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)

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)

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)

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)

In this paper, we address the question whether a mechanistic approach can account for evolutionary causes. The last decade has seen a major attempt to account for natural selection as a mechanism. Nevertheless, we stress the relevance of broadening the debate by including the other evolutionary causes inside the mechanistic approach, in order to be a legitimate conceptual framework on the same footing as other approaches to evolutionary theory. We analyse the current debate on natural selection as a mechanism, and (...) extend it to the rest of the evolutionary causes. We focus on three approaches that we call the stochastic view, the functional view, and the minimalist view. We argue that all of them are unable to account for evolutionary causes as mechanisms. It is concluded that the current mechanistic proposals cannot be accepted as a common framework for evolutionary causes. Finally, we outline some guidelines and requirements that any mechanistic proposal should meet in order to be applied to evolutionary theory. (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 causal nature of evolution is one of the central topics in the philosophy of biology. The issue concerns whether equations used in evolutionary genetics point to some causal processes or purely phenomenological patterns. To address this question the present article builds well-defined causal models that underlie standard equations in evolutionary genetics. These models are based on minimal and biologically plausible hypotheses about selection and reproduction, and generate statistics to predict evolutionary changes. The causal reconstruction of the evolutionary principles shows (...) adaptive evolution as a genuine causal process, where fitness and selection are both causes of evolution. 1 Introduction2 The Philosophical Puzzle3 Causal Models4 Causal Foundations of Evolutionary Genetics4.1 Univariate quantitative genetics model4.1.1 The causal graph4.1.2 Structural equations4.1.3 Deriving evolutionary transition functions4.2 One-locus population genetics model5 Evolution as a Causal Process5.1 Selection as a causal process5.2 Causes of evolutionary change6 Conclusion. (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)

The causal nature of evolution is one of the central topics in the philosophy of biology. The issue concerns whether equations used in evolutionary genetics point to some causal processes or purely phenomenological patterns. To address this question the present article builds well-defined causal models that underlie standard equations in evolutionary genetics. These models are based on minimal and biologically plausible hypotheses about selection and reproduction, and generate statistics to predict evolutionary changes. The causal reconstruction of the evolutionary principles shows (...) adaptive evolution as a genuine causal process, where fitness and selection are both causes of evolution. 1 Introduction2 The Philosophical Puzzle3 Causal Models4 Causal Foundations of Evolutionary Genetics4.1 Univariate quantitative genetics model4.1.1 The causal graph4.1.2 Structural equations4.1.3 Deriving evolutionary transition functions4.2 One-locus population genetics model5 Evolution as a Causal Process5.1 Selection as a causal process5.2 Causes of evolutionary change6 Conclusion. (shrink)

Really statistical explanation is a hitherto neglected form of noncausal scientific explanation. Explanations in population biology that appeal to drift are RS explanations. An RS explanation supplies a kind of understanding that a causal explanation of the same result cannot supply. Roughly speaking, an RS explanation shows the result to be mere statistical fallout.

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)

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)

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)

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.

The concept of developmental constraint was at the heart of developmental approaches to evolution of the 1980s. While this idea was widely used to criticize neo-Darwinian evolutionary theory, critique does not yield an alternative framework that offers evolutionary explanations. In current Evo-devo the concept of constraint is of minor importance, whereas notions as evolvability are at the center of attention. The latter clearly defines an explanatory agenda for evolutionary research, so that one could view the historical shift from ‘developmental constraint’ (...) towards ‘evolvability’ as the move from a concept that is a mere tool of criticism to a concept that establishes a positive explanatory project. However, by taking a look at how the concept of constraint was employed in the 1980s, I argue that developmental constraint was not just seen as restricting possibilities (‘constraining’), but also as facilitating morphological change in several ways. Accounting for macroevolutionary transformation and the origin of novel form was an aim of these developmental approaches to evolution. Thus, the concept of developmental constraint was part of a positive explanatory agenda long before the advent of Evo-devo as a genuine scientific discipline. In the 1980s, despite the lack of a clear disciplinary identity, this concept coordinated research among paleontologists, morphologists, and developmentally inclined evolutionary biologists. I discuss the different functions that scientific concepts can have, highlighting that instead of classifying or explaining natural phenomena, concepts such as ‘developmental constraint’ and ‘evolvability’ are more important in setting explanatory agendas so as to provide intellectual coherence to scientific approaches. The essay concludes with a puzzle about how to conceptually distinguish evolvability and selection. (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)

Recently, a number of philosophers of biology have endorsed views about random drift that, we will argue, rest on an implicit assumption that the meaning of concepts such as drift can be understood through an examination of the mathematical models in which drift appears. They also seem to implicitly assume that ontological questions about the causality of terms appearing in the models can be gleaned from the models alone. We will question these general assumptions by showing how the same equation (...) — the simple 2 = p2 + 2pq + q2 — can be given radically different interpretations, one of which is a physical, causal process and one of which is not. This shows that mathematical models on their own yield neither interpretations nor ontological conclusions. Instead, we argue that these issues can only be resolved by considering the phenomena that the models were originally designed to represent and the phenomena to which the models are currently applied. When one does take those factors into account, starting with the motivation for Sewall Wright’s and R.A. Fisher’s early drift models and ending with contemporary applications, a very different picture of the concept of drift emerges. On this view, drift is a term for a set of physical processes, namely, indiscriminate sampling processes. (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)

