Abstract
Darwinism is one of several research traditions in evolutionary biology. I identify it, both before and after its unification with genetics, with Darwin’s theory of descent by natural selection from a common ancestor. Other traditions include saltationism/mutationism, Lamarckism, and evolutionary developmentalism (“evo-devo”). I argue that Darwinism’s continued dominance in evolutionary science reflects its proven ability to interact productively with these other traditions, an ability impressed on it by its founder’s example. Evolution by sudden leaps (saltations) is alien to the spirit of Darwinism, but Darwinism advanced its own agenda by incorporating and subverting saltationist themes. Similarly, Lamarckism’s belief in the heritability of acquired characteristics has been discredited, but some of the facts to which it seems congenial reappear in genetic Darwinism as phenotypic plasticity and niche construction. These examples help assess challenges to Darwinism’s hegemony currently arising from the role of regulatory genes and epigenetic factors in development. Rather than executing already entrenched genetic programs and relying on chance mutation to initiate evolutionary change, the developmental process appears to generate heritable variations that ab initio respond to environmental factors in an adaptive way.
Notes
- 1.
Biology, evolutionary biology, evolutionary theory, and philosophy of biology differ but form a continuum. The center of gravity in this chapter is between the second and the third of these forms of inquiry, with glances at the fourth.
- 2.
“Neo-Darwinism” originally referred to August Weismann’s (1834–1914) belief that natural selection working exclusively on germ-line heritability is the sole cause of evolution. “Hard heredity” is a necessary condition for the Darwinism of the Modern Evolutionary Synthesis, but the Synthesis rejects the “all-sufficiency (Allmacht) of natural selection” and takes a population-level view of evolutionary processes.
- 3.
Lamarck used the term publicly for the first time in 1802. In Latin, “biologia” occurred in the 1760s in the philosophical works of Christian Wolff and his disciples but with a different meaning.
- 4.
Texts from letters to and from Darwin are cited by their identifying numbers in the Darwin Correspondence Project, https://www.darwinproject.ac.uk/letters
- 5.
For Darwin, adaptations evolve by natural selection in order to and because they perform biological functions. He included the functionalist but anti-evolutionist Cuvier on his short list of heroes but not the incipiently evolutionist but structuralist Geoffroy (Darwin to Ogle January 17, 1882, #13622). His other heroes, Aristotle and Linnaeus, were also non-evolutionary functionalists.
- 6.
- 7.
The phase “factors of [organic] evolution” was first used in Herbert Spencer (1887); the list of candidates is still growing. “Creative factor” was probably due to the influence of Henri Bergson’s Creative Evolution (1907), even if Morgan, Dobzhansky, and others ascribed evolutionary innovation and direction to factors other than Bergson’s inner-driven, intuitively apprehended source of change (élan vital) (Loison and Herring 2017).
- 8.
Fisher proved mathematically that under certain conditions natural selection can favor heterozygotes, but, unlike Dobzhansky, he did not assign an evolutionary function to this scenario (Fisher 1930). Dobzhansky’s encounter with French evolutionists may have been a source of the distinction he drew between adaptations to specific environments (which can be traps) and heterotic adaptations for adapting (Loison and Herring 2017). As a young man, Dobzhansky read Bergson. He devoted his career to showing that natural selection can explain tendencies that Bergson’s followers ascribed to an inner drive that can be philosophically intuited but not experimentally proven. When he argued that, “Nothing in biology makes sense except in the light of evolution,” Dobzhansky’s overt target was creationism, but his point has wider significance (Dobzhansky 1973). In contrast to traits that evolutionary and non-evolutionary observers alike can agree are adaptive—the differently shaped beaks of the finches Darwin found on neighboring islands in the Galapagos, for example—evolutionary history’s most important adaptations cannot even be seen by pre- and anti-evolutionary biologists, let alone be explained by them.
- 9.
Intelligent design creationists have glommed on to neutral mutation and evenly ticking molecular clocks as reasons for disputing not just natural selection but evolution itself. Not surprisingly, they have been loath to take note of complications suggesting the workings of natural selection after all (Hofmann 2017).
- 10.
A caveat. In the interwar period, a talented circle of French geneticists, many with training in and funding from other countries, began experiments in physiological genetics. This work, conducted in research institutes, positioned Boris Ephrussi, André Lwoff, Francois Jacob, and Jacques Monod (who with Jacob discovered the lac operon, the first regulatory genetic mechanism to be understood) to take the lead in studies of gene regulation in the 1960s, in the process restoring the decisive importance of experiment (Burian and Gayon 1999; Loison and Herring 2017). At that time, molecular geneticists in America were still preoccupied with nailing down the genetic code and finding the mechanisms and pathways of protein production.
- 11.
American neo-Lamarckism, prominent in the nineteenth century, did not last far into the twentieth. Unlike their French counterparts, American evolutionary biologists in the interwar period embraced genetic determinism and negative eugenics—eugenics aimed at preventing the supposed unfit from reproducing in contrast to the positive eugenics that flourished in the United Kingdom, which aimed at breeding a fitter governing class—in ways that tended to support the racism with which the United States still struggles (Kevles 1985). Dobzhansky worked with American anthropologists to develop a version of the Modern Synthesis that opposed all three: genetic determinism, eugenics, and racism (Jackson and Depew 2017).
- 12.
“It takes an enormous amount of biological machinery for genes to be expressed; exactly which parts of the genome are processed depends on specific settings and structure of that machinery” (Burian and Kampourakis 2013: 613).
- 13.
- 14.
Classical epigenesis and contemporary epigenetics do not refer to the same thing but do have historical connections. Neo-Darwinism reduced the scope of nongenetic forms of heritability, such as cytoplasmic inheritance, almost to zero. Those who defended the latter, notably C. H. Waddington, referred to all aspects of inheritance as “epigenetic.” The epigenome includes genes but goes beyond them. By stressing the ontogenetic locus in which an array of reproductive factors interact as “developmental resources,” current advocates of the evolutionary significance of epigenetic modifications contest whether DNA is the sole carrier of biological heritability. Seeing residues of preformationism lurking in the notion of molecules that carry and transmit “information,” they sometime call for a new form of preformationism’s ancient antagonist, epigenesis (Oyama et al. 2001).
- 15.
Embedding organisms in ecological systems brings into view the lawful thermodynamic imperatives to which ecological systems must conform. In thermodynamically open, far from equilibrium systems, variation and selection of efficient dissipative pathways is inevitable. These physical and chemical imperatives permit, or even encourage, the emergence of developmental systems in which variation and selection take specifically biological forms. A lesson favorable to integrating evo-devo and Darwinism is that adaptive natural selection properly so called can take place only in developmental systems, which in turn are entrained with the environments by which they are co-defined.
- 16.
Coyne and Orr (2004) summarize the methods and results of speciation research. That Coyne is an opponent of expanded, extended, or new syntheses is not unconnected with his understandable desire to defend real achievements of which he is a direct heir and contributor (Coyne and Orr 1989; Coyne 2009).
- 17.
Lamarkism co-opted by Darwinism yet again!
- 18.
My thanks to Richard Delisle, Jean-Baptiste Grodwohl, Jim Hofmann, and Bruce Weber for helping me improve this chapter.
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Depew, D.J. (2017). Darwinism in the Twentieth Century: Productive Encounters with Saltation, Acquired Characteristics, and Development. In: Delisle, R. (eds) The Darwinian Tradition in Context. Springer, Cham. https://doi.org/10.1007/978-3-319-69123-7_4
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