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Richard Lewontin, dialectical materialism, the relationship between evolutionary biology and Marxism

Originally published: Science for the People by Erik Svensson (November 3, 2021 )  | - Posted Nov 04, 2021

On July 4, 2021, the great population geneticist and evolutionary biologist Richard Lewontin passed away at the age of 92. Lewontin left a strong political legacy in the United States in addition to his scientific one. He shared this political legacy with his fellow Marxist and comrade Richard Levins, who passed away five years ago and who was memorialized in an article titled “The People’s Scientist” in Jacobin in 2016.1 In 1985, the two co-authored a famous but also controversial book, The Dialectical Biologist. Published in the last phase of the Cold War and during the Reagan era, The Dialectical Biologist was a brave attempt to promote Marxism as a useful philosophy to biologists during a politically difficult time for those on the left. Levins and Lewontin forcefully argued that Marxism in general, and dialectical materialism in particular, could enrich evolutionary thinking and help us better understand the complexities of the natural world.

Dialectical materialism combines two philosophical traditions: historical materialism formulated by Karl Marx and Friedrich Engels, and dialectics, an idealist philosophy formulated by German philosopher Georg Willhelm Friedrich Hegel. Marx and Engels rejected Hegel’s idealism, but incorporated his dialectical philosophy into their historical materialist theory of how societies change. They argued that while human societies developed through configurations of productive forces and changes in economic and material conditions (including access to natural resources), materialism alone is not enough to understand human societies. Marx and Engels’s intervention was the addition of history, which reflects the internal contradictions within societies: class struggles between lords and peasants under feudalism and between workers and capitalists under capitalism. Whereas historical materialism reflects that all human intellectual activities and achievements ultimately have a material basis and that history does not unfold according to some unknown and mysterious spiritual force or pre-determined plan, dialectics complements this materialist insight and emphasizes the inner contradictions of all material systems ultimately needed for history to take place at all. In other words, although there might be general rules and natural laws shaping the history of both humans and other parts of the natural world, inherent contradictions in all material systems generate historical contingencies. This idea is something that most evolutionary biologists are familiar with, and the arcs of neither history nor biology are therefore inevitable. Both dialectics and materialism were equally important in the philosophy of Marx and Engels, and this philosophy had an influence on later followers, including Levins and Lewontin.

Dialectical materialism was therefore an attempt to synthesize and combine two philosophies with different origins. The reason Marx and Engels gave for this integration of these two traditions is that dialectics without materialism runs the risk of becoming naïve idealism, whereas materialism without dialectics runs the risk of becoming naïve reductionism. Levins and Lewontin contrasted their dialectical materialist perspective to the two popular but idealistic currents among bourgeois intellectuals: vitalism and reductionism.2 A large part of Levin’s and Lewontin’s inspiration came from Engels’s classic text Dialectics of Nature, which stimulated their attempt to reintroduce dialectical materialism into Western science.

In Dialectics of Nature, Engels put forward some rather speculative ideas that, although not always upheld by later research, revealed a brilliant and creative approach and showed how dialectics could provide a fruitful way of thinking about the evolutionary process. Among Engels’s more radical contributions was to suggest that the human hand and bipedalism had coevolved with the evolution of a large brain. Engels argued that these two traits were mutually reinforcing during evolution: as the brain became more capable of solving difficult problems, the hand was enabled to take on more tasks, which resulted in a selective feedback loop that drove the evolution of an even more advanced cognitive ability of the brain, and so forth.

Here we see the intellectual precursor of an idea that Lewontin and Levins developed further in The Dialectical Biologist: that different traits are not independent of each other, but rather quasi-independent and many times co-selected for a common function. Organisms are thus not the mere collection of a series of independent “traits,” and the whole (the organism) is clearly more than the sum of its parts (the organism’s traits). To quote Levins and Lewontin: “Every part or activity of an organism acts as environment for other parts” (p. 58), which means that an important part of the selective environment of every trait in an organism are other traits of that organism, as well as the need for traits to function together with each other. Furthermore, Levins and Lewontin emphasize that “…interdependence permits survival when the parts function well, but in pathological conditions, produces pervasive disaster” (p. 58) and that “… different organs or processes that have little direct interaction may be bound together ecologically by their common adaptive significance” (p. 60), underscoring that organisms are co-adapted systems, where the sum is more than the parts and traits are co-selected to improve organismal survival and reproduction in challenging natural environments.

