Even after Willard 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. In this article, I aim to reconstruct (a) his neglected account of the analytic-synthetic distinction and the revisability of analytic sentences, (b) its connection to his inferentialist account of meaning, and (c) his response to Quine. While Sellars’s account of the revisability of analytic sentences bears certain similarities to 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 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)

Since, it has become clear that individuality is not to be considered as a given, but rather as something which needs to be explained. How has individuality emerged through evolution, and how has it subsequently been maintained? In particular, why is it that multicellular organisms appeared and persisted, despite the obvious interest of each cell of favoring its own replication? Several biologists see the immune system as one of the key components for explaining the maintenance of multicellular organisms’ individuality. Indeed, (...) the immune system exerts a constant surveillance on all the constituents of the organism, including “cheaters” like tumor cells, which favor their own replication at the expense of the whole organism. In most cases, the immune system eliminates those cheaters. This is the “immune surveillance” hypothesis, first suggested by Burnet and Thomas. In this paper, I account for recent findings on immune surveillance in order to determine the precise role of the immune system in the emergence and maintenance of individuality. This investigation gives rise to a critical question for the domain of biological individuality: should immunity be considered as something unique to multicellular organisms? If it is indeed unique, should the multicellular organism be considered as an individual to a higher degree than any other living entity? If it is not unique, what precisely are the equivalents of the immune system in other living individuals? (shrink)

A crucial question for a process view of life is how to identify a process and how to follow it through time. The genidentity view can contribute decisively to this project. It says that the identity through time of an entity X is given by a well-identified series of continuous states of affairs. Genidentity helps address the problem of diachronic identity in the living world. This chapter describes the centrality of the concept of genidentity for David Hull and proposes an (...) extension of Hull’s view to the ubiquitous phenomenon of symbiosis. Finally, using immunology as a key example, it shows that the genidentity view suggests that the main interest of a process approach is epistemological rather than ontological and that its principal claim is one of priority, namely that processes precede and define things, and not vice versa. (shrink)

The traditional practice of establishing morphological types and investigating morphological organization has found new support from evolutionary developmental biology (evo-devo), especially with respect to the notion of body plans. Despite recurring claims that typology is at odds with evolutionary thinking, evo-devo offers mechanistic explanations of the evolutionary origin, transformation, and evolvability of morphological organization. In parallel, philosophers have developed non-essentialist conceptions of natural kinds that permit kinds to exhibit variation and undergo change. This not only facilitates a construal of species (...) and higher taxa as natural kinds, but also broadens our perspective on the diversity of kinds found in biology. There are many different natural kinds relevant to the investigative and explanatory aims of evo-devo, including homologues and developmental modules. (shrink)

Reduction and reductionism have been central philosophical topics in analytic philosophy of science for more than six decades. Together they encompass a diversity of issues from metaphysics and epistemology. This article provides an introduction to the topic that illuminates how contemporary epistemological discussions took their shape historically and limns the contours of concrete cases of reduction in specific natural sciences. The unity of science and the impulse to accomplish compositional reduction in accord with a layer-cake vision of the sciences, the (...) seminal contributions of Ernest Nagel on theory reduction and how they strongly conditioned subsequent philosophical discussions, and the detailed issues pertaining to different accounts of reduction that arise in both physical and biological science (e.g., limit-case and part-whole reduction in physics, the difference-making principle in genetics, and mechanisms in molecular biology) are explored. The conclusion argues that the epistemological heterogeneity and patchwork organization of the natural sciences encourages a pluralist stance about reduction. (shrink)

There is an active research program currently underway, which treats cancer progression as an evolutionary process. This contribution investigates the ways that cancer progression is like and unlike evolution in other contexts. The aim is to take a multi-level perspective on cancer, investigating the levels at which selection may be acting, the unit or target of selection, the relative roles of selection and drift, and the idea that cancer progression may be a by-product of selection at other levels of organization.

