Radboud Universiteit Nijmegen, 06 september 2010.

Taxonomies of living things and the methods used to produce them changed little with the institutionalization of evolutionary thinking in biology. Instead, the relationships expressed in existing taxonomies were merely reinterpreted as the result of evolution, and evolutionary concepts were developed to justify existing methods. I argue that the delay of the Darwinian Revolution in biological taxonomy has resulted partly from a failure to distinguish between two fundamentally different ways of ordering identified by Griffiths : classification and systematization. Classification consists (...) of ordering entities into classes, groups defined by the attributes of their members; in contrast, systematization consists of ordering entities into systems, more inclusive entities whose existence depends on some natural process through which their parts are related. Evolutionary, or phylogenetic, systematics takes evolutionary descent to be the natural process of interest in biological taxonomy. I outline a general framework for a truly phylogenetic systematics and examine some of its consequences. (shrink)

W&S correctly ask if groups can be like individuals in the harmony and cooperation of their parts, but in their answer, they ignore the importance of the difference between genetically related and unrelated components, and also misconstrue the import of the Hutterites.

Selection type theories solve adaptation problems. Natural selection, clonal selection for antibody production, and selective theories of higher brain function are examples. An abstract characterization of typical selection processes is generated by analyzing and extending previous work on the nature of natural selection. Once constructed, this abstraction provides a useful tool for analyzing the nature of other selection theories and may be of use in new instances of theory construction. This suggests the potential fruitfulness of research to find other theory (...) types and construct their abstractions. (shrink)

This paper examines conceptual issues that arise in applications of Darwinian natural selection to cultural systems. I argue that many criticisms of cultural selectionist models have been based on an over-detailed reading of the analogy between biological and cultural units of selection. I identify five of the most powerful objections to cultural selection theory and argue that none cuts to its heart. Some objections are based on mistaken assumptions about the simplicity of the mechanisms of biological heredity. Other objections are (...) attributable, rather, to mistaken inferences from observations of biological subject matter to what is essential in natural selection. I argue that such features are idiosyncratic of biological systems, but not essential for natural selection. My arguments throughout are illustrated by examples from biological and cultural evolution, and counter-factual illustrations from the history of theoretical biology. Introduction Cultural Selection Theory First Objection: Lamarckianism Second Objection: Genotype–Phenotype Distinction Third Objection: Common Hereditary Architecture Fourth Objection: Biological Analogue for Cultural Units 6.1 Regarding strict analogues 6.2 Regarding the trait analogue 6.3 Regarding the virus analogue Fifth Objection: Environmental Interaction Conclusion CiteULike Connotea Del.icio.us What's this? (shrink)

Biologists and philosophers have long recognized the importance of species, yet species concepts serve two masters, evolutionary theory on the one hand and taxonomy on the other. Much of present-day evolutionary and systematic biology has confounded these two roles primarily through use of the biological species concept. Theories require entities that are real, discrete, irreducible, and comparable. Within the neo-Darwinian synthesis, however, biological species have been treated as real or subjectively delimited entities, discrete or nondiscrete, and they are often capable (...) of being decomposed into other, smaller units. Because of this, biological species are generally not comparable across different groups of organisms, which implies that the ontological structure of evolutionary theory requires modification. Some biologists, including proponents of the biological species concept, have argued that no species concept is universally applicable across all organisms. Such a view means, however, that the history of life cannot be embraced by a common theory of ancestry and descent if that theory uses species as its entities.These ontological and biological difficulties can be alleviated if species are defined in terms of evolutionary units. The latter are irreducible clusters of reproductively cohesive organisms that are diagnosably distinct from other such clusters. Unlike biological species, which can include two or more evolutionary units, these phylogenetic species are discrete entities in space and time and capable of being compared from one group to the next. (shrink)

Do multi-level selection explanations of the evolution of social traits deepen the understanding provided by single-level explanations? Central to the former is a mathematical theorem, the multi-level Price decomposition. I build a framework through which to understand the explanatory role of such non-empirical decompositions in scientific practice. Applying this general framework to the present case places two tasks on the agenda. The first task is to distinguish the various ways of suppressing within-collective variation in fitness, and moreover to evaluate their (...) biological interest. I distinguish four such ways: increasing retaliatory capacity, homogenising assortment, and collapsing either fitness structure or character distribution to a mean value. The second task is to discover whether the third term of the Price decomposition measures the effect of any of these hypothetical interventions. On this basis I argue that the multi-level Price decomposition has explanatory value primarily when the sharing-out of collective resources is `subtractable'. Thus its value is more circumscribed than its champions Sober and Wilson (1998) suppose. (shrink)

