In this paper we first briefly review Bell's (1964, 1966) Theorem to see how it invalidates any deterministic "hidden variable" account of the apparent indeterminacy of quantum mechanics (QM). Then we show that quantum uncertainty, at the level of DNA mutations, can "percolate" up to have major populational effects. Interesting as this point may be it does not show any autonomous indeterminism of the evolutionary process. In the next two sections we investigate drift and natural selection as the locus of (...) autonomous biological indeterminacy. Here we conclude that the population-level indeterminacy of natural selection and drift are ultimately based on the assumption of a fundamental indeterminacy at the level of the lives and deaths of individual organisms. The following section examines this assumption and defends it from the determinists' attack. Then we show that, even if one rejects the assumption, there is still an important reason why one might think evolutionary theory (ET) is autonomously indeterministic. In the concluding section we contrast the arguments we have mounted against a deterministic hidden variable account of ET with the proof of the impossibility of such an account of QM. (shrink)

Millstein [Bio. Philos. 17 (2002) 33] correctly identies a serious problem with the view that natural selection and random drift are not conceptually distinct. She offers a solution to this problem purely in terms of differences between the processes of selection and drift. I show that this solution does not work, that it leaves the vast majority of real biological cases uncategorized. However, I do think there is a solution to the problem she raises, and I offer it here. My (...) solution depends on solving the biological analogue of the reference class problem in probability theory and on the reality of individual fitnesses. (shrink)

We argue that a fashionable interpretation of the theory of natural selection as a claim exclusively about populations is mistaken. The interpretation rests on adopting an analysis of fitness as a probabilistic propensity which cannot be substantiated, draws parallels with thermodynamics which are without foundations, and fails to do justice to the fundamental distinction between drift and selection. This distinction requires a notion of fitness as a pairwise comparison between individuals taken two at a time, and so vitiates the interpretation (...) of the theory as one about populations exclusively. (shrink)

Among the liveliest disputes in evolutionary biology today are disputes concerning the role of chance in evolution--more specifically, disputes concerning the relative evolutionary importance of natural selection vs. so-called "random drift". The following discussion is an attempt to sort out some of the broad issues involved in those disputes. In the first half of this paper, I try to explain the differences between evolution by natural selection and evolution by random drift. On some common construals of "natural selection", those two (...) modes of evolution are completely indistinguishable. Even on a proper construal of "natural selection", it is difficult to distinguish between the "improbable results of natural selection" and evolution by random drift. In the second half of this paper, I discuss the variety of positions taken by evolutionists with respect to the evolutionary importance of random drift vs. natural selection. I will then consider the variety of issues in question in terms of a conceptual distinction often used to describe the rise of probabilistic thinking in the sciences. I will argue, in particular, that what is going on here is not, as might appear at first sight, just another dispute about the desirability of "stochastic" vs. "deterministic" theories. Modern evolutionists do not argue so much about whether evolution is stochastic, but about how stochastic it is. (shrink)

Recently advocates of the propensity interpretation of fitness have turned critics. To accommodate examples from the population genetics literature they conclude that fitness is better defined broadly as a family of propensities rather than the propensity to contribute descendants to some future generation. We argue that the propensity theorists have misunderstood the deeper ramifications of the examples they cite. These examples demonstrate why there are factors outside of propensities that determine fitness. We go on to argue for the more general (...) thesis that no account of fitness can satisfy the desiderata that have motivated the propensity account. (shrink)

The central point of this essay is to demonstrate the incommensurability of ‘Darwinian fitness’ with the numeric values associated with reproductive rates used in population genetics. While sometimes both are called ‘fitness’, they are distinct concepts coming from distinct explanatory schemes. Further, we try to outline a possible answer to the following question: from the natural properties of organisms and a knowledge of their environment, can we construct an algorithm for a particular kind of organismic life-history pattern that itself will (...) allow us to predict whether a type in the population will increase or decrease relative to other types? Introduction Darwinian fitness Reproductive fitness and genetical models of evolution The models of reproductive fitness 4.1 The Standard Viability Model 4.2 Frequency-dependent selection 4.3 Fertility models 4.4 Overlapping generations Fitness as outcome 5.1 Fitness as actual increase in type 5.2 Fitness as expected increase in type 5.2.1 Expected increase within a generation 5.2.2 Expected increase between generations 5.2.3 Postponed reproductive fitness effects The book-keeping problem Conclusion. (shrink)

The central point of this essay is to demonstrate the incommensurability of ‘Darwinian fitness’ with the numeric values associated with reproductive rates used in population genetics. While sometimes both are called ‘fitness’, they are distinct concepts coming from distinct explanatory schemes. Further, we try to outline a possible answer to the following question: from the natural properties of organisms and a knowledge of their environment, can we construct an algorithm for a particular kind of organismic life-history pattern that itself will (...) allow us to predict whether a type in the population will increase or decrease relative to other types? 1. Introduction2. Darwinian fitness3. Reproductive fitness and genetical models of evolution4. The models of reproductive fitness4.1 The Standard Viability Model4.2 Frequency-dependent selection4.3 Fertility models4.4 Overlapping generations5. Fitness as outcome5.1 Fitness as actual increase in type5.2 Fitness as expected increase in type5.2.1 Expected increase within a generation5.2.2 Expected increase between generations5.2.3 Postponed reproductive fitness effects6. The book-keeping problem7. Conclusion. (shrink)