The theory of evolution settled at what was thought to be its definitive form after the affiliation of Darwin’s theory with the new science of genetics. This historical event explains not only the success but also the vulnerability of evolutionary theory. The close affinity with genetics helped to provide the tools required for managing phylogenetic evolution, which was controlled by the molecular machinery of the genome, localized in most cells of each individual. This setup worked well for organizing the basics (...) of Darwinism but it faltered when applied to the evolution of culture. This is so because, as I argue in this article, under the given conditions the active support of a system of similar complexity as the genome would be required. At some point in the process of evolution the large brains of primates may have begun experimenting with phenotypic equivalents of genomic functions like memory, adaptation, fitness, processing information, and organizing replication. In this way the brain may have weakened its ties to the genome and by exploiting its new status have become, so to speak, for the phenotype what the genome is for the genotype, thus preparing the stage on which the complexity of human behavior might unfold. (shrink)

Evolutionary biology is a field currently animated by much discussion concerning its conceptual foundations. On the one hand, we have supporters of a classical view of evolutionary theory, whose backbone is provided by population genetics and the so-called Modern Synthesis (MS). On the other hand, a number of researchers are calling for an Extended Synthe- sis (ES) that takes seriously both the limitations of the MS (such as its inability to incorporate developmental biology) and recent empirical and theoretical research on (...) issues such as evolvability, modularity, and self-organization. In this article, I engage in an in-depth commentary of an influential paper by population geneticist Michael Lynch, which I take to be the best defense of the MS-population genetics position published so far. I show why I think that Lynch’s arguments are wanting and propose a modification of evolutionary theory that retains but greatly expands on population genetics. (shrink)

Debates concerning the units and levels of selection have persisted for over fifty years. One major question in this literature is whether units and levels of selection are genuine, in the sense that they are objective features of the world, or merely reflect the interests and goals of an observer. Scientists and philosophers have proposed a range of answers to this question. This Element introduces this literature and proposes a novel contribution. It defends a realist stance and offers a way (...) of delineating genuine levels of selection by invoking the notion of a functional unit. (shrink)

I extend work from Krakauer et al. (2020), who propose a conception of individuality as the capacity to propagate information through time. From this conception, they develop information-theoretic measures. I identify several shortcomings with these measures—in particular, that they are associative rather than causal. I rectify this shortcoming by deriving a causal information-theoretic measure of individuality. I then illustrate how this measure can be implemented and extended in the context of evolutionary transitions in individuality.

Embodied and extended cognition is a relatively new paradigm within cognitive science that challenges the basic tenet of classical cognitive science, viz. cognition consists in building and manipulating internal representations. Some of the pioneers of embodied cognitive science have claimed that this new way of conceptualizing cognition puts pressure on epistemological and ontological realism. In this paper I will argue that such anti-realist conclusions do not follow from the basic assumptions of radical embodied cognitive science. Furthermore I will show that (...) one can develop a form of realism that reflects rather than just accommodates the core principles of non-representationalist embodied cognitive science. (shrink)

While the existence of human cognitive and behavioral diversity is now widely recognized, it is not yet well established how to explain this diversity. In particular, it is still unclear how to determine whether any given instance of human cognitive and behavioral diversity is due to a common psychology that is merely “triggered” differently in different bio-cultural environments, or whether it is due to deeply and fundamentally different psychologies. This paper suggests that, to answer this question, we need to employ (...) subtle theoretical considerations of theory choice—especially the consideration of the complexity-weighted differential predictive successes of the two accounts. To make this clearer, the paper develops a novel analysis of the observed differences in human sharing dispositions. (shrink)

While the existence of human cognitive and behavioral diversity is now widely recognized, it is not yet well established how to explain this diversity. In particular, it is still unclear how to determine whether any given instance of human cognitive and behavioral diversity is due to a common psychology that is merely “triggered” differently in different bio-cultural environments, or whether it is due to deeply and fundamentally different psychologies. This paper suggests that, to answer this question, we need to employ (...) subtle theoretical considerations of theory choice—especially the consideration of the complexity-weighted differential predictive successes of the two accounts. To make this clearer, the paper develops a novel analysis of the observed differences in human sharing dispositions. (shrink)

Biological market theory has in recent years become an important part of the social evolutionist’s toolkit. This article discusses the explanatory potential and pitfalls of biological market theory in the context of big picture accounts of the evolution of human cooperation and morality. I begin by assessing an influential account that presents biological market dynamics as a key driver of the evolution of fairness norms in humans. I argue that this account is problematic for theoretical, empirical, and conceptual reasons. After (...) mapping the evidential and explanatory limits of biological market theory, I suggest that it can nevertheless fill a lacuna in an alternative account of hominin evolution. Trade on a biological marketplace can help explain why norm-based cooperation did not break down when our late-Pleistocene ancestors entered new, challenging social and economic environments. (shrink)

