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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