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)

Recently philosophers of biology have argued over whether or not Newtonian mechanics provides a useful analogy for thinking about evolutionary theory. For philosophers, the canonical presentation of this analogy is Sober's. Matthen and Ariew and Walsh, Lewins, and Ariew argue that this analogy is deeply wrong-headed. Here I argue that the analogy is indeed useful, however, not in the way it is usually interpreted. The Newtonian analogy depends on having the proper analogue of Newton's First Law. That analogue is what (...) McShea and Brandon call the Zero Force Evolutionary Law. According to the ZFEL, change, not stasis, is the default state of evolutionary systems. (shrink)

In their 2010 book, Biology’s First Law, D. McShea and R. Brandon present a principle that they call ‘‘ZFEL,’’ the zero force evolutionary law. ZFEL says (roughly) that when there are no evolutionary forces acting on a population, the population’s complexity (i.e., how diverse its member organisms are) will increase. Here we develop criticisms of ZFEL and describe a different law of evolution; it says that diversity and complexity do not change when there are no evolutionary causes.

The cis-regulatory hypothesis is one of the most important claims of evolutionary developmental biology. In this paper I examine the theoretical argument for cis-regulatory evolution and its role within evolutionary theorizing. I show that, although the argument has some weaknesses, it acts as a useful example for the importance of current scientific debates for science education.

On the classical account of evolution by natural selection found in Lewontin and many subsequent authors, ENS is conceived as involving three key ingredients: phenotypic variation, fitness differences, and heredity. Through the analysis of three problem cases involving heredity, I argue that the classical conception is substantially flawed, showing that heredity is not required for selection. I consider further problems with the classical account of ENS arising from conflations between three distinct senses of the central concept of ‘fitness’ and offer (...) an alternative to the classical conception of ENS involving the interaction of distinct evolutionary mechanisms. (shrink)

The Hardy–Weinberg equilibrium has been argued by Sober, Stephens and others to represent the zero-force state for evolutionary biology understood as a theory of forces. I investigate what it means for a model to involve forces, developing an explicit account by defining what the zero-force state is in a general theoretical context. I use this account to show that Hardy–Weinberg equilibrium is not the zero-force state in biology even in the contexts in which it applies, and argue based on this (...) that drift should not be understood as an evolutionary force. (shrink)

The explanatory role of natural selection is one of the long-term debates in evolutionary biology. Nevertheless, the consensus has been slippery because conceptual confusions and the absence of a unified, formal causal model that integrates different explanatory scopes of natural selection. In this study we attempt to examine two questions: (i) What can the theory of natural selection explain? and (ii) Is there a causal or explanatory model that integrates all natural selection explananda? For the first question, we argue that (...) five explananda have been assigned to the theory of natural selection and that four of them may be actually considered explananda of natural selection. For the second question, we claim that a probabilistic conception of causality and the statistical relevance concept of explanation are both good models for understanding the explanatory role of natural selection. We review the biological and philosophical disputes about the explanatory role of natural selection and formalize some explananda in probabilistic terms using classical results from population genetics. Most of these explananda have been discussed in philosophical terms but some of them have been mixed up and confused. We analyze and set the limits of these problems. (shrink)

Fitness is a central but notoriously vexing concept in evolutionary biology. The propensity interpretation of fitness is often regarded as the least problematic account for fitness. It ties an individual's fitness to a probabilistic capacity to produce offspring. Fitness has a clear causal role in evolutionary dynamics under this account. Nevertheless, the propensity interpretation faces its share of problems. We discuss three of these. We first show that a single scalar value is an incomplete summary of a propensity. Second, we (...) argue that the widespread method of “abstracting away” environmental idiosyncrasies by averaging over reproductive output in different environments is not a valid approach when environmental changes are irreversible. Third, we point out that expanding the range of applicability for fitness measures by averaging over more environments or longer time scales (so as to ensure environmental reversibility) reduces one's ability to distinguish selectively relevant differences among individuals because of mutation and eco‐evolutionary feedbacks. This series of problems leads us to conclude that a general value of fitness that is both explanatory and predictive cannot be attained. We advocate for the use of propensity‐compatible methods, such as adaptive dynamics, which can accommodate these difficulties. (shrink)

This article offers three contrasting cases of the use of neutrality and drift in molecular evolution. In the first, neutrality is assumed as a simplest case for modeling. In the second and third, concepts of drift and neutrality are developed within the context of population genetics testing and the development and application of the molecular clock.

