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An Argument for Pluralism in the Biological Sciences Dartmouth College Dartmouth Digital Commons Open Dartmouth: Published works by Dartmouth faculty Faculty Work 12-2006 Integration without Unification: An Argument for Pluralism in the Biological Sciences Sandra D. Mitchell University of Pittsburgh Michael R. Dietrich Dartmouth College Follow this and additional works at: https://digitalcommons.dartmouth.edu/facoa Part of the Biology Commons, and the Evolution Commons Dartmouth Digital Commons Citation Mitchell, Sandra D. and Dietrich, Michael R., "Integration without Unification: An Argument for Pluralism in the Biological Sciences" (2006). Open Dartmouth: Published works by Dartmouth faculty. 2077. https://digitalcommons.dartmouth.edu/facoa/2077 This Article is brought to you for free and open access by the Faculty Work at Dartmouth Digital Commons. It has been accepted for inclusion in Open Dartmouth: Published works by Dartmouth faculty by an authorized administrator of Dartmouth Digital Commons. For more information, please contact [email protected]. vol. 168, supplement the american naturalist december 2006 Integration without Unification: An Argument for Pluralism in the Biological Sciences Sandra D. Mitchell1,* and Michael R. Dietrich2,† 1. Department of History and Philosophy of Science, University of ological practice? Is it a mark of the immaturity of bio- Pittsburgh, Pittsburgh, Pennsylvania 15260; logical theory? Will today’s pluralism give way to a grand 2. Department of Biological Sciences, Dartmouth College, unified theory of biology in the image of Newtonian phys- Hanover, New Hampshire 03755 ics? Or does pluralism indicate a healthy competition among biologists about what is the singular true account of why a system behaves the way it does? Or is it something altogether different? abstract: In this article, we consider the tension between unifi- In this article, we will consider the tension between cation and pluralism in biological theory. We begin with a consid- unification and pluralism in biological theory. We begin eration of historical efforts to establish a unified understanding of with a consideration of historical efforts to create uni- evolution in the neo-Darwinian synthesis. The fragmentation of the fication in biology on a large scale through the neo- evolutionary synthesis by molecular evolution suggests the limita- tions of the general unificationist ideal for biology but not necessarily Darwinian synthesis. The fragmentation of the evolution- for integrating explanations. In the second half of this article, we ary synthesis with the rise of molecular evolution suggests defend a specific variety of pluralism that allows for the integration the limitations of the unificationist ideal for biology. In required for explanations of complex phenomena without unification the second half of this article, we will defend a specific on a large scale. variety of pluralism that allows for the integration required for explanations of complex phenomena without unifi- Keywords: integration, unification, pluralism, evolutionary synthesis, cation on a large scale. molecular evolution. The Fragmentation of the Evolutionary Synthesis Biology studies complex, developing, and evolving organ- isms and populations. Developing sea urchins, division of Theodosius Dobzhansky’s well-worn declaration that labor in social insect colonies, and predator-prey inter- “nothing in biology makes sense except in the light of actions, for example, are all compositionally and dynam- evolution” was a testament to his faith in the unifying ically complex. Their behavior depends on multiple levels power of the evolutionary synthesis (Dobzhansky 1973). of organization and multiple causal components. Addi- From the late 1930s, Dobzhansky, Ernst Mayr, and G. G. tionally, ever since Niko Tinbergen’s articulation of four Simpson organized a loose array of biologists interested questions and Ernst Mayr’s distinction between ultimate in evolution into members of a new discipline of evolu- and proximate causes, there is general recognition of mul- tionary biology. The circumstances of this neo-Darwinian tiple levels of analysis (function, cause, development, evo- or evolutionary synthesis are still a matter of debate among lution) or questions (why and how) that can be brought historians, but certainly a unified understanding of the to bear to explain a biological property or behavior. This foundational principles of evolutionary biology was the generates a plurality of models and explanations aiming goal of the synthesis’s architects (Smocovitis 1996). As to get scientific traction on the “blooming buzzing con- Dobzhansky put it, biology without evolution “becomes fusion” that constitutes life (James 1911, p. 50). But how a pile of sundry facts