Proponents of the standard evolutionary biology paradigm explain human “altruism” in terms of either nepotism or strict reciprocity. On that basis our underlying nature is reduced to a function of inclusive fitness: human nature has to be totally selfish or nepotistic. Proposed here are three possible paths to giving costly aid to nonrelatives, paths that are controversial because they involve assumed pleiotropic effects or group selection. One path is pleiotropic subsidies that help to extend nepotistic helping behavior (...) from close family to nonrelatives. Another is “warfare”—if and only if warfare recurred in the Paleolithic. The third and most plausible hypothesis is based on the morally based egalitarian syndrome of prehistoric hunter-gatherers, which reduced phenotypic variation at the within-group level, increased it at the between-group level, and drastically curtailed the advantages of free riders. In an analysis consistent with the fundamental tenets of evolutionary biology, these three paths are evaluated as explanations for the evolutionary development of a rather complicated human social nature. (shrink)

In social evolution theory, biological individuals are often represented on the model of rational agents, that is, as if they were ‘seeking’ to maximize their own (expected) reproductive success. In the 1990s, important criticisms of this mode of thinking were made by Brian Skyrms ([1994], [1996]) and Elliott Sober ([1998]), who both argued that ‘rational agent’ models can lead to incorrect predictions when there are positive correlations between individuals’ phenotypes. In this article, I argue that one model of rational choice—namely, (...) Savage’s model ([1954])—can actually be vindicated in evolutionary biology, provided that the pay-offs are computed in inclusive fitness terms. I also show that the use of this model is better avoided when pay-offs are non-additive, or when certain causal influences (due to manipulative behaviours) affect the outcome of natural selection. The result is a partial rehabilitation of this mode of thinking, conditional on both the additivity of the pay-off structure and the absence of any form of manipulation or coercion. 1 Introduction2 When Natural Selection and Rational Deliberation Part Ways3 A Simple Solution: Redefining the Pay-offs in Inclusive Fitness Terms4 Sober on Inclusive Fitness Maximization5 Inclusive Fitness with Non-additive Pay-offs6 Causal Influences and the Savage– Hamilton Model6.1 Reciprocity and partner choice6.2 Coercion and manipulation7 ConclusionAppendix. (shrink)

Altruism is a central concept in evolutionary biology. Evolutionary biologists still disagree about its meaning (E.O. Wilson 2005; Fletcher et al. 2006; D.S. Wilson 2008; Foster et al. 2006a, b; West et al. 2007a, 2008). Semantic disagreement appears to be quite robust and not easily overcome by attempts at clarification, suggesting that substantive conceptual issues lurk in the background. Briefly, group selection theorists define altruism as any trait that makes altruists losers to selfish traits within groups, and (...) makes groups of altruists fitter than groups of non-altruists. Inclusive fitness theorists reject a definition based on within- and between-group fitness. Traits are altruistic only if they cause a direct and absolute fitness loss to the donor. The latter definition is more restrictive and rejects as cases of altruism behaviors that are accepted by the former. Fletcher and Doebeli (2009) recently proposed a simple, direct and individually based fitness approach, which they claim returns to first principles: carriers of the genotype of interest “must, on average, end up with more net direct fitness benefits than average population members.” This seductively simple proposal uses the concept of assortment to explain how diverse kinds of altruists end up on average with more net fitness than their non-altruistic rivals. In this paper I shall argue that their approach implies a new concept of altruism that contrasts with and improves on the concept of the inclusive fitness approach. (shrink)

ABSTRACT In social evolution theory, biological individuals are often represented on the model of rational agents, that is, as if they were ‘seeking’ to maximize their own reproductive success. In the 1990s, important criticisms of this mode of thinking were made by Brian Skyrms and Elliott Sober, who both argued that ‘rational agent’ models can lead to incorrect predictions when there are positive correlations between individuals’ phenotypes. In this article, I argue that one model of rational choice—namely, Savage’s model —can (...) actually be vindicated in evolutionary biology, provided that the pay-offs are computed in inclusive fitness terms. I also show that the use of this model is better avoided when pay-offs are non-additive, or when certain causal influences affect the outcome of natural selection. The result is a partial rehabilitation of this mode of thinking, conditional on both the additivity of the pay-off structure and the absence of any form of manipulation or coercion. _1_ Introduction _2_ When Natural Selection and Rational Deliberation Part Ways _3_ A Simple Solution: Redefining the Pay-offs in Inclusive Fitness Terms _4_ Sober on Inclusive Fitness Maximization _5_ Inclusive Fitness with Non-additive Pay-offs _6_ Causal Influences and the Savage– Hamilton Model _6.1_ Reciprocity and partner choice _6.2_ Coercion and manipulation _7_ Conclusion Appendix. (shrink)

Reciprocal altruism was originally formulated in terms of individual selection and most theorists continue to view it in this way. However, this interpretation of reciprocal altruism has been challenged by Sober and Wilson (1998). They argue that reciprocal altruism (as well as all other forms of altruism) evolves by the process of group selection. In this paper, we argue that the original interpretation of reciprocal altruism is the correct one. We (...) accomplish this by arguing that if fitness attaches to (at minimum) entire life cycles, then the kind of fitness exchanges needed to form the group-level in such situations is not available. Reciprocal altruism is thus a result of individual selection and when it evolves, it does so because it is individually advantageous. (shrink)