One controversy about the existence of so called evolutionary forces such as natural selection and random genetic drift concerns the sense in which such “forces” can be said to interact. In this paper I explain how natural selection and random drift can interact. In particular, I show how population-level probabilities can be derived from individual-level probabilities, and explain the sense in which natural selection and drift are embodied in these population-level probabilities. I argue that whatever causal character the individual-level probabilities (...) have is then shared by the population-level probabilities, and that natural selection and random drift then have that same causal character. Moreover, natural selection and drift can then be viewed as two aspects of probability distributions over frequencies in populations of organisms. My characterization of population-level probabilities is largely neutral about what interpretation of probability is required, allowing my approach to support various positions on biological probabilities, including those which give biological probabilities one or another sort of causal character. ‡This paper has benefited from feedback on and discussions of this and earlier work. I want to thank André Ariew, Matt Barker, Lindley Darden, Patrick Forber, Nancy Hall, Mohan Matthen, Samir Okasha, Jeremy Pober, Robert Richardson, Alex Rosenberg, Eric Seidel, Denis Walsh, and Bill Wimsatt. †To contact the author, please write to: Department of Philosophy, University of Alabama at Birmingham, HB 414A, 900 13th Street South, Birmingham, AL 35294-1260; e-mail: [email protected]. (shrink)

The present edition provides a detailed and accessible discussion ofhis theories and adds an account of the immediate responses to the book on publication.

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.

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.

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)

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 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)

In this paper, it is argued that selection and drift might be distinct. This contradicts recent arguments by Brandon (forthcoming) and Matthen and Ariew (2002) that such a distinction “violates sound probabilistic thinking” (Matthen and Ariew 2002, 62). While their arguments might be valid under certain assumptions, they overlook a possible way to make sense of the distinction. Whether this distinction makes sense, I argue, depends on the source of probabilities in natural selection. In particular, if the probabilities used in (...) defining fitness values are at least partly a result of abstracting from or ignoring certain features of the environment, then selection and drift might in fact be causally distinct. (shrink)

The propensity interpretation of fitness (PIF) is commonly taken to be subject to a set of simple counterexamples. We argue that three of the most important of these are not counterexamples to the PIF itself, but only to the traditional mathematical model of this propensity: fitness as expected number of offspring. They fail to demonstrate that a new mathematical model of the PIF could not succeed where this older model fails. We then propose a new formalization of the PIF that (...) avoids these (and other) counterexamples. By producing a counterexample-free model of the PIF, we call into question one of the primary motivations for adopting the statisticalist interpretation of fitness. In addition, this new model has the benefit of being more closely allied with contemporary mathematical biology than the traditional model of the PIF. (shrink)

The concept of "fitness" is a notion of central importance to evolutionary theory. Yet the interpretation of this concept and its role in explanations of evolutionary phenomena have remained obscure. We provide a propensity interpretation of fitness, which we argue captures the intended reference of this term as it is used by evolutionary theorists. Using the propensity interpretation of fitness, we provide a Hempelian reconstruction of explanations of evolutionary phenomena, and we show why charges of circularity which have been levelled (...) against explanations in evolutionary theory are mistaken. Finally, we provide a definition of natural selection which follows from the propensity interpretation of fitness, and which handles all the types of selection discussed by biologists, thus improving on extant definitions. (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)

The latter half of the twentieth century has been marked by debates in evolutionary biology over the relative significance of natural selection and random drift: the so-called “neutralist/selectionist” debates. Yet John Beatty has argued that it is difficult, if not impossible, to distinguish the concept of random drift from the concept of natural selection, a claim that has been accepted by many philosophers of biology. If this claim is correct, then the neutralist/selectionist debates seem at best futile, and at worst, (...) meaningless. I reexamine the issues that Beatty raises, and argue that random drift and natural selection, conceived as processes, can be distinguished from one another. (shrink)

A hitherto neglected form of explanation is explored, especially its role in population genetics. “Statistically abstractive explanation” (SA explanation) mandates the suppression of factors probabilistically relevant to an explanandum when these factors are extraneous to the theoretical project being pursued. When these factors are suppressed, the explanandum is rendered uncertain. But this uncertainty traces to the theoretically constrained character of SA explanation, not to any real indeterminacy. Random genetic drift is an artifact of such uncertainty, and it is therefore wrong (...) to reify it as a cause of evolution or as a process in its own right. *Received July 2009. †To contact the author, please write to: Department of Philosophy, University of Toronto, 170 St. George St., Toronto, ON M5R 2M8, Canada; e‐mail: [email protected]. (shrink)

