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The Structure of Evolutionary Theory (Gould, 2002)
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GouldStructureofevolutionarytheory.jpg

Gould discusses Darwin and evolution, noting that Darwin was the first to name evolution by natural selection as the creative force of evolutionary change.

p.14 Darwin insisted that his admittedly weak and negative force of natural selection could, nonetheless, under certain assumptions (later proved valid) about the nature of variation, act as the positive mechanism of evolutionary novelty - that is, could "create the fit" as well as eliminate the unfit - by slowly accumulating the positive effects of favorable variations through innumerable generations.

p.60 Darwin recognized that his weak and negative force [JLJ - from p.14, natural selection], although surely a vera causa (true cause), could only play this creative role if variation met three crucial requirements: copious in extent, small in range of departure from the mean, and isotropic (or undirected towards adaptive needs of the organism). I would argue that Darwin's most brilliant intellectual move lay in his accurate identification, through the logical needs of his theory and not from any actual knowledge of heredity's mechanism, of these three major attributes of variation - because he recognized that natural selection could not otherwise operate as a creative force in the evolution of novelties.

p.61-62 Nearly all scientific revolutions originate as replacements and refutations of previous explanatory schemes, not as pure additions to a former state of acknowledged ignorance.

p.95 To impress readers with the power of natural selection, Darwin continually stressed the cumulative effect of small changes. He reserved his best literary lines, his finest metaphors, for this linchpin of his argument - as in this familiar passage: "It may be said that natural selection is daily and hourly scrutinizing, throughout the world, every variation, even the slightest; rejecting that which is bad, preserving and adding up all that is good; silently and insensibly working, whenever and wherever opportunity offers, at the improvement of each organic being in relation to its organic and inorganic conditions of life. We see nothing of these slow changes in progress, until the hand of time has marked the long lapse of ages" (1859, p. 84).

p.137-141 The following kind of incident has occurred over and over again, ever since Darwin. An evolutionist, browsing through some pre-Darwinian tome in natural history, comes upon a description of natural selection. Aha, he says; I have found something important, a proof that Darwin wasn't original. Perhaps I have even discovered a source of direct and nefarious pilfering by Darwin! In the most notorious of these claims, the great anthropologist and writer Loren Eiseley thought that he had detected such an anticipation in the writings of Edward Blyth. Eiseley laboriously worked through the evidence that Darwin had read (and used) Blyth's work and, making a crucial etymological mistake along the way, finally charged that Darwin may have pinched the central idea for his theory from Blyth. He published his case in a long article (Eiseley, 1959), later expanded by his executors into a posthumous volume entitled "Darwin and the Mysterious Mr. X" (1979).

Yes, Blyth had discussed natural selection, but Eiseley didn't realize—thus committing the usual and fateful error in this common line of argument—that all good biologists did so in the generations before Darwin. Natural selection ranked as a standard item in biological discourse—but with a crucial difference from Darwin's version—the usual interpretation invoked natural selection as part of a larger argument for created permanency.

Only two exceptions have been noted to this generality—both in the domain of anomalies that prove the rule. The Scottish fruit grower Patrick Matthew (in 1831) and the Scottish-American physician William Charles Wells (in 1813, published in 1818) spoke of natural selection as a positive force for evolutionary change, but neither recognized the significance of his speculation. Matthew buried his views in the appendix to a work entitled "Naval Timber and Arboriculture"; Wells published his conjecture in a concluding section, treating the origin of human races, to a paper on the medical case of a piebald woman. He presented this paper to the Royal Society in 1813, but only published it as he lay dying in 1818—as a subsidiary to his two famous essays on the origin of dew, and on why we see but one image with two eyes.

Matthew, still alive and vigorously kicking when Darwin published the Origin, wrote to express his frustration at Darwin's non-citation. Darwin offered some diplomatic palliation in the historical introduction added to later editions of the Origin, while professing, with ample justice, that he had meant no malice, but had simply never encountered Matthew's totally forgotten and inauspiciously located speculation. He responded to Matthew's ire in the Gardener's Chronicle for April 21, 1860: "I freely acknowledge that Mr. Matthew has anticipated by many years the explanation which I have offered of the origin of species, under the name of natural selection. I think that no one will feel surprised that neither I, nor apparently any other naturalist, has heard of Mr. Matthew's views, considering how briefly they are given, and that they appeared in the Appendix to a work on Naval Timber and Arboriculture."

Wells' article is particularly intriguing, if only for an antiquarian footnote, in the context of this book's focus on supraorganismal levels of selection. Although Wells has often been cited as a precursor, very few citationists have read his paper, and have therefore simply assumed that he spoke of natural selection by Darwin's route of advantages to individuals within populations. In fact, as I discovered (Gould, 1983a), Wells attributes racial differentiation in skin color to group selection among populations.

