Several analyses of biological function — for example, those of Williams, Millikan, and Kitcher — identify an item’s function with what natural selection designed it to do. Allen and Bekoff have disagreed, claiming that natural design is a special case of biological function. I argue that Allen and Bekoff’s account of natural design is unduly restrictive and that it fails to mark a principled distinction between function and design. I distinguish two approaches to the phenomenon of natural design — the (...) “trait-centered” approach of Allen and Bekoff and the “organism-centered” approach — and defend the latter. When design is understood according to the organism-centered approach, biological function and design are co-instantiated phenomena. (shrink)

In this paper I defend an etiological theory of biological functions (according to which the proper function of a trait is the effect for which it was selected by natural selection) against three objections which have been influential. I argue, contrary to Millikan, that it is wrong to base our defense of the theory on a rejection of conceptual analysis, for conceptual analysis does have an important role in philosophy of science. I also argue that biology requires a normative notion (...) of a "proper function", and that a normative notion is not ahistorical. (shrink)

I defend the historical definition of "function" originally given in my Language, Thought and Other Biological Categories (1984a). The definition was not offered in the spirit of conceptual analysis but is more akin to a theoretical definition of "function". A major theme is that nonhistorical analyses of "function" fail to deal adequately with items that are not capable of performing their functions.

Biological functions are dispositions or effects a trait has which explain the recent maintenance of the trait under natural selection. This is the "modern history" approach to functions. The approach is historical because to ascribe a function is to make a claim about the past, but the relevant past is the recent past; modern history rather than ancient.

How can we think about things in the outside world? There is still no widely accepted theory of how mental representations get their meaning. In light of pioneering research, Nicholas Shea develops a naturalistic account of the nature of mental representation with a firm focus on the subpersonal representations that pervade the cognitive sciences.

Ruth Garrett Millikan presents a strikingly original account of how we get to grips with the world in thought. Her question is Kant's 'How is knowledge possible?', answered from a contemporary naturalist standpoint. We begin with an understanding of what the world is like prior to cognition, then develop a theory of cognition within that world.

INTRODUCTION I What is teleology? If you ever look closely at an ants' nest, you will see an intricate network of pathways and chambers teeming with ...

This book is a critical survey of and guidebook to the literature on biological functions. It ties in with current debates and developments, and at the same time, it looks back on the state of discourse in naturalized teleology prior to the 1970s. It also presents three significant new proposals. First, it describes the generalized selected effects theory, which is one version of the selected effects theory, maintaining that the function of a trait consists in the activity that led to (...) its differential persistence or reproduction in a population, and not merely its differential reproduction. Secondly, it advances “within-discipline pluralism” (as opposed to between-discipline pluralism) a new form of function pluralism, which emphasizes the coexistence of function concepts within diverse biological sub-disciplines. Lastly, it provides a critical assessment of recent alternatives to the selected effects theory of function, namely, the weak etiological theory and the systems-theoretic theory. The book argues that, to the extent that functions purport to offer causal explanations for the existence of a trait, there are no viable alternatives to the selected effects view. -/- The debate about biological functions is still as relevant and important to biology and philosophy as it ever was. Recent controversies surrounding the ENCODE Project Consortium in genetics, the nature of psychiatric classification, and the value of ecological restoration, all point to the continuing relevance to biology of philosophical discussion about the nature of functions. In philosophy, ongoing debates about the nature of biological information, intentionality, health and disease, mechanism, and even biological trait classification, are closely related to debates about biological functions. (shrink)

This paper reviews the debate on the notion of biological function and on functional explanation as this takes place in philosophy. It describes the different perspectives, issues, intuitions, theories and arguments that have emerged. The author shows that the debate has been too heavily influenced by the concerns of a naturalistic philosophy of mind and argues that in order to improve our understanding of biology the attention should be shifted from the study of intuitions to the study of the actual (...) practice of biological inquiry. (shrink)

The Nature of Selection is a straightforward, self-contained introduction to philosophical and biological problems in evolutionary theory. It presents a powerful analysis of the evolutionary concepts of natural selection, fitness, and adaptation and clarifies controversial issues concerning altruism, group selection, and the idea that organisms are survival machines built for the good of the genes that inhabit them. "Sober's is the answering philosophical voice, the voice of a first-rate philosopher and a knowledgeable student of contemporary evolutionary theory. His book merits (...) broad attention among both communities. It should also inspire others to continue the conversation."-Philip Kitcher, Nature "Elliott Sober has made extraordinarily important contributions to our understanding of biological problems in evolutionary biology and causality. The Nature of Selection is a major contribution to understanding epistemological problems in evolutionary theory. I predict that it will have a long lasting place in the literature."-Richard C. Lewontin. (shrink)

