The philosophy of biology is witnessing an increasing enthusiasm for what can be called “individuals thinking”. Individuals thinking is a perspective on the metaphysics of biological entities according to which conceiving of them as individuals rather than kinds enables us to expose ongoing metaphysical debates as focusing on the wrong question, and to achieve better accounts of the metaphysics of biological entities. In this paper, I examine two cases of individuals thinking, the claim that species are individuals and the claim (...) that life on Earth is an individual. I argue that these claims fail to do the metaphysical work that one would want them to do. I highlight problems with the specific claims as well as with the general notion of ‘individual’, and argue that naturalistic metaphysicians of biology should think of the metaphysical status of theoretical entities, such as species and life, as fundamentally theory-dependent. This implies a metaphysical pluralism, that allows that in some theories species, life, and other such entities may feature as individuals, whereas in others they may feature as kinds. (shrink)

Fitness is a central concept in evolutionary theory. Just as it is central to biological evolution, so, it seems, it should be central to cultural evolutionary theory. But importing the biological fitness concept to CET is no straightforward task—there are many features unique to cultural evolution that make this difficult. This has led some theorists to argue that there are fundamental problems with cultural fitness that render it hopelessly confused. In this essay, we defend the coherency of cultural fitness against (...) those who call it into doubt. (shrink)

We argue that the economy of nature constitutes an invocation of structure in the biological sciences, one largely missed by philosophers of biology despite the turn in recent years toward structural explanations throughout the philosophy of science. We trace a portion of the history of this concept, beginning with the theologically and economically grounded work of Linnaeus, moving through Darwin’s adaptation of the economy of nature and its reconstitution in genetic terms during the first decades of the Modern Synthesis. What (...) this historical case study reveals, we argue, is a window into the shifting landscape of the explanatory and ontic uses of structural concepts. In Linnaeus, the economy of nature has both ontic and explanatory import; in Darwin the ontic and explanatory aspects start to come apart ; and finally, in the Modern Synthesis, the economy of nature is replaced by the conceptual toolkit of population genetics, the structural elements of which are nearly entirely explanatory. Having traced a historical trajectory of structural concepts that moves from an ontic formulation to an increasingly explanatory one, we conclude by outlining some insights for structural realism. (shrink)

Fitness is a central theoretical concept in evolutionary theory. Despite its importance, much debate has occurred over how to conceptualize and formalize fitness. One point of debate concerns the roles of organismic and trait fitness. In a recent addition to this debate, Elliott Sober argues that trait fitness is the central fitness concept, and that organismic fitness is of little value. In this paper, by contrast, we argue that it is organismic fitness that lies at the bases of both the (...) conceptual role of fitness, as well as its role as a measure of evolutionary dynamics. (shrink)

The biophilosophic justification for the idea that “brain death” is death needs to support two claims: that what dies in human death is a human organism, not merely a psychological entity distinct from it; that total brain failure signifies the end of the human organism as a whole. Defenders of brain death typically assume without argument that the first claim is true and argue for the second by defending the “integrative unity” rationale. Yet the integrative unity rationale has fallen on (...) hard times. In this article, I give reasons for why we should think of ourselves as organisms, and why the “fundamental work” rationale put forward by the 2008 President’s Council is better than the integrative unity rationale, despite persistent objections to it. (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)

Both Aristotle and Ernst Mayr present theories of dual explanation in biology, with proximal, clearly physical explanations and more distal, biology-specific explanations. Aristotle’s presentation of final cause explanations in Posterior Analytics relates final causes to the necessary material, formal, and efficient causes that mediate them. Johnson and Leunissen demonstrate the problematic nature of historical and recent interpretations and open the door for a new interpretation consistent with modern evolutionary theory. Mayr’s differentiation of proximate and ultimate/evolutionary causes provides a key to (...) appropriating Aristotle for modern use, if care is taken to differentiate Mayr’s proximate/distal and mechanical/evolutionary distinctions from his intentional/non-intentional distinction. Building on Mayr, and noting his appropriation of ancient and recent commentary, a strong case can be made for nested explanation, wherein evolutionary explanations are instantiated in systems of mechanical explanations. Biological concepts of organism, gene, function, etc., are etiologically prior to mechanical explanations of function, but temporally posterior to mechanical explanations of historical adaptation. Such an analysis sheds light on recent arguments within evolutionary biology while highlighting the importance of epistemology in contemporary science. (shrink)

