The notion that natural selection is a process of fitness maximization gets a bad press in population genetics, yet in other areas of biology the view that organisms behave as if attempting to maximize their fitness remains widespread. Here I critically appraise the prospects for reconciliation. I first distinguish four varieties of fitness maximization. I then examine two recent developments that may appear to vindicate at least one of these varieties. The first is the ‘new’ interpretation of Fisher's fundamental theorem (...) of natural selection, on which the theorem is exactly true for any evolving population that satisfies some minimal assumptions. The second is the Formal Darwinism project, which forges links between gene frequency change and optimal strategy choice. In both cases, I argue that the results fail to establish a biologically significant maximization principle. I conclude that it may be a mistake to look for universal maximization principles justified by theory alone. A more promising approach may be to find maximization principles that apply conditionally and to show that the conditions were satisfied in the evolution of particular traits. (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.

The biological functions debate is a perennial topic in the philosophy of science. In the first full-length account of the nature and importance of biological functions for many years, Justin Garson presents an innovative new theory, the 'generalized selected effects theory of function', which seamlessly integrates evolutionary and developmental perspectives on biological functions. He develops the implications of the theory for contemporary debates in the philosophy of mind, the philosophy of medicine and psychiatry, the philosophy of biology, and biology itself, (...) addressing issues ranging from the nature of mental representation to our understanding of the function of the human genome. Clear, jargon-free, and engagingly written, with accessible examples and explanatory diagrams to illustrate the discussion, his book will be highly valuable for readers across philosophical and scientific disciplines. (shrink)

Issues concerning scientific explanation have been a focus of philosophical attention from Pre- Socratic times through the modern period. However, recent discussion really begins with the development of the Deductive-Nomological (DN) model. This model has had many advocates (including Popper 1935, 1959, Braithwaite 1953, Gardiner, 1959, Nagel 1961) but unquestionably the most detailed and influential statement is due to Carl Hempel (Hempel 1942, 1965, and Hempel & Oppenheim 1948). These papers and the reaction to them have structured subsequent discussion concerning (...) scientific explanation to an extraordinary degree. After some general remarks by way of background and orientation (Section 1), this entry describes the DN model and its extensions, and then turns to some well-known objections (Section 2). It next describes a variety of subsequent attempts to develop alternative models of explanation, including Wesley Salmon's Statistical Relevance (Section 3) and Causal Mechanical (Section 4) models and the Unificationist models due to Michael Friedman and Philip Kitcher (Section 5). Section 6 provides a summary and discusses directions for future work. (shrink)

Samir Okasha offers a critical study of agential thinking in biology, where evolved organisms are seen as agents pursuing a goal. He examines the justification for transposing concepts from rational humans to the biological world, and considers whether agential thinking is mere anthropomorphism or plays a more intellectual role in the science.

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.

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)

Philosophy of biology is often said to have emerged in the last third of the twentieth century. Prior to this time, it has been alleged that the only authors who engaged philosophically with the life sciences were either logical empiricists who sought to impose the explanatory ideals of the physical sciences onto biology, or vitalists who invoked mystical agencies in an attempt to ward off the threat of physicochemical reduction. These schools paid little attention to actual biological science, and as (...) a result philosophy of biology languished in a state of futility for much of the twentieth century. The situation, we are told, only began to change in the late 1960s and early 1970s, when a new generation of researchers began to focus on problems internal to biology, leading to the consolidation of the discipline. In this paper we challenge this widely accepted narrative of the history of philosophy of biology. We do so by arguing that the most important tradition within early twentieth-century philosophy of biology was neither logical empiricism nor vitalism, but the organicist movement that flourished between the First and Second World Wars. We show that the organicist corpus is thematically and methodologically continuous with the contemporary literature in order to discredit the view that early work in the philosophy of biology was unproductive, and we emphasize the desirability of integrating the historical and contemporary conversations into a single, unified discourse. (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)

Natural selection comes in degrees. Some biological traits are subjected to stronger selective force than others, selection on particular traits waxes and wanes over time, and some groups can only undergo an attenuated kind of selective process. This has downstream consequences for any notions that are standardly treated as binary but depend on natural selection. For instance, the proper function of a biological structure can be defined as what caused that structure to be retained by natural selection in the past. (...) We usually think of proper functions in binary terms: storing bile is a function of the gall bladder, but making stones is not. However, if functions arise through natural selection, and natural selection comes in degrees, then a binary approach to proper functions is in tension with the biological facts. In order to resolve this tension, we need to revise our standard accounts of proper function. In particular, we may have to seriously consider the possibility that functions themselves come in degrees, in spite of the ramifications this will have for the way we speak about functions and related concepts such as dysfunction, disease, and teleosemantic content. (shrink)

