The book presents a new way of understanding Darwinism and evolution by natural selection, combining work in biology, philosophy, and other fields.

Biological species have been treated traditionally as spatiotemporally unrestricted classes. If they are to perform the function which they do in the evolutionary process, they must be spatiotemporally localized individuals, historical entities. Reinterpreting biological species as historical entities solves several important anomalies in biology, in philosophy of biology, and within philosophy itself. It also has important implications for any attempt to present an "evolutionary" analysis of science and for sciences such as anthropology which are devoted to the study of single (...) species. (shrink)

The Limits of the Self, will be essential reading for anyone interested in the definition of biological individuality and the understanding of the immune system.

Biological theory demands a clear organism concept, but at present biologists cannot agree on one. They know that counting particular units, and not counting others, allows them to generate explanatory and predictive descriptions of evolutionary processes. Yet they lack a unified theory telling them which units to count. In this paper, I offer a novel account of biological individuality, which reconciles conflicting definitions of ‘organism’ by interpreting them as describing alternative realisers of a common functional role, and then defines individual (...) organisms as essentially possessing some mechanisms that play this role. (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)

Putative counterexamples to the Principle of Identity of Indiscernibles (PII) are notoriously inconclusive. I establish ground rules for debate in this area, offer a new response to such counterexamples for friends of the PII, but then argue that no response is entirely satisfactory. Finally, I undermine some positive arguments for PII.

Drawing on recent advances in evolutionary biology, prominent scholars return to the question posed in a pathbreaking book: how evolution itself evolved.

Our intuitive assumption that only organisms are the real individuals in the natural world is at odds with developments in cell biology, ecology, genetics, evolutionary biology, and other fields. Although organisms have served for centuries as nature’s paradigmatic individuals, science suggests that organisms are only one of the many ways in which the natural world could be organized. When living beings work together—as in ant colonies, beehives, and bacteria-metazoan symbiosis—new collective individuals can emerge. In this book, leading scholars consider the (...) biological and philosophical implications of the emergence of these new collective individuals from associations of living beings. The topics they consider range from metaphysical issues to biological research on natural selection, sociobiology, and symbiosis. -/- The contributors investigate individuality and its relationship to evolution and the specific concept of organism; the tension between group evolution and individual adaptation; and the structure of collective individuals and the extent to which they can be defined by the same concept of individuality. These new perspectives on evolved individuality should trigger important revisions to both philosophical and biological conceptions of the individual. -/- Contributors: Frédéric Bouchard, Ellen Clarke, Jennifer Fewell, Andrew Gardner, Peter Godfrey-Smith, Charles J. Goodnight, Matt Haber, Andrew Hamilton, Philippe Huneman, Samir Okasha, Thomas Pradeu, Scott Turner, Minus van Baalen. (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)

What makes a biological entity an individual? Jack Wilson shows that past philosophers have failed to explicate the conditions an entity must satisfy to be a living individual. He explores the reason for this failure and explains why we should limit ourselves to examples involving real organisms rather than thought experiments. This book explores and resolves paradoxes that arise when one applies past notions of individuality to biological examples beyond the conventional range and presents an analysis of identity and persistence. (...) The book's main purpose is to bring together two lines of research, theoretical biology and metaphysics, which have dealt with the same subject in isolation from one another. Wilson explains an alternative theory about biological individuality which solves problems which cannot be addressed by either field alone. He presents a more fine-grained vocabulary of individuation based on diverse kinds of living things, allowing him to clarify previously muddled disputes about individuality in biology. (shrink)

It is a common complaint that antireductionist arguments are primarily negative. Here I describe an alternative nonreductionist epistemology based on considerations taken from multidisciplinary research in biology. The core of this framework consists in seeing investigation as coordinated around sets of problems (problem agendas) that have associated criteria of explanatory adequacy. These ideas are developed in a case study, the explanation of evolutionary innovations and novelties, which demonstrates the applicability and fruitfulness of this nonreductionist epistemological perspective. This account also bears (...) on questions of conceptual change and theory structure in philosophy of science. †To contact the author, please write to: Department of Philosophy, University of Minnesota, 831 Heller Hall, 271 19th Ave. S., Minneapolis, MN 55455; e‐mail: aclove@umn.edu. (shrink)