References in Darwin’s Origin of Species to competition between units of selection at and above the level of individual organisms are enumerated. In many cases these references clearly speak of natural selection and do not support the view that Darwin thought selection only occurred at the level of the individual organism. Darwin did see organismal selection as the main process by which varieties were created but he also espoused what is here termed community and varietal selection. He saw no essential (...) difference between varieties and species and the references show that he also believed that selection could operate at the species level. (shrink)

We have demonstrated, using the Cantor dust method, that the statistical distribution of appearance and disappearance of rodents species (Arvicolid rodent radiation in Europe) follows power laws strengthening the evidence for a fractal structure set. Self-similar laws have been used as model for the description of a huge number of biological systems. With Nottale we have shown that log-periodic behaviors of acceleration or deceleration can be applied to branching macroevolution, to the time sequences of major evolutionary leaps (global life tree, (...) sauropod and theropod dinosaurs postural structures, North American fossil equids, rodents, primates and echinoderms clades and human ontogeny). The Scale-Relativity Theory has others biological applications from linear with fractal behavior to non-linear and from classical mechanics to quantum mechanics. (shrink)

This chapter offers a review of standard views about the requirements for natural selection to shape evolution and for the sorts of ‘units’ on which selection might operate. It then summarizes traditional arguments for genic selectionism, i.e., the view that selection operates primarily on genes (e.g., those of G. C. Williams, Richard Dawkins, and David Hull) and traditional counterarguments (e.g., those of William Wimsatt, Richard Lewontin, and Elliott Sober, and a diffuse group based on life history strategies). It then offers (...) a series of responses to the arguments, based on more contemporary considerations from molecular genetics, offered by Carmen Sapienza. A key issue raised by Sapienza concerns the degree to which a small number of genes might be able to control much of the variation relevant to selection operating on such selectively critical organs as hearts. The response to these arguments suggests that selection acts on many levels at once and that sporadic selection, acting with strong effects, can act successively on different key traits (and genes) while maintaining a balance among many potentially conflicting demands faced by organisms within an evolving lineage. (shrink)

There is no question that the constituents of cells and organisms are joined together by the part-whole relation. Genes are part of cells, and cells are part of organisms. Species taxa, however, have traditionally been conceived of, not as wholes with parts, but as classes with members. But why does the relation change abruptly from part-whole to class-membership above the level of organisms? Ghiselin, Hull and others have argued that it doesn't. Cells and organisms are cohesive mereological sums, and since (...) species taxa are like cells and organisms in the relevant respects, they, too, are cohesive mereological sums. I provide further reasons in support of the thesis that species are mereological sums. I argue, moreover, that the advocate of this thesis is committed to a form of pluralism with respect to the species concept. (shrink)

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)

Despite hundreds of definitions, no consensus exists on a definition of life or on the closely related and problematic definitions of the organism and death. These problems retard practical and theoretical development in, for example, exobiology, artificial life, biology and evolution. This paper suggests improving this situation by basing definitions on a theory of a generalized particle hierarchy. This theory uses the common denominator of the “operator” for a unified ranking of both particles and organisms, from elementary particles to animals (...) with brains. Accordingly, this ranking is called “the operator hierarchy”. This hierarchy allows life to be defined as: matter with the configuration of an operator, and that possesses a complexity equal to, or even higher than the cellular operator. Living is then synonymous with the dynamics of such operators and the word organism refers to a select group of operators that fit the definition of life. The minimum condition defining an organism is its existence as an operator, construction thus being more essential than metabolism, growth or reproduction. In the operator hierarchy, every organism is associated with a specific closure, for example, the nucleus in eukaryotes. This allows death to be defined as: the state in which an organism has lost its closure following irreversible deterioration of its organization. The generality of the operator hierarchy also offers a context to discuss “life as we do not know it”. The paper ends with testing the definition’s practical value with a range of examples. (shrink)

When is conceptual change so significant that we should talk about a new theory, not a new version of the same theory? We address this problem here, starting from Gould’s discussion of the individuation of the Darwinian theory. He locates his position between two extremes: ‘minimalist’—a theory should be individuated merely by its insertion in a historical lineage—and ‘maximalist’—exhaustive lists of necessary and sufficient conditions are required for individuation. He imputes the minimalist position to Hull and attempts a reductio : (...) this position leads us to give the same ‘name’ to contradictory theories. Gould’s ‘structuralist’ position requires both ‘conceptual continuity’ and descent for individuation. Hull’s attempt to assimilate into his general selectionist framework Kuhn’s notion of ‘exemplar’ and the ‘semantic’ view of the structure of scientific theories can be used to counter Gould’s reductio , and also to integrate structuralist and population thinking about conceptual change. (shrink)

