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The paper works towards an account of explanatory integration in biology, using as a case study explanations of the evolutionary origin of novelties-a problem requiring the integration of several biological fields and approaches. In contrast to the idea that fields studying lower level phenomena are always more fundamental in explanations, I argue that the particular combination of disciplines and theoretical approaches needed to address a complex biological problem and which among them is explanatorily more fundamental varies with the problem pursued. (...) Solving a complex problem need not require theoretical unification or the stable synthesis of different biological fields, as items of knowledge from traditional disciplines can be related solely for the purposes of a specific problem. Apart from the development of genuine interfield theories, successful integration can be effected by smaller epistemic units (concepts, methods, explanations) being linked. Unification or integration is not an aim in itself, but needed for the aim of solving a particular scientific problem, where the problem's nature determines the kind of intellectual integration required. (shrink) | |
Many biological investigations are organized around a small group of species, often referred to as ‘model organisms’, such as the fruit fly Drosophila melanogaster. The terms ‘model’ and ‘modelling’ also occur in biology in association with mathematical and mechanistic theorizing, as in the Lotka–Volterra model of predator-prey dynamics. What is the relation between theoretical models and model organisms? Are these models in the same sense? We offer an account on which the two practices are shown to have different epistemic characters. (...) Theoretical modelling is grounded in explicit and known analogies between model and target. By contrast, inferences from model organisms are empirical extrapolations. Often such extrapolation is based on shared ancestry, sometimes in conjunction with other empirical information. One implication is that such inferences are unique to biology, whereas theoretical models are common across many disciplines. We close by discussing the diversity of uses to which model organisms are put, suggesting how these relate to our overall account. 1 Introduction2 Volterra and Theoretical Modelling3 Drosophila as a Model Organism4 Generalizing from Work on Model Organisms5 Phylogenetic Inference and Model Organisms6 Further Roles of Model Organisms6.1 Preparative experimentation6.2 Model organisms as paradigms6.3 Model organisms as theoretical models6.4 Inspiration for engineers6.5 Anchoring a research community7 Conclusion. (shrink) | |
Kinds that share historical properties are dubbed “historical kinds” or “etiological kinds,” and they have some distinctive features. I will try to characterize etiological kinds in general terms and briefly survey some previous philosophical discussions of these kinds. Then I will take a closer look at a few case studies involving different types of etiological kinds. Finally, I will try to understand the rationale for classifying on the basis of etiology, putting forward reasons for classifying phenomena on the basis of (...) diachronic features, thereby making a provisional case for considering at least some etiological kinds to be natural kinds. (shrink) | |
This essay analyzes and develops recent views about explanation in biology. Philosophers of biology have parted with the received deductive-nomological model of scientific explanation primarily by attempting to capture actual biological theorizing and practice. This includes an endorsement of different kinds of explanation (e.g., mathematical and causal-mechanistic), a joint study of discovery and explanation, and an abandonment of models of theory reduction in favor of accounts of explanatory reduction. Of particular current interest are philosophical accounts of complex explanations that appeal (...) to different levels of organismal organization and use contributions from different biological disciplines. The essay lays out one model that views explanatory integration across different disciplines as being structured by scientific problems. I emphasize the philosophical need to take the explanatory aims pursued by different groups of scientists into account, as explanatory aims determine whether different explanations are competing or complementary and govern the dynamics of scientific practice, including interdisciplinary research. I distinguish different kinds of pluralism that philosophers have endorsed in the context of explanation in biology, and draw several implications for science education, especially the need to teach science as an interdisciplinary and dynamic practice guided by scientific problems and explanatory aims. (shrink) | |
Evolutionary developmental biology (evo-devo) is considered a ‘mechanistic science,’ in that it causally explains morphological evolution in terms of changes in developmental mechanisms. Evo-devo is also an interdisciplinary and integrative approach, as its explanations use contributions from many fields and pertain to different levels of organismal organization. Philosophical accounts of mechanistic explanation are currently highly prominent, and have been particularly able to capture the integrative nature of multifield and multilevel explanations. However, I argue that evo-devo demonstrates the need for a (...) broadened philosophical conception of mechanisms and mechanistic explanation. Mechanistic explanation (in terms of the qualitative interactions of the structural parts of a whole) has been developed as an alternative to the traditional idea of explanation as derivation from laws or quantitative principles. Against the picture promoted by Carl Craver, that mathematical models describe but usually do not explain, my discussion of cases from the strand of evo-devo which is concerned with developmental processes points to qualitative phenomena where quantitative mathematical models are an indispensable part of the explanation. While philosophical accounts have focused on the actual organization and operation of mechanisms, properties of developmental mechanisms that are about how a mechanism reacts to modifications are of major evolutionary significance, including robustness, phenotypic plasticity, and modularity. A philosophical conception of mechanisms is needed that takes into account quantitative changes, transient entities and the generation of novel types of entities, feedback loops and complex interaction networks, emergent properties, and, in particular, functional-dynamical aspects of mechanisms, including functional (as opposed to structural) organization and distributed, system-wide phenomena. I conclude with general remarks on philosophical accounts of explanation. (shrink) | |