Schaffner’s model of theory reduction has played an important role in philosophy of science and philosophy of biology. Here, the model is found to be problematic because of an internal tension. Indeed, standard antireductionist external criticisms concerning reduction functions and laws in biology do not provide a full picture of the limits of Schaffner’s model. However, despite the internal tension, his model usefully highlights the importance of regulative ideals associated with the search for derivational, and embedding, deductive relations among mathematical (...) structures in theoretical biology. A reconstructed Schaffnerian model could therefore shed light on mathematical theory development in the biological sciences and on the epistemology of mathematical practices more generally. *Received November 2006; revised March 2009. †To contact the author, please write to: Philosophy Department, University of California, Santa Cruz, 1156 High St., Santa Cruz, CA 95064; e‐mail: [email protected]. (shrink)

Two controversies exist regarding the appropriate characterization of hierarchical and adaptive evolution in natural populations. In biology, there is the Wright-Fisher controversy over the relative roles of random genetic drift, natural selection, population structure, and interdemic selection in adaptive evolution begun by Sewall Wright and Ronald Aylmer Fisher. There is also the Units of Selection debate, spanning both the biological and the philosophical literature and including the impassioned group-selection debate. Why do these two discourses exist separately, and interact relatively little? (...) We postulate that the reason for this schism can be found in the differing focus of each controversy, a deep difference itself determined by distinct general styles of scientific research guiding each discourse. That is, the Wright-Fisher debate focuses on adaptive process, and tends to be instructed by the mathematical modeling style, while the focus of the Units of Selection controversy is adaptive product, and is typically guided by the function style. The differences between the two discourses can be usefully tracked by examining their interpretations of two contested strategies for theorizing hierarchical selection: horizontal and vertical averaging. (shrink)

The dangers of character reification for cladistic inference are explored. The identification and analysis of characters always involves theory-laden abstraction—there is no theory-free “view from nowhere.” Given theory-ladenness, and given a real world with actual objects and processes, how can we separate robustly real biological characters from uncritically reified characters? One way to avoid reification is through the employment of objectivity criteria that give us good methods for identifying robust primary homology statements. I identify six such criteria and explore each (...) with examples. Ultimately, it is important to minimize character reification, because poor character analysis leads to dismal cladograms, even when proper phylogenetic analysis is employed. Given the deep and systemic problems associated with character reification, it is ironic that philosophers have focused almost entirely on phylogenetic analysis and neglected character analysis. (shrink)

In a recent paper, Brandon and Nijhout argue against genic selectionism—the thesis, roughly, that evolutionary processes are best understood from the gene’s-eye point of view—by presenting a case in which genic models of selection allegedly make predictions that conflict with the (correct) predictions of higher-level genotypic selection models. Their argument, if successful, would refute the widely held belief that genic models and higher-level models are predictively equivalent. Here, I argue that Brandon and Nijhout fail to demonstrate that the models make (...) incompatible predictions. (shrink)

Okasha claims at the outset of his book "Evolution and the Levels of Selection" that the Price equation lays bare the fundamentals underlying all selection phenomena. However, the thoroughness of his subsequent analysis of multi-level selection theories leads him to abandon his fundamentalist commitments. At critical points he invokes cost benefit analyses that sometimes favors the Price approach and sometimes the contextual approach, sometimes favors MLS1 and sometimes MLS2. And although he doesn’t acknowledge it, even the Price approach breaks down (...) into a family of alternative equations that parse the causes in different ways, none of which is uniquely correct and none of which achieves the ultimate isolation of effects due to what Okasha believes are the fundamental causes. I argue that his book provides good reason to re-conceive our understanding of evolutionary theorizing in terms of a toolbox view and to stop subjecting the analyses of evolutionary concepts to a universalist standard. (shrink)