The normal and the pathological are terms used for structures, activities, individual or collective situations proper to living beings and especially to man. The relation of a fact and a norm is its positive or negative value. Can the assessment of behaviours be reduced to noting a necessity? Is a living being's disease a fact similar to universal attraction? The author maintains that diseases are not merely predetermined effects, but are revealing of a normative regulation proper to living beings and (...) man. (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)

Individuals of many animal species are said to have a personality. It has been shown that some individuals are bolder than other individuals of the same species, or more sociable or more aggressive. In this paper, we analyse what it means to say that an animal has a personality. We clarify what an animal personality is, that is, its ontology, and how different personality concepts relate to each other, and we examine how personality traits are identified in biological practice. Our (...) analysis shows that biologists often study specific personality traits, such as boldness, sociability or aggressiveness, rather than personalities in general. We claim that personality traits are best understood as dispositions and that they are operationally defined in terms of certain sets of behaviours, which are studied in specific experimental set-ups. Furthermore, we develop an integrative philosophical account that specifies and formalises three criteria for identifying personality traits, which are used in biological practice. For an individual animal to have a personality trait it must, first, behave differently than others. Second, these behavioural differences must be stable over a certain time, and third, they must be consistent in different contexts. (shrink)

What are the conditions under which one biological object is a part of another biological object? This paper answers this question by developing a general, systematic account of biological parthood. I specify two criteria for biological parthood. Substantial Spatial Inclusionrequires biological parts to be spatially located inside or in the region that the natural boundary of t he biological whole occupies. Compositional Relevance captures the fact that a biological part engages in a biological process that must make a necessary contribution (...) to a condition that is minimally sufficient to one or more of the characteristic behaviors of the biological whole. Instead of emphasizing the diversity of part-whole relations in the biological world, this paper asks what biological part-whole relations have in common and what constrains their existence, in general. After presenting the two criteria for biological parthood I discuss in how far my account can cope with hard cases (e.g., redundant parts) and I reveal the merits and limits of monism. (shrink)

Two major clarifications have greatly abetted the understanding and fruitful expansion of the theory of natural selection in recent years: the acknowledgment that interactors, not replicators, constitute the causal unit of selection; and the recognition that interactors are Darwinian individuals, and that such individuals exist with potency at several levels of organization (genes, organisms, demes, and species in particular), thus engendering a rich hierarchical theory of selection in contrast with Darwin’s own emphasis on the organismic level. But a piece of (...) the argument has been missing, and individuals at levels distinct from organisms have been denied potency (although granted existence within the undeniable logic of the theory), because they do not achieve individuality with the same devices used by organisms and therefore seem weak by comparison. We show here that different features define Darwinian individuality across scales of size and time. In particular, species-individuals may develop few emergent features as direct adaptations. The interactor approach works with emergent fitnesses, not with emergent features; and species, as a consequence of their different mechanism for achieving individuality (reproductive exclusivity among subparts, that is, among organisms), express many effects from other levels. Organisms, by contrast, suppress upwardly cascading effects, because the organismic style of individuality (by functional integration of subparts) does not permit much competition or differential reproduction of parts from within. Species do not suppress the operation of lower levels; such effects therefore become available as exaptations conferring emergent fitness—a primary source of the different strength that species achieve as effective Darwinian individuals in evolution. (shrink)

Individuals are a prominent part of the biological world. Although biologists and philosophers of biology draw freely on the concept of an individual in articulating both widely accepted and more controversial claims, there has been little explicit work devoted to the biological notion of an individual itself. How should we think about biological individuals? What are the roles that biological individuals play in processes such as natural selection (are genes and groups also units of selection?), speciation (are species individuals?), and (...) organismic development (do genomes code for organisms)? Much of our discussion here will focus on organisms as a central kind of biological individual, and that discussion will raise broader questions about the nature of the biological world, for example, about its complexity, its organization, and its relation to human thought. (shrink)

This paper analyzes what it means for philosophy of science to be normative. It argues that normativity is a multifaceted phenomenon rather than a general feature that a philosophical theory either has or lacks. It analyzes the normativity of philosophy of science by articulating three ways in which a philosophical theory can be normative. Methodological normativity arises from normative assumptions that philosophers make when they select, interpret, evaluate, and mutually adjust relevant empirical information, on which they base their philosophical theories. (...) Object normativity emerges from the fact that the object of philosophical theorizing can itself be normative, such as when philosophers discuss epistemic norms in science. Metanormativity arises from the kind of claims that a philosophical theory contains, such as normative claims about science as it should be. Distinguishing these three kinds of normativity gives rise to a nuanced and illuminating view of how philosophy of science can be normative. (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)

