Richard Lewontin as Master Dialectician

November 23, 2021

Richard Lewontin as Master Dialectician

Rest in Power, Dick

By Rasmus Grønfeldt Winther

Lewontin Memorial Collection

Art by Jake Thrasher
On July 4, 2021, Richard C. Lewontin died at age 92. It is hard to overestimate this intellectual giant’s influence on evolutionary biology and on the public face and politics of science. Lewontin was larger than life, and while he is no longer among the living, his spirit endures through his far-reaching influence. His opus is vast and diverse, ranging from technical population genetics to general philosophy of science to science communication and public polemics. Lewontin trained or interacted with a significant fraction of population geneticists and philosophers of biology. Indeed, having studied in detail a good portion of his vast opus; having read extensively in evolutionary theory, genetics, ecology, and philosophy of science and biology; and having corresponded and talked at length with him, I know there have been few minds in biology and philosophy as deep, sharp, and incisive as his. The pain of his loss is still acute.

Lewontin’s influence on me was exceptionally rich. In 2003, out of the blue, I received, by post, a three-page letter from Lewontin. He had read my first single-authored article, “Darwin on Variation and Heredity,”1 and wanted to comment on it. (Later, he gave me express permission to use the letter as I wished, though I’ve not made it public before today.) I read this letter as a PhD student about to defend his dissertation, which is to say, I read it with incredulity. The article, according to Lewontin, “has made me think a lot and to reconsider some things that I have said and written in the past about Darwin.”

In the first half of the letter, he offered an inspired lecture on canalization and mutation rate increase due to environmental influences. In the second half, he accepted the analysis of my article, acknowledging the limitations of his framing “the alienation of external causation and internal factors” as “the fundamental epistemological break introduced by Darwin”:

I have not, in the past, sufficiently appreciated the ambiguities and contradictions in Darwin’s entire corpus. Especially, I had never really thought about how his theory of pangenesis carried with it the inherent contradiction with the standard view of natural selection operating on random variation.

In the last page of the letter, he poetically philosophized about the “perpetual problem” of projecting onto path-breaking scientists “an unambiguous and logical system that leads ineluctably to the modern understanding.” He called out Stephen J (“Steve”) Gould as illustrative of this danger and wrote that Gould:

…was particularly guilty of constructing a Darwin to suit his purposes, and he made me conscious of this recreation of historical figures in a modern light in the history of science. What I had not seen before reading your paper is how much that I myself have done this while pretending to be more careful.

Lewontin noted that the problems of (1) papering over “the ambiguities and contradictions” of the corpus of a key scientist and (2) “recreat[ing]” and rereading historical figures “in a modern light” (partly, what historians call Whiggishness) went back “at least to Hegel’s lectures” and “would make a wonderful subject for a book.” In just this letter, considered as a microcosmos, we see Lewontin’s simultaneous technical prowess, grand philosophical vision, and gentle encouragement of the next generations.

I want to take a cue from Lewontin’s suggestion—not to say invitation—in his letter to me, and pay homage both to the diversity of his oeuvre, and specifically to those ambiguities and contradictions that promise possible future findings. In fact, Lewontin, together with Richard Levins, his co-author and co-thinker of many years, developed a subtle analysis of dialectics, which builds on—but moves beyond—Karl Marx’s dialectical materialism, and draws explicitly from Friedrich Engels’s perhaps more abstract analysis of dialectical thinking.

In the last, methodological chapter of The Dialectical Biologist, Levins and Lewontin observe:

…the most central concept in dialectical thought [is] the principle of contradiction. For some, contradiction is an epistemic principle only. It describes how we come to understand the world by a history of antithetical theories that, in contradiction to each other and in contradiction to observed phenomena, lead to a new view of nature. Kuhn’s (1962) theory of scientific revolution has some of this flavor of continual contradiction and resolution, giving way to new contradiction. For others, contradiction is not only epistemic but political as well, the contradiction between classes being the motive power of history. Thus contradiction becomes an ontological property at least of human social existence. For us, contradiction is not only epistemic and political, but ontological in the broadest sense. Contradictions between forces are everywhere in nature, not only in human social institutions.2

For many of us, a “contradiction” implies an error, a weakness, or unproductive thinking. A contradiction seems to be a mistake. Lewontin (and Levins) argue that, on the contrary, there is rich life and potential growth in the contradictions existing in our thinking and knowledge, our political and social structures, and, indeed, in nature and the world itself.

