Online research in philosophy

To evaluate Hume's thesis that causal claims are always empirical, I consider three kinds of causal statement: ?e1 caused e2 ?, ?e1 promoted e2 ?, and ?e1 would promote e2 ?. Restricting my attention to cases in which ?e1 occurred? and ?e2 occurred? are both empirical, I argue that Hume was right about the first two, but wrong about the third. Standard causal models of natural selection that have this third form are a priori mathematical truths. Some are obvious, others less so. Empirical work on natural selection takes the form of defending causal claims of the first two types. I provide biological examples that illustrate differences among these three kinds of causal claim.

The explanatory role of natural selection is one of the long-term debates in evolutionary biology. Nevertheless, the consensus has been slippery because conceptual confusions and the absence of a unified, formal causal model that integrates different explanatory scopes of natural selection. In this study we attempt to examine two questions: (i) What can the theory of natural selection explain? and (ii) Is there a causal or explanatory model that integrates all natural selection explananda? For the first question, we argue that five explananda have been assigned to the theory of natural selection and that four of them may be actually considered explananda of natural selection. For the second question, we claim that a probabilistic conception of causality and the statistical relevance concept of explanation are both good models for understanding the explanatory role of natural selection. We review the biological and philosophical disputes about the explanatory role of natural selection and formalize some explananda in probabilistic terms using classical results from population genetics. Most of these explananda have been discussed in philosophical terms but some of them have been mixed up and confused. We analyze and set the limits of these problems.

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.

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.

Group selection is said to occur when the traits of groups that systematically out-reproduce competing groups eventually come to characterize the species. Evolutionists have long disputed over the degree to which group selection is effective—that is, over the degree to which social group characteristics can be attributed to selection on these characteristics. The intractability of this controversy arises from three ambiguities in the natural selection metaphor that manifest themselves when that metaphor is shifted to the group level: (1) uncertainty about what constitutes the analogue for "flock" in the group level metaphor; (2) uncertainty about how to identify the group "parents" of offspring groups; and (3) uncertainty about what constitutes a group trait for the purposes of group selection. When group selection is specified as a theory about the evolution of emergent properties of groups through differential group productivity mediated by quantitative inheritance of group traits, these ambiguities disappear.

Hierarchical expansions of the theory of natural selection exist in two distinct bodies of thought in evolutionary biology, the group selection and the species selection traditions. Both traditions share the point of view that the principles of natural selection apply at levels of biological organization above the level of the individual organism. This leads them both to considermultilevel selection situations, where selection is occurring simultaneously at more than one level. Impeding unification of the theoretical approaches of the multilevel selection traditions are the different goals of investigators in the different subdisciplines and the different types of data potentially available for analysis. We identify two alternative approaches to multilevel situations, which we termmultilevel selection [1] andmultilevel selection [2]. Of interest in the former case are the effects of group membership onindividual fitnesses, and in the latter the tendencies for the groups themselves to go extinct or to found new groups (i.e., group fitnesses). We argue that: neither represents the entire multilevel selection process; both are aspects of any multilevel selection situation; and both are legitimate approaches, suitable for answering different questions. Using this formalism, we show that: multilevel selection [2] does not require emergent group properties in order to provide an explanatory mechanism of evolutionary change; multilevel selection [1] is usually more appropriate for neontological group selection studies; and species selection is most fruitfully considered from the point of view of multilevel selection [2]. Finally we argue that the effect hypothesis of macroevolution, requiring, in selection among species, both the absence of group effects on organismic fitness (multilevel selection [1]), and the direct determination of species fitnesses by those of organisms, is untestable with paleontological data. Furthermore, the conditions for the effect hypothesis to hold are extremely restrictive and unlikely to apply to the vast majority of situations encountered in nature.

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.

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.

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

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.