Does natural selection act primarily on individual organisms, on groups, on genes, or on whole species? The question of levels of selection - on which biologists and philosophers have long disagreed - is central to evolutionary theory and to the philosophy of biology. Samir Okasha's comprehensive analysis gives a clear account of the philosophical issues at stake in the current debate.
We consider the question: under what circumstances can the concept of adaptation be applied to groups, rather than individuals? Gardner and Grafen (2009, J. Evol. Biol.22: 659–671) develop a novel approach to this question, building on Grafen's ‘formal Darwinism’ project, which defines adaptation in terms of links between evolutionary dynamics and optimization. They conclude that only clonal groups, and to a lesser extent groups in which reproductive competition is repressed, can be considered as adaptive units. We re-examine the conditions under (...) which the selection–optimization links hold at the group level. We focus on an important distinction between two ways of understanding the links, which have different implications regarding group adaptationism. We show how the formal Darwinism approach can be reconciled with G.C. Williams’ famous analysis of group adaptation, and we consider the relationships between group adaptation, the Price equation approach to multi-level selection, and the alternative approach based on contextual analysis. (shrink)
Biologists and philosophers of biology typically regard essentialism about speciesas incompatible with modern Darwinian theory. Analytic metaphysicians such asKripke, Putnam and Wiggins, on the other hand, believe that their essentialist thesesare applicable to biological kinds. I explore this tension. I show that standard anti-essentialist considerations only show that species do not have intrinsic essential properties. I argue that while Putnam and Kripke do make assumptions that contradict received biological opinion, their model of natural kinds, suitably modified, is partially applicable to (...) biological species. However, Wiggins'' thesis that organisms belong essentially to their species is untenable, given modern species concepts. I suggest that Putnam''s, Kripke''s and Wiggins'' errors stem from adopting an account of the point of scientific classification which implies that relationally-defined kinds are likely to be of little value, an account which is inapplicable to biology. (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)
Philosophy of biology is a vibrant and growing field. From initial roots in the metaphysics of species (Ghiselin, Hull), questions about whether biology has laws of nature akin to those of physics (Ruse, Hull), and discussions of teleology and function (Grene 1974, Brandon 1981), the field has grown since the 1970s to include a vast range of topics. Over the last few decades, philosophy has had an important impact on biology, partly through following the model of engagement with science that (...) was set by first-wave philosophers of biology like Marjorie Grene, Morton Beckner, David Hull, William Wimsatt and others. Today some parts of philosophy of biology are indistinguishable from theoretical biology. This is due in part to the impetus provided by second-wave philosophers of biology like James Griesemer, John Beatty, William Bechtel, Robert Brandon, Elisabeth Lloyd, and Elliott Sober. Indeed, philosophers have been instrumental in establishing theoretical biology as a field by collaborating with scientists, publishing in science journals, and by taking up conceptual questions at the heart of the biological enterprise. (shrink)
A number of recent biologists have used multi-level selection theory to help explain the major transitions in evolution. I argue that in doing so, they have shifted from a ‘synchronic’ to a ‘diachronic’ formulation of the levels of selection question. The implications of this shift in perspective are explored, in relation to an ambiguity in the meaning of multi-level selection. Though the ambiguity is well-known, it has never before been discussed in the context of the major transitions.
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)
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. (shrink)
Elliott Sober and Ken Waters both raise interesting and difficult challenges for various aspects of the position I set out in Evolution and the Levels of the Selection. I am grateful to them for their penetrating criticisms of my work, and find myself in agreement with many of their points.