On the basis of findings from developmental biology, some researchers have argued that evolutionary theory needs to be significantly updated. Advocates of such a “developmental update” have, among other things, suggested that we need to re-conceptualize units of selection, that we should expand our view of inheritance to include environmental as well as genetic and epigenetic factors, that we should think of organisms and their environment as involved in reciprocal causation, and that we should reevaluate the rates of evolutionary change. (...) However, many of these same conclusions could be reached on the basis of other evidence, namely from microbiology. In this paper, I ask why microbiological evidence has not had a similarly large influence on calls to update biological theory, and argue that there is no principled reason to focus on developmental as opposed to microbiological evidence in support of these revisions to evolutionary theory. I suggest that the focus on developmental biology is more likely attributable to historical accident. I will also discuss some possible room for overlap between developmental and microbiology, despite the historical separation of these two subdisciplines. (shrink)

Many models of evolution are implicitly causal processes. Features such as causal feedback between evolutionary variables and evolutionary processes acting at multiple levels, though, mean that conventional causal models miss important phenomena. We develop here a general theoretical framework for analyzing evolutionary processes drawing on recent approaches to causal modeling developed in the machine-learning literature, which have extended Pearls do-calculus to incorporate cyclic causal interactions and multilevel causation. We also develop information-theoretic notions necessary to analyze causal information dynamics in our (...) framework, introducing a causal generalization of the Partial Information Decomposition framework. We show how our causal framework helps to clarify conceptual issues in the contexts of complex trait analysis and cancer genetics, including assigning variation in an observed trait to genetic, epigenetic and environmental sources in the presence of epigenetic and environmental feedback processes, and variation in fitness to mutation processes in cancer using a multilevel causal model respectively, as well as relating causally-induced to observed variation in these variables via information theoretic bounds. In the process, we introduce a general class of multilevel causal evolutionary processes which connect evolutionary processes at multiple levels via coarse-graining relationships. Further, we show how a range of fitness models can be formulated in our framework, as well as a causal analog of Prices equation, clarifying the relationships between realized/probabilistic fitness and direct/indirect selection. Finally, we consider the potential relevance of our framework to foundational issues in biology and evolution, including supervenience, multilevel selection and individuality. Particularly, we argue that our class of multilevel causal evolutionary processes, in conjunction with a minimum description length principle, provides a conceptual framework in which identification of multiple levels of selection may be reduced to a model selection problem. (shrink)

A notion of delegated causality is introduced here. This subtle kind of causality is dual to interventional causality. Delegated causality elucidates the causal role of dynamical systems at the “edge of chaos”, explicates evident cases of downward causation, and relates emergent phenomena to Gödel’s incompleteness theorem. Apparently rich implications are noticed in biology and Chinese philosophy. The perspective of delegated causality supports cognitive interpretations of self-organization and evolution.

This essay offers some reflections on Samir Okasha’s new monograph Agents and Goals in Evolution, his style of doing philosophy, and the broader philosophy of nature project of trying to make sense of agency and rationality as natural phenomena.

This paper introduces the reconstitutor as a comprehensive unit of heredity within the context of evolutionary research. A reconstitutor is the structure resulting from a set of relationships between different elements or processes that are actively involved in the recreation of a specific phenotypic variant in each generation regardless of the biomolecular basis of the elements or whether they stand in a continuous line of ancestry. Firstly, we justify the necessity of introducing the reconstitutor by showing the limitations of other (...) evolutionary conceptions of the unit of heredity, such as the replicator, the reproducer, and the Darwinian individual. We argue that these conceptions are based on the requirement of lineage formation, which we argue to be unnecessary for the existence of evolutionary heredity. In the second part, we introduce the reconstitutor, which we base on the concept of Stability of Traits, and illustrate how it covers cases of hereditary phenomena not covered by the previous accounts. Secondly, we illustrate how the reconstitutor could serve as a platform to rethink ecological inheritance and other forms of inheritance that have been recently introduced under the song/singer model of evolution. (shrink)

Darwin provided us with a powerful theoretical framework to explain the evolution of living systems. Natural selection alone, however, has sometimes been seen as insufficient to explain the emergence of new levels of selection. The problem is one of “circularity” for evolutionary explanations: how to explain the origins of Darwinian properties without already invoking their presence at the level they emerge. That is, how does evolution by natural selection commence in the first place? Recent results in experimental evolution suggest a (...) way forward: Paul Rainey and his collaborators have shown that Darwinian properties could be exogenously imposed via what they call “ecological scaffolding.” This could solve the “black box” dilemma faced by Darwinian explanations of new levels of organization. Yet, despite “scaffolding” recently becoming a popular term in the study of cognition, culture, and evolution, the concept has suffered from vagueness and ambiguity. This article aims to show that scaffolding can be turned into a proper scientific concept able to do explanatory work within the context of the major evolutionary transitions. Doing so will allow us to once again extend the scope of the Darwinian model of evolution by natural selection. (shrink)

