Scientific conflicts often stem from differences in the conceptual framework through which scientists view and understand their own field. In this chapter, I analyze the ontological and methodological assumptions of three traditions in evolutionary biology, namely, Ernst Mayr’s population thinking, the gene-centered view of the Modern Syn thesis, and the Extended Evolutionary Synthesis. Each of these frameworks presupposes a different account of "evolutionary causes," and this discrepancy prevents mutual understanding and objective evaluation in the recent contention surrounding the EES. From (...) this perspective, the chapter characterizes the EES research program as an attempt to introduce causal structures beyond genes as additional units of evolution, and compares its research methodology and objectives with those of the traditional MS framework. (shrink)

We assess the arguments for recognising functionally integrated multispecies consortia as genuine biological individuals, including cases of so-called ‘holobionts’. We provide two examples in which the same core biochemical processes that sustain life are distributed across a consortium of individuals of different species. Although the same chemistry features in both examples, proponents of the holobiont as unit of evolution would recognize one of the two cases as a multispecies individual whilst they would consider the other as a compelling case of (...) ecological dependence between separate individuals. Some widely used arguments in support of the ‘holobiont’ concept apply equally to both cases, suggesting that those arguments have misidentified what is at stake when seeking to identify a new level of biological individuality. One important aspect of biological individuality is evolutionary individuality. In line with other work on the evolution of individuality, we show that our cases can be distinguished by focusing on the fitness alignment between the partners of the consortia. We conclude that much of the evidence currently presented for the ubiquity and importance of multi-species individuals is simply not to the point, at least unless the issue of biological individuality is firmly divorced from the question of evolutionary individuality. (shrink)

Yes, fitness is the central concept of evolutionary biology, but it is an elusive concept. Almost everyone who looks at it seriously comes out in a different place.

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)

This Ph.D. thesis provides a pilosophical account of the structure of the evolutionary synthesis of the 1930s and 40s. The first, more historical part analyses how classical genetics came to be integrated into evolutionary thinking, highlighting in particular the importance of chromosomal mapping of Drosophila strains collected in the wild by Dobzansky, but also the work of Goldschmidt, Sumners, Timofeeff-Ressovsky and others. The second, more philosophical part attempts to answer the question wherein the unity of the synthesis consisted. I argue (...) that it exemplifies a weak of form of explanatory unification. (shrink)

Philosophy of science is positioned to make distinctive contributions to cognitive science by providing perspective on its conceptual foundations and by advancing normative recommendations. The philosophy of science I embrace is naturalistic in that it is grounded in the study of actual science. Focusing on explanation, I describe the recent development of a mechanistic philosophy of science from which I draw three normative consequences for cognitive science. First, insofar as cognitive mechanisms are information‐processing mechanisms, cognitive science needs an account of (...) how the representations invoked in cognitive mechanisms carry information about contents, and I suggest that control theory offers the needed perspective on the relation of representations to contents. Second, I argue that cognitive science requires, but is still in search of, a catalog of cognitive operations that researchers can draw upon in explaining cognitive mechanisms. Last, I provide a new perspective on the relation of cognitive science to brain sciences, one which embraces both reductive research on neural components that figure in cognitive mechanisms and a concern with recomposing higher‐level mechanisms from their components and situating them in their environments. (shrink)

No âmbito do debate sobre as unidades envolvidas nos processos seletivos, a controvérsia sobre a possibilidade de comportamentos genuinamente altruístas tem um lugar destacado. Partindo de uma posição declaradamente pluralista, discutir-se-ão, neste artigo, algumas questões relevantes para esse tópico. Em primeiro lugar, o altruísmo será concebido como uma propriedade fenotípica dos grupos biológicos, e não apenas de seus membros. Essa caraterização levará, em um segundo momento, à discussão sobre a relação entre grupos em sociedades complexas, como é o caso das (...) sociedades humanas. Nesse contexto, será esboçado um modelo sobre como os comportamentos altruístas e os comportamentos egoístas se solapam em níveis diferentes das dinâmicas sociais. Finalmente, será sugerido um método de análise dirigido a determinar quais forças evolutivas entram em jogo, e sobre que tipo de comportamento, em casos empíricos concretos. In the debate on the units of selection, the controversy about the possibility of genuinely altruistic behavior has a prominent place. By assuming a pluralistic position, some relevant issues on the topic will be discussed in this paper. Firstly, altruism will be conceived as a phenotypic property of biological groups, not just of their members. Secondly, this characterization will be applied to the discussion on the relationship between groups in complex societies, as humans ones. In that context, a model about how altruistic and egoistic behaviors overlap at different levels of social dynamics will be outlined. Finally, a method aimed at determining what kinds of evolutionary forces influence behavior in empirical cases will be suggested. (shrink)