The idea that traits function and evolve together, rather than being selected on in isolation, is an important insight promoted by Levins and Lewontin and supported by many evolutionary biologists today. One particularly relevant area of research has been to understand the genetic, genomic, and evolutionary consequences of correlational selection, that is, selection for certain character combinations over others.3 Such correlational selection can promote the evolution of genetic and developmental integration between different traits, even if they have different genetic and developmental origins, and thereby provides a link between ecology, evolution, genomics, and molecular biology.4 Thus, traits can become linked through selection for a common ecological function, even if they are governed by different sets of genes, provided that certain trait and gene combinations are more favorable to organismal survival than others. In short: organisms are more than the sum of their parts (traits) but represent successful co-adapted combinations of traits that are co-selected.

Science and Scientists under Capitalism

Scientists in general, including many evolutionary biologists, typically view themselves as apolitical and cold objective observers of the natural world. Many scientists tend to shy away from anything that risks undermining that illusory self-image. This is quite evident in Western capitalist countries, where the organization of modern science largely reflects the dominant bourgeois ideology, with its emphasis of state and market interests and the need for immediate and commercially useful results, and where politicians typically promote an atmosphere of uncritical celebration of competition among scientists as the only way to distribute grants and research resources. Levins and Lewontin, on the other hand, took an opposite stance: they argued that it was an advantage to be explicit about their philosophical and political views, rather than hiding under false objectivity, like many Western bourgeois intellectuals tend to do.

While science as a general human knowledge-producing enterprise clearly can reveal objective facts about our natural world, individual scientists are seldom (if ever) entirely objective and impartial (and should not even pretend that they are). In chapter 8 of The Dialectical Biologist, The Commoditization of Science,” Levins and Lewontin explored how modern science is both constrained by, and grew out of capitalism, and they noted several tendencies that have become amplified ever since and that are still operating in academia today: elitism, commoditization, separation of thinking from feeling, and increasing demands that scientific discoveries be quantifiable and ideally commercialized. To quote the first sentence of chapter 8 of the book, “modern science is a product of capitalism.”5 Levins and Lewontin further argued that the traditional view of science in the West is built on a strict Cartesian and reductionist worldview where nature and organisms are portrayed as well-functioning clock-like machines that can only be understood by breaking down the organisms into their constituent parts. A dialectical perspective, in contrast to such Cartesian reductionism, views organisms as more than the sum of their parts and emphasizes that traits typically function together, rather than in isolation. In short: organisms are not Cartesians machines, but products of a long evolutionary history, where their traits are quasi-independent and often subject to co-selection.

A good example of the failure of the Cartesian reductionist approach is the Human Genome Project (HGP), launched with the promise that once we knew the entire genetic code of all humans, we would be able to understand all human diseases and develop so-called “personalized medicine” based on the genetic makeup of every individual. More than three decades after the launching of HGP, we have only been able to explain a minor fraction, often only 5 to 10 percent of the variation in some of many important human diseases. The failure is most likely due to interactions among genes and between genes and environmental factors ignored by Genome Wide Association Studies (GWAS) and the search for single genes of major effect.6 These insights from the last decades of human genomic research are broadly consistent with the issues that Levins and Lewontin raised in their still highly relevant critique of naïve reductionism in molecular biology.

Lewontin and Evolutionary Biology

The Cartesian reductionist worldview criticized by Levins and Lewontin can still be seen in contemporary biology, including the research approach from some British evolutionary biologists like Alan Grafen and Andy Gardner, and particularly in Grafen’s “Formal Darwinism-project.”7 These biologists, alongside many other contemporary behavioral ecologists, are mainly interested in organismal adaptations, and not so much in the evolutionary processes and history behind such adaptations. Neither are these British evolutionary biologists very interested in developmental, genetic, or other constraints on adaptations. Their research approach is largely built on the intellectual foundations of British population geneticist Ronald Fisher, developed in his 1930 book The Genetical Theory of Natural Selection.8 The Fisherian worldview is one in which natural selection is an omnipresent force that operates on populations of very large sizes and where selection is thus largely unopposed by random factors and stochasticity. As a consequence of this, organisms are expected to be predominantly shaped by the optimizing force of natural selection, and few maladaptive features or constraints are expected to interfere with a “perfect” organismal adaptive design.

