Biological agency has received much attention in recent philosophy of biology. But what is the motivation for introducing talk of agency into biology and what is meant by “agent”? Two distinct motivations can be discerned. The first is that thinking of organisms as agents helps to articulate what is distinctive about organisms vis-à-vis other biological entities. The second is that treating organisms as agent-like is a useful heuristic for understanding their evolved behavior. The concept of agent itself may be understood (...) in at least four different ways: minimal agent, intelligent agent, rational agent, and intentional agent. Which understanding is most appropriate depends on which of the two motivations we are concerned with. (shrink)

The puzzle of how altruism can evolve has been at the center of recent debates over Hamilton’s Rule, inclusive fitness, and kin-selection. In this paper, I use recent debates over altruism and Hamilton’s legacy as an example to illustrate a more general problem in evolutionary theory that has philosophical significance; I attempt to explain this significance and to draw a variety of conclusions about it. The problem is that specific behaviours and general concepts of organism agency and intentionality are defined (...) in terms of concepts of evolutionary “costs” and “benefits,” and these terms have determined the role that agency should play in evolutionary explanation. However, costs, benefits, and agency are not only or even best conceived through evolutionary effects in a biological context. The paper proceeds as follows: first, I explain how the issue of agency relates to the evolutionary puzzle of altruism. Next, I discuss how questions about agency have figured in recent debates over Hamilton’s legacy. In the final section, I argue that Denis Walsh’s “situated Darwinism,” which attempts to return the organism to central status in biological explanation, offers a more productive route for thinking about different forms of costs, benefits, and agency. Finally, I argue that the upshot of all this is that there may be many different, and equally valid, ways to express what organisms are doing and how they are behaving based on different currencies of cost and benefit—even if these may stand in some tension. I illustrate this through returning to the case of altruism and using examples to show that even in non-humans there can be many forms of altruism, even if they are not all biological altruism as defined in the conventional evolutionary terms. (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)

The ‘phenotypic gambit,’ the assumption that we can ignore genetics and look at the fitness of phenotypes to determine the expected evolutionary dynamics of a population, is often used in evolutionary game theory. However, as this paper will show, an overlooked genotype to phenotype map can qualitatively affect evolution in ways the phenotypic approach cannot predict or explain. This gives us reason to believe that, even in the long-term, correspondences between phenotypic predictions and dynamical outcomes are not robust for all (...) plausible assumptions regarding the underlying genetics of traits. This paper shows important ways in which the phenotypic gambit can fail and how to proceed with evolutionary game theoretic modeling when it does. (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)

The primary challenge for generating a useful scientific definition of life comes from competing concepts of biological activity and our failure to make them explicit in our models. I set forth a three-part scheme for characterizing definitions of life, identifying a binary , a range , and a preference . The three components together form a proper activity in biology . To be clear, I am not proposing that proper activity be adopted as the best definition of life or even (...) as a desirable definition for life. Instead, I am arguing that some notion of proper activity already exists within common scientific definitions. By making the implicit elements explicit, the notion can be analyzed to see whether it is useful and appropriate in the context of the biological sciences. (shrink)

This paper provides a philosophical analysis of the Price equation and its role in evolutionary theory. Traditional models in population genetics postulate simplifying assumptions in order to make the models mathematically tractable. On the contrary, the Price equation implies a very specific way of theorizing, starting with assumptions that we think are true and then deriving from them the mathematical rules of the system. I argue that the Price equation is a generalization-sketch, whose main purpose is to provide a unifying (...) framework for researchers, helping them to develop specific models. The Price equation plays this role because, like other scientific principles, shows features as abstractness, unification and invariance. By underwriting this special role for the Price equation some recent disputes about it could be diverted. (shrink)

The dependence on history of both present and future dynamics of life is a common intuition in biology and in humanities. Historicity will be understood in terms of changes of the space of possibilities as well as by the role of diversity in life’s structural stability and of rare events in history formation. We hint to a rigorous analysis of “path dependence” in terms of invariants and invariance preserving transformations, as it may be found also in physics, while departing from (...) the physico-mathematical analyses. The idea is that the invariant traces of the past under organismal or ecosystemic transformations contribute to the understanding of present and future states of affairs. This yields a peculiar form of unpredictability in biology, at the core of novelty formation: the changes of observables and pertinent parameters may depend also on past events. In particular, in relation to the properties of synchronic measurement in physics, the relevance of diachronic measurement in biology is highlighted. This analysis may a fortiori apply to cognitive and historical human dynamics, while allowing to investigate some general properties of historicity in biology. (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)

