In this paper I develop a concept of behavioural ecological individuality. Using findings from a case study which employed qualitative methods, I argue that individuality in behavioural ecology should be defined as phenotypic and ecological uniqueness, a concept that is operationalised in terms of individual differences such as animal personality and individual specialisation. This account make sense of how the term “individuality” is used in relation to intrapopulation variation in behavioural ecology. The concept of behavioural ecological individuality can sometimes be (...) used to identify individuals. It also shapes research agendas and methodological choices in behavioural ecology, leading researchers to account for individuals as sources of variation. Overall, this paper draws attention to a field that has been largely overlooked in philosophical discussions of biological individuality and highlights the importance of individual differences and uniqueness for individuality in behavioural ecology. (shrink)

Elselijn Kingma argues that, in cases of mammalian placental pregnancy, the foster (roughly, the post-implantation embryo/foetus) is part of the gravida (the pregnant organism). But she does not consider the possibility of proper overlap. I show that this generates a number of serious problems for her argument and trace the oversight to a quite general issue within the literature on biological individuality. Doing so provides an opportunity to pull apart and clarify the relations between some importantly distinct questions concerning organismality (...) and organismic parthood, and to identify the implications that this has for those who want to draw on this literature when making metaphysical or mereological arguments. (shrink)

This paper examines the metaphysics of interdependence in the work of the Chinese Buddhist Fazang. The dominant approach of this metaphysics interprets it as a species of metaphysical coherentism wherein everything depends upon everything else, no individual is more fundamental than any other, and so reality itself is non-well-founded in the sense that chains of dependence never terminate. I argue, to the contrary, that Fazang's metaphysics is better interpreted as a novel variety of foundationalism. I argue, as well, using set- (...) and graph-theoretic techniques, that there is a consistent way to model this alternative interpretation, and that this model differs in significant ways from a coherentist model. (shrink)

Both concepts of the holobiont and the immune system are at the heart of an ongoing scientific and philosophical examination concerning questions of the organism’s individuality and identity as well as the relations between organisms and their environment. Examining the holobiont, the question of boundaries and individuality is challenging because it is both an assemblage of organisms with physiological cohesive aspects. I discuss the concept of immunity and the immune system function from the holobiont perspective. Because of the host-microbial close (...) relations of codependence and interdependence, the holobiont is more often than not confused with the host, as the host is the domain in which this entity exists. I discuss the holobiont unique ecological characteristics of microbial assemblages connected to a host in a network of interactions in which the host is one of the organisms in the community but also its landscape. Therefore, I suggest viewing the holobiont as a host-ecosystem and discuss the implication of such a view on the concept of immunity and the meaning of protection. Furthermore, I show that viewing the holobiont as a host ecosystem opens the possibility of using the same ecological definition of boundaries and immunity dealing with an ecological system. Thus, the holobiont’s boundaries and immunity are defined by the persistence of its complex system of interactions integrating existing and new interactions. This way of thinking presents a notion of immunity that materializes as the result of the complex interdependence relations between the different organisms composing the holobiont similar to that of an ecosystem. Taking this view further, I discuss the notion of immunogenicity that is ontologically heterogeneous with various causal explanations of the processes of tolerance and targeted immune response. Finally, I discuss the possible conceptualization of already existing and new biomedical practices. (shrink)

Some philosophers of biology state that the metaphysical status of biological species is context determined by the use different branches of biology make of their corresponding proper names, so that one and the same biological species can be both an individual and a natural kind. In this paper, I aim to undermine the idea, often associated with the present thesis, according to which the debate about the metaphysical status of biological species should be deflated, since it would be possible to (...) translate every sentence from natural-kind talk to individual-talk and vice versa. I offer a charitable interpretation of the principle grounding such an idea and show how it is in tension with independent theoretical biological notions. (shrink)

