Adaptations can occur at different hierarchical levels, but it can be difficult to identify the level of adaptation in specific cases. A major problem is that selection at a lower level can filter up, creating the illusion of selection at a higher level. We use optimality modeling of the volvocine algae to explore the emergence of genuine group adaptations. We find that it is helpful to develop an explicit model for what group fitness would be in the absence of group-level (...) relationships between traits and group fitness. We call this “counterfactual fitness,” because in many actual cases of interest there are group-level relationships. Once counterfactual fitness is modeled, the difference between effects that filter up and genuine group selection is explicit and so, therefore, is the distinction between apparent and genuine group adaptations. We call the latter group-specific adaptations. Recognizing group-specific adaptations is important because only group-specific adaptations would cause the lower-level units to be maladapted if they were to leave the group and enter a global cell-level population. Thus, as group-specific adaptations evolve, they create selective pressure for increased cohesiveness and individuality of groups. This article suggests that group-specific adaptations could be present in the simplest, earliest branching colonial volvocine species, which do not have distinct specialized cells. The article also makes predictions about the kind of empirical evidence needed to support or refute the hypothesis that a particular trait is a group-level adaptation. (shrink)

A number of scholars have recently defended the claim that there is a close connection between the evolutionary biological notion of fitness and the economic notion of utility: both are said to refer to an organism’s success in dealing with its environment, and both are said to play the same theoretical roles in their respective sciences. However, an analysis of two seemingly disparate but in fact structurally related phenomena—‘niche construction’ (the case where organisms change their environment to make it fit (...) to their needs) and ‘adaptive preferences’ (the case where agents change their wants to make them fit to what the world has given them)—shows that one needs to be very careful about the postulation of this sort of fitness–utility connection. Specifically, I here use the analysis of these two phenomena to establish when connecting fitness and utility is and is not possible. (shrink)

In this paper, I consider the recent resurgence of “evolutionary economics”—the idea that evolutionary theory can be very useful to push forward key debates in economics—and assess the extent to which it rests on a plausible foundation. To do this, I first distinguish two ways in which evolutionary theory can, in principle, be brought to bear on an economic problem—namely, evidentially and heuristically—and then apply this distinction to the three major hypotheses that evolutionary economists have come to defend: the implausibility (...) of rational choice theory as an account of economic rationality, the idea that firms are autonomous economic agents, and the need for a more dynamic, less equilibrium-focused economic methodology. In each of these cases, I conclude negatively: the relevant evolutionary considerations neither suggest interesting and novel hypotheses to investigate further nor are backed up by the needed data to constitute genuine evidence. I end by distinguishing this criticism of evolutionary economics from others that have been put forward in the literature: in particular, I make clear that, unlike those of other critics, the arguments of this paper are based on epistemic—not structural—considerations and therefore leave more room for a plausible form of evolutionary economics to come about in the future. (shrink)

I distinguish two versions of kin selection theory—a purely genetic version and a version that also appeals to cultural forms of cooperation —and present an argument in favor of using the former when it comes to accounting for the evolution of cooperation in non-human organisms. Specifically, I first show that both GKST and WKST are equally mathematically coherent—they can both be derived from the Price equation—but not necessarily equally empirically plausible, as they are based on different assumptions about the inheritance (...) system underlying the cooperative phenotype. Given this, I then, second, present a model selection theoretic argument in favor of GKST over WKST. This argument is based on the fact that, in non-human cases, the former theory is likely to be as empirically successful as WKST, while containing fewer degrees of freedom. I end by defending both the intrinsic importance of this argument and its relevance to the discussion surrounding the “gene’s eye view of evolution.”. (shrink)

Two questions are raised for Grafen’s formal darwinism project of aligning evolutionary dynamics under natural selection with the optimization of phenotypes for individuals of a population. The first question concerns mean fitness maximization during frequency-dependent selection; in such selection regimes, not only is mean fitness typically not maximized but it is implausible that any parameter closely related to fitness is being maximized. The second question concerns whether natural selection on inclusive fitness differences can be regarded as individual selection or whether (...) it leads to a departure from the central motivation that led to the formal darwinism project, viz., to show that “Darwinian” evolution through individual selection leads to “good design” or phenotypic adaptation through trait optimization. (shrink)

