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)

I argue that the propensity interpretation of fitness, properly understood, not only solves the explanatory circularity problem and the mismatch problem, but can also withstand the Pandora’s box full of problems that have been thrown at it. Fitness is the propensity (i.e., probabilistic ability, based on heritable physical traits) for organisms or types of organisms to survive and reproduce in particular environments and in particular populations for a specified number of generations; if greater than one generation, “reproduction” includes descendants of (...) descendants. Fitness values can be described in terms of distributions of propensities to produce varying number of offspring and can be modeled for any number of generations using computer simulations, thus providing both predictive power and a means for comparing the fitness of different phenotypes. Fitness is a causal concept, most notably at the population level, where fitness differences are causally responsible for differences in reproductive success. Relative fitness is ultimately what matters for natural selection. (shrink)

The “land community” (or “biotic community”) that features centrally in Aldo Leopold’s Land Ethic has typically been equated with the concept of “ecosystem.” Moreover, some have challenged this central Leopoldean concept given the multitude of meanings of the term “ecosystem” and the changes the term has undergone since Leopold’s time (see, e.g., Shrader-Frechette 1996). Even one of Leopold’s primary defenders, J. Baird Callicott, asserts that there are difficulties in identifying the boundaries of ecosystems and suggests that we recognize that their (...) boundaries are determined by scientific questions ecologists pose (Callicott 2013). I argue that we need to rethink Leopold’s concept of land community in the following ways. First, we should recognize that Leopold’s views are not identical to those of his contemporaries (e.g., Clements, Elton), although they resemble those of some subsequent ecologists, including some of our contemporaries (e.g., O’Neill 2001, Post et al. 2007, Hastings and Gross 2012). Second, the land community concept does not map cleanly onto the concept of “ecosystem”; it also incorporates elements of the “community” concept in community ecology by emphasizing the interactions between organisms and not just the matter/energy flow of the ecosystem concept. Third, the boundary question can be illuminated by considering some of the recent literature on the nature of biological individuals (in particular, Odenbaugh 2007; Hamilton, Smith, and Haber 2009; Millstein 2009), focusing on concentrations of causal relations as determinative of the boundaries of the land community qua individual. There are challenges to be worked out, particularly when the interactions of community members do not map cleanly onto matter/energy flows, but I argue that these challenges can be resolved. The result is a defensible land community concept that is ontologically robust enough to be a locus of moral obligation while being consistent with contemporary ecological theory and practice. (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)

Essay review of 'Darwinian Populations and Natural Selection'.

Scientists use models to understand the natural world, and it is important not to conflate model and nature. As an illustration, we distinguish three different kinds of populations in studies of ecology and evolution: theoretical, laboratory, and natural populations, exemplified by the work of R.A. Fisher, Thomas Park, and David Lack, respectively. Biologists are rightly concerned with all three types of populations. We examine the interplay between these different kinds of populations, and their pertinent models, in three examples: the notion (...) of “effective” population size, the work of Thomas Park on /Tribolium/ populations, and model-based clustering algorithms such as /Structure/. Finally, we discuss ways to move safely between three distinct population types while avoiding confusing models and reality. (shrink)

In a large and impressive body of published work, Quayshawn Spencer has meticulously articulated and defended a metaphysical project aimed at resuscitating a biological conception of race—one free from many of the pitfalls of biological essentialism. If successful, such a project would be highly rewarding, since it would provide a compelling response to philosophers who have denied the genuine existence of race while avoiding the very dangers that they sought to avoid. The aim of this paper is to subject those (...) moves to careful scrutiny and thereby appraise the prospects for a new biologism about race. (shrink)

Selectionist evolutionary theory has often been faulted for not making novel predictions that are surprising, risky, and correct. I argue that it in fact exhibits the theoretical virtue of predictive capacity in addition to two other virtues: explanatory unification and model fitting. Two case studies show the predictive capacity of selectionist evolutionary theory: parallel evolutionary change in E. coli, and the origin of eukaryotic cells through endosymbiosis.