The science philosopher Tim Lewens has labelled this research approach as the “Neo-Paleyan”9 evolutionary biology tradition in Britain, where selection is an omnipresent force that refines the tiniest details of organisms, making them perfectly adapted to their environments. The term Neo-Paleyan refers to the intellectual legacy of Pastor William Paley, a natural theologian living in eighteenth century England who saw design everywhere in nature and took such design as evidence for the existence of the Designer, an almighty God. Paley’s most famous analogy of design in nature was that if one would find a functioning clock on a beach, there must be a designer who has produced this clock. In the narrow intellectual research tradition of Neo-Paleyan evolutionary biologists, there is little room for historical legacies, evolutionary constraints, and maladaptive features of organisms, because natural selection is the only evolutionary force that matters in nature.

An extreme example of the Neo-Paleyan research tradition is illustrated in the views of another British evolutionary biologist and popular science writer Richard Dawkins. Dawkins, unlike Paley, is an atheist, but he has declared his great admiration of Paley in his popular science book The Blind Watchmaker. In this well-written book, Dawkins seems to almost replace God with Darwin’s principle of Natural Selection, and repeatedly emphasizes the perfection of organisms, which are viewed as almost “designed” by natural selection. Another famous book by Dawkins, The Selfish Gene (1976), further illustrates the Cartesian reductionist perspective and the Neo-Paleyan view of organisms as well-functioning machines. In this book and others, Dawkins reduces organisms to temporary survival machines of their genes. Organisms are not interesting in their own right, except as temporary vehicles for their genes, called replicators, which are the only important evolutionary units.10 In this ultra-reductionist view, the genetic level is the only level that matters in evolution and is sufficient to explain phenotypic change at the level of the organism.

The evolutionary biology tradition represented by Lewontin and some other North American biologists is very different from the ultra-adaptationist and ultra-Darwinist intellectual tradition represented by the Neo-Paleyan biologists in the behavioral ecology tradition in Great Britain. In North America, by contrast, a different research tradition developed in population genetics that was based more on the ideas of Sewall Wright, who emphasized not only natural selection, but also the role of randomness and stochasticity in evolution, particularly the role of genetic drift.11 Lewontin was largely a product of this Wrightian tradition in population genetics, where the interaction between natural selection, genetic drift, and stochasticity was the main focus, in contrast to the British Fisherian and Neo-Paylean tradition’s focus on natural selection alone.

Lewontin has also repeatedly emphasized that processes and interactions between different parts in nature are as important as the parts themselves. To understand complex systems such as organisms, populations, or ecosystems, it is not enough to divide up the systems into their component parts (e.g. genes or environments). One must also analyze the interactions between the parts (e.g. how genes and environments interact). Specifically, it is Lewontin’s view that in order to understand organisms, we must recognize that genes do not act in isolation, but are expressed in different environmental contexts, and that one aspect of the environment is actually other genes in the organism. The technical terms for these phenomena are gene-by-environment interaction and epistasis. These phenomena are seldom acknowledged by the Neo-Paleyans, who are mainly interested in global adaptive solutions, rather than in geographical variation, historical contingencies, or how different populations and species have evolved different adaptive “solutions” to similar ecological “problems.”

In Lewontin’s perspective, environmental context and history matters, and the organism plays a central role in evolution. Lewontin repeatedly emphasized that the fitness of organisms and the selective advantage of gene variants (alleles) was unlikely to be constant, but changed over time as environmental and genetic conditions changed. For instance, a well known evolutionary process is frequency-dependent selection, in which the fitness of a gene variant depends on its frequency in the population.12 A gene variant could be associated with high fitness when it is rare, and frequency-dependent selection would maintain genetic variation in the population. Conversely, a gene variant could give its bearers high fitness when it is common; then it would spread through the population and replace other existing genetic variants. Frequency-dependent selection is important because the fitness of the organism is not solely a result of its own genotype, but an emergent property of the interaction between the genotype and the genetic environment it finds itself in, i.e. the frequency of its own genotype in the population. Frequency-dependent selection therefore illustrates the dialectical principle that the whole is more than the sum of the parts; that genotype and environment are both needed to correctly infer fitness; that it is impossible and meaningless to assign a constant fitness value to a particular genotype. Fitness is highly context-dependent upon the local gene-frequency environment.

