Darwin's Untimely Burial - Despite reports to the contrary, the theory of natural selection remains very much alive

This interesting article by Stephen Jay Gould was originally written for Natural History in October 1976.

This interesting article by Stephen Jay Gould was originally written for Natural History in October 1976 and posted on the web here.

In one of the numerous movie versions of A Christmas Carol, Ebenezer Scrooge, mounting the steps to visit his dying partner, Jacob Marley, encounters a dignified gentleman sitting on a landing. "Are you the doctor?" Scrooge inquires. "No," replies the man, "I'm the undertaker; ours is a very competitive business." The cutthrought world of intellectuals must rank a close second, and few events attract more notice than a proclamation that popular ideas have died. Darwin's theory of natural selection has been a perennial candidate for burial.

Charles Robert Darwin at the age of 51
Charles Robert Darwin at the age of 51

Tom Bethell held the most recent wake in a piece called "Darwin's Mistake (Harper's, February 1976): "Darwin's theory, I believe, is on the verge of collapse....Natural selection was quietly abandoned, even by his most ardent supporters, some years ago." News to me, and I, although I wear the Darwinian label with some pride, am not among the most ardent defenders of natural selection. I recall Mark Twain's famous response to a premature obituary: "The reports of death are greatly exaggerated."

Bethell's argument has a curious ring for most practicing scientists. We are always ready to watch a theory fall under the impact of new data, but we do not expect a great and influential theory to collapse from a logical error in its formulation. Virtually every empirical scientist has a touch of the Philistine. Scientists tend to ignore academic philosophy as an empty pursuit. Surely, any intelligent person can think straight by intuition. Yet Bethell cites no data at all in sealing the coffin of natural selection, only an error in Darwin's reasoning: "Darwin made a mistake sufficiently serious to undermine his theory. And that mistake has only recently been recognized as such... At one point in his argument, Darwin was mislead."

Although I will try to refute Bethell, I also deplore the unwillingness of scientists to explore seriously the logical structure of arguments. Much of what passes for evolutionary theory is as vacuous as Bethell claims. Many great theories are held together by chains of dubious metaphor and analogy. Bethell has correctly identified the hogwash surrounding evolutionary theory. But we differ in one fundamental way: for Bethell, Darwinian theory is rotten to the core; I find a pearl of great price at the center.

Natural selection is the central concept of Darwinian theory-the fittest survive and spread their favored traits through populations. Natural selection is defined by Spencer's phrase "survival of the fittest," but what does this famous bit of jargon really mean? Who are the fittest? And how is "fitness" defined? We often read that fitness involves no more than "differential reproductive success"-the production of more surviving offspring than other competing members of the population. Whoa! cries Bethell, as many others have before him. This formulation defines fitness in terms of survival only. The crucial phrase of natural selection means no more than "the survival of those who survive"-a vacuous tautology. (A tautology is a phrase-like "my father is a man"-contain no information in the predicate ("a man") not inherent in the subject my ("my father"). Tautologies are fine as definitions, but not as testable scientific statements-there can be nothing to test in a statement true by definition.)

But how could Darwin have made such a monumental, two bit mistake? Even his sincerest critics have never accused him of crass stupidity. Obviously, Darwin must have tried to define fitness differently-to find a criterion for fitness independent of mere survival. Darwin did propose an independent criterion, but Bethell argues quite correctly that he relied upon analogy to establish it, a dangerous and slippery strategy. One might think that the first chapter of such a revolutionary book as origin of Species would deal with cosmic questions and general concerns. It doesn't. It's about pigeons. Darwin devotes most of his first forty pages to " artificial selection" of favored traits by animal breeders. For here an independent criterion surely operates. The pigeon fancier knows what he wants. The fittest are not defined by their survival. They are, rather, allowed to survive because they possess desired traits.

The principle of natural selection depends upon the validity of an analogy with artificial selection. We must be able, like the pigeon fancier, to identify the fittest beforehand, not only by their subsequent survival. But nature is not an animal breeder; no preordained purpose regulates the history of life. In nature, any traits possessed by survivors must be counted as "more evolved"; in artificial selection, "superior" traits are defined before breeding even begins. Later evolutionists, Bethell argues, recognize the failure of Darwin's analogy and redefined "fitness" as mere survival. But they did not realize that they had undermined the logical structure of Darwin's central postulate. Nature provides no independent criterion of fitness; thus, natural selection is tautological.