Both physiological and evolutionary criteria of biological individuality are underpinned by the idea that an individual is a functionally integrated whole. However, a precise account of functional integration has not been provided so far, and current notions are not developed in the details, especially in the case of composite systems. To address this issue, this paper focuses on the organisational dimension of two representative associations of prokaryotes: biofilms and the endosymbiosis between prokaryotes. Some critical voices have been raised against the (...) thesis that biofilms are biological individuals. Nevertheless, it has not been investigated which structural and functional obstacles may prevent them from being fully integrated physiological or evolutionary units. By contrast, the endosymbiotic association of different species of prokaryotes has the potential for achieving a different type of physiological integration based on a common boundary and interlocked functions. This type of association had made it possible, under specific conditions, to evolve endosymbionts into fully integrated organelles. This paper therefore has three aims: first, to analyse the organisational conditions and the physiological mechanisms that enable integration in prokaryotic associations; second, to discuss the organisational differences between biofilms and prokaryotic endosymbiosis and the types of integration they achieve; finally, to provide a more precise account of functional integration based on these case studies. (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)

Um argumento recorrente contra a liberalização do aborto parte do pressuposto de que, desde o momento da fertilização, seres humanos são indivíduos (no sentido de serem algo que necessariamente ocorre em uma entidade apenas). Nesse artigo, argumento que esse não é um caso a partir da possibilidade de geminação monozigótica e do fato de identidades serem necessárias. Defendo as premissas do argumento e discuto as possíveis interpretações de sua conclusão. Argumento que a interpretação completa da conclusão desse argumento está em (...) concordância com a conclusão de um argumento semelhante, que parte de noção de organismo. -/- ENGLISH: A common pro-life argument assumes that humans are individuals (in the sense that they necessarily occur in one entity only) from the moment of fertilization. In this paper, argue that this is not the case because of the possibility of monozygotic twinning and the necessity of identities. I will defend the premises of the argument and discuss the possible interpretations of its conclusion. I argue that the correct interpretation of the conclusion of this argument is in agreement with the conclusion of a similar arguments from the notion of organism. (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)

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)

Marquis’ future-like-ours argument against the morality of abortion assumes animalism—a family of theories according to which we are animals. Such an assumption is theoretically useful for various reasons, e.g., because it provides the theoretical underpinning for a reply to the contraception-abstinence objection. However, the connection between the future-like-ours argument and one popular version of animalism can prove lethal to the former, or so I argue in this paper.

Is the mammalian embryo/fetus a part of the organism that gestates it? According to the containment view, the fetus is not a part of, but merely contained within or surrounded by, the gestating organism. According to the parthood view, the fetus is a part of the gestating organism. This paper proceeds in two stages. First, I argue that the containment view is the received view; that it is generally assumed without good reason; and that it needs substantial support if it (...) is to be taken seriously. Second, I argue that the parthood view derives considerable support from a range of biological and physiological considerations. I tentatively conclude in favour of the parthood view, and end by identifying some of the interesting questions it raises. (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)

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)

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)

In this paper I argue that it is finally time to move beyond the Nagelian framework and to break new ground in thinking about epistemic reduction in biology. I will do so, not by simply repeating all the old objections that have been raised against Ernest Nagel’s classical model of theory reduction. Rather, I grant that a proponent of Nagel’s approach can handle several of these problems but that, nevertheless, Nagel’s general way of thinking about epistemic reduction in terms of (...) theories and their logical relations is entirely inadequate with respect to what is going on in actual biological research practice. (shrink)

Darwin’s classic ‘Origin of Species’ (Darwin 1859) described forces of selection acting upon individuals, but there remains a great deal of controversy about what exactly the status and definition of a biological individual is. Recently some authors have argued that the individual is dispensable – that an inability to pin it down is not problematic because little rests on it anyway. The aim of this paper is to show that there is a real problem of biological individuality, and an urgent (...) need to arbitrate over the current plethora of solutions to it. (shrink)

The category of ‘organism’ has an ambiguous status: is it scientific or is it philosophical? Or, if one looks at it from within the relatively recent field or sub-field of philosophy of biology, is it a central, or at least legitimate category therein, or should it be dispensed with? In any case, it has long served as a kind of scientific “bolstering” for a philosophical train of argument which seeks to refute the “mechanistic” or “reductionist” trend, which has been perceived (...) as dominant since the 17th century, whether in the case of Stahlian animism, Leibnizian monadology, the neo-vitalism of Hans Driesch, or, lastly, of the “phenomenology of organic life” in the 20th century, with authors such as Kurt Goldstein, Maurice Merleau-Ponty, and Georges Canguilhem. In this paper I try to reconstruct some of the main interpretive ‘stages’ or ‘layers’ of the concept of organism in order to critically evaluate it. How might ‘organism’ be a useful concept if one rules out the excesses of ‘organismic’ biology and metaphysics? Varieties of instrumentalism and what I call the ‘projective’ concept of organism are appealing, but perhaps ultimately unsatisfying. (shrink)