Another important and related point emphasized repeatedly by Lewontin is organism-environment interaction, which was beautifully illustrated in chapter 3 of The Dialectical Biologist, “The Organism as the Subject and Object of Evolution.” Here, Levin and Lewontin applied dialectical thinking and their Marxist philosophical perspective more explicitly than anywhere else in the book. The main point is that organisms are not simply passive subjects to external evolutionary forces like natural selection, but organisms are also active subjects in their own right. Specifically, organisms do not simply enter empty ecological niches, but they actively construct their niches, thereby influencing the selection pressures operating on themselves. As an example, consider thermoregulation in some ectothermic (“cold-blooded”) organisms like lizards, where the body temperature is largely determined by external ambient temperature.13 One might envision lizards as passive objects of their external environmental circumstances, freezing to death during cold spells and dying of overheating during hot spells. This is not what actually happens in nature. Instead, lizards actively seek out warmer environmental patches under cold weather conditions, and cooler spots during hot weather conditions. The lizards therefore actively counteract the selection pressures operating on themselves, by maintaining homeostasis and keeping favorable body temperatures. This is an excellent example of the central role of organisms in the evolutionary process that Lewontin highlighted. Chapter 3 gave rise to the term “niche construction” that has germinated lively debates in the evolutionary biology research community about the intricate relationships between organisms, their environment, and evolution.14 More generally, Levins and Lewontin emphasized reciprocal causation in evolution, involving selective feedbacks where a cause would later become an effect (and vice versa) in chains of causation,15 exemplified by reciprocal interactions between, for example, predators and prey in the process of co-evolution.16

Evolutionary Biology and Marxism: Natural Bedfellows?

A dialectical perspective fits very well with a worldview where nature is seen not as a static phenomenon but rather as a series of dynamic equilibria that change as the balance of various counteracting forces change. Lewontin was not alone; he was not the only great evolutionary biologist who openly identified as a Marxist. As a matter of fact, the fields of evolutionary biology and population genetics have had many high-profile, openly Marxist scientists, at least during parts of their life. Among these are population geneticists J. B. S. Haldane and John Maynard Smith (Haldane’s student), paleontologist Stephen Jay Gould, and science philosopher Elliott Sober. It is interesting, and maybe not a coincidence, that these great scientists were all interested in the balance of forces, either in population genetics (Haldane, Maynard Smith, Lewontin, Sober), at the macroevolutionary level (Gould) or in ecology (Levins).

John Maynard Smith, for instance, was a committed Marxist and member of the Communist Party of Great Britain, as was his mentor Haldane, although both of them later left the party. Smith was also sympathetic to The Dialectical Biologist and reviewed it favourably, and he and Lewontin remained close friends and colleagues during their entire careers.17 Perhaps population and evolutionary geneticists, with their main focus and research interests in the evolutionary process rather than evolutionary outcomes such as adaptations, are more prone than other biologists, such as behavioral ecologists, to incorporate dialectical thinking in their research programs. This interpretation is strengthened by the fact that philosopher Elliott Sober—known for his classic book The Nature of Selection-Evolutionary Theory in Philosophical Focus (1994)who spent some time in Lewontin’s laboratory, has openly identified as a Marxist and defended Marxism in 1994, after the collapse of the Berlin Wall and the East Bloc.18 Sober’s work explores how different forces such as mutation, selection, recombination, and gene flow interact in the evolutionary process. It remains an excellent overview of the philosophical implications of population genetics, and one can also see signs of dialectical thinking, despite this term not being mentioned explicitly in this classic text. Natural selection is often opposed by other evolutionary forces such as mutations and genetic drift, a common focus of much population genetics research. This dynamic view of the evolutionary process is seen very little in Dawkins’s static adaptationism and among Neo-Paleyan evolutionary biologists in Great Britain, but for Marxists, it is a natural perspective on both society and nature.

More recent examples of ecologists and evolutionary biologists who implicitly or explicitly use dialectical thinking in their research include theoretical biologist Sergey Gavrilets and former population ecologist and current quantitative historian Peter Turchin, who were both born in Russia and exposed to Marxist philosophy growing up in the Soviet Union. Gavrilets has taken an explicitly dialectical approach when modelling the evolutionary transition from a hierarchical social system in our close relatives, the great apes, to the more egalitarian hunter-gatherer societies in our own species, Homo sapiens.19 In Gavrilets’s models, human egalitarianism and social norms arise as emergent results from within-group conflicts, where subordinates form alliances to prevent dominant “bullies” from monopolizing resources. To Gavrilets, “the same forces that shape the emergence of highly despotic groups dialectically create conditions for the evolution of counterdominant coalitionary behavior and psychology.”20 Moreover, the evolutionary transition to egalitarianism could have been very fast and proceeded in the form of a rapid phase transition driven by a positive feedback loop, where larger and cooperative alliances grow even larger, resulting in an egalitarian revolution.21