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? (shrink)
We critically examine a number of aspects of Grafen’s ‘formal Darwinism’ project. We argue that Grafen’s ‘selection-optimality’ links do not quite succeed in vindicating the working assumption made by behavioural ecologists and others—that selection will lead organisms to exhibit adaptive behaviour—since these links hold true even in the presence of strong genetic and developmental constraints. However we suggest that the selection-optimality links can profitably be viewed as constituting an axiomatic theory of fitness. Finally, we compare Grafen’s project with Fisher’s ‘fundamental (...) theorem of natural selection’, and we speculate about whether Grafen’s results can be extended to a game-theoretic setting. (shrink)
John Harsanyi and John Rawls both used the veil of ignorance thought experiment to study the problem of choosing between alternative social arrangements. With his, Harsanyi tried to show that the veil of ignorance argument leads inevitably to utilitarianism, an argument criticized by Sen, Weymark and others. A quite different use of the veil-of-ignorance concept is found in evolutionary biology. In the cell-division process called meiosis, in which sexually reproducing organisms produce gametes, the chromosome number is halved; when meiosis is (...) fair, each gene has only a fifty percent chance of making it into any gamete. This creates a Mendelian veil of ignorance, which has the effect of aligning the interests of all the genes in an organism. This paper shows how Harsanyi's version of the veil-of-ignorance argument can shed light on Mendelian genetics. There turns out to be an intriguing biological analogue of the impartial observer theorem that is immune from the Sen/Weymark objections to Harsanyi's original. (shrink)
What is science? Is there a real difference between science and myth? Is science objective? Can science explain everything? This Very Short Introduction provides a concise overview of the main themes of contemporary philosophy of science. Beginning with a short history of science to set the scene, Samir Okasha goes on to investigate the nature of scientific reasoning, scientific explanation, revolutions in science, and theories such as realism and anti-realism. He also looks at philosophical issues in particular sciences, including the (...) problem of classification in biology, and the nature of space and time in physics. The final chapter touches on the conflicts between science and religion, and explores whether science is ultimately a good thing. (shrink)
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  (...) 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)
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. (shrink)
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 .. (shrink)
In this paper, I explore Larry Laudan's and Jarrett Leplin's recent claim that empirically equivalent theories may be differentially confirmed. I show that their attempt to prise apart empirical equivalence and epistemic parity commits them to two principles of confirmation that Hempel demonstrated to be incompatible.
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. (shrink)
I examine the argument that scientific theories are typically 'underdetermined' by the data, an argument which has often been used to combat scientific realism. I deal with two objections to the underdetermination argument: (i) that the argument conflicts with the holistic nature of confirmation, and (ii) that the argument rests on an untenable theory/data dualism. I discuss possible responses to both objections, and argue that in both cases the proponent of underdetermination can respond in ways which are individually plausible, but (...) that the best response to the first objection conflicts with the best response to the second. Consequently underdetermination poses less of a problem for scientific realism than has often been thought. (shrink)
Much of biological and economic theorizing takes place by modeling, the indirect study of real-world phenomena by the construction and examination of models. Books and articles about biological and economic theory are often books and articles about models, many of which are highly idealized and chosen for their explanatory power and analytical convenience rather than for their fit with known data sets. Philosophers of science have recognized these facts and have developed literatures about the nature of models, modeling, idealization, as (...) well as testing of models and explanation by models, for both biology and economics. The impetus for this special issue came from our recognition that there is remarkably little overlap between the “modeling in biology” and “modeling in economics” literatures, despite many of the same themes appearing in these literatures. (shrink)
Advocates of the "strong programme" in the sociology of knowledge have argued that, because scientific theories are "underdetermined" by data, sociological factors must be invoked to explain why scientists believe the theories they do. I examine this argument, and the responses to it by J.R. Brown (1989) and L. Laudan (1996). I distinguish between a number of different versions of the underdetermination thesis, some trivial, some substantive. I show that Brown's and Laudan's attempts to refute the sociologists' argument fail. Nonetheless, (...) the sociologists' argument falls to a different criticism, for the version of the underdetermination thesis that the argument requires, has not been shown to be true. (shrink)
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. (shrink)
In their recent book, Elliott Sober and David Wilson (1998) argue that evolutionary biologists have wrongly regarded kinship as the exclusive means by which altruistic behavior can evolve, at the expense of other mechanisms. I argue that Sober and Wilson overlook certain genetical considerations which suggest that kinship is likely to be a more powerful means for generating complex altruistic adaptations than the alternative mechanisms they propose.