A dialectic underscores this potentiality by describing a productive disjunction or paradox that captures or contains two contradictory—but profound—poles, concepts, or truths. Lewontin well knew that engaging in dialectical thinking allows us to build a rich conceptual space, with an immense universe of possible thoughts and actions. After all, each dialectic provides one spectrum or dimension of this space. Given his analysis, as well as his recognition that not sufficiently appreciating the ambiguities and contradictions of key thinkers such as Darwin limited our understanding of their work, he also clearly recognized the importance of the future study and continued innovation that can occur within this conceptual and semantic space.

Further, while a dialectic is sometimes limited to the identification of the complex tension between two poles of a spectrum, Lewontin implicitly argued for a “thirdness,” a unique and emergent co-development and co-evolution of the dialectical poles. He pointed out that an “aspect of contradiction is the interpenetration of seemingly mutually exclusive categories.”3 In other words, the opposite poles of a dialectic interpenetrate and thereby provide a unique, third, perspective and research program on the spectrum.

Lewontin preferred to speak of interpenetration of a dialectic—e.g., organism versus environment, parts versus wholes, and subject versus object—rather than synthesis. The choice may have also been informed by the baggage of “synthesis,” which, despite its powerful Hegelian roots, Lewontin likely associated with the modern neo-Darwinian synthesis of his dissertation advisor Theodosius Dobzhansky and with E.O. Wilson’s sociobiological “new synthesis” research program.

Given my own interests, which have been comprehensively and deeply informed by Lewontin’s work, I seek to identify the potential growth contingent on three of his richest interpenetrative dialectics and productive contradictions:

  1. An early enthusiasm for molecular reductionism followed by philosophical, integrative anti-reductionism;
  2. An anti-reductionism, which simultaneously emphasizes organismic agency and organisms as the “subject” of development and evolution, while also highlighting a hierarchy of integrative levels of organization above and below the organism;
  3. An insistence on gene interaction throughout his career and an initial enthusiasm for—followed by silence on—multi-level selection (which manifests in Lewontin’s consistent denial of “horizontal averaging” and an initial vociferous denial but eventual silence on “vertical averaging”).

Ultimately, I reflexively repeat the investigation of “the ambiguities and contradictions [of an] entire corpus,” which Lewontin modeled for me in his unexpected response to my work on Darwin. I seek, in so doing, to both celebrate and contextualize Lewontin, while also gesturing to the gains we might accrue in evolutionary biology and the philosophy of science by resisting our desire to paper over the ambiguities and contradictions of this major public face of science.

1. An early enthusiasm for molecular reductionism followed by philosophical, integrative anti-reductionism.

Lewontin completed his PhD in his mid-twenties at Columbia University under the mentorship of Theodosius Dobzhansky, one of the foremost evolutionary geneticists at the time. As is often the case, Lewontin received from his mentor not only content and knowledge, but also ways of thinking and general paradigms and research approaches. (Although relevant, here is not the place to review, for instance, the so-called classical versus balance debate between, respectively, H. J. Muller and Dobzhansky).4 For the young Lewontin, perhaps the key question of experimental population genetics was “at what proportion of his loci will the average individual in a population be heterozygous?”5

After moving to the University of Chicago, Lewontin published two pathbreaking articles with molecular geneticist John (“Jack”) Hubby. Based on an earlier molecular strategy developed by Hubby, Lewontin devised an “application” of this molecular strategy for assessing both allele variation in a population, and the typical amount of heterozygosity in individuals, within and among populations.6 It is hard to overestimate the influence of this work on the field of evolutionary genetics.7 Evolutionary genetics was now on track to adopt an experimental, reductionist strategy, which started producing the much-needed data that Lewontin felt was missing from theoretical population genetic research: “with the discoveries of molecular biology about the relationship between genes and proteins, the possibility of an unambiguous characterization of genetic variation among individuals was opened.”8 This was data on allele frequencies, and allele frequency changes, within and across populations. While measurement of reductive, genetic phenomena need not imply in-principle reductionism, measurement is not ontologically innocent. It implies assumptions about how the world works and what is worth measuring.