One powerful and influential approach to mental representation analyses representation in terms of biological functions, and biological functions in terms of histories of natural selection. This “teleosemantic” package, however, faces a familiar challenge. Surely representation depends only on the present-day structures of cognitive systems, and not on their historical provenance. “Swampman” drives the point home. Suppose a bolt of lightning creates an intrinsic duplicate of a human being in a steamy tropic swamp; will not this creature be representing its surroundings, (...) despite its lack of any selectional history? In this paper I shall answer this challenge by showing how a proper appreciation of the structure of natural kinds in general, and of mental representation in particular, implies that selectional histories are indeed built into the nature of mental representation. In particular, I shall address a recent argument by Peter Schulte against this general line of argument. (shrink) | |
In this chapter I lay out a notion of philosophical naturalism that aligns with pragmatism. It is developed and illustrated by a presentation of my views on natural kinds and my theory of concepts. Both accounts reflect a methodological naturalism and are defended not by way of metaphysical considerations, but in terms of their philosophical fruitfulness. A core theme is that the epistemic interests of scientists have to be taken into account by any naturalistic philosophy of science in general, and (...) any account of natural kinds and scientific concepts in particular. I conclude with general methodological remarks on how to develop and defend philosophical notions without using intuitions. (shrink) | |
The comparative method grants epistemic access to the biological past. Comparing lineages provides empirical traction on both hypotheses about particular lineages and models of trait evolution. Understanding this evidential role is important. Although philosophers have recently turned their attention to relations of descent, little work exists exploring the status of evidence from convergences. I argue that, where they exist, convergences play a central role in the confirmation of adaptive hypotheses. I focus on ‘analogous inferences’, show how such inferences ought to (...) be analysed and suggest three methods for strengthening their evidential weight. 1 Introduction2 Analogous Inferences2.1 Adaptive explanations and analogies2.2 Analogous inferences2.3 Scope, grain, and specificity3 Parallel Modelling, Integrated Explanations, and Convergent Modelling3.1 Parallel modelling3.2 Integrated explanations3.1 Convergent modelling4 Conclusion. (shrink) | |
Ethnographic analogy, the use of comparative data from anthropology to inform reconstructions of past human societies, has a troubled history. Archaeologists often express concern about, or outright reject, the practice—and sometimes do so in problematically general terms. This is odd, as the use of comparative data in archaeology is the same pattern of reasoning as the ‘comparative method’ in biology, which is a well-developed and robust set of inferences which play a central role in discovering the biological past. In pointing (...) out this continuity, I argue that there is no ‘special pleading’ on the part of archaeologists in this regard: biologists must overcome analogous epistemic difficulties in their use of comparative data. I then go on to emphasize the local, empirically tractable ways in which particular ethnographic analogies may be licensed. (shrink) | |
I argue that there is a tension between the claim that at least some kinds in the special sciences are multiply realized and the claim that the reason why kinds are prized by science is that they enter into a variety of different empirical generalizations. Nevertheless, I show that this tension ceases in the case of ‘cultural homologues’—such as specific ideologies, religions, and folk wisdom. I argue that the instances of such special science kinds do have several projectable properties in (...) common due to their shared history of reproduction, and that the social learning involved means that we should also expect these kinds to be multiply realized. (shrink) | |
My aim in this paper is twofold. First, I provide an analysis of the notion of cognitive homology. In contrast with the well-known concept of structural homology in biology—defined as the same structure in different animals regardless of form and function—the notion of cognitive homology captures the idea that the basic cognitive contribution of a given homologous brain structure tends to remain stable over long evolutionary time scales. Second, I argue that this notion provides a powerful conceptual tool for the (...) study of cognition. Since a cognitive homology will often consist of an evolutionarily conserved relationship between a homologous brain structure and its basic cognitive contribution, such structure–function mappings can be conceived as basic building blocks of human cognition. (shrink) | |