"Modern History" versions of the etiological theory claim that in order for a trait X to have the proper function F, individuals with X must have been recently favored by natural selection for doing F (Godfrey-Smith 1994; Griffiths 1992, 1993). For many traits with prototypical proper functions, however, such recent selection may not have occurred: traits may have been maintained due to lack of variation or due to selection for other effects. I examine this flaw in Modern History accounts and (...) offer an alternative etiological theory, the Continuing Usefulness account, which appears to avoid such problems. (shrink)

In response to Germain argument that evolution by natural selection has a limited explanatory power in cancer, Lean and Plutynski have recently argued that many adaptations in cancer only make sense at the tumor level, and that cancer progression mirrors the major evolutionary transitions. While we agree that selection could potentially act at various levels of organization in cancers, we argue that tumor-level selection is unlikely to actually play a relevant role in our understanding of the somatic evolution of human (...) cancers. (shrink)

argue that natural selection does not explain the genotypic arid phenotypic properties of individuals. On this view, natural selection explains the adaptedness of individuals, not by explaining why the individuals that exist have the adaptations they do, but rather by explaining why the individuals that exist are the ones with those adaptations. This paper argues that this ‘Negative’ view of natural selection ignores the fact that natural selection is a cumulative selection process. So understood, it explains how the genetic sequences (...) that individuals inherit and that are responsible for their complex (and co-adapted) adaptations first arose in the gene-pool. (shrink)

This paper argues that a minimal notion of function and a notion of normal-proper function are used in explaining how bodies and brains operate. Neither is Cummins’ notion, as originally defined, and yet his is often taken to be the clearly relevant notion for such an explanatory context. This paper also explains how adverting to normal-proper functions, even if these are selected functions, can play a significant scientific role in the operational explanations of complex systems that physiologists and neurophysiologists provide, (...) despite a lack of relevant causal efficacy on the part of such functions. (shrink)

In this paper, we develop an organizational account that defines biological functions as causal relations subject to closure in living systems, interpreted as the most typical example of organizationally closed and differentiated self-maintaining systems. We argue that this account adequately grounds the teleological and normative dimensions of functions in the current organization of a system, insofar as it provides an explanation for the existence of the function bearer and, at the same time, identifies in a non-arbitrary way the norms that (...) functions are supposed to obey. Accordingly, we suggest that the organizational account combines the etiological and dispositional perspectives in an integrated theoretical framework. IntroductionDispositional ApproachesEtiological TheoriesBiological Self-maintenance Closure, teleology, and normativityOrganizational differentiationFunctions C1: Contributing to the maintenance of the organization C2: Producing the functional trait Implications and Objections Functional versus useful Dysfunctions, side effects, and accidental contributionsProper functions and selected effectsReproductionRelation with other ‘unitarian’ approachesConclusions. (shrink)

This collection of essays serves both as an introduction to Ruth Millikan’s much-discussed volume Language, Thought, and Other Biological Categories and as an extension and application of Millikan’s central themes, especially in the philosophy of psychology. The title essay discusses meaning rationalism and argues that rationality is not in the head, indeed, that there is no legitimate interpretation under which logical possibility and necessity are known a priori. In other essays, Millikan clarifies her views on the nature of mental representation, (...) explores whether human thought is a product of natural selection, examines the nature of behavior as studied by the behavioral sciences, and discusses the issues of individualism in psychology, psychological explanation, indexicality in thought, what knowledge is, and the realism/antirealism debate. Table of Contents Preface Acknowledgments Introduction 1 In Defense of Proper Functions 2 Propensities, Exaptations, and the Brain 3 Thoughts without Laws 4 Biosemantics 5 On Mentalese Orthography, Part 1 6 Compare and Contrast Dretske, Fodor, and Millikan on Teleosemantics 7 What Is Behavior? A Philosophical Essay on Ethology and Individualism in Psychology, Part 1 8 The Green Grass Growing All Around: A Philosophical Essay on Ethology and Individualism in Psychology, Part 2 9 Explanation in Biopsychology 10 Metaphysical Antirealism? 11 Truth Rules, Hoverflies, and the Kripke-Wittgenstein Paradox 12 Naturalist Reflections on Knowledge 13 The Myth of the Essential Indexical 14 White Queen Psychology; or, The Last Myth of the Given References Index. (shrink)