Later in his career, Lewontin presented a public face of opposition to many (new) reductionist molecular strategies. The Preface of The Dialectical Biologist took issue with the “Cartesian Reductionism” implied by “the extraordinary successes of mechanistic reductionist molecular biology.”9 Moreover, in the Conclusion, Levins and Lewontin “reject[ed] the molecular euphoria that has led many universities to shift biology to the study of the smallest units, dismissing population, organismic, evolutionary, and ecological studies as forms of ‘stamp collecting’ and allowing museum collections to be neglected.”10 Molecular strategies, successes, and euphoria included the sequencing of the entire human genome and the use of genetic information in forensics and identity-hunting, as well as, more generally, a reductionist faith in the explanatory and predictive power of genes to account for our body and mind. Indeed, Human Diversity (1982), The Dialectical Biologist (1985), Biology as Ideology (1993), and The Triple Helix (2002), defend an anti-reductionist paradigm in which bodies and minds emerge within a complex web of interaction of co-defined and co-entangled genes, environment, and organisms.11 Rather than turn to the genetic level for mapping, tracking, explaining, and predicting, Lewontin was now emphasizing the role of the organism and developmental system, of phenomena such as organismic plasticity, canalization, and robustness in the face of genotypic and environmental variation. The organism became the active subject of the evolutionary play, rather than the passive object of its constituent genes and surrounding environment.

Can molecular reductionism and organismic and systemic anti-reductionism (itself partly adopting holism) co-exist? Perhaps Lewontin was merely shifting scales of analysis, preferring larger and more “zoomed out” scales later in his career. Perhaps, like other great minds, he started his career as a technical wizard only to wrestle later with grand philosophical questions and paradigms (in part in response to work by the likes of E.O. Wilson and Richard Dawkins). Even so, reductionism and anti-reductionism each have a tendency to assert explanatory and predictive completeness—scientific reductionism does not cohabitate comfortably with philosophical, to not say ideological, anti-reductionism. Much remains to be learned from this grand, to not say eternal, scientific dialectic.

2. An anti-reductionism, which simultaneously emphasizes organismic agency and organisms as the “subject” of development and evolution, while also highlighting a hierarchy of integrative levels of organization above and below the organism.

Let’s look more deeply at Lewontin’s anti-reductionism, and a further implicit dialectic in it. A significant difference in explanatory, epistemic, and methodological resources exists in whether an anti-reductionist stance focuses on organisms per se or on a variety of hierarchical levels and processes, including organisms. Lewontin endorsed both, and it is not always clear which anti-reductionist ontology he preferred.

Turning first to the organism-centric stance, in Human Diversity, Lewontin contrasts three “models of the developmental process”: genes or environment as the main “determinant,” and one emphasizing organismic agency and robustness. Each model distinctly represents and understands the “interaction of causes in determining the variation among organisms.” The first two are both models of a sort of “factory production.” In the first, genetic blueprints are analogous to different car designs whereas, in the second, the choice of materials such as “wood, steel, or stone” are determinative for constructing a building, and are analogous to “environmental” input.12

In contrast:

Our [Lewontin’s third] model is not so much that of a factory as that of an artisan’s workshop. The cabinet maker knows what she is to produce, but, as she works, the materials, with their inevitable variation in texture and quality, begin to exert their own influences. The shape of the table legs emerges in part as a response to the influences of the materials. Moreover, because it is handiwork, there is some variation from leg to leg and from table to table simply because the worker is not in perfect control of the process. The relation between gene, environment, and organism is not one-to-one but many-to-many. Given the genes and the environment, one cannot predict the organism. Given the organism, we cannot infer its genotype or the environment in which it developed.13

The organism is an emergent and unique descriptive and prescriptive level. Recall also Chapter 3 of The Dialectical Biologist: “The Organism as Subject and Object of Evolution,” where Lewontin ascribed immense agency to organisms, including that organisms “modify their environments in several ways” and “transform the statistical structure of their environment,” and that “as any environmental factor impinges on the organism, the physical form of the signal changes” and even that “every part or activity of an organism acts as environment for other parts.”14 This is a powerful organism-centric approach to development, evolution, and ecology.

