This paper attempts to reconcile critics and defenders of inclusive fitness by constructing a synthesis that does justice to the insights of both. I argue that criticisms of the regression-based version of Hamilton’s rule, although they undermine its use for predictive purposes, do not undermine its use as an organizing framework for social evolution research. I argue that the assumptions underlying the concept of inclusive fitness, conceived as a causal property of an individual organism, are unlikely to be exactly true (...) in real populations, but they are approximately true given a specific type of weak selection that Hamilton took, on independent grounds, to be responsible for the cumulative assembly of complex adaptation. Finally, I reflect on the uses and limitations of “design thinking” in social evolution research. The debate about the foundations of inclusive fitness theory that has followed in the wake of Nowak, Tarnita and Wilson’s (1) critique has been remarkably polarizing. After several rounds of rebuttals and replies, there is still little evidence of any serious reconciliation between the theory’s critics (2-10) and its defenders (11-24). It doesn’t have to be this way. I believe that, on the main points of disagreement, it is possible to find a way forward that does justice to the insights of both camps. My aim in this paper is to find that way forward. (shrink)

The notion that natural selection is a process of fitness maximization gets a bad press in population genetics, yet in other areas of biology the view that organisms behave as if attempting to maximize their fitness remains widespread. Here I critically appraise the prospects for reconciliation. I first distinguish four varieties of fitness maximization. I then examine two recent developments that may appear to vindicate at least one of these varieties. The first is the ‘new’ interpretation of Fisher's fundamental theorem (...) of natural selection, on which the theorem is exactly true for any evolving population that satisfies some minimal assumptions. The second is the Formal Darwinism project, which forges links between gene frequency change and optimal strategy choice. In both cases, I argue that the results fail to establish a biologically significant maximization principle. I conclude that it may be a mistake to look for universal maximization principles justified by theory alone. A more promising approach may be to find maximization principles that apply conditionally and to show that the conditions were satisfied in the evolution of particular traits. (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)

Understanding modeling in biology requires understanding how biology is organized as a discipline and how this organization influences the research practices of biologists. Biology includes a wide range of sub-disciplines, such as cell biology, population biology, evolutionary biology, molecular biology, and systems biology among others. Biologists in sub-disciplines such as cell, molecular, and systems biology believe that the use of a few experimental models allows them to discover biological universals, whereas biologists in sub-disciplines such as ecology and evolutionary biology believe (...) that the use of many different experimental and mathematical models is necessary in order to do this. Many practitioners of both approaches misunderstand best practices of modeling, especially those related to model testing. We stress the need for biologists to better engage with best practices and for philosophers of biology providing normative guidance for biologists to better engage with current developments in biology. This is especially important as biology transitions from a “data-poor” to a “data-rich” discipline. If 21st century biology is going to capitalize on the unprecedented availability of ecological, evolutionary, and molecular data, of computational resources, and of mathematical and statistical tools, biologists will need a better understanding of what modeling is and can be. (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)