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Abstract

Genetic conflict occurs when one portion of the genome promotes its owntransmission at the expense of another portion. Genetic conflict is ubiquitous, and canhave significant evolutionary consequences. Meiotic drive is a kind of genetic conflictthat occurs when one allele manipulates normal processes during meiosis orgametogenesis to be overrepresented in the gametes. X-chromosomes carrying meioticdrivers are known as sex-ratio X-chromosomes (SR) because they manipulatespermatogenesis to sabotage the production of Y-bearing sperm, resulting in the carriermale siring only daughters. The North American fruit fly Drosophila neotestacea harborsan SR system found at steady frequencies of up to 30% in some populations. In thissystem, there are no known suppressors or drive and 99% of offspring produced by SRmales are daughters. In this dissertation, I investigate the molecular evolutionaryconsequences of drive in this system and identify candidate genes for the molecularmechanism. I show that SR carries large inversions and is significantly geneticallydifferentiated from the standard X-chromosome (ST), but there is evidence that gene flowoccurs between ST and SR. I find variation on SR is not present in distinct haplotypes,but is likely maintained through recombination in SR homozygous females. I thenidentified hundreds of transcripts that are differentially expressed between ST and SR inthe testes. Most of these transcripts are located on the X-chromosome, and differentialexpression begins early in spermatogenesis. I also found dozens of transcripts that showpatterns of nucleotide sequence differences between ST and SR that are suggestive ofpositive selection. I identified candidates for the mechanism of drive as differentiallyexpressed, testis-specific transcripts under putative positive selection. One of thesecandidates is a fast-evolving X-linked duplicate of the autosomal gene importin-a2, acritical part of the nuclear import pathway. At this locus and other candidates, I foundmolecular population genetic patterns consistent with involvement in the mechanism ofdrive: relaxed purifying selection in ST and positive selection in SR. Nuclear import isalso targeted by a meiotic driver in D. melanogaster, suggesting that this process may beparticularly susceptible to conflict.

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