Evolutionary theorists often talk as if natural selection were choosing the most adapted traits, or if organisms were deciding to do the most adaptive strategy. Moreover, the payoff of those decisions often depend on what others are doing, and since Hamilton (1964), biologists possess conceptual tools such as kin selection and inclusive fitness to make sense of outcomes of evolution in these contexts, even when they seem unadaptive (such as sterility). The link between selection and adaptation through which selection or (...) organisms can be seen as agents, as well as the scope and nature of Hamiltonian conceptions of social evolution, stimulated many formal elaborations (such as, initially, Fisher’s “Fundamental theorem of natural selection”), but also raise major philosophical issues about causation and statistics, and about rationality and adaptation or selection. Two recent philosophy books, Okasha’s Agents and goals in evolution, and Birch’s Philosophy of social evolution, tackle those question. This essay reflects on them in order to think of those two issues. After having reviewed the books, I try to sketch some philosophical lessons onto which they concur. (shrink)

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)

Hamilton’s Rule, named after the evolutionary biologist Bill Hamilton, and the related concepts of inclusive fitness and kin selection, have been the bedrock of the study of social evolution for the past half century. In ’The Philosophy of Social Evolution’, Jonathan Birch provides a comprehensive introduction to the conceptual foundations of the Hamiltonian view of social evolution, and a passionate defence of its enduring value in face of the recent high profile criticism. In this review essay, I first outline the (...) version of Hamilton’s Rule defended by Birch, dubbed Hamilton’s Rule General, and its derivation. With this in place, I then navigate through the fierce disagreements that Hamilton’s Rule generates among social evolution researchers and evaluate Birch’s central argument of the book that HRG serves as an organizing framework for social evolution research under which we can compare and interpret more detailed causal models. I then spend the remainder of the review discussing what I take to be three of the most exciting implications of Hamilton’s thinking raised by Birch: the extension of Hamilton’s Rule to mobile genetic elements, maximization of inclusive fitness models and the idea of adaptation as organism design, and the relationship between Hamilton’s approaches to social behaviour and the gene’s-eye view of evolution. (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)

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)

The assumption that natural selection alone is sufficient to explain not only which traits get fixed in a population/species, but also how they develop, has been questioned since Darwin’s times, and increasingly in the last decades. Alternative theories, linked to genetic and phenotypic processes, or to the theory of complex systems, have been proposed to explain the rise of the phenotypic variety upon which natural selection acts. In this article, we illustrate the current state of the issue and we propose (...) a logical space based on phenotypic robustness that allows a classification of evolutionary phenomena and can provide a framework for unifying all these accounts. (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)

Biologists explain organisms’ behavior not only as having been programmed by genes and shaped by natural selection, but also as the result of an organism’s agency: the capacity to react to environmental changes in goal-driven ways. The use of such ‘agential explanations’ reopens old questions about how justified it is to ascribe agency to entities like bacteria or plants that obviously lack rationality and even a nervous system. Is organismic agency genuinely ‘real’ or is it just a useful fiction? In (...) this paper we focus on two questions: whether agential explanations are to be interpreted ontically, and whether they can be reduced to non-agential explanations (thereby dispensing with agency). The Kantian approach we identify interprets agential explanations non-ontically, yet holds agency to be indispensable. Attributing agency to organisms is not to be taken literally in the way we attribute physical properties such as mass or acceleration, but nor is it a mere heuristic or predictive tool. Rather, it is an inevitable consequence of our own rational capacity: as long as we are rational agents ourselves, we cannot avoid seeing agency in organisms. (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 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)

I distinguish two roles for a fitness concept in the context of explaining cumulative adaptive evolution: fitness as a predictor of gene frequency change, and fitness as a criterion for phenotypic improvement. Critics of inclusive fitness argue, correctly, that it is not an ideal fitness concept for the purpose of predicting gene-frequency change, since it relies on assumptions about the causal structure of social interaction that are unlikely to be exactly true in real populations, and that hold as approximations only (...) given a specific type of weak selection. However, Hamilton took this type of weak selection, on independent grounds, to be responsible for cumulative assembly of complex adaptations. In this special context, I argue that inclusive fitness is distinctively valuable as a criterion for improvement and a standard for optimality. Yet to call inclusive fitness a criterion for improvement and a standard for optimality is not to make any claim about the frequency with which inclusive fitness optimization actually occurs in nature. This is an empirical question that cannot be settled by theory alone. I close with some reflections on the place of inclusive fitness in the long running clash between ‘causalist’ and ‘statisticalist’ conceptions of fitness. (shrink)

Hamilton introduced two conceptions of social fitness, which he called neighbour-modulated fitness and inclusive fitness. Although he regarded them as formally equivalent, a re-analysis of his own argument for their equivalence brings out two important assumptions on which it rests: weak additivity and actor's control. When weak additivity breaks down, neither fitness concept is appropriate in its original form. When actor's control breaks down, neighbour-modulated fitness may be appropriate, but inclusive fitness is not. Yet I argue that, despite its more (...) limited domain of application, inclusive fitness provides a distinctively valuable perspective on social evolution. (shrink)