Game theory has a prominent role in evolutionary biology, in particular in the ecological study of various phenomena ranging from conflict behaviour to altruism to signalling and beyond. The two central methodological tools in biological game theory are the concepts of Nash equilibrium and evolutionarily stable strategy. While both were inspired by a dynamic conception of evolution, these concepts are essentially static—they only show that a population is uninvadable, but not that a population is likely to evolve. In this article, (...) we argue that a static methodology can lead to misleading views about dynamic evolutionary processes. We advocate, instead, a more pluralistic methodology, which includes both static and dynamic game theoretic tools. Such an approach provides a more complete picture of the evolution of strategic behaviour. 1 Introduction2 The Equilibrium Methodology3 Common Interest Signalling3.1 Lewis’s signalling game3.2 Static analysis3.3 Dynamic analysis4 The Sir Philip Sidney Game4.1 Static analysis4.2 Other equilibria4.3 Dynamic analysis5 Related Literature6 Static and Dynamic Approaches. (shrink)

Signaling games provide basic insights into some fundamental questions concerning the explanation of meaning. They can be analyzed in terms of rational choice theory and in terms of evolutionary game theory. It is argued that an evolutionary approach provides better explanations for the emergence of simple communication systems. To substantiate these arguments, I will look at models similar to those of Skyrms (2000) and Komarova and Niyogi (2004) and study their dynamical properties. My results will lend partial support to the (...) thesis that evolution leads to communication. In general, states of partial communication may evolve with positive probability under standard evolutionary dynamics. However, unlike states of perfect communication, they are unstable relative to neutral drift. (shrink)

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)

Brian Skyrms offers a fascinating demonstration of how fundamental signals are to our world. He uses various scientific tools to investigate how meaning and communication develop. Signals operate in networks of senders and receivers at all levels of life, transmitting and processing information. That is how humans and animals think and interact.

Preface by Daniel C. Dennett Beginning with a general theory of function applied to body organs, behaviors, customs, and both inner and outer representations, ...

The signaling theory of religion has many claimed virtues, but these are not necessarily all realizable at the same time. Modeling choices involve trade-offs, and the available options here have not traditionally been well understood. This paper offers an overview of signaling theory relevant to the signaling theory of religion, arguing for a narrow, “core” reading of it. I outline a broad taxonomy of the choices on offer for signaling models, and examples of how previous and potential approaches to modeling (...) religious signaling meet or fail to meet the initial promise of the theory. A pluralist approach to religious signaling seems possible, but this would require a high level of detail and specificity with respect to both formal models and target systems. (shrink)

The handicap principle has come under significant challenge both from empirical studies and from theoretical work. As a result, a number of alternative explanations for honest signaling have been proposed. This paper compares the evolutionary plausibility of one such alternative, the "hybrid equilibrium," to the handicap principle. We utilize computer simulations to compare these two theories as they are instantiated in Maynard Smith's Sir Philip Sidney game. We conclude that, when both types of communication are possible, evolution is unlikely to (...) lead to handicap signaling and is far more likely to result in the partially honest signaling predicted by hybrid equilibrium theory. (shrink)

Costly signalling theory has become a common explanation for honest communication when interests conflict. In this paper, we provide an alternative explanation for partially honest communication that does not require significant signal costs. We show that this alternative is at least as plausible as traditional costly signalling, and we suggest a number of experiments that might be used to distinguish the two theories.

The concept of fitness began its career in biology long before evolutionary theory was mathematized. Fitness was used to describe an organism’s vigor, or the degree to which organisms “fit” into their environments. An organism’s success in avoiding predators and in building a nest obviously contribute to its fitness and to the fitness of its offspring, but the peacock’s gaudy tail seemed to be in an entirely different line of work. Fitness, as a term in ordinary language (as in “physical (...) fitness”) and in its original biological meaning, applied to the survival of an organism and its offspring, not to sheer reproductive output (Paul ////; Cronin 1991). Darwin’s separation of natural from sexual selection may sound odd from a modern perspective, but it made sense from this earlier point of view. (shrink)