Once upon a time in evolutionary theory, everything happened for the best. Predators killed only the old or the sick. Pecking orders and other dominance hierarchies minimized wasteful conflict within the group. Male displays ensured that only the best and the fittest had mates. In the culmination of this tradition, Wynne-Edwards argued that many species have mechanisms that ensure groups do not over-exploit their resource base. The “central function” of territoriality in birds and other higher animals is “of limiting the (...) numbers of occupants per unit area of habitat”. Species with dominance hierarchies, species with lekking breeding systems, and species with communal breeding regulate their populations. These social mechanisms have population regulation as their “underlying primary function”. Wynne-Edwards argued that these mechanisms evolve through group selection. Populations without such mechanisms are apt to go extinct by eroding their own resource base. (shrink)

The authors demonstrate that unselfish behavior is in fact an important feature of both biological and human nature. Their book provides a panoramic view of altruism throughout the animal kingdom--from self-sacrificing parasites to the human capacity for selflessness--even as it explains the evolutionary sense of such behavior.

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

A simple and general criterion is derived for the evolution of altruism when individuals interact in pairs. It is argued that the treatment of this problem in kin selection theory and in game theory are special cases of this general criterion.

The proper treatment of correlation in evolutionary game theory has unexpected connections with recent philosophical discussions of the theory of rational decision. The Logic of Decision (Jeffrey 1983) provides the correct framework for correlated evolutionary game theory and a variant of "ratifiability" is the appropriate generalization of "evolutionarily stable strategy". The resulting theory unifies the treatment of correlation due to kin, population viscosity, detection, signaling, reciprocal altruism, and behavior-dependent contexts. It is shown that (1) a strictly dominated strategy may be (...) selected, and (2) under conditions of perfect correlation a strictly efficient strategy must be selected. (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)

Two alternative statistical approaches to modelling multi-level selection in nature, both found in the contemporary biological literature, are contrasted. The simple covariance approach partitions the total selection differential on a phenotypic character into within-group and between-group components, and identifies the change due to group selection with the latter. The contextual approach partitions the total selection differential into different components, using multivariate regression analysis. The two approaches have different implications for the question of what constitutes group selection and what does not. (...) I argue that the contextual approach is theoretically preferable. This has important implications for a number of issues in the philosophical debate about the levels of selection. Introduction Group selection and the covariance formulation of selection The contextual approach A modification of the simple covariance approach Consequences: frameshifting and additivity 5.1 Frameshifting 5.2 Additivity Conclusion. (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)

Recent years have seen increased interest in the question of whether it is possible to provide an evolutionary game-theoretic explanation for certain kinds of social norms. I sketch a proof of a general representation theorem for a large class of evolutionary game-theoretic models played on a social network, in hope that this will contribute to a greater understanding of the long-term evolutionary dynamics of such models, and hence the evolution of social norms.

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)

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

Proponents of genic selectionism have claimed that evolutionary processes normally viewed as selection on individuals can be "represented" as selection on alleles. This paper discusses the relationship between mathematical questions about the formal requirements upon state spaces necessary for the representation of different types of evolutionary processes and causal questions about the units of selection in such processes.

Brian Skyrms, author of the successful Evolution of the Social Contract has written a sequel. The book is a study of ideas of cooperation and collective action. The point of departure is a prototypical story found in Rousseau's A Discourse on Inequality. Rousseau contrasts the pay-off of hunting hare where the risk of non-cooperation is small but the reward is equally small, against the pay-off of hunting the stag where maximum cooperation is required but where the reward is so much (...) greater. Thus, rational agents are pulled in one direction by considerations of risk and in another by considerations of mutual benefit. Written with Skyrms's characteristic clarity and verve, this intriguing book will be eagerly sought out by students and professionals in philosophy, political science, economics, sociology and evolutionary biology. (shrink)

Altruism is generally understood to be behavior that benefits others at a personal cost to the behaving individual. However, within evolutionary biology, different authors have interpreted the concept of altruism differently, leading to dissimilar predictions about the evolution of altruistic behavior. Generally, different interpretations diverge on which party receives the benefit from altruism and on how the cost of altruism is assessed. Using a simple trait-group framework, we delineate the assumptions underlying different interpretations and show how they relate to one (...) another. We feel that a thorough examination of the connections between interpretations not only reveals why different authors have arrived at disparate conclusions about altruism, but also illuminates the conditions that are likely to favor the evolution of altruism. (shrink)

In this paper Wimsatt's analysis of units of selection is taken as defining the units of selection question. A definition of levels of selection is offered and it is shown that the levels of selection question is quite different from the units of selection question. Some of the relations between units and levels are briefly explored. It is argued that the levels of selection question is the question relevant to explanatory concerns, and it is suggested that it is the question (...) relevant to ontological concerns. (shrink)