Jonathan Birch

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.

I consider some hitherto unexplored examples of teleological language in the sciences. In explicating these examples, I aim to show (a) that such language is not the sole preserve of the biological sciences, and (b) that not all such talk is reducible to the ascription of functions. In chemistry and biochemistry, scientists explaining molecular rearrangements and protein folding talk informally of molecules rearranging “in order to” maximize stability. Evolutionary biologists, meanwhile, often speak of traits evolving “in order to” optimize some fitness-relevant variable. I argue that in all three contexts such locutions are best interpreted as shorthands for more detailed explanations which, were we to spell them out in full, would show that the relevant process would robustly converge towards the same end-point despite variation in initial conditions. This suggests that, in biology, such talk presupposes a substantial form of adaptationism. The upshot is that such shorthands may be more applicable in the physical sciences than the biological.

Altruistic deception (or the telling of “white lies”) is common in humans. Does it also exist in non-human animals? On some definitions of deception, altruistic deception is impossible by definition, whereas others make it too easy by counting useful-but-ambiguous information as deceptive. I argue for a definition that makes altruistic deception possible in principle without trivializing it. On my proposal, deception requires the strategic exploitation of a receiver by a sender, where “exploitation” implies that the sender elicits a behaviour in the receiver that is beneficial in a different type of situation and is expressed only because the signal raises the probability, from the receiver’s standpoint, of that type of situation. I then offer an example of a real signal that is deceptive in this sense, and yet potentially altruistic (and certainly cooperative): the purr call of the pied babbler. Fledglings associate purr calls with food, and adults exploit this learned association, in the absence of food, to lead fledglings away from predators following an alarm call. I conclude by considering why altruistic deception is apparently so rare in non-human animals.

This PhD dissertation examines the conceptual and theoretical foundations of the most general and most widely used framework for understanding social evolution, W. D. Hamilton's theory of kin selection. While the core idea is intuitive enough (when organisms share genes, they sometimes have an evolutionary incentive to help one another), its apparent simplicity masks a host of conceptual subtleties, and the theory has proved a perennial source of controversy in evolutionary biology. To move towards a resolution of these controversies, we need a careful and rigorous analysis of the philosophical foundations of the theory. My aim in this work is to provide such an analysis. I begin with an examination of the concepts behavioural ecologists employ to describe and classify types of social behaviour. I stress the need to distinguish concepts that are often conflated: for example, we need to distinguish simple cooperation from collaboration in collective tasks, behaviours from strategies, and control from manipulation and coercion. I proceed from here to the formal representation of kin selection via George R. Price’s covariance selection mathematics. I address a number of interpretative issues the Price formalism raises, including the vexed question of whether kin selection theory is ‘formally equivalent’ to multi-level selection theory. In the second half of the dissertation, I assess the uses and limits of Hamilton’s rule for the evolution of social behaviour; I provide a precise statement of the conditions under which the rival neighbour-modulated fitness and inclusive fitness approaches in contemporary kin selection theory are equivalent (and describe cases in which they are not); and I criticize recent formal attempts to establish the controversial claim that kin selection leads to organisms behaving as if maximizing their inclusive fitness.

The hunt for a biologically respectable definition for the folk concept of innateness is still on. I defend Ariew’s Canalization account of innateness against the criticisms of Griffiths and Machery, but highlight the remaining flaws in this proposal. I develop a new analysis based on the notion of environmental induction. A trait is innate, I argue, iff it is not environmentally induced. I augment this definition with a novel analysis of environmental induction that draws on the contrastive nature of causal explanation. Whether a trait is environmentally induced, I argue, depends on a context sensitive contrast class. I argue that a “Noninduction” analysis of innateness allows the concept an explanatory role in biology. I show how my proposal co-opts the successes of the Canalization account whilst avoiding its pitfalls, and I account for why biologists associate a range of disparate properties with innateness.

When two agents engage in a joint action, such as rowing together, they exercise joint know-how. But what is the relationship between the joint know-how of the two agents and the know-how each agent possesses individually? I construct an “active mutual enablement” account of this relationship, according to which joint know-how arises when each agent knows how to predict, monitor, and make failure-averting adjustments in response to the behaviour of the other agent, while actively enabling the other to make such adjustments. I defend the AME account from three objections, and I then use this account as the platform for an examination of the reducibility of joint know-how to joint propositional knowledge. A summative account of joint propositional knowledge is incompatible with the reduction of joint know-how to joint propositional knowledge, whereas a distributive account is not. I close by highlighting some open questions the AME account brings into view concerning the evolutionary origin and scaling up of joint know-how.

