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The weaker sex is heterogametic: 75 years of Haldane's rule.

J . B. S. HALDANE (1892-1964) made his greatest contributions to biology in the area of theoretical population genetics, especially the dynamics of natural selection. Along with R. A. FISHER and SEWALL WRIGHT, he is considered one of the founding fathers of population genetics and a major player in the development of the modern synthetic theory of evolution. This work alone identifies WANE as one of the great scientists of the twentieth century, but he also made important contributions in an amazing variety of areas including enzyme kinetics (the BRIGGS-HALDANE equation), biochemical genetics, physiology, biostatistics, linkage (the first mapping function), mutation rates in humans, and the origin of life (WHITE 1965; CROW 1992). He also noticed and described an interesting evolutionary pattern that came to be known as HALDANE’S rule. One of HALDANE’s earliest papers (1922) is a compilation of data from the literature on sex-specific inviability and sterility in hybrids between animal species. An analysis of 45 Lepidopteran, 10 bird, and six mammalian crosses, as well as a few other types of animal hybrids, led him to formulate a “rule” with only one or two known exceptions: “When in the F1 offspring of two different animal races one sex is absent, rare, or sterile, that sex is the heterozygous [heterogametic] sex.” In birds and Lepidopterans, females (known to be heterogametic) were more often inviable or sterile, while in mammals the heterogametic males were more often affected. In his 1922 paper, HALDANE stated an earlier suggestion of A. H. STURTEVANT that a deficiency of one sex might be due to inviability or to sexual transformation. He noted that STURTEVANT had demonstrated lethality of male larvae in a Drosophila cross,  while  GOLDSCHMIDT and HARRISON had shown partial sexual transformation in moth hybrids. He recognized that genetic incompatibilities may result in inviability in hybrids, but did not offer a clear explanation of why the heterogametic sex would be affected more often. With regard to sexual transformation, he suggested that an imbalance of sexdetermining factors would be less serious in the homogametic sex, since “the effect of the two 2