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)

The units or levels of selection debate concerns the question of what kind of biological systems are stable enough that part of their evolution is a result of the process of natural selection acting at their level. Traditionally, the debate has concerned at least two different, though related, questions: the question of the level at which interaction with the environment occurs, and the question of the level at which reproduction occurs. In recent years, biologists and philosophers have discussed a new (...) aspect of this debate, namely the possibility that certain multi-species consortia formed by a host and its microbiome may act as a unit of selection. This thesis, however, has not been without criticism, as it is doubtful that such consortia could meet the conditions required to achieve the degree of stability that would allow them to experience natural selection. The purpose of this paper is to systematically examine such criticisms and to defend the thesis that the holobiont/hologenome can act as a genuine level of selection both in the form of an interactor and in the form of a reproducer. To do so, it will be argued that the microbiome should be characterized in functional rather than taxonomic terms. (shrink)

The hologenome concept of evolution is a hypothesis about the evolution of animals and plants. It asserts that the evolution of animals and plants was partially triggered by their interactions with their symbiotic microbiomes. In that vein, the hologenome concept posits that the holobiont (animal host + symbionts of the microbiome) is a unit of selection. -/- The hologenome concept has been severely criticized on the basis that selection on holobionts would only be possible if there were a tight transgenerational (...) host-genotype-to-symbiont-genotype connection. As our current evidence suggests that this is not the case for most of the symbiont species that compose the microbiome of animals and plants, the opportunity for holobiont selection is very low in relation to the opportunity for selection on each of the species that compose the host microbiome. Therefore, holobiont selection will always be disrupted ‘from below’, by selection on each of the species that compose the microbiome. -/- This thesis constitutes a conceptual effort to defend philosophically the hologenome concept. I argue that the criticism according to which holobiont selection requires tight transgenerational host-genotype-to-symbiont-genotype connection is grounded on a metaphysical view of the world according to which the biological hierarchy needs to be nested, such that each new level of selection includes every entity from below. Applied to hologenomes, it entails that the hologenome is a collection of genomes, and selection of hologenomes is assumed to entail cospeciation of the host with the species that constitute its microbiome. -/- Against that interpretation, I propose the ‘stability of traits’ account, according to which hologenome evolution is the result of the action of natural selection in a non-nested hierarchical world. In that vein, hologenome evolution does not entail cospeciation, and thus it does not require tight transgenerational host-genotype-to-symbiont-genotype connection. By embracing a multilevel selection perspective, I argue that hologenome evolution results from the simultaneous action of natural selection on each of the lineages that compose the microbiome, and on the assemblage composed by the host genome plus the functional traits of its microbiome. Hologenome selection occurs when the evolution of the traits of the microbiome result from their effects on the fitness of the host, and it can take the form of multilevel selection 1, or multilevel selection 2. In both cases, hologenome selection entails the evolution of microbiome traits, as well as evolution of the host genome, rather than cospeciation of lineages. (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)

Although contemporary metaphysics has recently undergone a neo-Aristotelian revival wherein dispositions, or capacities are now commonplace in empirically grounded ontologies, being routinely utilised in theories of causality and modality, a central Aristotelian concept has yet to be given serious attention – the doctrine of hylomorphism. The reason for this is clear: while the Aristotelian ontological distinction between actuality and potentiality has proven to be a fruitful conceptual framework with which to model the operation of the natural world, the distinction between (...) form and matter has yet to similarly earn its keep. In this chapter, I offer a first step toward showing that the hylomorphic framework is up to that task. To do so, I return to the birthplace of that doctrine - the biological realm. Utilising recent advances in developmental biology, I argue that the hylomorphic framework is an empirically adequate and conceptually rich explanatory schema with which to model the nature of organisms. (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)

