from this inferring infection prevalence and phenotype in historical populations. Comparison of contemporary and museum samples revealed profound change in four of five populations examined. Two populations become extremely female biased, associated with spread of the male-killer bacterium. One evolved from extremely female biased to a sex ratio near parity, resulting from the infection losing male-killing activity. The final population fluctuated widely in sex ratio, associated with varying frequency of the male killer. We conclude that asynchronous invasion and decline of sex-ratio distorters combines with the evolution of host suppressors to produce a rapidly changing mosaic of sex ratio. As a consequence, the reproductive ecology of the host species is likely to be fundamentally altered over short time scales [6]. Further, the study demonstrates the utility of museum specimens as “silent witnesses” of evolutionary change

Disrupting the timing of Wolbachia-induced male-killing

Several lineages of maternally inherited symbionts have evolved the ability to kill infected females' sons, a phenomenon known as male-killing. Male-killing varies in its timing, from early (death during embryogenesis) to late (mortality of late larval instars). Following the observation that treatment of male-killer infected adult females Hypolimnas bolina with tetracycline, a bacteriostatic antibiotic, produces a delay in the timing of male death, we hypothesized that early male-killers possess the ability to kill males through bacterial activity outside of embryogenesis. We verified this hypothesis by showing that treatment of surviving larvae with the bacteriocidal antibiotic rifampicin rescues males. This discounted the hypothesis that delayed death occurred due to postponed effects of toxins produced at earlier stages, and thus supported the importance of bacterial activity in the larval phase in delayed male-killing. These results argue against the view that early male-killing is achieved by specifically targeting an early developmental process within the sex determination pathway