The dangers of character reification for cladistic inference are explored. The identification and analysis of characters always involves theory-laden abstraction—there is no theory-free “view from nowhere.” Given theory-ladenness, and given a real world with actual objects and processes, how can we separate robustly real biological characters from uncritically reified characters? One way to avoid reification is through the employment of objectivity criteria that give us good methods for identifying robust primary homology statements. I identify six such criteria and explore each (...) with examples. Ultimately, it is important to minimize character reification, because poor character analysis leads to dismal cladograms, even when proper phylogenetic analysis is employed. Given the deep and systemic problems associated with character reification, it is ironic that philosophers have focused almost entirely on phylogenetic analysis and neglected character analysis. (shrink) | |
The Mind-Brain Identity Theory lived a short life as a respectable philosophical position in the late 1950s, until Hilary Putnam developed his famous argument on the multiple realizability of mental states. The argument was, and still is, taken as the definitive demonstration of the falsity of Identity Theory and the foundation on which contemporary functionalist computational cognitive science was to be grounded. In this paper, in the wake of some contemporary philosophers, we reopen the case for Identity Theory and offer (...) a solution to the problem of multiple realizabilty. The solution is based on the necessity, at the time of establishing identity relations, of appealing to the notions of “homology” and “analogy” developed in the nineteenth century by Richard Owen. We also suggest that these notions are useful in order to correct certain shortcomings of some recent attempts at rebutting the Multiple Realizability argument. (shrink) | |
While philosophers tend to consider a single type of causal history, biologists distinguish between two kinds of causal history: evolutionary history and developmental history. This essay studies the peculiarity of development as a criterion for the individuation of biological traits and its relation to form, function, and evolution. By focusing on examples involving serial homologies and genetic reprogramming, we argue that morphology (form) and function, even when supplemented with evolutionary history, are sometimes insufficient to individuate traits. Developmental mechanisms bring in (...) a novel aspect to the business of classification—identity of process-type—according to which entities are type-identical across individuals and natural kinds in virtue of the fact that they form and develop through similar processes. These considerations bear important metaphysical implications and have potential applications in several areas of philosophy. (shrink) | |
Although classical evolutionary theory, i.e., population genetics and the Modern Synthesis, was already implicitly ‘gene-centred’, the organism was, in practice, still generally regarded as the individual unit of which a population is composed. The gene-centred approach to evolution only reached a logical conclusion with the advent of the gene-selectionist or gene’s eye view in the 1960s and 1970s. Whereas classical evolutionary theory can only work with (genotypically represented) fitness differences between individual organisms, gene-selectionism is capable of working with fitness differences (...) among genes within the same organism and genome. Here, we explore the explanatory potential of ‘intra-organismic’ and ‘intra-genomic’ gene-selectionism, i.e., of a behavioural-ecological ‘gene’s eye view’ on genetic, genomic and organismal evolution. First, we give a general outline of the framework and how it complements the—to some extent—still ‘organism-centred’ approach of classical evolutionary theory. Secondly, we give a more in-depth assessment of its explanatory potential for biological evolution, i.e., for Darwin’s ‘common descent with modification’ or, more specifically, for ‘historical continuity or homology with modular evolutionary change’ as it has been studied by evolutionary developmental biology (evo-devo) during the last few decades. In contrast with classical evolutionary theory, evo-devo focuses on ‘within-organism’ developmental processes. Given the capacity of gene-selectionism to adopt an intra-organismal gene’s eye view, we outline the relevance of the latter model for evo-devo. Overall, we aim for the conceptual integration between the gene’s eye view on the one hand, and more organism-centred evolutionary models (both classical evolutionary theory and evo-devo) on the other. (shrink) | |
George McGhee’s book “Convergent Evolution: limited forms most beautiful” provides an extensive survey of biological convergence. This paper has two main aims. First, it examines the theoretical claims McGhee makes about convergent evolution—specifically criticizing his use of a total morphospace to understand contingency and his assumption that functional constraints are non-contingent. Second, it sketches a group of important conceptual challenges facing researchers interested in convergence. | |
Although classical evolutionary theory, i.e., population genetics and the Modern Synthesis, was already implicitly ‘gene-centred’, the organism was, in practice, still generally regarded as the individual unit of which a population is composed. The gene-centred approach to evolution only reached a logical conclusion with the advent of the gene-selectionist or gene’s eye view in the 1960s and 1970s. Whereas classical evolutionary theory can only work with fitness differences between individual organisms, gene-selectionism is capable of working with fitness differences among genes (...) within the same organism and genome. Here, we explore the explanatory potential of ‘intra-organismic’ and ‘intra-genomic’ gene-selectionism, i.e., of a behavioural-ecological ‘gene’s eye view’ on genetic, genomic and organismal evolution. First, we give a general outline of the framework and how it complements the—to some extent—still ‘organism-centred’ approach of classical evolutionary theory. Secondly, we give a more in-depth assessment of its explanatory potential for biological evolution, i.e., for Darwin’s ‘common descent with modification’ or, more specifically, for ‘historical continuity or homology with modular evolutionary change’ as it has been studied by evolutionary developmental biology during the last few decades. In contrast with classical evolutionary theory, evo-devo focuses on ‘within-organism’ developmental processes. Given the capacity of gene-selectionism to adopt an intra-organismal gene’s eye view, we outline the relevance of the latter model for evo-devo. Overall, we aim for the conceptual integration between the gene’s eye view on the one hand, and more organism-centred evolutionary models on the other. (shrink) | |
