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)

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)

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

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 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)

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

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)

This paper takes a critical look at the idea that evolutionary theory is a statistical theory. It argues that despite the strong instrumental motivation for statistical theories, they are not necessary to explain deterministic systems. Biological evolution is fundamentally a result of deterministic processes. Hence, a statistical theory is not necessary for describing the evolutionary forces of genetic drift and natural selection, nor is it needed for describing the fitness of organisms. There is a computational advantage to the statistical theory (...) of population genetics, but population genetics succeeds only by eliminating causes from its account of evolutionary change. (shrink)

High-fidelity cultural transmission, rather than brute intelligence, is the secret of our species’ success, or so many cultural evolutionists claim. It has been selected because it ensures the spread, stability and longevity of beneficial cultural traditions, and it supports cumulative cultural change. To play these roles, however, fidelity must be a causally-efficient property of cultural transmission. This is where the grain problem comes in and challenges the explanatory potency of fidelity. Assessing the degree of fidelity of any episode or mechanism (...) of cultural transmission always depends upon an investigator’s choice of grain of description at which cultural traditions are being studied. The fidelity of cultural transmission then appears to be relative to the granularity at which one approaches cultural variation, and since there is a multiplicity of grains of description by which the same tradition can be studied, there results a multiplicity of measures of fidelity for a same event or mechanism of cultural transmission. If this is correct, because fidelity is always relative to the grain of description dictated by the local and specific research interests of the investigator, then there seems to be no fact of the matter as to whether cultural transmission is faithful or not, independently from a researcher’s framework of analysis. The aims of this paper are to offer a conceptual clarification of the grain problem in cultural evolution, to assess its causes, to unpack its epistemological implications, and to examine its reach and consequences for a science of cultural evolution. (shrink)

In this paper, I argue that what counts as the proper function of a trait is a matter of the de facto perspective that the biological system, itself, possesses on what counts as proper functioning for that trait. Unlike non-perspectival accounts, internal perspectivalism does not succumb to generality problems. But unlike external perspectivalism, internal perspectivalism can provide a fully naturalistic, mind-independent grounding of proper function and natural norms. The attribution of perspectives to biological systems is intended to be neither metaphorical (...) nor anthropomorphic: I do not mean to imply that such systems thereby must possess agency, cognition, intentions, concepts, or mental or psychological states. Instead, such systems provide the grounding for norms of performance when they internally enforce their own standard of (i.e., their own perspective on) what constitutes proper functioning or malfunctioning. By operating with a fixed, determinate level of generality, such systems provide the basis for an account of proper function that is immune to generality problems. (shrink)

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)

We argue that Brandon and Carson's (1996) "The Indeterministic Character of Evolutionary Theory" fails to identify any indeterminism that would require evolutionary theory to be a statistical or probabilistic theory. Specifically, we argue that (1) their demonstration of a mechanism by which quantum indeterminism might "percolate up" to the biological level is irrelevant; (2) their argument that natural selection is indeterministic because it is inextricably connected with drift fails to join the issue with determinism; and (3) their view that experimental (...) methodology in botany assumes indeterminism is both false and incompatible with the commitment to discoverable causal mechanisms underlying biological processes. We remain convinced that the probabilism of the theory of evolution is epistemically, not ontologically, motivated. (shrink)

This volume explores questions about conceptual change from both scientific and philosophical viewpoints by analyzing the recent history of evolutionary developmental biology. It features revised papers that originated from the workshop "Conceptual Change in Biological Science: Evolutionary Developmental Biology, 1981-2011" held at the Max Planck Institute for the History of Science in Berlin in July 2010. The Preface has been written by Ron Amundson. In these papers, philosophers and biologists compare and contrast key concepts in evolutionary developmental biology and their (...) development since the original, seminal Dahlem conference on evolution and development held in Berlin in 1981. Many of the original scientific participants from the 1981 conference are also contributors to this new volume and, in conjunction with other expert biologists and philosophers specializing on these topics, provide an authoritative, comprehensive view on the subject. Taken together, the papers supply novel perspectives on how and why the conceptual landscape has shifted and stabilized in particular ways, yielding insights into the dynamic epistemic changes that have occurred over the past three decades. This volume will appeal to philosophers of biology studying conceptual change, evolutionary developmental biologists focused on comprehending the genesis of their field and evaluating its future directions, and historians of biology examining this period when the intersection of evolution and development rose again to prominence in biological science. (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)

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 work consists of two parts. Part I will be a contribution to a philo- sophical discussion of the nature of causal explanation. It will present my contrastive counterfactual theory of causal explanation and show how it can be used to deal with a number of problems facing theories of causal explanation. Part II is a contribution to a discussion of the na- ture of interest explanation in social studies of science. The aim is to help to resolve some controversies (...) concerning interest explanation by explicating the concept of interest and its explanatory uses by using the account of explanation developed in Part I. (shrink)

When philosophers of physics explore the nature of chance, they usually look to quantum mechanics. When philosophers of biology explore the nature of chance, they usually look to microevolutionary phenomena, such as mutation or random drift. What has been largely overlooked is the role of chance in macroevolution. The stochastic models of paleobiology employ conceptions of chance that are similar to those at the microevolutionary level, yet different from the conceptions of chance often associated with quantum mechanics and Laplacean determinism.

