Does natural selection act primarily on individual organisms, on groups, on genes, or on whole species? The question of levels of selection - on which biologists and philosophers have long disagreed - is central to evolutionary theory and to the philosophy of biology. Samir Okasha's comprehensive analysis gives a clear account of the philosophical issues at stake in the current debate.

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.]. (shrink)

We experimentally investigated the influence of context-based biases, such as prestige and popularity, on the preferences for quotations. Participants were presented with random quotes associated to famous or unknown authors, or with random quotes presented as popular, i.e. chosen by many previous participants, or unpopular. To exclude effects related to the content of the quotations, all participants were subsequently presented with the same quotations, again associated to famous and unknown authors, or presented as popular or unpopular. Overall, our results showed (...) that context-based biases had no, or limited, effect in determining participants’ choices. Quotations preferred for their content were preferred in general, despite the contextual cues to which they were associated. We conclude discussing how our results fit with the well-known phenomenon of the spread and success of misattributed quotations, and we draw some more general implications for cultural evolution research. (shrink)

Acknowledgments 1. Culture Is Essential 2. Culture Exists 3. Culture Evolves 4. Culture Is an Adaptation 5. Culture Is Maladaptive 6. Culture and Genes Coevolve 7. Nothing about Culture Makes Sense except in the Light of Evolution.

Jon Elster and Daniel Little have criticized social scientists for appealing to a mechanism of social selection in functional explanations of social practices. Both believe that there is no such mechanism operative in the social world. I develop and defend an account of functional explanation in which a mechanism of social selection figures centrally. In addition to developing an account of social selection, I clarify what functional hypotheses purport to claim, and re-examine the role of agents’ intentions in functional explanations (...) in an effort to show why a mechanism of social selection is indispensable to adequate functional explanations. (shrink)

Woodward's long awaited book is an attempt to construct a comprehensive account of causation explanation that applies to a wide variety of causal and explanatory claims in different areas of science and everyday life. The book engages some of the relevant literature from other disciplines, as Woodward weaves together examples, counterexamples, criticisms, defenses, objections, and replies into a convincing defense of the core of his theory, which is that we can analyze causation by appeal to the notion of manipulation.

This paper attempts to elucidate three characteristics of causal relationships that are important in biological contexts. Stability has to do with whether a causal relationship continues to hold under changes in background conditions. Proportionality has to do with whether changes in the state of the cause “line up” in the right way with changes in the state of the effect and with whether the cause and effect are characterized in a way that contains irrelevant detail. Specificity is connected both to (...) David Lewis’ notion of “influence” and also with the extent to which a causal relation approximates to the ideal of one cause–one effect. Interrelations among these notions and their possible biological significance are also discussed. (shrink)

The theoretical status of ‘niche construction’ in evolution is intensely debated. Here we substantiate the reasons for different interpretations. We consider two concepts of niche construction brought to bear on evolutionary theory; one that emphasizes how niche construction contributes to selection and another that emphasizes how it contributes to development and inheritance. We explain the rationale for claims that selective and developmental niche construction motivate conceptual change in evolutionary biology and the logic of those who reject these claims. Our analysis (...) shows how the contention arises from alternative assumptions regarding the causal independence of the processes that generate variation, differential fitness and inheritance. (shrink)

I defend a radical interpretation of biological populations—what I call population pluralism—which holds that there are many ways that a particular grouping of individuals can be related such that the grouping satisfies the conditions necessary for those individuals to evolve together. More constraining accounts of biological populations face empirical counter-examples and conceptual difficulties. One of the most intuitive and frequently employed conditions, causal connectivity—itself beset with numerous difficulties—is best construed by considering the relevant causal relations as ‘thick’ causal concepts. I (...) argue that the fine-grained causal relations that could constitute membership in a biological population are huge in number and many are manifested by degree, and thus we can construe population membership as being defined by massively multidimensional constructs, the differences between which are largely arbitrary. I end by showing that positions in two recent debates in theoretical biology depend on a view of biological populations at odds with the pluralism defended here. (shrink)

