Human knowledge is a phenomenon whose roots extend from the cultural, through the neural and the biological and finally all the way down into the Precambrian “primordial soup.” The present paper reports an attempt at understanding this Greater System of Knowledge (GSK) as a hierarchical nested set of selection processes acting concurrently on several different scales of time and space. To this end, a general selection theory extending mainly from the work of Hull and Campbell is introduced. The perhaps most (...) drastic change from previous similar theories is that replication is revealed as a composite function consisting of what is referred to as memory and synthesis. This move is argued to drastically improve the fit between theory and human-related knowledge systems. The introduced theory is then used to interpret the subsystems of the GSK and their interrelations. This is done to the end of demonstrating some of the new perspectives offered by this view. (shrink)

Mesoudi et al. argue that the current inability to identify the means by which cultural traits are acquired does not debilitate their project to draw clear parallels between cultural and biological evolution. However, I suggest that cultural phenomena may be accounted for by biological processes, unless we can identify a cultural “genotype” that carries information from person to person independently of genes. (Published Online November 9 2006).

In their 2010 book, Biology’s First Law, D. McShea and R. Brandon present a principle that they call ‘‘ZFEL,’’ the zero force evolutionary law. ZFEL says (roughly) that when there are no evolutionary forces acting on a population, the population’s complexity (i.e., how diverse its member organisms are) will increase. Here we develop criticisms of ZFEL and describe a different law of evolution; it says that diversity and complexity do not change when there are no evolutionary causes.

If temperature does not vary from one generation from to the next but its value is crucial for the development of particular phenotypic characteristics, a long-term change in its value may trigger major evolutionary changes of the organism. If a bird's nest maintains the critical temperature, then a statement that the bird is the nest's way of making another nest is as helpful as accounts couched in terms of genes' intentions. However, the language of intentions rests on different evidence and (...) assumptions from causal language and the languages are not interchangeable. Understanding ontogeny in causal terms requires explanations that are made as simple as possible but not so simple that they become completely unrealistic. (shrink)

When one takes the evolution of operant behavior as prototype, one sees that the term replication is too tied to the peculiarities of genetic evolution. A more general term is recurrence. The important problem raised by recurrence is not “information” but relationship: deciding when two occurrences belong to the same lineage. That is solved by looking at common environmental effects.

Ernst Mayr''s distinction between ultimate and proximate causes is justly considered a major contribution to philosophy of biology. But how did Mayr come to this philosophical distinction, and what role did it play in his earlier scientific work? I address these issues by dividing Mayr''s work into three careers or phases: 1) Mayr the naturalist/researcher, 2) Mayr the representative of and spokesman for evolutionary biology and systematics, and more recently 3) Mayr the historian and philosopher of biology. If we want (...) to understand the role of the proximate/ultimate distinction in Mayr''s more recent career as a philosopher and historian, then it helps to consider hisearlier use of the distinction, in the course of his research, and in his promotion of the professions of evolutionary biology and systematics. I believe that this approach would also shed light on some other important philosophical positions that Mayr has defended, including the distinction between essentialism: and population thinking. (shrink)

Richard Lewontin's (1970) early work on the units of selection initiated the conceptual and theoretical investigations that have led to the hierarchical perspective on selection that has reached near consensus status today. This paper explores other aspects of his work, work on what he termed continuity and quasi-independence, that connect to contemporary explorations of modularity in development and evolution. I characterize such modules and argue that they are the true units of selection in that they are what evolution by natural (...) selection individuates, selects among, and transforms. (shrink)

Sober (1992) has recently evaluated Brandon's (1982, 1990; see also 1985, 1988) use of Salmon's (1971) concept of screening-off in the philosophy of biology. He critiques three particular issues, each of which will be considered in this discussion.

The question "what is (are) the unit(s) of selection" can be interpreted in three different ways. These interpretations are discussed and it is shown that they prompt different answers; such units are shown to be individuals in the context of the given interpretation. One of these interpretations is argued, by examples, not always to have an unambiguously correct answer. An alternative approach to this question is sketched.

Sober and Lewontin's critique of genic selectionism is based upon the principle that a unit of selection should make a context‐independent contribution to fitness. Critics have effectively shown that this principle is flawed. In this paper I show that the context independence principle is an instance of a more general principle for characterizing causes,called the contextual unanimity principle. I argue that this latter principle, while widely accepted, is erroneous. What is needed is to replace the approach to causality characterized by (...) the contextual unanimity criterion with an approach based on the concept of causal mechanism. After sketching such an approach, I show how it can be used to shed light on the units of selection problem. (shrink)

Group-structured populations, of the kind prominent in discussions of multilevel selection, are contrasted with ‘neighbor-structured’ populations. I argue that it is a necessary condition on multilevel description of a selection process that there should be a nonarbitrary division of the population into equivalence classes (or an approximation to this situation). The discussion is focused via comparisons between two famous problem cases involving group structure (altruism and heterozygote advantage) and two neighbor-structured cases that resemble them. Conclusions are also drawn about the (...) role of correlated interaction in the evolution of altruism. 1 Introduction 2 Two Kinds of Population Structure 3 Objections and Replies 4 Particles on a Line 5 Conclusion Appendix: Neighborhoods and Selection CiteULike Connotea Del.icio.us What's this? (shrink)

