Group Selection

Critiques G. Edelman's Theory of Neuronal Group Selection , from the perspectives of social constructionism, contemporary theory of metaphor, and existential-phenomenological psychology. This theory provides a contemporary biological view of consciousness and the self. Edelman's notion of consciousness as purely biological, and his attempt to ground intentionality in the body, are reductionistic. It is suggested that phenomenological descriptions must be taken into the center of the problem of consciousness. An understanding of the relation of the experiential basis of metaphor and (...) its ensuing ontological status, can offer a view of the influence of particular biological metaphors which could help to eliminate their reification in psychology and induce their critique. 2012 APA, all rights reserved). (shrink)

In evolutionary biology and ecology, ontological and epistemological perspectives based on the replicator and the interactor have become the background that makes it possible to transcend traditional biological levels of organization and to achieve a unified view of evolution in which replication and interaction are fundamental operating processes. Using the transactional perspective proposed originally by John Dewey and Arthur Fisher Bentley, a new ontological and methodological category is proposed here: the transactor. The transactional perspective, based on the concept of the (...) transactor, bridges the dichotomy between organisms and environment that characterizes the interactional perspective on evolution and provides epistemological support for the emergentist, systemic view of evolutionary and developmental processes. (shrink)

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

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

In this paper I argue against the claim, recently put forward by some philosophers of biology and evolutionary biologists, that there can be two or more ontologically distinct levels of selection. I show by comparing the fitness of individuals with that of collectives of individuals in the same environment and over the same period of time – as required to decide if one or more levels of selection is acting in a population – that the selection of collectives is a (...) by-product of selection at the individual level; thus, talking about two or more levels of selection represents merely a different perspective on one and the same process. (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.

Do multi-level selection explanations of the evolution of social traits deepen the understanding provided by single-level explanations? Central to the former is a mathematical theorem, the multi-level Price decomposition. I build a framework through which to understand the explanatory role of such non-empirical decompositions in scientific practice. Applying this general framework to the present case places two tasks on the agenda. The first task is to distinguish the various ways of suppressing within-collective variation in fitness, and moreover to evaluate their (...) biological interest. I distinguish four such ways: increasing retaliatory capacity, homogenising assortment, and collapsing either fitness structure or character distribution to a mean value. The second task is to discover whether the third term of the Price decomposition measures the effect of any of these hypothetical interventions. On this basis I argue that the multi-level Price decomposition has explanatory value primarily when the sharing-out of collective resources is `subtractable'. Thus its value is more circumscribed than its champions Sober and Wilson (1998) suppose. (shrink)

The concept of adaptation is pivotal to modern evolutionary thinking, but it has long been the subject of controversy, especially in respect of the relative roles of selection versus constraints in explaining the traits of organisms. This paper tackles a different problem for the concept of adaptation: its interpretation in light of multilevel selection theory. In particular, I arbitrate a dispute that has broken out between the proponents of rival perspectives on multilevel adaptations. Many experts now say that multilevel and (...) kin selection views are mathematically equivalent to one another—that the mathematical accounting of evolution can be carried out at any hierarchical level one chooses. But what does this formal equivalence imply - are they equivalent in other ways too? I show here that significant conceptual non-equivalence has survived: the two sides commit to different views regarding how much selection has to act at a level before we can call traits at that level adaptations; about whether policing mechanisms are adaptations, and about whether non-organisms can bear adaptations. (shrink)

Evolutionary anthropology has focused on the origins of war, or rather ethnocentricity, because it epitomizes the problem of group selection, and because war may itself have been the main agent of group selection. The neo-Darwinian synthesis in biology has explained how ethnocentricity might evolve by group selection, and the distinction between evoked culture and adopted culture, suggested by the emerging synthesis in evolutionary psychology, has explained how it might be transmitted. Ethnocentric mechanisms could have evolved by genetic selection in ancestral (...) hominids, or through the interaction of genetic and cultural selection in modern humans, or both. The existence of similar behaviors in chimpanzees and the parallel development of early human societies around the globe are arguments for such inherited mechanisms. There may have been some adaptive breakthroughs in purely cultural evolution, but this process does not seem likely to produce long-term Darwinian adaptations because of the prolificity of cultural traits. Warfare may once have been a major agent of group selection, but the rates of extinction among human groups are so slow as to render this improbable since the rise of state-level societies, whose warfare tends to become a cyclical balance-of-power situation. Perhaps the most serious implication of current evolutionary thought is that the individualistic model of culture common in the social sciences and humanities is outmoded, and should be replaced by a new model that recognizes the organismic nature of human societies. (shrink)