In both biology and the human sciences, social groups are sometimes treated as adaptive units whose organization cannot be reduced to individual interactions. This group-level view is opposed by a more individualistic one that treats social organization as a byproduct of self-interest. According to biologists, group-level adaptations can evolve only by a process of natural selection at the group level. Most biologists rejected group selection as an important evolutionary force during the 1960s and 1970s but (...) a positive literature began to grow during the 1970s and is rapidly expanding today. We review this recent literature and its implications for human evolutionary biology. We show that the rejection of group selection was based on a misplaced emphasis on genes as “replicators” which is in fact irrelevant to the question of whether groups can be like individuals in their functional organization. The fundamental question is whether social groups and other higher-level entities can be “vehicles” of selection. When this elementary fact is recognized, group selection emerges as an important force in nature and what seem to be competing theories, such as kin selection and reciprocity, reappear as special cases of group selection. The result is a unified theory of natural selection that operates on a nested hierarchy of units.The vehicle-based theory makes it clear that group selection is an important force to consider in human evolution. Humans can facultatively span the full range from self-interested individuals to “organs” of group-level “organisms.” Human behavior not only reflects the balance between levels of selection but it can also alter the balance through the construction of social structures that have the effect of reducing fitness differences within groups, concentrating natural selection (and functional organization) at the group level. These social structures and the cognitive abilities that produce them allow group selection to be important even among large groups of unrelated individuals. (shrink)

Sober and Wilson's enthusiasm for a multi-level perspective in evolutionary biology leads to conceptualizations which appropriate all sources of bio-altruistic traits as products of ‘group’ selection. The key biological issue is whether genes enhancing one sub-population's viability in competition with others can thrive, despite inducing some members to lose fitness in intra-group terms. The case for such selection amongst primates remains unproven. Flexible social loyalties required prior evolution of subjective self-definition and self-identification with others. But normative readiness for (...) truly group-serving sacrifices of ego-interests presupposes entitative conceptions of in-groups as collective social units -- only made possible with the emergence of human symbolic language. (shrink)

The question on how the diverse forms of cooperative behavior in humans and nonhuman animals could have evolved under the pressure of natural selection has been a challenge for evolutionary biology ever since Darwin himself. In this chapter, we briefly review and summarize results from the last 50 years of research on human and nonhuman cooperativeness from a theoretical (biology) and an experimental perspective (experimental economics). The first section presents six concepts from theoretical biology able to explain a variety (...) of forms of cooperativeness which evolved in many different species. These are kin selection, mutualism, reciprocity, green-beard altruism, costly signaling, and cultural group selection. These considerations are complemented by two short examples of evolved cooperative behavior, one from microbiology and one from ethology. The second main section focuses on recent experimental research on human cooperativeness. We present a brief review of factors known to impact individual human decision-making in social dilemmas, most prominently communication, punishment, reputation, and assortment. Our conclusion then draws attention to tasks for further research in this area. (shrink)

Phenomena like meat sharing in hunter-gatherers, altruistic self-sacrifice in intergroup conflicts, and contribution to the production of public goods in laboratory experiments have led to the development of numerous theories trying to explain human prosocial preferences and behavior. Many of these focus on direct and indirect reciprocity, assortment, or (cultural) group selection. Here, I investigate analytically how genetic relatedness changes the incentive structure of that paradigmatic game which is conventionally used to model and experimentally investigate collective action problems: the (...) public goods game. Using data on contemporary hunter-gatherer societies I then estimate a threshold value determining when biological altruism turns into maximizing inclusive fitness in this game. I find that, on average, contributing no less than about 40% of individual fitness to public goods production still is an optimal strategy from an inclusive fitness perspective under plausible socio-ecological conditions. (shrink)

In both biology and the human sciences, social groups are sometimes treated as adaptive units whose organization cannot be reduced to individual interactions. This group-level view is opposed by a more individualistic one that treats social organization as a byproduct of self-interest. According to biologists, group-level adaptations can evolve only by a process of natural selection at the group level. Most biologists rejected group selection as an important evolutionary force during the 1960s and 1970s but (...) a positive literature began to grow during the 1970s and is rapidly expanding today. We review this recent literature and its implications for human evolutionary biology. We show that the rejection of group selection was based on a misplaced emphasis on genes as "replicators" which is in fact irrelevant to the question of whether groups can be like individuals in their functional organization. The fundamental question is whether social groups and other higher-level entities can be "vehicles" of selection. When this elementary fact is recognized, group selection emerges as an important force in nature and what seem to be competing theories, such as kin selection and reciprocity, reappear as special cases of group selection. The result is a unified theory of natural selection that operates on a nested hierarchy of units. The vehicle-based theory makes it clear that group selection is an important force to consider in human evolution. Humans can facultatively span the full range from self-interested individuals to "organs" of group-level "organisms." Human behavior not only reflects the balance between levels of selection but it can also alter the balance through the construction of social structures that have the effect of reducing fitness differences within groups, concentrating natural selection (and functional organization) at the group level. These social structures and the cognitive abilities that produce them allow group selection to be important even among large groups of unrelated individuals. (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)