A set of constraints forces trade-offs which prevent us from achieving the best possible definitions of the ‘level’ and ‘unit’ of natural selection. This set consists in decisions concerning conflicting pre-analytic intuitions in problematic cases, the relative roles of various conceptual resources in the definitions, which facts need to be accounted for using the definitions, how the relation between selection and evolution orients the definitions, and the relation between the level and unit concepts. Systematic reconstruction and evaluation of leading analyses (...) along these dimensions favors a new functional analysis over Williams’ consequentialist analysis, Sober’s causal analysis, and Dawkins’ teleological analysis. (shrink)

The theory of evolution is supported by the theory of genetics, which provides a single causal mechanism to explain the activities of replicators and interactors. A common misrepresentation of the theory of evolution, however, is that interaction (involving interactors), and transmission (involving replicators), are distinct causal processes. Sandra Mitchell (1987) is misled by this. I discuss why only a single causal mechanism is working in evolution and why it is sufficient. Further, I argue that Mitchell's mistaken view of the causal (...) mechanism in evolution prevents her from resolving the conflict between Dawkins and Brandon. I conclude that the unit-of-selection question remains very much alive. (shrink)

The theory of evolution by natural selection is, perhaps, the crowning intellectual achievement of the biological sciences. There is, however, considerable debate about which entity or entities are selected and what it is that fits them for that role. This article aims to clarify what is at issue in these debates by identifying four distinct, though often confused, concerns and then identifying how the debates on what constitute the units of selection depend to a significant degree on which of these (...) four questions a thinker regards as central. (shrink)

David Hull's analysis of conceptual change in science, as presentedin his book, Science as a Process (1988), provides a useful framework for understanding one of the scientific controversies in which he actively and constructively intervened, the units of selectiondebates in evolutionary biology. What follows is a brief overview ofthose debates and some reflections on them.

The conflation of two fundamentally distinct issues has generated serious confusion in the philosophical and biological literature concerning the units of selection. The question of how a unit of selection of defined, theoretically, is rarely distinguished from the question of how to determine the empirical accuracy of claims--either specific or general--concerning which unit(s) is undergoing selection processes. In this paper, I begin by refining a definition of the unit of selection, first presented in the philosophical literature by William Wimsatt, which (...) is grounded in the structure of natural selection models. I then explore the implications of this structural definition for empirical evaluation of claims about units of selection. I consider criticisms of this view presented by Elliott Sober--criticisms taken by some (for example, Mayo and Gilinsky 1987) to provide definitive damage to the structuralist account. I shall show that Sober has misinterpreted the structuralist views; he knocks down a straw man in order to motivate his own causal account. Furthermore, I shall argue, Sober's causal account is dependent on the structuralist account that he rejects. I conclude by indicating how the refined structural definition can clarify which sorts of empirical evidence could be brought to bear on a controversial case involving units of selection. (shrink)

In The Levels of Selection (Brandon, 1984), Robert Brandon provides a suggestive but ultimately unsuccessful attempt to use the probabilistic notion ofscreening off in providing a schema for dealing with an aspect of the units of selection question in the philosophy of biology. I characterize that failure, and suggest a revision and expansion of Brandon's account which addresses its key shortcoming.

The controversy regarding the unit of selection is fundamentally a dispute about what is the correct causal structure of the process of evolution by natural selection and its ontological commitments. By characterizing the process as consisting of two essential steps--interaction and transmission--a singular answer to the unit question becomes ambiguous. With such an account on hand, two recent defenses of competing units of selection are considered. Richard Dawkins maintains that the gene is the appropriate unit of selection and Robert Brandon, (...) in response, argues that the individual organism is better suited to the role. This paper argues that by making explicit the underlying questions that each of these views addresses, the apparent conflict can be resolved. Furthermore, such a resolution allows for a more complete and realistic understanding of the process of evolution by natural selection. (shrink)

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.

The levels of selection problem was central to Maynard Smith’s work throughout his career. This paper traces Maynard Smith’s views on the levels of selection, from his objections to group selection in the 1960s to his concern with the major evolutionary transitions in the 1990s. The relations between Maynard Smith’s position and those of Hamilton and G.C. Williams are explored, as is Maynard Smith’s dislike of the Price equation approach to multi-level selection. Maynard Smith’s account of the ‘core Darwinian principles’ (...) is discussed, as is his debate with Sober and Wilson (1998) over the status of trait-group models, and his attitude to the currently fashionable concept of pluralism about the levels of selection. (shrink)

This paper compares two well-known arguments in the units of selection literature, one due to , the other due to . Both arguments concern the legitimacy of averaging fitness values across contexts and making inferences about the level of selection on that basis. The first three sections of the paper shows that the two arguments are incompatible if taken at face value, their apparent similarity notwithstanding. If we accept Sober and Lewontin's criterion for when averaging genic fitnesses across diploid genotypes (...) is illegitmate, we cannot accept Sober and Wilson's criterion for when averaging individual fitnesses across groups is illegitimate, and vice versa. The final section suggests a possible way of reconciling the two arguments, by invoking an ambiguity in the concept of genic selection. (shrink)

Activity anorexia illustrates selection of behavior at the biological, behavioral, and neural levels. Based on evolutionary history, food depletion increases the reinforcement value of physical activity that, in turn, decreases the reinforcement effectiveness of eating – resulting in activity anorexia. Neural opiates participate in the selection of physical activity during periods of food depletion.