But especially with respect to the Dialectical View that Levins and Lewontin explicitly contrasted with Cartesian Reductionism, anti-reductionism focused less on a single emergent level, such as the organism, and more on a rich, emergent, and non-pre-existing hierarchy of shifting and dynamic processes and objects. According to their Dialectic View, objects are “internally heterogeneous at every level,”15 parts do not pre-exist the wholes they can be taken to compose, wholes are emphasized over parts,16 and “change is a characteristic of all systems and all aspects of all systems.”17 Such a multi-faceted Dialectical View emphasizes interconnection, heterogeneity, and historicity of many parts and wholes, at many levels.18 The individual organism is found only at one such level, among many.

Should anti-reductionism in biology focus on the organism or on more diffuse, fractal, and heterogeneous objects and processes, at any number of (perhaps themselves non-preexisting) levels of organization? This remains a fascinating question and quandary for anti-reductionists.

3. An insistence on gene interaction, and an initial enthusiasm for—followed by silence on—multi-level selection (which manifests in Lewontin’s consistent denial of “horizontal averaging” and an initial vociferous denial but eventual silence on “vertical averaging”).

In 1970, Lewontin published an article that had an enormous influence on the “levels of selection” debate in evolutionary theory and the philosophy of biology.19 In this piece, he outlined the three principles necessary for evolution by natural selection: (1) phenotypic variation, (2) differential fitness, and (3) heritability of fitness. He argued that “the generality” of these principles implied “that any entities in nature that have variation, reproduction, and heritability may evolve.”20 In the article, Lewontin defended multi-level selection, but at the same time, he chastised others for agreeing with multi-level selection in the absence of good evidence for it.

His article is indeed a meticulous presentation of many distinct lines of evidence for selection occurring among units at a variety of levels: “molecules, cells, populations, species, and [ecological] communities.”21 Toward the conclusion, he even asserted that “at yet higher levels, the species and the community, natural selection obviously must occur.”22 Given contemporary ecological catastrophes, the potential for which was only barely adumbrated in the 1970s, Lewontin was also prescient in concluding the piece thus: “man, by his rapid transformation of habitats, creates the conditions for efficient natural selection of species and species relationships, on a scale never before existent in evolution.”23

The scientific and philosophical literature that flowed from this article is immense and fascinating. Here I wish to make a simple point. In a paper I co-authored with Michael J. Wade, the mentor for my master’s degree in evolutionary biology, we distinguish between two debates in genetics: on the one hand, between those who defend versus deny vertical averaging, that is, averaging across the fitness of units at different levels of selection; and on the other hand, between those who defend versus deny horizontal averaging, that is, averaging across (parts of) the genomic context—across background genes.24 Lewontin initially enthusiastically opposed vertical averaging, arguing that natural selection genuinely and powerfully occurs among units at many different levels in the evolutionary hierarchy. After 1970, however, Lewontin seems to have basically abandoned, and remained mostly silent about, multi-level selection, certainly in solo-authored work.25 That is, he focused primarily on selection at the organismic level. This seems part and parcel of the organism-centric approach explored in the second dialectic above.

However, Lewontin always critiqued horizontal averaging. He fiercely insisted that different genes and their respective alleles interacted with one another in their collective effects on fitness. We cannot, in general, according to Lewontin, average across the genetic context of a single locus to theorize or measure the “additive” effect of one gene at a time. Evolution does not work as Dawkins’s “Oxford oarsmen” thought experiment would have it—we cannot determine the objective best (or worst) oarsman (or two or three…) by averaging all possible oarsmen with all possible partners in all possible environments.26 Indeed, the title of the last chapter of The Genetic Basis of Evolutionary Change is “The Genome as the Unit of Selection”—the entire genome was now selected! This chapter built on earlier technical, co-authored, key works.27 Indeed, in the penultimate and longest chapter of the 1974 book, “The Paradox of Variation,” Lewontin declaimed that the population genetics “theory machine… cannot transform into a finished product the great volume of raw material that has [now] been provided. The entire relationship between the theory and the facts needs to be reconsidered.”28 He felt that one of the gravest problems of the population genetic theory machine was that “the usual treatment of the genome as a collection of single loci ignores both the physiological interaction between loci and the fact that the genes are arranged on chromosomes and therefore do not segregate independently.”29