The idea that natural selection can operate on cultural as well as genetic variation is central to recent theories of cultural evolution. This raises an overarching question: how much of traditional evolutionary theory, which was formulated in population-genetic terms, can survive intact once the possibility of cultural inheritance is taken into account? This question is addressed in relation to R. A. Fisher’s “fundamental theorem” of natural selection. Though Fisher’s theorem may appear to be an essentially genetic result, a version of (...) the theorem can be derived that applies to any evolving population, with an arbitrary inheritance mechanism. However, I argue that the biological significance of this “generalized” version of the fundamental theorem is less than that of the Fisherian original. (shrink)

Social evolution theory conventionally takes an externalist explanatory stance, treating observed cooperation as explanandum and the positive assortment of cooperative behaviour as explanans. We ask how the circumstances bringing about this positive assortment arose in the first place. Rather than merely push the explanatory problem back a step, we move from an externalist to an interactionist explanatory stance, in the spirit of Lewontin and the Niche Construction theorists. We develop a theory of ‘social niche construction’ in which we consider biological (...) entities to be both the subject and object of their own social evolution. Some important cases of the evolution of cooperation have the side-effect of causing changes in the hierarchical level at which the evolutionary process acts. This is because the traits that act to align the fitness interests of particles in a collective can also act to diminish the extent to which those particles are bearers of heritable fitness variance, while augmenting the extent to which collectives of such particles are bearers of heritable fitness variance. In this way, we can explain upward transitions in the hierarchical level at which the Darwinian machine operates in terms of particle-level selection, even though the outcome of the process is a collective-level selection regime. Our theory avoids the logical and metaphysical paradoxes faced by other attempts to explain evolutionary transitions. (shrink)

Both Popper and van Fraassen have used evolutionary analogies to defend their views on the aim of science, although these are diametrically opposed. By employing Price's equation in an illustrative capacity, this paper considers which view is better supported. It shows that even if our observations and experimental results are reliable, an evolutionary analogy fails to demonstrate why conjecture and refutation should result in: (1) the isolation of true theories; (2) successive generations of theories of increasing truth-likeness; (3) empirically adequate (...) theories; or (4) successive generations of theories of increasing proximity to empirical adequacy. Furthermore, it illustrates that appeals to induction do not appear to help. It concludes that an evolutionary analogy is only sufficient to defend the notion that the aim of science is to isolate a particular class of false theories, namely those that are empirically inadequate. (shrink)

Holobionts, consisting of a host and diverse microbial symbionts, function as distinct biological entities anatomically, metabolically, immunologically, and developmentally. Symbionts can be transmitted from parent to offspring by a variety of vertical and horizontal methods. Holobionts can be considered levels of selection in evolution because they are well-defined interactors, replicators/reproducers, and manifestors of adaptation. An initial mathematical model is presented to help understand how holobionts evolve. The model offered combines the processes of horizontal symbiont transfer, within-host symbiont proliferation, vertical symbiont (...) transmission, and holobiont selection. The model offers equations for the population dynamics and evolution of holobionts whose hologenomes differ in gene copy number, not in allelic or loci identity. The model may readily be extended to include variation among holobionts in the gene identities of both symbionts and host. (shrink)

The history of biological systematics documents a continuing tension between classifications in terms of nested hierarchies congruent with branching diagrams (the ‘Tree of Life’) versus reticulated relations. The recognition of conflicting character distribution led to the dissolution of the scala naturae into reticulated systems, which were then transformed into phylogenetic trees by the addition of a vertical axis. The cladistic revolution in systematics resulted in a representation of phylogeny as a strictly bifurcating pattern (cladogram). Due to the ubiquity of character (...) conflict—at the genetic or morphological level, or at any level in between—some characters will necessarily have to be discarded ( qua noise) in favor of others in support of a strictly bifurcating phylogenetic tree. Pattern analysts will seek maximal congruence in the distribution of characters (ultimately of any kind) relative to a branching tree-topology; process explainers will call such tree-topologies into question by reference to incompatible evolutionary processes. Pattern analysts will argue that process explanations must not be brought to bear on pattern reconstruction; process explainers will insist that the reconstructed pattern requires a process explanation to become scientifically relevant, i.e., relevant to evolutionary theory. The core question driving the current debate about the adequacy of the ‘Tree of Life’ metaphor seems to be whether the systematic dichotomization of the living world is an adequate representation of the complex evolutionary history of global biodiversity. In ‘Questioning the Tree of Life’, it seems beneficial to draw at least four conceptual distinctions: pattern reconstruction versus process explanation as different epistemological approaches to the study of phylogeny; open versus closed systems as expressions of different kinds of population (species) structures; phylogenetic trees versus cladograms as representations of evolutionary processes versus patterns of relationships; and genes versus species as expressions of different levels of causal integration and evolutionary transformation. (shrink)

The goal of forming a science of intentional behavior requires a more richly detailed account of symbolic systems than is assumed by the authors. Cultural systems are not simply the equivalent in the ideational domain of culture of the purported Baldwin Effect in the genetic domain. © 2014 Cambridge University Press.

