The sex-ratio polymorphism (SR) of Drosophila pseudoobscura provides a particularly interesting system for study since it appears to be maintained in nature by a balance between and selection. The trait is a sex-linked condition which causes males to produce predominantly X-bearing sperm. Approximately 99% of progeny of their mates are females. Since corresponding value for normal males is 50%, SR chrolaosome may enjoy nearly a two-fold advantage in transmission over standard X chromosome (ST). A two-fold advantage is an extremely powerful force in evolutionary terms. Herein lies problem: despite advantage, SR is maintained in apparent equilibrium through much of southern portion of species range (Dobzhansky, 1958). In cage populations, SR is rapidly reduced in frequency or eliminated (Wallace, 1948; Anderson, 1968), and although Wallace found some of SR genotypes, particularly SR/SR females, to be at a selective disadvantage, fitnesses he measured were insufficient to account for its elimination (Edwards, 1961). Policansky and Ellison (1970) proposed a solution: since mechanism of SR involves elimination of half of spermatids per spermatid bundle in SR males, they suggested that SR males transfer fewer sperm and therefore fertilize fewer eggs than ST males. Policansky (1974) provided experimental evidence of a fertility deficiency of SR males and suggested that SR-mated females run out of sperm and remate sooner than ST-mated females. He concluded that the SR X-chromosome is not driven (i.e., has no advantage by virtue of aberrant segregation ratios) with respect to normal X-chromosome. It should be noted that this use of term is somewhat broader than definition originally provided by Sandler and Novitski (1957) since it includes fertility differences between SR and ST males in addition to effect of segregation ratios. To sharpen distinction between and fitness parameters, term meiotic drive will be used in this paper to denote advantage which aberrant segregation ratios would produce in absence of any fitness differences between SR and ST males. Thus any differences in fertility, mating success or sperm displacement will be considered fitness parameters even if differences are direct results of mechanism of the. SR trait. I attempted to study fertility differences between SR and ST males, but was able to demonstrate fertility disadvantage to SR males only under two conditions: when males were very young (20-33 h from eclosion) or after their supply of sperm had been previously reduced by repeated matings (Beckenbach, 1978). I also found that females inseminated by males whose supply of sperm had been partially depleted by previous matings showed no reduction in either fecundity or egg hatchability early in their reproductive period, compared to more heavily inseminated females. Thus fertility deficiency proposed by Policansky and Ellison is conditional at best, and populational consequences of any deficiency are minimized by pattern of fertilized egg production of females. Thus on basis of information obtained by studying singly-mated females, deficiency is insufficient to account for failure of SR to increase in frequency in nature or to cause its rapid elimination from experimental populations. However, picture may be radically different if multiple mating of females is allowed.
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