Samir Okasha

In models of multi-level selection, the property of Darwinian fitness is attributed to entities at more than one level of the biological hierarchy, e.g. individuals and groups. However, the relation between individual and group fitness is a controversial matter. Theorists disagree about whether group fitness should always, or ever, be defined as total (or average) individual fitness. This paper tries to shed light on the issue by drawing on work in social choice theory, and pursuing an analogy between fitness and utility. Social choice theorists have long been interested in the relation between individual and social utility, and have identified conditions under which social utility equals total (or average) individual utility. These ideas are used to shed light on the biological problem.

An evolutionary basis for Bayesian rationality is suggested, by considering how natural selection would operate on an organism’s ‘policy’ for choosing an action depending on an environmental signal. It is shown that the evolutionarily optimal policy, as judged by the criterion of maximal expected reproductive output, is the policy that, for each signal, picks an action that maximizes conditional expected output given that signal. This suggests a possible route by which Bayes-rational creatures might have evolved.

This paper provides a philosophical analysis of the ongoing controversy surrounding R.A. Fisher's famous ‘fundamental theorem’ of natural selection. The difference between the ‘traditional’ and ‘modern’ interpretations of the theorem is explained. I argue that proponents of the modern interpretation have captured Fisher's intended meaning correctly and shown that the theorem is mathematically correct, pace the traditional consensus. However, whether the theorem has any real biological significance remains an unresolved issue. I argue that the answer depends on whether we accept Fisher's non-standard notion of environmental change, on which the theorem rests; arguments for and against this notion are explored. I suggest that there is a close link between Fisher's fundamental theorem and the modern ‘gene's eye’ view of evolution. Introduction What Does the Fundamental Theorem Say? Key Concepts Explained Alleged Significance of the FTNS Traditional versus Modern Interpretations of the FTNS The Modern Interpretation Illustrated Fisher's Concept of ‘Environmental Change’ Causality and the Modern Interpretation The Significance of the FTNS Re-considered Appendix CiteULike    Connotea    Del.icio.us    What's this?

The ‘levels of selection’ question is one of the most fundamental in evolutionary biology, for it arises directly from the logic of Darwinism. As is well-known, the principle of natural selection is entirely abstract; it says that any entities satisfying certain conditions will evolve by natural selection, whatever those entities are. (These conditions are: variability, associated fitness differences, and heritability (cf. Lewontin 1970).) This fact, when combined with the fact that the biological world is hierarchically structured, i.e. smaller biological units are nested within larger ones, gives rise to the levels of selection question. For in principle, entities at many different hierarchical levels, above and below that of the ‘individual organism’, (e.g. gene, chromosome, cell, kin group, colony, lineage, species) can satisfy the requirements for Darwinian evolution. This possibility has long been recognised by biologists, from Darwin himself to contemporary proponents of ‘multi-level selection’; and there exist numerous biological phenomena which suggest that it has actually occurred

The problem of how to make optimal choices in the face of risk arises in both economics/decision theory and also evolutionary biology; in the former, ‘optimal’ means utility maximizing, while in the latter it means fitness maximizing. This article explores the links, thematic and formal, between the economic and evolutionary theories of optimal choice in risky situations, with particular reference to the relationship between utility and fitness. It is argued that the link is strongest between evolution and ‘nonexpected’ utility theory, rather than traditional expected utility theory.

The levels of selection problem was central to Maynard Smith’s work throughout his career. This paper traces Maynard Smith’s views on the levels of selection, from his objections to group selection in the 1960s to his concern with the major evolutionary transitions in the 1990s. The relations between Maynard Smith’s position and those of Hamilton and G.C. Williams are explored, as is Maynard Smith’s dislike of the Price equation approach to multi-level selection. Maynard Smith’s account of the ‘core Darwinian principles’ is discussed, as is his debate with Sober and Wilson (1998) over the status of trait-group models, and his attitude to the currently fashionable concept of pluralism about the levels of selection.

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.

This paper compares two well-known arguments in the units of selection literature, one due to , the other due to . Both arguments concern the legitimacy of averaging fitness values across contexts and making inferences about the level of selection on that basis. The first three sections of the paper shows that the two arguments are incompatible if taken at face value, their apparent similarity notwithstanding. If we accept Sober and Lewontin's criterion for when averaging genic fitnesses across diploid genotypes is illegitmate, we cannot accept Sober and Wilson's criterion for when averaging individual fitnesses across groups is illegitimate, and vice versa. The final section suggests a possible way of reconciling the two arguments, by invoking an ambiguity in the concept of genic selection.

Evolutionary biology is striking for its ability to explain a large and diverse range of empirical phenomena on the basis of a few general theoretical principles. This article offers a philosophical perspective on the way that evolutionary biology has come to achieve such impressive generality, by focusing on “the strategy of endogenization”. This strategy involves devising evolutionary explanations for biological features that were originally part of the background conditions, or scaffolding, against which such explanations take place. Where successful, the strategy moves biology a step closer to the ideal of explaining as much as possible from evolutionary first principles. The strategy of endogenization is illustrated through a series of biological examples, historical and recent, and its philosophical implications are explored.