Such rapid phase transitions driven by positive feedback loops were discussed by Engels in Dialectics of Nature, and later in The Dialectical Biologist as exemplifying the dialectical principle of “the transformation of quantity into quality.” The most well known example of this is water moving from a liquid state to a gas state when the boiling temperature is reached. Similar ideas about how slow, gradual change can cross certain critical thresholds above which rapid major change is triggered have been put forward in several areas of modern evolutionary biology, including the ideas of “punctuated equilibrium” in paleontology and “tipping points” in speciation research.22

The “Lysenko Affair” and the Politicization of Science

Richard Lewontin’s Marxism was not aimed at pushing political dogmas, but rather at a critical analysis of science, aiming to improve the state of affairs and pointing to hidden non-scientific and ideological assumptions that practicing scientists were unaware of. In his critique of science under capitalism, Lewontin certainly did not deny the many successes of the Cartesian reductionist approach, and in The Dialectical Biologist, even describes evolutionary game theory as a spectacular success of bourgeois economics. In a recent obituary for Lewontin, Andrew Berry and Dmitri Petrov stated that his Marxism was arguably a key component of his scientific success” and that “he was dogmatically non dogmatic.”23

Nevertheless, liberal technocrats and bourgeois critics of Marxism often claim that science should be kept entirely free from politics and ideology, because otherwise the consequences would be disastrous. As a case in point, they often point to the “Lysenko Affair” in the Stalin-era Soviet Union. Readers are probably already aware of the horrific state terror directed toward Mendelian geneticists under Stalin’s regime, which promoted the “barefoot” scientist Trofim Lysenko to a high position. Lysenko denounced Mendelian genetics as “undialectical” and idealistic, instead promoting a Lamarckian hereditary theory which proposed that plants could acquire traits during their life-time, and then transmit these traits to their offspring. Lysenko was not, however, a rigorous scientist, and his experiments failed to support his claim that acquired characteristics could be inherited. Lysenkoism clearly had a negative effect on the development of agricultural science and genetics in the Soviet Union, and many Marxists and communists in the West lost their illusions about the Soviet Union, communism, and Marxism as a result of these tragic events.

In chapter 7 of The Dialectical Biologist, Levins and Lewontin critically dissected Lysenko’s vulgarization of Marxism and his use of superficial dialectical arguments. They concluded that Lamarckism cannot be considered a refuted hereditarian theory or a valid alternative to Mendelian inheritance, as it was completely abandoned as a plausible hypothesis following the development of modern genetics. For this reason, they described the Lysenko movement as a failed attempt at a scientific revolution, but which reflected some real tensions and class-based conflicts in Soviet society neglected by the Western liberals who characterized it as a movement created from above by the Communist Party, Stalin, and other political leaders. Thus, Levins and Lewontin did not only criticize Lysenko and his followers; they also criticized the simplistic views within and outside of the Soviet Union who tried to explain away Lysenkoism by a broad stroke reduction to mere authoritarianism. Levins and Lewontin argued that Lysenkoism was a genuine populist movement, albeit one built on erroneous assumptions about how heredity and biology work. Lysenkoism, then, should not be understood as a natural outcome of Marxist science, but rather as a vulgarization of Marxism. At the same time, they issued an important warning that “unless Marxism examines its failures, they will be repeated.”24

Our Dialectical Future: Ecosocialism or Climate Breakdown

I did not know Lewontin personally, although I briefly interacted with him in Tennessee in 2001, during a meeting arranged by the Society for the Study of Evolution (SSE). Lewontin’s work and ideas were always inspiring to me and many of my colleagues in evolutionary biology. Growing up in Sweden, a small country strongly influenced by the British behavioral ecology research tradition, with its strong emphasis solely on organismal adaptation, it took me a while to discover Lewontin’s work. Before this discovery, I struggled hard to reconcile my socialist political sympathies and my Marxist identity with my professional research identity as an evolutionary biologist. I tried to keep these identities apart—not always successfully—because identities in separation were, and still are, the ideals in the British behavioral ecology and evolutionary biology tradition that has passed down to my generation. When I discovered Lewontin, I realized that being a Marxist could actually be an advantage for a scientist. Marxists see problems differently from liberal intellectuals, who often have great difficulties appreciating other perspectives than the strict Cartesian and reductionist view of organisms as machines.