I defend a radical interpretation of biological populations—what I call population pluralism—which holds that there are many ways that a particular grouping of individuals can be related such that the grouping satisfies the conditions necessary for those individuals to evolve together. More constraining accounts of biological populations face empirical counter-examples and conceptual difficulties. One of the most intuitive and frequently employed conditions, causal connectivity—itself beset with numerous difficulties—is best construed by considering the relevant causal relations as ‘thick’ causal concepts. I (...) argue that the fine-grained causal relations that could constitute membership in a biological population are huge in number and many are manifested by degree, and thus we can construe population membership as being defined by massively multidimensional constructs, the differences between which are largely arbitrary. I end by showing that positions in two recent debates in theoretical biology depend on a view of biological populations at odds with the pluralism defended here. 1 Introduction2 Biological Population, Broad and Narrow3 Difficulties with Narrow Biological Population Conditions3.1 Against the genealogical condition3.2 Against the conspecificity condition3.3 Against the proximity condition3.4 Against the typology condition4 Causal Connectivity5 Massively Multidimensional Population Constructs6 Population Uniqueness and Natural Selection6.1 Statisticalism and its discontents6.2 Price at what price?7 Conclusion. (shrink)

This article introduces and defends a probabilistic measure of the explanatory power that a particular explanans has over its explanandum. To this end, we propose several intuitive, formal conditions of adequacy for an account of explanatory power. Then, we show that these conditions are uniquely satisfied by one particular probabilistic function. We proceed to strengthen the case for this measure of explanatory power by proving several theorems, all of which show that this measure neatly corresponds to our explanatory intuitions. Finally, (...) we briefly describe some promising future projects inspired by our account. (shrink)

Recent debates about memetics have revealed some widespread misunderstandings about Darwinian approaches to cultural evolution. Drawing from these debates, this paper disputes five common claims: (1) mental representations are rarely discrete, and therefore models that assume discrete, gene-like particles (i.e., replicators) are useless; (2) replicators are necessary for cumulative, adaptive evolution; (3) content-dependent psychological biases are the only important processes that affect the spread of cultural representations; (4) the “cultural fitness” of a mental representation can be inferred from its successful (...) transmission; and (5) selective forces only matter if the sources of variation are random. We close by sketching the outlines of a unified evolutionary science of culture. (shrink)

Human cooperation is highly unusual. We live in large groups composed mostly of non-relatives. Evolutionists have proposed a number of explanations for this pattern, including cultural group selection and extensions of more general processes such as reciprocity, kin selection, and multi-level selection acting on genes. Evolutionary processes are consilient; they affect several different empirical domains, such as patterns of behavior and the proximal drivers of that behavior. In this target article, we sketch the evidence from five domains that bear on (...) the explanatory adequacy of cultural group selection and competing hypotheses to explain human cooperation. Does cultural transmission constitute an inheritance system that can evolve in a Darwinian fashion? Are the norms that underpin institutions among the cultural traits so transmitted? Do we observe sufficient variation at the level of groups of considerable size for group selection to be a plausible process? Do human groups compete, and do success and failure in competition depend upon cultural variation? Do we observe adaptations for cooperation in humans that most plausibly arose by cultural group selection? If the answer to one of these questions is “no,” then we must look to other hypotheses. We present evidence, including quantitative evidence, that the answer to all of the questions is “yes” and argue that we must take the cultural group selection hypothesis seriously. If culturally transmitted systems of rules that limit individual deviance organize cooperation in human societies, then it is not clear that any extant alternative to cultural group selection can be a complete explanation. (shrink)

Fitness is a central concept in evolutionary theory. Just as it is central to biological evolution, so, it seems, it should be central to cultural evolutionary theory. But importing the biological fitness concept to CET is no straightforward task—there are many features unique to cultural evolution that make this difficult. This has led some theorists to argue that there are fundamental problems with cultural fitness that render it hopelessly confused. In this essay, we defend the coherency of cultural fitness against (...) those who call it into doubt. (shrink)

