Inlay

## Introduction Dialectical thinking in biology has famously been put forward by the eponymous book of Levins and Lewontin (1985). Much of the book was dedicated to the interpenetration between organisms and their environments. In particular, chapter 3, “The Organism as the Subject and Object of Evolution”, originally published by Lewontin (1983), explored how organisms simultaneously change and respond to their environments. This contrasted with two one-way causal models that commonly infuse biology: (1) on developmental timescales the phenotype is determined by the genotype, and (2) on evolutionary timescales the phenotype is determined by the environment, through the selection of genotypes. Levins and Lewontin’s take on the reciprocal causation between organisms and their environments contributed to the elaboration of dialectical accounts of development (e.g. Oyama, Gray, and Griffiths 2001; Sultan 2015) and of selection pressures (e.g. Odling-Smee, Laland, and Feldman 2003; Svensson 2018). However, one of Lewontin’s most cited and influential contributions is strikingly absent from the Dialectical Biologist. In “The Units of Selection”, Lewontin (1970) argues that Darwinian evolution by natural selection can in principle apply to any level of biological organisation, provided that heritable phenotypic variability cause differences in fitness. This was a major step towards the emergence of what is called today Multi-Level Selection (MLS) theory (e.g. Okasha 2006; Wilson and Sober 1989), according to which several levels of biological organization can simultaneously be the target of selection. ## Parts and wholes The first and second principles lie at the core of MLS theory, according to which properties of parts result, in part, from selection pressures exerted on the wholes they form. In the case of the evolution of altruism1 for instance, behaving altruistically is an individual property that is negatively selected at short time scales by individual-level selection inside each group, and positively selected at longer time scales during selection among groups. Selection among groups occurs because containing many altruists is a beneficent group-level property. MLS theory is at odds with the naive version of holism which asserted that individual traits evolved for the preservation of the species (“Arterhaltend”, Lorenz 1963), in line with the old concept of Balance of Nature. As Dawkins (1981) ironically put: “Should we then not expect lions to refrain from killing antelopes, ‘for the good of the mammals’ ?”. As indicated by its name, MLS theory considers that several levels of selection are acting simultaneously, with no predetermined outcome. In contrast to MLS theory, according to the “gene-eyes view” (Ågren 2021) championed by Dawkins (1976) and others, selection acts at the level of genes only. Altruistic alleles can be selected if they spread more efficiently indirectly through the bearer’s relatives (“kin selection”), following Hamilton’s rule (Hamilton 1964). According to the gene-eyes view, the group-level property “containing many altruists” is therefore explained by selection at the level of genes. Even the individual-level property “behaving altruistically” is explained by lower-level processes at the gene-level. ## Subjects and objects According to the third principle, subjects and objects are interchangeable. The usual theory of evolution by natural selection instead favours linear causation, where populations of organisms respond to external selection pressures dictated by the environment. Selection pressures are conceived as independent causes that remain unaltered during the process of adaptation. Standard evolutionary theory has therefore been qualified as “externalist” (Godfrey-Smith 1996; Laland et al. 2013). Dialectical accounts of biology like the Developmental Systems Theory and Niche Construction Theory (Odling-Smee, Laland, and Feldman 2003; Oyama, Gray, and Griffiths 2001), instead rely on the reciprocal causality between organisms and their environments. MLS theory also reach the conclusion that selective forces act both as subjects and objects. In the Descent of Man, Darwin famously provided the first sketch of what is called today group selection. He wrote: “It must not be forgotten that although a high standard of morality gives but a slight or no advantage to each individual man and his children over the other men of the same tribe, yet that an increase in the number of well-endowed men and an advancement in the standard of morality will certainly give an immense advantage to one tribe over another. A tribe including many members who, from possessing in a high degree the spirit of patriotism, fidelity, obedience, courage, and sympathy, were always ready to aid one another, and to sacrifice themselves for the common good, would be victorious over most other tribes; and this would be natural selection.” (Darwin 1871) ## Change The fourth principle claims that change must be sought in “all aspects of all systems”, that processes are ontologically prior to objects. The idea that natural selection is both a cause and an effect, as outlined in the previous section, reveals that natural selection itself is a historical product subject to change, especially in a MLS context where higher levels of selection modify previous selection regimes. ## Struggle of opposites Levins and Lewontin’s fifth principle stresses that processes are fueled by actions of opposing forces, paraphrasing Lenin (1925): “Development is the struggle of opposites.” and inspired by the law of the unity and conflict of opposites formulated by Engels (1873-1886). Although this has not been formulated explicitly by advocates of MLS theory, the struggle of opposites is one one its central aspects, as in the example provided by Lewontin (1970) and summarized by Williams (1966) as follows: “There is a series of alleles symbolized by the in house-mouse populations that produces a marked distortion of the segregation ratio of sperm. As much as 95 per cent of the sperm of a heterozygous male may bear such a gene, and only 5 per cent bear the wild-type allele. This marked selective advantage is opposed by other adverse effects in the homozygotes, either an embryonic lethality or male sterility. Such characters as lethality, sterility, and measurable segregation ratios furnish an excellent opportunity for calculating the effect of selection as a function of gene frequency in hypothetical populations. Such calculations, based on a deterministic model of selection, indicate that these alleles should have certain equilibrium frequencies in the populations in which they occur. Studies of wild populations, however, consistently give frequencies below the calculated values. Lewontin concludes that the deficiency must be ascribed to some force in opposition to genic selection, and that group selection is the likely force. He showed that by substituting a stochastic model of natural selection, so as to allow for a certain rate of fixation of one or another allele in family groups and small local populations, he could account for the observed low frequencies of the t-alleles.” ## Quantity and quality Although they don’t incorporate it in their “official” list of principles, Levins and Lewontin also mention on several occasions Engels’s law of the passage of quantitative changes into qualitative changes, it self derived from Hegel. Trotsky previously noted that the Darwinian theory of the origin of species is an instance of this law, where the continuous transformation of species can lead to the formation of a new species: “This brilliant biologist [i.e., Darwin], while showing how small quantitative deviations accumulate and yield a completely new biological ‘quality,’ in this way explaining the origin of species, applied without being conscious of it, the methods of dialectical materialism in the area of organic life. The Hegelian law of the transition from quantity into quality found in Darwin a brilliant, although philosophically unenlightened application” (Trotsky 1999) ## Conclusion I do not contend that the similarities between several dialectical aspects of social history and MLS theory outlined here correspond to deep ontological homologies. It would for instance make little sense to hold that the state of dual power in 1917 is precisely the same thing as dual fitness during an ETI. 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