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82<br />

CHARLES DARWIN AND THE ORIGIN OF SPECIES<br />

done. In 1977, the American paleontologist Stephen Jay Gould<br />

(1941–2002) wrote, ‘‘Our understanding of genetic mutation suggests<br />

that Darwin was right in maintaining that variation is not predirected<br />

in favorable ways. Evolution is a mixture of chance and necessity—<br />

chance at the level of variation, necessity in the working of selection.’’<br />

7 What historians of science call the Neo-Darwinist synthesis<br />

had occurred.<br />

Three major trends contributed to the reemergence of Darwin’s<br />

theories as a major component of the explanation for evolution. The<br />

first of these was population genetics, the study of the spread of species<br />

based on mathematics. The American scientist Sewall Wright<br />

(1889–1988) and the British scientists Ronald Aylmer Fisher (1890–<br />

1962) and J. B. S. Haldane (1892–1964) are the best-known proponents<br />

of this approach to evolution. In his book The Genetical Theory<br />

of Natural Selection (1930), Fisher suggested that genes spread<br />

through a population as discrete units; the more adaptive a gene<br />

became the faster it would increase through a population. As genes<br />

were the source of mutation, a ‘‘good’’ gene, one that adapted the<br />

best, would produce increasingly useful mutations. Haldane demonstrated<br />

that the spread of a ‘‘good’’ gene could occur much more rapidly<br />

than Fisher thought. He drew attention to examples such as the<br />

peppered moth. Between 1850 and 1900, the darker form of the<br />

moth began to dominate the species because it could hide from predators<br />

in the soot, which was a common feature of cities in Britain.<br />

Wright demonstrated that the interaction of genes was much more<br />

complex than Fisher and Haldane suggested. Multiple genes might<br />

affect one characteristic in an organism. Thus, the mutation in one<br />

gene could produce a large range of variations in a species because<br />

of its interaction with other genes. Through natural selection, the<br />

most adaptive of these variations would survive—just as Darwin had<br />

predicted.<br />

Fisher, Haldane, and Wright’s work was highly theoretical. The<br />

second major trend was the ‘‘translation’’ of the theories of population<br />

geneticists into experiments and laws that naturalists could use<br />

in their research. The American zoologist Theodosius Dobzhansky<br />

(1900–1975) was the most important contributor to this trend. In<br />

Genetics and the Origin of Species (1937), Dobzhansky explained that<br />

the application of the laws of genetics to small populations showed<br />

how a mutation could occur and spread. Like Darwin, he argued that<br />

what occurred on a microlevel could occur on a macrolevel. Mutation<br />

ultimately led to transmutation or speciation.<br />

Other scientists had been working on the connection between<br />

genetics and evolution. The Russian population geneticist Sergei

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