Humans often behave altruistically towards strangers with no chance of reciprocation. From an evolutionary perspective, this is puzzling. The evolution of altruistic cooperative behavior—in which an organism’s action reduces its fitness and increases the fitness of another organism —only makes sense when it is directed at genetically related organisms or when one can expect the favor to be returned. Therefore, evolutionary theorists such as Sober and Wilson have argued that we should revise Neo-Darwininian evolutionary theory. They argue that human (...) class='Hi'>altruism evolved through group selection in which groups of altruists were naturally selected because they had a comparative advantage over other groups. Wilson and Sober’s hypothesis attracted followers but is rejected by most of their peers. The heated debate between advocates and critics of group selection often suffers from a lack of conceptual clarity. In response, I set out to clearly distinguish ‘genetic’ from ‘cultural’ group selection and argue that the latter does not face the potentially debilitating problems plaguing the former. I defend the claim that human altruistic dispositions evolved through cultural group selection and gene-culture coevolution and offer empirical evidence in support. I also argue that actual altruistic behavior often goes beyond the kind of behavior humans have evolved to display. Conscious and voluntary reasoning processes, I show, have an important role in altruistic behavior. This is often overlooked in the scientific literature on human altruism. (shrink)

It’s recently been argued that biological fitness can’t change over the course of an organism’s life as a result of organisms’ behaviors. However, some characterizations of biological function and biological altruism tacitly or explicitly assume that an effect of a trait can change an organism’s fitness. In the first part of the paper, I explain that the core idea of changing fitness can be understood in terms of conditional probabilities defined over sequences of events in an organism’s life. The (...) result is a notion of “conditional fitness” which is static but which captures intuitions about apparent behavioral effects on fitness. The second part of the paper investigates the possibility of providing a systematic foundation for conditional fitness in terms of spaces of sequences of states of an organism and its environment. I argue that the resulting “organism–environment history conception” helps unify diverse biological perspectives, and may provide part of a metaphysics of natural selection. (shrink)

Inclusive fitness theory provides conditions for the evolutionary success of a gene. These conditions ensure that the gene is selfish in the sense of Dawkins (The selfish gene, Oxford University Press, Oxford, 1976): genes do not and cannot sacrifice their own fitness on behalf of the reproductive population. Therefore, while natural selection explains the appearance of design in the living world (Dawkins in The blind watchmaker: why the evidence of evolution reveals a universe without design, W. W. Norton, (...) New York, 1996), inclusive fitness theory does not explain how. Indeed, Hamilton’s rule is equally compatible with the evolutionary success of prosocial altruistic genes and antisocial predatory genes, whereas only the former, which account for the appearance of design, predominate in successful organisms. Inclusive fitness theory, however, permits a formulation of the central problem of sociobiology in a particularly poignant form: how do interactions among loci induce utterly selfish genes to collaborate, or to predispose their carriers to collaborate, in promoting the fitness of their carriers? Inclusive fitness theory, because it abstracts from synergistic interactions among loci, does not answer this question. Fitness-enhancing collaboration among loci in the genome of a reproductive population requires suppressing alleles that decrease, and promoting alleles that increase the fitness of its carriers. Suppression and promotion are effected by regulatory networks of genes, each of which is itself utterly selfish. This implies that genes, and a fortiori individuals in a social species, do not maximize inclusive fitness but rather interact strategically in complex ways. It is the task of sociobiology to model these complex interactions. (shrink)

The adequacy of currently popular accounts of the genetic basis for psychological altruism, including inclusive fitness (kin selection), reciprocal altruism, sociality, and group selection, is questioned. Problems exist both with the evidence cited as supporting these accounts and with the relevance of the accounts to what is being explained. Based on the empathy-altruism hypothesis, a more plausible account is proposed: generalized parental nurturance. It is suggested that four evolutionary developments combined to provide a genetic (...) basis for psychological altruism. First is the evolution in mammals of parental nurturance. Second is the evolution in humans (and possibly a few other species) of the ability to see others as sentient, intentional agents and, thereby, to recognize other's needs, even subtle ones. Third is the evolution in humans of tender, empathic emotions as an important component of parental nurturance. Fourth is the evolution in humans of cognitive capacities that make it possible to generalize tender, empathic feelings and, thereby, altruism beyond offspring. (shrink)

The biological world is full of phenomena that seem to run counter to Darwin's insight that natural selection can lead to the appearance of design. For instance, why do organisms in some species divide reproductive labor? The existence of non-reproducing organisms in such 'eusocial' species looks to be at odds with an evolutionary theory which posits traits exist because they help organisms survive and reproduce. What is the evolutionary advantage of an insect being distasteful to its predators? The distastefulness (...) appears designed to deter predators, but can only affect the predator's actions when the insect is eaten; it is hard to see how such a trait could be passed on. This Element will cover the shared foundations of evolutionary explanations for these and other seemingly puzzling phenomena, focusing on the concepts of inclusive fitness and kin selection. (shrink)

