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.

Biologists studying ecology and evolution use the term “population” in many different ways. Yet little philosophical analysis of the concept has been done, either by biologists or philosophers, in contrast to the voluminous literature on the concept of “species.” This is in spite of the fact that “population” is arguably a far more central concept in ecological and evolutionary studies than “species” is. The fact that such a central concept has been employed in so many different ways is potentially problematic (...) for the reason that inconsistent usages (especially when the usage has not been made explicit) might lead to false controversies in which disputants are simply talking past one another. However, the inconsistent usages are not the only, or even the most important reason to examine the concept. If any set of organisms is legitimately called a “population,” selection and drift processes become purely arbitrary, too. Moreover, key ecological variables, such as abundance and distribution, depend on a nonarbitrary way of identifying populations. I sketch the beginnings of a population concept, drawing inspiration from the Ghiselin-Hull individuality thesis, and show why some alternative approaches are nonstarters. (shrink)

I defend a view of the species category, pluralistic realism, which is designed to do justice to the insights of many different groups of systematists. After arguing that species are sets and not individuals, I proceed to outline briefly some defects of the biological species concept. I draw the general moral that similar shortcomings arise for other popular views of the nature of species. These shortcomings arise because the legitimate interests of biology are diverse, and these diverse interests are reflected (...) in different legitimate approaches to the classification of organisms. In the final section, I show briefly how the pluralistic approach can help to illuminate some areas of biological and philosophical dispute. (shrink)

We distinguish dynamical and statistical interpretations of evolutionary theory. We argue that only the statistical interpretation preserves the presumed relation between natural selection and drift. On these grounds we claim that the dynamical conception of evolutionary theory as a theory of forces is mistaken. Selection and drift are not forces. Nor do selection and drift explanations appeal to the (sub-population-level) causes of population level change. Instead they explain by appeal to the statistical structure of populations. We briefly discuss the implications (...) of the statistical interpretation of selection for various debates within the philosophy of biologythe `explananda of selection' debate and the `units of selection' debate. (shrink)

There are two competing interpretations of the modern synthesis theory of evolution: the dynamical (also know as ‘traditional’) and the statistical. The dynamical interpretation maintains that explanations offered under the auspices of the modern synthesis theory articulate the causes of evolution. It interprets selection and drift as causes of population change. The statistical interpretation holds that modern synthesis explanations merely cite the statistical structure of populations. This paper offers a defense of statisticalism. It argues that a change in trait frequencies (...) in a population can be attributed only to selection or drift against the background of a particular statistical description of the population. The traditionalist supposition that selection and drift are description‐independent causes of population change leads the dynamical interpretation into a dilemma: it must face a contradiction or accept the loss of explanatory power. (shrink)

In evolutionary biology changes in population structure are explained by citing trait fitness distribution. I distinguish three interpretations of fitness explanations—the Two‐Factor Model, the Single‐Factor Model, and the Statistical Interpretation—and argue for the last of these. These interpretations differ in their degrees of causal commitment. The first two hold that trait fitness distribution causes population change. Trait fitness explanations, according to these interpretations, are causal explanations. The last maintains that trait fitness distribution correlates with population change but does not cause (...) it. My defense of the Statistical Interpretation relies on a distinctive feature of causation. Causes conform to the Sure Thing Principle. Trait fitness distributions, I argue, do not. *Received July 2009; revised October 2009. †To contact the author, please write to: Department of Philosophy/Institute for the History, Philosophy of Science and Technology, University of Toronto, Victoria College, 91 Charles Street West, Toronto, ON M5S 1K7, Canada; e‐mail: [email protected]. (shrink)

Traditionally a scientific theory is viewed as based on universal laws of nature that serve as axioms for logical deduction. In analyzing the logical structure of evolutionary biology, Elisabeth Lloyd argues that the semantic account is more appropriate and powerful. This book will be of interest to biologists and philosophers alike.

Recently, several philosophers have challenged the view that evolutionary theory is usefully understood by way of an analogy with Newtonian mechanics. Instead, they argue that evolutionary theory is merely a statistical theory. According to this alternate approach, natural selection and random genetic drift are not even causes, much less forces. I argue that, properly understood, the Newtonian analogy is unproblematic and illuminating. I defend the view that selection and drift are causes in part by attending to a pair of important (...) distinctions—that between process and product and that between natural selection and fitness. (shrink)

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.