The theory of evolution by natural selection is, perhaps, the crowning intellectual achievement of the biological sciences. There is, however, considerable debate about which entity or entities are selected and what it is that fits them for that role. This article aims to clarify what is at issue in these debates by identifying four distinct, though often confused, concerns and then identifying how the debates on what constitute the units of selection depend to a significant degree on which of these (...) four questions a thinker regards as central. (shrink)

This paper focuses on physiological integration in multicellular systems, a notion often associated with biological individuality, but which has not received enough attention and needs a thorough theoretical treatment. Broadly speaking, physiological integration consists in how different components come together into a cohesive unit in which they are dependent on one another for their existence and activity. This paper argues that physiological integration can be understood by considering how the components of a biological multicellular system are controlled and coordinated in (...) such a way that their activities can contribute to the maintenance of the system. The main implication of this perspective is that different ways of controlling their parts may give rise to multicellular organizations with different degrees of integration. After defining control, this paper analyses how control is realized in two examples of multicellular systems located at different ends of the spectrum of multicellularity: biofilms and animals. It focuses on differences in control ranges, and it argues that a high degree of integration implies control exerted at both medium and long ranges, and that insofar as biofilms lack long-range control (relative to their size) they can be considered as less integrated than other multicellular systems. It then discusses the implication of this account for the debate on physiological individuality and the idea that degrees of physiological integration imply degrees of individuality. (shrink)

En este trabajo ofrezco una interpretación de la concepción heideggeriana del organismo biológico presente en sus cursos de 1929. Para ello, primero enmarcaré la discusión dentro de la interpretación heideggeriana del naturalismo y su manera de entender la relación entre filosofía y ciencia. Luego, analizaré su interpretación de la embriología y la ecología mediante la cual Heidegger esboza aportes originales a la filosofía de la biología, especialmente en torno al problema de la identidad del organismo biológico. Finalmente, mostraré cómo esta (...) interpretación supone una profunda crítica a las concepciones sustancialistas del organismo a favor de una concepción relacional y procesual. (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)

Organisms receded from view in much of twentieth-century biology, only to undergo a sort of renaissance at the start of the twenty-first. The story of why this should be so is complicated and fascinating, but belongs primarily to the history of biology. On the other hand, to the extent that it is so, a question naturally arises: what, after all, are organisms? This question has a long and complicated history of its own, both within and without of biology; an investigation (...) of this history yields some guidance as to how organisms might yet be conceived today. One suggestion borne of these investigations is this: organisms are, for better or worse, normatively delineated unities. (shrink)

It is now widely accepted that microorganisms play many important roles in the lives of plants and animals. Every macroorganism has been shaped in some way by microorganisms. The recognition of the ubiquity and importance of microorganisms has led some to argue for a revolution in how we understand biological individuality and the primary units of natural selection. The term “holobiont” was introduced as a name for the biological unit made up by a host and all of its associated microorganisms, (...) and much of this new debate about biological individuality has focused on whether holobionts are integrated individuals or communities. In this paper, I show how parts of the holobiont can span both characterizations. I argue that most holobionts share more affinities with communities than they do with organisms, and that, except for maybe in rare cases, holobionts do not meet the criteria for being organisms, evolutionary individuals, or units of selection. (shrink)

The problem of plant individuality is something which has vexed botanists throughout the ages, with fashion swinging back and forth from treating plants as communities of individuals (Darwin 1800 ; Braun and Stone 1853 ; Münch 1938 ) to treating them as organisms in their own right, and although the latter view has dominated mainstream thought most recently (Harper 1977 ; Cook 1985 ; Ariew and Lewontin 2004 ), a lively debate conducted mostly in Scandinavian journals proves that the issues (...) are far from being resolved (Tuomi and Vuorisalo 1989b ; Fagerström 1992 ; Pan and Price 2001 ). In this paper I settle the matter once and for all, by showing which elements of each side are correct. (shrink)