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)
This volume explores from multiple perspectives the subtle and interesting relationship between the theory of rational choice and Darwinian evolution. In rational choice theory, agents are assumed to make choices that maximize their utility; in evolution, natural selection 'chooses' between phenotypes according to the criterion of fitness maximization. So there is a parallel between utility in rational choice theory and fitness in Darwinian theory. This conceptual link between fitness and utility is mirrored by the interesting parallels between formal models of (...) evolution and rational choice. The essays in this volume, by leading philosophers, economists, biologists and psychologists, explore the connection between evolution and rational choice in a number of different contexts, including choice under uncertainty, strategic decision making and pro-social behaviour. They will be of interest to students and researchers in philosophy of science, evolutionary biology, economics and psychology. (shrink)
The idea that natural selection can operate on cultural as well as genetic variation is central to recent theories of cultural evolution. This raises an overarching question: how much of traditional evolutionary theory, which was formulated in population-genetic terms, can survive intact once the possibility of cultural inheritance is taken into account? This question is addressed in relation to R. A. Fisher’s “fundamental theorem” of natural selection. Though Fisher’s theorem may appear to be an essentially genetic result, a version of (...) the theorem can be derived that applies to any evolving population, with an arbitrary inheritance mechanism. However, I argue that the biological significance of this “generalized” version of the fundamental theorem is less than that of the Fisherian original. (shrink)
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. (shrink)
In a recent article I compared the problem of theory choice, in which scientists must choose between competing theories, with the problem of social choice, in which society must choose between competing social alternatives. I argued that the formal machinery of social choice theory can be used to shed light on the problem of theory choice in science, an argument that has been criticized by Michael Morreau and Jacob Stegenga. This article replies to Morreau’s and Stegenga’s criticisms.
It is widely agreed that Hume's description of human inductive reasoning is inadequate. But many philosophers think that this inadequacy in no way affects the force of Hume's argument for the unjustifiability of inductive reasoning. I argue that this constellation of opinions contains a serious tension, given that Hume was not merely pointing out that induction is fallible. I then explore a recent diagnosis of where Hume's sceptical argument goes wrong, due to Elliott Sober. Sober argues that Hume committed a (...) quantifier-shift fallacy, i.e. inferred a statement of ?? form from one of ?? form. The implications of this diagnosis for the traditional problem of induction are briefly examined. (shrink)
Many philosophers argue that Bayesian epistemology cannot help us with the traditional Humean problem of induction. I argue that this view is partially but not wholly correct. It is true that Bayesianism does not solve Hume’s problem, in the way that the classical and logical theories of probability aimed to do. However I argue that in one important respect, Hume’s sceptical challenge cannot simply be transposed to a probabilistic context, where beliefs come in degrees, rather than being a yes/no matter.
Marc Lange has criticized my assertion that relative to a Bayesian conception of inductive reasoning, Hume's argument for inductive scepticism cannot be run. I reply that the way in which Lange suggests one should run the Humean argument in a Bayesian framework ignores the fact that in Bayesian models of learning from experience, the domain of an agent's probability measure is exogenously determined. I also show that Lange is incorrect to equate probability distributions which 'support inductive inferences' with probability distributions (...) which assign probability to contingent propositions/events. (shrink)
An evolutionary basis for Bayesian rationality is suggested, by consid- ering 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 which for each signal, chooses an action that maximizes conditional expected output given that signal. An organism using such a policy is behaving as if it were a Bayesian agent with (...) probabilistic beliefs about the states of nature, that it updates by conditionalization, and whose choice behaviour obeys expected util- ity maximization. This suggests a possible route by which Bayes-rational creatures might have evolved. (shrink)
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. (shrink)
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. (shrink)
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. (shrink)