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)

E. O. Wilson (1974: 54) describes the problem that social organisms pose: “On what bases do we distinguish the extremely modified members of an invertebrate colony from the organs of a metazoan animal?” This framing of the issue has inspired many to look more closely at how groups of organisms form and behave as emergent individuals. The possible existence of “superorganisms” test our best intuitions about what can count and act as genuine biological individuals and how we should study them. (...) As we will discuss, colonies of certain organisms display many of the properties that we usually reserve only to individual organisms. Although there is good reason to believe that many social insects form genuine emergent biological individuals, the conclusion offered here is of a slightly different sort. I will argue that to understand some social insects' interactions and the emergent traits they give rise to, it may be helpful to shift our understanding from a community-level approach to an ecosystem-level approach. I will argue that viewing certain insect colonies (termites) as parts of ecosystems allows us to better understand some of the adaptations that have emerged from their evolution. (shrink)

One of intuitions driving the acceptance of a neat structured tree of life is the assumption that organisms and the lineages they form have somewhat stable spatial and temporal boundaries. The phenomenon of symbiosis shows us that such ‘fixist’ assumptions does not correspond to how the natural world actually works. The implications of lateral gene transfer (LGT) have been discussed elsewhere; I wish to stress a related point. I will focus on lateral function transfer (LFT) and will argue, using examples (...) of what many would call ‘superorganisms’, that the emergence of symbiotic individuals revives the importance of functional and adaptationist thinking in how we conceptualize the lineages of biological individuals. The consequence of the argument is that, if we really want to hold onto tree of life thinking, we had better accept that new saplings appear and disappear all the time. (shrink)

We argue that a fashionable interpretation of the theory of natural selection as a claim exclusively about populations is mistaken. The interpretation rests on adopting an analysis of fitness as a probabilistic propensity which cannot be substantiated, draws parallels with thermodynamics which are without foundations, and fails to do justice to the fundamental distinction between drift and selection. This distinction requires a notion of fitness as a pairwise comparison between individuals taken two at a time, and so vitiates the interpretation (...) of the theory as one about populations exclusively. (shrink)

Following Wallace’s suggestion, Darwin framed his theory using Spencer’s expression “survival of the fittest”. Since then, fitness occupies a significant place in the conventional understanding of Darwinism, even though the explicit meaning of the term ‘fitness’ is rarely stated. In this paper I examine some of the different roles that fitness has played in the development of the theory. Whereas the meaning of fitness was originally understood in ecological terms, it took a statistical turn in terms of reproductive success throughout (...) the 20th Century. This has lead to the ever-increasing importance of sexually reproducing organisms and the populations they compose in evolutionary explanations. I will argue that, moving forward, evolutionary theory should look back at its ecological roots in order to be more inclusive in the type of systems it examines. Many biological systems can only be satisfactorily accounted for by offering a non-reproductive account of fitness. This argument will be made by examining biological systems with very small or transient population structures. I argue this has significant consequences for how we define Darwinism, increasing the significance of survival over that of reproduction. (shrink)

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)

Biologists studying ecology and evolution use the term “population” in many different ways. Yet little philosophical analysis of the concept has been done, either by biologists or philosophers, in contrast to the voluminous literature on the concept of “species.” This is in spite of the fact that “population” is arguably a far more central concept in ecological and evolutionary studies than “species” is. The fact that such a central concept has been employed in so many different ways is potentially problematic (...) for the reason that inconsistent usages (especially when the usage has not been made explicit) might lead to false controversies in which disputants are simply talking past one another. However, the inconsistent usages are not the only, or even the most important reason to examine the concept. If any set of organisms is legitimately called a “population,” selection and drift processes become purely arbitrary, too. Moreover, key ecological variables, such as abundance and distribution, depend on a nonarbitrary way of identifying populations. I sketch the beginnings of a population concept, drawing inspiration from the Ghiselin-Hull individuality thesis, and show why some alternative approaches are nonstarters. (shrink)

We distinguish dynamical and statistical interpretations of evolutionary theory. We argue that only the statistical interpretation preserves the presumed relation between natural selection and drift. On these grounds we claim that the dynamical conception of evolutionary theory as a theory of forces is mistaken. Selection and drift are not forces. Nor do selection and drift explanations appeal to the (sub-population-level) causes of population level change. Instead they explain by appeal to the statistical structure of populations. We briefly discuss the implications (...) of the statistical interpretation of selection for various debates within the philosophy of biologythe `explananda of selection' debate and the `units of selection' debate. (shrink)