How many individuals are present where we see a pregnant individual? Within a substance ontological framework, there are exactly two possible answers to this question. The standard answer—two individuals—is typically championed by scholars endorsing the predominant Containment View of pregnancy, according to which the foetus resides in the gestating organism like in a container. The alternative answer—one individual—has recently found support in the Parthood View, according to which the foetus is a part of the gestating organism. Here I propose a (...) third answer: a pregnant individual is neither two individuals nor one individual but something in between one and two. This is because organisms are better understood as processes than as substances. With a special focus on the Parthood View, I explain why a Process View of pregnancy, according to which a pregnant individual is a bifurcating hypercomplex process, surpasses the substance ontological approaches. (shrink)

Our understanding of the universe has grown rapidly in recent decades. We’ve discovered evidence of water in nearby planets, discovered planets outside our solar system, mapped the genomes of thousands of organisms, and probed the very origins and limits of life. The scientific perspective of life-as-it-could-be has expanded in part by research in astrobiology, synthetic biology, and artificial life. In the face of such scientific developments, we argue there is an ever-growing need for universal biology, life-as-it-must-be, the multidisciplinary study of (...) non-contingent aspects of life as guided by biological theory and constrained by the universe. We present three distinct but connected ways of universalizing biology—with respect to characterizing aspects of life everywhere, with respect to the explanatory scope of biological theory, and with respect to extending biological insights to the structure of nonbiological entities. For each of these, we sketch the theoretical goals and challenges, as well as give examples of current research that might be labeled universal biology. (shrink)

The concept of superorganism has a mixed reputation in biology—for some it is a convenient way of discussing supra-organismal levels of organization, and for others, little more than a poetic metaphor. Here, I show that a considerable step forward in the understanding of superorganisms results from a thorough review of the supra-organismal levels of organization now known to exist among the “unicellular” protists. Limiting the discussion to protists has enormous advantages: their bodies are very well studied and relatively simple, and (...) they exhibit an enormous diversity of anatomies and lifestyles. This allows for unprecedented resolution in describing forms of supra-organismal organization. Here, four criteria are used to differentiate loose, incidental associations of hosts with their microbiota from “actual” superorganisms: obligatory character, specific spatial localization of microbiota, presence of attachment structures and signs of co-evolution in phylogenetic analyses. Three groups—that have never before been described in the philosophical literature—merit special attention: Symbiontida, Oxymonadida and Parabasalia. Specifically, it is argued that in certain cases—for Bihospites bacati and Calkinsia aureus, Streblomastix strix, Joenia annectens and Mixotricha paradoxa and Kentrophoros —it is fully appropriate to describe the whole protist-microbiota assocation as a single organism and its elements as “tissues” or, arguably, even “organs”. To account for this level of biological complexity, I propose the term “structured superorganism”. (shrink)

Biological individuals are usually defined by evolutionists through a reference to natural selection. This article looks for a concept of individuality that would hold at the same time for organisms and for communities or ecosystems, the latter being unaffected by natural selection. In the wake of Simon’s notion of “quasi-independence,” I elaborate a concept of “weak individuality” defined by probabilistic connections between sub-entities, read off our knowledge of their interactions. This formal scheme of connections allows one to infer what are (...) the individuals in the domain addressed by the theory of the interactions. The article argues that if ecosystems do not have strong individuality, they still possess a weak individuality, ecological theories providing the values of the variables in the formula for individuality. (shrink)

Several advocates of the lively field of “metaphysics of science” have recently argued that a naturalistic metaphysics should be based solely on current science, and that it should replace more traditional, intuition-based, forms of metaphysics. The aim of the present paper is to assess that claim by examining the relations between metaphysics of science and general metaphysics. We show that the current metaphysical battlefield is richer and more complex than a simple dichotomy between “metaphysics of science” and “traditional metaphysics”, and (...) that it should instead be understood as a three dimensional “box”, with one axis distinguishing “descriptive metaphysics” from “revisionary metaphysics”, a second axis distinguishing a priori from a posteriori metaphysics, and a third axis distinguishing “commonsense metaphysics”, “traditional metaphysics” and “metaphysics of science”. We use this three-dimensional figure to shed light on the project of current metaphysics of science, and to demonstrate that, in many instances, the target of that project is not defined with enough precision and clarity. (shrink)