The concept of fitness, central to population genetics and to the synthetic theory of evolution, is discussed. After a historical introduction on the origin of this concept, the current meaning of it in population genetics is examined: a cause of the selective process and its quantification. Several difficulties arise for its exact definition. Three adequacy criteria for such a definition are formulated. It is shown that it is impossible to formulate an adequate definition of fitness respecting these criteria. The propensity (...) definition of fitness is presented and rejected. Finally it is argued that fitness is a conceptual device, a useful tool, only for descriptive purposes of selective processes, changing from case to case, and thus devoid of any substantial physical counterpart. Any attempt to its reification is an apport to the metaphysical load evolutionary theory has inherited from Natural Theology. (shrink)

Among philosophers, controversy over the notion of drift in population genetics is ongoing. This is at least partly because the notion of drift has an ambiguous usage among population geneticists. My goal in this paper is to explicate the causal dimension of drift, to say what causal influences are responsible for the stochasticity in population genetics models. It is commonplace for population genetics to oppose the influence of selection to that of drift, and to consider how the dynamics of populations (...) are altered when each has greater or lesser influence. I define the causes that are referred to as drift when researchers speak this way. Introduction Populations and Variant Types The Cause–Effect Ambiguity of Drift Non-directional Factors in Population Genetics How N ev Is Used in Population Genetics Causal Conceptions of Drift 6.1 The Millstein/Beatty conception of drift 6.2 Rosenberg and Bouchard: Drift as initial conditions NINPICs 7.1 Why drift is instituted by NINPICs 7.2 How NINPICS work 7.3 NINPICs and random sampling 7.4 Independent sampling and effective population size 7.5 Variance in progeny number 7.6 Population effects of NINPICs NINPICs and the Stochastic Character of Selection Theory Conclusion Appendix CiteULike Connotea Del.icio.us What's this? (shrink)

Do the sciences aim to uncover the structure of nature, or are they ultimately a practical means of controlling our environment? In Instrumental Biology, or the Disunity of Science, Alexander Rosenberg argues that while physics and chemistry can develop laws that reveal the structure of natural phenomena, biology is fated to be a practical, instrumental discipline. Because of the complexity produced by natural selection, and because of the limits on human cognition, scientists are prevented from uncovering the basic structure of (...) biological phenomena. Consequently, biology and all of the disciplines that rest upon it--psychology and the other human sciences--must aim at most to provide practical tools for coping with the natural world rather than a complete theoretical understanding of it. (shrink)

How do fitness and natural selection relate to other evolutionary factors like architectural constraint, mode of reproduction, and drift? In one way of thinking, drawn from Newtonian dynamics, fitness is one force driving evolutionary change and added to other factors. In another, drawn from statistical thermodynamics, it is a statistical trend that manifests itself in natural selection histories. It is argued that the first model is incoherent, the second appropriate; a hierarchical realization model is proposed as a basis for a (...) statistical treatment. It emerges that natural selection does not cause evolution; it just is evolution. The theory incorporates relations of statistical correlation, but not the kind of causation found in fundamental physical processes. (shrink)

Evolutionary theory is awash with probabilities. For example, natural selection is said to occur when there is variation in fitness, and fitness is standardly decomposed into two components, viability and fertility, each of which is understood probabilistically. With respect to viability, a fertilized egg is said to have a certain chance of surviving to reproductive age; with respect to fertility, an adult is said to have an expected number of offspring.1 There is more to evolutionary theory than the theory of (...) natural selection, and here too one finds probabilistic concepts aplenty. When there is no selection, the theory of neutral evolution says that a gene’s chance of eventually reaching fixation is 1/(2N), where N is the number of organisms in the generation of the diploid population to which the gene belongs. The evolutionary consequences of mutation are likewise conceptualized in terms of the probability per unit time a gene has of changing from one state to another. The examples just mentioned are all “forwarddirected” probabilities; they describe the probability of later events, conditional on earlier events. However, evolutionary theory also uses “backwards probabilities” that describe the probability of a cause conditional on its effects; for example, coalescence theory allows one to calculate the expected number of generations in the past that the genes in the present generation find their most recent common ancestor. (shrink)

The concept of fitness began its career in biology long before evolutionary theory was mathematized. Fitness was used to describe an organism’s vigor, or the degree to which organisms “fit” into their environments. An organism’s success in avoiding predators and in building a nest obviously contribute to its fitness and to the fitness of its offspring, but the peacock’s gaudy tail seemed to be in an entirely different line of work. Fitness, as a term in ordinary language (as in “physical (...) fitness”) and in its original biological meaning, applied to the survival of an organism and its offspring, not to sheer reproductive output (Paul ////; Cronin 1991). Darwin’s separation of natural from sexual selection may sound odd from a modern perspective, but it made sense from this earlier point of view. (shrink)