The debate about the levels of selection has been one of the most controversial both in evolutionary biology and in philosophy of science. Okasha’s book makes the sort of contribution that simply will not be able to be ignored by anyone interested in this field for many years to come. However, my interest here is in highlighting some examples of how Okasha goes about discussing his material to suggest that his book is part of an increasingly interesting trend that sees (...) scientists and philosophers coming together to build a broadened concept of “theory” through a combination of standard mathematical treatments and conceptual analyses. Given the often contentious history of the relationship between philosophy and science, such trend cannot but be welcome. (shrink)

The debate about the levels of selection has been one of the most controversial both in evolutionary biology and in philosophy of science. Okasha’s book makes the sort of contribution that simply will not be able to be ignored by anyone interested in this field for many years to come. However, my interest here is in highlighting some examples of how Okasha goes about discussing his material to suggest that his book is part of an increasingly interesting trend that sees (...) scientists and philosophers coming together to build a broadened concept of “theory” through a combination of standard mathematical treatments and conceptual analyses. Given the often contentious history of the relationship between philosophy and science, such trend cannot but be welcome. (shrink)

Individual and group selection are usually conceived as opposed evolutionary processes. Though cases of synergy are occasionally recognized, the evolutionary importance of synergy is largely ignored. However, synergy is the plausible explanation for the evolution of collectives as higher level individuals i.e., collectives acting as adaptive units, e.g., genomes and colonies of social insects. It rests on the suppression of the predictable tendency of evolutionary units to benefit at the expense of other units or of the wholes they contribute to (...) build. It plausibly explains human cooperation and morality: the molding of human groups into adaptive units. (shrink)

Natural selection is an important force that shapes the evolution of all living things by determining which individuals contribute the most descendents to future generations. The biological unit upon which selection acts has been the subject of serious debate, with reasonable arguments made on behalf of populations, individuals, individual phenotypic characters and, finally, individual genes themselves. In this essay, I argue that the usual unit of selection is the gene. There are powerful logical arguments in favor of this conclusion, as (...) well as many real-world examples. I also explore the possibility that epigenetic differences between individuals may be heritable between generations. Although few such examples exist, epigenetic differences provide an exciting source of potentially heritable variation that may allow rapid evolutionary change to occur, perhaps in response to environmental influences. (shrink)

On the basis of distinctions between those properties of entities that can be defined without reference to other entities and those that (in different ways) cannot, this note argues that non-trivial forms of frequency-dependent selection of entities should be interpreted as selection occurring at a level higher than that of those entities. It points out that, except in degenerately simple cases, evolutionary game-theoretic models of selection are not models of individual selection. Similarly, models of genotypic selection such as heterosis cannot (...) be legitimately interpreted as models of genic selection. The analysis presented here supports the views that: (i) selection should be viewed as a multi-level process; (ii) upper-level selection is ubiquitous; (iii) kin selection should be viewed as a type of group selection rather than individual selection; and (iv) inclusive fitness is not an individual property. (shrink)

In an important article, Kim Sterelny and Philip Kitcher (1988) challenge the common assumption that for any biological phenomenon requiring a selectionist explanation, it is possible to identify a uniquely correct account of the relevant selection process. They argue that selection events can be modeled in any of a number of different, equally correct ways. They call their view 'Pluralism,' and explicitly connect it with various antirealist positions in the philosophy of science. I critically evaluate Sterelny and Kitcher's Pluralism along (...) with its attendant antirealist theses. In particular, I argue that there are serious problems with their pluralistic antirealism regarding units of selection. By correctly diagnosing these problems a more adequate position can be constructed. I defend such a position, which I designate Inclusive Hierarchical Monism, and show how it captures the important virtues of Sterelny and Kitcher's approach while avoiding its problems. (shrink)

In recent years philosophers have attempted to clarify the units of selection controversy in evolutionary biology by offering conceptual analyses of the term 'unit of selection'. A common feature of many of these analyses is an emphasis on the claim that units of selection are entities exhibiting heritable variation in fitness. In this paper I argue that the demand that units of selection be characterized in terms of heritability is unnecessary, as well as undesirable, on historical, theoretical, and philosophical grounds. (...) I propose a positive account of the proper referent of the term 'unit of selection', distinguishing between the processes of evolution and phenotypic selection. The main result of this analysis is greater clarity about the conceptual structure of evolutionary theory. (shrink)

I discuss two subjects in Samir Okasha’s excellent book, Evolution and the Levels of Selection. In consonance with Okasha’s critique of the conventionalist view of the units of selection problem, I argue that conventionalists have not attended to what realists mean by group, individual, and genic selection. In connection with Okasha’s discussion of the Price equation and contextual analysis, I discuss whether the existence of these two quantitative frameworks is a challenge to realism.

Brandon ([1982] 1984, 1990) has argued that Salmon's (1971) concept of screening-off can be used to characterize (i) the idea that natural selection acts directly on an organism's phenotype, only indirectly on its genotype, and (ii) the biological problem of the levels of selection. Brandon also suggests (iii) that screening-off events in a causal chain are better explanations than the events they screen off. This paper critically evaluates Brandon's proposals.