Because it is hard to find a more poetic and thrilling defense of gene interaction in the population genetics literature, Lewontin earns the last word on this dialectic of how to approach vertical versus horizontal averaging:

The fitness at a single locus ripped from its interactive context is about as relevant to real problems of evolutionary genetics as the study of the psychology of individuals isolated from their social context is to an understanding of man’s sociopolitical evolution. In both cases context and interaction are not simply second-order effects to be superimposed on a primary monadic analysis. Context and interaction are of the essence.30

Each of these three dialectics are central to core knowledge, ontology, and methodology of evolutionary theory. The first is about overarching research strategies, and how we should think and do science: can we, and should we try to, reduce phenomena and processes to basic “atoms,” as the Cartesian View would have it or must we engage in science that looks at many levels of organization, and at the whole—at interactions among parts of a whole—as Lewontin (and Levins) preferred? The second dialectic is, in effect, a subset of the first—it is a zoomed in query about how we should imagine the anti-reductionist worldview: should organisms be given pride of place in explanatory and causal analyses of development and evolution, or should we be more ontologically democratic and distributed? The third dialectic can be seen as related to the second, but really gets to the technical heart of evolutionary theory: what is the nature of selection? Can selection operate above or below the standard owners of fitness—organismic individuals? It is, again, a testament to Lewontin’s brilliance that his work became central to discussions of both vertical and horizontal averaging, even if he fairly early on closed shop on the former.

Recently, as I copy edited my book Our Genes, the breadth and depth of Lewontin’s influence on my work became incredibly clear.31 I’ve been shaped by his critique of capitalism; his reflections on the role of competition-centric ideology in Darwinism, sociobiology, and evolutionary psychology; and the outstanding question of whether any ideology can ever speak from an absolute position.32 Lewontin’s corpus of technical contributions and breakthroughs is rich and diverse. It is also filled with “ambiguities and contradictions.” In fact, while these dialectics, which are so rich in future investigative possibilities, make it possible for us to construct a Lewontin to suit our purposes, they also resist any easy recreation.

Thank you, Dick. Rest in Power.

—

Acknowledgements 

Carlos López Beltrán, John Novembre, and Marie Raffn provided constructive feedback. Science for the People editors Edward Millar, Jake Ogata Bernstein, Claire Ramsay, and Calvin Wu helped make this a stronger piece. Lynn Chiu facilitated my initial contact with Science for the People. Molly Gage and Lucas McGranahan edited expertly. Needless to say, I will forever remain grateful to Dick Lewontin, and everything he represents.

About the Author

Rasmus Grønfeldt Winther is Professor of Humanities at the University of California, Santa Cruz. He works in the philosophy of science and philosophy of biology, and has interests in epistemology and political philosophy, Latin American history and philosophy, cartography and GIS, the oceans, and science in general. Recent publications include “A Beginner’s Guide to the New Population Genomics of Homo sapiens: Origins, Race, and Medicine” in The Harvard Review of Philosophy; “Mapping the Deep Blue Oceans” in The Philosophy of GIS (Springer); “¿El ombligo del mundo? Mapas y contramapas” in Revista de la Universidad de México; When Maps Become the World (University of Chicago Press, 2020); and Our Genes: A Philosophical Perspective on Human Evolutionary Genomics (Cambridge University Press, 2022).