Biological fitness is a foundational concept in the theory of natural selection. Natural selection is often defined in terms of fitness differences as “any consistent difference in fitness (i.e., survival and reproduction) among phenotypically different biological entities” (Futuyma 1998, 349). And in Lewontin’s (1970) classic articulation of the theory of natural selection, he lists fitness differences as one of the necessary conditions for evolution by natural selection to occur. Despite this foundational position of fitness, there remains much debate over the (...) nature of fitness, especially whether fitness differences can truly be said to cause evolutionary change. In recent years these debates have crystalized into two camps: (1) causalists, who see fitness differences as being one of the causes of evolutionary change, and (2) statisticalists, who deny the causal efficacy of fitness and instead hold that “fitness is a mere statistical, noncausal property of trait types” (Walsh 2010, 148). (shrink)

The emergence of individuality during the evolutionary transition from single cells to multicellularity poses a range of problems. A key issue is how variation in lower‐level individuals generates a corporate (collective) entity with Darwinian characteristics. Of central importance to this process is the evolution of a means of collective reproduction, however, the evolution of a means of collective reproduction is not a trivial issue, requiring careful consideration of mechanistic details. Calling upon observations from experiments, we draw attention to proto‐life cycles (...) that emerge via unconventional routes and that transition, in single steps, individuality to higher levels. One such life cycle arises from conflicts among levels of selection and invokes cheats as a primitive germ line: it lays the foundation for collective reproduction, the basis of a self‐policing system, the selective environment for the emergence of development, and hints at a plausible origin for a soma/germ line distinction. (shrink)

This paper reviews Rosenberg’s and McShea’s textbook in philosophy of biology, entitled Philosophy of Biology. A Contemporary Introduction. I insist on the excellent quality of this textbook, then I turn to more critical comments, which deal mainly with what philosophy of biology is, and what it should be.

Biological individuality is a major topic of discussion in biology and philosophy of biology. Recently, several objections have been raised against traditional accounts of biological individuality, including the objections of monism, theory-centrism, ahistoricity, disciplinary isolationism, and the multiplication of conceptual uncertainties. In this introduction, I will examine the current philosophical landscape about biological individuality, and show how the contributions gathered in this special issue address these five objections. Overall, the aim of this issue is to offer a more diverse, unifying, (...) and scientifically informed conception of what a biological individual is. (shrink)

The definition of biological individuality is one of the most discussed topics in philosophy of biology, but current debate has focused almost exclusively on evolution-based accounts. Moreover, several participants in this debate consider the notions of a biological individual and an organism as equivalent. In this paper, I show that the debates would be considerably enriched and clarified if philosophers took into account two elements. First, physiological fields are crucial for the understanding of biological individuality. Second, the category of biological (...) individuals should be divided into two subcategories: physiological individuals and evolutionary individuals, which suggests that the notions of organism and biological individual should not be used interchangeably. I suggest that the combination of an evolutionary and a physiological perspective will enable biologists and philosophers to supply an account of biological individuality that will be both more comprehensive and more in accordance with scientific practices. (shrink)

What does human evolutionary theory reveal about the origins of human nature and the constraints it imposes on human cognition, behavior, and society? “The whole field of human evolution is pregnant with philosophical questions of great interest”, Michael Ruse concludes in the final passage of The Philosophy of Human Evolution. This engaging and eminently readable romp through the philosophical landscape of human evolution fills a significant niche in the existing literature. There are numerous scientific texts surveying historical and contemporary problems (...) in the field of human evolution, and there are many philosophical texts exploring conceptual and methodological problems in evolutionary theory. Ruse interweaves scientific and philosophical work on human evolution with the latest work in biological theory to produce a unique and timely book, one that addresses a range of important topics that rarely find their way into works of this genre. With a relentless lucidity and charming, f .. (shrink)