The Nature of Selection is a straightforward, self-contained introduction to philosophical and biological problems in evolutionary theory. It presents a powerful analysis of the evolutionary concepts of natural selection, fitness, and adaptation and clarifies controversial issues concerning altruism, group selection, and the idea that organisms are survival machines built for the good of the genes that inhabit them. "Sober's is the answering philosophical voice, the voice of a first-rate philosopher and a knowledgeable student of contemporary evolutionary theory. (...) His book merits broad attention among both communities. It should also inspire others to continue the conversation."-Philip Kitcher, Nature "Elliott Sober has made extraordinarily important contributions to our understanding of biological problems in evolutionary biology and causality. The Nature of Selection is a major contribution to understanding epistemological problems in evolutionary theory. I predict that it will have a long lasting place in the literature."-Richard C. Lewontin. (shrink)

The group selection controversy is about whether natural selection ever operates at the level of groups, rather than at the level of individual organisms. Traditionally, group selection has been invoked to explain the existence of altruistic behaviour in nature. However, most contemporary evolutionary biologists are highly sceptical of the hypothesis of group selection, which they regard as biologically implausible and not needed to explain the evolution of altruism anyway. But in their recent book, Elliot Sober (...) and David Sloan Wilson [1998] argue that the widespread opposition to group selection is founded on conceptual confusion. The theories that have been propounded as alternatives to group selection are actually group selection in disguise, they maintain. I examine their arguments for this claim, and John Maynard Smith's arguments against it. I argue that Sober and Wilson arrive at a correct position by faulty reasoning. In the final section, I examine the issue of how to apply the principle of natural selection at different levels of the biological hierarchy, which underlies the dispute between Sober and Wilson and Maynard Smith. (shrink)

Group selection is one acknowledged mechanism for the evolution of altruism. It is well known that for altruism to spread by natural selection, interactions must be correlated; that is, altruists must tend to associate with one another. But does group selection itself require correlated interactions? Two possible arguments for answering this question affirmatively are explored. The first is a bad argument, for it rests on a product/process confusion. The second is a more subtle argument, whose (...) validity (or otherwise) turns on issues concerning the meaning of multi-level selection and how it should be modelled. A cautious defence of the second argument is offered. Introduction Multi-level selection and the evolution of altruism Price's equation and multi-level selection Contextual analysis and multi-level selection The neighbour approach Recapitulation and conclusion. (shrink)

According to the standard formulation, natural selection requires variation, differential fitness, and heritability. I argue that this formulation is inadequate because it fails to distinguish natural selection from artificial selection, intelligent design, forward-looking orthogenetic selection, and adaptation via the selection of nonrandom variation. I suggest adding a _no teleology_ condition. The no teleology condition says that the evolutionary process is not guided toward an endpoint represented in the mind of an agent, variation is produced randomly with respect to (...) adaptation, and selection pressures are not forward looking. (shrink)

This essay addresses recent claims about the compatibility of the sociobiological theory of reciprocal altruism with standard Western formulations of the Golden Rule. Derek Parfit claims that the theory of reciprocal altruism teaches us to be “reciprocal altruists,” who benefit only those people from whom we can reasonably expect benefits in the future. The Golden Rule, on the other hand, teaches us to benefit anyone regardless of their intention or ability to return the favor, or (...) as Parfit puts it, the Golden Rule teaches us to be “suckers.” I argue that this distinction is founded on a misconception of the nature of the theory of reciprocal altruism, which is sociobiological as opposed to moral, and that this distinction accordingly confuses is with ought. Sociobiological theories may explain underlying psychological motivations in individuals (and perhaps even in populations), but these theories do not prescribe any sort of moral behavior. Furthermore, the theory of reciprocal altruism does not imply mental states of which agents are aware. The unconscious motivations assumed by this theory are in fact compatible with certain formulations of the Golden Rule; I will accordingly argue for the view that certain words with moral content related to the Golden Rule—such as “altruism” and “selfishness”—exist only insofar as they are social tools, which can further the self-interests of an individual in any group. (shrink)

This chapter contains section titled: Kin Selection Reciprocity Group Selection Coercion Mutualism Byproduct Mutualism Local Interactions References.