Bethell then moves to two important corollaries of his major argument. First, if fitness only means survival, then how can natural selection be a "creative" force, as Darwinians insist. Natural selection can only tell us how that "a given type of animal became more numerous"; it cannot explain "how one type of animal gradually changed into another." Secondly, why were Darwin and other eminent Victorians so sure that mindless nature could be compared with conscious selection by breeders. Bethell argues that the cultural climate of triumphant industrial capitalism had defined any change as inherently progressive. Mere survival in nature could only be for the good: "It is beginning to look as though what Darwin really discovered was nothing more than the Victorian propensity to believe in progress."

I believe that Darwin was right and that Bethell and his colleagues are mistaken: criteria of fitness independent of survival can be applied to nature and have been used consistently by evolutionists. But let me first admit that Bethell's criticism applies to much of the technical literature in evolutionary theory, especially to the abstract mathematical treatments that consider evolution only as an alteration in numbers, not as a change in quality. These studies do assess fitness only in terms of differential survival. What else can be done with abstract models that traced the relative successes of hypothetical genes A and B in populations that exist only on computer tape? Nature, however, is not limited by the calculations of theoretical geneticists. In nature, A's "superiority" over B will be expressed as differential survival, but it is not defined by it-or, at least, it better not be so defined, lest Bethell et al. triumph and Darwin surrender.

My defense of Darwin is neither startling, novel, nor profound. I merely assert that Darwin was justified in analogizing natural selection with animal breeding. In artificial selection, a breeder's desire represents a "change in environment" for a population. In this new environment, certain traits are superior a priori; (they survive and spread by our breeders choice, but this is a result of their fitness, not a definition of it). In nature, Darwinian evolution is also a response to changing environments. Now, the key point: certain morphological, physiological, and behavioral traits should be superior a priori as designs for living in new environments. These traits confer fitness by an engineer's criterion on a good design, not by the empirical fact of their survival and spread. It got colder before the woolly mammoth evolved its shaggy coat.

Why does this issue agitate evolutionists so much? OK, Darwin was right: superior design in changed environments is an independent criterion of fitness. So what? Did anyone ever seriously propose that the poorly designed shall triumph? Yes, in fact, many did. In Darwin's day many rival evolutionary theories asserted that the fitness (best designed) must perish. One popular notion it-that theory of radical life cycles-was championed by a former inhabitant of the office I now occupy, the great American paleontologist Alpheus Hyatt. Hyatt claimed that evolutionary lineages, like individuals, had cycles of youth, maturity, old age, and death (extinction). Decline and extinction are programmed into the history of species. As maturity yields to old age, the best-designed individuals die and the hobbled, deformed creatures of phyletic senility take over.

Another anti-Darwinian notion, the but he theory of orthogenesis, held that certain trends, once initiated, could not be halted, even though they must lead to extinction caused by increasing inferior design. Many nineteenth-century evolutionist did (perhaps a majority) held that Irish elks became extinct because they could not halt their evolutionary increase in antler size; thus, they died-caught in trees or bowed (literally) in the mire. Likewise, the demise of saber-toothed "tigers" was often attributed to canine teeth grown so long that the poor cats couldn't open their jaws wide enough to use them.

Thus, it is not true, as Bethell claims, that any traits possessed by survivors must be designed as fitter. "Survival of the fittest" is not a tautology. It is also not the only imaginable or reasonable reading of the evolutionary record. It as testable. It had rivals that failed under the weight of contrary evidence and changing attitudes about the nature of life. It has rivals that may succeed, at least in limiting its scope.

