I examine recent debates in the philosophy of biology over the determinism or indeterminism of the evolutionary process, focusing on two papers in particular: Glymour 2001 and Stamos 2001. I argue that neither of these papers succeeds in making the case for the indeterminism of the evolutionary process, and suggest that what is needed is a detailed analysis of the causal processes at every level from the quantum mechanical to the evolutionary.

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

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

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

In this paper I argue that we can best make sense of the practice of experimental evolutionary biology if we see it as investigating contingent, rather than lawlike, regularities. This understanding is contrasted with the experimental practice of certain areas of physics. However, this presents a problem for those who accept the Logical Positivist conception of law and its essential role in scientific explanation. I address this problem by arguing that the contingent regularities of evolutionary biology have a limited range (...) of nomic necessity and a limited range of explanatory power even though they lack the unlimited projectibility that has been seen by some as a hallmark of scientific laws. (shrink)

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

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

This volume focuses on various questions concerning the interpretation of probability and probabilistic reasoning in biology and physics. It is inspired by the idea that philosophers of biology and philosophers of physics who work on the foundations of their disciplines encounter similar questions and problems concerning the role and application of probability, and that interaction between the two communities will be both interesting and fruitful. In this introduction we present the background to the main questions that the volume focuses on (...) and summarize the highlights of the individual contributions. (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)

It’s recently been argued that biological fitness can’t change over the course of an organism’s life as a result of organisms’ behaviors. However, some characterizations of biological function and biological altruism tacitly or explicitly assume that an effect of a trait can change an organism’s fitness. In the first part of the paper, I explain that the core idea of changing fitness can be understood in terms of conditional probabilities defined over sequences of events in an organism’s life. The result (...) is a notion of “conditional fitness” which is static but which captures intuitions about apparent behavioral effects on fitness. The second part of the paper investigates the possibility of providing a systematic foundation for conditional fitness in terms of spaces of sequences of states of an organism and its environment. I argue that the resulting “organism–environment history conception” helps unify diverse biological perspectives, and may provide part of a metaphysics of natural selection. (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)

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)

Since the last decades of the twentieth and the beginning of the twenty-first century, the use of metaphysics by philosophers when approaching conceptual problems in biology has increased. Some philosophers call this tendency in philosophy of biology ‘Metaphysics of Biology’. In this paper, I aim at characterizing Metaphysics of Biology by paying attention to the diverse ways philosophers use metaphysics when addressing conceptual problems in biology. I will claim that there are two different modes of doing Metaphysics of Biology, namely (...) Metaphysics for Biology and Metaphysics in Biology. (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)

In "The Indeterministic Character of Evolutionary Theory: No 'Hidden Variables Proof' But No Room for Determinism Either," Brandon and Carson (1996) argue that evolutionary theory is statistical because the processes it describes are fundamentally statistical. In "Is Indeterminism the Source of the Statistical Character of Evolutionary Theory?" Graves, Horan, and Rosenberg (1999) argue in reply that the processes of evolutionary biology are fundamentally deterministic and that the statistical character of evolutionary theory is explained by epistemological rather than ontological considerations. In (...) this paper I focus on the topic of mutation. By focusing on some of the theory and research on this topic from early to late, I show how quantum indeterminism hooks up to point mutations (via tautomeric shifts, proton tunneling, and aqueous thermal motion). I conclude with a few thoughts on some of the wider implications of this topic. (shrink)

This paper explores whether natural selection, a putative evolutionary mechanism, and a main one at that, can be characterized on either of the two dominant conceptions of mechanism, due to Glennan and the team of Machamer, Darden, and Craver, that constitute the “new mechanistic philosophy.” The results of the analysis are that neither of the dominant conceptions of mechanism adequately captures natural selection. Nevertheless, the new mechanistic philosophy possesses the resources for an understanding of natural selection under the rubric.

We review Evolution and the Levels of Selection by Samir Okasha. This important book provides a cohesive philosophical framework for understanding levels-of-selections problems in biology. Concerning evolutionary transitions, Okasha proposes that three stages characterize the shift from a lower level of selection to a higher one. We discuss the application of Okasha’s three-stage concept to the evolutionary transition from unicellularity to multicellularity in the volvocine green algae. Okasha’s concepts are a provocative step towards a more general understanding of the major (...) evolutionary transitions; however, the application of certain ideas to the volvocine model system is not straightforward. (shrink)

Evolutionary models can explain the dynamics of populations, how genetic, genotypic, or phenotypic frequencies change with time. Models incorporating chance, or drift, predict specific patterns of change. These are illustrated using classic work on blood types by Cavalli-Sforza and his collaborators in the Parma Valley of Italy, in which the theoretically predicted patterns are exhibited in human populations. These data and the models display properties of ensembles of populations. The explanatory problem needs to be understood in terms of how likely (...) an observed change, in either a population or an ensemble, would be under drift alone; this is fundamentally a matter of chance. Understood in this way, issues of drift and chance undercut most recent philosophical, but not biological, discussions of the role of "genetic drift.". (shrink)

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

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

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)

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)

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)

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)

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)

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.