From mitochondria to meerkats, the natural world is full of spectacular examples of social behaviour. In the early 1960s W. D. Hamilton changed the way we think about how such behaviour evolves. He introduced three key innovations - now known as Hamilton's rule, kin selection, and inclusive fitness - and his pioneering work kick-started a research program now known as social evolution theory. This is a book about the philosophical foundations and future prospects of that program. [Note: only the Introduction is available to download.]

Animal welfare scientists face an acute version of the problem of inductive risk, since they must choose whether to affirm attributions of mental states to animals in advisory contexts, knowing their decisions hold consequences for animal welfare. In such contexts, the burden of proof should be sensitive to the consequences of error, but a framework for setting appropriate burdens of proof is lacking. Through reflection on two cases—pain and cognitive enrichment—I arrive at a tentative framework based on the principle of expected welfare maximization. I then discuss the limitations of this framework and the questions it leaves open.

Is there any way to reconcile the adaptationist’s image of natural selection as an engine of optimality with the more complex image of its dynamics we get from population genetics? This has long been an important strand in the controversy surrounding adaptationism, yet debate has been hampered by a tendency to conflate various different ways of thinking about maximization. Here I distinguish four varieties of maximization principle. I then discuss the logical relations between these varieties, arguing that, although they may seem similar at face value, none entails any of the others. I then turn briefly to the status of each variety, arguing that, while each type of maximization principle faces serious problems, the problems are subtly different for each type. In the last section, I reflect on what is at stake in the debate.

In debates about animal sentience, the precautionary principle is often invoked. The idea is that when the evidence of sentience is inconclusive, we should “give the animal the benefit of the doubt” or “err on the side of caution” in formulating animal protection legislation. Yet there remains confusion as to whether it is appropriate to apply the precautionary principle in this context, and, if so, what “applying the precautionary principle” means in practice regarding the burden of proof for animal sentience. Here I construct a version of the precautionary principle tailored to the question of animal sentience together with a practical framework for implementing it. I explain and defend the key features of this framework, argue that it is well-aligned with current practice in animal welfare science, and consider and reject a number of influential counterarguments to the use of precautionary reasoning in this area.

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 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 and its defenders. 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.

Various results show the ‘formal equivalence’ of kin and group selectionist methodologies, but this does not preclude there being a real and useful distinction between kin and group selection processes. I distinguish individual- and population-centred approaches to drawing such a distinction, and I proceed to develop the latter. On the account I advance, the differences between kin and group selection are differences of degree in the structural properties of populations. A spatial metaphor provides a useful framework for thinking about these differences: kin and group selection may be conceptualized as large, overlapping regions of K - G space. I then consider some implications of the account, defend it from possible objections, and further argue that the structural features characteristic of both kin and group selection may recur at multiple levels of biological organization.

Nick Bostrom’s ‘Simulation Argument’ purports to show that, unless we are confident that advanced ‘posthuman’ civilizations are either extremely rare or extremely rarely interested in running simulations of their own ancestors, we should assign significant credence to the hypothesis that we are simulated. I argue that Bostrom does not succeed in grounding this constraint on credence. I first show that the Simulation Argument requires a curious form of selective scepticism, for it presupposes that we possess good evidence for claims about the physical limits of computation and yet lack good evidence for claims about our own physical constitution. I then show that two ways of modifying the argument so as to remove the need for this presupposition fail to preserve the original conclusion. Finally, I argue that, while there are unusual circumstances in which Bostrom’s selective scepticism might be reasonable, we do not currently find ourselves in such circumstances. There is no good reason to uphold the selective scepticism the Simulation Argument presupposes. There is thus no good reason to believe its conclusion.

An influential argument due to Elliott Sober, subsequently strengthened by Denis Walsh and Joel Pust, moves from plausible premises to the bold conclusion that natural selection cannot explain the traits of individual organisms. If the argument were sound, the explanatory scope of selection would depend, surprisingly, on metaphysical considerations concerning origin essentialism. I show that the Sober-Walsh-Pust argument rests on a flawed counterfactual criterion for explanatory relevance. I further show that a more defensible criterion for explanatory relevance recently proposed by Michael Strevens lends support to the view that natural selection can be relevant to the explanation of individual traits.