It is widely assumed that selection history accounts of function can support a fully reductive naturalization of functional properties. I argue that this assumption is false. A problem with the alternative causal role account of function in this context is that it invokes the teleological notion of a goal in analysing real function. The selection history account, if it is to have reductive status, must not do the same. But attention to certain cases of selection history in biology, specifically those (...) involving meiotic drive, shows that selection historical explanations are available in the case of items without any plausible function. Making contributions to goals, here fitness, must be introduced as an additional constraint in the analysis, either explicitly, or implicitly by appeal to individual functions or the selection-of/selection-for distinction. (shrink)

One of the major developments in cancer research in recent years has been the construction of models that treat cancer as a cellular population subject to natural selection. We expand on this idea, drawing upon multilevel selection theory. Cancer is best understood in our view from a multilevel perspective, as both a by-product of selection at other levels of organization, and as subject to selection at several levels of organization. Cancer is a by-product in two senses. First, cancer cells co-opt (...) signaling pathways that are otherwise adaptive at the organismic level. Second, cancer is also a by-product of features distinctive to the metazoan lineage: cellular plasticity and modularity. Applying the multilevel perspective in this way permits one to explain transitions in complexity and individuality in cancer progression. Our argument is a reply to Germain’s scepticism towards the explanatory relevance of natural selection for cancer. The extent to which cancer fulfills the conditions for being a paradigmatic Darwinian population depends on the scale of analysis, and the details of the purported selective scenario. Taking a multilevel perspective clarifies some of the complexities surrounding how to best understand the relevance of evolutionary thinking in cancer progression. (shrink)

Arguably, the most widely endorsed account of normative functions in philosophy of biology is an etiological theory that holds that the function of current traits is fixed by the past selection history of other traits of that type. The earlier formulations of this “selected-effects” theory had trouble accommodating vestigial traits. In order to remedy these difficulties, the influential recent selection or modern history selected-effects theory was introduced. This paper expands upon and strengthens the argument that this theory has trouble stemming (...) from recent “no variation” cases. In addition, several influential arguments for the necessity of including a selection requirement in a theory of normative biological functions are contested. It is suggested that accounting for biological functions in certain areas of biology does not require adverting to selection. (shrink)

Some properties are causally relevant for a certain effect, others are not. In this paper we describe a problem for our understanding of this notion and then offer a solution in terms of the notion of a program explanation.

The etiological approach to ‘proper functions’ in biology can be strengthened by relating it to Robert Cummins' general treatment of function ascription. The proper functions of a biological trait are the functions it is assigned in a Cummins-style functional explanation of the fitness of ancestors. These functions figure in selective explanations of the trait. It is also argued that some recent etiological theories include inaccurate accounts of selective explanation in biology. Finally, a generalization of the notion of selective explanation allows (...) an analysis of the proper functions of human artifacts. (shrink)

Elliott Sober''s selection for/selection of distinction has been widely used to clarify the idea that some properties of organisms are side-effects of selection processes. It has also been used, however, to choose between different descriptions of an evolutionary product when assigning biological functions to that product. We suggest that there is a characteristic error in these uses of the distinction. Complementary descriptions of function are misrepresented as mutually excluding one another. This error arises from a failure to appreciate that selection (...) processes can be described at several different theoretical levels. (shrink)

The aim of this paper is to assess the relevance of somatic evolution by natural selection to our understanding of cancer development. I do so in two steps. In the first part of the paper, I ask to what extent cancer cells meet the formal requirements for evolution by natural selection, relying on Godfrey-Smith’s (2009) framework of Darwinian populations. I argue that although they meet the minimal requirements for natural selection, cancer cells are not paradigmatic Darwinian populations. In the second (...) part of the paper, I examine the most important examples of adaptation in cancer cells. I argue that they are not significant accumulations of evolutionary changes, and that as a consequence natural selection plays a lesser role in their explanation. Their explanation, I argue, is best sought in the previously existing wiring of the healthy cells. (shrink)