If I am right, how can Bethell claim, "Darwin, I suggest, is in the process of being discarded, but perhaps in difference to the venerable old gentleman, resting comfortably in Westminster Abbey next to Sir Isaac Newton, it is being done as discreetly and as gently as possible with a minimum of publicity." I'm afraid I must say that Bethell has not been quite fair in his report of prevailing opinion. He cites the gadflies C. H, Waddington and H. J. Muller as though they epitomized a consensus. He never mentions the leading selectionists of our present generation-E. O. Wilson or D. Janzen, for example. And he quotes the architects of neo-Darwinism-Dobzhansky, Simpson, Mayr, and J. Huxley-only to ridicule their metaphors on the "creativity" of natural selection. (I am not claiming that Darwinism should be cherished because it is still popular; I am enough of a gadfly to believe that uncriticized consensus is a sure sign of impending trouble. I merely report that, for better or worse, Darwinism is alive and thriving, despite Bethell's obituary.)

But why was natural selection compared to a composer by Dobzhansky; to a poet by Simpson; to a sculptor by Mayr; and to, of all people, Mr. Shakespeare by Julian Huxley? I won't defend the choice of metaphors, but I will uphold the intent, namely, to illustrate the essence of Darwinism-the creativity of natural selection. Natural selection has a place in all anti-Darwinian theories that I know. It is cast in a negative role as an executioner, a headsman for the unfit (while the fit arise by such non-Darwinian mechanisms as the inheritance of acquired characters or direct induction of favorable variation by the environment). The essence of Darwinism lies in its claim that natural selection creates the fit. Variation is ubiquitous and random in direction. It supplies the raw material only. Natural selection directs the course of the evolutionary change. It preserves favorable variants and build fitness gradually. In fact, since artist fashion their creations from the raw material of notes, words, and stone, the metaphors to not strike me as inappropriate. Since Bethell does not accept a criterion of fitness independent of mere survival, he can hardly grant a creative role to natural selection.

According to Bethell, Darwin's concept of natural selection as a creative force can be no more than an illusion encouraged by the social and political climate of his times. In the throes of Victorian optimism in imperial Britain, change seemed to be inherently progressive; why not equate survival in nature with increasing fitness in the nontautological sense of improved design.

I am a strong advocate of the general argument that "truth" as preached by scientists often turns out to be no more than prejudice inspired by prevailing social and political beliefs. I have devoted several essays to this theme because I believe that helps to "demystify" the practice of science by showing its similarity to all creative human activity. But the truth of the general argument does not validate any specific application, and I maintain that Bethell's application is badly misinformed.

Darwin did two very separate things: he convinced the scientific world that evolution had occurred and he proposed the theory of natural selection as its mechanism. I am quite willing to admit that the common equation of evolution with progress made Darwin's first claim more palatable to his contemporaries. But Darwin failed in the second quest during his own lifetime. The theory of natural selection did not triumph until the 1940s. It's Victorian unpopularity, in my view, lay primarily in its denial of general progress as inherent in the workings of evolution. Natural selection is a theory of local adaptation to changing environments. It proposes no perfecting principles, no guarantee of general improvement; in short, no reason for general approbation in a political climate favoring inmate progress in nature.

Darwin's independent criterion of fitness is, indeed, "improved design," but not "improved" in the cosmic sense that contemporary Britain favored. To Darwin, improved meant only "better designed for immediate, local environment." Local environments change consistently: they get colder or hotter, wetter or drier, more grassy or more forested. Evolution by natural selection is no more than a tracking of these changing environments by differential preservation of organisms better designed to live in them: hair on a mammoth is not progressive in any cosmic sense. Natural selection can produce a trend that tempts us to think of more general progress-increase in brain size does not characterize the evolution of group after group of mammals. But big brains have their uses in local environments; they do not marked intrinsic trends to higher states. And Darwin delighted in showing that local adaptation opted produces "degeneration" in design-anatomical simplification in parasites, for example.

If natural selection is not a doctrine of progress, then its popularity cannot reflect the politics that Bethell invokes. If the theory of natural selection contains an independent criterion of fitness, then it is not tautological. I maintain, perhaps naïvely, that its current, unabated popularity must have something to do with its success in explaining the admittedly imperfect information we now possess about evolution. I rather suspect that we'll have Charles Darwin to kick around for some time.

[Stephen Jay Gould, "Darwin's Untimely Burial," Natural History 85 (Oct. 1976): 24-30.]