This year’s topic is “Genomics and Philosophy of Race.” Different researchers might work on distinct subsets of the six thematic clusters below, which are neither mutually exclusive nor collectively exhaustive: (1) Concepts of ‘Race’; (2) Mathematical Modeling of Human History and Population Structure; (3) Data and Technologies of Human Genomics; (4) Biological Reality of Race; (5) Racialized Selves in a Global Context; (6) Pragmatic Consequences of ‘Race Talk’ among Biologists.

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

I defend a radical interpretation of biological populations—what I call population pluralism—which holds that there are many ways that a particular grouping of individuals can be related such that the grouping satisfies the conditions necessary for those individuals to evolve together. More constraining accounts of biological populations face empirical counter-examples and conceptual difficulties. One of the most intuitive and frequently employed conditions, causal connectivity—itself beset with numerous difficulties—is best construed by considering the relevant causal relations as ‘thick’ causal concepts. I (...) argue that the fine-grained causal relations that could constitute membership in a biological population are huge in number and many are manifested by degree, and thus we can construe population membership as being defined by massively multidimensional constructs, the differences between which are largely arbitrary. I end by showing that positions in two recent debates in theoretical biology depend on a view of biological populations at odds with the pluralism defended here. 1 Introduction2 Biological Population, Broad and Narrow3 Difficulties with Narrow Biological Population Conditions3.1 Against the genealogical condition3.2 Against the conspecificity condition3.3 Against the proximity condition3.4 Against the typology condition4 Causal Connectivity5 Massively Multidimensional Population Constructs6 Population Uniqueness and Natural Selection6.1 Statisticalism and its discontents6.2 Price at what price?7 Conclusion. (shrink)

Millstein (2009) argues against conceptual pluralism with respect to the definition of “population,” and proposes her own definition of the term. I challenge both Millstein's negative arguments against conceptual pluralism and her positive proposal for a singular definition of population. The concept of population, I argue, does not refer to a natural kind; populations are constructs of biologists variably defined by contexts of inquiry.

Millstein argues against conceptual pluralism with respect to the definition of “population,” and proposes her own definition of the term. I challenge both Millstein’s negative arguments against conceptual pluralism and her positive proposal for a singular definition of population. The concept of population, I argue, does not refer to a natural kind; popula tions are constructs of biologists variably defined by contexts of inquiry.

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)

In this article I show that population geneticists are acknowledging a kind of biological population that has hitherto been unappreciated by philosophers. The new population talk occurs when population geneticists call continent-level human genetic clusters ‘populations’ in population structure research. My theory is that the kind of population being referred to is the K population, which is, roughly, a biological population whose members are united by common genomic ancestry and in which population membership is graded. After presenting and defending the (...) theory, I show that the K population is indeed a kind of biological population. Finally, I address likely objections. (shrink)

In the early 2000s, Esteban Burchard and his colleagues defended a controversial route to the view that there’s a racial classification of people that’s useful in medicine. The route, which I call ‘Burchard’s route,’ is arguing that there’s a racial classification of people that’s useful in medicine because, roughly, there’s a racial classification with medically relevant genetic differentiation :1170–1175, 2003). While almost all scholars engaged in this debate agree that there’s a racial classification of people that’s useful in medicine in (...) some way, there’s tremendous controversy over whether any racial scheme is useful in medicine because there are medically relevant genetic differences among those races : 564–565, 2016). The goal of this paper will be to show that Burchard’s route is basically correct. However, I will use a slightly different argument than Burchard et al.’s in order to provide a firmer foundation for the thesis, both metaphysically and genetically. I begin by reviewing Burchard’s route and its critics. Second, I present an original argument for establishing Burchard et al.’s conclusion using a Burchard-like route. I call it ‘Spencer’s route’. I reply to major objections along the way, and I end with a summary. (shrink)