The evolutionary problem of the units of selection has elicited a good deal of conceptual work from philosophers. We review this work to determine where the issues now stand.

Genic selectionism holds that all selection can be understood as operating on particular genes. Critics (and conventional biological wisdom) insist that this misrepresents the actual causal structure of selective phenomena at higher levels of biological organization, but cannot convincingly defend this intuition. I argue that the real failing of genic selectionism is pragmatic – it prevents us from adopting the most efficient corpus of causal laws for predicting and intervening in the course of affairs – and I offer a Pragmatic (...) account of causation itself which ultimately bears out the claim that genic selectionism misrepresents the causal structure of selective contexts. (shrink)

The Units of Selection debate is a dispute about the causes of population change. I argue that it is generated by a particular `dynamical'' interpretation of natural selection theory, according to which natural selection causes differential survival and reproduction of individuals and natural selection explanations cite these causes. I argue that the dynamical interpretation is mistaken and offer in outline an alternative, `statistical'' interpretation, according to which natural selection theory is a fancy kind of `bookkeeping''. It explains by citing the (...) statistical structure of a population and not by citing the causes of survival and reproduction. From the perspective of the statistical interpretation there is no substantive Units of Selection issue. (shrink)

A correct analysis of hierarchical selection processes must specify 1) the objects that succeed differentially as units, and 2) the properties that provide the causal bases for differential success. Here I illustrate how failing to recognize the units/bases distinction creates a contradiction in Elliott Sober's recent account of selection. A revised criterion for units of selection is developed and applied to examples at several biological levels. Criteria for bases of selection are discussed in terms of the degree of context-dependence and (...) directness of a property's effect on the success of units. The significance of previous work by Sober, Wimsatt and Brandon is thereby clarified. (shrink)

This paper distinguishes and critiques several forms of pluralism about the levels of selection, and introduces a novel way of thinking about the biological properties and processes typically conceptualized in terms of distinct levels. In particular, "levels" should be thought of as being entwined or fused. Since the pluralism discussed is held by divergent theorists, the argument has implications for many positions in the debate over the units of selection. And since the key points on which the paper turns apply (...) beyond this specific issue, the paper may prove of general interest in thinking about the metaphysics of science. (shrink)

I investigate how theoretical assumptions, pertinent to different perspectives and operative during the modeling process, are central in determining how nature is actually taken to be. I explore two different models by Michael Turelli and Steve Frank of the evolution of parasite-mediated cytoplasmic incompatility, guided, respectively, by Fisherian and Wrightian perspectives. Since the two models can be shown to be commensurable both with respect to mathematics and data, I argue that the differences between them in the (1) mathematical presentation of (...) the models, (2) explanations, and (3) objectified ontologies stem neither from differences in mathematical method nor the employed data, but from differences in the theoretical assumptions, especially regarding ontology, already present in the respective perspectives. I use my "set up, mathematically manipulate, explain, and objectify" (SMEO) account of the modeling process to track the model-mediated imposition of theoretical assumptions. I conclude with a discussion of the general implications of my analysis of these models for the controversy between Fisherian and Wrightian perspectives. (shrink)

The application of phylogenetic methods to cultural variation raises questions about how cultural adaption works and how it is coupled to cultural transmission. Cultural group selection is of particular interest in this context because it depends on the same kinds of mechanisms that lead to tree-like patterns of cultural variation. Here, we review ideas about cultural group selection relevant to cultural phylogenetics. We discuss why group selection among multiple equilibria is not subject to the usual criticisms directed at group selection, (...) why multiple equilibria are a common phenomena, and why selection among multiple equilibria is not likely to be an important force in genetic evolution. We also discuss three forms of group competition and the processes that cause populations to shift from one equilibrium to another and create a mutation-like process at the group level. (shrink)

Consider the paradox of altruism: the existence of truly altruistic behaviors is difficult to reconcile with an evolutionary theory which holds that 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, even setting aside difficulties about understanding the relationship between altruistic behaviors and morality, 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 out and concludes by offering suggestions why one seems best. (shrink)

This article is primarily a study of the group selection controversy, with special emphasis on the period from 1962 to the present, and the rise of inclusive fitness theory. Interest is focused on the relations between individual fitness theory and other fitness theories and on the methodological imperatives used in the controversy over the status of these theories. An appendix formalizes the notion of "assertive part" which is used in the informal discussion of the methodological imperatives elicited from the controversy.

argues that correlated interactions are necessary for group selection. His argument turns on a particular procedure for measuring the strength of selection, and employs a restricted conception of correlated interaction. It is here shown that the procedure in question is unreliable, and that while related procedures are reliable in special contexts, they do not require correlated interactions for group selection to occur. It is also shown that none of these procedures, all of which employ partial regression methods, are reliable when (...) correlated interactions of a specific kind arise, and it is argued that such correlated interactions will likely be ubiquitous in natural populations. Introduction Process and Product Fitness, Mean Fitness, and Phenotypic Change Correlated Interactions Causation Implications CiteULike Connotea Del.icio.us What's this? (shrink)