For decades Darwinian processes were framed in the form of the Lewontin conditions: reproduction, variation and differential reproductive success were taken to be sufficient and necessary. Since Buss and the work of Maynard Smith and Szathmary biologists were eager to explain the major transitions from individuals to groups forming new individuals subject to Darwinian mechanisms themselves. Explanations that seek to explain the emergence of a new level of selection, however, cannot employ properties that would already have to exist on that (...) level for selection to take place. Recently, Hammerschmidt et al. provided a ‘bottom-up’ experiment corroborating much of the theoretical work Paul Rainey has done since 2003 on how cheats can play an important role in the emergence of new Darwinian individuals on a multicellular level. The aims of this paper are twofold. First, I argue for a conceptual shift in perspective from seeing cheats as a ‘problem’ that needs to be solved for multi-cellularity to evolve to the very ‘key’ for the evolution of multicellularity. Secondly, I illustrate the consequences of this shift for both theoretical and experimental work, arguing for a more prominent role of ecology and the multi-level selection framework within the debate then they currently occupy. (shrink)

In his 2009 monograph, Darwinian Populations and Natural Selection, Peter Godfrey-Smith accuses biologists of demonstrating ‘Darwinian Paranoia’ when they engage in what he dubs ‘agential thinking’. But as Daniel Dennett points out, he offers neither an illuminating set of examples nor an extended argument for this assertion, deeming it to be a brilliant propaganda stroke against what is actually a useful way of thinking. Compared to the dangers of teleological thinking in biology, the dangers of agential thinking have unfortunately rarely (...) been discussed. Drawing on recent work by Samir Okasha, I attempt to remedy this omission, through analyzing the nature of agential thinking, and providing a philosophical treatment of the unexamined dangers in this peculiar, yet tempting way of thinking. (shrink)

Beatrice Han argues that the theories of subjection (determinism: structure) and subjectivation (freedom: agency) are the “the blind spot of Foucault's work:” to the very end of his life, in being transcendental and historical theories, respectively, they were in irresolvable conflict. In part I, I have argued that Foucault encourages us to situate the theories of the subject in an un-thematized reach for a metaphysics of realism which, in effect, was to ground his uncertain complementary reach for a naturalist conduct (...) of research. In part II I also argue that it is this fundamental feature of Foucault's Foucault that drives his returns to Kant, the purpose of which is to resolve the conflicting theories of the subject, and thus to solve his Giddensian problem of structure and creativity. Locating the returns and their purpose in my context of Science for Humanism and the recovery of human agency, I ultimately argue that Foucault's two special returns to Kant in order to solve his structure/agency problem led to two unfortunate solutions. The resort to Baudelair's aesthetic subject is a failed solution in so far as it regresses to being a pre-noumenal conception of the subject. The subsequent mere reinstatement of Kant's subject as causally empowered, minus the noumenalism, is nothing more than a reclamation of Kant's conception. Only the reconstruction of Foucault's realism permits us to assert that he could have moved beyond the reclamation of human agency to its recovery. (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)

This paper offers a paleobiological perspective on the debate concerning the possible use of biotechnology to bring back extinct species. One lesson from paleobiology is that extinction selectivity matters in addition to extinction rates and extinction magnitude. Combining some of Darwin’s insights about artificial selection with the theory of species selection that paleobiologists developed in the 1970s and 1980s provides a useful context for thinking about de-extinction. Using recent work on the prioritization of candidate species for de-extinction as a test (...) case, the paper argues that de-extinction would be a form of artificial species selection in which humans influence which species persist vs. go extinct. This points to a serious gap in our ethical theory: Much work has been done to clarify the value of biological diversity, but we also need theoretical guidance for decisions that amount to species sorting, and that will shape the macroevolutionary future. (shrink)