Although natural philosophers have long been interested in individuality, it has been of interest to contemporary philosophers of biology because of its role in different aspects of evolutionary biology. These debates include whether species are individuals or classes, what counts as a unit of selection, and how transitions in individuality occur evolutionarily. Philosophical analyses are often conducted in terms of metaphysics (“what is an individual?”), rather than epistemology (“how can and do researchers conceptualize individuals so as to address some of (...) their scientific goals?”). We review several philosophical distinctions in order to shift attention from metaphysics to epistemology. Many controversies involve epistemological differences rather than metaphysical disagreement. This implies that a pluralist stance about individuality in biology is warranted and has metaphysical consequences because the pluralism emerges from the diversity of scientific interests that investigate the complexity of living phenomena. (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)

Following a previous elaboration of the concept of weak individuality and some examples of its instances in ecology and biology, the article focuses on general features of the concept, arguing that in any ontological field individuals are understood on the basis of our knowledge of interactions, through the application of these general formulas for extracting individuals from interactions. Then, the specificities of the individuality in the sense of this weak concept are examined in ecology; I conclude by addressing the differences (...) between ecosystems and organisms as they appear in the viewpoint of such concept. (shrink)

The concept of genidentity has been proposed as a way to better understand identity through time, especially in physics and biology. The genidentity view is utterly anti-substantialist in so far as it suggests that the identity of X through time does not presuppose whatsoever the existence of a permanent “core” or “substrate” of X. Yet applications of this concept to real science have been scarce and unsatisfying. In this paper, our aim is to show that a well-defined concept of functional (...) genidentity can be crucial to shed light on identity through time in classical physics and especially in biology. Finally, we show that understanding identity on the basis of continuity suggests a move towards an ontology of processes. (shrink)

No comprehensive theory of development is available yet. Traditionally, we regard the development of animals as a sequence of changes through which an adult multicellular animal is produced, starting from a single cell which is usually a fertilized egg, through increasingly complex stages. However, many phenomena that would not qualify as developmental according to these criteria would nevertheless qualify as developmental in that they imply nontrivial (e.g., non degenerative) changes of form, and/or substantial changes in gene expression. A broad, comparative (...) approach is badly needed. In the Cnidaria, for example, even the boundary between generations is problematic. Describing their life cycle in terms of metagenesis (alternation between polyp generation and medusa generation) or in terms of metamorphosis (polyp as larva or juvenile) are matters of semantics more than biology. The life cycle of other metazoans, described in textbooks in terms of larva-to-adult metamorphosis, is hardly different from a typical metagenetic life cycle of cnidarians. This applies to holometabolous insects and to marine invertebrates like sea urchins, where most of the larval cells are discarded at metamorphosis. The uncertain temporal and spatial boundaries of individual development are also shown by the widespread lack of a strict correspondence between adult and mature. A comprehensive theory of development should start with a zero principle of “developmental inertia,” corresponding to an indeterminate local self-perpetuation of cell-level dynamics. Indeterminate growth, scale-invariance, segmentation, and regeneration provide examples of developmental dynamics close to that. (shrink)

Two problems related to the biological identity of living beings are faced: the who-problem (which are the biological properties making that living being unique and different from the others?); the persistence-problem (what does it take for a living being to persist from a time to another?). They are discussed inside a molecular biology framework, which shows how epigenetics can be a good ground to provide plausible answers. That is, we propose an empirical solution to the who-problem and to the persistence-problem (...) on the basis of the new perspectives opened by a molecular understanding of epigenetic processes. In particular, concerning the former, we argue that any living being is the result of the epigenetic processes that have regulated the expression of its genome; concerning the latter, we defend the idea that the criterion for the persistence of its identity is to be indicated in the continuity of those epigenetic processes. We also counteract possible objections, in particular (1) whether our approach has something to say at a metaphysical level; (2) how it could account for the passage from the two phenotypes of the parental gametes to the single phenotype of the zygote; (3) how it could account for the identity of derivatives of one living being that continue to live disjoined from that original living being; (4) how it could account for higher mental functions. (shrink)

An individual, as this term will be understood here, is an entity to which the concepts of unity and identity fully and determinately apply. That is to say, an entity x is an individual just in case x determinately counts as one entity and x has a determinate identity. Many philosophers tacitly assume that all entities are individuals in the foregoing sense, and indeed that it is a necessary truth that they are. But this can certainly be disputed. It is, (...) very arguably, both logically and metaphysically possible for there to be nonindividuals. The aim of this chapter is to clarify and explore both the notion of an individual and that of a nonindividual, to propose a typology of entities based on the individual/nonindividual distinction, and to illustrate the potential of these notions for application within and across the sciences. (shrink)

This paper does not intend to provide an exhaustive account of Canguilhem's thinking. It will focus on his philosophical approach to the biological sciences.