Mainstream teleosemantics is the view that mental representation should be understood in terms of biological functions, which, in turn, should be understood in terms of selection processes. One of the traditional criticisms of teleosemantics is the problem of novel contents: how can teleosemantics explain our ability to represent properties that are evolutionarily novel? In response, some have argued that by generalizing the notion of a selection process to include phenomena such as operant conditioning, and the neural selection that underlies it, (...) we can resolve this problem. Here, we do four things: we develop this suggestion in a rigorous way through a simple example, we draw on recent neurobiological research to support its empirical plausibility, we defend the move from a host of objections in the literature, and we sketch how the picture can be extended to help us think about more complex “conceptual” representations and not just perceptual ones. (shrink)

Theories of function are conventionally divided up into historical and ahistorical ones. Proponents of ahistorical theories often cite the ahistoricity of their accounts as a major virtue. Here, I argue that none of the mainstream “ahistorical” accounts are actually ahistorical. All of them embed, implicitly or explicitly, an appeal to history. In Boorse’s goal-contribution account, history is latent in the idea of statistical-typicality. In the propensity theory, history is implicit in the idea of a species’ natural habitat. In the causal (...) role theory, history is required for making sense of dysfunction. I elaborate some consequences for the functions debate. (shrink)

Despite the voluminous literature on biological functions produced over the last 40 years, few philosophers have studied the concept of function as it is used in neuroscience. Recently, Craver (forthcoming; also see Craver 2001) defended the causal role theory against the selected effects theory as the most appropriate theory of function for neuroscience. The following argues that though neuroscientists do study causal role functions, the scope of that theory is not as universal as claimed. Despite the strong prima facie superiority (...) of the causal role theory, the selected effects theory (when properly developed) can handle many cases from neuroscience with equal facility. It argues this by presenting a new theory of function that generalizes the notion of a ‘selection process’ to include processes such as neural selection, antibody selection, and some forms of learning—that is, to include structures that have been differentially retained as well as those that have been differentially reproduced. This view, called the generalized selected effects theory of function, will be defended from criticism and distinguished from similar views in the literature. (shrink)

A common misunderstanding of the selected effects theory of function is that natural selection operating over an evolutionary time scale is the only functionbestowing process in the natural world. This construal of the selected effects theory conflicts with the existence and ubiquity of neurobiological functions that are evolutionary novel, such as structures underlying reading ability. This conflict has suggested to some that, while the selected effects theory may be relevant to some areas of evolutionary biology, its relevance to neuroscience is (...) marginal. This line of reasoning, however, neglects the fact that synapses, entire neurons, and potentially groups of neurons can undergo a type of selection analogous to natural selection operating over an evolutionary time scale. In the following, I argue that neural selection should be construed, by the selected effect theorist, as a distinct type of function-bestowing process in addition to natural selection. After explicating a generalized selected effects theory of function and distinguishing it from similar attempts to extend the selected effects theory, I do four things. First, I show how it allows one to identify neural selection as a distinct function-bestowing process, in contrast to other forms of neural structure formation such as neural construction. Second, I defend the view from one major criticism, and in so doing I clarify the content of the view. Third, I examine drug addiction to show the potential relevance of neural selection to neuroscientific and psychological research. Finally, I endorse a modest pluralism of function concepts within biology. (shrink)

I present and defend the generalized selected effects theory (GSE) of function. According to GSE, the function of a trait consists in the activity that contributed to its bearer’s differential reproduction, or differential retention, within a population. Unlike the traditional selected effects (SE) theory, it does not require that the functional trait helped its bearer reproduce; differential retention is enough. Although the core theory has been presented previously, I go significantly beyond those presentations by providing a new argument for GSE (...) and defending it from a recent objection. I also sketch its implications for teleosemantics and philosophy of medicine. (shrink)

We argue that ecology in general and biodiversity and ecosystem function research in particular need an understanding of functions which is both ahistorical and evolutionarily grounded. A natural candidate in this context is Bigelow and Pargetter’s evolutionary forward-looking account which, like the causal role account, assigns functions to parts of integrated systems regardless of their past history, but supplements this with an evolutionary dimension that relates functions to their bearers’ ability to thrive and perpetuate themselves. While Bigelow and Pargetter’s account (...) focused on functional organization at the level of organisms, we argue that such an account can be extended to functional organization at the community and ecosystem levels in a way that broadens the scope of the reconciliation between ecosystem ecology and evolutionary biology envisioned by many BEF researchers. By linking an evolutionary forward-looking account of functions to the persistence-based understanding of evolution defended by Bouchard and others, and to the theoretical research on complex adaptive systems, we argue that ecosystems, by forming more or less resilient assemblages, can evolve even while they do not reproduce and form lineages. We thus propose a Persistence Enhancing Propensity account of role functions in ecology to account for this overlap of evolutionary and ecological processes. (shrink)