The neutral and nearly neutral theories of molecular evolution are sometimes characterized as theories about drift alone, where drift is described solely as an outcome, rather than a process. We argue, however, that both selection and drift, as causal processes, are integral parts of both theories. However, the nearly neutral theory explicitly recognizes alleles and/or molecular substitutions that, while engaging in weakly selected causal processes, exhibit outcomes thought to be characteristic of random drift. A narrow focus on outcomes obscures the (...) significant role of weakly selected causal processes in the nearly neutral theory. (shrink)

There are several different ways in which chance affects evolutionary change. That all of these processes are called “random genetic drift” is in part a due to common elements across these different processes, but is also a product of historical borrowing of models and language across different levels of organization in the biological hierarchy. A history of the concept of drift will reveal the variety of contexts in which drift has played an explanatory role in biology, and will shed light (...) on some of the philosophical controversy surrounding whether drift is a cause of evolutionary change. (shrink)

Denis Walsh has written a striking new defense in this journal of the statisticalist (i.e., noncausalist) position regarding the forces of evolution. I defend the causalist view against his new objections. I argue that the heart of the issue lies in the nature of nonadditive causation. Detailed consideration of that turns out to defuse Walsh’s ‘description‐dependence’ critique of causalism. Nevertheless, the critique does suggest a basis for reconciliation between the two competing views. *Received December 2009; revised December 2009. †To contact (...) the author, please write to: Department of Philosophy, 599 Lucas Hall, One University Boulevard, University of Missouri, St. Louis, MO 63121; e‐mail: [email protected]. (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)

Recent work on the heat-shock protein Hsp90 by Rutherford and Lindquist (1998) has been included among the pieces of evidence taken to show the essential role of developmental processes in evolution; Hsp90 acts as a buffer against phenotypic variation, allowing genotypic variation to build. When the buffering capacity of Hsp90 is altered (e.g., in nature, by mutation or environmental stress), the genetic variation is "revealed," manifesting itself as phenotypic variation. This phenomenon raises questions about the genetic variation before and after (...) what I will call a "revelation event": Is it neutral, nearly neutral, or non-neutral (i.e., strongly deleterious or strongly advantageous)? Moreover, what kinds of evolutionary processes do we take to be at work? Rutherford and Lindquist (1998) focus on the implications of non-neutral variation and selection. Later work by Queitsch, Sangster, and Lindquist (2002) and Sangster, Lindquist, and Queitsch (2004) raises the possibility that Hsp90 buffering may play the role that was played by drift in Sewall Wright's shifting balance model, permitting transition from one adaptive peak to another. However, Ohta (2002) suggests that much of this variation may be nearly neutral, which in turn, would imply a strong role for drift as well as selection. The primary goal of this paper is to illuminate the alternative scenarios and the processes operating in each. At the end, I raise the possibility of a synthesis between evo-devo and nearly neutral evolution. (shrink)

Recently, much philosophical discussion has centered on the best way to characterize the concepts of random drift and natural selection, and, in particular, whether selection and drift can be conceptually distinguished (Beatty, 1984; Brandon, 2005; Hodge, 1983, 1987; Millstein, 2002, 2005; Pfeifer, 2005; Shanahan, 1992; Stephens, 2004). These authors all contend, to a greater or lesser degree, that their concepts make sense of biological practice. So it should be instructive to see how the concepts of drift and selection were distinguished (...) by the disputants in a high-profile debate; debates such as these often force biologists to take a more philosophical turn, discussing the concepts at issue in greater detail than usual. Moreover, it is important to consider a debate where the disputants are actually trying to apply the models of population genetics to natural populations; only then can their proper interpretations become fully apparent. (Indeed, I contend that some of the philosophical confusion has arisen because authors have considered only the models themselves, and not the phenomena that the models are attempting to represent). A prime candidate for just such a case study is what Provine (1986) has termed “The Great Snail Debate,” that is, the debates over the highly polymorphic land snails Cepaea nemoralis and C. hortensis in the 1950s and early 1960s. These studies represent one of the best, if not the best, of the early attempts to demonstrate drift in natural populations. (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)