This paper analyzes recent attempts to reject reproduction with lineage formation as a necessary condition for evolution by means of natural selection :560–570, 2008; Stud Hist Philos Sci Part C Stud Hist Philos Biol Biomed Sci 42:106–114, 2011; Bourrat in Biol Philos 29:517–538, 2014; Br J Philos Sci 66:883–903, 2015; Charbonneau in Philos Sci 81:727–740, 2014; Doolittle and Inkpen in Proc Natl Acad Sci 115:4006–4014, 2018). Building on the strengths of these attempts and avoiding their pitfalls, it is argued that (...) a robust formulation of evolution by natural selection without reproduction can be established. The main contribution of this paper is a reformulation of Lewontin’s three principles stating that minimal evolution by natural selection occurs when two conditions are met in a population: fitness-related variation and memory. Paradigmatic evolution by natural selection, which can generate adaptations, takes place when an additional condition is present, namely regeneration. (shrink)

We develop a cultural evolutionary theory of the origins of prosocial religions and apply it to resolve two puzzles in human psychology and cultural history: (1) the rise of large-scale cooperation among strangers and, simultaneously, (2) the spread of prosocial religions in the last 10–12 millennia. We argue that these two developments were importantly linked and mutually energizing. We explain how a package of culturally evolved religious beliefs and practices characterized by increasingly potent, moralizing, supernatural agents, credible displays of faith, (...) and other psychologically active elements conducive to social solidarity promoted high fertility rates and large-scale cooperation with co-religionists, often contributing to success in intergroup competition and conflict. In turn, prosocial religious beliefs and practices spread and aggregated as these successful groups expanded, or were copied by less successful groups. This synthesis is grounded in the idea that although religious beliefs and practices originally arose as nonadaptive by-products of innate cognitive functions, particular cultural variants were then selected for their prosocial effects in a long-term, cultural evolutionary process. This framework (1) reconciles key aspects of the adaptationist and by-product approaches to the origins of religion, (2) explains a variety of empirical observations that have not received adequate attention, and (3) generates novel predictions. Converging lines of evidence drawn from diverse disciplines provide empirical support while at the same time encouraging new research directions and opening up new questions for exploration and debate. (shrink)

In our response to the 27 commentaries, we refine the theoretical claims, clarify several misconceptions of our framework, and explore substantial disagreements. In doing so, we show that our framework accommodates multiple historical scenarios; debate the historical evidence, particularly about “pre-Axial” religions; offer important details about cultural evolutionary theory; clarify the termprosociality;and discuss proximal mechanisms. We review many interesting extensions, amplifications, and qualifications of our approach made by the commentators.

argue that natural selection does not explain the genotypic arid phenotypic properties of individuals. On this view, natural selection explains the adaptedness of individuals, not by explaining why the individuals that exist have the adaptations they do, but rather by explaining why the individuals that exist are the ones with those adaptations. This paper argues that this ‘Negative’ view of natural selection ignores the fact that natural selection is a cumulative selection process. So understood, it explains how the genetic sequences (...) that individuals inherit and that are responsible for their complex (and co-adapted) adaptations first arose in the gene-pool. (shrink)

Cognitive gadgets puts forward an ambitious claim: language, mindreading, and imitation evolved by cultural group selection. Defending this claim requires more than Heyes' spirited and effective critique of nativist claims. The latest human “cognitive gadgets,” such as literacy, did not spread through cultural group selection. Why should social cognition be different? The book leaves this question pending. It also makes strong assumptions regarding cultural evolution: it is moved by selection rather than transformation; it relies on high‐fidelity imitation; it requires specific (...) cognitive adaptations to cultural learning. Each of these assumptions raises crucial yet unaddressed difficulties. (shrink)

We suggest that human culture exhibits key Darwinian evolutionary properties, and argue that the structure of a science of cultural evolution should share fundamental features with the structure of the science of biological evolution. This latter claim is tested by outlining the methods and approaches employed by the principal subdisciplines of evolutionary biology and assessing whether there is an existing or potential corresponding approach to the study of cultural evolution. Existing approaches within anthropology and archaeology demonstrate a good match with (...) the macroevolutionary methods of systematics, paleobiology, and biogeography, whereas mathematical models derived from population genetics have been successfully developed to study cultural microevolution. Much potential exists for experimental simulations and field studies of cultural microevolution, where there are opportunities to borrow further methods and hypotheses from biology. Potential also exists for the cultural equivalent of molecular genetics in “social cognitive neuroscience,” although many fundamental issues have yet to be resolved. It is argued that studying culture within a unifying evolutionary framework has the potential to integrate a number of separate disciplines within the social sciences. (shrink)