Michael Ghiselin and David Hull’s individuality thesis is that biological species are individuals. Philosophers often treat the term “individual” as synonymous with “mereological sum” and characterize it in terms of mereology. It is easy to see how the biological project has been interpreted as a mereological one. This chapter argues that this is a mistake, that biological part/whole relations often violate the axioms of mereology. Conflating these projects confuses the central issues at stake in both, and makes the job of (...) evaluating either unnecessarily burdensome. Clarifying this helps identify the genuine metaphysical implications of the individuality thesis, which serves as an exemplar of scientifically informed metaphysics. (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.

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.

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)

Why is there a specific problem with biological individuality? Because the living realm contains a wide range of exotic particular concrete entities that do not easily match our ordinary concept of an individual. Slime moulds, dandelions, siphonophores are among the Odd Entities that excite the ontological zeal of the philosophers of biology. Most of these philosophers, however, seem to believe that these Odd Cases oblige us to refine or revise our common concept of an individual. They think, explicitly or tacitly, (...) that to be a living, evolutionary entity is to be a living individual. In this paper, we explore an alternative proposal: the variety and oddity of the forms of the living realm might be ontologically regimented through an increase in the categorial complexity of the living realm, by admitting, beside living individuals, living non-individuals or by acknowledging, more generally, that the evolutionary development of the living forms is not necessarily a process of building individuals, that life is not necessarily individuals-oriented. We claim that, from an ontological point of view, the spectacle of the living realm obliges us to take aggregativity seriously. (shrink)

One of the most remarkable features of mid-to-late twentieth century analytic philosophy is the revival of interest in the traditional problems of metaphysics. However, given the long period of neglect from which metaphysics in only now emerging, it is perhaps not surprising that philosophers in the analytic tradition are still finding their metaphysical feet, particularly on meta-metaphysical and methodological matters. Thus contemporary metaphysicians find themselves in an exciting but ultimately unstable position: We are convinced that metaphysical questions are worth pursing, (...) but we still lack a convincing story to offer our detractors about how we conduct our business qua metaphysicians. This essay is an attempt to provide such a story. The core recommendation is that we need not re-invent the discipline out of whole cloth if we are willing to learn from the past masters, particularly the Scholastics. What ultimately emerges is a defense of Scholastic metaphysical principles based on contemporary evolutionary biology. (shrink)

New mechanistic philosophy has not examined explanations in ecology although they are based extensively on describing mechanisms responsible for phenomena under scrutiny. This chapter uses the example of research on the shrub Lonicera maackii (Amur honeysuckle) to scrutinize individual-level mechanisms that are generally accepted and used in ecology and confronts them with the minimal account of mechanisms. Individual-level mechanisms are for a phenomenon, are hierarchical, and absent entities play a role in their functioning. They are distinguished by the role played (...) by properties in determining activities and organization. The chapter also considers the experimental methods for discovery of individual-level mechanisms, the possibility of group-level mechanisms in ecology, and suggests further research problems. (shrink)

The question What is an individual? goes back beyond Aristotle’s discussion of substance to the Ionians’ preoccupation with the paradox of change -- the fact that if anything changes it must stay the same. Mere reflection on this fact and the common-sense notion of a countable thing yields a concept of a “minimal individual”, which is particular (a logical matter) specific (a taxonomic matter), and unique (an evaluative empirical matter). Individuals occupy space, and therefore might be dislodged. Even minimal individuals, (...) therefore (Strawsonian individual or Aristotelian substance) already contain the potential for competition or conflict. What is added by biology to this basic notion? It emerges from some recent work on the evolution of metazoan animals that individuals as we know them are minimal individuals towhich four features have been added, and which appear to be inseparable: differentiated multicellularity; sexual reproduction; segregation of germ from somatic cells; and obligatory death. Whether or not individuals are to be counted as units of selection, they are not the beneficiaries of natural selection. (shrink)