Individuals come into the world with talents that differ greatly in their marketability. These talents command a wide variety of rents; from millions of dollars per film for a movie star, to just a few thousands of dollars per annum for a fast-food joint worker. How should we distribute income fairly given these disparities? That is the topic of Kristi Olson’s excellent new book, The Solidarity Solution: Principles for a Fair Income Distribution. 1 1 The question is not new, but (...) Olson’s treatment is among the most comprehensive and well-argued one can find. The manuscript is well researched, scholarly, thorough, packed full of arguments and low on fluff. It is well worth the time of anyone interested in fairness in the sphere of work and income. (shrink)

1. IntroductionIn What Biological Functions Are and Why They Matter, Justin Garson offers a novel theory of biological functions, the generalized selected effects (GSE) theory.1 He presents the theory in a clear and comprehensive way, defends it against various objections and applies it to different areas of philosophy, including the philosophy of psychiatry, the debate about mechanisms and the debate about teleosemantic theories of mental content.2Like other proponents of the aetiological approach to functions, Garson maintains that a trait’s biological functions (...) are grounded in its selectional history. However, his GSE theory differs from standard versions of the aetiological approach – so-called selected effects (SE) theories – in a crucial respect: while SE theories assume that all biological functions must ultimately be grounded in phylogenetic processes of natural selection or other (closely analogous) processes of differential reproduction (Millikan 1984: 15–82, Neander 1991, Godfrey-Smith 1994), the GSE theory allows that functions may also be grounded in processes of differential retention (see §2 for details). Garson forcefully argues that this broader account is theoretically well-motivated and has significant advantages over standard SE accounts when it comes to accommodating certain important classes of ontogenetically determined functions. (shrink)

Two recently popular metaphilosophical movements, formal philosophy and experimental philosophy, promote what seem to be conflicting methodologies. Nonetheless, I argue that the two can be mutually supportive. I propose an experimentally-informed variation on explication, a powerful formal philosophical tool introduced by Carnap. The resulting method, which I call “experimental explication,” provides the formalist with a means of responding to explication's gravest criticism. Moreover, this method introduces a philosophically salient, positive role for survey-style experiments while steering clear of several objections that (...) critics of “positive experimental philosophy” raise. Thus, it provides the experimentalist with a more defensible example of how empirical work can have positive philosophical import. For these reasons, experimental explication should appeal to experimental philosophers and formal philosophers alike. (shrink)

While evolutionary thinking is increasingly becoming popular in fields of investigation outside the biological sciences, it remains unclear how helpful it is there and whether it actually yields good explanations of the phenomena under study. Here we examine the ontology of a recent approach to applying evolutionary thinking outside biology, the generalized Darwinism approach proposed by Geoffrey Hodgson and Thorbjørn Knudsen. We examine the ontology of populations in biology and in GD, and argue that biological evolutionary theory sets ontological criteria (...) that GD fails to meet. We suggest two options to revise the population concept in GD: reformulating the concept in terms of inheritance and reproduction such that it comes to pick out individuals similar to evolving populations, or trying to build an adequate population concept on a principle of differential retention instead of differential reproduction. 1 Introduction2 Generalized Darwinism2.1 What is generalized Darwinism?2.2 Darwinian principles3 The Ontology of Generalized Darwinism: What Are Populations?3.1 The population concept of generalized Darwinism3.2 The population concept in evolutionary theory4 Locating Evolving Systems in Generalized Darwinism5 Conclusion. (shrink)

This paper analyzes recent attempts to reject reproduction with lineage formation as a necessary condition for evolution by means of natural selection :560–570, 2008; Stud Hist Philos Sci Part C Stud Hist Philos Biol Biomed Sci 42:106–114, 2011; Bourrat in Biol Philos 29:517–538, 2014; Br J Philos Sci 66:883–903, 2015; Charbonneau in Philos Sci 81:727–740, 2014; Doolittle and Inkpen in Proc Natl Acad Sci 115:4006–4014, 2018). Building on the strengths of these attempts and avoiding their pitfalls, it is argued that (...) a robust formulation of evolution by natural selection without reproduction can be established. The main contribution of this paper is a reformulation of Lewontin’s three principles stating that minimal evolution by natural selection occurs when two conditions are met in a population: fitness-related variation and memory. Paradigmatic evolution by natural selection, which can generate adaptations, takes place when an additional condition is present, namely regeneration. (shrink)

