Organisms can be treated as optimizers when there is consensus among their genes about what is best to be done, but genomic consensus is often lacking, especially in interactions among kin because kin share some genes but not others. Grafen adopts a majoritarian perspective in which an individual’s interests are identified with the interests of the largest coreplicon of its genome, but genomic imprinting and recombination factionalize the genome so that no faction may predominate in some interactions among kin. Once (...) intragenomic conflicts are recognized, the individual organism can be conceptualized as an arbiter among competing interests within a collective. Organismal adaptation can be recognized without phenotypes being optimized. (shrink)

This paper uses the framework of Formal Darwinism (FD) to evaluate organism-centric critiques of the Modern Synthesis (MS). The first section argues that the FD project reconciles two kinds of selective explanations in biology. Thus it is not correct to say that the MS neglects organisms—instead, it explains organisms’ design, as argued in the second section. In the third section I employ a concept of the organism derived from Kant that has two aspects: the parts presupposing the whole, and the (...) productivity of the parts in relation to the whole. The first aspect corresponds to the “design” that FD explains, whereas the second aspect is something about which the MS is largely silent. (shrink)

Individual-as-maximizing agent analogies result in a simple understanding of the functioning of the biological world. Identifying the conditions under which individuals can be regarded as fitness maximizing agents is thus of considerable interest to biologists. Here, we compare different concepts of fitness maximization, and discuss within a single framework the relationship between Hamilton’s (J Theor Biol 7:1–16, 1964) model of social interactions, Grafen’s (J Evol Biol 20:1243–1254, 2007a) formal Darwinism project, and the idea of evolutionary stable strategies. We distinguish cases (...) where phenotypic effects are additive separable or not, the latter not being covered by Grafen’s analysis. In both cases it is possible to define a maximand, in the form of an objective function ϕ(z), whose argument is the phenotype of an individual and whose derivative is proportional to Hamilton’s inclusive fitness effect. However, this maximand can be identified with the expression for fecundity or fitness only in the case of additive separable phenotypic effects, making individual-as-maximizing agent analogies unattractive (although formally correct) under general situations of social interactions. We also feel that there is an inconsistency in Grafen’s characterization of the solution of his maximization program by use of inclusive fitness arguments. His results are in conflict with those on evolutionary stable strategies obtained by applying inclusive fitness theory, and can be repaired only by changing the definition of the problem. (shrink)

I argue that Grafen’s formal darwinism project could profitably incorporate a gene’s-eye view, as informed by the major transitions framework. In this, instead of the individual being assumed to maximise its inclusive fitness, genes are assumed to maximise their inclusive fitness. Maximisation of fitness at the individual level is not a straightforward concept because the major transitions framework shows that there are several kinds of biological individual. In addition, individuals have a definable fitness, exhibit individual-level adaptations and arise in a (...) major transition, only to the extent that the inclusive-fitness interests of genes within them coincide. Therefore, as others have suggested, the fundamental level at which fitness is maximised is the gene level. Previous reconciliations of the concepts of gene-level fitness and individual-level fitness implicitly recognise this point. Adaptations always maximise the fitness of their causative genes, but may be simple or complex. Simple adaptations may be controlled by single genes and be maladaptive at higher levels, whereas complex adaptations are controlled by multiple genes and rely on those genes having coinciding fitness interests at a higher level, for a given trait. (shrink)

This paper is a commentary on the focal article by Grafen and on earlier papers of his on which many of the results of this focal paper depend. Thus it is in effect a commentary on the “formal Darwinian project”, the focus of this sequence of papers. Several problems with this sequence are raised and discussed. The first of these concerns fitness maximization. It is often claimed in these papers that natural selection leads to a maximization of fitness and that (...) this view is claimed in Fisher’s “fundamental theorem of natural selection”. These claims are refuted, and various incorrect statements about the meaning and interpretation of the fundamental theorem of natural selection, in this sequence and in other papers by other authors, are discussed. Next, much of the work in this sequence rests on the first Price equation. In the deterministic (infinite population) case this equation is no more than the standard classical equation relating to changes in gene frequencies. In the stochastic case the equation gives the change in gene frequencies as the sum of two terms (the second of which vanishes in the deterministic case). These two terms are of essentially equal importance in the situation considered in the focal article, yet one of Grafen’s results ignores the second term in the stochastic analysis. This is associated with a wavering between deterministic and stochastic analyses and the use of the Price fitness concept and the classical fitness concept. These comments cast doubts on Grafen’s optimization theory. (shrink)

The Formal Darwinism project probes the connections between the dynamics of natural selection and the design of organisms. Here, I explain why this work should be of interest to philosophers, arguing that it is the natural development in a long-running scholarly enquiry into the meaning of life. I then review some of my own work which has applied the tools of Formal Darwinism to address issues concerning the units of adaptation in social evolution, leading to a deeper understanding of the (...) adaptation of individual organisms. Finally, I sketch some directions Formal Darwinism to explore beyond the biological sciences, with a focus upon cosmology. (shrink)