This paper takes a close look at the role of behavior in the “major transitions” in evolution—events during which inheritance and development, and therefore the process of adaptation by natural selection, are reorganized at a new level of compositional hierarchy—and at the requirements for sufficiently explaining these important events in the history of life. I argue that behavior played a crucial role in driving at least some of the major transitions. Because behavioral interactions can become stably organized in novel ways (...) on timescales faster than the lifetime of an organism, behavior can lead the way into a transition—becoming organized at the new level prior to inheritance and development. It is widely acknowledged that behavioral plasticity can play an important role in evolution; environmental novelty can elicit novel behavior that may feed back on the evolutionary process through niche selection or cultural inheritance, for example (Jablonka and Lamb 2005). I argue here that not just novel behaviors but novel forms of behavioral organization (distributed or hierarchical control that produces functional coherence) can emerge, binding the evolutionary fates of a group of organisms which were previously independent in terms of behavior as well as reproduction, and leading the way into a transition to an aggregative or “higher-level” mode of reproduction. (shrink)

Contemporary biological research has suggested that some host–microbiome multispecies systems (referred to as “holobionts”) can in certain circumstances evolve as unique biological individual, thus being a unit of selection in evolution. If this is so, then it is arguably the case that some biological adaptations have evolved at the level of the multispecies system, what we call hologenomic adaptations. However, no research has yet been devoted to investigating their nature, or how these adaptations can be distinguished from adaptations at the (...) species-level (genomic adaptations). In this paper, we cover this gap by investigating the nature of hologenomic adaptations. By drawing on the case of the evolution of sanguivory diet in vampire bats, we argue that a trait constitutes a hologenomic adaptation when its evolution can only be explained if the holobiont is considered the biological individual that manifests this adaptation, while the bacterial taxa that bear the trait are only opportunistic beneficiaries of it. We then use the philosophical notions of emergence and inter-identity to explain the nature of this form of individuality and argue why it is special of holobionts. Overall, our paper illustrates how the use of philosophical concepts can illuminate scientific discussions, in the trend of what has recently been called metaphysics of biology. (shrink)

Bourrat and Griffiths :33, 2018) have recently argued that most of the evidence presented by holobiont defenders to support the thesis that holobionts are evolutionary individuals is not to the point and is not even adequate to discriminate multispecies evolutionary individuals from other multispecies assemblages that would not be considered evolutionary individuals by most holobiont defenders. They further argue that an adequate criterion to distinguish the two categories is fitness alignment, presenting the notion of fitness boundedness as a criterion that (...) allows divorcing true multispecies evolutionary individuals from other multispecies assemblages and provides an adequate criterion to single out genuine evolutionary multispecies assemblages. A consequence of their criterion is that holobionts, as conventionally defined by hologenome defenders, are not evolutionary individuals except in very rare cases, and for very specific host-symbiont associations. This paper is a critical response to Bourrat and Griffiths’ arguments and a defence of the arguments presented by holobiont defenders. Drawing upon the case of the hologenomic basis of the evolution of sanguivory in vampire bats, I argue that Bourrat and Griffiths overlook some aspects of the biological nature of the microbiome that justifies the thesis that holobionts are evolutionarily different to other multispecies assemblages. I argue that the hologenome theory of evolution should not define the hologenome as a collection of genomes, but as the sum of the host genome plus some traits of the microbiome which together constitute an evolutionary individual, a conception I refer to as the stability of traits conception of the hologenome. Based on that conception I argue that the evidence presented by holobiont defenders is to the point, and supports the thesis that holobionts are evolutionary individuals. In this sense, the paper offers an account of the holobiont that aims to foster a dialogue between hologenome advocates and hologenome critics. (shrink)

If there is a single discipline of science calling the basic concepts of biology into question, it is without doubt microbiology. Indeed, developments in microbiology have recently forced us to rethink such fundamental concepts as the organism, individual, and genome. In this paper I show how microorganisms are changing our understanding of natural aggregations and develop the concept of a Darwinian population to embrace these discoveries. I start by showing that it is hard to set the boundaries of a Darwinian (...) population, and I suggest thinking of a Darwinian population as a relative property of a Darwinian individual. Then I argue, in contrast to the commonly held view, that Darwinian populations are multispecies units, and that in order to accept the multispecies account of Darwinian populations we have to separate fitness from natural selection. Finally, I show how all these ideas provide a theoretical framework leading to a more precise understanding of the ecology of endosymbiosis than is afforded by poetic metaphors such as ‘slavery’. (shrink)