The article addresses the question whether culture evolves in a Lamarckian manner. I highlight three central aspects of a Lamarckian concept of evolution: the inheritance of acquired characteristics, the transformational pattern of evolution, and the concept of directed changes. A clear exposition of these aspects shows that a system can be a Darwinian variational system instead of a Lamarckian transformational one, even if it is based on inheritance of acquired characteristics and/or on Lamarckian directed changes. On this basis, I apply (...) the three aspects to culture. Taking for granted that culture is a variational system, based on selection processes, I discuss in detail the senses in which cultural inheritance can be said to be Lamarckian and in which sense problem solving, a major factor in cultural change, leads to directed variation. (shrink)

The idea that cultural evolution exhibits variation, competition, and inheritance and therefore can be studied by adjusting the Darwinian theory of evolution by natural selection is an attractive one. It has been argued by a number of authors (e.g., Campbell 1960; Monod 1970; Dawkins 1976; Cavalli-Sforza and Feldman 1981; Boyd and Richerson 1985; Durham 1991; Aunger 2002; Mesoudi et al. 2004) and pursued in a variety of ways, some (Dawkins and memeticists) staying close to the Darwinian model, others (e.g., Boyd, (...) Richerson, and their collaborators) being more innovative. We agree that there are relevant analogies between biological and cultural evolution and, in particular, that cultural items do exhibit variation, competition, and cumulative modification. On the other hand, we believe that a proper understanding of the mechanisms of cultural propagation drawing on the work of cognitive and social scientists (see Sperber and Hirschfeld 1999 for a review) contradicts the idea that culture exhibits inheritance in the strict sense needed for the theory of evolution by natural selection to apply straightforwardly to it. If so, it will take more than adjusting the Darwinian model to be faithful to the Darwinian inspiration. (shrink)

This paper examines conceptual issues that arise in applications of Darwinian natural selection to cultural systems. I argue that many criticisms of cultural selectionist models have been based on an over-detailed reading of the analogy between biological and cultural units of selection. I identify five of the most powerful objections to cultural selection theory and argue that none cuts to its heart. Some objections are based on mistaken assumptions about the simplicity of the mechanisms of biological heredity. Other objections are (...) attributable, rather, to mistaken inferences from observations of biological subject matter to what is essential in natural selection. I argue that such features are idiosyncratic of biological systems, but not essential for natural selection. My arguments throughout are illustrated by examples from biological and cultural evolution, and counter-factual illustrations from the history of theoretical biology. Introduction Cultural Selection Theory First Objection: Lamarckianism Second Objection: Genotype–Phenotype Distinction Third Objection: Common Hereditary Architecture Fourth Objection: Biological Analogue for Cultural Units 6.1 Regarding strict analogues 6.2 Regarding the trait analogue 6.3 Regarding the virus analogue Fifth Objection: Environmental Interaction Conclusion CiteULike Connotea Del.icio.us What's this? (shrink)

In The Secret of Our Success, Joseph Henrich claims that human beings are unique—different from all other animals—because we engage in cumulative cultural evolution. It is the technological and social products of cumulative cultural evolution, not the intrinsic rationality or ‘smartness’ of individual humans, that enable us to live in a huge range of different habitats, and to dominate most of the creatures who share those habitats with us. We are sympathetic to this general view, the latest expression of the (...) ‘California school’s’ view of cultural evolution, and impressed by the lively and interesting way that Henrich handles evidence from anthropology, economics, and many fields of biology. However, because we think it is time for cultural evolutionists to get down to details, this essay review raises questions about Henrich’s analysis of both the cognitive processes and the selection processes that contribute to cumulative cultural evolution. In the former case, we argue that cultural evolutionists need to make more extensive use of cognitive science, and to consider the evidence that mechanisms of cultural learning are products as well as processes of cultural evolution. In the latter case, we ask whether the California school is really serious about selection, or whether it is offering a merely ‘kinetic’ view of cultural evolution, and, assuming the former, outline four potential models of cultural selection that it would be helpful to distinguish more clearly. (shrink)