We develop an account of laboratory models, which have been central to the group selection controversy. We compare arguments for group selection in nature with Darwin's arguments for natural selection to argue that laboratory models provide important grounds for causal claims about selection. Biologists get information about causes and cause-effect relationships in the laboratory because of the special role their own causal agency plays there. They can also get information about patterns of effects and antecedent conditions in nature. But to (...) argue that some cause is actually responsible in nature, they require an inference from knowledge of causes in the laboratory context and of effects in the natural context. This process, cause detection, forms the core of an analogical argument for group selection. We discuss the differing roles of mathematical and laboratory models in constructing selective explanations at the group level and apply our discussion to the units of selection controversy to distinguish between the related problems of cause determination and evaluation of evidence. Because laboratory models are at the intersection of the two problems, their study is crucial for framing a coherent theory of explanation for evolutionary biology. (shrink)

Groups, individuals, and evolutionary restraints : the making of the contemporary debate over group selection Content Type Journal Article Pages 1-14 DOI 10.1007/s10539-011-9255-5 Authors Andrew Hamilton, Center for Biology and Society, School of Life Sciences, Arizona State University, Tempe, AZ 85287-4501 USA Christopher C. Dimond, Center for Biology and Society, School of Life Sciences, Arizona State University, Tempe, AZ 85287-4501 USA Journal Biology and Philosophy Online ISSN 1572-8404 Print ISSN 0169-3867.

We consider the Stag Hunt in terms of Maynard Smith’s famous Haystack model. In the Stag Hunt, contrary to the Prisoner’s Dilemma, there is a cooperative equilibrium besides the equilibrium where every player defects. This implies that in the Haystack model, where a population is partitioned into groups, groups playing the cooperative equilibrium tend to grow faster than those at the non-cooperative equilibrium. We determine under what conditions this leads to the takeover of the population by cooperators. Moreover, we compare (...) our results to the case of an unstructured population and to the case of the Prisoner’s Dilemma. Finally, we point to some implications our findings have for three distinct ideas: Ken Binmore’s group selection argument in favor of the evolution of efficient social contracts, Sewall Wright’s Shifting Balance theory, and the equilibrium selection problem of game theory. (shrink)

Abstract. Scientists have long puzzled over how homosexual orientation has evolved, given the assumed low relative fitness of homosexual individuals compared to heterosexual individuals. A number of theoretical models for the evolution of homosexuality have been postulated including balance polymorphism, "Fertile females", hypervariability of DNA sequences, kin selection, and "parental manipulation". In this paper, I propose a new group-selection model for the evolution of homosexuality which offers two advantages over existing models: (1) its non-assumption of genetic determinism, and (2) its (...) lack of dependency on an inefficient altruism relation and family dynamics theory. (shrink)

The capacity to engage with art is a human universal present in all cultures and just about every individual human. This indicates that this capacity is evolved. In this Critical Notice of Denis Dutton's The Art Instinct, I discuss various evolutionary scenarios and their consequences. Dutton and I both reject the "spandrel" approach that originates from the work of Gould and Lewontin. Dutton proposes, following work of Geoffrey Miller, that art is sexually selected--that art-production is a sign of a fit (...) genome in males. I argue that while assortative mating may well have had a role in the evolution of "the art instinct", group selection is a better explanation. I also take issue with Dutton's "cluster concept" approach to defining art, and argue that it is a universal and essential characteristic of art that it is appreciated both for what it expresses and for the way that it expresses. It thus requires a reflexive capacity that is not operative in the appreciation of sport spectacles and pornography. (shrink)

The key problem in the controversy over group selection is that of defining a criterion of group selection that identifies a distinct causal process that is irreducible to the causal process of individual selection. We aim to clarify this problem and to formulate an adequate model of irreducible group selection. We distinguish two types of group selection models, labeling them type I and type II models. Type I models are invoked to explain differences among groups in their respective rates of (...) production of contained individuals. Type II models are invoked to explain differences among groups in their respective rates of production of distinct new groups. Taking Elliott Sober's model as an exemplar, we argue that although type I models have some biological importance--they force biologists to consider the role of group properties in influencing the fitness of organisms--they fail to identify a distinct group-level causal selection process. Type II models if properly framed, however, do identify a group-level causal selection process that is not reducible to individual selection. We propose such a type II model and apply it to some of the major candidates for group selection. (shrink)

The aim of this paper is to make an unlikely connection between the old question about the meaning of life and some important concepts in philosophy of biology. More precisely, I argue that while biology is unlikely to help us to figure out the meaning of life, the fact that this question has been considered to be such a crucial one could be explained with the help of some consideration of our evolutionary past. I argue that if there is evidence (...) for group selection in the course of human evolution, this may explain not the meaning of life but rather the reason why we are preoccupied with this question. First, I examine what group selection is and what role it played in human evolution. After surveying the evidence for the claim that in the course of human evolution we lived in isolated group societies, I analyse what influence this social structure had on our present psychological dispositions, including our quest for the meaning of life. (shrink)

