Perhaps the most readable and accessible of the great works of scientific imagination, The Origin of Species sold out on the day it was published in 1859. Theologians quickly labeled Charles Darwin the most dangerous man in England, and, as the Saturday Review noted, the uproar over the book quickly "passed beyond the bounds of the study and lecture-room into the drawing-room and the public street." Yet, after reading it, Darwin's friend and colleague T. H. Huxley had a different reaction: (...) "How extremely stupid not to have thought of that." Based largely on Darwin's experience as a naturalist while on a five-year voyage aboard H.M.S. Beagle, The Origin of Species set forth a theory of evolution and natural selection that challenged contemporary beliefs about divine providence and the immutability of species. A landmark contribution to philosophical and scientific thought, this edition also includes an introductory historical sketch and a glossary Darwin later added to the original text. Charles Darwin grew up considered, by his own account, "a very ordinary boy, rather below the common standard of intellect." A quirk of fate kept him from the career his father had deemed appropriate--that of a country parson--when a botanist recommended Darwin for an appointment as a naturalist aboard H.M.S. Beagle from 1831 to 1836. Darwin is also the author of the five-volume work Zoology of the Voyage of the Beagle (1839) and The Descent of Man (1871). (shrink)

We have argued elsewhere that: (A) Natural selection is not a cause of evolution. (B) A resolution-of-forces (or vector addition) model does not provide us with a proper understanding of how natural selection combines with other evolutionary influences. These propositions have come in for criticism recently, and here we clarify and defend them. We do so within the broad framework of our own “hierarchical realization model” of how evolutionary influences combine.

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.

This paper provides a philosophical analysis of the ongoing controversy surrounding R.A. Fisher's famous ‘fundamental theorem’ of natural selection. The difference between the ‘traditional’ and ‘modern’ interpretations of the theorem is explained. I argue that proponents of the modern interpretation have captured Fisher's intended meaning correctly and shown that the theorem is mathematically correct, pace the traditional consensus. However, whether the theorem has any real biological significance remains an unresolved issue. I argue that the answer depends on whether (...) we accept Fisher's non-standard notion of environmental change, on which the theorem rests; arguments for and against this notion are explored. I suggest that there is a close link between Fisher's fundamental theorem and the modern ‘gene's eye’ view of evolution. Introduction What Does the Fundamental Theorem Say? Key Concepts Explained Alleged Significance of the FTNS Traditional versus Modern Interpretations of the FTNS The Modern Interpretation Illustrated Fisher's Concept of ‘Environmental Change’ Causality and the Modern Interpretation The Significance of the FTNS Re-considered Appendix CiteULike Connotea Del.icio.us What's this? (shrink)

Recent arguments concerning the nature of causation in evolutionary theory, now often known as the debate between the 'causalist' and 'statisticalist' positions, have involved answers to a variety of independent questions – definitions of key evolutionary concepts like natural selection, fitness, and genetic drift; causation in multi-level systems; or the nature of evolutionary explanations, among others. This Element offers a way to disentangle one set of these questions surrounding the causal structure of natural selection. Doing so (...) allows us to clearly reconstruct the approach that some of these major competing interpretations of evolutionary theory have to this causal structure, highlighting particular features of philosophical interest within each. Further, those features concern problems not exclusive to the philosophy of biology. Connections between them and, in two case studies, contemporary metaphysics and philosophy of physics demonstrate the potential value of broader collaboration in the understanding of evolution. (shrink)

Written by leading theorists and empirical researchers, this book presents new ways of addressing the old question: Why did religion first emerge and then continue to evolve in all human societies? The authors of the book--each with a different background across the social sciences and humanities -- assimilate conceptual leads and empirical findings from anthropology, evolutionary biology, evolutionary sociology, neurology, primate behavioral studies, explanations of human interaction and group dynamics, and a wide range of religious scholarship to construct a deeper (...) and more powerful explanation of the origins and subsequent evolutionary development of religions than can be currently found in what is now vast literature. While explaining religion has been a central question in many disciplines for a long time, this book draws upon a much wider array of literatures to develop a robust and cross-disciplinary analysis of religion. The book remains true to its subtitle by emphasizing an array of both biological and sociocultural forms of selection dynamics that are fundamental to explaining religion as a universal institution in human societies. In addition to Darwinian selection, which can explain the biology and neurology of religion, the book outlines a set of four additional types of sociocultural natural selection that can explain fill out the explanation of why religion first emerged as an institutional system in human societies, and why it has continued to evolve over the last 200,000 years of societal evolution. These sociocultural forms of natural selection are labeled by the names of the early sociologists who first emphasized them; and these can be seen as a necessary supplement to the type of natural selection theorized by Charles Darwin. Explanations of religion that remain in the shadow cast by Darwin's great insights will, it is argued, remain narrow and incomplete when explaining a robust sociocultural phenomena like religion. (shrink)