Medical classification systems aim to provide a manageable taxonomy for sorting diagnoses into their proper classes. The question, this paper wants to critically examine, is how to correctly systematise diseases within classification systems that are applied in a variety of different settings. ICD and DSM , the two major classification systems in medicine and psychiatry, will be the main subjects of this paper; however, the arguments are not restricted to these classification systems but point out general methodological and epistemological challenges (...) of classifying diseases for differing purposes. Deciding what qualifies as a disease to be included into a classification system as well as choosing a specific validator for correctly systematising diseases is complicated because the broad applicability of medical classification systems simultaneously appears as aim and challenge. Drawing upon the case study of classifying Alzheimer’s disease, this paper will address three dilemmas in designing ‘good’ medical classification systems. They are due to general epistemological problems of medicine, such as the relationship between individual manifestations of diseases and the necessity of building groups in order to scientifically elucidate causes of diseases. Moreover, they involve pragmatic issues of designing usable classifications that allow for easily discriminating between classes of diseases, restricting, however, the completeness of disease representations. This paper wants to trace how the choice of certain validators is unavoidably value-laden and deeply intertwined with epistemological assumptions of how different uses relate to each other, resulting either in a prioritisation of (constrained) coherence or of (vague) pluralistic connectibility. (shrink)

Despite the radical importance of embryology in the development of organi- cism, developmental biology remains philosophically underexplored as a theoretical and empirical resource to clarify the nature of organisms. This paper discusses how embryology can help develop the organisational definition of the organism as a differentiated, function- ally integrated, and autonomous system. I distinguish two conceptions of development in the organisational tradition that yield two different conceptions of the organism: the life- history view claims that organisms can be considered as (...) such during their whole ontogeny; the constitutive view distinguishes two periods in the life history, a period of generation and a period of self-maintenance of a constitutive organisation. Arguing in favour of the con- stitutive view, it will be claimed that the organisational criteria for the definition of organ- ism (i.e., differentiation, functional integration, and autonomy) can only be applied to the developmental system when it has entered the period of self-maintenance of a constitutive organisation. under the light of current research in developmental biology, it is possible to make explicit how organisms come to be as organisms. To this end, I explore key onto- genetic events that help us clarify the core aspects of animal organisation and allow us to identify the developmental stage that marks the ontological transition between an organism in potency and an organism in actuality. The structure of this ontogenetic unfolding paral- lels the conceptual structure of the very notion of organism; the generation of the being of a particular organism parallels its definition. (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)

I propose an account of generous action in the Pāli Buddhist tradition, whereby generous actions are instances of giving in which the donor has esteem for the recipient of their giving. The account differs from recent Anglophone accounts of generous action. These tend to construe generous actions as instances of a donor freely offering a gift to the recipient for the sake of benefiting the recipient. Unlike the Buddhist account I propose, these accounts do not require donors to esteem their (...) recipient. Accordingly, I also offer a partial account of esteem, whereby one esteems another only if they refrain from noticing the other’s faults and they encounter the other as someone who is superior in virtue and goodness. Taken together, the Buddhist accounts of generous action and esteem offer insight into certain ways in which different philosophical traditions tend to characterize generous action. (shrink)

Convincing disputes about explanatory reductionism in the philosophy of biology require a clear and precise understanding of what a reductive explanation in biology is. The central aim of this book is to provide such an account by revealing the features that determine the reductive character of a biological explanation. Chapters I-IV provide the ground, on which I can then, in Chapter V, develop my own account of explanatory reduction in biology: Chapter I reveals the meta-philosophical assumptions that underlie my analysis (...) of reduction, Chapter II introduces the previous debate about reduction in the philosophy of biology by pointing out the most crucial lessons one should learn from this debate, Chapter III critically discusses the two perspectives on explanatory reduction that have been proposed in the philosophy of biology so far, and Chapter IV clarifies in what ways the issue of reduction is entangled with the issue of explanation. In Chapter V I present my own account of explanatory reduction. My main claim is that reductive explanations in the biological sciences possess three main characteristics: they explain the behavior of a biological system S, first, exclusively in terms of factors that are located on a lower level of organization than S, second, by focusing on factors that are internal to S , and third, by describing the genuine parts of S only as “parts in isolation”. This account is ontic because it traces the reductivity of an explanation back to certain relations that exist in the world. (shrink)

