Biologists studying complex causal systems typically identify some factors as causes and treat other factors as background conditions. For example, when geneticists explain biological phenomena, they often foreground genes and relegate the cellular milieu to the background. But factors in the milieu are as causally necessary as genes for the production of phenotypic traits, even traits at the molecular level such as amino acid sequences. Gene-centered biology has been criticized on the grounds that because there is parity among causes, the (...) “privileging” of genes reflects a reductionist bias, not an ontological difference. The idea that there is an ontological parity among causes is related to a philosophical puzzle identified by John Stuart Mill: what, other than our interests or biases, could possibly justify identifying some causes as the actual or operative ones, and other causes as mere background? The aim of this paper is to solve this conceptual puzzle and to explain why there is not an ontological parity among genes and the other factors. It turns out that solving this puzzle helps answer a seemingly unrelated philosophical question: what kind of causal generality matters in biology? (shrink)

This book explains the relationship between intelligence and environmental complexity, and in so doing links philosophy of mind to more general issues about the relations between organisms and environments, and to the general pattern of 'externalist' explanations. The author provides a biological approach to the investigation of mind and cognition in nature. In particular he explores the idea that the function of cognition is to enable agents to deal with environmental complexity. The history of the idea in the work of (...) Dewey and Spencer is considered, as is the impact of recent evolutionary theory on our understanding of the place of mind in nature. (shrink)

According to many biologists, explaining the evolution of morphological novelty and behavioral innovation are central endeavors in contemporary evolutionary biology. These endeavors are inherently multidisciplinary but also have involved a high degree of controversy. One key source of controversy is the definitional diversity associated with the concept of evolutionary novelty, which can lead to contradictory claims (a novel trait according to one definition is not a novel trait according to another). We argue that this diversity should be interpreted in light (...) of a different epistemic role played by the concept of evolutionary novelty—the structuring of a problem space or setting of an explanatory agenda—rather than the concept’s capacity to categorize traits as novel. This distinctive role is consistent with the definitional diversity and shows that the concept of novelty benefits ongoing investigation by focusing attention on answering different questions related to comprehending the origins of novelty. A review of recent theoretical and empirical work on evolutionary novelty confirms this interpretation. (shrink)

Woodward's long awaited book is an attempt to construct a comprehensive account of causation explanation that applies to a wide variety of causal and explanatory claims in different areas of science and everyday life. The book engages some of the relevant literature from other disciplines, as Woodward weaves together examples, counterexamples, criticisms, defenses, objections, and replies into a convincing defense of the core of his theory, which is that we can analyze causation by appeal to the notion of manipulation.

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)

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.

Homology is a biological sameness relation that is purported to hold in the face of changes in form, composition, and function. In spite of the centrality and importance of homology, there is no consensus on how we should understand this concept. The two leading views of homology, the genealogical and developmental accounts, have significant shortcomings. We propose a new account, the hierarchical-dependency account of homology, which avoids these shortcomings. Furthermore, our account provides for continuity between special, general, and serial homology.

Stephen Jay Gould argued that replaying the “tape of life” would result in a radically different evolutionary outcome. Some biologists and philosophers, however, have pointed to convergent evolution as evidence for robust replicability in macroevolution. These authors interpret homoplasy, or the independent origination of similar biological forms, as evidence for the power of natural selection to guide form toward certain morphological attractors, notwithstanding the diversionary tendencies of drift and the constraints of phylogenetic inertia. In this paper, I consider the implications (...) of homoplasy for the debate over the nature of macroevolution. I argue that once the concepts of contingency and convergence are fleshed out, it becomes clear that many instances of homoplasy fail to negate Gould’s overarching thesis, and may in fact support a Gouldian view of life. My argument rests on the distinction between parallelism and convergence, which I defend against a recent challenge from developmental biology. I conclude that despite the difficulties in defining and identifying parallelism, the concept remains useful and relevant to the contingency controversy insofar as it underscores the common developmental origins of iterated evolution. (shrink)

This paper attempts to elucidate three characteristics of causal relationships that are important in biological contexts. Stability has to do with whether a causal relationship continues to hold under changes in background conditions. Proportionality has to do with whether changes in the state of the cause “line up” in the right way with changes in the state of the effect and with whether the cause and effect are characterized in a way that contains irrelevant detail. Specificity is connected both to (...) David Lewis’ notion of “influence” and also with the extent to which a causal relation approximates to the ideal of one cause–one effect. Interrelations among these notions and their possible biological significance are also discussed. (shrink)