To evaluate Hume's thesis that causal claims are always empirical, I consider three kinds of causal statement: ?e1 caused e2 ?, ?e1 promoted e2 ?, and ?e1 would promote e2 ?. Restricting my attention to cases in which ?e1 occurred? and ?e2 occurred? are both empirical, I argue that Hume was right about the first two, but wrong about the third. Standard causal models of natural selection that have this third form are a priori mathematical truths. Some are (...) obvious, others less so. Empirical work on natural selection takes the form of defending causal claims of the first two types. I provide biological examples that illustrate differences among these three kinds of causal claim. (shrink)

An influential argument due to Elliott Sober, subsequently strengthened by Denis Walsh and Joel Pust, moves from plausible premises to the bold conclusion that natural selection cannot explain the traits of individual organisms. If the argument were sound, the explanatory scope of selection would depend, surprisingly, on metaphysical considerations concerning origin essentialism. I show that the Sober-Walsh-Pust argument rests on a flawed counterfactual criterion for explanatory relevance. I further show that a more defensible criterion for explanatory relevance (...) recently proposed by Michael Strevens lends support to the view that natural selection can be relevant to the explanation of individual traits. (shrink)

In this article I argue that we should pay extra attention to the ecological dimension of natural selection. By this I mean that we should view natural selection primarily as acting on the outcomes of the interactions organisms have with their environment, which influences their relative reproductive output. A consequence of this view is that natural selection is not sensitive to what system of inheritance ensures reoccurrences of organism-environment interactions over generations. I end by (...) showing the consequences of this view when looking at how processes like niche construction and the Baldwin effect relate to natural selection. (shrink)

This is the first 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. Here I focus 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 being responsible for the presence of the variation it acts upon, for example by directing the course of variation. 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)

This paper analyzes recent attempts to reject reproduction with lineage formation as a necessary condition for evolution by means of natural selection :560–570, 2008; Stud Hist Philos Sci Part C Stud Hist Philos Biol Biomed Sci 42:106–114, 2011; Bourrat in Biol Philos 29:517–538, 2014; Br J Philos Sci 66:883–903, 2015; Charbonneau in Philos Sci 81:727–740, 2014; Doolittle and Inkpen in Proc Natl Acad Sci 115:4006–4014, 2018). Building on the strengths of these attempts and avoiding their pitfalls, it is (...) argued that a robust formulation of evolution by natural selection without reproduction can be established. The main contribution of this paper is a reformulation of Lewontin’s three principles stating that minimal evolution by natural selection occurs when two conditions are met in a population: fitness-related variation and memory. Paradigmatic evolution by natural selection, which can generate adaptations, takes place when an additional condition is present, namely regeneration. (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)

Can selection explain why individuals have the traits they do? This question has generated significant controversy. I will argue that the debate encompasses two separable aspects, to detrimental effect: (1) the role of selection in explaining the origin and evolution of biological traits and (2) the implications this may have for explaining why individuals have the traits they do. (1) can be settled on the basis of evolutionary theory while (2) requires additional, extra-scientific assumptions. By making a distinction (...) between traits affected by a single factor and traits affected by multiple factors I show that selection can, under certain conditions, help explain the origin of traits. Resolving the first aspect enables us to critically assess the various incompatible and independent philosophical commitments made within the second aspect of the debate. (shrink)

Commemorate[s] the centennial of the meeting of the Linnean Society, July 1, 1858, and the paper of Charles Darwin and Alfred Russel Wallace which were read there.

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)

Natural selection is one of the fundamental keys of evolutionary biology, which is as much as saying of almost all life sciences. However, the effort to assess its true meaning has been involved in endless controversies almost from the beginning. The rethinking of natural selection through an abstract scheme with three ingredients - −population, environment and interaction between them− could significantly contribute to clarify this debate.

