I argue that there are at least four different ways in which the term ‘function’ is used in connection with the study of living organisms, namely: function as activity, function as biological role, function as biological advantage, and function as selected effect. Notion refers to what an item does by itself; refers to the contribution of an item or activity to a complex activity or capacity of an organism; refers to the value for the organism of an item having a (...) certain character rather than another; refers to the way in which a trait acquired and has maintained its current share in the population. The recognition of a separate notion of function as biological advantage solves the problem of the indeterminate reference situation that has been raised against a counterfactual analysis of function, and emphasizes the importance of counterfactual comparison in the explanatory practice of organismal biology. This reveals a neglected problem in the philosophy of biology, namely that of accounting for the insights provided by counterfactual comparison. (shrink)

Kenneth F. Schaffner compares the practice of biological and medical research and shows how traditional topics in philosophy of science—such as the nature of theories and of explanation—can illuminate the life sciences. While Schaffner pays some attention to the conceptual questions of evolutionary biology, his chief focus is on the examples that immunology, human genetics, neuroscience, and internal medicine provide for examinations of the way scientists develop, examine, test, and apply theories. Although traditional philosophy of science has regarded scientific discovery—the (...) questions of creativity in science—as a subject for psychological rather than philosophical study, Schaffner argues that recent work in cognitive science and artificial intelligence enables researchers to rationally analyze the nature of discovery. As a philosopher of science who holds an M.D., he has examined biomedical work from the inside and uses detailed examples from the entire range of the life sciences to support the semantic approach to scientific theories, addressing whether there are "laws" in the life sciences as there are in the physical sciences. Schaffner's novel use of philosophical tools to deal with scientific research in all of its complexity provides a distinctive angle on basic questions of scientific evaluation and explanation. (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)

How do minds emerge from developing brains? According to the representational features of cortex are built from the dynamic interaction between neural growth mechanisms and environmentally derived neural activity. Contrary to popular selectionist models that emphasize regressive mechanisms, the neurobiological evidence suggests that this growth is a progressive increase in the representational properties of cortex. The interaction between the environment and neural growth results in a flexible type of learning: minimizes the need for prespecification in accordance with recent neurobiological evidence (...) that the developing cerebral cortex is largely free of domain-specific structure. Instead, the representational properties of cortex are built by the nature of the problem domain confronting it. This uniquely powerful and general learning strategy undermines the central assumption of classical learnability theory, that the learning properties of a system can be deduced from a fixed computational architecture. Neural constructivism suggests that the evolutionary emergence of neocortex in mammals is a progression toward more flexible representational structures, in contrast to the popular view of cortical evolution as an increase in innate, specialized circuits. Human cortical postnatal development is also more extensive and protracted than generally supposed, suggesting that cortex has evolved so as to maximize the capacity of environmental structure to shape its structure and function through constructive learning. (shrink)

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)

"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 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)

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)

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.

Many areas of science develop by discovering mechanisms and role functions. Cummins' (1975) analysis of role functions-according to which an item's role function is a capacity of that item that appears in an analytic explanation of the capacity of some containing system-captures one important sense of "function" in the biological sciences and elsewhere. Here I synthesize Cummins' account with recent work on mechanisms and causal/mechanical explanation. The synthesis produces an analysis of specifically mechanistic role functions, one that uses the characteristic (...) active, spatial, temporal, and hierarchical organization of mechanisms to add precision and content to Cummins' original suggestion. This synthesis also shows why the discovery of role functions is a scientific achievement. Discovering a role function (i) contributes to the interlevel integration of multilevel mechanisms, and (ii) provides a unique, contextual variety of causal/mechanical explanation. (shrink)

Recently something close to a consensus about the best way to naturalize the notion of biological function appears to be emerging. Nonetheless, teleological notions in biology remain controversial. In this paper we provide a naturalistic analysis for the notion of natural design. Many authors assume that natural design should be assimilated directly to function. Others find the notion problematic because it suggests that evolution is a directed process. We argue that both of these views are mistaken. Our naturalistic account does (...) not simply equate design with function. We argue that the distinction between function and design is important for understanding the evolution of the physical and behavioral traits of organisms. (shrink)

The concept of mechanism is analyzed in terms of entities and activities, organized such that they are productive of regular changes. Examples show how mechanisms work in neurobiology and molecular biology. Thinking in terms of mechanisms provides a new framework for addressing many traditional philosophical issues: causality, laws, explanation, reduction, and scientific change.

I shall begin with the "anti-psychiatry" view that the lack of a physical basis excludes many familiar mental disorders from the category of "illness". My response to this argument will be that anti-psychiatrists are probably right to hold that most mental disorders do not involve any physical disorder, but that they are wrong to conclude from this that these mental disorders are not illnesses.

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.

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

This is a short overview of the biological functions debate in philosophy. While it was fairly comprehensive when it was written, my short book ​A Critical Overview of Biological Functions has largely supplanted it as a definitive and up-to-date overview of the debate, both because the book takes into account new developments since then, and because the length of the book allowed me to go into substantially more detail about existing views.