After the discovery of the structure of DNA in 1953, scientists working in molecular biology embraced reductionism—the theory that all complex systems can be understood in terms of their components. Reductionism, however, has been widely resisted by both nonmolecular biologists and scientists working outside the field of biology. Many of these antireductionists, nevertheless, embrace the notion of physicalism—the idea that all biological processes are physical in nature. How, Alexander Rosenberg asks, can these self-proclaimed physicalists also be antireductionists? With clarity and (...) wit, Darwinian Reductionism navigates this difficult and seemingly intractable dualism with convincing analysis and timely evidence. In the spirit of the few distinguished biologists who accept reductionism—E. O. Wilson, Francis Crick, Jacques Monod, James Watson, and Richard Dawkins—Rosenberg provides a philosophically sophisticated defense of reductionism and applies it to molecular developmental biology and the theory of natural selection, ultimately proving that the physicalist must also be a reductionist. (shrink)

This contribution to the adaptationism debate elaborates the nature of constraints and their importance in evolutionary explanation and argues that the adaptationism debate should be limited to the issue of how to privilege causes in evolutionary explanation. I argue that adaptationist explanations are deeply conceptually dependent on developmental constraints, and explanations that appeal to constraints are dependant on the results of natural selection. I suggest these explanations should be integrated into the framework of historical causal explanation. Each strategy explicitly appeals (...) to some aspect of the evolutionary process, while implicitly appealing to others. Thus, adaptationists and anti-adaptationists can offer complementary causal explanations of the same explanandum. This eliminates much of the adaptationism debate and explains why its adversaries regularly agree with each other more than they would like. The adaptationism issue that remains is a species of the general issue of how to privilege causes in explanation. I show how a proposed solution to this general problem might be brought to bear on evolutionary explanations, and investigate some difficulties that might arise due to the nature of the evolutionary process. (shrink)

Philosophers of evolutionary biology favor the so-called etiological concept of function according to which the function of a trait is its evolutionary purpose, defined as the effect for which that trait was favored by natural selection. We term this the selected effect (SE) analysis of function. An alternative account of function was introduced by Robert Cummins in a non-evolutionary and non-purposive context. Cummins''s account has received attention but little support from philosophers of biology. This paper will show that a similar (...) non-purposive concept of function, which we term causal role (CR) function, is crucial to certain research programs in evolutionary biology, and that philosophical criticisms of Cummins''s concept are ineffective in this scientific context. Specifically, we demonstrate that CR functions are a vital and ineliminable part of research in comparative and functional anatomy, and that biological categories used by anatomists are not defined by the application of SE functional analysis. Causal role functions are non-historically defined, but may themselves be used in an historical analysis. Furthermore, we show that a philosophical insistence on the primary of SE functions places practicing biologists in an untenable position, as such functions can rarely be demonstrated (in contrast to CR functions). Biologists who study the form and function of organismal design recognize that it is virtually impossible to identify the past action of selection on any particular structure retrospectively, a requirement for recognizing SE functions. (shrink)

A number of authors have pointed to “convergent evolution” as evidence for the central role of natural selection in shaping predictable trajectories of macroevolution. However, there are numerous conceptual and empirical difficulties that arise in broadly appealing to the frequency of homoplasy as evidence for a non-contingently constrained adaptational design space. Most important is the need to distinguish between convergent (externally constrained) and parallel (internally constrained) evolution, and to consider how the respective frequencies of these significantly different sources of homoplasy (...) affect a strong adaptationist view of life. In this paper, I critically evaluate Simon Conway Morris’s use of the homoplasy literature to support his argument for a non-contingent, counterfactually stable account of macroevolutionary pattern. In so doing, I offer a conception of parallelism which avoids the charge that it differs from convergence merely in degree and not in kind. I argue that although organisms sharing a homoplastic trait will also share varying degrees of homology, it is the underlying developmental homology with respect to the generators directly causally responsible for the homoplastic event that defines parallel evolution and non-arbitrarily distinguishes it from convergence. The notion of “screening-off” is used to distinguish the proximal generators of a homoplastic trait from its more distal genetic causes (such as a master control gene). (shrink)