Flipping convention on its head, Eric Dietrich argues that science uncovers awe-inspiring, enduring mysteries, while religion, regarded as the source for such mysteries, is a biological phenomenon. Just like spoken language, Dietrich shows that religion is an evolutionary adaptation. Science is the source of perplexing yet beautiful mysteries, however natural the search for answers may be to human existence. _Excellent Beauty_ undoes our misconception of scientific inquiry as an executioner of beauty, making the case that science has won the battle (...) with religion so thoroughly it can now explain why religion persists. The book also draws deep lessons for human flourishing from the very existence of scientific mysteries. It is these latter wonderful, completely _public_ truths that constitute some strangeness in the proportion, revealing a universe worthy of awe and wonder. (shrink)

In this landmark volume, an international group of scientists has synthesized their collective expertise and insight into a newly unified vision of insect ...

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)

Two major clarifications have greatly abetted the understanding and fruitful expansion of the theory of natural selection in recent years: the acknowledgment that interactors, not replicators, constitute the causal unit of selection; and the recognition that interactors are Darwinian individuals, and that such individuals exist with potency at several levels of organization (genes, organisms, demes, and species in particular), thus engendering a rich hierarchical theory of selection in contrast with Darwin’s own emphasis on the organismic level. But a piece of (...) the argument has been missing, and individuals at levels distinct from organisms have been denied potency (although granted existence within the undeniable logic of the theory), because they do not achieve individuality with the same devices used by organisms and therefore seem weak by comparison. We show here that different features define Darwinian individuality across scales of size and time. In particular, species-individuals may develop few emergent features as direct adaptations. The interactor approach works with emergent fitnesses, not with emergent features; and species, as a consequence of their different mechanism for achieving individuality (reproductive exclusivity among subparts, that is, among organisms), express many effects from other levels. Organisms, by contrast, suppress upwardly cascading effects, because the organismic style of individuality (by functional integration of subparts) does not permit much competition or differential reproduction of parts from within. Species do not suppress the operation of lower levels; such effects therefore become available as exaptations conferring emergent fitness—a primary source of the different strength that species achieve as effective Darwinian individuals in evolution. (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)

My general aim is to clarify the foundational difference between Stephen Jay Gould and Richard Dawkins concerning what biological entities are the units of selection in the process of evolution by natural selection. First, I recapitulate Gould’s central objection to Dawkins’s view that genes are the exclusive units of selection. According to Gould, it is absurd for Dawkins to think that genes are the exclusive units of selection when, after all, genes are not the exclusive interactors: those agents directly engaged (...) with, directly impacted by, environmental pressures. Second, I argue that Gould’s objection still goes through even when we take into consideration Sterelny and Kitcher’s defense of gene selectionism in their admirable paper “The Return of the Gene.” Third, I propose a strategy for defending Dawkins that I believe obviates Gould’s objection. Drawing upon Elisabeth Lloyd’s careful taxonomy of the various understandings of the unit of selection at play in the philosophy of biology literature, my proposal involves realizing that Dawkins endorses a different understanding of the unit of selection than Gould holds him to, an understanding that does not require genes to be the exclusive interactors. (shrink)

This paper argues that the contextual approach to natural selection does not offer an estimation of the contributions of individual and group selection to evolutionary change in multi-level selection scenarios, and that this is so because the term “group selection”, as defined by the contextual approach, does not refer to a process taking place at the group level. In the contextual analysis framework, this term simply denotes an evolutionary change that takes place due to the fact that, overall, individual types (...) do not share similar contexts or environments, and the only way to claim that such an evolutionary change is a result of selection is by admitting that “group selection” is in fact a kind of frequency-dependent selection, i.e. a selection process taking place at the individual level. Therefore, under the names “individual selection” and “group selection”, the contextual approach actually isolates two aspects of the relation between individual types and their environment, and not two distinct levels of selection. (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 notion of structure of evolution is proposed to capture what it means to say that two situations exhibit the same or similar constellations of factors affecting evolution. The key features of holobiont evolution and the hologenome theory are used to define a holobiont structure of evolution. Finally, Cultural Group Selection, a set of hypotheses regarding the evolution of human cognition, is shown to match the holobiont structure closely though not perfectly.