some of them interesting or curious are we to understand the pluralism of contemporary bi- but making no meaningful picture as a whole” (Dob- zhansky 1973, p. 125). * E-mail: [email protected]. While historians rightly emphasize the discipline- † E-mail: [email protected]. building work of getting paleontologists and geneticists to Am. Nat. 2006. Vol. 168, pp. S73–S79. ᭧ 2006 by The University of Chicago. communicate and agree about the processes of evolution, 0003-0147/2006/1680S6-41915$15.00. All rights reserved. the hallmark of the evolutionary synthesis in terms of S74 The American Naturalist biological theory was the developing consensus about the made his modest proposal that most allele substitutions and primacy of natural selection (Provine 1988; Cain 1993; polymorphisms do not substantially affect an organism’s fit- Smocovitis 1996). The rising panselectionist belief that ness and are governed, not by positive or balancing selection, natural selection was the most important mechanism of but by random drift. (Kondrashov 2005, p. 1106) evolutionary biology was dubbed the “hardening” of the synthesis by Stephen Jay Gould (Gould 1983). A hardened While Kondrashov’s fractured fairy tale approach is com- synthesis provided a foil for biologists’ intent of setting pelling, the challenge of neutrality to panselectionism war- themselves apart from the “evolutionary orthodoxy.” rants more careful scrutiny. While, in fact, there was never complete agreement among First, Kimura, King, and Jukes initially encouraged this evolutionary biologists about the importance and power idea of opposition. When King and Jukes titled their article of natural selection, there was enough agreement to make “Non-Darwinian Evolution,” they were deliberately trying divergence from the perceived orthodoxy seem revolu- to set their work apart from that of the neo-Darwinian tionary. Motoo Kimura took advantage of these percep- synthesis. This oppositional approach certainly contrib- tions in his advocacy of the neutral theory of molecular uted to the polarization of the neutralist-selectionist con- evolution, beginning in 1968 (Kimura 1968; Dietrich troversy in the early 1970s. This sense of polarization ex- 1994). tended beyond debates about the relative importance of As molecular biology rose in prominence in the 1960s, drift and selection to the place of molecular evolution molecular evolution emerged from the intersection of relative to evolutionary biology in general. While molec- fields such as evolutionary genetics, systematics, biochem- ular biologists described evolutionary biology as old- istry, molecular biology, anthropology, and microbiology. fashioned and compared it to stamp collecting, evolution- As biologists began to compare biochemical similarities ary biologists attacked molecular biology as a fad and a and differences in order to make evolutionary inferences, bandwagon. Molecular evolution arose right in the middle they began to entertain the idea that not all molecular of this larger turf war (Dietrich 1998; Hagen 1999). From changes had been subject to natural selection (Zuckerkandl the first conferences on molecular evolution in 1962, mo- and Pauling 1965; Dietrich 1994). The possibility of neu- lecular evolutionists such as Emile Zuckerkandl had been tral mutations had been previously posited by Dobzhansky clashing with architects of the synthesis, especially G. G. and others but was not taken seriously as an alternative Simpson. Zuckerkandl was interested in using molecular to natural selection (Dobzhansky 1955). When Motoo Ki- comparisons to infer phylogenetic relationships. His ad- mura, Jack King, and Tom Jukes began to advocate the vocacy with Linus Pauling of the molecular clock and his importance of neutral mutations and random drift in 1968 disagreement with Simpson’s primate phylogeny drove a and 1969, attitudes about drift and neutrality began to wedge between molecular and morphological approaches shift (Kimura 1968; King and Jukes 1969). Many molecular to evolution and systematics. When Kimura, Alan Wilson, evolutionists with biochemical training were willing to ac- and others began to argue that the apparent constancy of cept neutral mutations on the basis of detected biochem- the molecular clock could be explained by the neutral ical differences and molecular variability, but many biol- theory, the clock controversies became enmeshed with the ogists trained in evolutionary biology and steeped in the neutralist-selectionist controversies, and the split between evolutionary synthesis were much more skeptical (Dietrich neutrality and selection became identified with a split be- 1998). tween evolution as it occurred at the molecular and mor- The controversy between the neutralists and selection- phological levels (Dietrich
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