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)

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)

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)

I argue that results from foraging theory give us good reason to think some evolutionary phenomena are indeterministic and hence that evolutionary theory must be probabilistic. Foraging theory implies that random search is sometimes selectively advantageous, and experimental work suggests that it is employed by a variety of organisms. There are reasons to think such search will sometimes be genuinely indeterministic. If it is, then individual reproductive success will also be indeterministic, and so too will frequency change in populations of (...) organisms employing such search. (shrink)

The traditional view of evolutionary theory asserts that we can usefully understand natural selection, drift, mutation, migration, and the system of mating as forces that cause evolutionary change. Recently, Denis Walsh and Robert Brandon have objected to this view. Walsh argues that the traditional view faces a fatal dilemma and that the force analogy must be rejected altogether. Brandon accepts the force analogy but argues that drift, rather than the Hardy-Weinberg law, is the best candidate for a zero-force law. Here (...) I defend the traditional view against these objections. (shrink)

Evolutionary biology, indeed any science that attempts to reconstruct prehistory, faces practical limitations on available data. These limitations create the problem of contrast failure: specific observations may fail to discriminate between rival evolutionary hypotheses. Assessing the risk of contrast failure provides a way to evaluate testing protocols in evolutionary science. Here I will argue that part of the methodological critique in the Spandrels paper involves diagnosing contrast failure problems. I then distinguish the problem of contrast failure from the more familiar (...) philosophical problem of underdetermination, and demonstrate how contrast failure arises in scientific practice with an investigation into Lewontin and White’s estimation of an adaptive landscape. (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)

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)

I respond to Brandon's (2005) criticisms of my earlier (2002) essay. I argue that (1) biologists are inconsistent in their use of the terms 'selection' and 'drift' -- vacillating between 'process' and 'outcome' -- but that the process-oriented definitions I defend make better sense of the neutralist/selectionist debate; (2) Brandon's purported demonstration that there is no qualitative difference between drift and selection as processes begs the question against my account; and (3) biologists (e.g., Kimura) have argued for genuinely neutral variants. (...) Whether any such variants actually exist is an empirical question. However, the philosophical question at hand is conceptual, not empirical. (shrink)

Evo-Devo exhibits a plurality of scientific “cultures” of practice and theory. When are the cultures acting—individually or collectively—in ways that actually move research forward, empirically, theoretically, and ethically? When do they become imperialistic, in the sense of excluding and subordinating other cultures? This chapter identifies six cultures – three /styles/ (mathematical modeling, mechanism, and history) and three /paradigms/ (adaptationism, structuralism, and cladism). The key assumptions standing behind, under, or within each of these cultures are explored. Characterizing the internal structure of (...) the cultures is necessary for understanding how they collaborate or compete, and how they are fragmented or integrated, in the rich interdisciplinary /trading zone/ (Galison 1997) of Evo-Devo. Evo-Devo is an important example of how science can progress through a radical plurality of perspectives and cultures. (shrink)

One central tenet of the Modern Evolutionary Synthesis , and the consensus view among biologists until now, is that all genetic mutations occur by “chance” or at “random” with respect to adaptation. However, the discovery of some molecular mechanisms enhancing mutation rate in response to environmental conditions has given rise to discussions among biologists, historians and philosophers of biology about the “chance” vs “directed” character of mutations . In fact, some argue that mutations due to a particular kind of mutator (...) mechanisms challenge the Modern Synthesis because they are produced when and where needed by the organisms concerned. This paper provides a defense of the Modern Synthesis’ consensus view about the chance nature of all genetic mutations by reacting to Jablonka and Lamb’s analysis of genetic mutations and the explicit Lamarckian flavor of their arguments. I argue that biologists can continue to talk about chance mutations according to what I call and define as the notion of “evolutionary chance,” which I claim is the Modern Synthesis’ consensus view and a reformulation of Darwin’s most influential idea of “chance” variation. Advances in molecular genetics are therefore significant but not revolutionary with respect to the Modern Synthesis’ paradigm. (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)