Science is a dynamic process in which the assimilation of new phenomena, perspectives, and hypotheses into the scientific corpus takes place slowly. The apparent disunity of the sciences is the unavoidable consequence of this gradual integration process. Some thinkers label this dynamical circumstance a ‘crisis’. However, a retrospective view of the practical results of the scientific enterprise and of science itself, grants us a clear view of the unity of the human knowledge seeking enterprise. This book provides many arguments, case (...) studies and examples in favor of the unity of science. These contributions touch upon various scientific perspectives and disciplines such as: Physics, Computer Science, Biology, Neuroscience, Cognitive Psychology, and Economics. (shrink)

The position and role of humanity in the world of life is examined in the light of the ontological structure of life itself. This problem is approached by considering the possible units of life representing various modes of life phenomena. I argue that the only meaningful unit of life without interposing some special external conditions is “global life” framed in a star‐planet system. Any other possible unit of life exhibited by various kinds of individuals is conditional in the sense that (...) it would leave out an essential part as “co‐life.” The relationship between human being and the global life should be understood in this general scheme of individual and global life. It is emphasized, however, that human being occupies a unique position in global life in the sense that humanity can promote either a cancerous situation or a healthy higher‐order enhancement of the global life. (shrink)

Genes are thought to have evolved from long-lived and multiply-interactive molecules in the early stages of the origins of life. However, at that stage there were no replicators, and the distinction between interactors and replicators did not yet apply. Nevertheless, the process of evolution that proceeded from initial autocatalytic hypercycles to full organisms was a Darwinian process of selection of favourable variants. We distinguish therefore between Neo-Darwinian evolution and the related Weismannian and Central Dogma divisions, on the one hand, and (...) the more generic category of Darwinian evolution on the other. We argue that Hull’s and Dawkins’ replicator/interactor distinction of entities is a sufficient, but not necessary, condition for Darwinian evolution to take place. We conceive the origin of genes as a separation between different types of molecules in a thermodynamic state space, and employ a notion of reproducers. (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)

Memes are an idea whose time has come, again, and again, and again, but which has never really made it beyond metaphor. Anthropologist Robert Aunger’s book 'The Electric Meme' is a new attempt to take it to the next stage, setting up a research program with proper models and theoretical entities. He succeeds partially, with some contributions to the logic of replication, but in the end, his proposal for the substrate of memes is a non-solution to a central problem of (...) memetics. (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)

What are species? One popular answer is that species are individuals. Here I develop another approach to thinking about species, an approach based on the notion of a lineage. A lineage is a sequence of reproducing entities, individuated in terms of its components. I argue that one can conceive of species as groups of lineages, either organism lineages or population lineages. Conceiving of species as groups of lineages resolves the problems that the individual conception of species is supposed to resolve. (...) It has added the virtue of focusing attention on the characteristic of species that is most relevant to understanding their role in evolutionary processes, namely, the lineage structure of species. (shrink)

The Darwinian concept of natural selection was conceived within a set of Newtonian background assumptions about systems dynamics. Mendelian genetics at first did not sit well with the gradualist assumptions of the Darwinian theory. Eventually, however, Mendelism and Darwinism were fused by reformulating natural selection in statistical terms. This reflected a shift to a more probabilistic set of background assumptions based upon Boltzmannian systems dynamics. Recent developments in molecular genetics and paleontology have put pressure on Darwinism once again. Current work (...) on self-organizing systems may provide a stimulus not only for increased problem solving within the Darwinian tradition, especially with respect to origins of life, developmental genetics, phylogenetic pattern, and energy-flow ecology, but for deeper understanding of the very phenomenon of natural selection itself. Since self-organizational phenomena depend deeply on stochastic processes, self-organizational systems dynamics advance the probability revolution. In our view, natural selection is an emergent phenomenon of physical and chemical selection. These developments suggest that natural selection may be grounded in physical law more deeply than is allowed by advocates of the autonomy of biology, while still making it possible to deny, with autonomists, that evolutionary explanations can be modeled in terms of a deductive relationship between laws and cases. We explore the relationship between, chance, self-organization, and selection as sources of order in biological systems in order to make these points. (shrink)