In this paper, I argue that the increasing data about non-genetic inheritance requires the construction of a new conceptual framework that should complement the inclusive approaches already discussed in the literature. More precisely, I hold that this framework should be epistemologically relevant for evolutionary biologists in capturing the limits of extended inheritance and in reassessing the boundaries of biological systems that transmit traits to their offspring. I outline the first elements of an organizational account of extended inheritance. In this account, (...) the category of inherited factors is neither restricted to genes nor extended to stable resources related to trans-generational similarities. Instead, it includes persisting constitutive elements appearing as difference makers for heterogeneous organizational constraints, namely for heterogeneous constitutive parts whose specific role is to harness flows of matter and energy across generations of clearly delimited extended organized systems. This both inclusive and restrictive framework opens an additional way to apprehend how extended inheritance may affect evolutionary trajectories. (shrink)

Natural selection is an extremely powerful process – so powerful, in fact, that it is often tempting to deploy it in explaining phenomena as wide-ranging as the persistence of blue eyes, the origins or persistence of religious belief, or, the history of science. One long-standing debate among both critics and advocates of Darwin’s concerns the scope of Darwinian explanations, and how we are to draw the line. Peter Godfrey-Smith’s Darwinian Populations and Natural Selection is a detailed examination of this question. (...) The book explores the criteria for what may count as a “Darwinian population,” by which Godfrey-Smith means, which collections of entities have the capacity to undergo evolution via natural selection (p. 6). Drawing upon his answer to this question, Godfrey-Smith examines and provides his own solution to the following long-standing debates in philosophy of biology: (a) the twin problems of reproduction and individuation of biological entities, (b) the persistent “gene’s eye view,” (c) the levels and units of selection problem, and (d) the evolution of cultural artifacts and behaviors. (shrink)

With an increasing number of ways to ‘assist’ reproduction, some bioethicists have started to wonder what it takes to become a genetic parent. It is widely agreed that sharing genes is not enough to substantiate the parent–offspring relation, but what is? Without a better understanding of the concept of reproduction, our thinking about parent–offspring relations and the ethical issues surrounding them risk being unprincipled. Here, I address that problem by offering a principled account of reproduction—the Overlap, Development and Persistence account—which (...) I believe best captures the meaning of ‘genetic parenthood’. (shrink)

It is an unfortunate fact of academic life that there is a sharp divide between science and philosophy, with scientists often being openly dismissive of philosophy, and philosophers being equally contemptuous of the naivete ́ of scientists when it comes to the philosophical underpinnings of their own discipline. In this paper I explore the possibility of reducing the distance between the two sides by introducing science students to some interesting philosophical aspects of research in evolutionary biology, using biological theories of (...) the origin of religion as an example. I show that philosophy is both a discipline in its own right as well as one that has interesting implications for the understanding and practice of science. While the goal is certainly not to turn science students into philoso- phers, the idea is that both disciplines cannot but benefit from a mutual dialogue that starts as soon as possible, in the classroom. (shrink)

The debate about the levels of selection has been one of the most controversial both in evolutionary biology and in philosophy of science. Okasha’s book makes the sort of contribution that simply will not be able to be ignored by anyone interested in this field for many years to come. However, my interest here is in highlighting some examples of how Okasha goes about discussing his material to suggest that his book is part of an increasingly interesting trend that sees (...) scientists and philosophers coming together to build a broadened concept of “theory” through a combination of standard mathematical treatments and conceptual analyses. Given the often contentious history of the relationship between philosophy and science, such trend cannot but be welcome. (shrink)

The propensity interpretation of fitness (PIF) is commonly taken to be subject to a set of simple counterexamples. We argue that three of the most important of these are not counterexamples to the PIF itself, but only to the traditional mathematical model of this propensity: fitness as expected number of offspring. They fail to demonstrate that a new mathematical model of the PIF could not succeed where this older model fails. We then propose a new formalization of the PIF that (...) avoids these (and other) counterexamples. By producing a counterexample-free model of the PIF, we call into question one of the primary motivations for adopting the statisticalist interpretation of fitness. In addition, this new model has the benefit of being more closely allied with contemporary mathematical biology than the traditional model of the PIF. (shrink)