Philosophers are approaching a consensus that biological individuality, including evolutionary individuality, comes in degrees. Graded evolutionary individuality presents a puzzle when juxtaposed with another widely embraced view: that evolutionary individuality follows from being a selectable member of a Darwinian population. Population membership is, on the orthodox view, a bivalent condition, so how can members of Darwinian populations vary in their degree of individuality? This article offers a solution to the puzzle, by locating difference in degree of evolutionary individuality at the (...) level of population lineages, some of which are more Darwinian than others. In doing so, it sheds light on graded individuality in overlapping and nested population lineages, such as those that arise in multilevel selection and symbiotic collectives. (shrink)

Biologists and philosophers have offered differing concepts of biological race. That is, they have offered different candidates for what a biological correlate of race might be; for example, races might be subspecies, clades, lineages, ecotypes, or genetic clusters. One thing that is striking about each of these proposals is that they all depend on a concept of population. Indeed, some authors have explicitly characterized races in terms of populations. However, including the concept of population into concepts of race raises three (...) puzzles, all having to do with time. In this paper, I extend the causal interactionist population concept (CIPC) by introducing some simple assumptions about how to understand populations through time. These assumptions help to shed light on the three puzzles, and in the process show that if we want to understand races in terms of populations, we will need to revise our concept(s) of race. (shrink)

In “‘Population’ is Not a Natural Kind of Kinds,” Jacob Stegenga argues against the claim that the concept of “population” is a natural kind and in favor of conceptual pluralism, ostensibly in response to two papers of mine (Millstein 2009, 2010). Pluralism is often an attractive position in the philosophy of science. It certainly is a live possibility for the concept of population in ecology and evolutionary biology, and I welcome the opportunity to discuss the topic further. However, I argue (...) that the case for conceptual pluralism has not yet been made. In what follows, I first clarify the issues at stake before taking up the topic of conceptual pluralism and responding to Stegenga’s criticisms of the causal interactionist population concept. (shrink)

This paper presents an account of the biological populations that can undergo paradigmatic natural selection. I argue for, and develop Peter Godfrey-Smith’s claim that reproductive competition is a core attribute of such populations. However, as Godfrey-Smith notes, it is not the only important attribute. I suggest what the missing element is, co-opting elements of Alan Templeton’s notion of exchangeability. The final framework is then compared to two recent discussions regarding biological populations proposed by Roberta Millstein and Jacob Stegenga.

Natural selection comes in degrees. Some biological traits are subjected to stronger selective force than others, selection on particular traits waxes and wanes over time, and some groups can only undergo an attenuated kind of selective process. This has downstream consequences for any notions that are standardly treated as binary but depend on natural selection. For instance, the proper function of a biological structure can be defined as what caused that structure to be retained by natural selection in the past. (...) We usually think of proper functions in binary terms: storing bile is a function of the gall bladder, but making stones is not. However, if functions arise through natural selection, and natural selection comes in degrees, then a binary approach to proper functions is in tension with the biological facts. In order to resolve this tension, we need to revise our standard accounts of proper function. In particular, we may have to seriously consider the possibility that functions themselves come in degrees, in spite of the ramifications this will have for the way we speak about functions and related concepts such as dysfunction, disease, and teleosemantic content. (shrink)

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)

Methodological individualism has a long, successful, and controversial track record in the social sciences. Its record in ecology is much shorter but proving as successful and controversial with so-called individual-based models. Distinctions and debates about methodological individualism in social sciences clarify the commitments of this general, individualistic approach to modeling ecological phenomena and show that there is a lot recommending it. In particular, a representational priority on individual organisms yields a cogent albeit deflationary account of ecological emergence and helps reveal (...) how quite disparate models and theories in ecology might be unified. (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)