Although sciences are often conceptualized in terms of theory confirmation and hypothesis testing, an equally important dimension of scientific reasoning is the structure of problems that guide inquiry. This problem structure is evident in several concepts central to evolutionary developmental biology (Evo-devo)—constraints, modularity, evolvability, and novelty. Because problems play an important role in biological practice, they should be included in biological pedagogy, especially when treating the issue of scientific controversy. A key feature of resolving controversy is synthesizing methodologies from different (...) biological disciplines to generate empirically adequate explanations. Concentrating on problem structure illuminates this interdisciplinarity in a way that is often ignored when science is taught only from the perspective of theory or hypothesis. These philosophical considerations can assist life science educators in their continuing quest to teach biology to the next generation. -/- . (shrink) | |
There have been several recent attempts to think about psychological kinds as homologies. Nevertheless, there are serious epistemic challenges for individuating homologous psychological kinds, or cognitive homologies. Some of these challenges are revealed when we look at competing claims of cognitive homology. This paper considers two competing homology claims that compare human anger with putative aggression systems of nonhuman animals. The competition between these hypotheses has been difficult to resolve in part because of what I call the boundary problem: boundaries (...) between instances of psychological kinds (e.g., anger and fear) cannot be directly observed. Thus, there are distinctive difficulties for individuating psychological kinds across lineages. I draw four conclusions from this case study: First, recent evidence from the neuroscience of fear suggests that one of the proposed homologies involves a straightforward conflation of anger and fear. Second, this conflation arises because of the boundary problem. Third, there is an implicit constraint on the operational criteria that is easy to overlook in the psychological case. In this case, ignoring the constraint is part of the problem. Fourth, this is a clear case in which knowledge of homology cannot be accumulated piecemeal. Identifying homologs of human anger requires identifying homologs of fear. (shrink) | |
To naturalize religion, we must identify what religion is, and what aspects of it we are trying to explain. In this paper, religious social institutional behavior is the explanatory target, and an explanatory hypothesis based on shared primate social dominance psychology is given. The argument is that various religious features, including the high status afforded the religious, and the high status afforded to deities, are an expression of this social dominance psychology in a context for which it did not evolve: (...) high-density populations made possible by agriculture. (shrink) | |
One of the central aims of science is explanation: scientists seek to uncover why things happen the way they do. This chapter addresses what kinds of explanations are formulated in biology, how explanatory aims influence other features of the field of biology, and the implications of all of this for biology education. Philosophical treatments of scientific explanation have been both complicated and enriched by attention to explanatory strategies in biology. Most basically, whereas traditional philosophy of science based explanation on derivation (...) from scientific laws, there are many biological explanations in which laws play little or no role. Instead, the field of biology is a natural place to turn for support for the idea that causal information is explanatory. Biology has also been used to motivate mechanistic accounts of explanation, as well as criticisms of that approach. Ultimately, the most pressing issue about explanation in biology may be how to account for the wide range of explanatory styles encountered in the field. This issue is crucial, for the aims of biological explanation influence a variety of other features of the field of biology. Explanatory aims account for the continued neglect of some central causal factors, a neglect that would otherwise be mysterious. This is linked to the persistent use of models like evolutionary game theory and population genetic models, models that are simplified to the point of unreality. These explanatory aims also offer a way to interpret many biologists’ total commitment to one or another methodological approach, and the intense disagreements that result. In my view, such debates are better understood as arising not from different theoretical commitments, but commitments to different explanatory projects. Biology education would thus be enriched by attending to approaches to biological explanation, as well as the unexpected ways that these explanatory aims influence other features of biology. I suggest five lessons for teaching about explanation in biology that follow from the considerations of this chapter. (shrink) | |