I do not wish to make overly much of this point, as "precursoritis" is the bane of historiography; yet I am tickled by the ironic tidbit, in the light of later orthodoxy, that the first formulation of natural selection went forward in the supraorganismic mode. The point should not be overstressed, if only because Wells reached this alternative by the fallacious argument that favorable variants could not spread within populations. Echoing Jenkins' later criticism of Darwin, Wells held that blending inheritance prevents the transformation of populations from within because advantageous variants "quickly disappear from the intermarriages of different families. Thus, if a very tall man be produced, he very commonly marries a woman much less than himself, and their progeny scarcely differs in size from their countrymen" (1818, pp. 434-435).

Populations must therefore be transformed by fortuitous spread and propagation within small and isolated groups: "In districts, however, of very small extent, and having little intercourse with other countries, an accidental difference in the appearance of the inhabitants will often descend to their late posterity" (p. 435). Change may then occur within an entire species by group selection among these differentiated populations:

Of the accidental varieties of man, which would occur among the first few and scattered inhabitants of the middle regions of Africa, some would be better fitted than the others to bear the diseases of the country. This race would consequently multiply, while the others would decrease, not only from their inability to sustain the attacks of disease, but from their incapacity of contending with their more vigorous neighbors. The color of this vigorous race I take for granted…would be dark. But the same disposition to form varieties still existing, a darker and a darker race would in the course of time occur, and as the darkest would be the best fitted for the climate, this would at length become the most prevalent, if not the only race, in the particular country in which it had originated (pp. 435-436).

Note Wells' unquestioned assumption that our original color must have been white, and that dark skin could only arise as a modification of the type. As a final interesting footnote, Wells denied (probably wrongly) that dark skin could be adaptive in itself, and argued for its establishment in Africa as a result of noncausal correlation with unknown physiological mechanisms for protection against tropical disease. Thus, Wells presents an "internalist" explanation based on what Darwin would later call "correlation of growth." With this argument about channels, and his basic claim for group selection, Wells' departure from Darwin's later preferences lie very much in the spirit of modern critiques, though for reasons that we would now reject (as if our anachronistic judgment mattered).

Natural selection, in this [its common] negative formulation, acted only to preserve the type, constant and inviolate, by eliminating extreme variants and unfit individuals who threatened to degrade the essence of created form. Paley himself presents the following variant of this argument, doing so to refute (in later pages) a claim that modern species preserve the good designs winnowed from a much broader range of initial creations after natural selection had eliminated the less viable forms: "The hypothesis teaches, that every possible variety of being hath, at one time or other, found its way into existence (by what cause or in what manner is not said), and that those which were badly formed, perished" (Paley, 1803, pp. 70-71).

Darwin's theory therefore cannot be equated with the simple claim that natural selection operates. Nearly all his colleagues and predecessors accepted this postulate. Darwin, in his characteristic and radical way, grasped that this standard mechanism for preserving the type could be inverted, and then converted into the primary cause of evolutionary change. Natural selection obviously lies at the center of Darwin's theory, but we must recognize, as Darwin's second key postulate, the claim that natural selection acts as the creative force of evolutionary change. The essence of Darwinism cannot reside in the mere observation that natural selection operates—for everyone had long accepted a negative role for natural selection in eliminating the unfit and preserving the type.

We have lost this context and distinction today, and our current perspective often hampers an understanding of the late 19th century literature and its preoccupations. Anyone who has read deeply in this literature knows that no argument inspired more discussion, while no Darwinian claim seemed more vulnerable to critics, than the proposition that natural selection should be viewed as a positive force, and therefore as the primary cause of evolutionary change. The "creativity of natural selection"—the phrase generally used in Darwin's time as a shorthand description of the problem—set the cardinal subject for debate about evolutionary mechanisms during Darwin's lifetime and throughout the late 19th century.

Non-Darwinian evolutionists did not deny the reality, or the operationality, of natural selection as a genuine cause stated in the most basic or abstract manner—in the form that I called the "syllogistic core" on page 125 (still used as the standard pedagogical device for teaching the "bare bones" logic of Darwinism in general and introductory college courses). They held, rather, that natural selection, as a headsman or executioner, could only eliminate the unfit, while some other cause must play the positive role of constructing the fit.

For example, Charles Lyell—whom Darwin convinced about the factuality of evolution but who never (much to Darwin's sadness and frustration) accepted the mechanism of natural selection—admitted that he had become stymied on the issue of creativity. He could understand, he wrote in his fifth journal on the "species question" in March, 1860, how natural selection might act like two members of the "Hindoo Triad"—like Vishnu the preserver and Siva the destroyer, but he simply could not grasp how such a force could also work like Brahma, the creator (in Wilson, 1970, p. 369).