In this article, I consider the term “environment” in various claims and models by evolutionists and ecologists. I ask whether “environment” is amenable to a philosophical explication, in the same way some key terms of evolutionary theorizing such as “fitness,” “species,” or more recently “population” have been. I will claim that it cannot. In the first section, I propose a typology of theoretical terms, according to whether they are univocal or equivocal, and whether they have been the object of formal (...) or conceptual attempts of clarification. “Environment” will appear to be in a position similar to “population,” yet almost no extant attempt has been made to make sense of its meaning and reference. In the second section, I will present several theoretical claims or issues that refer to “environment” in apparently very diverse ways, but always supposing a contrastive term, which is not always the same. The third section directly considers models in evolution and in ecology, and asks “where” in them is the environment term. The fourth section proposes that “environment” refers to a distinction between a varying and an invariant term, but shows that this does not exhaust its meaning, which requires making more conceptual differences. Then, I take on the suggestion that there might be two “environment” terms, one in ecology and one in evolution, but show that is not the case. Finally I center on the specific notion of “complex environment,” which is the object of several research programs, and propose a typology of “complex environments” across theories. (shrink)

What is it for an individual thing in the natural world—a rock, a mouse, a family or a planet—to be made of other things—crystals, organs, animals, soil, water, or dirt? Rocks, mice, families and planets are composites, but how are we to understand the relation that holds between these composites and their component parts? My aim is to offer a new account of this relation, which I shall call corporeal composition. A central claim of my account is that corporeal composition (...) is grounded in causal relations between components. I will show how this claim, combined with a minimalist approach to causation—that causation is not one thing, but many—entails a position I call compositional minimalism—that composition is not one thing, but many. (shrink)

I criticize some arguments against the causal interpretability of population genetics put forward by Denis Walsh ([2007], [2010]). In particular, I seek to undermine the contention that population genetics exhibits frame of reference relativity or subjectivity with respect to its formal representations. I also show that classical population genetics does not fall foul of some criteria for causal representation put forward by James Woodward ([2003]), although those criteria do undermine some causalist stances. 1 Introduction2 Modularity3 The Crucially Important Point4 The (...) Gillespie Case: Density-Dependent Selection5 Conclusion. (shrink)

Mainstream teleosemantics is the view that mental representation should be understood in terms of biological functions, which, in turn, should be understood in terms of selection processes. One of the traditional criticisms of teleosemantics is the problem of novel contents: how can teleosemantics explain our ability to represent properties that are evolutionarily novel? In response, some have argued that by generalizing the notion of a selection process to include phenomena such as operant conditioning, and the neural selection that underlies it, (...) we can resolve this problem. Here, we do four things: we develop this suggestion in a rigorous way through a simple example, we draw on recent neurobiological research to support its empirical plausibility, we defend the move from a host of objections in the literature, and we sketch how the picture can be extended to help us think about more complex “conceptual” representations and not just perceptual ones. (shrink)

A common misunderstanding of the selected effects theory of function is that natural selection operating over an evolutionary time scale is the only functionbestowing process in the natural world. This construal of the selected effects theory conflicts with the existence and ubiquity of neurobiological functions that are evolutionary novel, such as structures underlying reading ability. This conflict has suggested to some that, while the selected effects theory may be relevant to some areas of evolutionary biology, its relevance to neuroscience is (...) marginal. This line of reasoning, however, neglects the fact that synapses, entire neurons, and potentially groups of neurons can undergo a type of selection analogous to natural selection operating over an evolutionary time scale. In the following, I argue that neural selection should be construed, by the selected effect theorist, as a distinct type of function-bestowing process in addition to natural selection. After explicating a generalized selected effects theory of function and distinguishing it from similar attempts to extend the selected effects theory, I do four things. First, I show how it allows one to identify neural selection as a distinct function-bestowing process, in contrast to other forms of neural structure formation such as neural construction. Second, I defend the view from one major criticism, and in so doing I clarify the content of the view. Third, I examine drug addiction to show the potential relevance of neural selection to neuroscientific and psychological research. Finally, I endorse a modest pluralism of function concepts within biology. (shrink)