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)

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 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)

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)

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)

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)

There is surprisingly little philosophical work on conceptually spelling out the difference between the traits on which natural selection may be said to act (e.g. “having a high running speed”) and mere circumstantial traits (e.g. “happening to be in the path of a forest fire”). I label this issue the “selectable traits problem” and, in this paper, I propose a solution for it. I first show that, contrary to our first intuition, simply equating selectable traits with heritable ones is not (...) an adequate solution. I then go on to argue that two recent philosophical solutions to this problem—due to Peter Godfrey-Smith and Pierrick Bourrat—are unconvincing because they cannot accommodate frequency-dependent selection. The way out of this difficulty is, I argue, to accept that extrinsic properties dependent on relations between intrinsic properties of the population members should also count as selectable traits. I then show that my proposal is legitimized by more than the simple accommodation of frequency-dependent selection. (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)

The notion of fitness is usually equated to reproductive success. However, this actualist approach presents some difficulties, mainly the explanatory circularity problem, which have lead philosophers of biology to offer alternative definitions in which fitness and reproductive success are distinguished. In this paper, we argue that none of these alternatives is satisfactory and, inspired by Mumford and Anjum’s dispositional theory of causation, we offer a definition of fitness as a causal dispositional property. We argue that, under this framework, the distinctiveness (...) that biologists usually attribute to fitness—namely, the fact that fitness is something different from both the physical traits of an organism and the number of offspring it leaves—can be explained, and the main problems associated with the concept of fitness can be solved. Firstly, we introduce Mumford and Anjum's dispositional theory of causation and present our definition of fitness as a causal disposition. We explain in detail each of the elements involved in our definition, namely: the relationship between fitness and the functional dispositions that compose it, the emergent character of fitness, and the context-sensitivity of fitness. Finally, we explain how fitness and realized fitness, as well as expected and realized fitness are distinguished in our approach to fitness as a causal disposition. (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)

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)

‘Statisticalists’ argue that the individual interactions of organisms taken together constitute natural selection. On this view, natural selection is an aggregated effect of interactions rather than some added cause acting on populations. The statisticalists’ view entails that natural selection and drift are indistinguishable aggregated effects of interactions, so that it becomes impossible to make a difference between them. The present paper attempts to make sense of the difference between selection and drift, given the main insights of statisticalism; basically, it will (...) disentangle the various kinds of indistinguishability between selection and drift that happen within biology, by examining the epistemological and metaphysical nature of the distinction between selection and drift. It will be based on a ‘difference-making account’ of selection. The first section will explicate the inscrutability of selection and drift, its various types in the statisticalist writings, and its implications. The second section specifies concepts of natural selection and drift in the difference making account of selection I am using, and shows that one can derive from this the statistical signatures of selection and drift. On this basis I focus on one sort of indistinguishability issue about selection and drift, which I call epistemic opacity, and explain why it mostly affects small populations. The last section explains why epistemic opacity does not raise an genuine epistemic problem for evolutionary biology. (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)

This chapter contains sections titled: Highlights of Past Literature Current Work Future Work.

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

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)

Population genetics attempts to measure the influence of the causes of evolution, viz., mutation, migration, natural selection, and random genetic drift, by understanding the way those causes change the genetics of populations. But how does it accomplish this goal? After a short introduction, we begin in section (2) with a brief historical outline of the origins of population genetics. In section (3), we sketch the model theoretic structure of population genetics, providing the flavor of the ways in which population genetics (...) theory might be understood as incorporating causes. In sections (4) and (5) we discuss two specific problems concerning the relationship between population genetics and evolutionary causes, viz., the problem of conceptually distinguishing natural selection from random genetic drift, and the problem of interpreting fitness. In section (6), we briefly discuss the methodology and key epistemological problems faced by population geneticists in uncovering the causes of evolution. Section (7) of the essay contains concluding remarks. (shrink)

Darwin's theory of evolution by natural selection provided the first, and only, causal-mechanistic account of the existence of adaptations in nature. As such, it provided the first, and only, scientific alternative to the “argument from design”. That alone would account for its philosophical significance. But the theory also raises other philosophical questions not encountered in the study of the theories of physics. Unfortunately the concept of natural selection is intimately intertwined with the other basic concepts of evolutionary theory—such as the (...) concepts of fitness and adaptation —that are themselves philosophically controversial. Fortunately we can make considerable headway in getting clear on natural selection without solving all of those outstanding problems. (shrink)

Proceedings of the Pittsburgh Workshop in History and Philosophy of Biology, Center for Philosophy of Science, University of Pittsburgh, March 23-24 2001 Session 4: Evolutionary Indeterminism.