Since the 1950s, the common view of development has been internalist: development is seen as the result of the unfolding of potentialities already present in the egg cell. In this article, I show that this view is incorrect because of the crucial influence of the environment on development. I focus on a fascinating example, that of the role played by symbioses in development, especially bacterial symbioses, a phenomenon found in virtually all organisms (plants, invertebrates, and vertebrates). I claim that we (...) must consequently modify our conception of the boundaries of the developing entity, and I show how immunology can help us in accomplishing this task. I conclude that the developing entity encompasses many elements traditionally seen as “foreign,” while I reject the idea that there is no possible distinction between the organism and its environment. (shrink)

[Przekład] Pogląd głoszący, że układ odpornościowy rozróżnia to, co swoje, od tego, co obce, był jednym z centralnych założeń immunologii w drugiej połowie XX wieku. Pogląd ten miał wpływ na projekty eksperymentalne i interpretowanie danych. Jednakże w obliczu nowych dowodów empirycznych konieczny jest w immunologii nowy aparat konceptualny.

In this paper, I address the question of what the Developmental Systems Theory (DST) aims at explaining. I distinguish two lines of thought in DST, one which deals specifically with development, and tries to explain the development of the individual organism, and the other which presents itself as a reconceptualization of evolution, and tries to explain the evolution of populations of developmental systems (organism-environment units). I emphasize that, despite the claiming of the contrary by DST proponents, there are two very (...) different definitions of the ‘developmental system’, and therefore DST is not a unified theory of evolution and development. I show that the DST loses the most interesting aspects of its reconceptualization of development when it tries to reconceptualize evolutionary theory. I suggest that DST is about development per se, and that it fails at offering a new view on evolution. (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...

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)

One of Leibniz’s most original ideas is his conception of the living individual as a hierarchical network of living beings whose relationships are essential to the proper functioning of its organic body. This idea is also valid to explain any existing order in nature that depends on the set of relationships of living beings that inhabit it. Both ideas are present in the conception of the natural world that Leibniz presents in his Monadology through his idea of biological infinitism. According (...) to this idea, nature consists of infinite theatres where living beings unfold their vital functions. Through this idea Leibniz defines both the biological complexity of nature and the living individual, which is in turn a portion of nature that unfolds from an infinite set of inferior living beings. The thesis that I defend in this work is that this Leibnizian understanding of the living individual and the natural complexity that includes infinite hierarchical levels of individuality has a marked ecological sense, as we would say today. This Leibnizian metaphysics of individuality that we could call biological is also interesting in light of the recent studies in the philosophy of biology. (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)

Individuality is an important concept in biology, yet there are many non-equivalent criteria of individuality expressed in different kinds of biological individuals. This article evaluates these different kinds in terms of their capacity to support explanatory generalizations over the systems they individuate. Viewing the problem of individuality from this perspective promotes a splitting strategy in which different kinds make different epistemic trade-offs that suit them for different explanatory roles. I argue that evolutionary individuals, interpreted as forming a functional kind, face (...) difficulties of individuation and explanatory power that are mitigated by relying on more structurally based properties and non-evolutionary kinds. 1Introduction2Kinds of Biological Individuals3Evolutionary Individuals and Functional Kinds4Evolutionary versus Non-evolutionary Kinds of Individuals 4.1Physiological individuality4.2Ecological individuality4.3Developmental individuality5Conclusion. (shrink)

Viruses have been virtually absent from philosophy of biology. In this editorial introduction, we explain why we think viruses are philosophically important. We focus on six issues, and we show how they relate to classic questions of philosophy of biology and even general philosophy.

A recent development in biology has been the growing acceptance that holobionts, entities comprised of symbiotic microbes and their host organisms, are widespread in nature. There is agreement that holobionts are evolved outcomes, but disagreement on how to characterize the operation of natural selection on them. The aim of this paper is to articulate the contours of the disagreement. I explain how two distinct foundational accounts of the process of natural selection give rise to competing views about evolutionary individuality.