I argue in opposition to Sam Rakover that the current lack of fully adequate theories of the subjective and qualitative aspects of mind does not justify the adoption of what he calls “methodological dualism” (Rakover, this issue). Scientific understanding of consciousness requires the continuation of attempts to explain it in terms of the neural mechanisms that support it. It would be premature to adopt a methodological stance that could foreclose on the possibility of more reductionistic approaches. The effects of such (...) a stance would be especially pernicious insofar as methodological dualism lacks rigorous methods for testing specific models. (shrink)

Few areas of scientific investigation have spawned more alternative approaches than animal behavior: comparative psychology, ethology, behavioral ecology, sociobiology, behavioral endocrinology, behavioral neuroscience, neuroethology, behavioral genetics, cognitive ethology, developmental psychobiology---the list goes on. Add in the behavioral sciences focused on the human animal, and you can continue the list with ethnography, biological anthropology, political science, sociology, psychology (cognitive, social, developmental, evolutionary, etc.), and even that dismal science, economics. Clearly, no reasonable-length chapter can do justice to such a varied collection. We (...) have opted therefore to focus on three of these subdisciplines and to provide a somewhat historical tour of them, mentioning along the way the philosophical points that are of particular interest to us, but allowing the development of these points to be limited only by the imaginations of our readers. For readers seeking a more-traditional historical survey, see Dewsbury (1984a, b) and Burghardt (1985a). Our chosen brief is to write about comparative psychology, ethology, and cognitive ethology, although other approaches, especially neuroscience, will be mentioned where appropriate. These sciences are philosophically significant because they are enmeshed in ancient philosophical questions about the nature of mind and purposeful action and about the differences between humans and other animals. These sciences are also clustered because of their attention to mechanistic explanations of individual animal behavior as opposed to attempting to capture regularities at a population level, such as the game-theoretic strategic models popular among behavioral ecologists. (shrink)

Methodological reductionists practice ‘wannabe reductionism’. They claim that one should pursue reductionism, but never propose how. I integrate two strains in prior work to do so. Three kinds of activities are pursued as “reductionist”. “Successional reduction” and inter-level mechanistic explanation are legitimate and powerful strategies. Eliminativism is generally ill-conceived. Specific problem-solving heuristics for constructing inter-level mechanistic explanations show why and when they can provide powerful and fruitful tools and insights, but sometimes lead to erroneous results. I show how traditional metaphysical (...) approaches fail to engage how science is done. The methods used do so, and support a pragmatic and non-eliminativist realism. (shrink)

Many standard philosophical accounts of scientific practice fail to distinguish between modeling and other types of theory construction. This failure is unfortunate because there are important contrasts among the goals, procedures, and representations employed by modelers and other kinds of theorists. We can see some of these differences intuitively when we reflect on the methods of theorists such as Vito Volterra and Linus Pauling on the one hand, and Charles Darwin and Dimitri Mendeleev on the other. Much of Volterra's and (...) Pauling's work involved modeling; much of Darwin's and Mendeleev's did not. In order to capture this distinction, I consider two examples of theory construction in detail: Volterra's treatment of post-WWI fishery dynamics and Mendeleev's construction of the periodic system. I argue that modeling can be distinguished from other forms of theorizing by the procedures modelers use to represent and to study real-world phenomena: indirect representation and analysis. This differentiation between modelers and non-modelers is one component of the larger project of understanding the practice of modeling, its distinctive features, and the strategies of abstraction and idealization it employs. (shrink)

