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Eunho Suh Biology of Wolbachia and Speciation Department of Entomology
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Photo by Stphen L. Dobson (http://microbewiki.kenyon.ed u/index.php/Image:Dapi.gif) Wolbachia sp. in ovarium cells of Rhagoletis cerasi (cherry fruit fly). Photo by S. Bluemel (http://microbewiki.kenyon.edu/index.php/Image:Wg1_image2.jpg)
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Over 16 % of insects Coleoptera, Diptera, Hemiptera, Homoptera, Hymenoptera, Lepidoptera, Orthoptera 35 % of terrestrial isopods 6 of 16 species of spider mites and 4 of 7 species of predatory mites 9 of 10 species of filarial nematodes (R. Stouthamer et al. 1999 ; JH Werren 1997) Distribution of Wolbachia
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Taxonomy and Phylogeny Bacteria Proteobacteria - subdivision Rickettsiales Rickettsiaceae Wolbachieae Wolbachia A & B groups : insects, mite, and crustaceans : 21 subgroups C & D groups : filarial nematodes E group : spring tail (Folsomia candida) F group : termites
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Phylogenetic Tree of Wolbachia
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(A. Jeyaprakash and M. A. Hoy 2000)
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Phenotypic Effects on Hosts Cytoplasmic Incompatibility (CI) Parthenogenesis Inducing (PI) Feminization Male Killing Other Phenomena
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Cytoplasmic Incompatibility (CI) Unidirectional CI
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Genetic Variance of CI Wolbachia 4 strains - mod + res + : wild type - mod + res - : modification, no rescue - mod - res + : no modification, rescue - mod - res - : no cytoplasmic incompatibility
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CI inducing mechanism of Wolbachia strains FM resc+mod+ resc+mod- offspring + + + + mod+ mod- resc- CI MF
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Factors Influencing CI Density of Wolbachia : high density of Wolbachia => strong CI expression, high frequency of transmission Genotype of host : genotype specific Wolbachia strains => different CI expressions or other phenomena in different host genotypes Strains of Wolbachia : different Wolbachia strains => different CI expression in same host => bidirectional CI
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Parthenogenesis Inducing (PI) Infected females produces infected females without fertilization Gamete duplication : no segregation in metaphase in first mitotic division => Diploid Haplodiploid specific (Hymenoptera spp.)
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Feminization of Genetic Males Disruption of forming androgenic gland to develop to males Feminized males produces offspring Some isopod species and two Lepidopteran species (Ostrinia furnacalis, O. scapulalis)
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Male Killing Secondary female biased sex ratios Wolbachia kills male progeny during embryogenesis Eliminating competition or providing resources to sibling females that feed dead brothers Two-spot lady bird(Aldalia bipunctata), African butterfly(Acraea encedon)
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Speciation : CI Wolbachia Sperm competition : infected sperm shows fitness advantage Tribolium confusum Longevity : Wolbachia popcorn over-replicates in host tissues => host mortality Drosophila melanogaster Host fitness effects - Positive and Negative : filarial nematode produce no progeny without Wolbachia reduction in fecundity of Trichogramma deion, T. pretiosum Other Phenomena
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Applications Biological Control Using CI Sterile male release => reduce reproductive potential of pest populations Bidirectionally incompatible Wolbachia strains => replace the existing population with less-harmful population of same species
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Speciation by Wolbachia Infectious Speciation - M. J. Wade 2001, Nature
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Evidence of Sepication by Wolbachia Wolbachia induced incompatibility precedes other hybrid incompatiblities in Nasonia - S. R. Bordenstein et al. 2001, Nature ;Bidrectional incompatibility Between Nasonia giraulti and N. longicornis No evidence of inviability and sterility among F1 hybrid females, and inviability and sterility of F2 hybrid males
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