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.

ORIGIN OF SPECIES. INTRODUCTION. When on board HMS 'Beagle,' as naturalist, I was ranch struck with certain facts in the distribution of the organic beings ...

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.

Woodward's long awaited book is an attempt to construct a comprehensive account of causation explanation that applies to a wide variety of causal and explanatory claims in different areas of science and everyday life. The book engages some of the relevant literature from other disciplines, as Woodward weaves together examples, counterexamples, criticisms, defenses, objections, and replies into a convincing defense of the core of his theory, which is that we can analyze causation by appeal to the notion of manipulation.

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)

David Hull's analysis of conceptual change in science, as presentedin his book, Science as a Process (1988), provides a useful framework for understanding one of the scientific controversies in which he actively and constructively intervened, the units of selectiondebates in evolutionary biology. What follows is a brief overview ofthose debates and some reflections on them.

Conflicting accounts of the role of mathematics in our physical theories can be traced to two principles. Mathematics appears to be both (1) theoretically indispensable, as we have no acceptable non-mathematical versions of our theories, and (2) metaphysically dispensable, as mathematical entities, if they existed, would lack a relevant causal role in the physical world. I offer a new account of a role for mathematics in the physical sciences that emphasizes the epistemic benefits of having mathematics around when we do (...) science. This account successfully reconciles theoretical indispensability and metaphysical dispensability and has important consequences for both advocates and critics of indispensability arguments for platonism about mathematics. (shrink)

The official model of explanation proposed by the logical empiricists, the covering law model, is subject to familiar objections. The goal of the present paper is to explore an unofficial view of explanation which logical empiricists have sometimes suggested, the view of explanation as unification. I try to show that this view can be developed so as to provide insight into major episodes in the history of science, and that it can overcome some of the most serious difficulties besetting the (...) covering law model. (shrink)

To explain the phenomena in the world of our experience, to answer the question “why?” rather than only the question “what?”, is one of the foremost objectives of all rational inquiry; and especially, scientific research in its various branches strives to go beyond a mere description of its subject matter by providing an explanation of the phenomena it investigates. While there is rather general agreement about this chief objective of science, there exists considerable difference of opinion as to the function (...) and the essential characteristics of scientific explanation. In the present essay, an attempt will be made to shed some light on these issues by means of an elementary survey of the basic pattern of scientific explanation and a subsequent more rigorous analysis of the concept of law and of the logical structure of explanatory arguments. (shrink)

This paper examines contemporary attempts to explicate the explanatory role of mathematics in the physical sciences. Most such approaches involve developing so-called mapping accounts of the relationships between the physical world and mathematical structures. The paper argues that the use of idealizations in physical theorizing poses serious difficulties for such mapping accounts. A new approach to the applicability of mathematics is proposed.

Biological theory demands a clear organism concept, but at present biologists cannot agree on one. They know that counting particular units, and not counting others, allows them to generate explanatory and predictive descriptions of evolutionary processes. Yet they lack a unified theory telling them which units to count. In this paper, I offer a novel account of biological individuality, which reconciles conflicting definitions of ‘organism’ by interpreting them as describing alternative realisers of a common functional role, and then defines individual (...) organisms as essentially possessing some mechanisms that play this role. (shrink)

Science depends on judgments of the bearing of evidence on theory. Scientists must judge whether an observation or the result of an experiment supports, disconfirms, or is simply irrelevant to a given hypothesis. Similarly, scientists may judge that, given all the available evidence, a hypothesis ought to be accepted as correct or nearly so, rejected as false, or neither. Occasionally, these evidential judgments can be made on deductive grounds. If an experimental result strictly contradicts a hypothesis, then the truth of (...) the data deductively entails the falsity of the hypothesis. In the great majority of cases, however, the connection between evidence and hypothesis is non-demonstrative, or inductive. In particular, this is so whenever a general hypothesis is inferred to be correct on the basis of the available data, since the truth of the data will not deductively entail the truth of the hypothesis. It always remains possible that the hypothesis is false even though the data are correct. (shrink)

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. (shrink)

Does mathematics ever play an explanatory role in science? If so then this opens the way for scientific realists to argue for the existence of mathematical entities using inference to the best explanation. Elsewhere I have argued, using a case study involving the prime-numbered life cycles of periodical cicadas, that there are examples of indispensable mathematical explanations of purely physical phenomena. In this paper I respond to objections to this claim that have been made by various philosophers, and I discuss (...) potential future directions of research for each side in the debate over the existence of abstract mathematical objects. (shrink)

Offering a new approach to scientific explanation, this book focuses initially on the explaining act itself.