Plant and animal species are information-processing entities of such complexity, integration, and adaptive competence that it may be scientifically fruitful to consider them intelligent. The possibility arises from the analogy between learning and evolution, and from recent developments in evolutionary science, psychology and cognitive science. Species are now described as spatiotemporally localized individuals in an expanded hierarchy of biological entities. Intentional and cognitive abilities are now ascribed to animal, human, and artificial intelligence systems that process information adaptively, and that manifest (...) problem-solving abilities. The structural and functional similarities between such systems and species are extensive, although they “are usually obscured by population-genetic metaphors that have nonetheless contributed much to our understanding of evolution.In this target article, I use Sewall Wright's notion of the “adaptive landscape” to compare the performances of evolving species to those of intelligent organisms. With regard to their adaptive achievements and the kinds of processes by which they are achieved, biological species compare very favorably to intelligent animals by virtue of interactions between populations and their environments, between ontogeny and phylogeny, and between natural, interdemic, organic, and species selection. Addressing the question of whether species are intelligent could help to refine our ideas about species, evolution, and intelligence, and could open new lines of empirical and theoretical inquiry in many disciplines. (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)

This paper uses a number of examples of diverse types and functions of models in evolutionary biology to argue that the demarcation between theory and practice, or "theory model" and "data model," is often difficult to make. It is shown how both mathematical and laboratory models function as plausibility arguments, existence proofs, and refutations in the investigation of questions about the pattern and process of evolutionary history. I consider the consequences of this for the semantic approach to theories and theory (...) confirmation. The paper attempts to reconcile the insights of both critics and advocates of the semantic approach to theories. (shrink)

Richard Levins’ distinction between aggregate, composed and evolved systems acquires new significance as we recognize the importance of mechanistic explanation. Criteria for aggregativity provide limiting cases for absence of organization, so through their failure, can provide rich detectors for organizational properties. I explore the use of failures of aggregativity for the analysis of mechanistic systems in diverse contexts. Aggregativity appears theoretically desireable, but we are easily fooled. It may be exaggerated through approximation, conditions of derivation, and extrapolating from some conditions (...) of decomposition illegtimately to others. Evolved systems particularly may require analyses under alternative complementary decompositions. Exploring these conditions helps us to better understand the strengths and limits of reductionistic methods. (shrink)

This paper compares two well-known arguments in the units of selection literature, one due to , the other due to . Both arguments concern the legitimacy of averaging fitness values across contexts and making inferences about the level of selection on that basis. The first three sections of the paper shows that the two arguments are incompatible if taken at face value, their apparent similarity notwithstanding. If we accept Sober and Lewontin's criterion for when averaging genic fitnesses across diploid genotypes (...) is illegitmate, we cannot accept Sober and Wilson's criterion for when averaging individual fitnesses across groups is illegitimate, and vice versa. The final section suggests a possible way of reconciling the two arguments, by invoking an ambiguity in the concept of genic selection. (shrink)

Two alternative statistical approaches to modelling multi-level selection in nature, both found in the contemporary biological literature, are contrasted. The simple covariance approach partitions the total selection differential on a phenotypic character into within-group and between-group components, and identifies the change due to group selection with the latter. The contextual approach partitions the total selection differential into different components, using multivariate regression analysis. The two approaches have different implications for the question of what constitutes group selection and what does not. (...) I argue that the contextual approach is theoretically preferable. This has important implications for a number of issues in the philosophical debate about the levels of selection. Introduction Group selection and the covariance formulation of selection The contextual approach A modification of the simple covariance approach Consequences: frameshifting and additivity 5.1 Frameshifting 5.2 Additivity Conclusion. (shrink)

For a number of years, the debate in evolutionary biology over the ’levels of selection’ has attracted intense interest from philosophers of science. The main question concerns the level of the biological hierarchy at which natural selection occurs. Does selection act on organisms, genes, groups, colonies, demes, species, or some combination of these? According to traditional Darwinian theory the answer is the organism -- it is the differential survival and reproduction of individual organisms that drives the evolutionary process. But there (...) are alternative views too, including pluralist views which say that the choice between different levels of selection is often a matter of perspective, not empirical fact. (shrink)