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)

I discuss two subjects in Samir Okasha’s excellent book, Evolution and the Levels of Selection. In consonance with Okasha’s critique of the conventionalist view of the units of selection problem, I argue that conventionalists have not attended to what realists mean by group, individual, and genic selection. In connection with Okasha’s discussion of the Price equation and contextual analysis, I discuss whether the existence of these two quantitative frameworks is a challenge to realism.

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)

Race is a prominent category in medicine. Epidemiologists describe how rates of morbidity and mortality vary with race, and doctors consider the race of their patients when deciding whether to test them for sickle‐cell anemia or what drug to use to treat their hypertension. At the same time, critics of racial classification say that race is not real but only an illusion or that race is scientifically meaningless. In this paper, I explain how race is used in medicine as a (...) proxy for genes that encode drug metabolizing enzymes and how a proper understanding of race calls into doubt the practice of treating race as a marker of any medically relevant genetic trait. (shrink)

While evolutionary thinking is increasingly becoming popular in fields of investigation outside the biological sciences, it remains unclear how helpful it is there and whether it actually yields good explanations of the phenomena under study. Here we examine the ontology of a recent approach to applying evolutionary thinking outside biology, the generalized Darwinism approach proposed by Geoffrey Hodgson and Thorbjørn Knudsen. We examine the ontology of populations in biology and in GD, and argue that biological evolutionary theory sets ontological criteria (...) that GD fails to meet. We suggest two options to revise the population concept in GD: reformulating the concept in terms of inheritance and reproduction such that it comes to pick out individuals similar to evolving populations, or trying to build an adequate population concept on a principle of differential retention instead of differential reproduction. 1 Introduction2 Generalized Darwinism2.1 What is generalized Darwinism?2.2 Darwinian principles3 The Ontology of Generalized Darwinism: What Are Populations?3.1 The population concept of generalized Darwinism3.2 The population concept in evolutionary theory4 Locating Evolving Systems in Generalized Darwinism5 Conclusion. (shrink)

Evolutionary processes such as natural selection and random drift are commonly regarded as causes of population-level change. We respond to a recent challenge that drift and selection are best understood as statistical trends, not causes. Our reply appeals to manipulation as a strategy for uncovering causal relationships: if you can systematically manipulate variable A to bring about a change in variable B, then A is a cause of B. We argue that selection and drift can be systematically manipulated to produce (...) different kinds of population-level change. They should therefore be regarded as causes. (shrink)

We critically examine Denis Walsh’s latest attack on the causalist view of fitness. Relying on Judea Pearl’s Sure-Thing Principle and geneticist John Gillespie’s model for fitness, Walsh has argued that the causal interpretation of fitness results in a reductio. We show that his conclusion only follows from misuse of the models, that is, (1) the disregard of the real biological bearing of the population-size parameter in Gillespie’s model and (2) the confusion of the distinction between ordinary probability and Pearl’s causal (...) probability. Properly understood, the models used by Walsh do not threaten the causalist view of fitness. (shrink)

Recent discussions in the philosophy of biology have brought into question some fundamental assumptions regarding evolutionary processes, natural selection in particular. Some authors argue that natural selection is nothing but a population-level, statistical consequence of lower-level events (Matthen and Ariew [2002]; Walsh et al. [2002]). On this view, natural selection itself does not involve forces. Other authors reject this purely statistical, population-level account for an individual-level, causal account of natural selection (Bouchard and Rosenberg [2004]). I argue that each of these (...) positions is right in one way, but wrong in another; natural selection indeed takes place at the level of populations, but it is a causal process nonetheless. (shrink)

We have argued elsewhere that: (A) Natural selection is not a cause of evolution. (B) A resolution-of-forces (or vector addition) model does not provide us with a proper understanding of how natural selection combines with other evolutionary influences. These propositions have come in for criticism recently, and here we clarify and defend them. We do so within the broad framework of our own “hierarchical realization model” of how evolutionary influences combine.

Elliott Sober and his defenders think of selection, drift, mutation, and migration as distinct evolutionary forces. This paper exposes an ambiguity in Sober's account of the force of selection: sometimes he appears to equate the force of selection with variation in fitness, sometimes with ‘selection for properties’. Sober's own account of fitness as a property analogous to life-expectancy shows how the two conceptions come apart. Cases where there is selection against variance in offspring number also show that selection and drift (...) cannot be distinguished in the way Sober hopes for. These issues have significance beyond the parochial matter of the coherence of Sober's system. There is no good principled answer to the question of which features of a population should count among the contributors to fitness. This means there is no non-arbitrary account of the nature of selection. (shrink)