This book brings together important essays by one of the leading philosophers of science at work today. Elisabeth A. Lloyd examines several of the central topics in philosophy of biology, including the structure of evolutionary theory, units of selection, and evolutionary psychology, as well as the Science Wars, feminism and science, and sexuality and objectivity. Lloyd challenges the current evolutionary accounts of the female orgasm and analyses them for bias. She also offers an innovative analysis of the concept of objectivity. (...) Lloyd analyses the structure of evolutionary theory and unlocks the puzzle of the units of selection debates into four distinct aspects, illuminating several mysteries in the biology literature. Central to all essays in this book is the author's abiding concern for evidence and empirical data. (shrink)

I argue that four of the fundamental claims of those calling themselves `genic pluralists'Philip Kitcher, Kim Sterelny, and Ken Watersare defective. First, they claim that once genic selectionism is recognized, the units of selection problems will be dissolved. Second, Sterelny and Kitcher claim that there are no targets of selection. Third, Sterelny, Kitcher, and Waters claim that they have a concept of genic causation that allows them to give independent genic causal accounts of all selection processes. I argue that each (...) one of these claims is either false or misleading. Moreover, the challenge that arises from the availability of genic causal accounts, namely, the inability to choose on rational grounds among genic and higher-level accounts, is unsupported. (shrink)

I admire Wilson & Sober's (W & S's) aim, to alert social scientists that group selection has risen from the ashqs, and to explicate its relevance to the behavioral sciences. Group selection has beenwidely misunderstood; furthermore, both authors have been instrumental in illuminating conceptual problems surrounding higher-level selection. Still, I find that this target article muddies the waters, primarily through its shifting and confused definition of a "vehicle" of selection. The fundamental problem is an ambiguity in the definition of "adaptation." (...) On the one hand, any evolutionary change that results from a selection process could be called an adaptation, by definition; I call this the "weak" view of adaptation. A "strong" view of adaptation, on the other hand, includes some notion of design - the evolution of a specific complex trait understood, in an engineering sense, to provide a mechanism favoring its owner's success in contributing to the evolutionary lineage. (shrink)

I address the controversy in evolutionary biology concerning which levels of biological entity (units) can and do undergo natural selection. I refine a definition of the unit of selection, first presented by William Wimsatt, that is grounded in the structure of natural selection models. I examine Elliott Sober's objection to this structural definition, the "homogeneous populations" problem; I find that neither the proposed definition nor Sober's own causal account can solve the problem. Sober, in his solution using his causal view, (...) imports precisely the information needed to make the structural definition effective. Finally, I indicate how the proposed definition can clarify which sorts of evidence could be brought to bear on the controversial case of the Myxoma virus. (shrink)

Most models of generational succession in sexually reproducing populations necessarily move back and forth between genic and genotypic spaces. We show that transitions between and within these spaces are usually hidden by unstated assumptions about processes in these spaces. We also examine a widely endorsed claim regarding the mathematical equivalence of kin-, group-, individual-, and allelic-selection models made by Lee Dugatkin and Kern Reeve. We show that the claimed mathematical equivalence of the models does not hold. *Received January 2007; revised (...) April 2008. †To contact the authors, please write to: Elisabeth Lloyd, Department of History and Philosophy of Science, 130 Goodbody Hall, Indiana University, Bloomington, IN 47405; e-mail: ealloyd@indiana.edu; Richard Lewontin, Department of Organismic and Evolutionary Biology, Harvard University, 26 Oxford Street, Cambridge, MA 02138; Marcus Feldman, Department of Biological Sciences, Stanford University, Stanford, CA 94305; e-mail: marc@charles.stanford.edu. (shrink)

this requirement for adaptations. Emergent characters are always potential adaptations. Not all selection processes produce adaptations, however. The key issue, in delineating a selection process, is the relationship between a character and fitness. The emergent character approach is more restrictive than alternative schemas that delineate selection..

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)

Review of Cooperation and its Evolution, edited by Kim Sterelny, Richard Joyce, Brett Calcott, and Ben Fraser.