Phenotype, whether conventional or extended, is defined as a reflectionof an underlying genotype. Adaptation and the natural selection thatfollows from it depends upon a progressively harmonious fit betweenphenotype and environment. There is in Richard Dawkins' notion ofthe extended phenotype a paradox that seems to undercut conventionalviews of adaptation, natural selection and adaptation. In a nutshell, ifthe phenotype includes an organism's environment, how then can theorganism adapt to itself? The paradox is resolvable through aphysiological, as opposed to a genetic, theory of (...) natural selection andadaptation. (shrink)

Recently, a number of philosophers of biology have endorsed views about random drift that, we will argue, rest on an implicit assumption that the meaning of concepts such as drift can be understood through an examination of the mathematical models in which drift appears. They also seem to implicitly assume that ontological questions about the causality of terms appearing in the models can be gleaned from the models alone. We will question these general assumptions by showing how the same equation (...) — the simple 2 = p2 + 2pq + q2 — can be given radically different interpretations, one of which is a physical, causal process and one of which is not. This shows that mathematical models on their own yield neither interpretations nor ontological conclusions. Instead, we argue that these issues can only be resolved by considering the phenomena that the models were originally designed to represent and the phenomena to which the models are currently applied. When one does take those factors into account, starting with the motivation for Sewall Wright’s and R.A. Fisher’s early drift models and ending with contemporary applications, a very different picture of the concept of drift emerges. On this view, drift is a term for a set of physical processes, namely, indiscriminate sampling processes. (shrink)

Philosophers of biology, along with everyone else, generally perceive life to fall into two broad categories, the microbes and macrobes, and then pay most of their attention to the latter. ‘Macrobe’ is the word we propose for larger life forms, and we use it as part of an argument for microbial equality. We suggest that taking more notice of microbes – the dominant life form on the planet, both now and throughout evolutionary history – will transform some of the philosophy (...) of biology’s standard ideas on ontology, evolution, taxonomy and biodiversity. We set out a number of recent developments in microbiology – including biofilm formation, chemotaxis, quorum sensing and gene transfer – that highlight microbial capacities for cooperation and communication and break down conventional thinking that microbes are solely or primarily single-celled organisms. These insights also bring new perspectives to the levels of selection debate, as well as to discussions of the evolution and nature of multicellularity, and to neo-Darwinian understandings of evolutionary mechanisms. We show how these revisions lead to further complications for microbial classification and the philosophies of systematics and biodiversity. Incorporating microbial insights into the philosophy of biology will challenge many of its assumptions, but also give greater scope and depth to its investigations. (shrink)

In this essay I first provide an analysis of various community concepts. Second, I evaluate two of the most serious challenges to the existence of communities—gradient and paleoecological analysis respectively—arguing that, properly understood, neither threatens the existence of communities construed interactively. Finally, I apply the same interactive approach to ecosystem ecology, arguing that ecosystems may exist robustly as well. ‡I would like to thank to the participants at the Ecology and Environmental Ethics Conference at the University of Utah, the Philosophy (...) of Ecology Conference hosted by the University of Brisbane, and those participants in a session at the Philosophy of Science Association Meeting in Vancouver, British Columbia for helpful discussions of this essay. Specific thanks go to Mark Colyvan, Greg Cooper, Steve Downes, Chris Elliott, Marc Ereshefsky, Paul Griffiths, Jesse Hendrikse, Greg Mikkelson, Anya Plutynski, Kate Ritchie, Sahotra Sarkar, Kim Sterelny, and Rob Wilson. †To contact the author, please write to: Department of Philosophy, Lewis and Clark College, 0615 SW Palatine Hill Road, Portland, OR 97219; e-mail: [email protected]. (shrink)

Recent discussions in the philosophy of biology have brought into question some fundamental assumptions regarding evolutionary processes, natural selection in particular. Some authors argue that natural selection is nothing but a population-level, statistical consequence of lower-level events (Matthen and Ariew [2002]; Walsh et al. [2002]). On this view, natural selection itself does not involve forces. Other authors reject this purely statistical, population-level account for an individual-level, causal account of natural selection (Bouchard and Rosenberg [2004]). I argue that each of these (...) positions is right in one way, but wrong in another; natural selection indeed takes place at the level of populations, but it is a causal process nonetheless. (shrink)

Throughout, this book questions our accepted definitions and biases, showing the self-reflective nature of scientific activity within society.

