Group Selection

The ‘managed-metabolism’ hypothesis suggests that a ‘cooperation barrier’ must be overcome if self-producing chemical organizations are to undergo the transition from non-life to life. This dynamical barrier prevents un-managed autocatalytic networks of molecular species from individuating into complex, cooperative organizations. The barrier arises because molecular species that could otherwise make significant cooperative contributions to the success of an organization will often not be supported within the organization, and because side reactions and other ‘free-riding’ processes will undermine cooperation. As a result, (...) the barrier seriously impedes the emergence of individuality, complex functionality and the transition to life. This barrier is analogous to the cooperation barrier that also impedes the emergence of complex cooperation at all levels of living organization. As has been shown at other levels of organization, the barrier can be overcome comprehensively by appropriate ‘management’. Management implements a system of evolvable constraints that can overcome the cooperation barrier by ensuring that beneficial co-operators are supported within the organization and by suppressing free-riders. In this way, management can control and manipulate the chemical processes of a collectively autocatalytic organization, producing novel processes that serve the interests of the organization as a whole and that could not arise and persist in an un-managed chemical organization. Management self-organizes because it is able to capture some of the benefits that are produced when its interventions promote cooperation, thereby enhancing productivity. Selection will therefore favour the emergence of managers that take over and manage chemical organizations so as to overcome the cooperation barrier. The managed-metabolism hypothesis demonstrates that if management is to overcome the cooperation barrier comprehensively, its interventions must be digitally coded. In this way, the hypothesis accounts for the two-tiered structure of all living cells in which a digitally-coded genetic apparatus manages an analogically-informed metabolism.GraphicalOpen image in new window. (shrink)

Evolution's Arrow argues that evolution is directional and progressive, and that this has major consequences for humanity. Without resort to teleology, the book demonstrates that evolution moves in the direction of producing cooperative organisations of greater scale and evolvability - evolution has organised molecular processes into cells, cells into organisms, and organisms into societies. The book founds this position on a new theory of the evolution of cooperation. It shows that self-interest at the level of the genes does not prevent (...) cooperation from increasing as evolution unfolds. Evolution progresses by discovering ways to build cooperative organisations out of self-interested individuals. The book also shows that evolution itself has evolved. Evolution has progressively improved the ability of evolutionary mechanisms to discover effective adaptations. And it has produced new and better mechanisms. Evolution's Arrow uses this understanding of the direction of evolution to identify the next great steps in the evolution of life on earth - the steps that humanity must take if we are to continue to be successful in evolutionary terms. A key step for humanity is to increase the scale and evolvability of our societies, eventually forming a unified and cooperative society on the scale of the planet. We must also transform ourselves psychologically to become self-evolving organisms - organisms that are able to escape their biological and cultural past by adapting in whatever directions are necessary to achieve future evolutionary success. (shrink)

Hayek’s social theory of evolution suggests that market liberal morality is adaptive for social groups. He justified the evolutionary superiority of market liberalism by asserting that groups operating under a market liberal morality would have a higher capacity to expand and reproduce than groups with alternative tribal moralities. Thus, market liberal groups would be favoured through cultural and genetic group selection. But in fact, market liberal morality reveals maladaptive tendencies and remains insufficiently powerful to create adaptive social groups. Hayek’s dismissal (...) of moral tribalism in favour of market liberal morality is found to underestimate the importance of tribal goals in the evolutionary system. (shrink)

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

I give concise derivations of Price's equation and the criteria for kin and group selection, prove that kin and group selection are equivalent, and discuss the controversies about altruism.

In a recent paper, Brandon and Nijhout argue against genic selectionism—the thesis, roughly, that evolutionary processes are best understood from the gene’s-eye point of view—by presenting a case in which genic models of selection allegedly make predictions that conflict with the (correct) predictions of higher-level genotypic selection models. Their argument, if successful, would refute the widely held belief that genic models and higher-level models are predictively equivalent. Here, I argue that Brandon and Nijhout fail to demonstrate that the models make (...) incompatible predictions. (shrink)

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

Since Gintis is a senior economist and I have read some of his previous books with interest, I was expecting some more insights into behavior. Sadly he makes the dead hands of group selection and phenomenology into the centerpieces of his theories of behavior, and this largely invalidates the work. Worse, since he shows such bad judgement here, it calls into question all his previous work. The attempt to resurrect group selection by his friends at Harvard, Nowak and Wilson, a (...) few years ago was one of the major scandals in biology in the last decade, and I have recounted the sad story in my article ‘Altruism, Jesus and the End of the World—how the Templeton Foundation bought a Harvard Professorship and attacked Evolution, Rationality and Civilization -- A review of E.O. Wilson 'The Social Conquest of Earth' (2012) and Nowak and Highfield ‘SuperCooperators’ (2012).’ Unlike Nowak, Gintis does not seem to be motivated by religious fanaticism, but by the strong desire to generate an alternative to the grim realities of human nature, made easy by the (near universal) lack of understanding of basic human biology and blank slateism of behavioral scientists, other academics, and the general public. -/- Gintis rightly attacks (as he has many times before) economists, sociologists and other behavioral scientists for not having a coherent framework to describe behavior. Of course the framework needed to understand behavior is an evolutionary one. Unfortunately he fails to provide one himself (according to his many critics and I concur), and the attempt to graft the rotten corpse of group selection onto whatever economic and psychological theories he has generated in his decades of work, merely invalidates his entire project. Although Gintis makes a valiant effort to understand and explain the genetics, like Wilson and Nowak, he is far from an expert, and like them, the math just blinds him to the biological impossibilities and of course this is the norm in science. As Wittgenstein famously noted on the first page of Culture and Value “There is no religious denomination in which the misuse of metaphysical expressions has been responsible for so much sin as it has in mathematics.” -/- It has always been crystal clear that a gene that causes behavior which decreases its own frequency cannot persist, but this is the core of the notion of group selection. Furthermore, it has been well known and often demonstrated that group selection just reduces to inclusive fitness (kin selection), which, as Dawkins has noted, is just another name for evolution by natural selection. Like Wilson, Gintis has worked in this arena for about 50 years and still has not grasped it, but after the scandal broke, it took me only 3 days to find, read and understand the most relevant professional work, as detailed in my article. It is mind boggling to realize that Gintis and Wilson were unable to accomplish this in nearly half a century. -/- I discuss the errors of group selection and phenomenology that are the norm in academia as special cases of the near universal failure to understand human nature that are destroying America and the world. -/- Those who wish to read all my articles please consult the ebook here Philosophy, Human Nature and the Collapse of Civilization -- Articles and Reviews 2006-2016 by Michael Starks 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)

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)

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)

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)

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.

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)

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)

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)

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

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)

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)

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..

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

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)

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)

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)

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)

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)

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)

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)

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)

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)

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)

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.

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)

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)

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.

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)

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)

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)

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