Andrew Bourke’s Principles of Social Evolution identifies three stages that characterize an evolutionary transition in individuality and deploys inclusive fitness theory to explain each stage. The third stage, social group transformation, has hitherto received relatively little attention from inclusive fitness theorists. In this review, I first discuss Bourke’s “virtual dominance” hypothesis for the evolution of the germ line. I then contrast Bourke’s inclusive fitness approach to the major transitions with the multi-level approach developed by Richard Michod, Samir Okasha and others. I suggest that, rather than choosing between these approaches, we should exploit the strengths of both. Finally, I stress the need for a firmer conceptual grasp of the nature of social group transformation.

Most of the contributions to Cooperation and Its Evolution grapple with the distinctive challenges presented by the project of explaining human sociality. Many of these puzzles have a ‘chicken and egg’ character: our virtually unparalleled capacity for large-scale cooperation is the product of psychological, behavioural, and demographic changes in our recent evolutionary history, and these changes are linked by complex patterns of reciprocal dependence. There is much we do not yet understand about the timing of these changes, and about the order in which different aspects of human social psychology (co-)evolved. In this review essay, I discuss four such puzzles the volume raises. These concern punishment and norm-psychology, moral judgement and the moral emotions, hierarchy and top-down coercion, and property rights and legal systems

In an incendiary 2010 Nature article, M. A. Nowak, C. E. Tarnita, and E. O. Wilson present a savage critique of the best-known and most widely used framework for the study of social evolution, W. D. Hamilton’s theory of kin selection. More than a hundred biologists have since rallied to the theory’s defence, but Nowak et al. maintain that their arguments ‘stand unrefuted’. Here I consider the most contentious claim Nowak et al. defend: that Hamilton’s rule, the core explanatory principle of kin selection theory, ‘almost never holds’. I first distinguish two versions of Hamilton’s rule in contemporary theory: a special version (HRS) that requires restrictive assumptions, and a general version (HRG) that does not. I then show that Nowak et al. are most charitably construed as arguing that HRS almost never holds, while HRG buys its generality at the expense of explanatory power. While their arguments against HRS are fairly uncontroversial, their arguments against HRG are more contentious, yet these have been largely overlooked in the ensuing furore. I consider the arguments for and against the explanatory value of HRG, with a view to assessing what exactly is at stake in the debate. I suggest that the debate hinges on issues concerning the causal interpretability of regression coefficients, and concerning the explanatory function Hamilton’s rule is intended to serve.

Skyrms, building on the work of Dretske, has recently developed a novel information-theoretic account of propositional content in simple signalling systems. Information-theoretic accounts of content traditionally struggle to accommodate the possibility of misrepresentation, and I show that Skyrms’s account is no exception. I proceed to argue, however, that a modified version of Skyrms’s account can overcome this problem. On my proposed account, the propositional content of a signal is determined not by the information that it actually carries, but by the information that it would carry at the nearest separating equilibrium of the underlying evolutionary dynamics. I show that this amended account yields reasonable ascriptions of false propositional content in a well-known formal model of the evolution of communication , and close with a discussion of the serious but perhaps not insuperable difficulties we face in applying the account to examples of signalling in the real world.

Working together creates mutual obligations. For example, the members of a football team owe it to each other to work hard for the good of the team. A player who doesn’t try hard enough, or who makes a costly mistake, lets the side down. When the team loses, feelings of responsibility, guilt, and shame ensue. Players ought to feel committed to the team and responsible for its failures. If they don’t, they deserve to be dropped. The idea at the heart of Michael Tomasello’s ‘natural history’ of human morality is that these oughts—the oughts of teamwork—were the first oughts.

Hamilton’s theory of kin selection is the best-known framework for understanding the evolution of social behavior but has long been a source of controversy in evolutionary biology. A recent critique of the theory by Nowak, Tarnita, and Wilson sparked a new round of debate, which shows no signs of abating. In this overview, we highlight a number of conceptual issues that lie at the heart of the current debate. We begin by emphasizing that there are various alternative formulations of Hamilton’s rule, including a general version, which is always true; an proximate version, which assumes weak selection; and a special version, which demands other restrictive assumptions. We then examine the relationship between the neighbor-modulated fitness and inclusive fitness approaches to kin selection. Finally, we consider the often-strained relationship between the theories of kin and multilevel selection.