This paper outlines a critique of the use of the genetic variance–covariance matrix (G), one of the central concepts in the modern study of natural selection and evolution. Specifically, I argue that for both conceptual and empirical reasons, studies of G cannot be used to elucidate so-called constraints on natural selection, nor can they be employed to detect or to measure past selection in natural populations – contrary to what assumed by most practicing biologists. I suggest that the search for (...) a general solution to the difficult problem of identifying causal structures given observed correlation’s has led evolutionary quantitative geneticists to substitute statistical modeling for the more difficult, but much more valuable, job of teasing apart the many possible causes underlying the action of natural selection. Hence, the entire evolutionary quantitative genetics research program may be in need of a fundamental reconsideration of its goals and how they correspond to the array of mathematical and experimental techniques normally employed by its practitioners. (shrink)

Evolutionary theory (ET) is teeming with probabilities. Probabilities exist at all levels: the level of mutation, the level of microevolution, and the level of macroevolution. This uncontroversial claim raises a number of contentious issues. For example, is the evolutionary process (as opposed to the theory) indeterministic, or is it deterministic? Philosophers of biology have taken different sides on this issue. Millstein (1997) has argued that we are not currently able answer this question, and that even scientific realists ought to remain (...) agnostic concerning the determinism or indeterminism of evolutionary processes. If this argument is correct, it suggests that, whatever we take probabilities in ET to be, they must be consistent with either determinism or indeterminism. This raises some interesting philosophical questions: How should we understand the probabilities used in ET? In other words, what is meant by saying that a certain evolutionary change is more or less probable? Which interpretation of probability is the most appropriate for ET? I argue that the probabilities used in ET are objective in a realist sense, if not in an indeterministic sense. Furthermore, there are a number of interpretations of probability that are objective and would be consistent with ET under determinism or indeterminism. However, I argue that evolutionary probabilities are best understood as propensities of population-level kinds. (shrink)

The four case studies on chance in evolution provide a rich source for further philosophical analysis. Among the issues raised are the following: Are there different conceptions of chance at work, or is there a common underlying conception? How can a given concept of chance be distinguished from other chance concepts and from nonchance concepts? How can the occurrence of a given chance process be distinguished empirically from nonchance processes or other chance processes? What role does chance play in evolutionary (...) theory? I argue that in order to answer these questions, a careful distinction between process and outcome must be made; however, the purpose of this essay is not to answer these questions definitively, but rather to elaborate on them and to provide a starting point for further discussion. (shrink)

The probability that the fitter of two alleles will increase in frequency in a population goes up as the product of N (the effective population size) and s (the selection coefficient) increases. Discovering the distribution of values for this product across different alleles in different populations is a very important biological task. However, biologists often use the product Ns to define a different concept; they say that drift “dominates” selection or that drift is “stronger than” selection when Ns is much (...) smaller than some threshold quantity (e.g., ½) and that the reverse is true when Ns is much larger than that threshold. We argue that the question of whether drift dominates selection for a single allele in a single population makes no sense. Selection and drift are causes of evolution, but there is no fact of the matter as to which cause is stronger in the evolution of any given allele. (shrink)

Several recent arguments by philosophers of biology have challenged the traditional view that evolutionary factors, such as drift and selection, are genuine causes of evolutionary outcomes. In the case of drift, advocates of the statistical theory argue that drift is merely the sampling error inherent in the other stochastic processes of evolution and thus denotes a mathematical, rather than causal, feature of populations. This debate has largely centered around one particular model of drift, the Wright–Fisher model, and this has contributed (...) to the plausibility of the statisticalists’ arguments. However, an examination of alternative, predictively inequivalent models shows that drift is a genuine cause that can be manipulated to change population outcomes. This case study illustrates the influence of methodological assumptions on ontological judgments, particularly the pernicious effect of focusing on a particular model at the expense of others and confusing its assumptions and idealizations for true claims about the phenomena being modeled. (shrink)