Evolutionary Transitions in Individuality have been responsible for the major transitions in levels of selection and individuality in natural history, such as the origins of prokaryotic and eukaryotic cells, multicellular organisms, and eusocial insects. The integrated hierarchical organization of life thereby emerged as groups of individuals repeatedly evolved into new and more complex kinds of individuals. The Social Protocell Hypothesis proposes that the integrated hierarchical organization of human culture can also be understood as the outcome of an ETI—one that produced (...) a “cultural organism” from a substrate of socially learned traditions that were contained in growing and dividing social communities. The SPH predicts that a threshold degree of evolutionary individuality would have been achieved by 2.0–2.5 Mya, followed by an increasing degree of evolutionary individuality as the ETI unfolded. We here assess the SPH by applying a battery of criteria—developed to assess evolutionary individuality in biological units—to cultural units across the evolutionary history of Homo. We find an increasing agreement with these criteria, which buttresses the claim that an ETI occurred in the cultural realm. (shrink)

Do multi-level selection explanations of the evolution of social traits deepen the understanding provided by single-level explanations? Central to the former is a mathematical theorem, the multi-level Price decomposition. I build a framework through which to understand the explanatory role of such non-empirical decompositions in scientific practice. Applying this general framework to the present case places two tasks on the agenda. The first task is to distinguish the various ways of suppressing within-collective variation in fitness, and moreover to evaluate their (...) biological interest. I distinguish four such ways: increasing retaliatory capacity, homogenising assortment, and collapsing either fitness structure or character distribution to a mean value. The second task is to discover whether the third term of the Price decomposition measures the effect of any of these hypothetical interventions. On this basis I argue that the multi-level Price decomposition has explanatory value primarily when the sharing-out of collective resources is `subtractable'. Thus its value is more circumscribed than its champions Sober and Wilson (1998) suppose. (shrink)

Microbes are generally thought of as unicellular organisms, but we know that many microbes live as parts of biofilms—complex, surface-attached microbial communities numbering millions of cells. Some authors have recently argued in favour of reconceiving biofilms as biological entities in their own right. In particular, some have claimed that multispecies biofilms are evolutionary individuals : 10126–10132 2015). Against this view, I defend the conservative consensus that selection acts primarily upon microbial cells.

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)

In Denis Walsh’s Organisms, Agency, and Evolution, he argues that new developments in the science of biology motivate a radical change to our metaphysical picture of life: what he calls ‘Situated Darwinism’. The central claim is that we should take the biological world to be at base about organisms, and organisms in a fundamentally teleological sense. We critically examine Walsh’s arguments and suggest further developments.

In this introductory article to the special issue on Multi-level semiosis we attempt to stage the background for qualifying the notion of “multi-levelness” when considering communication processes and semiosis in all life forms, i.e. from the cellular to the organismic level. While structures are organized hierarchically, communication processes require a kind of processual organization that may be better described as being heterarchical. Theoretically, the challenge arises in the temporal domain, that is, in the developmental and evolutionary dimension of dynamic semiotic (...) processes. We discuss the importance of this fundamental difference in order to explain how levels, domains and orders of magnitude, on the one hand, and synchronic and diachronic processes, on the other, contribute to the overall organization of every living being. To account for such multi-level organization, semiotic freedom is assumed to be a scalar property that endows living systems at different levels and domains with the capacity to ponder selectively the overall structural coherence and functional compatibility of their heterarchical processing, which is increasingly less conditioned by the underlying molecular determinism. (shrink)

In dicephalus conjoined twinning, it appears that two heads share a body; in cephalopagus, it appears that two bodies share a head. How many human animals are present in these cases? One answer is that there are two in both cases—conjoined twins are precisely that, conjoined twins. Another is that the number of humans corresponds to the number of bodies—so there is one in dicephalus and two in cephalopagus. I show that both of these answers are incorrect. Prominent accounts of (...) biological individuation, which treat the organism as an integrated whole, reveal that in these and other cases of conjoined twinning the ‘twins’ share a single human animal. This has a number of consequences for the debate about what we are. First, if animalism is true, individuals of our kind can be—and are—profoundly psychologically divided. Second, cephalopagus twinning does not divide animalism from its rivals, as has been claimed. Finally, animalists can reply to a vicious species of the ‘too many thinkers’ problem to which they are allegedly uniquely vulnerable. (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)