Philosophical discussions of biological classification have failed to recognise the central role of homology in the classification of biological parts and processes. One reason for this is a misunderstanding of the relationship between judgments of homology and the core explanatory theories of biology. The textbook characterisation of homology as identity by descent is commonly regarded as a definition. I suggest instead that it is one of several attempts to explain the phenomena of homology. Twenty years ago the ‘new experimentalist’ movement (...) in philosophy of science drew attention to the fact that many experimental phenomena have a ‘life of their own’: the conviction that they are real is not dependent on the theories used to characterise and explain them. I suggest that something similar can be true of descriptive phenomena, and that many homologies are phenomena of this kind. As a result the descriptive biology of form and function has a life of its own—a degree of epistemological independence from the theories that explain form and function. I also suggest that the two major ‘homology concepts’ in contemporary biology, usually seen as two competing definitions, are in reality complementary elements of the biological explanation of homology. (shrink)

In recent years, the concept of evolvability has been gaining in prominence both within evolutionary developmental biology (evo-devo) and the broader field of evolutionary biology. Despite this, there remains considerable disagreement about what evolvability is. This article offers a solution to this problem. I argue that, in focusing too closely on the role played by evolvability as an explanandum in evo-devo, existing philosophical attempts to clarify the evolvability concept have been overly narrow. Within evolutionary biology more broadly, evolvability offers a (...) robust explanation for the evolutionary trajectories of populations. Evolvability is an abstract, robust, dispositional property of populations, which captures the joint causal influence of their internal features on the outcomes of evolution (as opposed to the causal influence of selection, which is often characterized as external). When considering the nature of the physical basis of this disposition, it becomes clear that the many existing definitions of evolvability at play within evo-devo should be understood as capturing only aspects of a much broader phenomenon. 1 Introduction2 The Problem of Evolvability3 The Theoretical Role of Evolvability in Evolutionary Biology3.1 The explanatory targets of evolutionary biology3.2 Selection-based explanations3.3 Lineage explanations3.4 Evolvability-based explanations3.5 What properties must evolvability have?4 What Evolvability Really Is4.1 Making sense of ft4.2 Making sense of x and b5 What of the Limbs? The Power of E6 Conclusion. (shrink)

The philosophical theory of scientific explanation proposed here involves a radically new treatment of causality that accords with the pervasively statistical character of contemporary science. Wesley C. Salmon describes three fundamental conceptions of scientific explanation--the epistemic, modal, and ontic. He argues that the prevailing view is untenable and that the modal conception is scientifically out-dated. Significantly revising aspects of his earlier work, he defends a causal/mechanical theory that is a version of the ontic conception. Professor Salmon's theory furnishes a robust (...) argument for scientific realism akin to the argument that convinced twentieth-century physical scientists of the existence of atoms and molecules. To do justice to such notions as irreducibly statistical laws and statistical explanation, he offers a novel account of physical randomness. The transition from the "reviewed view" of scientific explanation to the causal/mechanical model requires fundamental rethinking of basic explanatory concepts. (shrink)

I adopt a cladistic view of species, and explore the possibility that there exists an equally valuable cladistic view of organismic traits. This suggestion seems to run counter to the stress on functional views of biological traits in recent work in philosophy and psychology. I show how the tension between these two views can be defused with a multilevel view of biological explanation. Despite the attractions of this compromise, I conclude that we must reject it, and adopt an essentially cladistic (...) conception of biological traits. (shrink)

In recent years, the concept of evolvability has been gaining in prominence both within evolutionary developmental biology (evo-devo) and the broader field of evolutionary biology. Despite this, there remains considerable disagreement about what evolvability is. This article offers a solution to this problem. I argue that, in focusing too closely on the role played by evolvability as an explanandum in evo-devo, existing philosophical attempts to clarify the evolvability concept have been overly narrow. Within evolutionary biology more broadly, evolvability offers a (...) robust explanation for the evolutionary trajectories of populations. Evolvability is an abstract, robust, dispositional property of populations, which captures the joint causal influence of their internal features on the outcomes of evolution (as opposed to the causal influence of selection, which is often characterized as external). When considering the nature of the physical basis of this disposition, it becomes clear that the many existing definitions of evolvability at play within evo-devo should be understood as capturing only aspects of a much broader phenomenon. 1 Introduction2 The Problem of Evolvability3 The Theoretical Role of Evolvability in Evolutionary Biology 3.1 The explanatory targets of evolutionary biology 3.2 Selection-based explanations 3.3 Lineage explanations 3.4 Evolvability-based explanations 3.5 What properties must evolvability have?4 What Evolvability Really Is 4.1 Making sense of ft 4.2 Making sense of x and b5 What of the Limbs? The Power of E6 Conclusion. (shrink)