This essay analyzes and develops recent views about explanation in biology. Philosophers of biology have parted with the received deductive-nomological model of scientific explanation primarily by attempting to capture actual biological theorizing and practice. This includes an endorsement of different kinds of explanation (e.g., mathematical and causal-mechanistic), a joint study of discovery and explanation, and an abandonment of models of theory reduction in favor of accounts of explanatory reduction. Of particular current interest are philosophical accounts of complex explanations that appeal (...) to different levels of organismal organization and use contributions from different biological disciplines. The essay lays out one model that views explanatory integration across different disciplines as being structured by scientific problems. I emphasize the philosophical need to take the explanatory aims pursued by different groups of scientists into account, as explanatory aims determine whether different explanations are competing or complementary and govern the dynamics of scientific practice, including interdisciplinary research. I distinguish different kinds of pluralism that philosophers have endorsed in the context of explanation in biology, and draw several implications for science education, especially the need to teach science as an interdisciplinary and dynamic practice guided by scientific problems and explanatory aims. (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)

Clatterbuck et al. (Biol Philos 28: 577–592, 2013) argue that there is no fact of the matter whether selection dominates drift or vice versa in any particular case of evolution. Their reasons are not empirically based; rather, they are purely conceptual. We show that their conceptual presuppositions are unmotivated, unnecessary and overly complex. We also show that their conclusion runs contrary to current biological practice. The solution is to recognize that evolution involves a probabilistic sampling process, and that drift is (...) a deviation from probabilistic expectation. We conclude that conceptually, there are no problems with distinguishing drift from selection, and empirically—as modern science illustrates—when drift does occur, there is a quantifiable fact of the matter to be discovered. (shrink)

ABSTRACTIn Darwinian Population and Natural Selection, Peter Godfrey-Smith brought the topic of natural selection back to the forefront of philosophy of biology, highlighting different issues surro...

The Darwinian theory of evolution is itself evolving and this book presents the details of the core of modern Darwinism and its latest developmental directions. The authors present current scientific work addressing theoretical problems and challenges in four sections, beginning with the concepts of evolution theory, its processes of variation, heredity, selection, adaptation and function, and its patterns of character, species, descent and life. The second part of this book scrutinizes Darwinism in the philosophy of science and its usefulness in (...) understanding ecosystems, whilst the third section deals with its application in disciplines beyond the biological sciences, including evolutionary psychology and evolutionary economics, Darwinian morality and phylolinguistics. The final section addresses anti-Darwinism, the creationist view and issues around teaching evolution in secondary schools. The reader learns how current experimental biology is opening important perspectives on the sources of variation, and thus of the very power of natural selection. This work examines numerous examples of the extension of the principle of natural selection and provides the opportunity to critically reflect on a rich theory, on the methodological rigour that presides in its extensions and exportations, and on the necessity to measure its advantages and also its limits. Scholars interested in modern Darwinism and scientific research, its concepts, research programs and controversies will find this book an excellent read, and those considering how Darwinism might evolve, how it can apply to the human sciences and other disciplines beyond its origins will find it particularly valuable. Originally produced in French (Les Mondes Darwiniens), the scope and usefulness of the book have led to the production of this English text, to reach a wider audience. This book is a milestone in the impressive penetration by Francophone scholars into the world of Darwinian science, its historiography and philosophy over the last two decades. Alex Rosenberg, R. Taylor Cole Professor of Philosophy, Duke University Until now this useful and comprehensive handbook has only been available to francophones. Thanks to this invaluable new translation, this collection of insightful and original essays can reach the global audience it deserves. Tim Lewens, University of Cambridge. (shrink)

Drift is often characterized in statistical terms. Yet such a purely statistical characterization is ambiguous for it can accept multiple physical interpretations. Because of this ambiguity it is important to distinguish what sorts of processes can lead to this statistical phenomenon. After presenting a physical interpretation of drift originating from the most popular interpretation of fitness, namely the propensity interpretation, I propose a different one starting from an analysis of the concept of drift made by Godfrey-Smith. Further on, I show (...) how my interpretation relates to previous attempts to make sense of the notion of expected value in deterministic setups. The upshot of my analysis is a physical conception of drift that is compatible with both a deterministic and indeterministic world. (shrink)