The aim of this paper is to make an unlikely connection between the old question about the meaning of life and some important concepts in philosophy of biology. More precisely, I argue that while biology is unlikely to help us to figure out the meaning of life, the fact that this question has been considered to be such a crucial one could be explained with the help of some consideration of our evolutionary past. I argue that if there is evidence (...) for group selection in the course of human evolution, this may explain not the meaning of life but rather the reason why we are preoccupied with this question. First, I examine what group selection is and what role it played in human evolution. After surveying the evidence for the claim that in the course of human evolution we lived in isolated group societies, I analyse what influence this social structure had on our present psychological dispositions, including our quest for the meaning of life. (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)

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)

We consider the question: under what circumstances can the concept of adaptation be applied to groups, rather than individuals? Gardner and Grafen (2009, J. Evol. Biol.22: 659–671) develop a novel approach to this question, building on Grafen's ‘formal Darwinism’ project, which defines adaptation in terms of links between evolutionary dynamics and optimization. They conclude that only clonal groups, and to a lesser extent groups in which reproductive competition is repressed, can be considered as adaptive units. We re-examine the conditions under (...) which the selection–optimization links hold at the group level. We focus on an important distinction between two ways of understanding the links, which have different implications regarding group adaptationism. We show how the formal Darwinism approach can be reconciled with G.C. Williams’ famous analysis of group adaptation, and we consider the relationships between group adaptation, the Price equation approach to multi-level selection, and the alternative approach based on contextual analysis. (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)

Within a laboratory experiment we investigate a principal-agent game in which agents may, first, self-select into a group task (GT) or an individual task (IT) and, second, choose work effort. In their choices of task and effort the agents have to consider pay contracts for both tasks as offered by the principal. The rational solution of the game implies that contract design may not induce agents to select GT and provide positive effort in GT. Furthermore it predicts equal behavior of (...) agents with different productivities. In contrast, considerations of trust, reciprocity and cooperation – the social-emotional model of behavior – suggest that contract design can influence the agents’ willingness to join groups and provide effort. We analyze the data by applying a two-step regression model (multinomial logit and tobit) and find that counter to the rational solution, contract design does influence both, task selection and effort choice. The principal can increase participation in work groups and can positively influence group performance. Larger payment increases the share of socially motivated agents in work groups. The selection effect is larger than the motivation effect. (shrink)

We live in interesting times. Two well-known biologists — E. O. Wilson and Richard Dawkins — and some of their well-known colleagues, who used to employ broadly similar selection models, now deeply disagree over the role of group selection in the evolution of eusociality (or so we argue). Yet they describe their models as interchangeable. As philosophers of biology, we wonder whether there is substantial (i.e., empirical) disagreement here at all, and, if there is, what is this disagreement about? We (...) argue that a substantial disagreement over the processes that caused eusociality best explains this debate, yet the common practice of using overarching definitions for “group selection” and “kin selection” renders empirical differences difficult to detect. We suggest Michael J. Wade’s use of these terms as a basis for models that reveal different selection processes. Wade’s models predict different outcomes for different processes and thus can be tested. (shrink)

In this article I summarize Friedrich Hayek’s cultural group selection theory and describe the evidence gathered by current cultural group selection theorists within the behavioral and social sciences supporting Hayek’s main assertions. I conclude with a few comments on Hayek and libertarianism.

The antipathy toward group selection expressed in the target article is puzzling because Laland et al.'s ideas dovetail neatly with modern group selection theory.

Wilson and Sober's (1994t) revival of group selection theory may have failed with some readers because its simple arithmetic foundation was obscured under the complexities of its presentation. When that uncontrovertible principle is uncovered, it broadens dramatically the fundamental motives that social scientists may impute to human nature and still be consistent with Darwinian evolutionary theory.

Human altruistic behavior has received a great deal of scientific attention over the past forty years. Altruistic-like behaviors found among insects and animals have illumined certain human behaviors, and the revival of interest in group selection has focused attention on how sacrificial altruism, although not adaptive for individuals, can be adaptive for groups. Curiously, at the same time that sociobiology has placed greater emphasis on the value of sacrificial altruism, Protestant ethics in America has moved away from it. While Roman (...) Catholic ethics has a longstanding tradition emphasizing an ordering of love, placing love of self second only to love for God, Protestant ethics in America has adopted a similar stance only recently, replacing a strong sacrificial ethic with one focusing on mutual regard for self and others. If sociobiology is correct about the significance of sacrificial altruistic behaviors for the survival of communities, this shift away from sacrificial agape by American Christianity may cut the community off from important resources for the development of a global ethic crucial for the survival of that faith community and humankind itself. (shrink)

Group selection is increasingly being viewed as an important force in human evolution. This paper examines the views of R.D. Alexander, one of the most influential thinkers about human behavior from an evolutionary perspective, on the subject of group selection. Alexander's general conception of evolution is based on the gene-centered approach of G.C. Williams, but he has also emphasized a potential role for group selection in the evolution of individual genomes and in human evolution. Alexander's views are internally inconsistent and (...) underestimate the importance of group selection. Specific themes that Alexander has developed in his account of human evolution are important but are best understood within the framework of multilevel selection theory. From this perspective, Alexander's views on moral systems are not the radical departure from conventional views that he claims, but remain radical in another way more compatible with conventional views. (shrink)

The evolution of the myxoma virus in Australia has been presented for many years as a test case for the hypothesis that group selection can function effectively `in the wild.' This paper critically examines the myxoma case, and argues that its failure as a test case for this hypothesis has broader implications for debates over the levels of selection.

