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.

This paper examines David Hull’s and Peter Godfrey-Smith’s accounts of biological individuality using the case of biofilms. Biofilms fail standard criteria for individuality, such as having reproductive bottlenecks and forming parent-offspring lineages. Nevertheless, biofilms are good candidates for individuals. The nature of biofilms shows that Godfrey-Smith’s account of individuality, with its reliance on reproduction, is too restrictive. Hull’s interactor notion of individuality better captures biofilms, and we argue that it offers a better account of biological individuality. However, Hull’s notion of (...) interactor needs more precision. We suggest some ways to make Hull’s notion of interactor and his account of individuality more precise. Generally, we maintain that biofilms are a good test case for theories of individuality, and a careful examination of biofilms furthers our understanding of biological individuality. (shrink)

This chapter develops the idea that the germ-soma split and the suppression of individual fitness differences within the corporate entity are not always essential steps in the evolution of corporate individuals. It illustrates some consequences for multilevel selection theory. It presents evidence that genetic heterogeneity may not always be a barrier to successful functioning as a higher-level individual. This chapter shows that levels-of-selection theorists are wrong to assume that the central problem in transitions is always that of minimizing within-group competition. (...) Evidence of intralevel conflict does not qualify as evidence against the existence of a higher level of selection. (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)

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

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)

John Dupr explores recent revolutionary developments in biology and considers their relevance for our understanding of human nature and society. He reveals how the advance of genetic science is changing our view of the constituents of life, and shows how an understanding of microbiology will overturn standard assumptions about the living world.

John Dupré explores recent revolutionary developments in biology and considers their relevance for our understanding of human nature and human society. Epigenetics and related areas of molecular biology have eroded the exceptional status of the gene and presented the genome as fully interactive with the rest of the cell. Developmental systems theory provides a space for a vision of evolution that takes full account of the fundamental importance of developmental processes. Dupré shows the importance of microbiology for a proper understanding (...) of the living world, and reveals how it subverts such basic biological assumptions as the organisation of biological kinds on a branching tree of life, and the simple traditional conception of the biological organism. (shrink)

The commentaries by Dennett, Sterelny, and Queller on Darwinian Populations and Natural Selection (DPNS) are so constructive that they make it possible to extend and improve the book’s framework in several ways. My replies will focus on points of disagreement, and I will pick a small number of themes and develop them in detail. The three replies below are mostly self-contained, except that all my comments about genes, discussed by all three critics, are in the reply to Queller. Agential views (...) of evolution, discussed by Queller and Dennett, are addressed in my reply to Dennett. (shrink)

Both biologists and philosophers often make use of simple verbal formulations of necessary and sufficient conditions for evolution by natural selection (ENS). Such summaries go back to Darwin's Origin of Species (especially the "Recapitulation"), but recent ones are more compact.1 Perhaps the most commonly cited formulation is due to Lewontin.2 These summaries tend to have three or four conditions, where the core requirement is a combination of variation, heredity, and fitness differences. The summaries are employed in several ways. First, they (...) are often used in pedagogical contexts, and in showing the coherence of evolutionary theory in response to attacks from outside biology. Second, they are important in discussions of extensions of evolutionary principles to new domains, such as cultural change. The summaries also have intrinsic scientific and philosophical interest as attempts to capture some core principles of evolutionary theory in a highly concise way. Despite their prominence, both the proper formulation and status of these summaries are unclear. Standard formulations are subject to counterexamples, and their relations to formal models of evolutionary change are not straightforward. Here I look closely at these verbal summaries, and at how they relate to formal models. Are the summaries merely rough approximations that have no theoretical role of their own? Perhaps they could operate as theoretical statements in Darwin's time, but have now been superseded by more exact treatments. (shrink)

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.

Authors frequently refer to gene-based selection in biological evolution, the reaction of the immune system to antigens, and operant learning as exemplifying selection processes in the same sense of this term. However, as obvious as this claim may seem on the surface, setting out an account of “selection” that is general enough to incorporate all three of these processes without becoming so general as to be vacuous is far from easy. In this target article, we set out such a general (...) account of selection to see how well it accommodates these very different sorts of selection. The three fundamental elements of this account are replication, variation, and environmental interaction. For selection to occur, these three processes must be related in a very specific way. In particular, replication must alternate with environmental interaction so that any changes that occur in replication are passed on differentially because of environmental interaction. One of the main differences among the three sorts of selection that we investigate concerns the role of organisms. In traditional biological evolution, organisms play a central role with respect to environmental interaction. Although environmental interaction can occur at other levels of the organizational hierarchy, organisms are the primary focus of environmental interaction. In the functioning of the immune system, organisms function as containers. The interactions that result in selection of antibodies during a lifetime are between entities (antibodies and antigens) contained within the organism. Resulting changes in the immune system of one organism are not passed on to later organisms. Nor are changes in operant behavior resulting from behavioral selection passed on to later organisms. But operant behavior is not contained in the organism because most of the interactions that lead to differential replication include parts of the world outside the organism. Changes in the organism's nervous system are the effects of those interactions. The role of genes also varies in these three systems. Biological evolution is gene-based (i.e., genes are the primary replicators). Genes play very different roles in operant behavior and the immune system. However, in all three systems, iteration is central. All three selection processes are also incredibly wasteful and inefficient. They can generate complexity and novelty primarily because they are so wasteful and inefficient. Key Words: evolution; immunology; interaction; operant behavior; operant learning; replication; selection; variation. (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.