Familiarity with Charles Darwin's treatise on evolution is essential to every well-educated individual. One of the most important books ever published--and a continuing source of controversy, a century and a half later--this classic of science is reproduced in a facsimile of the critically acclaimed first edition.

In The Origin of Species (1859) Darwin challenged many of the most deeply-held beliefs of the Western world. Arguing for a material, not divine, origin of species, he showed that new species are achieved by "natural selection." The Origin communicates the enthusiasm of original thinking in an open, descriptive style, and Darwin's emphasis on the value of diversity speaks more strongly now than ever. As well as a stimulating introduction and detailed notes, this edition offers a register of the many (...) writers referred to by Darwin in the text. (shrink)

Does natural selection act primarily on individual organisms, on groups, on genes, or on whole species? The question of levels of selection - on which biologists and philosophers have long disagreed - is central to evolutionary theory and to the philosophy of biology. Samir Okasha's comprehensive analysis gives a clear account of the philosophical issues at stake in the current debate.

Richard Lewontin's (1970) early work on the units of selection initiated the conceptual and theoretical investigations that have led to the hierarchical perspective on selection that has reached near consensus status today. This paper explores other aspects of his work, work on what he termed continuity and quasi-independence, that connect to contemporary explorations of modularity in development and evolution. I characterize such modules and argue that they are the true units of selection in that they are what evolution by natural (...) selection individuates, selects among, and transforms. (shrink)

Ecological fitness has been suggested to provide a unifying definition of fitness. However, a metric for this notion of fitness was in most cases unavailable except by proxy with differential reproductive success. In this article, I show how differential persistence of lineages can be used as a way to assess ecological fitness. This view is inspired by a better understanding of the evolution of some clonal plants, colonial organisms, and ecosystems. Differential persistence shows the limitation of an ensemblist noncausal understanding (...) of evolution. Causal explanations are necessary to understand the evolution by natural selection of these biological systems. †To contact the author write to: Department of Philosophy, University of Montreal, P.O. Box 6128, Station Centre‐Ville, Montreal, Quebec, H3C 3J7 Canada; e‐mail: [email protected]. (shrink)

Is the history of life a series of accidents or a drama scripted by selfish genes? Is there an “essential” human nature, determined at birth or in a distant evolutionary past? What should we conserve—species, ecosystems, or something else? -/- Informed answers to questions like these, critical to our understanding of ourselves and the world around us, require both a knowledge of biology and a philosophical framework within which to make sense of its findings. In this accessible introduction to philosophy (...) of biology, Kim Sterelny and Paul E. Griffiths present both the science and the philosophical context necessary for a critical understanding of the most exciting debates shaping biology today. The authors, both of whom have published extensively in this field, describe the range of competing views—including their own—on these fascinating topics. -/- With its clear explanations of both biological and philosophical concepts, Sex and Death will appeal not only to undergraduates, but also to the many general readers eager to think critically about the science of life. (shrink)

In both biology and the human sciences, social groups are sometimes treated as adaptive units whose organization cannot be reduced to individual interactions. This group-level view is opposed by a more individualistic one that treats social organization as a byproduct of self-interest. According to biologists, group-level adaptations can evolve only by a process of natural selection at the group level. Most biologists rejected group selection as an important evolutionary force during the 1960s and 1970s but a positive literature began to (...) grow during the 1970s and is rapidly expanding today. We review this recent literature and its implications for human evolutionary biology. We show that the rejection of group selection was based on a misplaced emphasis on genes as “replicators” which is in fact irrelevant to the question of whether groups can be like individuals in their functional organization. The fundamental question is whether social groups and other higher-level entities can be “vehicles” of selection. When this elementary fact is recognized, group selection emerges as an important force in nature and what seem to be competing theories, such as kin selection and reciprocity, reappear as special cases of group selection. The result is a unified theory of natural selection that operates on a nested hierarchy of units.The vehicle-based theory makes it clear that group selection is an important force to consider in human evolution. Humans can facultatively span the full range from self-interested individuals to “organs” of group-level “organisms.” Human behavior not only reflects the balance between levels of selection but it can also alter the balance through the construction of social structures that have the effect of reducing fitness differences within groups, concentrating natural selection (and functional organization) at the group level. These social structures and the cognitive abilities that produce them allow group selection to be important even among large groups of unrelated individuals. (shrink)

Biology lacks a central organism concept that unambiguously marks the distinction between organism and non-organism because the most important questions about organisms do not depend on this concept. I argue that the two main ways to discover useful biological generalizations about multicellular organization--the study of homology within multicellular lineages and of convergent evolution across lineages in which multicellularity has been independently established--do not require what would have to be a stipulative sharpening of an organism concept.

The authors demonstrate that unselfish behavior is in fact an important feature of both biological and human nature. Their book provides a panoramic view of altruism throughout the animal kingdom--from self-sacrificing parasites to the human capacity for selflessness--even as it explains the evolutionary sense of such behavior.

I discuss two subjects in Samir Okasha’s excellent book, Evolution and the Levels of Selection. In consonance with Okasha’s critique of the conventionalist view of the units of selection problem, I argue that conventionalists have not attended to what realists mean by group, individual, and genic selection. In connection with Okasha’s discussion of the Price equation and contextual analysis, I discuss whether the existence of these two quantitative frameworks is a challenge to realism.