Biologists who work on the pig (_Sus scrofa_) take advantage of its similarity to humans by constructing the inferential and material means to traffic data, information and knowledge across the species barrier. Their research has been funded due to its perceived value for agriculture and medicine. Improving selective breeding practices, for instance, has been a driver of genomics research. The pig is also an animal model for biomedical research and practice, and is proposed as a source of organs for cross-species (...) transplantation: xenotransplantation. Genomics research has informed transplantation biology, which has itself motivated developments in genomics. Both have generated models of correspondences between the genomes of pigs and humans. Concerning genomics, I detail how researchers traverse species boundaries to develop representations of the pig genome, alongside ensuring that such representations are sufficiently porcine. In transplantation biology, the representations of the genomes of humans and pigs are used to detect and investigate immunologically-pertinent differences between the two species. These key differences can then be removed, to ‘humanise’ donor pigs so that they can become a safe and effective source of organs. In both of these endeavours, there is a tension between practices that ‘humanise’ the pig (or representations thereof) through using resources from human genomics, and the need to ‘dehumanise’ the pig to maintain distinctions for legal, ethical and scientific reasons. This paper assesses the ways in which this tension has been managed, observing the differences between its realisations across comparative pig genomics and transplantation biology, and considering the consequences of this. (shrink) | |
A number of areas of biology raise questions about what is of value in the natural environment and how we ought to behave towards it: conservation biology, environmental science, and ecology, to name a few. Based on my experience teaching students from these and similar majors, I argue that the field of environmental ethics has much to teach these students. They come to me with pent-up questions and a feeling that more is needed to fully engage in their subjects, and (...) I believe some exposure to environmental ethics can help focus their interests and goals. I identify three primary areas in which environmental ethics can con- tribute to their education. The first is an examination of who (or what) should be considered to be part of our moral community (i.e., the community to whom we owe direct duties). Is it humans only? Or does it include all sentient life? Or all life? Or ecosystems considered holistically? Often, readings implicitly assume one or more of these answers; the goal is to make the student more sensitive to these implicit claims and to get them to think about the different reasons that support them. The second area, related to the first, is the application of the different answers concerning the extent of the ethical community to real environmental issues and problems. Students need to be aware of how the different answers concerning the moral community can imply conflicting answers for how we should act in certain cases and to think about ways to move toward conflict resolution. The third area in which environmental ethics can contribute is a more conceptual one, focusing on central concepts such as biodiversity, sustainability, species, and ecosystems. Exploring and evaluating various meanings of these terms will make students more reflective and thoughtful citizens and biologists, sensitive to the implications that different conceptual choices make. (shrink) | |
What is a larva, if it is not what survives of an ancestor's adult, compressed into a transient pre‐reproductive phase, as suggested by Haeckel's largely disreputed model of evolution by recapitulation? A recently published article hypothesizes that larva and adult of holometabolous insects are developmental expressions of two different genomes coexisting in the same animal as a result of an ancient hybridization event between an onychophoran and a primitive insect with eventless post‐embryonic development. More likely, however, larvae originated from late (...) embryonic or early post‐embryonic stages of ancestors with direct development. Evolutionary novelties would thus be intercalary rather than terminal, with respect to the ancestor's ontogenetic schedule. This scenario, supported by current research on holometabolous insects and marine invertebrates with complex life cycles, offers a serious alternative to the traditional scenario (‘what is early in ontogeny is also early in phylogeny’) underlying the current perception of the evolution of genetic regulatory networks. (shrink) No categories | |
Neural reuse theories should interest developmental psychologists because these theories can potentially illuminate the developmental relations among psychological characteristics observed across the lifespan. Characteristics that develop by exploiting pre-existing neural circuits can be thought of as developmental homologues. And, understood in this way, the homology concept that has proven valuable for evolutionary biologists can be used productively to study psychological/behavioral development. | |
I develop an account of homology and homoplasy drawing on their use in biological inference and explanation. Biologists call on homology and homoplasy to infer character states, support adaptationist explanations, identify evolutionary novelties and hypothesize phylogenetic relationships. In these contexts, the concepts must be understood phylogenetically and kept separate: as they play divergent roles, overlap between the two ought to be avoided. I use these considerations to criticize an otherwise attractive view defended by Gould, Hall, and Ramsey & Peterson. By (...) this view, homology and homoplasy can only be delineated qua some level of description, and some homoplasies (parallelisms) are counted as homologous. I develop an account which retains the first, but rejects the second, aspect of that view. I then characterize parallelisms and convergences in terms of their causal role. By the Strict Continuity account, homology and homoplasy are defined phylogenetically and without overlaps, meeting my restriction. Convergence and parallelisms are defined as two types of homoplasy: convergent homoplasies are largely constrained by external factors, while parallelisms are due to internal constraints. (shrink) | |