This article proposes two conditions to assess whether an entity at a level of description is a unit of selection qua interactor. These two conditions make it possible to (1) distinguish biologically relevant entities from arbitrary ones and (2) distinguish units that can _potentially_ enter a selection process from those that have already done so. I show that the classical approaches used in the literature on units and levels of selection do not fare well with respect to either or both (...) of these desiderata. (shrink)

Various results show the ‘formal equivalence’ of kin and group selectionist methodologies, but this does not preclude there being a real and useful distinction between kin and group selection processes. I distinguish individual- and population-centred approaches to drawing such a distinction, and I proceed to develop the latter. On the account I advance, the differences between kin and group selection are differences of degree in the structural properties of populations. A spatial metaphor provides a useful framework for thinking about these (...) differences: kin and group selection may be conceptualized as large, overlapping regions of K - G space. I then consider some implications of the account, defend it from possible objections, and further argue that the structural features characteristic of both kin and group selection may recur at multiple levels of biological organization. (shrink)

In this paper, I touch on the holotic structure of the ideas of unity, identity and finality, using the word “holotic” to refer to a theory that includes both partonomic (or mereological) wholes and taxonomic (or diairological) wholes. In the second section, I expound on two classifications of wholes and two classifications of the types of parts I deem relevant to the ideas of unity, identity and finality. In the third and fourth sections, I discuss how these three ideas acquire (...) different significance depending on the type of totalities and parts involved. (shrink)

In this work it is proposed that the conditions of identity for biological organisms are given by the following principle: for all organisms x and y, x = y if and only if x has been caused by the self-preserving activity of y. This principle determines both the inter-temporal identity of organisms and the identity of organism in different possible worlds. It unifies what can be supposed about conditions of identity coming from —at least— three different conceptions about the nature (...) of biological organisms, those of RobertWilson, Thomas Pradeu and Ellen Clarke. These conceptions are presented and compared. (shrink)

Philosophers of biology typically pose questions about individuation by asking “what is an individual?” For example, we ask, “what is an individual species”, “what is an individual organism”, and “what is an individual gene?” In the first part of this chapter, I present my account of the gene concept and how it is used in investigative practices in order to motivate a more pragmatic approach. Instead of asking “what is a gene?”, I ask: “how do biologists individuate genes?”, “for what (...) purposes?”, and “do their practices of individuating genes serve these purposes?" In the second part of this chapter, I propose that we use this approach when analyzing concepts of organisms and biological individuals. Following philosophical pragmatism, I argue that we should abstain from attempts to situate individuation of Darwinian individuals or of holobionts in a philosophy of nature. Instead, we should analyze practices of individuating organisms in terms of three-place relations between the world, ideas, and human purposes and actions. I conclude with three lessons: an ontological, an epistemological, and a meta-philosophical lesson, which I suggest, apply to philosophy of science generally and to philosophy and metaphysics at large. (shrink)

In this paper, I introduce the notion of ‘philosophy moves’: prominent tropes featured in contemporary academic philosophy. Moves are more than patterns – they are tools for advancing and enriching philosophical debates. By recognizing these patterns in the philosophical literature, we collect an ensemble of moves for deployment in novel contexts, each with the potential to forge new paths of philosophical investigation through a given topic. The moves featured in this paper are constructive and progressive, with the potential to push (...) past stalemates and open up a wider conceptual landscape. Following a presentation of five exemplars taken from the philosophy of biology literature, I show how different moves could be made within a current interdisciplinary debate. Overall, I aim to demonstrate that recognizing these patterns is insightful pedagogically and potentially useful when put to use as part of one’s philosophical methodology. (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)

Given the central place organisms occupy in Kant’s account of living nature, it might seem unlikely that his claims about biological wholes could be relevant to current debates over the problem of biological individuality. These debates acknowledge the multiple realizability of biological individuality in vastly different forms, including parts of organisms and complex groups of organisms at various levels of the biological hierarchy, sparking much controversy in attempts to characterize a biological individual. I argue that, far from being irrelevant to (...) this controversy, Kant’s account provides a key insight for addressing the multiple realizability problem. I show how the reciprocal causality between a self-organizing whole and its parts, which Kant thinks characterizes a natural end, is not limited to organisms but is exhibited by numerous types of beings in living nature. Self-organizing wholes of various kinds, and at various biological levels, may count as biological individuals, depending on the degree to which their functionally integrated parts are represented by reflective judgement as a natural end. (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)

