FOREWORD
Harriet Ritvo
Charles Darwin wrote On the Origin of Species in a hurry. He had, it was true, been formulating his ideas and arguments for several decades—since his round-the-world Beagle voyage of 1831-1836. These ideas and arguments had been slow to take definitive shape; Darwin had nurtured and reworked them, amassing evidence for what he projected to be a weighty magnum opus. Although he had shared his developing evolutionary speculations with his closest professional colleagues, Darwin was reluctant to publish them on several grounds. He was aware that his theory of evolution by natural selection (or descent with modification) was complex, that it rested on vast but not incontrovertible evidence, and that the chain of his reasoning was not uniformly strong. Further, his conclusions challenged not only the scientific assumptions of many fellow specialists but also the theological convictions of a much wider circle of fellow citizens.
In 1859, Darwin did not feel quite ready to expose his cherished theory to the harsh light of public scrutiny. In the introduction to the Origin he confessed that although his work on evolution by natural selection was “nearly finished,” he would need “two or three more years to complete it.” The Origin was, he suggested, merely a stopgap, a schematic “abstract” of a much longer and more fully supported treatise yet to come. He had been moved to preview his labors in this way, he explained, because his health was “far from strong” and, perhaps more importantly, because Alfred Russel Wallace, a younger naturalist working in isolation in southeast Asia, had sent a paper to the Linnean Society of London in which he “arrived at almost exactly the same general conclusions that I have on the origin of species.” If Darwin had not gone public with his theory at this point, he would have risked losing credit for the work of many years.
As its reception showed immediately and has continued to show, the Origin benefited from the succinctness imposed by circumstances. Darwin himself may have appreciated this point; at any rate, he never produced the massive treatise, although he repeatedly issued revised editions of the Origin. But he did not abandon his intention to buttress his initial schematic presentation with additional evidence. In the course of the next two decades he published several full-length elaborations of topics summarily discussed in the Origin: The Variation of Animals and Plants under Domestication; The Descent of Man, and Selection in Relation to Sex; and The Expression of the Emotions in Man and Animals. In addition to fleshing out the Origin, these subsequent studies bolstered its arguments and responded to questions raised by critical readers, especially pragmatic questions about the way that descent with modification actually operated.
In The Variation of Animals and Plants under Domestication, which appeared first in 1868 and in a revised edition in 1875, Darwin developed a theme to which he had accorded great rhetorical and evidentiary significance. He had begun the Origin with a description of artificial selection as practiced by farmers, stock breeders, and pet fanciers, thus using a reassuringly homely example—one recognizable by the general public as well as by members of the scientific community—to introduce the most innovative component of his evolutionary theory. In addition, domesticated animals and plants, because they were numerous and available for constant observation, provided a readily available body of evidence.
Reassuring as it was, the analogy between natural and artificial selection was far from perfect. The point of Darwin’s analogy was to make the idea of natural selection seem plausible by characterizing it as a grander version of a well-known process while emphasizing its efficiency and shaping power. He noted, for example, that some of the prize birds bred by London pigeon fanciers diverged so strikingly in size, plumage, beak shape, flying technique, vocalizations. bone structure, and many other attributes, that if they had been presented to an ornithologist as wild specimens, they would unquestionably have been considered to represent distinct species, perhaps even distinct genera. Darwin argued that if the relatively brief and constrained selective efforts of human breeders had produced such impressive results, it was likely that the more protracted and thorough-going efforts of nature would work still more efficaciously.
But as Darwin acknowledged, there were some fairly obvious reasons why the two processes might diverge. The superior power of natural selection—“Man can act only on external and visible characters: nature . . . can act on . . . the whole machinery of life. Man selects only for his own good; Nature only for that of the being which she tends” (Origin, chap. 5)—might constitute a difference of kind rather than of degree, as might the much greater stretches of time available for natural selection. Further, although the mechanism of the two processes appeared superficially similar, their outcomes tended to be rather different. Natural selection produced a constantly increasing and diversifying variety of forms; it never reversed or exactly repeated itself. Anyone familiar with artificial selection would have realized that, although new breeds were constantly being developed and although neither improved wheat nor improved cattle showed any tendency to revert to the condition of their aboriginal wild ancestors, the strains produced by human selection were neither as prolific nor as durable as those produced by nature. Indeed, the animals and plants celebrated as the noblest achievements of the breeder’s art were especially liable to delicacy and infertility. Highly bred strains, long isolated from others of their species to preserve their genealogical purity, far from serving as a springboard for further variation, often had to be revivified with infusions of less-rarefied blood. Yet any relaxation of reproductive boundaries threatened subsidence into the common run of conspecifics.
Darwin firmly connected Variation to the Origin by devoting its introduction to an overview of his theory of evolution by natural selection. In particular, the two volumes of Variation, cumbersomely organized and packed with zoological and botanical detail, addressed some of the difficulties inherent in the attractive but paradoxical analogy between natural selection and artificial selection. For selection of any sort to operate, diversity already had to exist. With wild populations living under natural conditions, however, diversity was difficult to discern. It was widely believed that a heightened propensity to vary (at least in ways obvious to human observers) was one of the few general characteristics that differentiated domestic animals as a group from their wild relatives. This point was conventionally illustrated with reference to coat color and design. American bison, for example, were, on the whole, brown, and all Burchell’s zebras shared similar black and white stripes. A single herd of either Bos tauras or Equus caballus (domestic cattle or horses), on the other hand, could display colors ranging from white through yellow, red, and brown to black, as well as a variety of spotted and blotched patterns.