In this essay, I examine the usage of the term “just-so story.” I attempt to show that just-so storytelling can be seen as an epistemic concept that, in various ways, tackles the epistemological and methodological problems relating to evolutionary explanations qua historical/narrative explanations. I identify two main, yet mutually exclusive, strategies of employing the concept of a just-so story: a negative strategy and a positive strategy. Subsequently, I argue that these strategies do not satisfactorily capture the core of the “original” (...) meaning advanced by Stephen Jay Gould and Richard Lewontin at the end of the 1970s. I revisit the foundation(s) of their anti-adaptationist critique in order to reframe it as a critique of distinctive methodological manners and epistemic maxims related to historical inquiry. Last but not least, I suggest that contemporary evolutionary thinkers have two conceptually different options: they can either adhere to the “original” meaning of the term “just-so story” or accept that “just-so story” is a term equivalent to “implausible narrative explanation.”. (shrink)

Recent debate on the nature of probabilities in evolutionary biology has focused largely on the propensity interpretation of fitness (PIF), which defines fitness in terms of a conception of probability known as “propensity”. However, proponents of this conception of fitness have misconceived the role of probability in the constitution of fitness. First, discussions of probability and fitness have almost always focused on organism effect probability, the probability that an organism and its environment cause effects. I argue that much of the (...) probability relevant to fitness must be organism circumstance probability, the probability that an organism encounters particular, detailed circumstances within an environment, circumstances which are not the organism’s effects. Second, I argue in favor of the view that organism effect propensities either don’t exist or are not part of the basis of fitness, because they usually have values close to 0 or 1. More generally, I try to show that it is possible to develop a clearer conception of the role of probability in biological processes than earlier discussions have allowed. (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)

Recent discussion of the statistical character of evolutionary theory has centered around two positions: (1) Determinism combined with the claim that the statistical character is eliminable, a subjective interpretation of probability, and instrumentalism; (2) Indeterminism combined with the claim that the statistical character is ineliminable, a propensity interpretation of probability, and realism. I point out some internal problems in these positions and show that the relationship between determinism, eliminability, realism, and the interpretation of probability is more complex than previously assumed (...) in this debate. Furthermore, I take some initial steps towards a more adequate account of the statistical character of evolutionary theory. (shrink)

Alexander Rosenberg (1994) claims that the omniscient viewpoint of the evolutionary process would have no need for the concept of random drift. However, his argument fails to take into account all of the processes which are considered to be instances of random drift. A consideration of these processes shows that random drift is not eliminable even given a position of omniscience. Furthermore, Rosenberg must take these processes into account in order to support his claims that evolution is deterministic and that (...) evolutionary biology is an instrumental science. (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)

Recent discussion of the statistical character of evolutionary theory has centered around two positions: Determinism combined with the claim that the statistical character is eliminable, a subjective interpretation of probability, and instrumentalism; Indeterminism combined with the claim that the statistical character is ineliminable, a propensity interpretation of probability, and realism. I point out some internal problems in these positions and show that the relationship between determinism, eliminability, realism, and the interpretation of probability is more complex than previously assumed in this (...) debate. Furthermore, I take some initial steps towards a more adequate account of the statistical character of evolutionary theory. (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)

In the recent philosophical literature, two questions have arisen concerning the status of natural selection: (1) Is it a population-level phenomenon, or is it an organism-level phenomenon? (2) Is it a causal process, or is it a purely statistical summary of lower-level processes? In an earlier work (Millstein, Br J Philos Sci, 57(4):627–653, 2006), I argue that natural selection should be understood as a population-level causal process, rather than a purely statistical population-level summation of lower-level processes or as an organism-level (...) causal process. In a 2009 essay entitled “Productivity, relevance, and natural selection,” Stuart Glennan argues in reply that natural selection is produced by causal pro- cesses operating at the level of individual organisms, but he maintains that there is no causal productivity at the population level. However, there are, he claims, many population-level properties that are causally relevant to the dynamics of evolution- ary processes. Glennan’s claims rely on a causal pluralism that holds that there are two types of causes: causal production and causal relevance. Without calling into question Glennan’s causal pluralism or his claims concerning the causal relevance of natural selection, I argue that natural selection does in fact exhibit causal production at the population level. It is true that natural selection does not fit with accounts of mechanisms that involve decomposition of wholes into parts, such as Glennan’s own. However, it does fit with causal production accounts that do not require decomposition, such as Salmon’s Mark Transmission account, given the extent to which populations act as interacting “objects” in the process of natural selection. (shrink)

This essay outlines one aspect of a larger collaboration with John Beatty and Alan Love.2 The project’s focus is philosophical, but for reasons that will become clear momentarily, the method of approach is historical. All three of us share the conviction that philosophical issues concerning the foundations of the sciences are often illuminated by investigating their history. It is my hope that this paper both provides support for that thesis, and illustrates it. The focal philosophical issue can be stated in (...) the form of a puzzle about contemporary discussions of the conceptual foundations of evolutionary theory. The role of the concepts of “chance” and “randomness” in that theory is a significant part of .. (shrink)