Life's Solution builds a persuasive case for the predictability of evolutionary outcomes. The case rests on a remarkable compilation of examples of convergent evolution, in which two or more lineages have independently evolved similar structures and functions. The examples range from the aerodynamics of hovering moths and hummingbirds to the use of silk by spiders and some insects to capture prey. Going against the grain of Darwinian orthodoxy, this book is a must read for anyone grappling with the meaning of (...) evolution and our place in the Universe. Simon Conway Morris is the Ad Hominen Professor in the Earth Science Department at the University of Cambridge and a Fellow of St. John's College and the Royal Society. His research focuses on the study of constraints on evolution, and the historical processes that lead to the emergence of complexity, especially with respect to the construction of the major animal body parts in the Cambrian explosion. Previous books include The Crucible of Creation (Getty Center for Education in the Arts, 1999) and co-author of Solnhofen (Cambridge, 1990). Hb ISBN (2003) 0-521-82704-3. (shrink)

There is still a great deal of debate over what counts as a constraint and about how to assess experimentally the relative importance of constraints and selection in evolutionary history. I will argue that the notion of a constraint on variation, and thus the selection-constraint distinction, depends on two specifications: (1) what counts as a variant -- constraints limit or bias the production of what? and (2) range of assessment -- over what range of times or conditions is the variation (...) assessed? Specifications 1 and 2 help us to understand empirical work on the relative importance of constraint and selection in evolution. (shrink)

This article argues that at least very many judgments of homology rest on prior attributions of selected‐effect (SE) function, and that many of the “parts” of biological systems that are rightly classified as homologous are constituted by (are so classified in virtue of) their consequence etiologies. We claim that SE functions are often used in the prior identification of the parts deemed to be homologous and are often used to differentiate more restricted homologous kinds within less restricted ones. In doing (...) so, we discuss recent criticism of this view that has been offered (especially that offered by Paul Griffiths). *Received July 2007; revised November 2008. †To contact the authors, please write to: Department of Philosophy, 201 West Duke Building, Box 90743, Durham, NC 27708; e‐mail: [email protected] or [email protected]. (shrink)

Editors' introduction to the special issue on homology (Biology and Philosophy Vol. 22, Issue 5, 2007).

I defend the view that many biological categories are defined by homology against a series of arguments designed to show that all biological categories are defined, at least in part, by selected function. I show that categories of homology are `abnormality inclusive'—something often alleged to be unique to selected function categories. I show that classifications by selected function are logically dependent on classifications by homology, but not vice-versa. Finally, I reject the view that biologists must use considerations of selected function (...) to abstract away from variation and pathology to form a canonical description of a class of biological systems. (shrink)

By linking the concepts of homology and morphological organization to evolvability, this paper attempts to (1) bridge the gap between developmental and phylogenetic approaches to homology and to (2) show that developmental constraints and natural selection are compatible and in fact complementary. I conceive of a homologue as a unit of morphological evolvability, i.e., as a part of an organism that can exhibit heritable phenotypic variation independently of the organism’s other homologues. An account of homology therefore consists in explaining how (...) an organism’s developmental constitution results in different homologues/characters as units that can evolve independently of each other. The explanans of an account of homology is developmental, yet the very explanandum is an evolutionary phenomenon: evolvability in a character-by-character fashion, which manifests itself in phylogenetic patterns as recognized by phylogenetic approaches to homology. While developmental constraints and selection have often been viewed as antagonistic forces, I argue that both are complementary as they concern different parts of the evolutionary process. Developmental constraints, conceived of as the presence of the same set of homologues across phenotypic change, pertain to how heritable variation can be generated in the first place (evolvability), while natural selection operates subsequently on the produced variation. (shrink)

Determining whether a homoplastic trait is the result of convergence or parallelism is central to many of the most important contemporary discussions in biology and philosophy: the relation between evolution and development, the importance of constraints on variation, and the role of contingency in evolution. In this article, I show that two recent attempts to draw a black-or-white distinction between convergence and parallelism fail, albeit for different reasons. Nevertheless, I argue that we should not be afraid of gray areas: a (...) clarified version of S. J. Gould's earlier account, based on a separation of underlying developmental mechanisms from the realized trait, still represents a useful approach. (shrink)