Are they needed? To be sure. The Darwinian industry, industrious though it is, has failed to provide texts of more than a handful of Darwin's books. If you want to know what Darwin said about barnacles (still an essential reference to cirripedists, apart from any historical importance) you are forced to search shelves, or wait while someone does it for you; some have been in print for a century; various reprints have appeared and since vanished." -Eric Korn,Times Literary Supplement Charles (...) Robert Darwin (1880-1882) has been widely recognized since his own time as one of the most influential writers in the history of Western thought. His books were widely read by specialists and the general public, and his influence had been extended by almost continuous public debate over the last 130 years. New York University Press' edition makes it possible for the first time to review Darwin's public literary output as a whole, plus his scientific journal articles, his private notebooks, and his correspondence. This is the first complete edition containing all of Darwin's published books, featuring definitive texts recording original paginations with Darwin's indexes retained. All illustrations and plates are presented, inclucing 82 color plates of birds and mammals and several folding maps and plates. The set also features a general introduction and index, and textural introductions in each volume. (shrink)

Stephen Jay Gould argued that replaying the ‘tape of life’ would result in radically different evolutionary outcomes. Recently, biologists and philosophers of science have paid increasing attention to the theoretical importance of convergent evolution—the independent origination of similar biological forms and functions—which many interpret as evidence against Gould’s thesis. In this paper, we examine the evidentiary relevance of convergent evolution for the radical contingency debate. We show that under the right conditions, episodes of convergent evolution can constitute valid natural experiments (...) that support inferences regarding the deep counterfactual stability of macroevolutionary outcomes. However, we argue that proponents of convergence have problematically lumped causally heterogeneous phenomena into a single evidentiary basket, in effect treating all convergent events as if they are of equivalent theoretical import. As a result, the ‘critique from convergent evolution’ fails to engage with key claims of the radical contingency thesis. To remedy this, we develop ways to break down the heterogeneous set of convergent events based on the nature of the generalizations they support. Adopting this more nuanced approach to convergent evolution allows us to differentiate iterated evolutionary outcomes that are probably common among alternative evolutionary histories and subject to law-like generalizations, from those that do little to undermine and may even support, the Gouldian view of life. (shrink)

Familiarity with Charles Darwin's treatise on evolution is essential to every well-educated individual. One of the most important books ever published--and a continuing source of controversy, a century and a half later--this classic of science is reproduced in a facsimile of the critically acclaimed first edition.

In The Origin of Species (1859) Darwin challenged many of the most deeply-held beliefs of the Western world. Arguing for a material, not divine, origin of species, he showed that new species are achieved by "natural selection." The Origin communicates the enthusiasm of original thinking in an open, descriptive style, and Darwin's emphasis on the value of diversity speaks more strongly now than ever. As well as a stimulating introduction and detailed notes, this edition offers a register of the many (...) writers referred to by Darwin in the text. (shrink)

Drift is to evolution as inertia is to Newtonian mechanics. Both are the "natural" or default states of the systems to which they apply. Both are governed by zero-force laws. The zero-force law in biology is stated here for the first time.

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)

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

This is the second of a two-part essay on the history of debates concerning the creativity of natural selection, from Darwin through the evolutionary synthesis and up to the present. In the first part, I focussed on the mid-late nineteenth century to the early twentieth, with special emphasis on early Darwinism and its critics, the self-styled “mutationists.” The second part focuses on the evolutionary synthesis and some of its critics, especially the “neutralists” and “neo-mutationists.” Like Stephen Gould, I consider the (...) creativity of natural selection to be a key component of what has traditionally counted as “Darwinism.” I argue that the creativity of natural selection is best understood in terms of selection initiating evolutionary change, and selection directing evolutionary change, for example by creating the variation that it subsequently acts upon. I consider the respects in which both of these claims sound non-Darwinian, even though they have long been understood by supporters and critics alike to be virtually constitutive of Darwinism. (shrink)

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

The modern synthesis in evolutionary biology is taken to be that period in which a consensus developed among biologists about the major causes of evolution, a consensus that informed research in evolutionary biology for at least a half century. As such, it is a particularly fruitful period to consider when reflecting on the meaning and role of chance in evolutionary explanation. Biologists of this period make reference to “chance” and loose cognates of “chance,” such as: “random,” “contingent,” “accidental,” “haphazard,” or (...) “stochastic.” Of course, what an author might mean by “chance” in any specific context varies. -/- In the following, we first offer a historiographical note on the synthesis. Second, we introduce five ways in which synthesis authors spoke about chance. We do not take these to be an exhaustive taxonomy of all possible ways in which chance meaningfully figures in explanations in evolutionary biology. These are simply five common uses of the term by biologists at this period. They will serve to organize our summary of the collected references to chance and the analysis and discussion of the following questions: • What did synthesis authors understand by chance? • How did these authors see chance operating in evolution? • Did their appeals to chance increase or decrease over time during the synthesis? That is, was there a “hardening” of the synthesis, as Gould claimed (1983)? (shrink)

The mathematical theory of communication.