Contemporary biology has inherited two key assumptions from the Modern Synthesis about the nature of population lineages: sexual reproduction is the exemplar for how individuals in population lineages inherit traits from their parents, and random mating is the exemplar for reproductive interaction. While these assumptions have been extremely fruitful for a number of fields, such as population genetics and phylogenetics, they are increasingly unviable for studying the full diversity and evolution of life. I introduce the “mixture” account of population lineages (...) that escapes these assumptions by dissolving the Modern Synthesis’s sharp line separating reproduction and development and characterizing reproductive integration in population lineages by the ephemerality of isolated subgroups rather than random mating. The mixture account provides a single criterion for reproductive integration that accommodates both sexual and asexual reproduction, unifying their treatment under Kevin de Queiroz’s generalized lineage concept of species. The account also provides a new basis for empirically assessing the effect of random mating as an idealization on the empirical adequacy of population genetic models. (shrink)

Multicellular organisms contain numerous symbiotic microorganisms, collectively called microbiomes. Recently, microbiomic research has shown that these microorganisms are responsible for the proper functioning of many of the systems of multicellular organisms. This has inclined some scholars to argue that it is about time to reconceptualise the organism and to develop a concept that would place the greatest emphasis on the vital role of microorganisms in the life of plants and animals. We believe that, unfortunately, there is a problem with this (...) suggestion, since there is no such thing as a universal concept of the organism which could constitute a basis for all biological sciences. Rather, the opposite is true: numerous alternative definitions exist. Therefore, comprehending how microbiomics is changing our understanding of organisms may be a very complex matter. In this paper we will demonstrate that this pluralism proves that claims about a change in our understanding of organisms can be treated as both true and untrue. Mainly, we assert that the existing concepts differ substantially, and that only some of them have to be reconsidered in order to incorporate the discoveries of microbiomics, while others are already flexible enough to do so. Taking into account the plurality of conceptualisations within different branches of modern biology, we will conduct our discussion using the developmental and the cooperation–conflict concepts of the organism. Then we will explain our results by referring to the recent philosophical debate on the nature of the concept of the organism within biology. (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)

Despite emerging attention to Indigenous philosophies both within and outside of feminism, Indigenous logics remain relatively underexplored and underappreciated. By amplifying the voices of recent Indigenous philosophies and literatures, I seek to demonstrate that Indigenous logic is a crucial aspect of Indigenous resurgence as well as political and ethical resistance. Indigenous philosophies provide alternatives to the colonial, masculinist tendencies of classical logic in the form of paraconsistent—many-valued—logics. Specifically, when Indigenous logics embrace the possibility of true contradictions, they highlight aspects of (...) the world rejected and ignored by classical logic and inspire a relational, decolonial imaginary. To demonstrate this, I look to biology, from which Indigenous logics are often explicitly excluded, and consider one problem that would benefit from an Indigenous, paraconsistent analysis: that of the biological individual. This article is an effort to expand the arenas in which allied feminists can responsibly take up and deploy these decolonial logics. (shrink)

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)

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)

Theoreticians that defend a form of realism regarding natural kinds minimally entertain the belief that the world features divisions into kinds and that the natural kind concept is a useful tool for philosophy of science. The objective of this paper is to challenge these assumptions. First, we challenge realism toward natural kinds by showing that the main arguments for their existence, which rely on the epistemic success of natural kinds, are unsatisfactory. Second, we show that, whether they exist or not, (...) natural kinds are expendable when it comes to describing and analyzing scientific explanations accurately. (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)