Two strategies for using a model as “null” are distinguished. Null modeling evaluates whether a process is causally responsible for a pattern by testing it against a null model. Baseline modeling measures the relative significance of various processes responsible for a pattern by detecting deviations from a baseline model. When these strategies are conflated, models are illegitimately privileged as accepted until rejected. I illustrate this using the neutral theory of ecology and draw general lessons from this case. First, scientists cannot (...) draw certain conclusions using null modeling. Second, these conclusions follow using baseline modeling, but doing so requires more evidence. (shrink)

There is a widespread philosophical interpretation of natural selection in evolutionary theory: natural selection, like mutation, migration, and drift are seen as forces that propel the evolution of populations. Natural selection is thus a population level causal process. This account has been challenged by the Statistics, claiming that natural selection is not a population level cause but rather a statistical feature of a population. This paper examines the nature of the aforementioned ontological debate and the nature of statistical explanations given (...) by population genetics. I claim that the Modern Synthesis provides good explanations of the changes in trait structure of populations without appealing to detailed causal information about the individual trajectories of the members of a population. (shrink)

Sober develops an account of adaptations on which they must have been positive causal factors for reproductive success. Glymour defends an account of a proper subset of adaptations—adaptations to particular environmental conditions—on which traits must interact in a special way with adapting conditions to cause reproductive success. These theories render conflicting judgments about which traits count as adaptations in some interesting cases. In this article I explore one such case and argue that we ought to replace the notion of adaptation (...) qua positive causal factor with a novel notion of adaptation. (shrink)

I define a concept of causal probability and apply it to questions about the role of probability in evolutionary processes. Causal probability is defined in terms of manipulation of patterns in empirical outcomes by manipulating properties that realize objective probabilities. The concept of causal probability allows us see how probabilities characterized by different interpretations of probability can share a similar causal character, and does so in such way as to allow new inferences about relationships between probabilities realized in different chance (...) setups. I clarify relations between probabilities and properties defined in terms of them, and argue that certain widespread uses of computer simulations in evolutionary biology show that many probabilities relevant to evolutionary outcomes are causal probabilities. This supports the claim that higher-level properties such as biological fitness and processes such as natural selection are causal properties and processes, contrary to what some authors have argued. (shrink)

Michael Scriven’s (1959) example of identical twins (who are said to be equal in fitness but unequal in their reproductive success) has been used by many philosophers of biology to discuss how fitness should be defined, how selection should be distinguished from drift, and how the environment in which a selection process occurs should be conceptualized. Here it is argued that evolutionary theory has no commitment, one way or the other, as to whether the twins are equally fit. This is (...) because the theory of natural selection is fundamentally about the fitnesses of traits, not the fitnesses of token individuals. A plausible philosophical thesis about supervenience entails that the twins are equally fit if they live in identical environments, but evolutionary biology is not committed to the thesis that the twins live in identical environments. Evolutionary theory is right to focus on traits, rather than on token individuals, because the fitnesses of token organisms (as opposed to their actual survivorship and degree of reproductive success) are almost always unknowable. This point has ramifications for the question of how Darwin’s theory of evolution and R. A. Fisher’s are conceptually different. (shrink)

Evolution in its contemporary meaning in biology typically refers to the changes in the proportions of biological types in a population over time (see the entry on the concept of evolution to 1872 for earlier meanings). As evolution is too large of a topic to address thoroughly in one entry, the primary goal of this entry is to serve as a broad overview of contemporary issues in evolution with links to other entries where more in-depth discussion can be found. The (...) entry begins with a brief survey of definitions of evolution, followed by a discussion of the different modes of evolution and related philosophical issues, and ends with a summary of other topics in the philosophy of evolution focusing particularly on topics covered in this encyclopedia. (shrink)

The 1981 Dahlem conference was a catalyst for contemporary evolutionary developmental biology (Evo-devo). This introductory chapter rehearses some of the details of the history surrounding the original conference and its associated edited volume, explicates the philosophical problem of conceptual change that provided the rationale for a workshop devoted to evaluating the epistemic revisions and transformations that occurred in the interim, explores conceptual change with respect to the concept of evolutionary novelty, and highlights some of the themes and patterns in the (...) different contributions to the present volume, Conceptual Change in Biology: Scientific and Philosophical Perspectives on Evolution and Development. (shrink)