Multiple biological groups, such as ant colonies, appear to have a noteworthy inefficiency: they contain vast amounts of redundant members that are not strictly needed to maintain the group. Philosophers and biologists have proposed that such inefficiency is illusory because redundancy enhances the resilience of groups when living under harsh conditions. Still, this proposal is unsatisfactory in different respects. First, it is too vague to account for when redundancy is selectively advantageous. Furthermore, it overlooks cases in which redundancy fails to (...) increase the resilience of groups. This paper offers an account of group redundancy that addresses these difficulties. Specifically, it advances the hypothesis that the mere presence of harsh conditions is not what drives the evolution of redundancy; rather, it is the fact organisms are often unable to predict when they will face harsh conditions. Redundancy enables groups to properly respond to unpredictable circumstances without resorting to the unreliable detection systems of their members. A better understanding of the phenomenon of redundancy is likely to impact other key issues in philosophy of science, including the evolution of cooperation and transitions in individuality, and the role of redundancy in complex systems, be they biological or not. (shrink)

Coen offers a unified explanation of natural selection, development, learning and cultural change, based on seven fundamental principles: population variation, persistence, reinforcement, competition, cooperation, combinatorial richness and recurrence. I discuss whether all seven principles are justified, successfully fit the four processes, encompass life processes only, and have any strong explanatory import. I find each of these claims doubtful.

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)

WITTGENSTEIN ON THE ARBITRARINESS OF GRAMMAR Michael N. Forster What is the nature of a conceptual scheme? Are there alternative conceptual schemes?

Ernst Mayr’s classical work on the nature of causation in biology has had a huge influence on biologists as well as philosophers. Although his distinction between proximate and ultimate causation recently came under criticism from those who emphasize the role of development in evolutionary processes, the formal relationship between these two notions remains elusive. Using causal graph theory, this paper offers a unified framework to systematically translate a given “proximate” causal structure into an “ultimate” evolutionary response, and illustrates evolutionary implications (...) of various kinds of causal mechanisms including epigenetic inheritance, maternal effects, and niche construction. These results not only reveal the essential interplay between proximate and ultimate causation in the study of evolution, but also provide a formal method to evaluate or discover non-standard or yet unknown evolutionary phenomena. (shrink)

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

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

Over the past decade philosophers of biology have discussed whether evolutionary theory is a causal theory or a phenomenological study of evolution based solely on the statistical features of a population. This article reviews this controversy from three aspects, respectively concerning the assumptions, applications, and explanations of evolutionary theory, with a view to arriving at a definite conclusion in each contention. In so doing I also argue that an implicit methodological assumption shared by both sides of the debate, namely the (...) overconfidence in conceptual analysis as a tool to understand the scientific theory, is the real culprit that has both generated the problem and precluded its solution for such a long time. (shrink)

Kin selection and multilevel selection are alternative approaches for studying the evolution of social behaviour, the relation between which has long been a source of controversy. Many recent theorists regard the two approaches as ultimately equivalent, on the grounds that gene frequency change can be correctly expressed using either. However, this shows only that the two are formally equivalent, not that they offer equally good causal representations of the evolutionary process. This article articulates the notion of an ‘adequate causal representation’ (...) using causal graphs, and then seeks to identify circumstances under which kin and multilevel selection do and do not satisfy the test of causal adequacy. 1 Introduction2 The KS and MLS Approaches2.1 The MLS decomposition2.2 The KS decomposition3 Equivalence and Causality4 Two Problem Cases4.1 The non-social trait case4.2 Genotypic selection with meiotic drive5 Casual Adequacy: A Graphical Approach5.1 The basic idea5.2 Graphs with individual and group variables5.3 Cases where KS is causally adequate5.4 Cases where MLS is causally adequate6 Discussion6.1 Relation to previous work. (shrink)

Evolutionary biology is striking for its ability to explain a large and diverse range of empirical phenomena on the basis of a few general theoretical principles. This article offers a philosophical perspective on the way that evolutionary biology has come to achieve such impressive generality, by focusing on “the strategy of endogenization”. This strategy involves devising evolutionary explanations for biological features that were originally part of the background conditions, or scaffolding, against which such explanations take place. Where successful, the strategy (...) moves biology a step closer to the ideal of explaining as much as possible from evolutionary first principles. The strategy of endogenization is illustrated through a series of biological examples, historical and recent, and its philosophical implications are explored. (shrink)

This paper contains replies to the reviews of my book by Steven Downes, Massimo Pigliucci and Deborah Shelton & Rick Michod.

