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.

With this manifesto, John Dupré systematically attacks the ideal of scientific unity by showing how its underlying assumptions are at odds with the central conclusions of science itself.

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)

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

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

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

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

This paper analyzes the generation and function of hitherto ignored or misrepresented interfield theories , theories which bridge two fields of science. Interfield theories are likely to be generated when two fields share an interest in explaining different aspects of the same phenomenon and when background knowledge already exists relating the two fields. The interfield theory functions to provide a solution to a characteristic type of theoretical problem: how are the relations between fields to be explained? In solving this problem (...) the interfield theory may provide answers to questions which arise in one field but cannot be answered within it alone, may focus attention on domain items not previously considered important, and may predict new domain items for one or both fields. Implications of this analysis for the problems of reduction and the unity and progress of science are mentioned. (shrink)

It is a challenge to explain how evolutionary altruism can evolve by the process of natural selection, since altruists in a group will be less fit than the selfish individuals in the same group who receive benefits but do not make donations of their own. Darwin proposed a theory of group selection to solve this puzzle. Very simply, even though altruists are less fit than selfish individuals within any single group, groups of altruists are more fit than groups of selfish (...) individuals. If a population is subdivided into many groups that vary in their altruistic tendencies, altruism will be favored at the level of selection among groups even as it is being disfavored at the level of selection among individuals within groups. Darwin’s scenario became the basis for a theoretical framework called multilevel selection theory. (shrink)

Developmental systems theory (DST) expands the unit of replication from genes to whole systems of developmental resources, which DST interprets in terms of cycling developmental processes. Expansion seems required by DST's argument against privileging genes in evolutionary and developmental explanations of organic traits. DST and the expanded replicator brook no distinction between biological and cultural evolution. However, by endorsing a single expanded unit of inheritance and leaving the classical molecular notion of gene intact, DST achieves only a nominal reunification of (...) heredity and development. I argue that an alternative conceptualization of inheritance denies the classical opposition of genetics and development while avoiding the singularity inherent in the replicator concept. It also yields a new unit--the reproducer--which genuinely integrates genetic and developmental perspectives. The reproducer concept articulates the non-separability of "genetic" and "developmental" roles in units of heredity, development, and evolution. DST reformulated in terms of reproducers rather than replicators preserves an empirically interesting distinction between cultural and biological evolution. (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)

Darwinians are realists about the force of selection, but there has been surprisingly little discussion about what form this realism should take. Arguments about the units of selection in general and genic selectionism in particular reveal two realist assumptions: (1) for any selection process, there is a uniquely correct identification of the operative selective forces and the level at which each impinges; and (2) selective forces must satisfy the Pareto-style requirement of probabilistic causation. I argue that both assumptions are false; (...) we must temper realism about the force of selection and revise the way we think about probabilistic causation. (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)

Sober (1984) presents an account of selection motivated by the view that one property can causally explain the occurrence of another only if the first plays a unique role in the causal production of the second. Sober holds that a causal property will play such a unique role if it is a population level cause of its effect, and on this basis argues that there is selection for a trait T only if T is a population level cause of survival (...) and reproductive success. Sterelny and Kitcher (1988) claim against Sober that some traits directly subject to selection will not satisfy the probabilistic condition on population level causation. In this paper I show that Sober has the resources to resist the Sterelny-Kitcher complaint, but I argue that not all traits that satisfy the probabilistic condition play the required unique role in the production of their effects. (shrink)

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

Several evolutionary biologists have used a parsimony argument to argue that the single gene is the unit of selection. Since all evolution by natural selection can be represented in terms of selection coefficients attaching to single genes, it is, they say, "more parsimonious" to think that all selection is selection for or against single genes. We examine the limitations of this genic point of view, and then relate our criticisms to a broader view of the role of causal concepts and (...) the dangers of reification in science. (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)

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)

In a recent article, Kim Sterelny and Philip Kitcher5 defend a version of genic selectionism and attempt to refute the criticisms I made of that doctrine. Their defense has two components. First, they find fault with the account I gave of the units-of-selection controversy-an account which uses the idea of probabilistic causality as a tool of explication. Second, they provide a positive account of their own of what that controversy concerns, one which they think allows genic selectionism to emerge as (...) a successful thesis. I believe that the position they sketch is mistaken, both in its general orientation and in its details. I believe that the Sterelny/ Kitcher position misunderstands what the biological question of the units of selection is about and that their criticisms of my own proposal are mistaken as well. (shrink)

One would think that after one hundred years of earnest work, Darwinians would know what it is that nature selects for. Yet while some Darwinians argue that nature selects only for genes, others insist that the units being selected for are organisms and groups of organisms. Darwinians have also had difficulty identifying the fundamental principles of their theory. Some think the basic principle is the survival of the fittest; others deny that this principle is an important part of their theory. (...) ;Philosophers of biology think that many of these fundamental issues could be resolved if we had a clearer understanding of evolutionary theory. Traditionally philosophers have tried to clarify scientific theories by formulating their central laws as universal statements in a formal language. This approach has been challenged recently by those who argue that theories should be thought of as sets of models, not as sets of universal statements. The central aim of my dissertation is to demonstrate how this shift in focus from statements to models can help resolve several long-standing problems associated with the theory of natural selection. ;In the first chapter, I identify the fundamental principles of Darwin's original theory of natural selection. I do so by ignoring Darwin's generalizations and focusing on the models of natural selection presented in the Origin as imaginary illustrations. I clarify Darwin's analogical argument for natural selection in the second chapter. My analysis is based upon the view that analogical reasoning involves using specific information about one model to infer specific information about isomorphic models. In the third chapter, I argue that the principle of the survival of the fittest is untestable and plays no role in specifying the models of natural selection. The fourth chapter concerns the units of selection controversy. I argue that many selection processes can be accounted for by more than one model and that the choice between them is often a pragmatic one. While the four chapters deal with different problems, they are related by their common perspective on the structure of scientific theories, a perspective which I suggest is largely responsible for their individual successes. (shrink)

"Additive variance in fitness" is an important concept in the formal apparatus of population genetics. Wimsatt and Lloyd have argued that this concept can also be used to decide the "unit of selection" in an evolutionary process. The paper argues that the proposed criteria of Wimsatt and Lloyd are ambiguous, and several interpretations of their views are presented. It is argued that none of these interpretations provide acceptable criteria for deciding units of selection. The reason is that additive variance in (...) fitness can be both a cause of evolution, but also a byproduct of selection at another level. (shrink)

It is shown that, for technical reasons, the additivity of variance criterion employed by Lloyd (1988) to define a unit of selection is, in almost all models of selection, inconsistent with the possibility that genes are sometimes not the unit of selection. A case when the latter view is particularly attractive is that of heterosis, and the additivity criterion is inadequate in even such an extreme case. The connection between that criterion and the so-called "fundamental theorem of natural selection" is (...) briefly explored. Skepticism is expressed about the value of measures such as variance in efforts to resolve any of the disputes about the "units of selection.". (shrink)