Canguilhem criticized the concept of “public health”: health and disease are concepts that only apply to individuals, taken as organic totalities. Their extension to a different level of organization is purely metaphorical. The importance assumed by epidemiology in the construction of our knowledge of the normal and the pathological does, however, call for reflection on the role and the status of the population level of organization in our approach to health phenomena. The entanglement of the biological and the social in (...) human life and in contemporary societies justifies this level of analysis for better understanding the complexity and the interaction of health determinants both at the level of individuals and their interactions and at that of the population. But is this population level just a useful level of analysis that makes it possible to bring to light the social determinants of health at the individual level, or does it rest instead on characteristics of the population that are irreducible to individual characteristics, but which are nevertheless important for understanding and taking action with respect to both population and individual health? Defending this second alternative, I show how the epidemiological point of view, and in particular that of social epidemiology, leads us to rethink the possibility of a concept of “population health” that is not reducible to the sum of individual instances of health. (shrink)

Given immunity’s general role in the organism’s economy—both in terms of its internal environment as well as mediating its external relations—immune theory has expanded its traditional formulation of preserving individual autonomy to one that includes accounting for nutritional processes and symbiotic relationships that require immune tolerance. When such a full ecological alignment is adopted, the immune system becomes the mediator of both defensive and assimilative environmental intercourse, where a balance of immune rejection and tolerance governs the complex interactions of the (...) organism’s ecological relationships. Accordingly, immunology, which historically had affiliated with the biology of individuals, now becomes a science concerned with the biology of communities. With this translocation, the ontological basis of the organism is undergoing a profound change. Indeed, the recent recognition of the ubiquity of symbiosis has challenged the traditional notions of biological individuality and requires a shift in the metaphysics undergirding biology, in which a philosophy of the organism must be characterized by ecological dialectics “all-the-way-down.”. (shrink)

This paper addresses the question of what organisms are and therefore what kinds of biological entities qualify as organisms. For some time now, the concept of organismality has been eclipsed by the notion of individuality. Biological individuals are those systems that are units of selection. I develop a conception of organismality that does not rely on evolutionary considerations, but instead draws on development and ecology. On this account, organismality and individuality can come apart. Organisms, in my view, are as Godfrey-Smith (...) puts it “essentially persisters.” I argue that persistence is underpinned by differentiation, integration, development, and the constitutive embeddedness of organisms in their worlds. I examine two marginal cases, the Portuguese Man O’ War and the honey bee colony, and show that both count as organisms in light of my analysis. Next, I examine the case of holobionts, hosts plus their microsymbionts, and argue that they can be counted as organisms even though they may not be biological individuals. Finally, I consider the question of whether other, less tightly integrated biological systems might also be treated as organisms. (shrink)

Organismal death is foundational to the evolution of life, and many biological concepts such as natural selection and life history strategy are so fashioned only because individuals are mortal. Organisms, irrespective of their organization, are composed of basic functional units—cells—and it is our understanding of cell death that lies at the heart of most general explanatory frameworks for organismal mortality. Cell death can be exogenous, arising from transmissible diseases, predation, or other misfortunes, but there are also endogenous forms of death (...) that are sometimes the result of adaptive evolution. These endogenous forms of death—often labeled programmed cell death, PCD—originated in the earliest cells and are maintained across the tree of life. Here, we consider two problematic issues related to PCD (and cell mortality generally). First, we trace the original discoveries of cell death from the nineteenth century and place current conceptions of PCD in their historical context. Revisions of our understanding of PCD demand a reassessment of its origin. Our second aim is thus to structure the proposed origin explanations of PCD into coherent arguments. In our analysis we argue for the evolutionary concept of PCD and the viral defense-immunity hypothesis for the origin of PCD. We suggest that this framework offers a plausible account of PCD early in the history of life, and also provides an epistemic basis for the future development of a general evolutionary account of mortality. (shrink)