Consider the paradox of altruism: the existence of truly altruistic behaviors is difficult to reconcile with evolutionary theory if natural selection operates only on individuals, since in that case individuals should be unwilling to sacrifice their own fitness for the sake of others. Evolutionists have frequently turned to the hypothesis of group selection to explain the existence of altruism; but group selection cannot explain the evolution of morality, since morality is a one-group phenomenon and (...) group selection is a many-group phenomenon. After spelling out just what the problem is, this paper discusses several ways of solving it. (shrink)

Description of the biologicalhierarchy of the organism has been extendedhere to included the evolutionary andecological sub-hierarchies with theirrespective levels in order to give a completehierarchical description of life. These newdescriptions include direction of formation,types of constraints, and dual levels. Constraints are produced at the macromolecularlevel of genes/proteins, some of which (a) aredescendent restraints which hold a hierarchytogether and others (b) interact horizontallywith selective agents at corresponding levelsof the niche. The organism is a dual levelconstrained by both the ecologicalsub-hierarchy (survival) and (...) evolutionarysub-hierarchy (fitness) while serving as thehighest level of the specializationsub-hierarchy, therefore, it is unique inbelonging to three sub-hierarchies. Organismsexperience birth, sometimes death, andparticipate in evolution. Birth of an organismis realized when a cell is removed from theconstraints of the organism and assumes itsown organismal constraints. Death occurs withthe cessation of protein synthesis becauseproteins and their products constitutecomponents at high levels of the hierarchy. Selection is of two types, natural selection,involving adaptive traits interacting withfeatures in a niche, and hierarchicalselection, requiring traits to be compatiblewith existing hierarchical constraints. (shrink)

Human intentional action, including the design and use of artifacts, involves the prior mental representation of the goal (end) and the means to achieve that goal. This representation is part of the efficient cause of the action, and thus can be used to explain both the action and the achievement of the end. This is intentional teleological explanation. More generally, teleological explanation that depends on the real existence of a representation of the goal (and the means to achieve it) can (...) be called representational teleological explanation. Such explanations in biology can involve both external representations (e.g., ideas in the mind of God) and internal representations (souls, vital powers, entelechies, developmental programs, etc.). However, another type of explanation of intentional action (or any other process) is possible. Given that an action achieving a result occurs, the action can be explained as fulfilling the necessary conditions (means) for that result (end), and, reciprocally, the result explained by the occurrence of those necessary conditions. This is conditional teleological explanation. For organisms, natural selection is often understood metaphorically as the designer, intentionally constructing them for certain ends. Unfortunately, this metaphor is often taken rather too literally, because it has been difficult to conceive of another way to relate natural selection to the process of evolution. I argue that combining a conditional teleological explanation of organisms and of evolution provides such an alternative. This conditional teleology can be grounded in existence or survival. Given that an organism exists, we can explain its existence by the occurrence of the necessary conditions for that existence. This principle of the 'conditions for existence' was introduced by Georges Cuvier in 1800, and provides a valid, conditional teleological method for explaining organismal structure and behavior. From an evolutionary perspective, the conditions for existence are the range of boundary conditions within which the evolutionary process must occur. Moreover, evolutionary change itself can be subjected to conditional teleological explanation, because natural selection theory is primarily a theory about the relation between the conditions for the existence of organisms and the conditions for the existence of traits in populations. I show that failure to distinguish representational from conditional teleological explanation has confused previous attempts to clarify the relation of teleology to biology. (shrink)

The study of biological altruism is a cornerstone of modern evolutionary biology. Associated with foundational issues about natural selection, it is often supposed that explaining altruism is key to understanding social behavior more generally. Typically, biological altruism is defined in purely effects-based, behavioral terms – as an interaction in which one organism contributes fitness to another, at its own expense. Crucially, such a definition isn’t meant to rest on psychological or intentional assumptions. We show that, appearances (...) and official definitions notwithstanding, the notion biological altruism carries a vestige of the psychological, intentional concept familiar to us from the human domain. In particular, definitions of altruism from Hamilton onwards presuppose an actor\recipient distinction – a distinction, so we argue, that has questionable biological grounding. We arrive at this conclusion step-by-step, first looking at several simple, “austere” definitions and their problems, and then critiquing the actor\recipient distinction directly. If successful, our arguments suggest that the category of biological altruism requires a significant rethink. (shrink)

In models of multi-level selection, the property of Darwinian fitness is attributed to entities at more than one level of the biological hierarchy, e.g. individuals and groups. However, the relation between individual and group fitness is a controversial matter. Theorists disagree about whether group fitness should always, or ever, be defined as total (or average) individual fitness. This paper tries to shed light on the issue by drawing on work in social choice theory, and pursuing an analogy between (...) fitness and utility. Social choice theorists have long been interested in the relation between individual and social utility, and have identified conditions under which social utility equals total (or average) individual utility. These ideas are used to shed light on the biological problem. (shrink)

I examine the relationship between evolutionary definitions of altruism that are based on fitness effects and psychological definitions that are based on the motives of the actor. I show that evolutionary altruism can be motivated by proximate mechanisms that are psychologically either altruistic or selfish. I also show that evolutionary definitions do rely upon motives as a metaphor in which the outcome of natural selection is compared to the decisions of a psychologically selfish (or altruistic) individual. Ignoring (...) the precise nature of both psychological and evolutionary definitions has obscured many important issues, including the biological roots of psychological altruism. (shrink)