According to the mainstream position in the bioethical definition of death debate, death is to be equated with the cessation of an organism. Given such a perspective, some bioethicists uphold the position that brain-dead patients are dead, while others claim that they are alive. Regardless of the specific opinion on the status of brain-dead patients, the mere bioethical concept of death, according to many bioethicists, has the merit of being unanimous and univocal, as well as grounded in biology. In the (...) present article, we challenge such a thesis. We provide evidence that theoretical biology operates with a plurality of equally valid organismic concepts, which imply different conclusions regarding the organismal status of a brain-dead patient. Moreover, the theoretical biology concepts of an organism are very distant from the view on an organism that appears by way of bioethicists theorizing on death. We conclude that if death is to be understood as the cessation of an organism, there is no single correct answer to the question of whether a brain-dead patient is alive or dead. (shrink)

This article defends lineage pluralism; the view that biological lineages are not a single, unified type of entity. I analyze aspects of evolutionary theory, phylogenetics, and developmental biology to show that these areas appeal to distinct notions of lineage. I formulate three arguments for lineage pluralism. These arguments undercut the main motivations for lineage monism; the view that biological lineages are a single, unified type of entity. Although this view is rarely made explicit, it is often assumed in philosophy and (...) biology. Hence, this article sheds light on this monistic assumption and shows why lineage pluralism should be adopted instead. (shrink)

The Problem of Biological Individuality is the problem of how to count organisms. Whilst counting organisms may seem easy, the biological world is full of difficult cases such as colonial siphonophores and aspen tree groves. One of the main solutions to the Problem of Biological Individuality is the Physiological Approach. Drawing on an argument made by Eric Olson in the personal identity debate, I argue that the Physiological Approach faces a metaphysical problem - the ‘Foetus Problem’. This paper illustrates how (...) metaphysics can contribute to debates about organisms in the philosophy of biology. (shrink)

The Problem of Biological Individuality is the problem of how to count organisms. Whilst counting organisms may seem easy, the biological world is full of difficult cases such as colonial siphonophores and aspen tree groves. One of the main solutions to the Problem of Biological Individuality is the Physiological Approach. Drawing on an argument made by Eric Olson in the personal identity debate, I argue that the Physiological Approach faces a metaphysical problem - the ‘Foetus Problem’. This paper illustrates how (...) metaphysics can contribute to debates about organisms in the philosophy of biology. (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)

Recently, philosophers have sought to determine the nature of individuals relevant to evolution by natural selection or evolutionary individuals. The Evolutionary Contingency Thesis is a claim about evolution that emphasizes the role of contingency or dependency relations and chance-based factors in how evolution unfolds. In this article, I argue that if we take evolutionary contingency seriously, then we should be pluralists about the types of individuals in selection.

Ecological communities are seldom, if ever, biological individuals. They lack causal boundaries as the populations that constitute communities are not congruent and rarely have persistent functional roles regulating the communities’ higher-level properties. Instead we should represent ecological communities indexically, by identifying ecological communities via the network of weak causal interactions between populations that unfurl from a starting set of populations. This precisification of ecological communities helps identify how community properties remain invariant, and why they have robust characteristics. This respects the (...) diversity and aggregational nature of these complex systems while still vindicating them as units worthy of investigation. (shrink)

I consider the relationship between scientific practice and the philosophical debate surrounding biological individuality. I argue for the sensitivity account, on which biologists do not require a resolution to the individuality debate. This view puts me in disagreement with much of the literature on biological individuality, where it has become common to claim that there is a relationship of dependence between biologists’ conceptions of individuality and the quality of their empirical work.

Synthetic biology promises to eliminate the distinction between biology and engineering by delivering a philosophically interesting new kind of entity: a biological organism that is wholly designed and constructed by humans. The possibility of such organisms raises interesting questions in three domains: the analysis of (1) biological functions, (2) engineering functions, and (3) health and disease. This paper identifies and systematically answers these questions. This does not only establish how we should think about functions and health and disease in synthetic (...) biological organisms, but it also reveals insights that are of broader relevance: (1) aetiological accounts of biological function need to omit or reinterpret reference to natural selection. This results in complete continuity between aetiological analyses of function in engineering and philosophy; (2) considering synthetic biology prompts interesting further questions about heritability, ancestry, and biological individuals; and (3) accounts of disease as biological dysfunction do not straightforwardly map onto our intuitive health and disease judgments regarding non‐human animals. In response to the latter point I examine three possible avenues, and tentatively defend one on behalf of the ‘disease as dysfunction’ theorist. (shrink)

Mammals are usually considered unproblematic as biological individuals. This article contends the opposite. Once we consider pregnancy, criteria for biological individuality are not easily applicab...