Enzyme directed genetic mechanisms causing random DNA sequence alterations are ubiquitous in both eukaryotes and prokaryotes. A number of molecular geneticist have invoked adaptation through natural selection to account for this fact, however, alternative explanations have also flourished. The population geneticist G.C. Williams has dismissed the possibility of selection for mutator activity on a priori grounds. In this paper, I attempt a refutation of Williams' argument. In addition, I discuss some conceptual problems related to recent claims made by microbiologists on (...) the adaptiveness of molecular variety generators in the evolution of prokaryotes. A distinction is proposed between selection for mutations caused by a mutator activity and selection for the mutator activity proper. The latter requires a concept of fitness different from the one commonly used in microbiology. (shrink)

The Units of Selection debate is a dispute about the causes of population change. I argue that it is generated by a particular `dynamical'' interpretation of natural selection theory, according to which natural selection causes differential survival and reproduction of individuals and natural selection explanations cite these causes. I argue that the dynamical interpretation is mistaken and offer in outline an alternative, `statistical'' interpretation, according to which natural selection theory is a fancy kind of `bookkeeping''. It explains by citing the (...) statistical structure of a population and not by citing the causes of survival and reproduction. From the perspective of the statistical interpretation there is no substantive Units of Selection issue. (shrink)

Crane envisions the speculative conjecture that intelligent civilizations might want and be able to produce black holes in the very far future. He implicitly suggests two main purposes of this enterprise: (i) energy production and (ii) universe production. We discuss those two options. The commentary is obviously highly speculative and should be read accordingly.

This paper surveys recent philosophy of biology. It aims to introduce outsiders to the field to the recent literature (which is reviewed in the footnotes) and the main recent debates. I concentrate on three of these: recent critiques of the replicator/vehicle distinction and its application to the idea of the gene as the unit of section; the recent defences of group selection and the idea that standard alternatives to group selection are in fact no more than a disguised form of (...) group selection; and recent ideas on the role of selection in evolution, especially the role of selection in structuring the large-scale history of life. The paper connects philosophy of biology to some more general problems in the philosophy of science, and concludes with a few suggestions about unfinished business. (shrink)

This paper evaluates and criticises the developmental systems conception of evolution and develops instead an extension of the gene's eye conception of evolution. We argue (i) Dawkin's attempt to segregate developmental and evolutionary issues about genes is unsatisfactory. On plausible views of development it is arbitrary to single out genes as the units of selection. (ii) The genotype does not carry information about the phenotype in any way that distinguishes the role of the genes in development from that other factors. (...) (iii) There is no simple and general causal criterion which distinguishes the role of genes in development and evolution. (iv) There is, however, an important sense in which genes but not every other developmental factor represent the phenotype. (v) The idea that genes represent features of the phenotype forces us to recognise that genes are not the only, or almost the only, replicators. Many mechanisms of replication are involved in both development and evolution. (vi) A conception of evolutionary history which recognises both genetic and non-genetic replicators, lineages of replicators and interactors has advantages over both the radical rejection of the replicator/interactor distinction and the conservative restriction of replication to genetic replication. (shrink)

The purpose of this paper is to test the contemporary concept of biological species against some of the problems caused by treating species as spatiotemporally extended entities governed by criteria of persistence, identity, etc. After outlining the general problem of symmetric division in natural objects, I set out some useful distinctions (section 1) and confirm that species are not natural kinds (section 2). Section 3 takes up the separate issue of species definition, focusing on the Biological Species Concept (BSC). Sections (...) 4 and 5 examine the matter of species identity over space and time respectively, as determined by the BSC. Both gradualistic and punctuated equilibrium models of speciation are discussed. In section 6 I argue that the BSC fails to determine adequate criteria for dealing with certain kinds of speciation. Section 7 moves speculatively beyond the BSC to a brief examination of alternatives. (shrink)

We review Evolution and the Levels of Selection by Samir Okasha. This important book provides a cohesive philosophical framework for understanding levels-of-selections problems in biology. Concerning evolutionary transitions, Okasha proposes that three stages characterize the shift from a lower level of selection to a higher one. We discuss the application of Okasha’s three-stage concept to the evolutionary transition from unicellularity to multicellularity in the volvocine green algae. Okasha’s concepts are a provocative step towards a more general understanding of the major (...) evolutionary transitions; however, the application of certain ideas to the volvocine model system is not straightforward. (shrink)