In 1935 geneticist Nikolai Timoféeff-Ressovsky, radiation physicist Karl G. Zimmer, and quantum physicist Max Delbrück published “On the Nature of Gene Mutation and Gene Structure,” known subsequently as the “Three-Man Paper.” This seminal paper advanced work on the physical exploration of the structure of the gene through radiation physics and suggested ways in which physics could reveal definite information about gene structure, mutation, and action. Representing a new level of collaboration between physics and biology, it played an important role in (...) the birth of the new field of molecular biology. The paper’s results were popularized for a wide audience in the What is Life? lectures of physicist Erwin Schrödinger in 1944. Despite its historical impact on the biological sciences, the paper has remained largely inaccessible because it was only published in a short-lived German periodical. Creating a Physical Biology makes the Three Man Paper available in English for the first time. Brandon Fogel’s translation is accompanied by an introductory essay by Fogel and Phillip Sloan and a set of essays by leading historians and philosophers of biology that explore the context, contents, and subsequent influence of the paper, as well as its importance for the wider philosophical analysis of biological reductionism. (shrink)

The Gouldian argument for evolutionary contingency found in Wonderful Life can be dissected into three premises: palaeontological, macro-evolutionary, and developmental. Discussions of evolutionary contingency have revolved primarily around the developmental. However, a shift in methodological practice and new palaeontological evidence subsequent to the book’s publication appears to threaten the palaeontological premise that asserts high Cambrian disparity, or, roughly, that morphological differences between the Cambrian species were high. This presents a prima facie problem: Did the Cambrian consist of enough anatomical variety (...) to fuel a different outcome upon a replay of the tape of life? I investigate the role that disparity plays in the Gouldian argument for evolutionary contingency and conclude that the reduced disparity view does not undermine the argument, per se. As for the macro-evolutionary premise, Gould never offers any explicit justification for the idea that species sorting was indiscriminate or ‘lottery-like’ (though he certainly gestures at it). Putting aside the exegetical, I consider certain empirical avenues for underwriting the macro-evolutionary premise and argue that the most plausible route is to appeal to the special status of mass extinctions, a move that is not entirely unfamiliar to macro-evolutionists. That is, mass extinctions can be good sources of contingency due to their being ‘chancy’ in at least three respects. In sum, I update the palaeontological and macro-evolutionary premises as to reconstrue the Gouldian argument for evolutionary contingency in its strongest form. (shrink)

Much philosophical progress has been made in elucidating the idea of evolutionary contingency in a recent re-burgeoning of the debate. However, additional progress has been impaired on three fronts. The first relates to its characterisation: the under-specification of various contingency claims has made it difficult to conceptually pinpoint the scope to which ‘contingency’ allegedly extends, as well as which biological forms are in contention. That is—there appears to be no systematic means with which to fully specify contingency claims which has (...) led to a tendency for authors to talk past each other. Secondly, on the matter of evidence, recent research has focused on the evidential import of convergent evolution which is taken to disconfirm the evolutionary contingency thesis. However, there has been a neglect of convergent evolution’s converse: ‘evolutionary idiosyncrasies’ or the singular evolution of certain forms, which I argue is evidentially supportive of evolutionary contingency. Thirdly, evolutionary contingency has often been claimed to vary in degrees and that the debate, itself, is a matter of ‘relative significance’. However, there has been no formal method of evaluating the strength of contingency and its relative significance in a particular domain. In this paper, I address all three issues by proposing a systematic means of fully specifying contingency theses with the concept of the modal range. Secondly, I propose an account of evolutionary idiosyncrasies, investigate the explanations for their occurrences, and, subsequently, spell out their significance with respect to the evolutionary contingency thesis. Finally, having been equipped with the evidential counterpart to convergent evolution, I shall sketch a likelihood framework for evaluating, precisely on the basis of a sequence of opposing data, the strength and relative significance of evolutionary contingency in a particular domain. With this in hand, the relative observations of idiosyncrasies and convergences can be informative of the strength and relative significance of contingency in any particular domain. (shrink)