The process of abstraction and concretisation is a label used for an explicative theory of scientific model-construction. In scientific theorising this process enters at various levels. We could identify two principal levels of abstraction that are useful to our understanding of theory-application. The first level is that of selecting a small number of variables and parameters abstracted from the universe of discourse and used to characterise the general laws of a theory. In classical mechanics, for example, we select position and (...) momentum and establish a relation amongst the two variables, which we call Newton’s 2nd law. The specification of the unspecified elements of scientific laws, e.g. the force function in Newton’s 2nd law, is what would establish the link between the assertions of the theory and physical systems. In order to unravel how and with what conceptual resources scientific models are constructed, how they function and how they relate to theory, we need a view of theory-application that can accommodate our constructions of representation models. For this we need to expand our understanding of the process of abstraction to also explicate the process of specifying force functions etc. This is the second principal level at which abstraction enters in our theorising and in which I focus. In this paper, I attempt to elaborate a general analysis of the process of abstraction and concretisation involved in scientific- model construction, and argue why it provides an explication of the construction of models of the nuclear structure. (shrink)

: This paper explores the calibration of laboratory models in population genetics as an experimental strategy for justifying experimental results and claims based upon them following Franklin (1986, 1990) and Rudge (1996, 1998). The analysis provided undermines Coyne et al.'s (1997) critique of Wade and Goodnight's (1991) experimental study of Wright's (1931, 1932) Shifting Balance Theory. The essay concludes by further demonstrating how this analysis bears on Diamond's (1986) claims regarding the weakness of laboratory experiments as evidence, and further how (...) the calibration strategy fits within Lloyd's (1987, 1988) account of the confirmation of ecological and evolutionary models. (shrink)

This article characterizes various species concepts in terms of set-theoretic models that license biological inferences and illustrates the logical connections among different species concepts. Species in this construal are abstract models, rather than biological or even tangible entities, and relate to individual organisms via representation, rather than the membership or mereological whole/part relationship. The proposal sheds new light on vexed issues of species and situates them within broader philosophical contexts of model selection, scientific representation, and scientific realism.

Plant and animal species are information-processing entities of such complexity, integration, and adaptive competence that it may be scientifically fruitful to consider them intelligent. The possibility arises from the analogy between learning and evolution, and from recent developments in evolutionary science, psychology and cognitive science. Species are now described as spatiotemporally localized individuals in an expanded hierarchy of biological entities. Intentional and cognitive abilities are now ascribed to animal, human, and artificial intelligence systems that process information adaptively, and that manifest (...) problem-solving abilities. The structural and functional similarities between such systems and species are extensive, although they “are usually obscured by population-genetic metaphors that have nonetheless contributed much to our understanding of evolution.In this target article, I use Sewall Wright's notion of the “adaptive landscape” to compare the performances of evolving species to those of intelligent organisms. With regard to their adaptive achievements and the kinds of processes by which they are achieved, biological species compare very favorably to intelligent animals by virtue of interactions between populations and their environments, between ontogeny and phylogeny, and between natural, interdemic, organic, and species selection. Addressing the question of whether species are intelligent could help to refine our ideas about species, evolution, and intelligence, and could open new lines of empirical and theoretical inquiry in many disciplines. (shrink)

Common monogenic genetic diseases, ones that have unexpectedly high frequencies in certain populations, have attracted a great number of conflicting evolutionary explanations. This paper will attempt to explain the mystery of why two particularly extensively studied common genetic diseases, Tay Sachs disease and cystic fibrosis, remain evolutionary mysteries despite decades of research. I review the most commonly cited evolutionary processes used to explain common genetic diseases: reproductive compensation, random genetic drift (in the context of founder effect), and especially heterozygote advantage. (...) The latter process has drawn a particularly large amount of attention, having so successfully explained the elevated frequency of sickle cell anemia in malaria-endemic areas. However, the empirical evidence for heterozygote advantage in other common genetic diseases is quite weak. I introduce and illustrate the significance of a hierarchy of genetic disease phenomena found within the genetic disease explanations, which include the phenomena: single mutation variants of a common genetic disease, single genetic diseases, and classes of diseases with related phenotypic effects. I demonstrate that some of the confusion over the explanations of common genetic diseases can be traced back to confusions over which phenomena are being explained. I proceed to briefly evaluate the existing evidence for two common human genetic diseases: Tay Sachs disease and cystic fibrosis. The above considerations will ultimately shed light on why these diseases’ evolutionary explanations remain so deeply unresolved after so such a great volume of research. (shrink)