Abstract
We trace the history of the Modern Evolutionary Synthesis, and of genetic Darwinism generally, with a view to showing why, even in its current versions, it can no longer serve as a general framework for evolutionary theory. The main reason is empirical. Genetical Darwinism cannot accommodate the role of development (and of genes in development) in many evolutionary processes. We go on to discuss two conceptual issues: whether natural selection can be the “creative factor” in a new, more general framework for evolutionary theorizing; and whether in such a framework organisms must be conceived as self-organizing systems embedded in self-organizing ecological systems.
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Notes
Scholars have searched in vain for thinkers who can be identified as actually asserting “Social Darwinism” as described in Hofstadter (1944) (Bannister 1970). This is the notion that success in an unconstrained free market economy proves an individual's biological fitness to survive in the struggle for existence together with the implication that constraints on free markets and charitable practices will result in the growing inability of a nation, race, or the whole species to preserve the fit. Failure to identify likely suspects who argued for this view does not imply, however, that social Darwinism is a negligible factor in the history of Darwinism. Social Darwinism is a discourse that circulated widely under the auspices of Progressive reformers, socialists of various stripes, and left liberal “fellow travelers,” all of whom followed Marx in thinking of Darwinism as Malthusianism projected onto all of living nature in order to legitimate laissez-faire economics. Marx protested that the supposed inability of working class people to restrain their appetites and to expand population beyond resources was nothing more than a slur. However that may be, Darwin was Malthusian to the core. For him the cause of the pressure that population increase puts on resources means that only some—the fit—can and will differentially survive. How Darwin applies this principle to humans is another question. He was no social Darwinian. But the issue is also moot. Among the advantages of twentieth century Darwinism is that fitness and unfitness are not defined by particular qualities, such as cunning aggression in the case of the supposedly fit or congenital degeneracy in the case of the unfit. (see Gayon 1998 for how the definition of fitness shifted.) Nor in twentieth century Darwinism does natural selection occur only at or near Malthusian limits. Unfortunately, however, these technical reformulations of the science of Darwinism are, after almost a century, still largely unknown in popular culture, which accordingly continues to circulate (and attack) nineteenth century images of a “social Darwinism” that no one ever held. Sometimes the complicity of Darwinians with racism, imperialism, or eugenics replaced “social Darwinism” as Hofstadter defined it in order to impute some sort of “social Darwinism” to actual Darwinians (Young 1985; Hawkins 1997; Crook 2007). But this strategy muddies the waters. It also draws attention away from the fact that Darwinism as it developed after 1937 has done everything it can to dissociate itself from its former ideological sins.
The term “neo-Darwinism” was first used to refer to Weismann’s rejection of the very possibility of the inheritance of acquired characteristics and his concomitant commitment (by presumed exclusion of a third alternative) to the “omnipotence of natural selection.” Darwin himself had been tolerant of both forms of inheritance. All twentieth century versions of Darwinism are neo-Darwinian in Weismann’s sense. But neo-Darwinism so construed includes the view that selection is a matter of eliminating the unfit as well as the quite different view of the Modern Synthesis that natural selection is a positive, even a creative, force that gradually shapes small genetic variations over trans-generational time into adaptive traits. Hence, to call the latter “neo-Darwinism” is to elide an important difference.
The popular science writing of Stephen Jay Gould was a major factor in reengaging the public in evolutionary discourse.
The mathematics of kin selection was worked out by W. Hamilton in the 1960s. The application of this formalism in the l970s to cooperation in ants by Wilson, the world’s greatest authority on these social insects, gave a huge boost to the prestige of kin selection theory and in turn to the selfish gene interpretation of it. Understandably, Wilson’s recent withdrawal of support for the kin selectionist interpretation of ants has been among the most eyebrow-arching events in recent evolutionary biology in view of the support it had earlier lent to selfish gene theory (Nowak et al. 2010.) It is worth noting, however, that Wilson had never been a selfish gene theorist. On the contrary, his training and inclinations were and are more oriented toward the “superorganism” concept.
Gould, no shrinking violet, responded by calling his opponents “Darwinian fundamentalists,” a term charged with negative connotations both religious and political.