The concept of fitness has generated a lot of discussion in philosophy of biology. There is, however, relative agreement about the need to distinguish at least two uses of the term: ecological fitness on the one hand, and population genetics fitness on the other. The goal of this paper is to give an explication of the concept of ecological fitness by providing a reconstruction of the theory of natural selection in which this concept was framed, that is, based (...) on the way the theory was put to use in Darwin’s main texts. I will contend that this reconstruction enables us to account for the current use of the theory of natural selection. The framework presupposed in the analysis will be that of metatheoretical structuralism. This framework will provide both a better understanding of the nature of ecological fitness and a more complete reconstruction of the theory. In particular, it will provide what I think is a better way of understanding how the concept of fitness is applied through heterogeneous cases. One of the major advantages of my way of thinking about natural selection theory is that it would not have the peculiar metatheoretical status that it has in other available views. I will argue that in order to achieve these goals it is necessary to make several concepts explicit, concepts that are frequently omitted in usual reconstructions. (shrink)

I argue that images of the notion of group, in correspondence with their social and political values, shape the debate over the evolution of altruism by group selection. Important aspects of this debate are empirical, and criteria can decide among a variety of selection processes. However, leading researchers undermine or reinterpret such tests, explaining the evolution of altruism on the basis of a single extreme metaphor of ‘group’ and a single inclusive selection process. I (...) shall argue that the extreme images for the notion of group are associated with ideologies that these researchers support or fear. Hence, the history of social and political uses of ‘group’ and ‘group selection’ can explain, at least in part, some of the empirical deficiencies of the debate, and why it has continued without resolution or dissolution. (shrink)

Natural selection is commonly seen not just as an explanation for adaptive evolution, but as the inevitable consequence of “heritable variation in fitness among individuals”. Although it remains embedded in biological concepts, such a formalisation makes it tempting to explore whether this precondition may be met not only in life as we know it, but also in other physical systems. This would imply that these systems are subject to natural selection and may perhaps be investigated in a biological (...) framework, where properties are typically examined in light of their putative functions. Here we relate the major questions that were debated during a three-day workshop devoted to discussing whether natural selection may take place in non-living physical systems. We start this report with a brief overview of research fields dealing with “life-like” or “proto-biotic” systems, where mimicking evolution by natural selection in test tubes stands as a major objective. We contend the challenge may be as much conceptual as technical. Taking the problem from a physical angle, we then discuss the framework of dissipative structures. Although life is viewed in this context as a particular case within a larger ensemble of physical phenomena, this approach does not provide general principles from which natural selection can be derived. Turning back to evolutionary biology, we ask to what extent the most general formulations of the necessary conditions or signatures of natural selection may be applicable beyond biology. In our view, such a cross-disciplinary jump is impeded by reliance on individuality as a central yet implicit and loosely defined concept. Overall, these discussions thus lead us to conjecture that understanding, in physico-chemical terms, how individuality emerges and how it can be recognised, will be essential in the search for instances of evolution by natural selection outside of living systems. (shrink)

Through a series of essays, the author discusses the conflict between cultural and biological evolution, covering intelligent design, gender differences, and the meaning of life while offering insight into the ethical aspects of civilization.

It is widely assumed that selection history accounts of function can support a fully reductive naturalization of functional properties. I argue that this assumption is false. A problem with the alternative causal role account of function in this context is that it invokes the teleological notion of a goal in analysing real function. The selection history account, if it is to have reductive status, must not do the same. But attention to certain cases of selection history in biology, specifically those (...) involving meiotic drive, shows that selection historical explanations are available in the case of items without any plausible function. Making contributions to goals, here fitness, must be introduced as an additional constraint in the analysis, either explicitly, or implicitly by appeal to individual functions or the selection-of/selection-for distinction. (shrink)

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

Recent years have seen a renewed debate over the importance of groupselection, especially as it relates to the evolution of altruism. Onefeature of this debate has been disagreement over which kinds ofprocesses should be described in terms of selection at multiple levels,within and between groups. Adapting some earlier discussions, we presenta mathematical framework that can be used to explore the exactrelationships between evolutionary models that do, and those that donot, explicitly recognize biological groups as fitness-bearing entities.We show a fundamental (...) set of mathematical equivalences between these twokinds of models, one of which applies a form of multi-level selectiontheory and the other being a form of ``individualism.'' However, we alsoargue that each type of model can have heuristic advantages over theother. Indeed, it can be positively useful to engage in a kind ofback-and-forth switching between two different perspectives on theevolutionary role of groups. So the position we defend is a``gestalt-switching pluralism.''. (shrink)

Both biologists and philosophers often make use of simple verbal formulations of necessary and sufficient conditions for evolution by natural selection (ENS). Such summaries go back to Darwin's Origin of Species (especially the "Recapitulation"), but recent ones are more compact.1 Perhaps the most commonly cited formulation is due to Lewontin.2 These summaries tend to have three or four conditions, where the core requirement is a combination of variation, heredity, and fitness differences. The summaries are employed in several ways. First, (...) they are often used in pedagogical contexts, and in showing the coherence of evolutionary theory in response to attacks from outside biology. Second, they are important in discussions of extensions of evolutionary principles to new domains, such as cultural change. The summaries also have intrinsic scientific and philosophical interest as attempts to capture some core principles of evolutionary theory in a highly concise way. Despite their prominence, both the proper formulation and status of these summaries are unclear. Standard formulations are subject to counterexamples, and their relations to formal models of evolutionary change are not straightforward. Here I look closely at these verbal summaries, and at how they relate to formal models. Are the summaries merely rough approximations that have no theoretical role of their own? Perhaps they could operate as theoretical statements in Darwin's time, but have now been superseded by more exact treatments. (shrink)