E. D. Cope, chief American critic and exponent of neo-Lamarckism, chose a sardonic title to highlight Darwin's supposedly fatal weakness in claiming a creative role for natural selection. He called his book The Origin of the Fittest (1887)—a parody on Darwin's "survival of the fittest," and a motto for what natural selection could not accomplish. Cope wrote: "The doctrines of 'selection' and 'survival' plainly do not reach the kernel of evolution, which is, as I have long since pointed out, the question of 'the origin of the fittest.' This omission of this problem from the discussion of evolution is to leave Hamlet out of the play to which he has given the name. The law by which structures originate is one thing; those by which they are restricted, directed, or destroyed, is another thing" (1887, p. 226).

We can understand the trouble that Darwin's contemporaries experienced in comprehending how selection could work as a creative force when we confront the central paradox of Darwin's crucial argument: natural selection makes nothing; it can only choose among variants originating by other means. How then can selection possibly be conceived as a "progressive," or "creative," or "positive" force?

In resolving this paradox, Darwin recognized his logical need, within the basic structure of his argument, to explicate the three main requirements and implications of an argument for selection's creativity: (1) the nature of variation; (2) the rate and continuity of change; (3) the meaning of adaptation. This interrelated set of assertions promotes natural selection from mere existence as a genuine, but secondary and negative, mechanism to domination as the primary cause of evolutionary change and pattern. This set of defenses for selection's creativity therefore ranks as the second of three essential postulates, or "minimal commitments" of Darwinian logic.

As the epitome of his own solution, Darwin admitted that his favored mechanism "made" nothing, but held that natural selection must be deemed "creative" (in any acceptable vernacular sense of the term) if its focal action of differential preservation and death could be construed as the primary cause for imparting direction to the process of evolutionary change. Darwin reasoned that natural selection can only play such a role if evolution obeys two crucial conditions: (1) if nothing about the provision of raw materials—that is, the sources of variation—imparts direction to evolutionary change; and (2) if change occurs by a long and insensible series of intermediary steps, each superintended by natural selection—so that "creativity" or "direction" can arise by the summation of increments.

Under these provisos, variation becomes raw material only—an isotropic sphere of potential about the modal form of a species. Natural selection, by superintending the differential preservation of a biassed region from this sphere in each generation, and by summing up (over countless repetitions) the tiny changes thus produced in each episode, can manufacture substantial, directional change. What else but natural selection could be called "creative," or direction-giving, in such a process? As long as variation only supplies raw material; as long as change accretes in an insensibly gradual manner; and as long as the reproductive advantages of certain individuals provide the statistical source of change; then natural selection must be construed as the directional cause of evolutionary modification... Variation, in short, must be copious, small in extent, and undirected.

p.143 natural selection can only operate in a creative manner if its cumulating force builds adaptation step by step from an isotropic pool of small-scale variability.

p.149 Selection becomes creative only if it can impart direction to evolution by superintending the slow and steady accumulation of favored subsets from an isotropic pool of variation. If gradualism does not accompany this process of change, selection must relinquish this creative role and Darwinism then fails as a creative source of evolutionary novelty.

p.150 if the tiny increment of each step remains inconsequential in itself, then creativity must reside in the summation of these steps into something substantial - and natural selection, in Darwin's theory, acts as the agent of accumulation.

p.158 To accept Darwin's full argument about the creativity of natural selection, one must buy into an entire conceptual world - a world where externalities direct, and internalities supply raw material but impose no serious constraint upon change; a world where the functional impetus for change comes first and the structural alteration of form can only follow.

p.342 Darwin's evolutionary critics encountered their greatest stumbling blocks in their inability to envision natural selection as a creative force.

p.479 Darwin... identifies natural selection as a force of local adaptation.

p.552 Occam devised his famous motto, "non sunt multiplicanda entia praeter necessitatem" (entities are not to be multiplied beyond necessity), as a weapon in this philosophical battle - an argument against the existence of an ideal platonic realm... Occam's razor, in its legitimate application, therefore operates as a logical principle about the complexity of argument, not as an empirical claim that nature must be maximally simple... The Lamarckian one-step route to adaptation, for example, operates more simply and directly than the Darwinian two-step process of variation and selection. But nature happens to follow Darwin's path.

p.695 Evolution continually recycles, in different and creative ways, many structures built for radically different initial reasons.

p.965 Technology is making skills and knowledge the only sources of sustainable strategic advantage.

p.966 Columbus did not succeed because he was lucky. He succeeded because he made the effort to set sail in a direction never before taken despite a lot of resistance from those around him.

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