We distinguish dynamical and statistical interpretations of evolutionary theory. We argue that only the statistical interpretation preserves the presumed relation between natural selection and drift. On these grounds we claim that the dynamical conception of evolutionary theory as a theory of forces is mistaken. Selection and drift are not forces. Nor do selection and drift explanations appeal to the (sub-population-level) causes of population level change. Instead they explain by appeal to the statistical structure of populations. We briefly discuss the implications (...) of the statistical interpretation of selection for various debates within the philosophy of biologythe `explananda of selection' debate and the `units of selection' debate. (shrink)

This paper develops a critical response to John Beatty’s recent (2006) engagement with Stephen Jay Gould’s claim that evolutionary history is contingent. Beatty identifies two senses of contingency in Gould’s work: an unpredictability sense and a causal dependence sense. He denies that Gould associates contingency with stochastic phenomena, such as drift. In reply to Beatty, this paper develops two main claims. The first is an interpretive claim: Gould really thinks of contingency has having to do with stochastic effects at the (...) level of macroevolution, and in particular with unbiased species sorting. This notion of contingency as macro-level stochasticity incorporates both the causal dependence and the unpredictability senses of contingency. The second claim is more substantive: Recent attempts by other scientists to put Gould’s claim to the test fail to engage with the hypothesis that species sorting sometimes resembles a lottery. Gould’s claim that random sorting is a significant macroevolutionary phenomenon remains an intriguing and largely untested live hypothesis about evolution. (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)

The Nature of Selection is a straightforward, self-contained introduction to philosophical and biological problems in evolutionary theory. It presents a powerful analysis of the evolutionary concepts of natural selection, fitness, and adaptation and clarifies controversial issues concerning altruism, group selection, and the idea that organisms are survival machines built for the good of the genes that inhabit them. "Sober's is the answering philosophical voice, the voice of a first-rate philosopher and a knowledgeable student of contemporary evolutionary theory. His book merits (...) broad attention among both communities. It should also inspire others to continue the conversation."-Philip Kitcher, Nature "Elliott Sober has made extraordinarily important contributions to our understanding of biological problems in evolutionary biology and causality. The Nature of Selection is a major contribution to understanding epistemological problems in evolutionary theory. I predict that it will have a long lasting place in the literature."-Richard C. Lewontin. (shrink)

In The Structure of Biological Science I argued that the theory of natural selection is a statistical theory for reasons much like those which makes thermodynamics a statistical theory. In particular, the theory claims that fitness differences are large enough and the life span of species long enough for increases in average fitness always to appear in the long run; and this claim, I held, is of the same form as the statistical version of the second law of thermodynamics.For the (...) latter law also makes a claim about the long run, and its statistical character is due to this claim: thermodynamic systems must in the long run approach an equilibrium level of organization that maximizes entropy. Over finite times, given local boundary conditions, an isolated mechanical system, like the molecules in a container of gas, may sometimes interact so as to move the entropy of the system further from, instead of closer to the equilbrium level. But given enough interacting bodies, and enough time, the system will always eventually move in the direction prescribed by the law. Thus, we can attach much higher probabilities to the prediction that non-equilibrium systems will reflect greater entropy in future periods than we can to predictions that they will move in the opposite direction. And as we increase the amount of time and the number of bodies interacting, the strength of the probability we can attach to the prediction becomes greater and greater. (shrink)

In The Structure of Biological Science I argued that the theory of natural selection is a statistical theory for reasons much like those which makes thermodynamics a statistical theory. In particular, the theory claims that fitness differences are large enough and the life span of species long enough for increases in average fitness always to appear in the long run; and this claim, I held, is of the same form as the statistical version of the second law of thermodynamics.For the (...) latter law also makes a claim about the long run, and its statistical character is due to this claim: thermodynamic systems must in the long run approach an equilibrium level of organization that maximizes entropy. Over finite times, given local boundary conditions, an isolated mechanical system, like the molecules in a container of gas, may sometimes interact so as to move the entropy of the system further from, instead of closer to the equilbrium level. But given enough interacting bodies, and enough time, the system will always eventually move in the direction prescribed by the law. Thus, we can attach much higher probabilities to the prediction that non-equilibrium systems will reflect greater entropy in future periods than we can to predictions that they will move in the opposite direction. And as we increase the amount of time and the number of bodies interacting, the strength of the probability we can attach to the prediction becomes greater and greater. (shrink)