Two questions are raised for Grafen’s formal darwinism project of aligning evolutionary dynamics under natural selection with the optimization of phenotypes for individuals of a population. The first question concerns mean fitness maximization during frequency-dependent selection; in such selection regimes, not only is mean fitness typically not maximized but it is implausible that any parameter closely related to fitness is being maximized. The second question concerns whether natural selection on inclusive fitness differences can be regarded as individual selection or whether (...) it leads to a departure from the central motivation that led to the formal darwinism project, viz., to show that “Darwinian” evolution through individual selection leads to “good design” or phenotypic adaptation through trait optimization. (shrink)

The broader context for the formal darwinism project established by two of the commentators, in terms of reconciling the Modern Synthesis with Darwinian arguments over design and in terms of links to other types of selection and design, is discussed and welcomed. Some overselling of the project is admitted, in particular of whether it claims to consider all organic design. One important fundamental question raised in two commentaries is flagged but not answered of whether design is rightly represented by an (...) optimisation program, and another from one commentary of whether the coreplicon dissolves in the face of multi-generational imprinting. Calls for the project to be extended to design at levels above and below the individual are considered sympathetically, but judged impractical at the high level of abstraction of the project. All claims of substantive technical error are emphatically rejected. Close technical readings are welcomed that, among other things, represent the project as 'axiomatizing fitness'. The prospects for the project are set out in the light of this highly varied set of commentaries. (shrink)

We consider the question: under what circumstances can the concept of adaptation be applied to groups, rather than individuals? Gardner and Grafen (2009, J. Evol. Biol.22: 659–671) develop a novel approach to this question, building on Grafen's ‘formal Darwinism’ project, which defines adaptation in terms of links between evolutionary dynamics and optimization. They conclude that only clonal groups, and to a lesser extent groups in which reproductive competition is repressed, can be considered as adaptive units. We re-examine the conditions under (...) which the selection–optimization links hold at the group level. We focus on an important distinction between two ways of understanding the links, which have different implications regarding group adaptationism. We show how the formal Darwinism approach can be reconciled with G.C. Williams’ famous analysis of group adaptation, and we consider the relationships between group adaptation, the Price equation approach to multi-level selection, and the alternative approach based on contextual analysis. (shrink)

The Formal Darwinism Project is an attempt to use mathematical theory to prove the claim that fitness maximization is the outcome of evolution in nature. Grafen’s (2014, p. 12) conclusion from this project is that “….there is a very general expectation of something close to fitness maximisation, which will convert into fitness-maximisation unless there are particular kinds of circumstances—and further, that fitness is the same quantity for all genetic architectures.” Grafen’s claim appears to mean to him that natural populations are (...) expected to contain individuals whose traits are optimal, i.e., any given trait outperforms all reasonable alternatives. I describe why Grafen’s attempt can never provide a meaningful expectation as to the ubiquity of optimal traits in nature. This is so because it is based upon a misconception of the relationship between theory and empirical analysis. Even if one could use theory in the way Grafen proposes, I describe how his theory is causally incomplete. Finally, I describe how Grafen’s conceptual framework is ambiguous. The Formal Darwinism Project has been inspired by “On The Origin of Species” by Darwin. The great lesson of this book was Darwin’s demonstration of the necessary dialog between theory and data, with each influencing and being influenced by the other. Grafen’s Formal Darwinism Project, an attempt to create understanding of nature by removing data from this dialog, reflects a failure to understand Darwin’s great lesson. (shrink)

In The Origin of Species (1859) Darwin challenged many of the most deeply-held beliefs of the Western world. Arguing for a material, not divine, origin of species, he showed that new species are achieved by "natural selection." The Origin communicates the enthusiasm of original thinking in an open, descriptive style, and Darwin's emphasis on the value of diversity speaks more strongly now than ever. As well as a stimulating introduction and detailed notes, this edition offers a register of the many (...) writers referred to by Darwin in the text. (shrink)

We critically examine a number of aspects of Grafen’s ‘formal Darwinism’ project. We argue that Grafen’s ‘selection-optimality’ links do not quite succeed in vindicating the working assumption made by behavioural ecologists and others—that selection will lead organisms to exhibit adaptive behaviour—since these links hold true even in the presence of strong genetic and developmental constraints. However we suggest that the selection-optimality links can profitably be viewed as constituting an axiomatic theory of fitness. Finally, we compare Grafen’s project with Fisher’s ‘fundamental (...) theorem of natural selection’, and we speculate about whether Grafen’s results can be extended to a game-theoretic setting. (shrink)

Understanding good design requires addressing the question of what units undergo natural selection, thereby becoming adapted. There is, therefore, a natural connection between the formal Darwinism project (which aims to connect population genetics with the evolution of design and fitness maximization) and levels of selection issues. We argue that the formal Darwinism project offers contradictory and confusing lines of thinking concerning level(s) of selection. The project favors multicellular organisms over both the lower (cell) and higher (social group) levels as the (...) level of adaptation. Grafen offers four reasons for giving such special status to multicellular organisms: (1) they lack appreciable within-organism cell selection, (2) they have multiple features that appear contrived for the same purpose, (3) they possess a set of phenotypes, and (4) they leave offspring according to their phenotypes. We discuss why these rationales are not compelling and suggest that a more even-handed approach, in which multicellular organisms are not assumed to have special status, would be desirable for a project that aims to make progress on the foundations of evolutionary theory. (shrink)