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Notes

  1. Rasmus G. Winther, “Darwin on Variation and Heredity,” Journal of the History of Biology 33, no. 3 (Winter 2000): 425–55, https://www.jstor.org/stable/4331610.
  2. Richard Levins and Richard Lewontin, The Dialectical Biologist (Cambridge, MA: Harvard University Press, 1985), 279 (emphasis added).
  3. Levins and Lewontin, The Dialectical Biologist, 283.
  4. John Beatty, “Weighing the Risks: Stalemate in the Classical/Balance Controversy,” Journal of the History of Biology 20, no. 3 (Autumn 1987), 289–319, https://www.jstor.org/stable/4331020; Richard C. Lewontin, “Twenty-Five Years Ago in Genetics: Electrophoresis in the Development of Evolutionary Genetics: Milestone or Millstone?” Genetics 128, no. 4 (August 1991): 657–662, https://doi.org/10.1093/genetics/128.4.657; Rasmus Grønfeldt Winther, “Lewontin (1972),” in Remapping Race in a Global Context, ed. Ludovica Lorusso and Rasmus Grønfeldt Winther (London: Routledge, 2022), https://arxiv.org/abs/2110.10945; John Novembre, “The Background and Legacy of Lewontin’s Apportionment of Diversity” (unpublished manuscript, October 11, 2022), Microsoft Word file.
  5. Richard C. Lewontin and John L. Hubby, “A Molecular Approach to the Study of Genic Heterozygosity in Natural Populations. II. Amount of Variation and Degree of Heterozygosity in Natural Populations of Drosophila pseudoobscura,” Genetics 54, no. 2 (August 1966): 595–609, https://doi.org/10.1093/genetics/54.2.595. See also John L. Hubby and Richard C. Lewontin, “A Molecular Approach to the Study of Genic Heterozygosity in Natural Populations. I. The Number of Alleles at Different Loci in Drosophila pseudoobscura,” Genetics 54, no. 2 (August 1966): 577–594, https://doi.org/10.1093/genetics/54.2.577.
  6. Lewontin and Hubby, “A Molecular Approach to the Study of Genic Heterozygosity,” 595.
  7. See Brian Charlesworth and Deborah Charlesworth, “Population Genetics from 1966 to 2016.” Heredity 118 (2017): 2–9, https://doi.org/10.1038/hdy.2016.55; Winther, “Lewontin (1972)”; Richard C. Lewontin, “The Apportionment of Human Diversity,” in Evolutionary Biology 6, ed. T. Dobzhansky, M. K. Hecht, and W. C. Steere  (New York: Springer, 1972), 381–398. Lewontin’s “The Apportionment” lacks sufficient detail regarding page numbers and table contents from its primary data sources: see Arthur E. Mourant, The Distribution of the Human Blood Groups (Oxford: Oxford University Press, 1954) and Eloise R. Giblett, Genetic Markers in Human Blood (Oxford and Edinburgh: Blackwell, 1969). Nor did Lewontin explain the relevant population genetic theory in this piece. I have done both of these in “Lewontin (1972),” as well as  recalculate his diversity apportionments, finding errors for all genes except one (P). My calculations show Lewontin’s grand average diversity apportionments of “within populations,” “among populations but within ‘races’,” and “among ‘races’” to actually be, respectively, 86%/7%/7%.
  8. Richard C. Lewontin, The Genetic Basis of Evolutionary Change (New York: Columbia University Press, 1974), ix.
  9. Levins and Lewontin, The Dialectical Biologist, vii.
  10. Levins and Lewontin, The Dialectical Biologist, 288.
  11. Richard C. Lewontin, Human Diversity (New York: Scientific American Library, 1982); Richard C. Lewontin, Biology as Ideology: The Doctrine of DNA (New York: Harper Perennial, 1993); Richard C. Lewontin, The Triple Helix: Gene, Organism, and Environment (Cambridge, MA: Harvard University Press, 2002).
  12. Lewontin, Human Diversity, 26–27.
  13. Richard C. Lewontin, Human Diversity, 27–28.
  14. Levins and Lewontin, The Dialectical Biologist, 54–58.
  15. Levins and Lewontin, The Dialectical Biologist, 272.
  16. Levins and Lewontin, The Dialectical Biologist, 273.