The self/non-self model, first proposed by F.M. Burnet, has dominated immunology for 60 years now. According to this model, any foreign element will trigger an immune reaction in an organism, whereas endogenous elements will not, in normal circumstances, induce an immune reaction. In this paper we show that the self/non-self model is no longer an appropriate explanation of experimental data in immunology, and that this inadequacy may be rooted in an excessively strong metaphysical conception of biological identity. We suggest that (...) another hypothesis, one based on the notion of continuity, gives a better account of immune phenomena. Finally, we underscore the mapping between this metaphysical deflation from self to continuity in immunology and the philosophical debate between substantialism and empiricism about identity. (shrink)

The main objective of immunology is to establish why and when an immune response occurs, that is, to determine a criterion of immunogenicity. According to the consensus view, the proper criterion of immunogenicity lies in the discrimination between self and nonself. Here we challenge this consensus by suggesting a simpler and more comprehensive criterion, the criterion of continuity. Moreover, we show that this criterion may be considered as an interpretation of the immune 'self'. We conclude that immunologists can continue to (...) speak of the self, provided that they admit that the self/nonself discrimination is not an adequate criterion of immunogenicity. (shrink)

The functional complexity, or the number of functions, of organisms hasfigured prominently in certain theoretical and empirical work inevolutionary biology. Large-scale trends in functional complexity andcorrelations between functional complexity and other variables, such assize, have been proposed. However, the notion of number of functions hasalso been operationally intractable, in that no method has been developedfor counting functions in an organism in a systematic and reliable way.Thus, studies have had to rely on the largely unsupported assumption thatnumber of functions can be (...) measured indirectly, by using number ofmorphological, physiological, and behavioral parts as a proxy. Here, amodel is developed that supports this assumption. Specifically, the modelpredicts that few parts will have many functions overlapping in them, andtherefore the variance in number of functions per part will be low. If so,then number of parts is expected to be well correlated with number offunctions, and we can use part counts as proxies for function counts incomparative studies of organisms, even when part counts are low. Alsodiscussed briefly is a strategy for identifying certain kinds of parts inorganisms in a systematic way. (shrink)

Biology lacks a central organism concept that unambiguously marks the distinction between organism and non-organism because the most important questions about organisms do not depend on this concept. I argue that the two main ways to discover useful biological generalizations about multicellular organization—the study of homology within multicellular lineages and of convergent evolution across lineages in which multicellularity has been independently established—do not require what would have to be a stipulative sharpening of an organism concept.

In this paper we argue that an operational organism concept can help to overcome the structural deficiency of mathematical models in biology. In our opinion, the structural deficiency of mathematical models lies mainly in our inability to identify functionally relevant biological characters in biological systems, and not so much in a lack of adequate mathematical representations of biological processes. We argue that the problem of character identification in biological systems is linked to the question of a properly formulated organism concept. (...) Lastly, we demonstrate how a decomposition of an organism into independent characters in the context of a specific biological process—such as adaptation by means of natural selection—depends on the dynamical properties and invariance conditions of the equations that describe this process. (shrink)

In this paper we argue that an operational organism concept can help to overcome the structural deficiency of mathematical models in biology. In our opinion, the structural deficiency of mathematical models lies mainly in our inability to identify functionally relevant biological characters in biological systems, and not so much in a lack of adequate mathematical representations of biological processes. We argue that the problem of character identification in biological systems is linked to the question of a properly formulated organism concept. (...) Lastly, we demonstrate how a decomposition of an organism into independent characters in the context of a specific biological process--such as adaptation by means of natural selection--depends on the dynamical properties and invariance conditions of the equations that describe this process. (shrink)