The assumption that natural selection alone is sufficient to explain not only which traits get fixed in a population/species, but also how they develop, has been questioned since Darwin’s times, and increasingly in the last decades. Alternative theories, linked to genetic and phenotypic processes, or to the theory of complex systems, have been proposed to explain the rise of the phenotypic variety upon which natural selection acts. In this article, we illustrate the current state of the issue and we propose (...) a logical space based on phenotypic robustness that allows a classification of evolutionary phenomena and can provide a framework for unifying all these accounts. (shrink)

Niche construction is a concept that captures a wide array of biological phenomena, from the environmental effects of metabolism to the creation of complex structures such as termite mounds and beaver dams. A central point in niche construction theory is that organisms do not just passively undergo developmental, ecological, or evolutionary processes, but are also active participants in them Evolution: From molecules to men, Cambridge University Press, Cambridge, 1983; Laland KN, Odling-Smee J, Feldman MW, In: KN Laland and T Uller (...) Evolutionary causation: Biological and philosophical reflections, MIT Press, Cambridge, MA, 2019). In this paper, we distinguish between two fundamentally different ways in which organisms are active participants: as agents and as contributors. Roughly, organisms act as agents when niche constructing effects are a result of a goal-directed behavior over which the organisms have some degree of control. Organisms act as contributors when the niche constructing effects do not arise from a goal to perform the constructive activity. As illustrative examples we discuss plants altering leaf-morphology to optimize light exposure as reported by Sultan and bacteria creating novel niches through excreting energy-rich metabolites. The difference between agential and contributional niche construction is important for understanding the different ways organisms can actively participate in development, ecology, and evolution. Additionally, this distinction can increase our understanding of how the capacity of agency is distributed across the tree of life and how agency influences developmental and evolutionary processes. (shrink)

Organismic agency is often understood as the capacity to produce goal-directed behavior. This paper proposes a new way of modelling agency, namely as a naturalized deliberation. Deliberative action is not directed towards a particular goal, but involves a process of weighing multiple goals and a choice for a particular combination of these. The underlying causal model is symmetry breaking, where the organism breaks symmetries present in the selective environment. Deliberation is illustrated though the phenomena of mate choice and bacterial chemotaxis.

Both biosemiotics and evolutionary epistemology are concerned with how knowledge evolves. (Applied) Evolutionary Epistemology thereby focuses on identifying the units, levels, and mechanisms or processes that underlie the evolutionary development of knowing and knowledge, while biosemiotics places emphasis on the study of how signs underlie the development of meaning. We compare the two schools of thought and analyze how in delineating their research program, biosemiotics runs into several problems that are overcome by evolutionary epistemologists. For one, by emphasizing signs, biosemiotics (...) needs to delineate a semiotic threshold, which is a problem not encountered by evolutionary epistemologists. Instead, the latter recognizes that all organisms are knowers that evolve knowledge, which they recognize to extend toward phenomena produced by organisms such as behavior, cognition, language, culture, science, and technology. Secondly, biosemiotics attempts at continuing adaptationist notions on how organisms relate to their environment, while especially Applied Evolutionary Epistemology comes to redefine the nature of the organism–environment relationship in such a way that it recognizes the spatiotemporal boundedness of existence, which in turn makes adaptationist accounts obsolete. (shrink)

In the last decades two different and apparently unrelated lines of research have increasingly connected mathematics and evolutionism. Indeed, on the one hand different attempts to formalize darwinism have been made, while, on the other hand, different attempts to naturalize logic and mathematics have been put forward. Those researches may appear either to be completely distinct or at least in some way convergent. They may in fact both be seen as supporting a naturalistic stance. Evolutionism is indeed crucial for a (...) naturalistic perspective, and formalizing it seems to be a way to strengthen its scientificity. The paper shows that, on the contrary, those directions of research may be seen as conflicting, since the conception of knowledge on which they rest may be undermined by the consequences of accepting an evolutionary perspective. (shrink)

Darwin’s claim that Natural Selection, through optimization of fitness, explains complex biological design has not yet been properly formalized. Alan Grafen’s Formal Darwinism Project aims at providing such a formalization and at demonstrating that fitness maximization is coherent with results from Population Genetics, usually interpreted as denying it. We suggest that Grafen’s proposal suffers from some limitations linked to its concept of design as optimized fitness. In order to overcome these limitations, we propose a classification of evolutionary facts based on (...) a bi-dimensional complexity space, which adds robustness to fitness as measure of the quality of a design. In this space, each point represents an organismic architecture, and movement between two points would be an evolutionary fact. Natural Selection explains movement along the fitness axis, while non-selective mechanisms explain movement on the robustness axis. Moreover, we propose a thermodynamic metaphor to draw parallelisms between notions of fitness and entropy, robustness and energy, and movement in this space and reversible and irreversible cycles. We argue that this metaphor illustrates different evolutionary processes usually overlooked by proposals of formalization of Darwinian theory, such Grafen’s Formal Darwinism Project. (shrink)