We address three fundamental questions: What does it mean for an entity to be living? What is the role of inter-organismic collaboration in evolution? What is a biological individual? Our central argument is that life arises when lineage-forming entities collaborate in metabolism. By conceiving of metabolism as a collaborative process performed by functional wholes, which are associations of a variety of lineage-forming entities, we avoid the standard tension between reproduction and metabolism in discussions of life – a tension particularly evident (...) in discussions of whether viruses are alive. Our perspective assumes no sharp distinction between life and non-life, and does not equate life exclusively with cellular or organismal status. We reach this conclusion through an analysis of the capabilities of a spectrum of biological entities, in which we include the pivotal case of viruses as well as prions, plasmids, organelles, intracellular and extracellular symbionts, unicellular and multicellular life-forms. The usual criterion for classifying many of the entities of our continuum as non-living is autonomy. This emphasis on autonomy is problematic, however, because even paradigmatic biological individuals, such as large animals, are dependent on symbiotic associations with many other organisms. These composite individuals constitute the metabolic wholes on which selection acts. Finally, our account treats cooperation and competition not as polar opposites but as points on a continuum of collaboration. We suggest that competitive relations are a transitional state, with multi-lineage metabolic wholes eventually outcompeting selfish competitors, and that this process sometimes leads to the emergence of new types or levels of wholes. Our view of life as a continuum of variably structured collaborative systems leaves open the possibility that a variety of forms of organized matter – from chemical systems to ecosystems – might be usefully understood as living entities. (shrink)

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)

Can selection explain why individuals have the traits they do? This question has generated significant controversy. I will argue that the debate encompasses two separable aspects, to detrimental effect: (1) the role of selection in explaining the origin and evolution of biological traits and (2) the implications this may have for explaining why individuals have the traits they do. (1) can be settled on the basis of evolutionary theory while (2) requires additional, extra-scientific assumptions. By making a distinction between traits (...) affected by a single factor and traits affected by multiple factors I show that selection can, under certain conditions, help explain the origin of traits. Resolving the first aspect enables us to critically assess the various incompatible and independent philosophical commitments made within the second aspect of the debate. (shrink)

With this manifesto, John Dupré systematically attacks the ideal of scientific unity by showing how its underlying assumptions are at odds with the central conclusions of science itself.

WE HAVE LEARNED in the preceding chapter that a revolutionary change of the species concept is in the making, a change which not only affects taxonomic procedure, but which also contributes considerably toward a better understanding of ...

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.

_This sweeping discussion of the philosophy of evolutionary biology is based on the revolutionary idea that species are not kinds of organisms but wholes composed of organisms._.

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

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)

One way to understand science is as a selection process. David Hull, one of the dominant figures in contemporary philosophy of science, sets out in this 2001 volume a general analysis of this selection process that applies equally to biological evolution, the reaction of the immune system to antigens, operant learning, and social and conceptual change in science. Hull aims to distinguish between those characteristics that are contingent features of selection and those that are essential. Science and Selection brings together (...) many of David Hull's most important essays on selection in one accessible volume. (shrink)

Traditionally, species have been treated as classes. In fact they may be considered individuals. The logical term “individual” has been confused with a biological synonym for “organism.” If species are individuals, then: 1) their names are proper, 2) there cannot be instances of them, 3) they do not have defining properties, 4) their constituent organisms are parts, not members. “ Species " may be defined as the most extensive units in the natural economy such that reproductive competition occurs among their (...) parts. Species are to evolutionary theory as firms are to economic theory: this analogy resolves many issues, such as the problems of “reality” and the ontological status of nomenclatorial types. (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)

How do fitness and natural selection relate to other evolutionary factors like architectural constraint, mode of reproduction, and drift? In one way of thinking, drawn from Newtonian dynamics, fitness is one force driving evolutionary change and added to other factors. In another, drawn from statistical thermodynamics, it is a statistical trend that manifests itself in natural selection histories. It is argued that the first model is incoherent, the second appropriate; a hierarchical realization model is proposed as a basis for a (...) statistical treatment. It emerges that natural selection does not cause evolution; it just is evolution. The theory incorporates relations of statistical correlation, but not the kind of causation found in fundamental physical processes. (shrink)

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. (shrink)