I defend a view of the species category, pluralistic realism, which is designed to do justice to the insights of many different groups of systematists. After arguing that species are sets and not individuals, I proceed to outline briefly some defects of the biological species concept. I draw the general moral that similar shortcomings arise for other popular views of the nature of species. These shortcomings arise because the legitimate interests of biology are diverse, and these diverse interests are reflected (...) in different legitimate approaches to the classification of organisms. In the final section, I show briefly how the pluralistic approach can help to illuminate some areas of biological and philosophical dispute. (shrink)

Judea Pearl has been at the forefront of research in the burgeoning field of causal modeling, and Causality is the culmination of his work over the last dozen or so years. For philosophers of science with a serious interest in causal modeling, Causality is simply mandatory reading. Chapter 2, in particular, addresses many of the issues familiar from works such as Causation, Prediction and Search by Peter Spirtes, Clark Glymour, and Richard Scheines. But philosophers with a more general interest in (...) causation will also profit from reading Pearl’s book, especially the material in chapters 7, 9, and 10, which is self-contained and less technical than other parts of the book. The present review is aimed primarily at readers of the second type. (shrink)

I criticize some arguments against the causal interpretability of population genetics put forward by Denis Walsh ([2007], [2010]). In particular, I seek to undermine the contention that population genetics exhibits frame of reference relativity or subjectivity with respect to its formal representations. I also show that classical population genetics does not fall foul of some criteria for causal representation put forward by James Woodward ([2003]), although those criteria do undermine some causalist stances. 1 Introduction2 Modularity3 The Crucially Important Point4 The (...) Gillespie Case: Density-Dependent Selection5 Conclusion. (shrink)

At least below the level of species, biological populations are not mind‐independent objects that scientists discover. Rather, biological populations are pragmatically constructed as objects of investigation according to the aims, interests, and values that inform particular research contexts. The relations among organisms that are constitutive of population‐level phenomena (e.g., mating propensity, genealogy, and competition) occur as matters of degree and so give rise to statistically defined open‐ended biological systems. These systems are rendered discrete units to satisfy practical needs and theoretical (...) preferences associated with specific contexts of investigation. While it may be possible to defend a realist position regarding biological relations among organisms, biological populations are “made” when contextual features determine which kinds and degrees of relations to privilege over others, and so how to bound genes in space and time. Consequently, the objectivity of population‐based approaches to species genome diversity cannot rest in the mind‐independence of populations themselves. (shrink)

Species pluralism gives us reason to doubt the existence of the species category. The problem is not that species concepts are chosen according to our interests or that pluralism and the desire for hierarchical classifications are incompatible. The problem is that the various taxa we call 'species' lack a common unifying feature.

In this sequence of philosophical essays about natural science, the author argues that fundamental explanatory laws, the deepest and most admired successes of modern physics, do not in fact describe regularities that exist in nature. Cartwright draws from many real-life examples to propound a novel distinction: that theoretical entities, and the complex and localized laws that describe them, can be interpreted realistically, but the simple unifying laws of basic theory cannot.

Causation is in trouble—at least as it is pictured in current theories in philosophy and in economics as well, where causation is also once again in fashion. In both disciplines the accounts of causality on offer are either modelled too closely on one or another favoured method for hunting causes or on assumptions about the uses to which causal knowledge can be put—generally for predicting the results of our efforts to change the world. The first kind of account supplies no (...) reason to think that causal knowledge, as it is pictured, is of any use; the second supplies no reason to think our best methods will be reliable for establishing causal knowledge. So, if these accounts are all there is to be had, how do we get from method to use? Of what use is knowledge of causal laws that we work so hard to obtain? (shrink)

La autora presenta algunas criticas generales al proyecto de reducir las leyes causales a probabilidades. Además, muestra que las leyes causales son imprescindibles para poder diferenciar las strategias efectivas de las que no lo son y da un criterio para considerar cuando podemos deducir causalidad a través de datos estadísticos.

We examine, from the partial structures perspective, two forms of applicability of mathematics: at the “bottom” level, the applicability of theoretical structures to the “appearances”, and at the “top” level, the applicability of mathematical to physical theories. We argue that, to accommodate these two forms of applicability, the partial structures approach needs to be extended to include a notion of “partial homomorphism”. As a case study, we present London's analysis of the superfluid behavior of liquid helium in terms of Bose‐Einstein (...) statistics. This involved both the introduction of group theory at the top level, and some modeling at the “phenomenological” level, and thus provides a nice example of the relationships we are interested in. We conclude with a discussion of the “autonomy” of London's model. (shrink)