Some cultural evolution researchers have argued for the importance of prestige bias as a systematic and widespread social learning bias, that structures human social learning and cultural transmission patterns. Broadly speaking, prestige bias accounts understand it as a bias towards copying ‘prestigious’ individuals. Prestige bias, along with other social learning biases, has been argued to pay a crucial role in allowing cumulative cultural selection to take place, thereby generating adaptations that are key to our success as a species. However, I (...) argue for skepticism about the plausibility and scope of a prestige bias account. I argue that although an account of prestige bias seems plausible or compelling on their face, it is committed to a particular view of the cognition underpinning the bias, and therefore to predictions regarding its flexibility and context-sensitivity. Given this, current empirical evidence gives us reason to doubt the explanatory value of a prestige bias account over a naive, goal-directed agent account. Additionally, the way that prestige is defined in empirical work is in tension with a general understanding of prestige, casting doubt upon its status as evidence of prestige bias. I examine two studies cited as evidence of prestige bias, arguing that in these cases we cannot clearly favour a prestige bias explanation over a goal-directed agent explanation. (shrink)

Recently advocates of the propensity interpretation of fitness have turned critics. To accommodate examples from the population genetics literature they conclude that fitness is better defined broadly as a family of propensities rather than the propensity to contribute descendants to some future generation. We argue that the propensity theorists have misunderstood the deeper ramifications of the examples they cite. These examples demonstrate why there are factors outside of propensities that determine fitness. We go on to argue for the more general (...) thesis that no account of fitness can satisfy the desiderata that have motivated the propensity account. (shrink)

Cultural evolution studies are characterized by the notion that culture evolves accordingly to broadly Darwinian principles. Yet how far the analogy between cultural and genetic evolution should be pushed is open to debate. Here, we examine a recent disagreement that concerns the extent to which cultural transmission should be considered a preservative mechanism allowing selection among different variants, or a transformative process in which individuals recreate variants each time they are transmitted. The latter is associated with the notion of “cultural (...) attraction”. This issue has generated much misunderstanding and confusion. We first clarify the respective positions, noting that there is in fact no substantive incompatibility between cultural attraction and standard cultural evolution approaches, beyond a difference in focus. Whether cultural transmission should be considered a preservative or reconstructive process is ultimately an empirical question, and we examine how both preservative and reconstructive cultural transmission has been studied in recent experimental research in cultural evolution. Finally, we discuss how the relative importance of preservative and reconstructive processes may depend on the granularity of analysis and the domain being studied. (shrink)

Tim Lewens aims to understand what it means to take an evolutionary approach to cultural change, and why it is that these approaches are sometimes treated with suspicion. While making a case for the value of evolutionary thinking for students of culture, he shows why the concerns of sceptics should not dismissed as mere prejudice, confusion, or ignorance. Indeed, confusions about what evolutionary approaches entail are propagated by their proponents, as well as by their detractors. By taking seriously the problems (...) faced by these approaches to culture, he shows how such approaches can be better formulated, where their most significant limitations lie, and how the tools of cultural evolutionary thinking might become more widely accepted. (shrink)

This book was first published in 1984, as the revised edition of a 1979 original. The text is composed of studies in a descending sequence from perfect rationality, through imperfect and problematical rationality, to irrationality. Specifically human rationality is characterized by its capacity to relate strategically to the future, in contrast to the myopic 'gradient climbing' of natural selection. There is trenchant analysis of some of the parallels proposed in this connection between the biological and the social sciences. In the (...) chapter on imperfect rationality the crucial notion is that of 'binding oneself', as Ulysses did before setting out to the Sirens, when weakness of will may prevent us from using our capacity for perfect rationality. The second half of the book deals with rational-actor theory, comparing its logical power and success to rival approaches, and with the varieties of irrationality expressed in contradictory beliefs and desires. (shrink)