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

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)

When Charles Darwin published his theory in 1859 the biological community gave very different receptions to the idea of evolution and to the theory of natural selection. Evolution was accepted as widely and rapidly as natural selection was rejected. Most biologists were ready to accept that evolution had occurred, but not that natural selection was its cause. They preferred other explanations of evolution, such as theories of big directed variation, or admitted that they did (...) not know its cause. Darwin himself never maintained that natural selection was the sole cause of evolution. He thought of it as one among several causes, and did not specify how much evolution had occurred by the natural selection of fortuitous variations, and how much by other factors such as the inherited effects of use and disuse. However, Darwin did maintain that natural selection was in principle capable of explaining all the observed properties of organisms. He did not think that there were some characteristics that were particularly likely to have evolved by natural selection, and other kinds that were not. Against this, many of this critics thought that there were characteristics that natural selection was particularly powerless to explain. Thus it could not account for characteristics that were detrimental, or those that seemed useless (such as species differences), or those that were of too little importance for natural selection to have favoured them. There were also characteristics of such complexity that it was unimaginable that natural selection could have built them up in tiny stages from fortuitous, undirected variants. The present essay will be concerned with just one, the last mentioned, of these kinds of characteristics. (shrink)

Thomas Henry Huxley recalled that after he had read Darwin’s Origin of Species, he had exclaimed to himself: “How extremely stupid not to have thought of that!” (Huxley,1900, 1: 183). It is a famous but puzzling remark. In his contribution to Francis Darwin’s Life and Letters of Charles Darwin, Huxley rehearsed the history of his engagement with the idea of transmutation of species. He mentioned the views of Robert Grant, an advocate of Lamarck, and Robert Chambers, who anonymously published Vestiges (...) of the Natural History of Creation (1844), which advanced a crude idea of transmutation. He also recounted his rejection of Agassiz’s belief that species were progressively replaced by the divine hand. He neglected altogether his friend Herbert Spencer’s early Lamarckian ideas about species development, which were also part of the long history of his encounters with the theory of descent. None of these sources moved him to adopt any version of the transmutation hypothesis. (shrink)

Recently some philosophers have emphasized a potentially irreconcilable conceptual antagonism between the statistical characterization of natural selection and the standard scientific discussion of natural selection in terms of forces and causes. Other philosophers have developed an account of the causal character of selectionist statements represented in terms of counterfactuals. I examine the compatibility between such statisticalism and counterfactually based causal accounts of natural selection by distinguishing two distinct statisticalist claims: firstly the suggested impossibility for (...) natural selection to be a cause acting upon populations and secondly the conceptualization that all evolutionary causes occur at the level of interactions between individual organisms. I argue that deriving the latter from the former involves supplementary assumptions concerning precisely what causation is. I critically examine two of these assumptions purportedly preventing natural selection being regarded as a cause: the locality claim and the modularity claim. I conclude that justifying the strongest version of statisticalism—i.e. evolutionary causation only occurs at the level of individual interactions between organisms—would require further metaphysical arguments that are likely to be deemed highly problematic. Additionally, I argue that such a metaphysical position would be considered incongruous with both our scientific and ordinary use of the concepts of causality and explanation as employed within our everyday epistemological framework. (shrink)

Aside from the Principia and occasional appearances of the Opticks , Newton' writings have remained largely inaccessible to students of philosophy, science, and literature as well as to other readers. This book provides a remedy with wide representation of the interests, problems, and diverse philosophic issues that preoccupied the greatest scientific mind of the seventeenth century. Grouped in sections corresponding to methods, principles, and theological considerations, these selections feature explanatory notes and cross-references to related essays.

Some naturalistic theories of consciousness give an essential role to teleology.1 This teleology is said to arise due to natural selection. Thus it is claimed that only certain states, namely, those that have been selected for by evolutionary pro- cesses because they contribute to (or once contributed to) an organism’s fitness, are conscious states. These theories look as if they are assigning a creative role to natural selection. If a state is conscious only if it has (...) been selected for, then selec- tion appears to be able to create a new feature of states, namely, their conscious nature. Yet, intuitively, natural selection cannot create anything. Natural selec- tion chooses certain features that already exist and makes them more (or less) prevalent in a population, but it cannot bring features into existence itself. Natu- ral selection can select for conscious states, but it cannot create them. This con- clusion has recently been argued for by Steven Horst (1999). If it is right, then teleological theories of conscious states should be rejected. A state cannot become a conscious experience in virtue of having been selected for by evolu- tionary process. (shrink)