One contentious debate in the philosophy of biology is that between the statisticalists and causalists. The former understand core evolutionary concepts like fitness and selection to be mere statistical summaries of underlying causal processes. In this view, evolutionary changes cannot be causally explained by selection or fitness. The causalist side, on the other hand, holds that populations can change in response to selection—one can cite fitness differences or driftability in causal explanations of evolutionary change. But, on the causalist side, it (...) is often not clear how, precisely, one should understand these causes. Thus, much more could be said about what sort of causes fitness and driftability are. In this paper, I borrow Dretske's distinction between structuring and triggering causes and I suggest that fitness and driftability are structuring causes of evolution. (shrink)

In this paper, I argue (contra some recent philosophical work) that an objective distinction between natural selection and drift can be drawn. I draw this distinction by conceiving of drift, in the most fundamental sense, as an individual-level phenomenon. This goes against some other attempts to distinguish selection from drift, which have argued either that drift is a population-level process or that it is a population-level product. Instead of identifying drift with population-level features, the account introduced here can explain these (...) population-level features based on a property that I label driftability. Additionally, this account shows that biology’s “first law”—the Principle of Drift (Brandon, J Phil 102(7):319–335 2006)—is not a foundational law, but is a consequence of driftability. (shrink)

These preprints were automatically compiled into a PDF from the collection of papers deposited in PhilSci-Archive in conjunction with the PSA 2012.

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)

Biologists and philosophers have been extremely pessimistic about the possibility of demonstrating random drift in nature, particularly when it comes to distinguishing random drift from natural selection. However, examination of a historical case-Maxime Lamotte's study of natural populations of the land snail, Cepaea nemoralis in the 1950s - shows that while some pessimism is warranted, it has been overstated. Indeed, by describing a unique signature for drift and showing that this signature obtained in the populations under study, Lamotte was able (...) to make a good case for a significant role for drift. It may be difficult to disentangle the causes of drift and selection acting in a population, but it is not impossible. (shrink)

This article elaborates on McShea and Brandon’s idea that drift is unlike the rest of the evolutionary factors because it is constitutive rather than imposed on the evolutionary process. I show that the way they spelled out this idea renders it inadequate and is the reason why it received some objections. I propose a different way in which their point could be understood, that rests on two general distinctions. The first is a distinction between the underlying mathematical apparatus used to (...) formulate a theory and a concept proposed by that theory. With the aid of a formal reconstruction of a population genetic model, I show that drift belongs to the first category. That is, that drift is constitutive of population genetics in the same sense that multiplication is constitutive in classical mechanics, or that circle is constitutive in Ptolemaic astronomy. The second distinction is between eliminating a concept from a theory and setting its value to zero. I will show that even though drift can be set to zero just like the rest of the evolutionary factors, eliminating drift is much harder than eliminating those other factors, since it would require changing the entire mathematical apparatus of standard population genetic theory. I conclude by drawing some other implications from the proposed formal reconstruction. (shrink)

Over the past fifteen years there has been a considerable amount of debate concerning what theoretical population dynamic models tell us about the nature of natural selection and drift. On the causal interpretation, these models describe the causes of population change. On the statistical interpretation, the models of population dynamics models specify statistical parameters that explain, predict, and quantify changes in population structure, without identifying the causes of those changes. Selection and drift are part of a statistical description of population (...) change; they are not discrete, apportionable causes. Our objective here is to provide a definitive statement of the statistical position, so as to allay some confusions in the current literature. We outline four commitments that are central to statisticalism. They are: 1. Natural Selection is a higher order effect; 2. Trait fitness is primitive; 3. Modern Synthesis (MS)-models are substrate neutral; 4. MS-selection and drift are model-relative. (shrink)

This article analyzes the view of evolutionary theory as a theory of forces. The analogy with Newtonian mechanics has been challenged due to the alleged mismatch between drift and the other evolutionary forces. Since genetic drift has no direction several authors tried to protect its status as a force: denying its lack of directionality, extending the notion of force and looking for a force in physics which also lacks of direction. I analyse these approaches, and although this strategy finally succeeds, (...) this discussion overlooks the crucial point on the debate between causalists and statisticalists: the causal status of evolutionary theory. (shrink)