This chapter presents a displacement of the organism as a privileged level of analysis in evolutionary biology. It is concerned with the ontology of biology systems, with particular reference to hierarchical organization. It argues that the concept of a rank-free hierarchy can be transposed to the major transitions hierarchy, with interesting consequences. This chapter shows that the idea of rank freedom makes good sense of a number of facets of the recent discussion of evolutionary transitions and multilevel selection. It suggests that the idea of rank freedom is already at work, implicitly, in much theorizing about evolutionary transitions and/or multilevel selection.

For a number of years, the debate in evolutionary biology over the ’levels of selection’ has attracted intense interest from philosophers of science. The main question concerns the level of the biological hierarchy at which natural selection occurs. Does selection act on organisms, genes, groups, colonies, demes, species, or some combination of these? According to traditional Darwinian theory the answer is the organism -- it is the differential survival and reproduction of individual organisms that drives the evolutionary process. But there are alternative views too, including pluralist views which say that the choice between different levels of selection is often a matter of perspective, not empirical fact.

My comments will focus on the second and third chapters of Sober’s book , which explore Darwin’s ideas about altruism, group selection and kin selection , and sex-ratio evolution . Sober makes a persuasive argument for his main claim: that Darwin was a subtler thinker on these topics than he is often taken to be. While there is much that I admire in Sober’s lucid discussion, I will focus on points of disagreement. Readers should note that this is not the first time that Sober and I have disagreed on these issues .Sober begins his chapter on group selection with a brief history of the modern debate on the topic, which began in earnest in the 1960s. One of the key figures in this debate was George Williams, whose book Adaptation and Natural Selection helped convince many biologists that group selection was unlikely to be an important factor in evolution. Sober is critical of many of Williams’ arguments, though ..

A number of recent philosophers, including Michael Williams, Barry Stroud and Donald Davidson, have argued that scepticism about the external world stems from the foundationalist assumption that sensory experience supplies the data for our beliefs about the world. In order to assess this thesis, I offer abrief characterisation of the logical form of sceptical arguments. I suggest that sceptical arguments rely on the idea that many of our beliefs about the world are ‘underdetermined’ by the evidence on which they are based. Drawing on this characterisation of scepticism, I argue that Williams, Stroud andDavidson are right to see the foundationalist assumption as essential to the sceptic’s argument, but wrong to think that scepticism is inevitable once that assumption is in place. By pursuing an analogy with some recent debates in the philosophy of science, I try to locate the additional assumptions which the sceptic must make, in order to derive her conclusion.

This paper investigates the role of the concept of group heritability in group selection theory, in relation to the well-known distinction between type 1 and type 2 group selection (GS1 and GS2). I argue that group heritability is required for the operation of GS1 but not GS2, despite what a number of authors have claimed. I offer a numerical example of the evolution of altruism in a multi-group population which demonstrates that a group heritability coefficient of zero is perfectly compatible with the successful operation of group selection in the GS2 sense. A diagnosis of why group heritability has wrongly been regarded as necessary for GS2 is suggested.

Verificationism has often seemed attractive to philosophers because of its apparent abilityto deliver us from scepticism. However, I argue that purely epistemological considerationsprovide insufficient reason for embracing verificationism over realism. I distinguish twotypes of sceptical problem: those that stem from underdetermination by the actual data,and those that stem from underdetermination by all possible data. Verificationismevades problems of the second sort, but is powerless in the face of problems of the firstsort. But problems of the first sort are equally pressing. Furthermore, there is some reasonto think that the two types of problem have a common origin. Thus the desire to avoidscepticism provides insufficient reason for adopting verificationism.

The idea that clades might be units of selection, defended by a number of biologists and philosophers of biology, is critically examined. I argue that only entities which reproduce, i.e. leave offspring, can be units of selection, and that a necessary condition of reproduction is that the offspring entity be able, in principle, to outlive its parental entity. Given that clades are monophlyetic by definition, it follows that clades do not reproduce, so it makes no sense to talk about a clade's fitness, so clade selection is impossible. Three possible responses to this argument are examined and found wanting.

Sceptical arguments in epistemology typically employ sceptical hypotheses, which are rivals to our everyday beliefs so constructed that they fit exactly the evidence on which those beliefs are based. There are two ways of using a sceptical hypothesis to undermine an everyday belief, giving rise to two distinct sorts of sceptical argument: underdetermination‐based and closure‐based. However, both sorts of argument, as usually formulated in the literature, fall foul of evidential holism, for they ignore the crucial role of background beliefs. An analogy with the philosophy of science makes this point explicit. There is no simple way to “holism proof” the two sceptical arguments.

Many philosophers have claimed that Bayesianism can provide a simple justification for hypothetico-deductive inference, long regarded as a cornerstone of the scientific method. Following up a remark of van Fraassen, we analyze a problem for the putative Bayesian justification of H-D inference in the case where what we learn from observation is logically stronger than what our theory implies. Firstly, we demonstrate that in such cases the simple Bayesian justification does not necessarily apply. Secondly, we identify a set of sufficient conditions for the mismatch in logical strength to be justifiably ignored as a "harmless idealization''. Thirdly, we argue, based upon scientific examples, that the pattern of H-D inference of which there is a ready Bayesian justification is only rarely the pattern that one actually finds at work in science. Whatever the other virtues of Bayesianism, the idea that it yields a simple justification of a pervasive pattern of scientific inference appears to have been oversold.