Building on work by Popper, Schweber, Nozick, Sober, and others in a still-growing literature, I explore here the conceptual kinship between Adam Smith''s ''invisible hand'' and Darwinian natural selection. I review the historical ties, and examine Ullman -Margalit''s ''constraints'' on invisible-hand accounts, which I later re-apply to natural selection, bringing home the close relationship. These theories share a ''parent'' principle, itself neither biological no politico-economic, that collective order and well-being can emerge parsimoniously from the dispersed action (...) of individuals. The invisible hand operates on ''memes'' the way natural selection operates on genes. Like Darwin''s concept, it brings together traditional opposites, ''nature'' and ''selection,'' forming a saltation-mitigating transition between biological instinct and full-blown conscious design. Herschel''s criterion of confirmation, which Darwin long strove to satisfy, is itself an invisible hand-like meme – a ''Midas effect'' revealing and rewarding the fittest theories, Darwin''s and Smith''s emphatically among them. (shrink)

Reification is a concept of critical theory that denotes certain problematic, habitualised forms of objectification. In this article, I examine whether the concept can be applied in environmental philosophy and what value it has for environmental critical theory. I begin by introducing the concept and the two senses in which reification of the non-human world has been discussed in the literature: first, denoting the misrecognition of others’ attitudes towards the natural world; and second, denoting a misconceived relationship (...) between humans and their environment. After this, I introduce two potentially new subjects of reification: non-human animals and non-human nature. I also discuss two phenomena that could count as reification: industrial meat production and the commodification of ecological systems. (shrink)

... Difficulty of distinguishing between Varieties and Species — Origin of Domestic ... and Origin— Principle of Selection anciently followed, its Effects— ...

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

It should be evident from the foregoing discussion that one man's natural selection is not necessarily the same as another man's. Why should this be so? How can two theories, which both Matthew and Darwin believed to be nearly identical, be so dissimilar? Apparently, neither Matthew nor Darwin understood the other's theory. Each man's viewpoint was colored by his own intellectual background and philosophical assumptions, and each read these into the other's ideas. The words sounded the same, so (...) they assumed the concepts must als be the same.123As Ghiselin has pointed out, historians attempting to evaluate Darwin's predecessors have been similarly blinded by a preoccupation with words, without regard to their proper context.124 In the case of Matthew, the practice of quoting only brief passages from the appendix to Naval Timber and Arboriculture, without relating them to the rest of his work, has suggested a greater resemblance to Darwin's theory than actually exists.It is clear, both from the use which Matthew made of his ideas and from the philosophical roots of his natural world view, that he could not have arrived at the concept of natural selection by the same thought process which Darwin employed. His discussion of natural selection is presented not as an argument, but as an axiom. No theory is proposed, no evidence marshaled to support it. Natural selection is stated as a fact, a Law of Nature, unquestioned, and presumably, unquestionable.Despite his clamor for recognition as the discoverer of natural selection, Matthew recognized and acknowledged this very fundamental difference between Darwin and himself. In a letter to the Gardener's Chronicle of May 12, 1860, he wrote:To me the conception of this law of Nature came intuitively as a self-evident fact, almost without an effort of concentrated thought. Mr. Darwin here seems to have more merit in the discovery than I have had—to me it did not appear a discovery. He seems to have worked it out by inductive reason, slowly and with due caution to have made his way synthetically from fact to fact onwards; while with me it was by a general glance at the scheme of Nature that I estimated this select production of species as an a priori recognisable fact—an axiom, requiring only to be pointed out to be admitted by unprejudiced minds of sufficient grasp.125In the same letter, Matthew maintained that his ideas had not been accepted because “the age was not ripe for such ideas.”126 Nor, he said, was the present age. He considered the inability of most of Darwin's critics to grasp his theory to be “incurable.” Yet he did not argue that natural selection should be accepted because of the evidence, but rather, that it should be accepted on faith:Belief here requires a certain grasp of mind. No direct proof of phenomena embracing so long a period of time is within the compass of short-lived man. To attempt to satisfy a school of ultra skeptics, who have a wonderfully limited power of perception of means to ends... would be labour in vain.... They could not be brought to conceive the purpose of a handsaw though they saw its action, if the whole individual building it assisted to construct were not presented complete before their eyes... Like a child looking upon the motion of a wheel in an engine they would only perceive and admire... without noticing its agency in... affecting the purposed end.127Here, then, is the final irony. In a passage urging acceptance of Darwin's theory, a theory which was to banish design and purpose from the natural world, we find echoes of Paley and of Providence.Loren Eiseley has lamented the fact that Matthew “did not bring his views into the open, because the amount of ground he was able to cover in a few paragraphs suggests that he might have been able to sustain a longer treatise.”128 Now that the intellectual and historical context of Matthew's ideas are known, this statement is no longer tenable. Matthew was not a scientist, and his books were not written as biological treatises. His discussions of natural selection were not attempts to “cover ground” in advancing a particular scientific theory, but were simply reflections of his own assumptions about the natural world.Furthermore, despite Matthew's acceptance of evolution and natural selection, his biological thought was basically conservative on points where Darwin's was radical. Where Matthew saw a series of stable worlds interrupted by violent upheavals, Darwin saw a continuous process of change in an ever-fluctuating world. Where Matthew conceived of species in terms of Aristotelian classes and essences, Darwin revolutionized our concept of species by treating them as populations. Where Matthew saw a world of design and beauty functioning according to natural laws laid down by benevolent Providence, Darwin abolished design and Providence from nature and ushered in a world which cycles ever onward according to laws of chance and probability.It is not even particularly useful to point to Matthew as evidence that evolution was “in the air” prior to 1859.129 His ideas did not represent the first wave of a coming revolution, but were the product of his own personal philosophical outlook, as expressed in the context of the biological thought of the 1830's. Matthew is important in the history of ideas, not simply because he accepted the concept of evolution or thought of something resembling natural selection, but because he did so without overthrowing, in his own mind, any of the basic philosophical assumptions which had underlain biological science since Aristotle. In recognizing Matthew's failure to do so, we are in a position to appreciate more fully the significance of the Darwinian Revolution. (shrink)