Overview The evolution of multicellularity raises questions regarding genomic and developmental commonalities and discordances, selective advantages and disadvantages, physical determinants of development, and the origins of morphological novelties. It also represents a change in the definition of individuality, because a new organism emerges from interactions among single cells. This volume considers these and other questions, with contributions that explore the origins and consequences of the evolution of multicellularity, addressing a range of topics, organisms, and experimental protocols. Each section focuses on (...) selected topics or particular lineages that present a significant insight or challenge. The contributors consider the fossil record of the paleontological circumstances in which animal multicellularity evolved; cooptation, recurrent patterns, modularity, and plausible pathways for multicellular evolution in plants; theoretical approaches to the amoebozoa and fungi (cellular slime molds having long provided a robust model system for exploring the evolution of multicellularity), plants, and animals; genomic toolkits of metazoan multicellularity; and philosophical aspects of the meaning of individuality in light of multicellular evolution. Contributors Maja Adamska, Argyris Arnellos, Juan A. Arias, Eugenio Azpeitia, Mariana Benítez, Adriano Bonforti, John Tyler Bonner, Peter L. Conlin, A. Keith Dunker, Salva Duran-Nebreda, Ana E. Escalante, Valeria Hernández-Hernández, Kunihiko Kaneko, Andrew H. Knoll, Stephan G. König, Daniel J. G. Lahr, Ottoline Leyser, Alan C. Love, Raul Montañez, Emilio Mora van Cauwelaert, Alvaro Moreno, Vidyanand Nanjundiah, Aurora M. Nedelcu, Stuart A. Newman, Karl J. Niklas, William C. Ratcliff, Iñaki Ruiz-Trillo, Ricard Solé . (shrink)

The notion of an individual and the related issues on individuation are topics that appear in the philosophical discussion ever since the antiquity. The idea of an individual thing is intuitively clear: an individual is something of a specific kind that is a unity, having its own identity, and being so that it is possible at least in principle to discern it from any other individual, even of similar species. But when we try to leave the intuitive realm and push (...) this idea to a logical analysis, we find a cluster of problems that are difficult to overcome within standard logico-mathematical contexts. In this work, we shall be concerned with some aspects of this intuitive concept of an individual and on some related facts about individuation taken from recent discussions that arose ever since the inception of quantum theory, pushing the discussion to a “logical” view, which in our opinion is still lacking in the usual debates on such issues. In the final part of the paper, we propose a metaphysics where the notion of identity is substituted, for some objects, by a weaker notion of indiscernibility, and we try to justify such a move. In most of the uses of the expression “quantum theory”, we shall not make explicit the distinction between the non-relativistic and the relativistic approaches, although they of course are quite different, for we think that the problems as we shall present them appear in both versions. But, as the text goes, the context will distinguish between them and these questions will become clear to the reader. (shrink)

Abstract“Development” suggests that there is something that is developing, or changing over time. We can ask about temporal boundaries of that developmental process, asking when development begins or ends and whether it has defined stages along the way, for example. We can ask about spatial boundaries as well: where does the developing object start and end? For this article, I ask about the boundary definition of the developing organism in particular. What is an individual organism, and what defines it as (...) the same organism as it changes over time? In particular, how has this been answered historically: how have researchers described and explained what an individual developing organism is? This article explores ideas and approaches especially starting in the late 19th century, and in particular looks at the role of “organization.”. (shrink)

Many aspects of biology, such as population genetics and senescence, are predicated on identifying individuals and generations. Conventional demarcations of individuals and generations, such as physiological autonomy, unicellular bottlenecks, and alternation of generation, are rife with problems. Do physically separated cuttings or plant ramets constitute separate individuals or generations? Are chimaeras one or more individuals? To resolve these problems, Clarke : 321–361, 2012) proposed that individuals are circumscribed by mechanisms that constrain heritable variance in fitness. Simultaneously, Gorelick and Heng : (...) 1088–1098 2011) showed that sex constrains heritable variance Therefore, for eukaryotes, meiosis and karyogamy provide a consistent way to demarcate individuals and generations. Epigenetic reset associated with meiosis and karyogamy rejuvenates the next generation, but not the parent that engaged in the sex act. Wholesale epigenetic resets that probably only occur with meiosis and karyogamy imply that monozygotic twins are two different individuals, but apomictic progeny are diffuse parts of one disaggregated individual. Mitotic heritability circumscribes an individual, whereas meiotic heritability demarcates new individuals and generations. (shrink)