Abstract
THE SHARPEST critique of Wynne-Edwards came from George C. Williams, in his 1966 book Adaptation and Natural Selection: A Critique of Some Current Evolutionary Thought. Williams was driven by a distaste for multi-level evolutionary thought that went back to his early years teaching at the University of Chicago in the mid-1950s.
The triggering event may have been a lecture by A. E. Emerson, a renowned ecologist and termite specialist. The lecture dealt with what Emerson termed beneficial death, an idea that included August Weismann’s theory that senescence was evolved to cull the old and impaired from populations so that fitter youthful individuals could take their place. My reaction was that if Emerson’s presentation was acceptable biology, I would prefer another calling. (quoted by Borello 2010, p. 107)
Likewise, Williams’s tone in a letter to David Lack indicates that he considered the idea of group selection rather ridiculous.
You probably had some trouble with the wording of your discussion of Wynne-Edwards. The subject requires great care to avoid the appearance of sarcasm or ridicule. I know that when I got to that part about the epideictic function of the vertical movement of plankton [Wynne-Edwards’s Chapter 16] I suddenly wondered if I had fallen for a really elaborate joke.” (quoted by Borello 2010, p. 111)
Williams insisted that adaptations
should be attributed to no higher a level of organization than is demanded by the evidence. In explaining adaptation, one should assume the adequacy of the simplest form of natural selection, that of alternative alleles in Mendelian populations, unless the evidence clearly shows that this theory does not suffice. (Williams 1966, pp. 4–5)
Conflating the principle of parsimony with the a focus on the small, Williams continued his argument in the following terms:
Various levels of adaptive organization, from the subcellular to the biospheric, might conceivably be recognized, but the principle of parsimony demands that we recognize adaptation at the level necessitated by the facts and no higher. It is my position that adaptation need almost never be recognized at any level above that of a pair of parents and associated offspring. (Williams 1966, p. 19)
Am I living in a beautiful vacuum?
R.E.M.
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- 1.
Note the contradiction with the passage quoted above regarding the instability of genotypes. That statement implies the inability of selection to act on individual organisms.
- 2.
Note that Williams’s definition retains some flexibility in its use of the phrase “need not be closely related” rather than “are not related”.
- 3.
Williams did allow that “closeness of relationship between individuals of a colony can sometimes be seriously questioned. Even though multiple queens are normally supposed to be sisters, they would inevitably be genetically different and produce genetically different offspring. Genotypic diversity within such sister-queen colonies would be significantly greater than in the population as a whole. If it could be shown that there are thoroughly unified insect societies that normally contain several unrelated reproductives, they could only be explained as biotic adaptations resulting from effective group selection. The kinship of the reproductives would be a difficult proposition to prove one way or the other, but it is an extremely important point” (Williams 1966, pp. 200–201). However, describing the communal behavior of a decidedly diploid and non-insect species, the California woodpecker, he wrote that he “would predict … the societies of the California woodpecker, of the social insects and of all other such organized groups, will be found to be based almost entirely on family relationship” (Williams 1966, p. 202, my italics). Again, given his definition, their interactions could not be described as group selection. It is hard to avoid the conclusion that Williams had decided the case before even entering the courtroom.
- 4.
In this context, note recent work reviewed by Scheffer et al. (2009, 2012) and by Jeff Gore’s group at MIT (Chen et al. 2014; Dai et al. 2012, 2013) on fluctuations preceding population collapse. There may indeed be “value” in regulating population fluctuations (though this does not mean, of course, that such regulation actually has evolved as a group-level trait).
- 5.
It would affect the amount of genetic diversity in the population, however.
- 6.
The ability of RNA to do this, discovered by Tom Cech in the self-splicing activity of RNA in Tetrahymena thermophila, led to the hypothesis of the “RNA world”, in which genetic information was initially encoded in RNA rather than DNA.
- 7.
I would argue, and I believe so would many other scientists, that natural selection does nothing of the kind. It favors fitness, it favors offspring who survive, even if they are a bit different from their parents. Indeed, natural selection sometimes favors an increase in mutation rate, as will be discussed below in Chap. 14.
- 8.
Panagrellus redivivus. They also live in book-binding glue.
- 9.
Okasha (2006) makes precisely this argument, referencing the work of Szathmáry and Demeter. “[W]hat Dawkins misses is that [the ‘ganging up’ of genes] in effect invokes group selection. From the selective point of view, replicating molecules combining themselves into compartments is strictly analogous to individual organisms combining themselves into colonies or groups (Szathmáry and Demeter 1987). But Dawkins is an implacable opponent of group selection, insisting on the impotence of selection for group advantage as an evolutionary mechanism. Clearly, Dawkins does not appreciate that evolutionary transitions necessarily involve selection at multiple levels.” (Okasha 2006, p. 222)
- 10.
- 11.
This statement is somewhat belied, or at least muddied, by the adoption, early in The Selfish Gene, of a definition of the gene in terms of its role as a unit of natural selection. “The definition I want to use,” Dawkins wrote, “comes from G. C. Williams. A gene is defined as any portion of chromosomal material that potentially lasts for enough generations to serve as a unit of natural selection” (Dawkins 1976, p. 28).
- 12.
Caddisflies will construct nests out of whatever material is available in their environment, whether small stones or plant matter, as shown in (Fig. 7.2). The artist Hubert Duprat “collaborates” with caddisflies by providing them with jewels and precious metals with which to build their houses.
- 13.
Such a transition might resemble that likely undergone by mitochondria and other organelles during the evolution of eukaryotic cells, as suggested by Lynn Margulis (whom Dawkins does not mention).
- 14.
This is not the case in aggregative multicellular organisms such as Dictyostelium, however (see Chap. 10).
- 15.
At a recent advertising festival at Cannes, Dawkins pointed out that internet memes do not mutate by chance, but produced by human attempts at creativity. (In 2016, we learned that they can be mutated for political purposes as well.)
- 16.
In another example, Dawkins rather acidly remarks, in terms that will not surprise readers of his more recent works, that “the meme for blind faith secures its own perpetuation by the simple unconscious expedient of discouraging rational inquiry” (Dawkins 1976, p. 198). Furthermore, in his review (“Caricature of Darwinism”) of The Selfish Gene in the March 17, 1977 issue of Nature, Lewontin remarked, with even lower pH, that Dawkins has failed to consider another, far more plausible explanation for the perpetuation of the idea of hellfire, namely that it is not self-perpetuating at all, but simply is “perpetuated by some people because it gives them power over other people” (Lewontin 1977).
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Bahar, S. (2018). The Vanishing Point Appears. In: The Essential Tension. The Frontiers Collection. Springer, Dordrecht. https://doi.org/10.1007/978-94-024-1054-9_7
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