In this paper, I try to develop an account of functions which might be of use to a biologist engaged in classifying and explaining natural phenomena. The most pressing difficulty facing such an account is the need to reconcile the normativity of function statements with their explanatory force. I consider two familiar accounts of function statements, offered by Andrew Woodfield and Larry Wright . I examine both accounts in search of the strongest possible formulation of each type of theory. I (...) fail to find a formulation of the ACT which is completely immune from counterexamples, but I do find a satisfactory formulation of the HT. In particular, I argue that the HT should incorporate a point central to the ACT that functions involve a means/end relationship between two devices. I then consider functional explanations: I argue that an ACT which holds that function statements are intrinsically explanatory cannot offer a satisfactory account. The account offered by Woodfield, who rejects this assumption, is adequate, though less attractive than the account suggested by the HT. I conclude by considering how the HT can account for functional norms. (shrink)

Formulations of the essential commitment of the etiological theory of functions have varied significantly, with some individual authors' formulations even varying from one place to another. The logical geography of these various formulations is different from what is standardly assumed; for they are not stylistic variants of the same essential commitment, but stylistic variants of two non-equivalent versions of the etiological theory. I distinguish these “strong” and “weak” versions of the etiological theory (which differ with respect to the role of (...) selection in their definitions of function), draw out their respective implications, and argue that the weak version is to be preferred to the strong. (shrink)

Following Wallace’s suggestion, Darwin framed his theory using Spencer’s expression “survival of the fittest”. Since then, fitness occupies a significant place in the conventional understanding of Darwinism, even though the explicit meaning of the term ‘fitness’ is rarely stated. In this paper I examine some of the different roles that fitness has played in the development of the theory. Whereas the meaning of fitness was originally understood in ecological terms, it took a statistical turn in terms of reproductive success throughout (...) the 20th Century. This has lead to the ever-increasing importance of sexually reproducing organisms and the populations they compose in evolutionary explanations. I will argue that, moving forward, evolutionary theory should look back at its ecological roots in order to be more inclusive in the type of systems it examines. Many biological systems can only be satisfactorily accounted for by offering a non-reproductive account of fitness. This argument will be made by examining biological systems with very small or transient population structures. I argue this has significant consequences for how we define Darwinism, increasing the significance of survival over that of reproduction. (shrink)

Ecological fitness has been suggested to provide a unifying definition of fitness. However, a metric for this notion of fitness was in most cases unavailable except by proxy with differential reproductive success. In this article, I show how differential persistence of lineages can be used as a way to assess ecological fitness. This view is inspired by a better understanding of the evolution of some clonal plants, colonial organisms, and ecosystems. Differential persistence shows the limitation of an ensemblist noncausal understanding (...) of evolution. Causal explanations are necessary to understand the evolution by natural selection of these biological systems. †To contact the author write to: Department of Philosophy, University of Montreal, P.O. Box 6128, Station Centre‐Ville, Montreal, Quebec, H3C 3J7 Canada; e‐mail: [email protected]. (shrink)

The goal of this essay is to put forward an original theory of artifact function, which takes on board the results of the debate on the notion of biological function and also accommodates the distinctive aspects of artifacts. More precisely, the paper develops and defends the Dual-Aspect Theory, which is a monist account according to which an artifact’s function depends on intentional and reproductive aspects. It is argued that this approach meets a set of theoretical and meta-theoretical desiderata and is (...) superior to alternative views. (shrink)

Ruth Millikan and others advocate theories which attempt to naturalize wide mental content (e.g. beliefs.

Preface ix 1 Meaning and the Means to an Understanding of Ends 2 Why Is an Eye? 21 3 Adaptationism and Engineering 39 4 On Five "-Isms" 67 5 Function, Selection, and Explanation 87 6 Deflating Function 119 7 Artifacts and Organisms 139 References 167 Index 177.

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

In this adventurous contribution to the project of combining philosophy and biology to understand the mind, Carolyn Price investigates what it means to say that mental states--like thoughts, wishes, and perceptual experiences--are about things in the natural world. Her insight into this deep philosophical problem offers a novel teleological account of intentional content, grounded in and shaped by a carefully constructed theory of functions. Along the way she defends her view from recent objections to teleological theories and indicates how it (...) might be applied to notable problems in the philosophy of mind. (shrink)