See Kevles and Hood (1992), for an early collection of debates about the prospects of the HGP. The book contains absurdly unrealistic prophecies by W. Gilbert and J. Watson himself, who had become Director of the HGP at the National Institutes of Health, as well as an early formulation of Keller’s skepticism.
References in the list of points (1)–(9) (as throughout this paper) are representative, not (by a considerable margin) exhaustive.
This fact can be seen perspicuously by considering the relationship between Dobzhansky and the developmental biologist Waddington. Waddington claimed that no evolutionary synthesis worthy of the name would be complete until it came back to the fundamental problem of development (Gilbert 1994, 1998). Dobzhansky did not disagree. He merely thought, probably wrongly, that the developmental genetics of the future would be subsumed under population genetics. Like Waddington, Dobzhansky also thought highly of another pioneering developmental geneticist, I. Schmalhausen. By using Schmalhausen’s ideas about development to express his and Waddington’s belief that natural selection is not only directional, but has evolved mechanisms for stabilizing normal development, Dobzhansky was in effect saying that natural selection applies uniquely to organisms considered as developmental systems. See Dobzhansky (1970), Gilbert (1994, 1998), and Depew (2010b). Most of the tension between the Modern Synthesis and developmental biology has been based on Mayr’s attack on another pioneering developmental geneticist, Goldschmidt. Gould, in his introduction to a reissue of Goldschmidt (1940), calls this judgment into question.
On the religious background of the adaptationist Oxford School of interpreting Darwinism, whose recent adepts have taken a decidedly atheistic turn, see England (2001).
The philosophers who urge the anti-causalist interpretation of statistics and probability in genetical Darwinism take at face value the claim of Dawkins and others that genocentrism is the most fully articulated version of genetical Darwinism. They are reacting to the overly causalist conception of what genes do in Dawkins interpretation. Still, they do impose a burden of proof on non-genocentric versions of the Modern Synthesis, even if it is a burden that can be met. We interpret Pigliucci and Kaplan (2007) and Pigliucci and Müller (2010) as meeting the burden.
We are distinguishing “full-scale” epigenesis from the more limited meaning of “epigenetic” that took hold after Weismann and is now common: everything that happens to an organism after the sequestration of germ cells. This meaning of “epigenetic” has more recently given rise to a third. The discovery that not only gene sequences but chemical markings of DNA (methylation) are heritable has led to the emergence of “epigenetics” considered as the study of these modifications. The third meaning presumes that Weismann’s sequestered germ cells are Dawkins’s “immortal replicators,” that is, coding sectors of DNA. Since methylation seems to defy the dogma that only DNA is heritable, it has been semantically assimilated to epigenesis in the second sense. Correctly understood, methylation and other strictly heritable, but non-DNA, developmental recourses should lead to restoration of the first, ancient, “full-scale” sense of epigenesis.
The sense of “Aristotelian” alluded to here might not even be Aristotle’s. See Depew (2008).
We have written on the possible ways in which self-organization can be related to natural selection (Depew and Weber 1995). Our list has been commented on, sometimes critically, by Richardson (2001), Swenson (2010), and Linde Medina (2010). We appreciate these responses. In particular, we appreciate Swenson’s point that claiming that natural selection is just self-organization in certain kinds of systems need not imply elimination or reduction of natural selection to something else (Swenson 2010). We have no dogmatic view to defend on this point. It seems clear enough to us that the issue will be decided by the progress of science itself, which includes discussion of conceptual issues.
This article is an English version of DARWINISMO: Il destino dell’evoluzione dopo la Sintesi Moderna; in Frontiere della biologia: Prospettive filosofiche sulle scienze della vita (Davis J, Michelini F, eds); Milan-Udine: Mimesis, in press. It appears with the permission of the editors.
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Depew, D.J., Weber, B.H. The Fate of Darwinism: Evolution After the Modern Synthesis. Biol Theory 6, 89–102 (2011). https://doi.org/10.1007/s13752-011-0007-1
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DOI: https://doi.org/10.1007/s13752-011-0007-1