The theories of inclusive fitness and multilevel selection provide alternative perspectives on social evolution. The question of whether these perspectives are of equal generality remains a divisive issue. In an analysis based on the Price equation, Queller argued (by means of a principle he called the separation condition) that the two approaches are subject to the same limitations, arising from their fundamentally quantitative-genetical character. Recently, van Veelen et al. have challenged Queller’s results, using this as the basis for a broader critique of the Price equation, the separation condition, and the very notion of inclusive fitness. Here we show that the van Veelen et al. model, when analyzed in the way Queller intended, confirms rather than refutes his original conclusions. We thereby confirm (i) that Queller’s separation condition remains a legitimate theoretical principle and (ii) that the standard inclusive fitness and multilevel approaches are indeed subject to the same limitations.

Inclusive fitness theory was not originally designed to explain the major transitions in evolution, but there is a growing consensus that it has the resources to do so. My aim in this paper is to highlight, in a constructive spirit, the puzzles and challenges that remain. I first consider the distinctive aspects of the cooperative interactions we see within the most complex social groups in nature: multicellular organisms and eusocial insect colonies. I then focus on one aspect in particular: the extreme redundancy these societies exhibit. I argue that extreme redundancy poses a distinctive explanatory puzzle for inclusive fitness theory, and I offer a potential solution which casts coercion as the key enabler. I suggest that the general moral to draw from the case is one of guarded optimism: while inclusive fitness is a powerful tool for understanding evolutionary transitions, it must be integrated within a broader framework that recognizes the distinctive problems such transitions present and the distinctive mechanisms by which these problems may be overcome

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.

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.

Evolution and Rationality marks the end of a three-year project, ‘Evolution, Cooperation, and Rationality’, directed at the University of Bristol by the book’s editors, Samir Okasha and Ken Binmore. The collection draws together the editors’ pick of the papers delivered at the conferences the project hosted, and covers a wide range of topics at the intersection of evolutionary theory and the social sciences. It is a splendid anthology: timely, interdisciplinary, thematically cohesive, and full of substantive and interesting disagreements between the contributors

Cooperation is rife in the microbial world, yet our best current theories of the evolution of cooperation were developed with multicellular animals in mind. Hamilton’s theory of inclusive fitness is an important case in point: applying the theory in a microbial setting is far from straightforward, as social evolution in microbes has a number of distinctive features that the theory was never intended to capture. In this article, I focus on the conceptual challenges posed by the project of extending Hamilton’s theory to accommodate the effects of gene mobility. I begin by outlining the basics of the theory of inclusive fitness, emphasizing the role that the concept of relatedness is intended to play. I then provide a brief history of this concept, showing how, over the past fifty years, it has departed from the intuitive notion of genealogical kinship to encompass a range of generalized measures of genetic similarity. I proceed to argue that gene mobility forces a further revision of the concept. The reason in short is that, when the genes implicated in producing social behaviour are mobile, we cannot talk of an organism’s genotype simpliciter; we can talk only of an organism’s genotype at a particular stage in its life cycle. We must therefore ask: with respect to which stage(s) in the life cycle should relatedness be evaluated? For instance: is it genetic similarity at the time of social interaction that matters to the evolution of social behaviour, or is it genetic similarity at the time of reproduction? I argue that, strictly speaking, it is neither of these: what really matters to the evolution of social behaviour is diachronic genetic similarity between the producers of fitness benefits at the time they produce them and the recipients of those benefits at the end of their life-cycle. I close by discussing the implications of this result. The main payoff is that it makes room for a possible new mechanism for the evolution of altruism in microbes that does not require correlated interaction among bearers of the genes for altruism. The importance of this mechanism in nature remains an open empirical question

The sense of fairness is a central aspect of human moral psychology. Intuitions about fairness lead to many widespread moral beliefs, such as the belief that the punishment should fit the crime or the belief that one deserves a fair share of what one has earned. In The Origins of Fairness, Nicolas Baumard sets out to shed light on the evolutionary origin of these intuitions. He argues that the human sense of fairness is innate and universal, and he offers an account of its evolution that highlights the role of bargaining in early human “cooperation markets.”