Elliott Sober and Ken Waters both raise interesting and difficult challenges for various aspects of the position I set out in Evolution and the Levels of the Selection. I am grateful to them for their penetrating criticisms of my work, and find myself in agreement with many of their points.

The levels of selection problem was central to Maynard Smith’s work throughout his career. This paper traces Maynard Smith’s views on the levels of selection, from his objections to group selection in the 1960s to his concern with the major evolutionary transitions in the 1990s. The relations between Maynard Smith’s position and those of Hamilton and G.C. Williams are explored, as is Maynard Smith’s dislike of the Price equation approach to multi-level selection. Maynard Smith’s account of the ‘core Darwinian principles’ (...) is discussed, as is his debate with Sober and Wilson (1998) over the status of trait-group models, and his attitude to the currently fashionable concept of pluralism about the levels of selection. (shrink)

My comments will focus on the second and third chapters of Sober’s book , which explore Darwin’s ideas about altruism, group selection and kin selection , and sex-ratio evolution . Sober makes a persuasive argument for his main claim: that Darwin was a subtler thinker on these topics than he is often taken to be. While there is much that I admire in Sober’s lucid discussion, I will focus on points of disagreement. Readers should note that this is not the (...) first time that Sober and I have disagreed on these issues .Sober begins his chapter on group selection with a brief history of the modern debate on the topic, which began in earnest in the 1960s. One of the key figures in this debate was George Williams, whose book Adaptation and Natural Selection helped convince many biologists that group selection was unlikely to be an important factor in evolution. Sober is critical of many of Williams’ arguments, though .. (shrink)

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)

In models of multi-level selection, the property of Darwinian fitness is attributed to entities at more than one level of the biological hierarchy, e.g. individuals and groups. However, the relation between individual and group fitness is a controversial matter. Theorists disagree about whether group fitness should always, or ever, be defined as total (or average) individual fitness. This paper tries to shed light on the issue by drawing on work in social choice theory, and pursuing an analogy between fitness and (...) utility. Social choice theorists have long been interested in the relation between individual and social utility, and have identified conditions under which social utility equals total (or average) individual utility. These ideas are used to shed light on the biological problem. (shrink)

“The Role of the Individual in Evolution” is a prescient yet neglected 1941 work by the 20th century’s most important paleontologist, George Gaylord Simpson. In a curious intermingling of explanation and critique, Simpson engages questions that would become increasingly fundamental in modern biological theory and philosophy. Did individuality, adaptation, and evolutionary causation reside at more than one level: the cell, the organism, the genetically coherent reproductive group, the social group, or some combination thereof? What was an individual, anyway? In this (...) introduction, we highlight two points in a wider historical context. First, recognizing the political context of Simpson’s writing profoundly deepens our understanding of the development of his science as the Modern Evolutionary Synthesis infused biology. Second, this story illuminates the emergence of debates around what would eventually come to be called multilevel selection theory. The organism-centered concept of biological individuality defended by Simpson is situated in relation to the then-emerging Synthesis, in which he was a renowned player, and also in relation to the views he opposed: the “metaphysical” ideas of paleontologists such as Henry Fairfield Osborn, who claimed that some evolutionary trends derived from potentialities already implanted in the germplasm; and the organicist ideas of Ralph W. Gerard and the Chicago School of ecologists, which he derided as all too congenial to totalitarianism. We find parallels between the ways that Simpson thought then about human individuality under totalitarianism and the way he thought about individuality in evolution; not that any causal relationship linked the two, but that commonalities of hierarchical structure exist between single entities and groups in both instances. We then trace the subsequent development of Simpson’s political and philosophical takes on the role of the individual in evolution through the 1960s and lightly sketch out the later fate of the organicist ideas of the Chicago School. Simpson’s paper, originally published in the Journal of the Washington Academy of Sciences, is available as supplementary material in the online version of this article, as part of the "Classics in Biological Theory" collection. (shrink)

