Drive and sperm: The evolution and genetics of male meiotic drive

Some selfish genetic elements in eukaryotic genomes have been harnessed to perform essential functions for their hosts, whereas others have gained transmission advantages at the expense of their hosts. Meiotic drive elements are particularly dramatic examples of the latter. Meiotic drive is the unequal—and thus non-Mendelian—transmission of alternative alleles or chromosomes from heterozygotes. Most drive elements achieve greater than Mendelian transmission from their carriers by excluding, impairing or killing competing alternative gametes. The best-characterized drive elements are the Segregation Distorter (SD) complex in the fruitfly Drosophila melanogaster and the t-haplotype in the Mus house mouse species. Despite being found in very distantly related species, both meiotic drive systems share genetic and chromosomal features that appear to characterize meiotic drive systems in general. For these, and all systems characterized to date, drive involves an interaction between at least two loci in which a trans-acting drive allele at one locus impairs transmission of sensitive alleles at a cis-acting target locus. In effect, the drive allele produces a toxin that incapacitates sperm bearing sensitive forms of the target. Drive chromosomes bear a driver and a drive-insensitive target, whereas non-drive chromosomes lack the driver and typically carry a drivesensitive target. This chapter reviews the genetics, molecular basis, and evolutionary history of several well-characterized drive systems, and considers the impact of drive on spermatogenesis and sperm competition, including potential evolutionary responses of organisms to drive.