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)

Recent attention to complex group-level behavior amongst bacteria has led some to conceive of multicellular clusters of bacteria as individuals. In this article, I assess these recent claims by first drawing a distinction between two concepts of individuality: physiological and evolutionary. I then survey cases that are representative of three different modes of growth: myxobacteria (surface-attached agglomerative growth), Bacillus subtilis (agglomerative growth not attached to a surface), and cyanobacteria (filamentous growth). A closer look at these cases indicates that multicellular individuality (...) among bacteria is remarkably complex. Physiologically, the three cases of multicellular clusters do not form physiological individuals. But matters are different when it comes to evolutionary individuality; although multicellular clusters that grow by agglomeration are not highly individuated, filament-forming groups achieve a relatively high degree of individuality. I also suggest that debates about bacterial multicellular individuality may have been obscured by a failure to see that selection on highly individuated groups is by no means the only mechanism to bring about the complex group-level behaviors that have led some to view bacteria as multicellular individuals. (shrink)

In this paper, I argue that attempts in the philosophy of biology to show that organisms are processes rather than substances fail. Despite what process ontologists have said, I argue that substance ontology is perfectly able to accommodate the dynamic nature of organisms, their ecological dependence, and their vague boundaries, and that their criticisms are not directed at substance ontology simpliciter, but only at specific (perhaps untenable) characterisations of substances. The paper ends by considering what a processual philosophy of biology (...) that is radically in conflict with an ontology of substances might look like. (shrink)

The organism is one of the fundamental concepts of biology and has been at the center of many discussions about biological individuality, yet what exactly it is can be confusing. The definition that we find generally useful is that an organism is a unit in which all the subunits have evolved to be highly cooperative, with very little conflict. We focus on how often organisms evolve from two or more formerly independent organisms. Two canonical transitions of this type—replicators clustered in (...) cells and endosymbiotic organelles within host cells—demonstrate the reality of this kind of evolutionary transition and suggest conditions that can favor it. These conditions include co-transmission of the partners across generations and rules that strongly regulate and limit conflict, such as a fair meiosis. Recently, much attention has been given to associations of animals with microbes involved in their nutrition. These range from tight endosymbiotic associations like those between aphids and Buchnera bacteria, to the complex communities in animal intestines. Here, starting with a reflection about identity through time, we consider the distinctions between these kinds of animal–bacteria interactions and describe the criteria by which a few can be considered jointly organismal but most cannot. (shrink)

In this article, I address the question of what Developmental Systems Theory aims at explaining. I distinguish two lines of thought in DST, one that deals specifically with development and tries to explain the development of the individual organism, and the other that presents itself as a reconceptualization of evolution and tries to explain the evolution of populations of developmental systems. I emphasize that, despite the claim of the contrary by DST proponents, there are two very different definitions of the (...) “developmental system,” and therefore DST is not a unified theory of evolution and development. I show that DST loses the most interesting aspects of its reconceptualization of development when it tries to reconceptualize evolutionary theory. I suggest that DST is about development per se, and that it fails at offering a new view on evolution. (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)

Though viruses have generally been characterized by their pathogenic and more generally harmful effects, many examples of mutualistic viruses exist. Here I explain how the idea of mutualistic viruses has been defended in recent virology, and I explore four important conceptual and practical consequences of this idea. I ask to what extent this research modifies the way scientists might search for new viruses, our notion of how the host immune system interacts with microbes, the development of new therapeutic approaches, and, (...) finally, the role played by the criterion of autonomy in our understanding of living things. Overall, I suggest that the recognition of mutualistic viruses plays a major role in a wider ongoing revision of our conception of viruses. (shrink)

Criticisms of the “container” model of pregnancy picturing female and embryo as separate entities multiply in various philosophical and scientific contexts during the last decades. In this paper, we examine how this model underlies received views of pregnancy in evolutionary biology, in the characterization of the transition from oviparity to viviparity in mammals and in the selectionist explanations of pregnancy as an evolutionary strategy. In contrast, recent evo-devo studies on eutherian reproduction, including the role of inflammation and new maternal cell (...) types, gather evidence in favor of considering pregnancy as an evolved relational novelty. Our thesis is that from this perspective we can identify the emergence of a newhistoricalindividual in evolution. In evo-devo, historical units are conceptualized as evolved entities which fulfill two main criteria, their continuous persistence and their non-exchangeability. As pregnancy can be individuated in this way, we contend that pregnant females are historical individuals. We argue that historical individuality differs from, and coexists with, other views of biological individuality as applied to pregnancy (the physiological, the evolutionary and the ecological one), but brings forward an important new insight which might help dissolve misguided conceptions. (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)