ORIGIN OF SPECIES. INTRODUCTION. When on board HMS 'Beagle,' as naturalist, I was ranch struck with certain facts in the distribution of the organic beings ...

Recent discussion of mechanism has suggested new approaches to several issues in the philosophy of science, including theory structure, causal explanation, and reductionism. Here, I apply what I take to be the fruits of the 'new mechanical philosophy' to an analysis of a contemporary debate in evolutionary biology about the role of natural selection in speciation. Traditional accounts of that debate focus on the geographic context of genetic divergence--namely, whether divergence in the absence of geographic isolation is possible (...) (or significant). Those accounts are at best incomplete, I argue, because they ignore the mechanisms producing divergence and miss what is at stake in the biological debate. I argue that the biological debate instead concerns the scope of particular speciation mechanisms which assign different roles to natural selection at various stages of divergence. The upshot is a new interpretation of the crux of that debate--namely, whether divergence with gene flow is possible (or significant) and whether the isolating mechanisms producing it are adaptive. (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)

The notion that natural selection is a process of fitness maximization gets a bad press in population genetics, yet in other areas of biology the view that organisms behave as if attempting to maximize their fitness remains widespread. Here I critically appraise the prospects for reconciliation. I first distinguish four varieties of fitness maximization. I then examine two recent developments that may appear to vindicate at least one of these varieties. The first is the ‘new’ interpretation of Fisher's (...) fundamental theorem of natural selection, on which the theorem is exactly true for any evolving population that satisfies some minimal assumptions. The second is the Formal Darwinism project, which forges links between gene frequency change and optimal strategy choice. In both cases, I argue that the results fail to establish a biologically significant maximization principle. I conclude that it may be a mistake to look for universal maximization principles justified by theory alone. A more promising approach may be to find maximization principles that apply conditionally and to show that the conditions were satisfied in the evolution of particular traits. (shrink)

In this paper, I am clarifying and defending my argument in favor of the claim that cumulative selection can explain adaptation provided that the environmental resources are limited. Further, elaborate on what this limitation of environmental resources means and why it is relevant for the explanatory power of natural selection.