How do we go about weighing evidence, testing hypotheses, and making inferences? The model of " inference to the best explanation " -- that we infer the hypothesis that would, if correct, provide the best explanation of the available evidence--offers a compelling account of inferences both in science and in ordinary life. Widely cited by epistemologists and philosophers of science, IBE has nonetheless remained little more than a slogan. Now this influential work has been thoroughly revised and updated, and features (...) a new introduction and two new chapters. Inference to the Best Explanation is an unrivaled exposition of a theory of particular interest in the fields both of epistemology and the philosophy of science. (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)

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)

How do we go about weighing evidence, testing hypotheses, and making inferences? According to the model of _Inference to the Best Explanation_, we work out what to infer from the evidence by thinking about what would actually explain that evidence, and we take the ability of a hypothesis to explain the evidence as a sign that the hypothesis is correct. In _Inference to the Best Explanation_, Peter Lipton gives this important and influential idea the development and assessment it deserves. The (...) second edition has been substantially enlarged and reworked, with a new chapter on the relationship between explanation and Bayesianism, and an extension and defence of the account of contrastive explanation. It also includes an expanded defence of the claims that our inferences really are guided by diverse explanatory considerations, and that this pattern of inference can take us towards the truth. This edition of _Inference to the Best Explanation_ has also been updated throughout and includes a new bibliography. (shrink)

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. (shrink)

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 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)

Kin selection and multilevel selection are alternative approaches for studying the evolution of social behaviour, the relation between which has long been a source of controversy. Many recent theorists regard the two approaches as ultimately equivalent, on the grounds that gene frequency change can be correctly expressed using either. However, this shows only that the two are formally equivalent, not that they offer equally good causal representations of the evolutionary process. This article articulates the notion of an ‘adequate causal representation’ (...) using causal graphs, and then seeks to identify circumstances under which kin and multilevel selection do and do not satisfy the test of causal adequacy. 1 Introduction2 The KS and MLS Approaches2.1 The MLS decomposition2.2 The KS decomposition3 Equivalence and Causality4 Two Problem Cases4.1 The non-social trait case4.2 Genotypic selection with meiotic drive5 Casual Adequacy: A Graphical Approach5.1 The basic idea5.2 Graphs with individual and group variables5.3 Cases where KS is causally adequate5.4 Cases where MLS is causally adequate6 Discussion6.1 Relation to previous work. (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)

Altruism has both an evolutionary and a psychological meaning. As the term is used in evolutionary theory, a trait is deemed altruistic if it reduces the fitness of the actor and enhances the fitness of someone else. In its psychological sense, the thesis that we have altruistic ultimate motives asserts that we care about the welfare of others, not just as a means of enhancing our own well-being, but as an end in itself. In Unto Others (hereafter UO), we consider (...) both evolutionary altruism (Part I) and psychological altruism (Part II) from an evolutionary perspective. We propose a multilevel selection theory to explain how altruism can evolve from group selection, based on Darwin’s solution to the problem. Our presentation of multilevel selection theory expands the range of topics beyond altruism in other respects as well. In Part I1 of UO, we evaluate the arguments, both conceptual and empirical, that philosophers and psychologists have advanced in an attempt to choose between psychological egoism and motivational pluralism. We conclude that these arguments are inconclusive. (shrink)

In an incendiary 2010 Nature article, M. A. Nowak, C. E. Tarnita, and E. O. Wilson present a savage critique of the best-known and most widely used framework for the study of social evolution, W. D. Hamilton’s theory of kin selection. More than a hundred biologists have since rallied to the theory’s defence, but Nowak et al. maintain that their arguments ‘stand unrefuted’. Here I consider the most contentious claim Nowak et al. defend: that Hamilton’s rule, the core explanatory principle (...) of kin selection theory, ‘almost never holds’. I first distinguish two versions of Hamilton’s rule in contemporary theory: a special version (HRS) that requires restrictive assumptions, and a general version (HRG) that does not. I then show that Nowak et al. are most charitably construed as arguing that HRS almost never holds, while HRG buys its generality at the expense of explanatory power. While their arguments against HRS are fairly uncontroversial, their arguments against HRG are more contentious, yet these have been largely overlooked in the ensuing furore. I consider the arguments for and against the explanatory value of HRG, with a view to assessing what exactly is at stake in the debate. I suggest that the debate hinges on issues concerning the causal interpretability of regression coefficients, and concerning the explanatory function Hamilton’s rule is intended to serve. (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)

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)

An updated and extended second edition supporting the findings of its well-known predecessor which claimed that courts employ common-sense notions of causation in determining legal responsibility.

Woodward's long awaited book is an attempt to construct a comprehensive account of causation explanation that applies to a wide variety of causal and explanatory claims in different areas of science and everyday life. The book engages some of the relevant literature from other disciplines, as Woodward weaves together examples, counterexamples, criticisms, defences, objections, and replies into a convincing defence of the core of his theory, which is that we can analyse causation by appeal to the notion of manipulation.

The conflation of two fundamentally distinct issues has generated serious confusion in the philosophical and biological literature concerning the units of selection. The questions of how a unit of selection is defined, theoretically, is rarely distinguished from the question of how to determine the empirical accuracy of claims--either specific or general--concerning which unit(s) are undergoing selection processes. In this paper, I begin by refining a definition of the unit of selection, first presented by William Wimsatt, that is grounded in the (...) structure of natural selection models. I then explore the implications of this definition for empirical evaluation of group selection models. I examine an objection to Wimsatt’s definition raised by Elliott Sober, and I conclude by indicating how the proposed refined definition can clarify which sorts of empirical evidence could be brought to bear on group selection controversies. (shrink)