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)

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)

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.

This thesis focuses on models of conceptual change in science and philosophy. In particular, I developed a new bootstrapping methodology for studying conceptual change, centered around the formalization of several popular models of conceptual change and the collective assessment of their improved formal versions via nine evaluative dimensions. Among the models of conceptual change treated in the thesis are Carnap’s explication, Lakatos’ concept-stretching, Toulmin’s conceptual populations, Waismann’s open texture, Mark Wilson’s patches and facades, Sneed’s structuralism, and Paul Thagard’s conceptual revolutions. (...) -/- In order to analyze and compare the conception of conceptual change provided by these different models, I rely on several historical reconstructions of episodes of scientific conceptual change. The historical episodes of scientific change that figure in this work include the emergence of the morphological concept of fish in biological taxonomies, the development of scientific conceptions of temperature, the Church-Turing thesis and related axiomatizations of effective calculability, the history of the concept of polyhedron in 17th and 18th century mathematics, Hamilton’s invention of the quaternions, the history of the pre-abstract group concepts in 18th and 19th century mathematics, the expansion of Newtonian mechanics to viscous fluids forces phenomena, and the chemical revolution. I will also present five different formal and informal improvements of four specific models of conceptual change. I will first present two different improvements of Carnapian explication, a formal and an informal one. My informal improvement of Carnapian explication will consist of a more fine-grained version of the procedure that adds an intermediate, third step to the two steps of Carnapian explication. I will show how this novel three-step version of explication is more suitable than its traditional two-step relative to handle complex cases of explications. My second, formal improvement of Carnapian explication will be a full explication of the concept of explication itself within the theory of conceptual spaces. By virtue of this formal improvement, the whole procedure of explication together with its application procedures and its pragmatic desiderata will be reconceptualized as a precise procedure involving topological and geometrical constraints inside the theory of conceptual spaces. My third improved model of conceptual change will consist of a formal explication of Darwinian models of conceptual change that will make vast use of Godfrey-Smith’s population-based Darwinism for targeting explicitly mathematical conceptual change. My fourth improvement will be dedicated instead to Wilson’s indeterminate model of conceptual change. I will show how Wilson’s very informal framework can be explicated within a modified version of the structuralist model-theoretic reconstructions of scientific theories. Finally, the fifth improved model of conceptual change will be a belief-revision-like logical framework that reconstructs Thagard’s model of conceptual revolution as specific revision and contraction operations that work on conceptual structures. -/- At the end of this work, a general conception of conceptual change in science and philosophy emerges, thanks to the combined action of the three layers of my methodology. This conception takes conceptual change to be a multi-faceted phenomenon centered around the dynamics of groups of concepts. According to this conception, concepts are best reconstructed as plastic and inter-subjective entities equipped with a non-trivial internal structure and subject to a certain degree of localized holism. Furthermore, conceptual dynamics can be judged from a weakly normative perspective, bound to be dependent on shared values and goals. Conceptual change is then best understood, according to this conception, as a ubiquitous phenomenon underlying all of our intellectual activities, from science to ordinary linguistic practices. As such, conceptual change does not pose any particular problem to value-laden notions of scientific progress, objectivity, and realism. At the same time, this conception prompts all our concept-driven intellectual activities, including philosophical and metaphilosophical reflections, to take into serious consideration the phenomenon of conceptual change. An important consequence of this conception, and of the analysis that generated it, is in fact that an adequate understanding of the dynamics of philosophical concepts is a prerequisite for analytic philosophy to develop a realistic and non-idealized depiction of itself and its activities. (shrink)

Contingency-theorists have gestured to a series of phenomena such as random mutations or rare Armageddon-like events as that which accounts for evolutionary contingency. These phenomena constitute a class, which may be aptly called the ‘sources of contingency’. In this paper, I offer a probabilistic conception of what it is to be a source of contingency and then examine two major candidates: chance variation and genetic drift, both of which have historically been taken to be ‘chancy’ in a number of different (...) senses. However,contrathe gesturing of contingency-theorists, chance variation and genetic drift are not always strong sources of contingency, as they can be non-chancy in at least one sense that opposes evolutionary contingency. The probabilistic conception offered herein allows for sources of contingency to appropriately vary in strength. To this end, I import Shannon’sinformation entropyas a statistical measure for systematically assessing the strength of a source of contingency, which is part and parcel of identifying sources of contingency. In brief, the higher the entropy, the greater the strength. This is also empirically significant because molecular, mutational, and replicative studies often contain sufficient frequency or probability data to allow for entropies to be calculated. In this way, contingency-theorists can evaluate the strength of a source of contingency in real-world cases. Moreover, the probabilistic conception also makes conceptual room for the converse of sources of contingency: ‘sources of directionality’, which ought to be recognised, as they can interact with genuine sources of contingency in undermining evolutionary contingency. (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)

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)