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)

The proper function of the heart is pumping the blood. According to what we call the type etiological view, this is because previous tokens of the type HEART were selected for pumping the blood. Nanay :412–431, 2010) argues that the type etiological view is viciously circular. He claims that the only plausible accounts of trait type individuation use proper functions, such that whenever the type etiological view is supplemented with a plausible account of trait type individuation, the result is a (...) view that uses proper functions to explain proper functions. We refine this objection, and argue that Nanay at most establishes a potentially benign definitional circularity. However, we show that the type etiological view’s reliance on types nevertheless generates a vicious regress. Hence the type etiological view is false. We reject dispositional and modal alternatives to the type etiological view for the reason that they either cannot accommodate malfunction, or do so at the cost of proliferation; and then formulate a novel token etiological view that overcomes both problems because it makes no reference to trait types. (shrink)

Academia is subdivided into separate disciplines, most of which are quite discrete. In this review I trace the interactions between two of these disciplines: biology and philosophy of biology. I concentrate on those topics that have the most extensive biological content: function, species, systematics, selection, reduction and development. In the final section of this paper I touch briefly on those issues that biologists and philosophers have addressed that do not have much in the way of biological content.

In this article, I propose a new way of thinking about natural necessity and a new way of thinking about biological laws. I suggest that much of the lack of progress in making a positive case for distinctively biological laws is that we’ve been looking for necessity in the wrong place. The trend has been to look for exceptionlessness at the level of the outcomes of biological processes and to build one’s claims about necessity off of that. However, as Beatty (...) (1995) observed, even when we are lucky enough to find a biological ‘rule’ of some sort, that rule is apt to be a victim of ‘the rule-breaking capabilities of evolutionary change’. If indeed no distinctively biological generalization—even an exceptionless one—is safe, we need to locate necessity elsewhere. A good place to start is, I think, precisely the point at which Beatty sees the possibility of lawhood as breaking down—namely, at the level of chances. 1 The ‘Necessity’ Objection to Biological Laws2 Necessary Chances2.1 Necessary chances: random drift2.2 Necessary chances: fitness3 But is it Biological?4 Conclusion. (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 work is a critical consideration of several arguments recently given by Elliott Sober that are aimed at undermining the Laplacean stance on probability in evolutionary theory. The Laplacean contends that the only objective probability an event has is the one assigned to it by a complete description of the relevant microparticles. Sober alleges a formal demonstration that the Laplacean stance on probability in evolutionary theory is inconsistent. But Sober’s argument contains a crucial lacuna, one that likely cannot be repaired (...) to yield the conclusion that Sober draws. He also argues that the Laplacean is committed to inferring semantic facts about probabilities from pragmatic facts about agents’ reasons for using probabilities. But Sober’s arguments against inferring semantic facts from pragmatic constraints would only undermine one basis for Laplaceanism. The Laplacean who formulates her position carefully need not leave herself vulnerable to Sober’s objection to inferring semantics from pragmatics. (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)

Does the brain behave indeterministically? I argue that accounting for ion channels, key functional units in the brain, requires indeterministic models. These models are probabilistic, so the brain does behave indeterministically in a weak sense. I explore the implications of this point for a stronger sense of indeterminism. Ultimately I argue that it is not possible, either empirically or through philosophical argument, to show that the brain is indeterministic in that stronger sense.

The paper provides a new critical perspective on the propensity interpretation of fitness, by investigating its relationship to the propensity interpretation of probability. Two main conclusions are drawn. First, the claim that fitness is a propensity cannot be understood properly: fitness is not a propensity in the sense prescribed by the propensity interpretation of probability. Second, this interpretation of probability is inessential for explanations proposed by the PIF in evolutionary biology. Consequently, interpreting the probabilistic dimension of fitness in terms of (...) propensities is neither a strong motivation in favor of this interpretation, nor a possible target for substantial criticism. (shrink)

In what follows, I suggest that it makes good sense to think of the truth of the probabilistic generalizations made in the life sciences as metaphysically grounded in stochastic mechanisms in the world. To further understand these stochastic mechanisms, I take the general characterization of mechanism offered by MDC :1–25, 2000) and explore how it fits with several of the going philosophical accounts of chance: subjectivism, frequentism, Lewisian best-systems, and propensity. I argue that neither subjectivism, frequentism, nor a best-system-style interpretation (...) of chance will give us what we need from an account of stochastic mechanism, but some version of propensity theory can. I then draw a few important lessons from recent propensity interpretations of fitness in order to present a novel propensity interpretation of stochastic mechanism according to which stochastic mechanisms are thought to have probabilistic propensities to produce certain outcomes over others. This understanding of stochastic mechanism, once fully fleshed-out, provides the benefits of allowing the stochasticity of a particular mechanism to be an objective property in the world, a property investigable by science, a way of quantifying the stochasticity of a particular mechanism, and a way to avoid a problematic commitment to the causal efficacy of propensities. (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)

Recently, philosophers of biology have debated the status of the evolutionary process: is it deterministic or indeterministic? I argue that there is insufficient reason to favor one side of the debate over the other, and that a more philosophically defensible position argues neither for the determinacy nor for the indeterminacy of the evolutionary process. In other words, I maintain that the appropriate stand to take towards the question of the determinism of the evolutionary process is agnosticism. I then suggest that (...) an examination of the phenomenon of developmental noise might yield a solution to the problem. (shrink)

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