Philosophers who study the problem of biological function often begin their deliberations by reflecting on the functions of parts of animals, or the behavior of animals. Applying theories of biological function to unconventional or borderline cases can help us to better evaluate and refine those theories. This is the case when we consider whether parts of transposable elements —bits of “selfish” DNA that move about within a host genome—have functions of their own, that is, whether the parts of TEs have (...) the function of helping the TE move about within the genome. Here I argue that whether or not the parts of TEs have functions depends crucially on whether collections of TEs form “populations,” by which I mean, here, a group of individuals of the same type that impact one another’s chances of persistence or multiplication, by impacting one another’s access to a shared resource. I think there is suggestive, but not conclusive, evidence that some TEs have functions of their own. Considering the problem of TE functionality, then, has value both for philosophy and for biology. (shrink)

I present and defend the generalized selected effects theory (GSE) of function. According to GSE, the function of a trait consists in the activity that contributed to its bearer’s differential reproduction, or differential retention, within a population. Unlike the traditional selected effects (SE) theory, it does not require that the functional trait helped its bearer reproduce; differential retention is enough. Although the core theory has been presented previously, I go significantly beyond those presentations by providing a new argument for GSE (...) and defending it from a recent objection. I also sketch its implications for teleosemantics and philosophy of medicine. (shrink)

As the debate about holism and reductionism in ecology has ebbed in the last twenty years, this article aims to reassess the traditional opposition between holistic and reductionist epistemologies during the development of population biology. The history of the notion of carrying capacity, the upper demographic limit of a viable population, will be analyzed as a paradigmatic case of the progressive imposition of reductionist strategies, from both an epistemological and a semantic point of view, since the middle of the twentieth (...) century. Then, Richard Looijen’s reduction of the carrying capacity concept to the niche partitioning theory will be assessed and rebuked for both empirical and logical reasons. Eventually, some recent “weak” and “hard” emergent conceptualizations of the notion of carrying capacity, in logistic map models or in coupled niche-population systems, will be presented in order to show how they call into question the nature and the use of the notion of carrying capacity as a predefined ecological limit. (shrink)

For a population to undergo evolution by natural selection, it is assumed that the constituents of the population form parent-offspring lineages, that is, that they must reproduce. I challenge this assumption by dividing the notion of reproduction into two subprocesses, that is, multiplication and inheritance, that produce parent-offspring lineages between the parts of a population, and I show that their population-level roles, generation and memory, respectively, can be effected by processes that do not rely on such local-level lineages. I further (...) argue that these two population-level processes, not local parent-offspring lineages, are necessary conditions for a population to undergo Darwinian evolution. (shrink)

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

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)

Darwinian evolution is a population-level phenomenon. This paper deals with a structural population concept within the framework of generalized Darwinism, resp. within a generalized theory of evolution. According to some skeptical authors, GD is in need of a valid population concept in order to become a practicable research program. Populations are crucial and basic elements of any evolutionary explanation—biological or cultural—and have to be defined as clearly as possible. I suggest the “causal interactionist population concept”, by R. Millstein for this (...) purpose, and I will try to embed the approach into a generalized evolutionary perspective by mathematically formalizing its key definitions. Using graph-theory, populations as described in the CIPC can serve as proper clusters of evolutionary classification based on the rates of interactions between their elements. I will introduce the concept of a cohesion index as a measurement of possible population candidates within a distribution of elements. The strength of this approach lies in its applicability and interactions are relatively easy to observe. Furthermore, problems of clustering tokens via typicality, e.g. their similarity in intrinsic key characteristics, can be avoided, because CIPC is a external approach. However, some formal problems and conceptual ambiguities occur within a simple version of this CI, which will be addressed in this paper as well as some possible applications. (shrink)

In this article I argue that we should pay extra attention to the ecological dimension of natural selection. By this I mean that we should view natural selection primarily as acting on the outcomes of the interactions organisms have with their environment, which influences their relative reproductive output. A consequence of this view is that natural selection is not sensitive to what system of inheritance ensures reoccurrences of organism-environment interactions over generations. I end by showing the consequences of this view (...) when looking at how processes like niche construction and the Baldwin effect relate to natural selection. (shrink)