Biologists, historians of biology, and philosophers of biology often ask what is it to be an individual, really. This book does not answer that question. Instead, it answers a much more interesting one: How do biologists individuate individuals? In answering that question, the authors explore why biologists individuate individuals, in what ways, and for what purposes. The cross-disciplinary, dialogical approach to answering metaphysical questions that is pursued in the volume may seem strange to metaphysicians who are not biologically focused, but (...) it is adroitly achieved by the editors. Scott Lidgard (a paleontologist and marine ecologist) and Lynn K. Nyhart (a historian of biology) orchestrate a dialogue among historians of biology, philosophers of biology, and practicing biologists over 10 chapters. These are followed by three reflective commentaries written to frame the different disciplinary perspectives and to highlight the historical, biological, and philosophical themes across the chapters. The result is a volume—in structure and in content—that has much to be generously commended. Biological individuality is a hotly discussed topic, but it is also part of a series of long-standing arguments within both the history and philosophy of biology (HPB) and metaphysics. Notable and fervent debates have centered on evolution and the units of selection, predominantly on Michael T. Ghiselin’s and David L. Hull’s notion of species as individuals, Peter Godfrey-Smith’s Darwinian individuals, and Ellen Clarke’s individuating mechanisms. Lately, it has encompassed non-Darwinian individuals, symbiotic associations like Thomas Pradeu’s immunological individuals, and John Dupré and Maureen A. O’Malley’s metabolic individuals.2 The present volume is curated in a way to introduce the reader to new research in HPB that articulates these debates as well as to introduce and engage in the study of further notions of biological individuality. But its aim is more than an introduction. As the subtitle suggests, it is also intended to give the reader insight into the working together of biologists, historians of biology, and philosophers of biology in figuring out how the notion of biological individuality is instantiated. As such, the problem-centered dialogue that results does more than talk through biological individuality. It shows how the different and often divergent goals of the authors’ disciplines shape not only how they think about individuality but how they communicate this thinking in reciprocal collaboration with others in different disciplines. … cont’d…. (shrink)

Immunology and its philosophy are a primary source for thinking about biological individuality. Through its discriminatory function, the immune system is believed to delineate organism and environment within one generation, thus defining the physiological individual. Based on the paradigmatic instantiations of immune systems, immune interactions and, thus, the physiological individual are believed to last only for one generation. However, in recent years, transgenerationally persisting immune responses have been reported in several phyla, but the consequences for physiological individuality have not yet (...) been explored. In this article, I will introduce an invertebrate immune system that is RNA-based and operates through a heritable silencing/licensing paradigm. I will discuss how such a perspective on immune systems can illuminate our conceptions of individuality. I will particularly introduce an account of immunological individuality that is not restricted to one generation. (shrink)

Evolution by natural selection has been extended to several supraorganismic levels, but whether it can apply to ecosystems remains controversial on two main counts. First, local ecosystems are loosely individuated, so that it is unclear how they manifest heredity and fitness. Second, even if they did, the meta-ecosystem formed by this population of local ecosystems will also suffer from a very low degree of cohesion, which will jeopardize any ENS. We suggest a way to overcome both issues, focusing on ecosystem (...) properties rather than on ecosystemic individuals. First, we follow recent theoretical developments which deny the centrality of reproduction in ENS. This leads us to merge heredity and fitness in a single process, in which a local ecosystem causally influences its neighbors, in a way responsible for their phenotypic similarity. Second, we suggest that the resulting meta-ecosystem need not meet the criteria of individuality. Instead, the evolving meta-ecosystem is best understood as a homeostatic property cluster, i.e., a set of phenotypes that together evolve by ENS at the ecosystem level. We illustrate this conceptual framework with a hypothetical example of local ecosystems that vary in terms of stability, productivity, diversity, and complementarity between species. Finally, we stress that our property-based account of ecosystemic ENS can help understand evolution at other biological levels, such as early life evolution and holobionts. (shrink)