Dieser Aufsatz ist eine Besinnung auf den Begriff Individuum, wie dieser in der Biologie gebraucht wird, und er beabsichtigt, den Inhalt dieses Begriffes theoretisch zu begründen. Es ist aber unmöglich für die nähere Bestimmung des Begriffes Individuum von der etymologischen Bedeutung „unteilbar” auszugehen, weil die biologischen Individuen nicht immer unteilbar sind. Beim Suchen nach einem richtigen Untersuchungsverfahren wird zunächst dargelegt, wie diese Untersuchung wesentlich ein Teil der theoretischen Biologie, also der Wissenschaft, ist, wenn man auch von deutlichen Zusammenhängen mit der (...) Philosophie und dem spontanen Denken sprechen kann. Deshalb wird die Methode, nach der wir vorangehen müssen, die induktive Methode sein, weil diese in der Wissenschaft mit Recht hoch eingeschätzt wird.Bei den grossen Vertretern der verschiedenen Richtungen in der Biologie, beiHaeckel, von Bertalanffy, Driesch undBergson, werden in ihre Auffassungen über Individuum und Individualität derartige Übereinstimmungen festgestellt, dass darin nach den Gesetzen der Induktion die wesentlichen Momente des biologischen Individuumbegriffes erkannt werden können. Ausserdem stimmen diese Momente mit dem Inhalt des Begriffs zusammen, wie der sich aus dem spontanen Denken ergibt.Diese Momente sind: Subsistenz, Einheit und Totalität, Tendenz nach einem geschlossenen System und schliesslich die Unmöglichkeit einer erschöpfenden Definition. In dieser Reihenfolge werden sie kritisch erörtert, und daraufhin werden folgende biologische Begriffsbestimmungen vorgeschlagen : Individualität ist die Tendenz des einzelnen Organismus nach ungeteilter subsistenter Totalität. Etwas ist Individuum in dem Masse, wo diese Tendenz verwirklicht ist.Durch einige Beispiele aus dem System und der Ontogenie wird die Anwendung dieses dynamischen Begriffes erläutert.Dans cette étude nous avons soumis à un examen l'idée d'individu, telle qu'elle est d'un usage courant en biologie. Nous avons cherché à fonder la théorie du contenu de cette idée. Pour en donner une définition plus exacte, nous ne pouvons pas prendre pour point de départ le sens étymologique d'„indivisible”, puisque les individus biologiques ne le sont pas toujours. En essayant d'abord d'établir la vraie méthode de notre recherche, nous avons expliqué que celle-ci est une branche de la biologie théorique, par conséquent de la science positive, bien qu'il y ait des rapports avec la philosophie et avec la simple pensée spontanée. Il nous faudra donc procéder d'après la méthode qui est surtout demandée par les sciences exactes, à savoir la méthode inductive.Chez les grands représentants des différents courants de la pensée en biologie, notamment chezHaeckel, von Bertalanffy, Driesch etBergson, nous rencontrons dans leurs conceptions sur l'individu et sur l'individualité des correspondances tellement frappantes que nous sommes autorisé à y voir, d'après les lois mêmes de l'induction, les éléments essentiels de l'idée d'individu biologique. Ces éléments s'accordent d'ailleurs avec le contenu de l'idée que nous nous en formons spontanément.Ces éléments sont les suivants: subsistance, unité et totalité, tendance à devenir un système clos, et enfin l'impossibilité d'une définition adéquate. Nous en avons fait la critique dans cet ordre. En suite nous avons proposé les définitions biologiques suivantes : l'individualité consiste en la tendance que possède un organisme naturellement isolé à devenir une totalité subsistante et non-divisée. On aura donc un individu dans la mesure que cette tendance a été réalisée.Enfin, nous avons illustré l'application de cette notion par quelques exemples empruntés au système et à l'ontogénie. (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)

Developmental systems theory (DST) expands the unit of replication from genes to whole systems of developmental resources, which DST interprets in terms of cycling developmental processes. Expansion seems required by DST's argument against privileging genes in evolutionary and developmental explanations of organic traits. DST and the expanded replicator brook no distinction between biological and cultural evolution. However, by endorsing a single expanded unit of inheritance and leaving the classical molecular notion of gene intact, DST achieves only a nominal reunification of (...) heredity and development. I argue that an alternative conceptualization of inheritance denies the classical opposition of genetics and development while avoiding the singularity inherent in the replicator concept. It also yields a new unit--the reproducer--which genuinely integrates genetic and developmental perspectives. The reproducer concept articulates the non-separability of "genetic" and "developmental" roles in units of heredity, development, and evolution. DST reformulated in terms of reproducers rather than replicators preserves an empirically interesting distinction between cultural and biological evolution. (shrink)

This work has been selected by scholars as being culturally important, and is part of the knowledge base of civilization as we know it. This work was reproduced from the original artifact, and remains as true to the original work as possible. Therefore, you will see the original copyright references, library stamps (as most of these works have been housed in our most important libraries around the world), and other notations in the work.This work is in the public domain in (...) the United States of America, and possibly other nations. Within the United States, you may freely copy and distribute this work, as no entity (individual or corporate) has a copyright on the body of the work.As a reproduction of a historical artifact, this work may contain missing or blurred pages, poor pictures, errant marks, etc. Scholars believe, and we concur, that this work is important enough to be preserved, reproduced, and made generally available to the public. We appreciate your support of the preservation process, and thank you for being an important part of keeping this knowledge alive and relevant. (shrink)

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)

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)

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

The question “What is an organism?”, formerly considered as essential in biology, has now been increasingly replaced by a larger question, “What is a biological individual?”. On the grounds that i) individuation is theory-dependent, and ii) physiology does not offer a theory, biologists and philosophers of biology have claimed that it is the theory of evolution by natural selection which tells us what counts as a biological individual. Here I show that one physiological field, immunology, offers a theory, which makes (...) possible a biological individuation based on physiological grounds. I give a new answer to the question of the individuation of an organism by linking together the evolutionary and the immunological approaches to biological individuation. (shrink)

There is no broadly accepted definition of ‘life.’ Suggested definitions face problems, often in the form of robust counter-examples. Here we use insights from philosophical investigations into language to argue that defining ‘life’ currently poses a dilemma analogous to that faced by those hoping to define ‘water’ before the existence of molecular theory. In the absence of an analogous theory of the nature of living systems, interminable controversy over the definition of life is inescapable.

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)