Rudolf Leuckart's 1851 pamphlet Ueber den Polymorphismus der Individuen (On the polymorphism of individuals) stood at the heart of naturalists' discussions on biological individuals, parts and wholes in mid-nineteenth-century Britain and Europe. Our analysis, which accompanies the first translation of this pamphlet into English, situates Leuckart's contribution to these discussions in two ways. First, we present it as part of a complex conceptual knot involving not only individuality and the understanding of compound organisms, but also the alternation of generations, the (...) division of labor in nature, and the possibility of finding general laws of the organic world. Leuckart's pamphlet is important as a novel attempt to give order to the strands of this knot. It also solved a set of key biological problems in a way that avoided some of the drawbacks of an earlier teleological tradition. Second, we situate the pamphlet within a longer trajectory of inquiry into part-whole relations in biology from the mid-eighteenth century to the present. We argue that biological individuality, along with the problem-complexes with which it engaged, was as central a problem to naturalists before 1859 as evolution, and that Leuckart's contributions to it left a long legacy that persisted well into the twentieth century. As biologists' interests in part-whole relations are once again on the upswing, the longue durée of this problem merits renewed consideration. (shrink)

Rudolf Leuckart’s 1851 pamphlet Ueber den Polymorphismus der Individuen stood at the heart of naturalists’ discussions on biological individuals, parts and wholes in mid-nineteenth-century Britain and Europe. Our analysis, which accompanies the first translation of this pamphlet into English, situates Leuckart’s contribution to these discussions in two ways. First, we present it as part of a complex conceptual knot involving not only individuality and the understanding of compound organisms, but also the alternation of generations, the division of labor in nature, (...) and the possibility of finding general laws of the organic world. Leuckart’s pamphlet is important as a novel attempt to give order to the strands of this knot. It also solved a set of key biological problems in a way that avoided some of the drawbacks of an earlier teleological tradition. Second, we situate the pamphlet within a longer trajectory of inquiry into part-whole relations in biology from the mid-eighteenth century to the present. We argue that biological individuality, along with the problem-complexes with which it engaged, was as central a problem to naturalists before 1859 as evolution, and that Leuckart’s contributions to it left a long legacy that persisted well into the twentieth century. As biologists’ interests in part-whole relations are once again on the upswing, the longue durée of this problem merits renewed consideration. (shrink)

According to a once influential view of selection, it consists of repeated cycles of replication and interaction. It has been argued that this view is wrong: replication is not necessary for evolution by natural selection. I analyze the nine most influential arguments for this claim and defend the replication–interaction conception of selection against these objections. In order to do so, however, the replication–interaction conception of selection needs to be modified significantly. My proposal is that replication is not the copying of (...) an entity, the replicator, but the copying of a property. Thus, we can have a replication process without there being a replicator that is being copied. (shrink)

One of the most deeply entrenched ideas in Popper's philosophy is the analogy between the growth of scientific knowledge and the Darwinian mechanism of natural selection. Popper gave his first exposition of these ideas very early on. In a letter to Donald Campbell, 1 Popper says that the idea goes back at least to the early thirties. 2 And he had a fairly detailed account of it in his "What is dialectic?", a talk given in 1937 and published in 1940: (...) 3 If we want to explain why human thought tends to try out every conceivable solution for any problem with which it is faced, then we can appeal to a highly general sort of regularity. The method by which a solution is approached is .. (shrink)

The contextual approach is a prominent framework for thinking about group selection. Here, I highlight ambiguity about what the contextual approach is. Then, I discuss problematic entailments the contextual approach has for what processes count as group selection—entailments more troublesome than typically noted. However, Sober and Wilson’s version of the Price approach, which is the main alternative to the contextual approach, is problematic too: it leads to an underappreciated paradox called the vanishing selection problem and thereby generates the wrong qualitative (...) account of whether group selection is occurring in a certain family of cases. In response, I develop an account of group selection that can deal with the counterexamples to both the contextual approach and the Price approach. I then discuss the role that contextual analysis can continue to play in the discussion of individual fitness and metapopulation evolution. (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)

Even after Quine’s critique of the analytic-synthetic distinction in “Two Dogmas of Empiricism,” Wilfrid Sellars maintained some forms of analyticity or truth in virtue of meaning. This paper aims to reconstruct his neglected account of the analytic-synthetic distinction and the revisability of analytic sentences, its connection to his inferentialist account of meaning, and his response to Quine. While Sellars’s account of how analytic sentences can be revised bears certain similarities with Carnap’s and Grice and Strawson’s accounts, it is still striking (...) in that it is part of his broader picture of how our language develops dynamically in our ongoing inquiry. I aim to show how it relates within his broader picture to his account of how the diachronic continuity in meaning, language, or conceptual framework can be preserved through a revision of analytic sentences, and to his evolutionary account of the development of our language or conceptual framework. (shrink)