-/- Because species names play an important role in scientific communication, it is more important that species be understood to be taxa than that they be equated with functional ecological or evolutionary entities. Although most biologists would agree that taxa are composed of organisms that share a unique common history, 2 major challenges remain in developing a species-as-taxa concept. First, grouping: in the face of genealogical discordance at all levels in the taxonomic hierarchy, how can we understand the nature of (...) taxa? Second, ranking: what criteria should be used to designate certain taxa in a nested series as being species? The grouping problem can be solved by viewing taxa as exclusive groups of organisms— sets of organisms that form a clade for a plurality of the genome (more than any conflicting set). However, no single objective criterion of species rank can be proposed. Instead, the species rank should be assigned by practitioners based on the semisubjective application of a set of species-ranking criteria. Although these criteria can be designed to yield species taxa that approximately match the ecological, evolutionary, and morphological entities that taxonomists have traditionally associated with the species rank, such a correspondence cannot be enforced without undermining the assumption that species are taxa. The challenge and art of monography is to use genealogical and other kinds of data to assign all organisms to one and only one species-ranked taxon. Various implications of the species-as-ranked-taxa view are discussed, including the synchronic nature of taxa, fossil species, the treatment of hybrids, and species nomenclature. I conclude that, although challenges remain, adopting the view that species are ranked taxa will facilitate a much-needed revolution in taxonomy that will allow it to better serve the biodiversity informatic needs of the 21st century. (shrink)

This paper brings needed clarity to the influential view that species are cohesive entities held together by gene flow, and then develops an empirical argument against that view: Neglected data suggest gene flow is neither necessary nor sufficient for species cohesion. Implications are discussed. ‡I'm grateful to Rob Wilson, Alex Rueger and Lindley Darden for important comments on earlier drafts, and to Joseph Nagel, Heather Proctor, Ken Bond, members of the DC History and Philosophy of Biology reading group, and audience (...) members at the November 2006 meeting of the PSA, for helpful comments or discussion. Social Sciences and Humanities Research Council of Canada fellowship 752-2005-1208 supported research. †To contact the author, please write to: Philosophy Department, University of Wisconsin–Madison, 5185 Helen C. White Hall, 600 North Park Street, Madison, WI 53706; e-mail: [email protected]. (shrink)

We argue for a new conventionalism about many kinds of evolutionary groups, including clades, cohesive units, and populations. This rejects a consensus, which says that given any one of the many legitimate grouping concepts, only objective biological facts determine whether a collection is such a group. Surprisingly, being any one kind of evolutionary group typically depends on which of many incompatible values are taken by suppressed variables. This is a novel pluralism underlying most any one group concept, rather than a (...) familiar pluralism claiming many concepts are legitimate. Consequently, we must help biological facts determine grouphood, even when given a single grouping concept. (shrink)

A far-reaching and influential view in evolutionary biology claims that species are cohesive units held together by gene flow. Biologists have recognized empirical problems facing this view; after sharpening the expression of the view, we present novel conceptual problems for it. At the heart of these problems is a distinction between two importantly different concepts of cohesion, what we call integrative and response cohesion. Acknowledging the distinction problematizes both the explanandum of species cohesion and the explanans of gene flow that (...) are central to the view we discuss. We conclude by tracing four broader implications for the study and conceptualization of species. (shrink)

The book presents a new way of understanding Darwinism and evolution by natural selection, combining work in biology, philosophy, and other fields.

It is often supposed that the spectacular successes of our modern mathematical sciences support a lofty vision of a world completely ordered by one single elegant theory. In this book Nancy Cartwright argues to the contrary. When we draw our image of the world from the way modern science works - as empiricism teaches us we should - we end up with a world where some features are precisely ordered, others are given to rough regularity and still others behave in (...) their own diverse ways. This patchwork makes sense when we realise that laws are very special productions of nature, requiring very special arrangements for their generation. Combining classic and newly written essays on physics and economics, The Dappled World carries important philosophical consequences and offers serious lessons for both the natural and the social sciences. (shrink)

This paper aims to illustrate one of the primary goals of the philosophy of biology⎯namely, the examination of central concepts in biological theory and practice⎯through an analysis of the concepts of population and metapopulation in evolutionary biology and ecology. I will first provide a brief background for my analysis, followed by a characterization of my proposed concepts: the causal interactionist concepts of population and metapopulation. I will then illustrate how the concepts apply to six cases that differ in their population (...) structure; this analysis will also serve to flesh out and defend the concepts a bit more. Finally, I will respond to some possible questions that my analysis may have raised and then conclude briefly. (shrink)