What changes when an evolutionary transition in individuality takes place? Many different answers have been given, in respect of different cases of actual transition, but some have suggested a general answer: that a major transition is a change in the extent to which selection acts at one hierarchical level rather than another. The current paper evaluates some different ways to develop this general answer as a way to characterise the property ‘evolutionary individuality’; and offers a justification of the option taken (...) in Clarke :413–435, 2013)—to define evolutionary individuality in terms of an object’s capacity to undergo selection at its own level. In addition, I suggest a method by which the property can be measured and argue that a problem which is often considered to be fatal to that method—the problem of ‘cross-level by-products’—can be avoided. (shrink)

Evolutionary transitions in individuality (ETIs) are events during which individuals at a given level of organization (particles) interact to form higher-level entities (collectives) which are then recognized as new individuals at that level. ETIs are intimately related to levels of selection, which, following Okasha, can be approached from two different perspectives. One, referred to as ‘synchronic’, asks whether selection occurs at the collective level while the partitioning of particles into collectives is taken for granted. The other, referred to as ‘diachronic’, (...) asks about the origins of the partitioning of particles into collectives. After having presented the two perspectives and a classical formalism used to deal with the levels-of-selection question in the literature, namely the multilevel version of the Price equation, I show that because this formalism treats the levels-of-selection question from a synchronic perspective, it is inadequate to explain ETIs. This is because a fundamental aspect of ETIs is the origin of collectives. From there, I develop a framework for levels of selection compatible with the diachronic perspective. This framework relies on the distinction between what I call, on the one hand, a ‘functional aggregative collective trait’, and on the other hand, a ‘functional non-aggregative collective trait’. After having presented this distinction, I implement it in the Price equation, leading to a new statistical partitioning of this equation which, I argue, represents a causal decomposition more relevant for ETIs. Finally, I exploit this partitioning to explain the critical stages of an ETI. In addition to its explanatory power, a measure of the degree of functional non-aggregativity could be used as a proxy for the degree of individuality of a collective. (shrink)

We assess the arguments for recognising functionally integrated multispecies consortia as genuine biological individuals, including cases of so-called ‘holobionts’. We provide two examples in which the same core biochemical processes that sustain life are distributed across a consortium of individuals of different species. Although the same chemistry features in both examples, proponents of the holobiont as unit of evolution would recognize one of the two cases as a multispecies individual whilst they would consider the other as a compelling case of (...) ecological dependence between separate individuals. Some widely used arguments in support of the ‘holobiont’ concept apply equally to both cases, suggesting that those arguments have misidentified what is at stake when seeking to identify a new level of biological individuality. One important aspect of biological individuality is evolutionary individuality. In line with other work on the evolution of individuality, we show that our cases can be distinguished by focusing on the fitness alignment between the partners of the consortia. We conclude that much of the evidence currently presented for the ubiquity and importance of multi-species individuals is simply not to the point, at least unless the issue of biological individuality is firmly divorced from the question of evolutionary individuality. (shrink)

The Darwinian theory of evolution is itself evolving and this book presents the details of the core of modern Darwinism and its latest developmental directions. The authors present current scientific work addressing theoretical problems and challenges in four sections, beginning with the concepts of evolution theory, its processes of variation, heredity, selection, adaptation and function, and its patterns of character, species, descent and life. The second part of this book scrutinizes Darwinism in the philosophy of science and its usefulness in (...) understanding ecosystems, whilst the third section deals with its application in disciplines beyond the biological sciences, including evolutionary psychology and evolutionary economics, Darwinian morality and phylolinguistics. The final section addresses anti-Darwinism, the creationist view and issues around teaching evolution in secondary schools. The reader learns how current experimental biology is opening important perspectives on the sources of variation, and thus of the very power of natural selection. This work examines numerous examples of the extension of the principle of natural selection and provides the opportunity to critically reflect on a rich theory, on the methodological rigour that presides in its extensions and exportations, and on the necessity to measure its advantages and also its limits. Scholars interested in modern Darwinism and scientific research, its concepts, research programs and controversies will find this book an excellent read, and those considering how Darwinism might evolve, how it can apply to the human sciences and other disciplines beyond its origins will find it particularly valuable. Originally produced in French (Les Mondes Darwiniens), the scope and usefulness of the book have led to the production of this English text, to reach a wider audience. This book is a milestone in the impressive penetration by Francophone scholars into the world of Darwinian science, its historiography and philosophy over the last two decades. Alex Rosenberg, R. Taylor Cole Professor of Philosophy, Duke University Until now this useful and comprehensive handbook has only been available to francophones. Thanks to this invaluable new translation, this collection of insightful and original essays can reach the global audience it deserves. Tim Lewens, University of Cambridge. (shrink)