Stephen Jay Gould argued that replaying the “tape of life” would result in a radically different evolutionary outcome. Some biologists and philosophers, however, have pointed to convergent evolution as evidence for robust replicability in macroevolution. These authors interpret homoplasy, or the independent origination of similar biological forms, as evidence for the power of natural selection to guide form toward certain morphological attractors, notwithstanding the diversionary tendencies of drift and the constraints of phylogenetic inertia. In this paper, I consider the implications (...) of homoplasy for the debate over the nature of macroevolution. I argue that once the concepts of contingency and convergence are fleshed out, it becomes clear that many instances of homoplasy fail to negate Gould’s overarching thesis, and may in fact support a Gouldian view of life. My argument rests on the distinction between parallelism and convergence, which I defend against a recent challenge from developmental biology. I conclude that despite the difficulties in defining and identifying parallelism, the concept remains useful and relevant to the contingency controversy insofar as it underscores the common developmental origins of iterated evolution. (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)

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)

‘Gouldian arguments’ appeal to the contingency of a scientific domain to establish that domain’s autonomy from some body of theory. For instance, pointing to evolutionary contingency, Stephen Jay Gould suggested that natural selection alone is insufficient to explain life on the macroevolutionary scale. In analysing contingency, philosophers have provided source-independent accounts, understanding how events and processes structure history without attending to the nature of those events and processes. But Gouldian Arguments require source-dependent notions of contingency. An account of contingency is (...) source-dependent when it is indexed to some pattern and some process. Positions like Gould’s do not turn on the mere fact of life’s contingency—that life’s shape could have been different due to its sensitivity to initial conditions, path-dependence or stochasticity. Rather, Gouldian arguments require that the contingency is due to particular kinds of processes: in this case, those which microevolutionary theory cannot account for. This source-dependent perspective clarifies both debates about the nature and importance of contingency, and empirical routes for testing Gould’s thesis. (shrink)

What is the relationship between evolutionary contingency and diversity? The evolutionary contingency thesis emphasizes dependency relations and chance as the hallmarks of evolution. While contingency can be destructive of, for example, the fragile and complex dynamics in an ecosystem, I will mainly focus on the productive or causal aspect of contingency for a particular sort of diversity. There are many sorts of diversities: Gould is most famous for his diversity-to-decimation model, which includes disparate body plans distinguishing different phyla. However, structural (...) diversity construed more broadly spans scales, such as organization in and among cells, structural arrangements and biomechanics on various scales, and even the profile of ancestor-descendent relationships or community structure of interactions within ecosystems. By focusing on stochastic processes in contingent evolution, I argue that contingency causes structural diversity. Specifically, I focus on the plurality of structural types of cells, genetic codes, and phyla diversity as case studies. (shrink)

It is generally thought that objective chances for particular events different from 1 and 0 and determinism are incompatible. However, there are important scientific theories whose laws are deterministic but which also assign non-trivial probabilities to events. The most important of these is statistical mechanics whose probabilities are essential to the explanations of thermodynamic phenomena. These probabilities are often construed as 'ignorance' probabilities representing our lack of knowledge concerning the microstate. I argue that this construal is incompatible with the role (...) of probability in explanation and laws. This is the 'paradox of deterministic probabilities'. After surveying the usual list of accounts of objective chance and finding them inadequate I argue that an account of chance sketched by David Lewis can be modified to solve the paradox of deterministic probabilities and provide an adequate account of the probabilities in deterministic theories like statistical mechanics. (shrink)

We offer a new argument for the claim that there can be non-degenerate objective chance (“true randomness”) in a deterministic world. Using a formal model of the relationship between different levels of description of a system, we show how objective chance at a higher level can coexist with its absence at a lower level. Unlike previous arguments for the level-specificity of chance, our argument shows, in a precise sense, that higher-level chance does not collapse into epistemic probability, despite higher-level properties (...) supervening on lower-level ones. We show that the distinction between objective chance and epistemic probability can be drawn, and operationalized, at every level of description. There is, therefore, not a single distinction between objective and epistemic probability, but a family of such distinctions. (shrink)

How, according to Darwin, does chance variation affect evolutionary outcomes? In his 1866 book, On the Various Contrivances by which British and Foreign Orchids are Fertilised by Insects, Darwin developed an argument that played an important role in his overall case for evolution by natural selection, as articulated in later editions of the Origin. This argument also figured significantly in Darwin's reflections on the theological dimensions of evolution by natural selection.