In Darwinism's Struggle for Survival Jean Gayon offers a philosophical interpretation of the history of theoretical Darwinism. He begins by examining the different forms taken by the hypothesis of natural selection in the nineteenth century and the major difficulties which it encountered, particularly with regard to its compatibility with the theory of heredity. He then shows how these difficulties were overcome during the seventy years which followed the publication of Darwin's Origin of Species, and he concludes by analysing (...) the major features of the genetic theory of natural selection, as it developed from 1920 to 1960. This rich and wide-ranging study will appeal to philosophers and historians of science and to evolutionary biologists. (shrink)

Here I advance two related evolutionary propositions. (1) Natural selection is most often considered to require competition between reproducing “individuals”, sometimes quite broadly conceived, as in cases of clonal, species or multispecies-community selection. But differential survival of non-competing and non-reproducing individuals will also result in increasing frequencies of survival-promoting “adaptations” among survivors, and thus is also a kind of natural selection. (2) Darwinists have challenged the view that the Earth’s biosphere is an evolved global homeostatic (...) system. Since there is only one biosphere, reproductive competition cannot have been involved in selection for such survival-promoting adaptations, they claim. But natural selection through survival could reconcile Gaia with evolutionary theory. (shrink)

Fisher’s ‘fundamental theorem of natural selection’ is notoriously abstract, and, no less notoriously, many take it to be false. In this paper, I explicate the theorem, examine the role that it played in Fisher’s general project for biology, and analyze why it was so very fundamental for Fisher. I defend Ewens (1989) and Lessard (1997) in the view that the theorem is in fact a true theorem if, as Fisher claimed, ‘the terms employed’ are ‘used strictly as defined’ (...) (1930, p. 38). Finally, I explain the role that projects such as Fisher’s play in the progress of scientific inquiry. (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)

This paper is a commentary on the focal article by Grafen and on earlier papers of his on which many of the results of this focal paper depend. Thus it is in effect a commentary on the “formal Darwinian project”, the focus of this sequence of papers. Several problems with this sequence are raised and discussed. The first of these concerns fitness maximization. It is often claimed in these papers that natural selection leads to a maximization of fitness (...) and that this view is claimed in Fisher’s “fundamental theorem of natural selection”. These claims are refuted, and various incorrect statements about the meaning and interpretation of the fundamental theorem of natural selection, in this sequence and in other papers by other authors, are discussed. Next, much of the work in this sequence rests on the first Price equation. In the deterministic (infinite population) case this equation is no more than the standard classical equation relating to changes in gene frequencies. In the stochastic case the equation gives the change in gene frequencies as the sum of two terms (the second of which vanishes in the deterministic case). These two terms are of essentially equal importance in the situation considered in the focal article, yet one of Grafen’s results ignores the second term in the stochastic analysis. This is associated with a wavering between deterministic and stochastic analyses and the use of the Price fitness concept and the classical fitness concept. These comments cast doubts on Grafen’s optimization theory. (shrink)

RESUMEN: El tema de este trabajo es la reconstrucción de la teoría de la selección natural darwiniana. Me propongo esbozar la ley fundamental de esta teoría de manera informal a partir de sus aplicaciones en El origen de las especies de Darwin y presentar sus conceptos fundamentales. Presentaré la red teórica de leyes especiales que surgen de la especialización de esta ley fundamental. Supondré el estructuralismo como marco metateórico. Señalaré también algunas consecuencias que mi propuesta tiene sobre ciertas discusiones (...) metateóricas en torno a la teoría y, finalmente, relacionaré mi reconstrucción con otras reconstrucciones disponibles.ABSTRACT: This paper is about the reconstruction of the Darwinian Theory of Natural Selection. My aim here is to outline the fundamental law of this theory in an informal way from its applications in The Origin of Species and to make explicit its fundamental concepts. I will introduce the theory-nets of special laws that arisefrom the specialization of the fundamental law. I will assume the metatheoretical structuralist frame. I will also point out many consequences that my proposal has about a few metatheoretical discussions around the theory and, finally, I will relate my propose to other reconstructions available. (shrink)

The author replies to H. Allen Orr's review of "Darwin's Dangrous Idea" (Boston Review, Summer 1996).

Fisher’s ‘fundamental theorem of natural selection’ is notoriously abstract, and, no less notoriously, many take it to be false. In this paper, I explicate the theorem, examine the role that it played in Fisher’s general project for biology, and analyze why it was so very fundamental for Fisher. I defend Ewens and Lessard in the view that the theorem is in fact a true theorem if, as Fisher claimed, ‘the terms employed’ are ‘used strictly as defined’. Finally, I (...) explain the role that projects such as Fisher’s play in the progress of scientific inquiry. (shrink)