Two controversies exist regarding the appropriate characterization of hierarchical and adaptive evolution in natural populations. In biology, there is the Wright-Fisher controversy over the relative roles of random genetic drift, natural selection, population structure, and interdemic selection in adaptive evolution begun by Sewall Wright and Ronald Aylmer Fisher. There is also the Units of Selection debate, spanning both the biological and the philosophical literature and including the impassioned groupselection debate. Why do these two discourses exist separately, and interact relatively little? (...) We postulate that the reason for this schism can be found in the differing focus of each controversy, a deep difference itself determined by distinct general styles of inquiry (e.g., Hacking 2002; Elwick 2007; Winther 2012, 2013) guiding each discourse. That is, the Wright-Fisher debate focuses on /adaptive process/, and tends to be instructed by the /mathematical modeling style/, while the focus of the Units of Selection controversy is /adaptive product/, and is typically guided by the /function style/. The differences between the two discourses can be usefully tracked by examining their interpretations of two contested strategies for theorizing hierarchical selection: /horizontal/ and /vertical/ averaging. (shrink)

The aim of this paper is to outline a typologyof selection processes, and show that differentsub-categories have different explanatorypower. The basis of this typology of selectionprocesses is argued to be the difference ofreplication processes involved in them. Inorder to show this, I argue that: 1.Replication is necessary for selection and 2.Different types of replication lead todifferent types of selection. Finally, it isargued that this typology is philosophicallysignificant, since it contrasts cases ofselection (on the basis of the replicationprocesses involved in them) (...) whereby selectioncauses adaptation – and, therefore, can beused in explanations of the (real or apparent)teleology of Nature – and cases in whichselection lacks such explanatory power. (shrink)

One critic complained that my argument was ‘philosophical’, as though that was sufficient condemnation. Philosophical or not, the fact is that neither he nor anybody else has found any flaw in what I said. And ‘in principle’ arguments such as mine, far from being irrelevant to the real world, can be more powerful than arguments based on particular factual research. My reasoning, if it is correct, tells us something important about life everywhere in the universe. Laboratory and field research can (...) tell us only about life as we have sampled it here. (The Selfish Gene (second edn, p322 in endnotes). (shrink)

In a recent Inquiry article Louis Pascal argues that the problem of massive starvation in the modern world is the result of a genetically-based human propensity to produce as many offspring as possible, regardless of ecological conditions. In this paper biological and anthropological objections to Pascal's thesis are discussed as well as the conclusions he draws from it. It is suggested that natural selection has produced humans who are flexible in their reproductive behavior in order to cope with rapidly changing (...) environments. The implications of both arguments for the population movement and the attempt to eliminate starvation are discussed. (shrink)

Two controversies exist regarding the appropriate characterization of hierarchical and adaptive evolution in natural populations. In biology, there is the Wright-Fisher controversy over the relative roles of random genetic drift, natural selection, population structure, and interdemic selection in adaptive evolution begun by Sewall Wright and Ronald Aylmer Fisher. There is also the Units of Selection debate, spanning both the biological and the philosophical literature and including the impassioned groupselection debate. Why do these two discourses exist separately, and interact relatively little? (...) We postulate that the reason for this schism can be found in the differing focus of each controversy, a deep difference itself determined by distinct general styles of inquiry (e.g., Hacking 2002; Elwick 2007; Winther 2012, 2013) guiding each discourse. That is, the Wright-Fisher debate focuses on /adaptive process/, and tends to be instructed by the /mathematical modeling style/, while the focus of the Units of Selection controversy is /adaptive product/, and is typically guided by the /function style/. The differences between the two discourses can be usefully tracked by examining their interpretations of two contested strategies for theorizing hierarchical selection: /horizontal/ and /vertical/ averaging. (shrink)

According to a once influential view of selection, it consists of repeated cycles of replication and interaction. It has been argued that this view is wrong: replication is not necessary for evolution by natural selection. I analyze the nine most influential arguments for this claim and defend the replication–interaction conception of selection against these objections. In order to do so, however, the replication–interaction conception of selection needs to be modified significantly. My proposal is that replication is not the copying of (...) an entity, the replicator, but the copying of a property. Thus, we can have a replication process without there being a replicator that is being copied. (shrink)

E. O. Wilson (1974: 54) describes the problem that social organisms pose: “On what bases do we distinguish the extremely modified members of an invertebrate colony from the organs of a metazoan animal?” This framing of the issue has inspired many to look more closely at how groups of organisms form and behave as emergent individuals. The possible existence of “superorganisms” test our best intuitions about what can count and act as genuine biological individuals and how we should study them. (...) As we will discuss, colonies of certain organisms display many of the properties that we usually reserve only to individual organisms. Although there is good reason to believe that many social insects form genuine emergent biological individuals, the conclusion offered here is of a slightly different sort. I will argue that to understand some social insects' interactions and the emergent traits they give rise to, it may be helpful to shift our understanding from a community-level approach to an ecosystem-level approach. I will argue that viewing certain insect colonies (termites) as parts of ecosystems allows us to better understand some of the adaptations that have emerged from their evolution. (shrink)