What are the prospects for a monistic view of biological individuality given the multiple epistemic roles the concept must satisfy? In this paper, I examine the epistemic adequacy of two recent accounts based on the capacity to undergo natural selection. One is from Ellen Clarke, and the other is by Peter Godfrey-Smith. Clarke’s position reflects a strong monism, in that she aims to characterize individuality in purely functional terms and refrains from privileging any specific material properties as important in their (...) own right. I argue that Clarke’s functionalism impairs the epistemic adequacy of her account compared to a middle-ground position taken by Godfrey-Smith. In comparing Clarke and Godfrey-Smith’s account, two pathways emerge to pluralism about biological individuality. The first develops from the contrast between functionalist and materialist approaches, and the second from an underlying temporal structure involved in using evolutionary processes to define individuality. (shrink)

I defend a radical interpretation of biological populations—what I call population pluralism—which holds that there are many ways that a particular grouping of individuals can be related such that the grouping satisfies the conditions necessary for those individuals to evolve together. More constraining accounts of biological populations face empirical counter-examples and conceptual difficulties. One of the most intuitive and frequently employed conditions, causal connectivity—itself beset with numerous difficulties—is best construed by considering the relevant causal relations as ‘thick’ causal concepts. I (...) argue that the fine-grained causal relations that could constitute membership in a biological population are huge in number and many are manifested by degree, and thus we can construe population membership as being defined by massively multidimensional constructs, the differences between which are largely arbitrary. I end by showing that positions in two recent debates in theoretical biology depend on a view of biological populations at odds with the pluralism defended here. (shrink)

An understanding of the factors behind the evolution of multicellularity is one of today’s frontiers in evolutionary biology. This is because multicellular organisms are made of one subset of cells with the capacity to transmit genes to the next generation and another subset responsible for maintaining the functionality of the organism, but incapable of transmitting genes to the next generation. The question arises: why do somatic cells sacrifice their lives for the sake of germline cells? How is germ/soma separation maintained? (...) One conventional answer refers to inclusive fitness theory, according to which somatic cells sacrifice themselves altruistically, because in so doing they enhance the transmission of their genes by virtue of their genetic relatedness to germline cells. In the present article we will argue that this explanation ignores the key role of policing mechanisms in maintaining the germ/soma divide. Based on the pervasiveness of the latter, we argue that the role of altruistic mechanisms in the evolution of multicellularity is limited and that our understanding of this evolution must be enriched through the consideration of coercion mechanisms. (shrink)

In this article I develop a big picture of the evolution of human cooperation, and contrast it to an alternative based on group selection. The crucial claim is that hominin history has seen two major transitions in cooperation, and hence poses two deep puzzles about the origins and stability of cooperation. The first is the transition from great ape social lives to the lives of Pleistocene cooperative foragers; the second is the stability of the social contract through the early Holocene (...) transition to complex hierarchical societies. The first of these transitions is driven, at least initially, by individual advantage: cooperation paid off for individual foragers, initially through mutualist interaction, then through reciprocation. This argument leads to a reanalysis of the role of violence and the nature of the freeriding threat to cooperation. But the conditions that select for cooperative individuals in the Pleistocene were eroded in the Pleistocene–Holocene transition. So we need an alternative account of the survival, and indeed the expansion, of cooperation in the Holocene. Group selection driven by intercommunal conflict really may well be central to this second transition. (shrink)

Here I reply to Jean Gayon's, Tim Lewens's, and Samir Okasha's comments on Did Darwin write the Origin backwards? The topics addressed include: Darwin's thinking that common ancestry is "evidentially prior" to natural selection; how Darwin uses phylogenetic trees to test hypotheses concerning natural selection; how group and indivdiual selection should be defined, and how each is related to the concept of adaptation.

I discuss two subjects in Samir Okasha’s excellent book, Evolution and the Levels of Selection. In consonance with Okasha’s critique of the conventionalist view of the units of selection problem, I argue that conventionalists have not attended to what realists mean by group, individual, and genic selection. In connection with Okasha’s discussion of the Price equation and contextual analysis, I discuss whether the existence of these two quantitative frameworks is a challenge to realism.