Nowak is (or was) a respected Harvard professor of mathematical biology with numerous well regarded publications. Sadly, he has chosen to launch an arrogant attack on science motivated by religious fervor. His recent actions show the evil consequences when universities accept money from religious groups, science journals are so awed by big names that they avoid proper peer review, and egos are permitted to get out of control. Most of this book is good, but it repeats the utterly misconceived attacks (...) on inclusive fitness in favor of group selection which he and his much more famous Harvard colleague E.O.Wilson have been making the last 5 years, so it is on that chapter of the book that I will concentrate. This takes us into the nature of evolution, the basics of scientific methodology, how math relates to science, what constitutes a law or a theory, and what attitudes to religion and generosity are appropriate as we inexorably approach the collapse of industrial civilization. -/- Those wishing a comprehensive up to date framework for human behavior from the modern two systems view may consult my article The Logical Structure of Philosophy, Psychology, Mind and Language as Revealed in Wittgenstein and Searle 59p(2016). For all my articles on Wittgenstein and Searle see my e-book ‘The Logical Structure of Philosophy, Psychology, Mind and Language in Wittgenstein and Searle 367p (2016). Those interested in all my writings in their most recent versions may consult my e-book Philosophy, Human Nature and the Collapse of Civilization - Articles and Reviews 2006-2016’ 662p (2016). -/- All of my papers and books have now been published in revised versions both in ebooks and in printed books. -/- Talking Monkeys: Philosophy, Psychology, Science, Religion and Politics on a Doomed Planet - Articles and Reviews 2006-2017 (2017) https://www.amazon.com/dp/B071HVC7YP. -/- The Logical Structure of Philosophy, Psychology, Mind and Language in Ludwig Wittgenstein and John Searle--Articles and Reviews 2006-2016 (2017) https://www.amazon.com/dp/B071P1RP1B. -/- Suicidal Utopian Delusions in the 21st century: Philosophy, Human Nature and the Collapse of Civilization - Articles and Reviews 2006-2017 (2017) https://www.amazon.com/dp/B0711R5LGX . (shrink)

Famous ant-man E.O. Wilson has always been one of my heroes --not only an outstanding biologist, but one of the tiny and vanishing minority of intellectuals who at least dares to hint at the truth about our nature that others fail to grasp, or insofar as they do grasp, studiously avoid for of political expedience. Sadly, he is ending his long career in a most sordid fashion as a party to an ignorant and arrogant attack on science motivated at least (...) in part by religious fervor. It shows the vile consequences when universities accept money from religious groups, science journals are so awed by big names that they avoid proper peer review, and when egos are permitted to get out of control. It takes us into the nature of evolution, the basics of scientific methodology, how math relates to science, what constitutes a theory, and even what attitudes to religion and generosity are appropriate as we inexorably approach the collapse of industrial civilization. -/- Those interested in all my writings in their most recent versions may download from this site my e-book ‘Philosophy, Human Nature and the Collapse of Civilization Michael Starks (2016)- Articles and Reviews 2006-2016’ by Michael Starks First Ed. 662p (2016). -/- All of my papers and books have now been published in revised versions both in ebooks and in printed books. -/- Talking Monkeys: Philosophy, Psychology, Science, Religion and Politics on a Doomed Planet - Articles and Reviews 2006-2017 (2017) https://www.amazon.com/dp/B071HVC7YP. -/- The Logical Structure of Philosophy, Psychology, Mind and Language in Ludwig Wittgenstein and John Searle--Articles and Reviews 2006-2016 (2017) https://www.amazon.com/dp/B071P1RP1B. -/- Suicidal Utopian Delusions in the 21st century: Philosophy, Human Nature and the Collapse of Civilization - Articles and Reviews 2006-2017 (2017) https://www.amazon.com/dp/B0711R5LGX . (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)

Climate stresses and monetary resources seem to lead to different collective adaptations. However, the reference to adaptation and to ambiguous collective dimensions appears premature; populations may entertain nothing more than shared adaptiveness. At this point, the intricacy of the underlying evolutionary processes (cultural selection, fitness-utility decoupling) very much obscures any diagnosis based on correlations.

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)

Phenomena like meat sharing in hunter-gatherers, self-sacrifice in intergroup conflicts, and voluntary contribution to public goods provision in laboratory experiments have led to the development of numerous theories on the evolution of altruistic in-group beneficial behavior in humans. Many of these theories abstract away from the effects of kinship on the incentives for public goods provision, though. Here, it is investigated 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 linear public goods game. Using recent anthropological data sets on relatedness in 61 contemporary hunter-gatherer and horticulturalist societies the relevant parameters of this model are then estimated. It turns out that the kinship patterns observed in these societies substantially reduce the negative effect of increasing group size on incentives for public goods provision. It is suggested, therefore, that renewed attention should be given to inclusive fitness theory in the context of public goods provision also in sizable groups, because its explanatory power with respect to this central problem in the evolution of human cooperativeness and altruism might have been substantially underrated. (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)