Understanding how cooperation evolves is central to explaining some core features of our biological world. Many important evolutionary events, such as the arrival of multicellularity or the origins of eusociality, are cooperative ventures between formerly solitary individuals. Explanations of the evolution of cooperation have primarily involved showing how cooperation can be maintained in the face of free-riding individuals whose success gradually undermines cooperation. In this paper I argue that there is a second, distinct, and less well explored, problem of cooperation (...) that I call the generation of benefit. Focusing on how benefit is generated within a group poses a different problem: how is it that individuals in a group can (at least in principle) do better than those who remain solitary? I present several different ways that benefit may be generated, each with different implications for how cooperation might be initiated, how it might further evolve, and how it might interact with different ways of maintaining cooperation. I argue that in some cases of cooperation, the most important underlying “problem” of cooperation may be how to generate benefit, rather than how to reduce conflict or prevent free-riding. (shrink)

Sometimes themes can be found in common across very different systems in which change occurs. Imre Lakatos developed a theory of change in science, and one involving entities visible at different levels. There are theories defended at a particular time, and there are also research programs, larger units that bundle together a sequence of related theories and within which many scientists may work. Research programs are competing higher-level units within a scientific field. Scientific change involves change within research programs, and (...) change in the ensemble of research programs present at a time, where some will be growing, some shrinking, some progressing, some degenerating. These are also themes in biological evolution. Recent biology has often found itself dealing with the relation between change at a level of "collectives" – such as organisms like us – and change at a lower level – the level of cells, genes, and other evolving parts. This work is continuous with an older discussion, one that arose when biological evolution was no more than a vague speculation, round the beginning of the 19th century. What is the living individual? What is the basic unit of life or living organization? Questions like this were pursued by Goethe, by Erasmus Darwin, the grandfather of Charles, and many others. Initially it was plants, especially, that were seen to raise these problems, and then newly described marine animals with strange life cycles. The discussion was influenced by the rise of the cell theory in the early 19th century, but some writers looked for individuals well below the level of the cell. (shrink)

Exploring whether clades can reproduce leads to new perspectives on general accounts of biological development and individuation. Here we apply James Griesemer's general account of reproduction to clades. Griesemer's account of reproduction includes a requirement for development, raising the question of whether clades may bemeaningfully said to develop. We offer two illustrative examples of what clade development might look like, though evaluating these examples proves difficult due to the paucity of general accounts of development. This difficulty, however, is instructive about (...) what a general account of development should look like and how it may usefully be applied to research problems (further suggesting a means for evaluating general accounts of development). Reproduction also requires individuation of parent and offspring. We argue that there is no special problem of individuating older and younger clades. The vagaries involved with determining when clades begin, mature, and end are precisely the same as those that arise when the same questions are asked of cells, organisms, or species. Though the question of clade reproduction and selection may still be open, the process of discovery presents new insights into old problems. (shrink)

Standard biological and philosophical treatments assume that dramatic genotypic or phenotypic change constitutes instantaneous speciation, and that barring such saltation, speciation is gradual evolutionary change in individual properties. Both propositions appear to be incongruent with standard theoretical perspectives on species themselves, since these perspectives are (a) non-pheneticist, and (b) tend to disregard intermediate cases. After reviewing certain key elements of such perspectives, it is proposed that species-membership is mediated by membership in a population. Species-membership depends, therefore, not on intrinsic characteristics (...) of an organism, but on relationship of an organism to others. A new definition of speciation is proposed in the spirit of this proposal. This definition implies that dramatic change is neither necessary nor sufficient for speciation. It also implies, surprisingly, that an organism can change species during its lifetime. (shrink)

We have argued elsewhere that: (A) Natural selection is not a cause of evolution. (B) A resolution-of-forces (or vector addition) model does not provide us with a proper understanding of how natural selection combines with other evolutionary influences. These propositions have come in for criticism recently, and here we clarify and defend them. We do so within the broad framework of our own “hierarchical realization model” of how evolutionary influences combine.

Views on the evolution of altruism based upon multilevel selection on structured populations pay little attention to the difference between fortuitous and deliberate processes leading to assortative grouping. Altruism may evolve when assortative grouping is fortuitously produced by forces external to the organism. But when it is deliberately produced by the same proximate mechanism that controls altruistic responses, as in humans, exploitation of altruists by selfish individuals is unlikely and altruism evolves as an individually advantageous trait. Groups formed with altruists (...) of this sort are special, because they are not affected by subversion from within. A synergistic process where altruism is selected both at the individual and at the group level can take place. (shrink)

There are two competing interpretations of the modern synthesis theory of evolution: the dynamical (also know as ‘traditional’) and the statistical. The dynamical interpretation maintains that explanations offered under the auspices of the modern synthesis theory articulate the causes of evolution. It interprets selection and drift as causes of population change. The statistical interpretation holds that modern synthesis explanations merely cite the statistical structure of populations. This paper offers a defense of statisticalism. It argues that a change in trait frequencies (...) in a population can be attributed only to selection or drift against the background of a particular statistical description of the population. The traditionalist supposition that selection and drift are description‐independent causes of population change leads the dynamical interpretation into a dilemma: it must face a contradiction or accept the loss of explanatory power. (shrink)