I first argue against Peter Singer's exciting thesis that the Prisoner's Dilemma explains why there could be an evolutionary advantage in making reciprocal exchanges that are ultimately motivated by genuine altruism over making such exchanges on the basis of enlightened long-term self-interest. I then show that an alternative to Singer's thesis — one that is also meant to corroborate the view that natural selection favors genuine altruism, recently defended by Gregory Kavka, fails as well. Finally, I (...) show that even granting Singer's and Kavka's claim about the selective advantage of altruism proper, it is doubtful whether that type of claim can be used in a particular sort of sociobiological argument against psychological egoism. (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)

What are the prospects for a monistic view of biological individuality given the multiple epistemic roles the concept must satisfy? In this paper, I examine the epistemic adequacy of two recent accounts based on the capacity to undergo natural selection. One is from Ellen Clarke, and the other is by Peter Godfrey-Smith. Clarke’s position reflects a strong monism, in that she aims to characterize individuality in purely functional terms and refrains from privileging any specific material properties as important in (...) their own right. I argue that Clarke’s functionalism impairs the epistemic adequacy of her account compared to a middle-ground position taken by Godfrey-Smith. In comparing Clarke and Godfrey-Smith’s account, two pathways emerge to pluralism about biological individuality. The first develops from the contrast between functionalist and materialist approaches, and the second from an underlying temporal structure involved in using evolutionary processes to define individuality. (shrink)

We argue that a fashionable interpretation of the theory of natural selection as a claim exclusively about populations is mistaken. The interpretation rests on adopting an analysis of fitness as a probabilistic propensity which cannot be substantiated, draws parallels with thermodynamics which are without foundations, and fails to do justice to the fundamental distinction between drift and selection. This distinction requires a notion of fitness as a pairwise comparison between individuals taken two at a time, and so vitiates the (...) interpretation of the theory as one about populations exclusively. (shrink)

What explains the distinctive features of human behavior? In this book, Stewart-Williams aims to answer this ambitious question. This book is an engaging addition to the already long list of recent attempts to provide an evolutionary explanation of human uniqueness. It is organized into six chapters, plus two appendices. These chapters address several key topics in evolutionary theory, sex differences and sexual behavior, altruism, and cultural evolution, albeit with varying degrees of detail and depth. These topics include sexual selection, (...) kin selection, Hamilton’s rule, reciprocal altruism, costly signaling theory, group selection, gene-centered views of evolution, inclusive fitness, proximate and ultimate evolutionary explanations, inbreeding avoidance, the Westermarck effect, jealousy, sperm competition, mating and parenting effort, cumulative cultural evolution, imitation and learning biases, evolutionary mismatch theories, and more. (shrink)

Multi-level selection can be understood via the Price equation or contextual analysis, which offer incompatible statistical decompositions of evolutionary change into components of group and individual selection. Okasha argued that each approach suffers from problem cases. I introduce further problem cases for the Price approach, arguing that it is appropriate for MLS 2 group selection but not MLS 1. I also show that the problem cases Okasha raises for contextual analysis can be resolved. For some such cases, however, (...) it emerges that there is no determinate answer to the question of how much of the total selective effect was due to group selection compared to individual selection. This suggests that when there is interaction between the effect of group character and individual character, one cannot separate selection into distinct ‘levels’ at all. (shrink)

For evolution by natural selection to occur it is classically admitted that the three ingredients of variation, difference in fitness and heredity are necessary and sufficient. In this paper, I show using simple individual-based models, that evolution by natural selection can occur in populations of entities in which neither heredity nor reproduction are present. Furthermore, I demonstrate by complexifying these models that both reproduction and heredity are predictable Darwinian products (i.e. complex adaptations) of populations initially lacking these two (...) properties but in which new variation is introduced via mutations. Later on, I show that replicators are not necessary for evolution by natural selection, but rather the ultimate product of such processes of adaptation. Finally, I assess the value of these models in three relevant domains for Darwinian evolution. (shrink)

The debate between the dynamical and the statistical interpretations of natural selection is centred on the question of whether all explanations that employ the concepts of natural selection and drift are reducible to causal explanations. The proponents of the statistical interpretation answer negatively, but insist on the fact that selection/drift arguments are explanatory. However, they remain unclear on where the explanatory power comes from. The proponents of the dynamical interpretation answer positively and try to reduce selection/drift arguments to (...) some of the most prominent accounts of causal explanation. In turn, they face the criticism raised by statisticalists that current accounts of causation have to be violated in some of their core conditions or otherwise used in a very loose manner in order to account for selection/drift explanations. We propose a reconciliation of both interpretations by conveying evolutionary explanations within the unificationist model of scientific explanation. Therefore, we argue that the explanatory power in natural selection arguments is a result of successful unification of individual- and population-level facts. A short case study based on research on sympatric speciation will be presented as an example of how population- and individual-level facts are unified to explain the morphological mosaic of bill shape in island scrub jays. (shrink)