Do evolutionary processes such as selection and random drift cause evolutionary change, or are they merely convenient ways of describing or summarizing it? Philosophers have lined up on both sides of this question. One recent defense (Reisman and Forber 2005) of the causal status of selection and drift appeals to a manipulability theory of causation. Yet, even if one accepts manipulability, there are still reasons to doubt that genetic drift, in particular, is genuinely causal. We will address two challenges to (...) treating drift as causal within a manipulation framework. We will argue that both challenges ultimately fail, but that they raise interesting and subtle issues about the nature of causation and the differences between selection and drift. ‡Thanks to audiences at the ‘PBDB1' Conference and the 2006 PSA Meeting for valuable feedback. †To contact the authors, please write to: Patrick Forber, Tufts University, Philosophy Department, Miner Hall, Medford, MA 02155; e-mail: [email protected]. (shrink)

In this paper I critically evaluate Reisman and Forber’s :1113–1123, 2005) arguments that drift and natural selection are population-level causes of evolution based on what they call the manipulation condition. Although I agree that this condition is an important step for identifying causes for evolutionary change, it is insufficient. Following Woodward, I argue that the invariance of a relationship is another crucial parameter to take into consideration for causal explanations. Starting from Reisman and Forber’s example on drift and after having (...) briefly presented the criterion of invariance, I show that once both the manipulation condition and the criterion of invariance are taken into account, drift, in this example, should better be understood as an individual-level rather than a population-level cause. Later, I concede that it is legitimate to interpret natural selection and drift as population-level causes when they rely on genuinely indeterministic events and some cases of frequency-dependent selection. (shrink)

Niche construction is the process of organisms changing themselves or their environment—or their relationship with their environment—in ways that affect the evolutionary trajectory of their population. These evolutionary trajectory changes are traditionally understood to be triggered by changes in selection pressures. Niche construction thus necessarily involves organisms altering selection pressures. In this paper, we argue that changes in selection pressures is not the only way organisms can influence the evolutionary futures of their population. We propose that organisms can also affect (...) drift probabilities, and that such changes should be considered niche construction. Drift probabilities can be modulated by altering population size or by affecting driftability (individual variance in possible reproductive outcomes). We consider both and provide examples of how niche construction can stabilize, increase, or dampen drift probabilities. Finally, we revisit and broaden the traditional definition of niche construction. We hold that organismic activities that modify drift probabilities should count as niche construction, even if selection pressures remain unaltered. (shrink)

Susan Mills and John Beatty proposed a propensity interpretation of fitness (1979) to show that Darwinian explanations are not circular, but they did not address the critics' chief complaint that the principle of the survival of the fittest is either tautological or untestable. I show that the propensity interpretation cannot rescue the principle from the critics' charges. The critics, however, incorrectly assume that there is nothing more to Darwin's theory than the survival of the fittest. While Darwinians all scoff at (...) this assumption, they do not agree about what role, if any, this principle plays in Darwin's theory of natural selection. I argue that the principle has no place in Darwin's theory. His theory does include the idea that some organisms are fitter than others. But greater reproductive success is simply inferred from higher fitness. There is no reason to embody this inference in the form of a special principle of the survival of the fittest. (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)

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)

The adequacy of Elliott Sober’s analogy between classical mechanics and evolutionary theory—according to which both theories explain via a zero-force law and a set of forces that alter the zero-force state—has been criticized from various points of view. I focus here on McShea and Brandon’s claim that drift shouldn’t be considered a force because it is not directional. I argue that there are a number of different theses that could be meant by this, and show that one of those theses—the (...) idea that drift cannot bias populations to be taken somewhere in the evolutionary space from one generation to the next—is actually false. Not only has this thesis been implicitly assumed in the discussion of the force analogy thus far, but it is also commonly found in a wider range of philosophical and biological texts. I argue that correcting this view, and the usual images associated with it, will thereby bring heuristic benefits that impact the force analogy discussion, but that also go beyond it. (shrink)

Work throughout the history and philosophy of biology frequently employs ‘chance’, ‘unpredictability’, ‘probability’, and many similar terms. One common way of understanding how these concepts were introduced in evolution focuses on two central issues: the first use of statistical methods in evolution (Galton), and the first use of the concept of “objective chance” in evolution (Wright). I argue that while this approach has merit, it fails to fully capture interesting philosophical reflections on the role of chance expounded by two of (...) Galton's students, Karl Pearson and W.F.R. Weldon. Considering a question more familiar from contemporary philosophy of biology—the relationship between our statistical theories of evolution and the processes in the world those theories describe—is, I claim, a more fruitful way to approach both these two historical actors and the broader development of chance in evolution. (shrink)