The debate between the dynamical and the statistical interpretations of natural selection is centred on the question of whether all explanations that employ the concepts of natural selection and drift are reducible to causal explanations. The proponents of the statistical interpretation answer negatively, but insist on the fact that selection/drift arguments are explanatory. However, they remain unclear on where the explanatory power comes from. The proponents of the dynamical interpretation answer positively and try to reduce selection/drift arguments to some of (...) the most prominent accounts of causal explanation. In turn, they face the criticism raised by statisticalists that current accounts of causation have to be violated in some of their core conditions or otherwise used in a very loose manner in order to account for selection/drift explanations. We propose a reconciliation of both interpretations by conveying evolutionary explanations within the unificationist model of scientific explanation. Therefore, we argue that the explanatory power in natural selection arguments is a result of successful unification of individual- and population-level facts. A short case study based on research on sympatric speciation will be presented as an example of how population- and individual-level facts are unified to explain the morphological mosaic of bill shape in island scrub jays. (shrink)

One hotly debated philosophical question in the analysis of evolutionary theory concerns whether or not evolution and the various factors which constitute it may profitably be considered as analogous to “forces” in the traditional, Newtonian sense. Several compelling arguments assert that the force picture is incoherent, due to the peculiar nature of genetic drift. I consider two of those arguments here – that drift lacks a predictable direction, and that drift is constitutive of evolutionary systems – and show that they (...) both fail to demonstrate that a view of genetic drift as a force is untenable. I go on to diagnose the reasons for the stubborn persistence of this problem, considering two open philosophical issues and offering some preliminary arguments in support of the force metaphor. (shrink)

Discussions of “chance” and related concepts are found throughout philosophical work on evolutionary theory. By drawing attention to three very commonly-recognized distinctions, I separate four independent concepts falling under the broad heading of “chance”: randomness, epistemic unpredictability, causal indeterminism, and probabilistic causal processes. Far from a merely semantic distinction, however, it is demonstrated that conflation of these obviously distinct notions has an important bearing on debates at the core of evolutionary theory, particularly the debate over the interpretation of fitness, natural (...) selection, and genetic drift. (shrink)

Science is about conceptualizing the natural world in a way that can be understood by human beings while at the same time reflecting as much as possible what we can empirically infer about how the world actually is. Among the crucial tools that allow scientists to formulate hypotheses and to contribute to a progressive understanding of nature are the use of imagery and metaphors, on the one hand, and the ability to assume certain starting points on which to build new (...) avenues of inquiry on the other hand. The premise of this work is that, in the words of philosopher of science Daniel Dennett, "There is no such thing as philosophy-free science; there is only science whose philosophical baggage is taken on board without examination." The purpose here is precisely to examine some of this philosophical baggage, and to see where such analysis may lead us. Obviously, it is not possible for a single project to take on science as a whole, or even an entire discipline such as physics, or ecology. Therefore, the core of this work is constituted by a series of eight case studies drawn from the field of evolutionary biology, with which I am particularly familiar as a practicing biologist. The hope is that combining expertise in both philosophy and science in one person, the resulting insights might be useful for the practicing scientist as well as because of their value as philosophical inquires. The dissertation is a combination of conceptual analysis and science criticism applied to specific questions in organismal biology, largely of an evolutionary nature, with the eight case studies grouped into three broad categories: Unexamined Baggage, Bad Habits, and Good and bad metaphors. (shrink)

I argue that the propensity interpretation of fitness, properly understood, not only solves the explanatory circularity problem and the mismatch problem, but can also withstand the Pandora’s box full of problems that have been thrown at it. Fitness is the propensity (i.e., probabilistic ability, based on heritable physical traits) for organisms or types of organisms to survive and reproduce in particular environments and in particular populations for a specified number of generations; if greater than one generation, “reproduction” includes descendants of (...) descendants. Fitness values can be described in terms of distributions of propensities to produce varying number of offspring and can be modeled for any number of generations using computer simulations, thus providing both predictive power and a means for comparing the fitness of different phenotypes. Fitness is a causal concept, most notably at the population level, where fitness differences are causally responsible for differences in reproductive success. Relative fitness is ultimately what matters for natural selection. (shrink)