  17. Levins and Lewontin, The Dialectical Biologist, 275.
  18. See also William C. Wimsatt, Re-Engineering Philosophy for Limited Beings: Piecewise Approximations to Reality (Cambridge, MA: Harvard University Press, 2007); Rasmus Grønfeldt Winther, “On the Dangers of Making Scientific Models Ontologically Independent: Taking Richard Levins’ Warnings Seriously,” Biology and Philosophy 21 (November 2016): 703–724, https://doi.org/10.1007/s10539-006-9053-7; Rasmus Grønfeldt Winther, “Part-Whole Science.” Synthese 178 (February 2011): 397–427, https://doi.org/10.1007/s11229-009-9647-0.
  19. Richard C. Lewontin, “The Units of Selection,” Annual Review of Ecology and Systematics 1 (1970): 1–18. https://www.jstor.org/stable/2096764.
  20. Lewontin, “The Units of Selection,” 1.
  21. Lewontin, “The Units of Selection,” 2.
  22. Lewontin, “The Units of Selection,” 15.
  23. Lewontin, “The Units of Selection,” 15-16.
  24. Rasmus Grønfeldt Winther, Michael J. Wade, and Christopher C. Dimond, “Pluralism in Evolutionary Controversies: Styles and Averaging Strategies in Hierarchical Selection Theories,” Biology & Philosophy 28, no. 6 (November 2013): 957–979, https://doi.org/10.1007/s10539-013-9378-y.
  25. I was fortunate to have studied the philosophy of biology under Godfrey-Smith and Lloyd for, respectively, my undergraduate and master’s thesis, and my PhD dissertation, which provided me with a fairly synoptic presentation of these philosophical issues: to simplify, Lloyd has always staunchly opposed vertical averaging, whereas Godfrey-Smith accepts many aspects of it, certainly implicitly. See also Peter Godfrey-Smith and Richard C. Lewontin, “The Dimension of Selection,” Philosophy of Science 60, no. 3 (September 1993): 373–395, https://www.jstor.org/stable/188081; Elisabeth A. Lloyd, Richard C. Lewontin, and Marcus W. Feldman, “The Generational Cycle of State Spaces and Adequate Genetical Representation,” Philosophy of Science 75, no. 2 (April 2008): 140–156, https://doi.org/10.1086/590196.
  26. Richard Dawkins, The Selfish Gene: 40th Anniversary Edition (Oxford: Oxford University Press, 2006), 38–39, 84–86.
  27. Richard C. Lewontin and Michael J. D. White, “Interaction Between Inversion Polymorphisms of Two Chromosome Pairs in the Grasshopper, Moraba scurra.” Evolution 14, no. 1 (March 1960): 116–129, https://doi.org/10.1111/j.1558-5646.1960.tb03061.x; R. C. Lewontin and Ken-ichi Kojima, “The Evolutionary Dynamics of Complex Polymorphisms.” Evolution 14, no. 4 (December 1960): 458–472, https://doi.org/10.1111/j.1558-5646.1960.tb03113.x; Ian Franklin and Richard C. Lewontin, “Is the Gene the Unit of Selection?” Genetics 65, no. 4 (August 1970): 707–734, https://doi.org/10.1093/genetics/65.4.707.
  28. Lewontin, The Genetic Basis of Evolutionary Change, 189.
  29. Lewontin, The Genetic Basis of Evolutionary Change, 271. See also Rasmus Grønfeldt Winther, When Maps Become the World (Chicago, IL: University of Chicago Press, 2020) for a discussion of how Lewontin imagined the mapping relation between genotype and phenotype.
  30. Lewontin, The Genetic Basis of Evolutionary Change, 318 (emphasis added).
  31. Rasmus Grønfeldt Winther, Our Genes: A Philosophical Perspective on Human Evolutionary Genomics (Cambridge, U.K.: Cambridge University Press, 2022).
  32. Furthermore, my book’s last chapter “The Conscious Universe: Genes in Complex Systems” is a kind of response to Lewontin’s technical critique of Analysis of Variance, which is “the most widely used and powerful statistical device in existence… [that] reifies… numerical components [of variance] as objective forces with actual physical interactions” (Levins and Lewontin, The Dialectical Biologist, 2). Even so, this chapter also endorses a dialectical, distributed, and interactive concept of causation.