Building upon a non-standard understanding of evolutionary process focusing on variation and persistence, I will argue that communities and ecosystems can evolve by natural selection as emergent individuals. Evolutionary biology has relied ever increasingly on the modeling of population dynamics. Most have taken for granted that we all agree on what is a population. Recent work has reexamined this perceived consensus. I will argue that there are good reasons to restrict the term “population” to collections of monophyletically related replicators and (...) interactors, which explains why many existing models in population biology exclude by definition many genuine evolving biological individuals such as communities and ecosystems. By studying a case of community evolution, we will see that it is variation that is important to evolutionary processes, not populations. Variation within a population is only one of many types of variation that can lead to evolution by natural selection. The upshot of focusing on variation is that cases of community and ecosystem adaptive change become tractable in evolutionary terms. I will show that complex emergent individuals such as communities and ecosystems cannot be fully accommodated by conventional population/reproduction models but can be accommodated by variation/persistence models. (shrink)

Through a critical engagement with Merleau-Ponty’s discussion of the concepts of nature, life, and behavior, and with contemporary accounts of animal groups, this article argues that animal groups exhibit sociality and that sociality is a fundamental ontological condition. I situate my account in relation to the superorganism and selfish individual accounts of animal groups in recent biology and zoology. I argue that both accounts are inadequate. I propose an alternative account of animal groups and animal sociality through a Merleau-Pontian inspired (...) definition of behavior. I criticize Merleau-Ponty’s individualistic prejudice, but show that his philosophy contains the resources necessary to overcome this bias. I define behavior as a holistic, ongoing, meaningful and Umwelt-oriented intrinsically configured expression of living forms of existence. By looking at cases of animal groups drawn from contemporary studies in zoology and behavioral ecology, I show that animal groups, in the fact that they behave, manifest themselves to be a fundamental form of existence, namely, the social form of existence. (shrink)

The impressive variation amongst biological individuals generates many complexities in addressing the simple-sounding question what is a biological individual? A distinction between evolutionary and physiological individuals is useful in thinking about biological individuals, as is attention to the kinds of groups, such as superorganisms and species, that have sometimes been thought of as biological individuals. More fully understanding the conceptual space that biological individuals occupy also involves considering a range of other concepts, such as life, reproduction, and agency. There has (...) been a focus in some recent discussions by both philosophers and biologists on how evolutionary individuals are created and regulated, as well as continuing work on the evolution of individuality. (shrink)

The biological sciences have always proven a fertile ground for philosophical analysis, one from which has grown a rich tradition stemming from Aristotle and flowering with Darwin. And although contemporary philosophy is increasingly becoming conceptually entwined with the study of the empirical sciences with the data of the latter now being regularly utilised in the establishment and defence of the frameworks of the former, a practice especially prominent in the philosophy of physics, the development of that tradition hasn’t received the (...) wider attention it so thoroughly deserves. This review will briefly introduce some recent significant topics of debate within the philosophy of biology, focusing on those whose metaphysical themes (in everything from composition to causation) are likely to be of wide-reaching, cross-disciplinary interest. (shrink)

Developmental systems theory (DST) is a wholeheartedly epigenetic approach to development, inheritance and evolution. The developmental system of an organism is the entire matrix of resources that are needed to reproduce the life cycle. The range of developmental resources that are properly described as being inherited, and which are subject to natural selection, is far wider than has traditionally been allowed. Evolution acts on this extended set of developmental resources. From a developmental systems perspective, development does not proceed according to (...) a preformed plan; what is inherited is much more than DNA; and evolution is change not only in gene frequencies, but in entire developmental systems. (shrink)

Recent debates between proponents of the modern evolutionary synthesis (the standard model in evolutionary biology) and those of a possible extended synthesis are a good example of the fascinating tangle among empirical, theoretical, and conceptual or philosophical matters that is the practice of evolutionary biology. In this essay, we briefly discuss two case studies from this debate, highlighting the relevance of philosophical thinking to evolutionary biologists in the hope of spurring further constructive cross-pollination between the two fields.