As Levins and Lewontin pointed out in The Dialectical Biologist, a dialectical perspective is not the only way to resolve contradictions in science, writing that Dialectical materialism is not, and has never been, a programmatic method for solving particular physical problems. Rather, dialectical analysis provides us with an overview and a set of warning signs against particular forms of dogmatism and narrowness of thought.”25 These points are more valid than ever before, and reveal undogmatic and dynamic mindsets of both Levins and Lewontin and the Marxism they represented.

Many of the ideas put forward by Lewontin under the umbrella of Marxism and dialectical materialism several decades ago have, once again, become highly relevant today in light of the unfolding climate crisis, and with all the societal feedback associated with it. Indeed, many evolutionary biologists do already unconsciously adopt a dialectical perspective in their research. By viewing natural systems as results of various counteracting forces that keep each other more or less in balance, we can better understand the dynamics of evolutionary change than we could if we viewed the world as composed of a number of static objects. One theorist working in Lewontin’s dialectical tradition is a Swedish environmental historian, climate activist, and ecosocialist: Andreas Malm. Malm has made many aware of the positive feedback driving climatic systems and the dialectical relationships between fossil capital, nature, and society.26 Such dialectical feedback loops are crucial knowledge in our times of rapid climate change that threatens the long-term survival of humanity. Humans are perhaps the best example of an organism that can construct (and destroy!) its own niche, emphasizing Lewontin’s perspective that organisms are not passive objects of natural external forces, but actively construct their niches and the selection pressures on themselves. Indeed, humans have been called the world’s greatest evolutionary force, and human activities strongly affect the evolution of other organisms with profound consequences for our own societies, and by extension our future survival prospects.27

In light of these bleak prospects for humanity, Malm, myself, and other ecosocialists therefore argue that the only way to prevent the forthcoming disaster is through a social revolution and a large-scale socialist transformation of society. Dialectical materialism reminds us that history is contingent and influenced by internal conditions and contradictions within societies. This philosophy can help us recognize humans as active agents capable of shaping our future, and realize that fossil capital must be abolished, and that workers and citizens must take control and socialize finance and the means of production.28 Ultimately, we should develop a democratically and centrally planned economy that is organized along scientific principles and in an environmentally sustainable way. The alternative to such a socialist transformation is horrific so even non-Marxists will soon realize that social revolution is our only remaining chance as a species to survive in the long run. Otherwise, we risk seeing our civilization collapse under the weight of accelerating climate breakdown. The options could be illustrated, to put it rather bluntly, by reformulating a famous quote from the socialist revolutionary Rosa Luxemburg (“Socialism or barbarism!”) to emphasize the stark choice we now face in our own time: “Socialism or extinction!”29


Erik Svensson is Professor of Evolutionary Ecology at the Department of Biology, Lund University, Sweden.

Notes:

  1. Pankaj Mehta, “The People’s Scientist,” Jacobin, January 22, 2016, www.jacobinmag.com.
  2. Brett Clark and Richard York, “Dialectical Nature: Reflections in Honor of the Twentieth Anniversary of Levins and Lewontin’s The Dialectical Biologist,” Monthly Review 57, no. 1 (May 2005), monthlyreview.org.
  3. Barry Sinervo and Erik Svensson, “Correlational Selection and the Evolution of Genomic Architecture,” Heredity 89 (2002): 948–955, doi.org; Erik Svensson et al., “Correlational Selection in the Age of Genomics,” Nature Ecology & Evolution 5, no. 5 (2021): 562–573, doi.org.
  4. Svensson et al., “Correlational Selection,” 562.
  5. Levins and Lewontin, The Dialectical Biologist, 197.
  6. Evan E. Eichler et al., “Missing Heritability and Strategies for Finding the Underlying Causes of Complex Disease,” Nature Reviews. Genetics 11, no. 6 (June 2010): 446–50, doi.org; Teri A. Manolio et al., “Finding the Missing Heritability of Complex Diseases,” Nature 461, no. 7265 (October 8, 2009): 747–53, doi.org.
  7. Alan Grafen, “The Formal Darwinism Project in Outline,” Biology & Philosophy 29 (March 2014): 155–174, doi.org.
  8. Ronald A. Fisher, The Genetical Theory of Natural Selection (Clarendon Press, 1930).
  9. Tim Lewens, “Neo-Paleyan biology,” Studies in History and Philosophy of Science Part C 76 (August 2019): 101185, doi.org.
  10. Richard Dawkins, “Replicators and Vehicles,” PhilPapers, accessed October 20, 2021, philpapers.org.
  11. William B. Provine, Sewall Wright and Evolutionary Biology (University of Chicago Press, 1986); Ernst Mayr and William B. Provine, The Evolutionary Synthesis: Perspectives on the Unification of Biology (Harvard University Press, 1998).
  12. Richard Lewontin, “A General Method for Investigating the Equilibrium of Gene Frequency in a Population,” Genetics 43, no 3 (May 1958): 419–434, doi.org.
  13. Raymond B. Huey, Paul E. Hertz, and B. Sinervo, “Behavioral Drive versus Behavioral Inertia in Evolution: A Null Model Approach,” The American Naturalist 161, no. 3 (March 2003): 357–366, doi.org.
  14. F. John Odling-Smee, Kevin N. Laland, and Marcus W. Feldman, Niche Construction: The Neglected Process in Evolution (New Jersey: Princeton University Press, 2003); Thomas C. Scott-Phillips et al., “The Niche Construction Perspective: A Critical Appraisal,” Evolution 68, no. 5 (May 2014): 1231–43, doi.org.
  15. Erik I. Svensson, “On Reciprocal Causation in the Evolutionary Process,” Evolutionary Biology 45 (2018): 1–14, doi.org.
  16. Svensson, “On Reciprocal Causation in the Evolutionary Process.”
  17. John Maynard Smith, “Molecules Are Not Enough,” in Did Darwin Get It Right? Essays on Games, Sex and Evolution, ed. John Maynard Smith (Springer-Verlag, 1988): 30–38, doi.org; John Maynard Smith, “Reconciling Marx and Darwin,” Evolution 55, no. 7 (July 2001): 1496–1498, www.jstor.org.
  18. Erik Olin Wright, Andrew Levine, and Elliott Sober, “Reconstructing Marxism: A Reply,” Science & Society 58, no. 1 (Spring, 1994): 53–60, www.jstor.org; Elliott Sober, The Nature of Selection: Evolutionary Theory in Philosophical Focus (University of Chicago Press, 1984).
  19. Sergey Gavrilets, Edgar A. Duenez-Guzman, and Michael D. Vose, “Dynamics of Alliance Formation and the Egalitarian Revolution,” PLoS ONE 3, no. 10 (October 2008): e3293, doi.org; Sergey Gavrilets, “On the Evolutionary Origins of the Egalitarian Syndrome,” Proceedings of the National Academy of Sciences 109, no. 35 (August 2012): 14069–74, doi.org.
  20. Gavrilets, “On the Evolutionary Origins of the Egalitarian Syndrome.”
  21. Gavrilets, Duenez-Guzman, and Vose, “Dynamics of Alliance Formation”.
  22. David Sepkoski, Rereading the Fossil Record: The Growth of Paleobiology as an Evolutionary Discipline (University of Chicago Press, 2012); Patrik Nosil et al., “Tipping Points in the Dynamics of Speciation,” Nature Ecology & Evolution 1 (Janurary 2017): 1–8, doi.org.
  23. Andrew Berry and Dmitri A. Petrov, “Richard C. Lewontin (1929–2021),” Science 373, no. 6556 (August 2021): 745, doi.org.
  24. Levins and Lewontin, The Dialectical Biologist, 166.
  25. Lewontin and Levins, The Dialectical Biologist, 191.
  26. Andreas Malm, “The Anthropocene Myth,” Jacobin, May 30, 2015, jacobinmag.com.
  27. Stephen R. Palumbi, “Humans as the World’s Greatest Evolutionary Force,” Science 293, no. 5536 (September 2001): 1786–90, doi.org; Andrew P. Hendry, Kiyoko M. Gotanda, and Erik I. Svensson, “Human Influences on Evolution, and the Ecological and Societal Consequences,” Philosophical Transactions of the Royal Society B 372, no. 1712 (January 2017): 10160028, doi.org.
  28. Andreas Malm, “The Climate Movement Must Disrupt the Normal Routines of Fossil Capital,” Jacobin, October 14, 2020, jacobinmag.com.
  29. Mark Montegriffo, “Yes, Socialism or Extinction is Exactly the Choice We Face,” Jacobin, April 9, 2020, jacobinmag.com.
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