The 'Evolution of Cooperation' addresses a simple yet age-old question; If living things evolve through competition, how can cooperation ever emerge? Despite the abundant evidence of cooperation all around us, there existed no purely naturalistic answer to this question until 1979, when Robert Axelrod famously ran a computer tournament featuring a standard game-theory exercise called The Prisoner's Dilemma. To everyone's surprise, the program that won the tournament, named Tit for Tat, was not only the simplest but the most "cooperative" entrant. (...) This unexpected victory proved that cooperation - one might even say altruism - is mathematically possible and therefore needs no hidden hand or divine agent to create and sustain it. A great roadblock to the understanding of all sorts of behavior was at last removed. The updated edition includes an extensive new chapter on cooperation in cancer cells and among terrorist organizations. (shrink)

The book presents a new way of understanding Darwinism and evolution by natural selection, combining work in biology, philosophy, and other fields.

Human migration is nonrandom. In small scale societies of the past, and in the modern world, people tend to move to wealthier, safer, and more just societies from poorer, more violent, less just societies. If immigrants are assimilated, such nonrandom migration can increase the occurrence of culturally transmitted beliefs, values, and institutions that cause societies to be attractive to immigrants. Here we describe and analyze a simple model of this process. This model suggests that long run outcomes depend on the (...) relative strength of migration and local adaptation. When local adaption is strong enough to preserve cultural variation among groups, cultural variants that make societies attractive always predominate, but never drive alternative variants to extinction. When migration predominates, outcomes depend both on the relative attractiveness of alternative variants and on the initial sizes of societies that provide and receive immigrants. (shrink)

Niche construction refers to the activities of organisms that bring about changes in their environments, many of which are evolutionarily and ecologically consequential. Advocates of niche construction theory (NCT) believe that standard evolutionary theory fails to recognize the full importance of niche construction, and consequently propose a novel view of evolution, in which niche construction and its legacy over time (ecological inheritance) are described as evolutionary processes, equivalent in importance to natural selection. Here, we subject NCT to critical evaluation, in (...) the form of a collaboration between one prominent advocate of NCT, and a team of skeptics. We discuss whether niche construction is an evolutionary process, whether NCT obscures or clarifies how natural selection leads to organismal adaptation, and whether niche construction and natural selection are of equivalent explanatory importance. We also consider whether the literature that promotes NCT overstates the significance of niche construction, whether it is internally coherent, and whether it accurately portrays standard evolutionary theory. Our disagreements reflect a wider dispute within evolutionary theory over whether the neo-Darwinian synthesis is in need of reformulation, as well as different usages of some key terms (e.g., evolutionary process). (shrink)

We make the case that the Prisoner’s Dilemma, notwithstanding its fame and the quantity of intellectual resources devoted to it, has largely failed to explain any phenomena of social scientific or biological interest. In the heart of the paper we examine in detail a famous purported example of Prisoner’s Dilemma empirical success, namely Axelrod’s analysis of WWI trench warfare, and argue that this success is greatly overstated. Further, we explain why this negative verdict is likely true generally and not just (...) in our case study. We also address some possible defenses of the Prisoner’s Dilemma. (shrink)

Group beneficial norms are common in human societies. The persistence of such norms is consistent with evolutionary game theory, but existing models do not provide a plausible explanation for why they are common. We show that when a model of imitation used to derive replicator dynamics in isolated populations is generalized to allow for population structure, group beneficial norms can spread rapidly under plausible conditions. We also show that this mechanism allows recombination of different group beneficial norms arising in..

Over the past several decades, we have argued that cultural evolution can facilitate the evolution of largescale cooperation because it often leads to more rapid adaptation than genetic evolution, and, when multiple stable equilibria exist, rapid adaptation leads to variation among groups. Recently, Lehmann, Feldman, and colleagues have published several papers questioning this argument. They analyze models showing that cultural evolution can actually reduce the range of conditions under which cooperation can evolve and interpret these models as indicating that we (...) were wrong to conclude that culture facilitated the evolution of human cooperation. In the main, their models assume that rates of cultural adaption are not.. (shrink)