This paper sets out asking what is to be gained from grounding the pursuit of a cosmopolitan morality in the evolutionary history of our morals, namely, by ascertaining some of the natural constraints under which normative ethical theory must operate. In Section II, I review two major forms of altruism: kin-based and reciprocal altruism. Experimental evidence is cited to support the view that biological altruism involves a carefully self-interested calculation to enhance adaptive fitness to the (...) social environment. In Section III, I explain that, in natural selection, these self-interested calculations about relatedness and reciprocation don’t happen consciously, but rather, almost instantaneously, as heuristics. These heuristics are advantageous to the organism and thus, over time, hardwired to its ‘emotional framework’. Emotional contagion, guilt, envy, and inequity aversion are covered as examples of the emotions at work in altruistic behavior. What can ethical deliberation on cosmopolitan morality adopt from a naturalistic examination of the evolution of altruism? According to situationists, what we do is best explained by inherent tendencies to respond to features of our situation. The pursuit of cosmopolitan morality via the evolutionary history of altruism reiterates this situationist conclusion, and seems to carry a recommendation for polity: to adjust our environment in response to findings about human nature as to better correspond with our proposed moral ends. (shrink)

The selected effects or ‘etiological’ theory of Proper function is a naturalistic and realist account of biological teleology. It is used to analyse normativity in philosophy of language, philosophy of mind, philosophy of medicine and elsewhere. The theory has been developed with a simple and intuitive view of natural selection. Traits are selected because of their positive effects on the fitness of the organisms that have them. These ‘selected effects’ are the Proper functions of the traits. (...) Proponents argue that this analysis of biological teleology has the unique advantage that the selected effect function of a trait is also a causal explanation of the trait: the trait exists because it performs this function. We show, however, that selected effect functions as currently defined explain the existence of traits only under highly restrictive assumptions about evolutionary dynamics. In many common scenarios in which traits evolve by natural selection, selected effect functions do not explain those traits. This is because definitions of selected effect function extract from any evolutionary scenario only the information that would be explanatorily relevant in the simple evolutionary scenario implicit in those definitions. When applied to more complex scenarios selected effect functions omit the key information that is explanatorily relevant in those scenarios. The assumptions required for selected effect functions to be explanatory are particularly unlikely to hold in the domain that its proponents care most about - the evolution of representation. A more adequate selected effects theory of Proper functions may be possible, but will require much greater attention to the structure of actual evolutionary explanations. (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)

A number of recent discussions have argued that George Price's equationfor representing evolutionary change is a powerful and illuminatingtool, especially in the context of debates about multiple levels ofselection. Our paper dissects Price's equation in detail, and comparesit to another statistical tool: the calculation and comparison ofaverage fitnesses. The relations between Price's equation and equationsfor evolutionary change using average fitness are closer than issometimes supposed. The two approaches achieve a similar kind ofstatistical summary of one generation of change, and they (...) achieve thisvia a similar loss of information about the underlying fitnessstructure. (shrink)

Unto Others (Sober and Wilson 1998) shows how the general principles of Multi-Level Selection (MLS) theory apply to selection at multiple levels of the biological hierarchy. It also argues for the existence of "genuine" evolutionary and psychological altruism. The authors’ views on altruism do not follow logically from principles of MLS, and their failure do disentangle these two themes undermines their otherwise excellent presentation of MLS theory. Rebuttal of the view that human nature is completely selfish depends not (...) on the prevalence of altruism but on the importance of group-advantageous traits that benefit both self and other group members without necessarily inflicting direct costs on outsiders. (shrink)

The selected effects or ‘etiological’ theory of Proper function is a naturalistic and realist account of biological teleology. It is used to analyse normativity in philosophy of language, philosophy of mind, philosophy of medicine, and elsewhere. The theory has been developed with a simple and intuitive view of natural selection. Traits are selected because of their positive effects on the fitness of the organisms that have them. These ‘selected effects’ are the Proper functions of the traits. (...) Proponents argue that this analysis of biological teleology has the unique advantage that the selected effect function of a trait is also a causal explanation of the trait: the trait exists because it performs this function. We show, however, that selected effect functions as currently defined explain the existence of traits only under highly restrictive assumptions about evolutionary dynamics. In many common scenarios in which traits evolve by natural selection, selected effect functions do not explain those traits. This is because definitions of selected effect function extract from any evolutionary scenario only the information that would be explanatorily relevant in the simple evolutionary scenario implicit in those definitions. When applied to more complex scenarios, selected effect functions omit the key information that is explanatorily relevant in those scenarios. The assumptions required for selected effect functions to be explanatory are particularly unlikely to hold in the domain that its proponents care most about—the evolution of representation. A more adequate selected effects theory of Proper functions may be possible, but will require much greater attention to the structure of actual evolutionary explanations. (shrink)