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)

The goal of this paper is to sketch and defend a new interpretation or 'theory' of objective chance, one that lets us be sure such chances exist and shows how they can play the roles we traditionally grant them. The account is 'Humean' in claiming that objective chances supervene on the totality of actual events, but does not imply or presuppose a Humean approach to other metaphysical issues such as laws or causation. Like Lewis (1994) I take the Principal Principle (...) (PP) to be the key to understanding objective chance. After describing the main features of Humean objective chance (HOC), I deduce the validity of PP for Humean chances, and end by exploring the limitations of Humean chance. (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)

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)

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)

There has recently been a renewed interest in the “force” interpretation of evolutionary biology. In this article, I present the general structure of the arguments for the force interpretation and identify a problem in its overly permissive conditions for being a Newtonian force. I then attempt a solution that (1) helps to illuminate the difference between forces and other types of causes and (2) makes room for random genetic drift as a force. In particular, I argue that forces are not (...) different in kind from other types of causes but rather that forces are situated on a continuum of causes distinguished by their unifying power. †To contact the author, please write to: Department of Philosophy, University of Wisconsin–Madison, 5185 Helen C. White Hall, Madison, WI 53706; e‐mail: [email protected]. (shrink)

This essay examines the claim “path dependence entails irreversibility” from the point of view of evolutionary biology. I argue that evolutionary irreversibility possesses many faces, sometimes conflicting with path dependence. I propose an account of path dependence that does not rely on irreversibility and explains why it more naturally coexists with the notion of (contingent) irreversibility developed by the Belgian paleontologist Louis Dollo. However, I argue that we should not conceive of this relationship as necessary.

Biologists in the last 50 years have increasingly emphasized the role of historical contingency in explaining the distribution and dynamics of biological systems. However, recent work in philosophy of biology has shown that historical contingency carries various interpretations and that we are still lacking a general understanding of historicity, i.e., a framework from which to interpret why and to what extent history matters in biological processes. Building from examples and analyses of the long-term experimental evolution (LTEE) project, this paper argues (...) that historicity possess three essential conditions: (1) multiple possible pasts, (2) multiple possible outcomes at a given instant, and (3) a relationship of causal dependence between these two sets. These criteria can be further specified in two general forms of historicity: dependence on initial conditions and path dependence. More attention is devoted to developing a rigorous account of the latter, which captures the type of historicity displayed by stochastic processes. This paper also highlights that it is often more productive to adopt an instant-relative approach and think in terms of degree of historicity instead of trying to maintain a rigid and absolute dichotomy between historical and ahistorical (completely convergent) processes. (shrink)

The world's foremost geneticist surveys the major developments in what is emerging as the most important single area of scientific inquiry in the twentieth century: biological theory of evolution.

Ideas about heredity and evolution are undergoing a revolutionary change. New findings in molecular biology challenge the gene-centered version of Darwinian theory according to which adaptation occurs only through natural selection of chance DNA variations. In Evolution in Four Dimensions, Eva Jablonka and Marion Lamb argue that there is more to heredity than genes. They trace four "dimensions" in evolution -- four inheritance systems that play a role in evolution: genetic, epigenetic, behavioral, and symbolic. These systems, they argue, can all (...) provide variations on which natural selection can act. Evolution in Four Dimensions offers a richer, more complex view of evolution than the gene-based, one-dimensional view held by many today. The new synthesis advanced by Jablonka and Lamb makes clear that induced and acquired changes also play a role in evolution.After discussing each of the four inheritance systems in detail, Jablonka and Lamb "put Humpty Dumpty together again" by showing how all of these systems interact. They consider how each may have originated and guided evolutionary history and they discuss the social and philosophical implications of the four-dimensional view of evolution. Each chapter ends with a dialogue in which the authors engage the contrarieties of the fictional "I.M.," or Ifcha Mistabra -- Aramaic for "the opposite conjecture" -- refining their arguments against I.M.'s vigorous counterarguments. The lucid and accessible text is accompanied by artist-physician Anna Zeligowski's lively drawings, which humorously and effectively illustrate the authors' points. (shrink)