Scientists, philosophers, and even the lay public commonly accept that schizophrenia stems from a biological or internal ‘dysfunction.’ However, this assessment is typically accompanied neither by well-defined criteria for determining that something is dysfunctional nor empirical evidence that schizophrenia satisfies those criteria. In the following, a concept of biological function is developed and applied to a neurobiological model of schizophrenia. It concludes that current evidence does not warrant the claim that schizophrenia stems from a biological dysfunction, and, in fact, that (...) unusual neural structures associated with schizophrenia may have functional or adaptive significance. The fact that current evidence is ambivalent between these two possibilities (dysfunction versus adaptive function) implies that schizophrenia researchers should be much more cautious in using the ‘dysfunction’ label than they currently are. This has implications for both psychiatric treatment as well as public perception of mental disorders. (shrink)

Both biologists and philosophers often make use of simple verbal formulations of necessary and sufficient conditions for evolution by natural selection (ENS). Such summaries go back to Darwin's Origin of Species (especially the "Recapitulation"), but recent ones are more compact.1 Perhaps the most commonly cited formulation is due to Lewontin.2 These summaries tend to have three or four conditions, where the core requirement is a combination of variation, heredity, and fitness differences. The summaries are employed in several ways. First, they (...) are often used in pedagogical contexts, and in showing the coherence of evolutionary theory in response to attacks from outside biology. Second, they are important in discussions of extensions of evolutionary principles to new domains, such as cultural change. The summaries also have intrinsic scientific and philosophical interest as attempts to capture some core principles of evolutionary theory in a highly concise way. Despite their prominence, both the proper formulation and status of these summaries are unclear. Standard formulations are subject to counterexamples, and their relations to formal models of evolutionary change are not straightforward. Here I look closely at these verbal summaries, and at how they relate to formal models. Are the summaries merely rough approximations that have no theoretical role of their own? Perhaps they could operate as theoretical statements in Darwin's time, but have now been superseded by more exact treatments. (shrink)

Human behavior is the joint product of (i) contingencies of survival responsible for natural selection, and (ii) contingencies of reinforcement responsible for the repertoires of individuals, including (iii) the special contingencies maintained by an evolved social environment. Selection by consequences is a causal mode found only in living things, or in machines made by living things. It was first recognized in natural selection: Reproduction, a first consequence, led to the evolution of cells, organs, and organisms reproducing themselves under increasingly diverse (...) conditions. The behavior functioned well, however, only under conditions similar to those under which it was selected.Reproduction under a wider range of consequences became possible with the evolution of processes through which organisms acquired behavior appropriate to novel environments. One of these, operant conditioning, is a second kind of selection by consequences: New responses could be strengthened by events which followed them. When the selecting consequences are the same, operant conditioning and natural selection work together redundantly. But because a species which quickly acquires behavior appropriate to an environment has less need for an innate repertoire, operant conditioning could replace as well as supplement the natural selection of behavior.Social behavior is within easy range of natural selection, because other members are one of the most stable features of the environment of a species. The human species presumably became more social when its vocal musculature came under operant control. Verbal behavior greatly increased the importance of a third kind of selection by consequences, the evolution of social environments or cultures. The effect on the group, and not the reinforcing consequences for individual members, is responsible for the evolution of culture. (shrink)

The central aim of this paper is to shed light on the nature of explanation in computational neuroscience. I argue that computational models in this domain possess explanatory force to the extent that they describe the mechanisms responsible for producing a given phenomenon—paralleling how other mechanistic models explain. Conceiving computational explanation as a species of mechanistic explanation affords an important distinction between computational models that play genuine explanatory roles and those that merely provide accurate descriptions or predictions of phenomena. It (...) also serves to clarify the pattern of model refinement and elaboration undertaken by computational neuroscientists. (shrink)

What Is Biodiversity? is a theoretical and conceptual exploration of the biological world and how diversity is valued. Maclaurin and Sterelny explore not only the origins of the concept of biodiversity, but also how that concept has been shaped by ecology and more recently by conservation biology. They explain the different types of biodiversity important in evolutionary theory, developmental biology, ecology, morphology and taxonomy and conclude that biological heritage is rich in not just one biodiversity but many. Maclaurin and Sterelny (...) also explore the case for the conservation of these biodiversities using option value theory, a tool borrowed from economics. (shrink)

We sketch a framework for building a unified science of cognition. This unification is achieved by showing how functional analyses of cognitive capacities can be integrated with the multilevel mechanistic explanations of neural systems. The core idea is that functional analyses are sketches of mechanisms , in which some structural aspects of a mechanistic explanation are omitted. Once the missing aspects are filled in, a functional analysis turns into a full-blown mechanistic explanation. By this process, functional analyses are seamlessly integrated (...) with multilevel mechanistic explanations. (shrink)

Biologists studying ecology and evolution use the term “population” in many different ways. Yet little philosophical analysis of the concept has been done, either by biologists or philosophers, in contrast to the voluminous literature on the concept of “species.” This is in spite of the fact that “population” is arguably a far more central concept in ecological and evolutionary studies than “species” is. The fact that such a central concept has been employed in so many different ways is potentially problematic (...) for the reason that inconsistent usages (especially when the usage has not been made explicit) might lead to false controversies in which disputants are simply talking past one another. However, the inconsistent usages are not the only, or even the most important reason to examine the concept. If any set of organisms is legitimately called a “population,” selection and drift processes become purely arbitrary, too. Moreover, key ecological variables, such as abundance and distribution, depend on a nonarbitrary way of identifying populations. I sketch the beginnings of a population concept, drawing inspiration from the Ghiselin-Hull individuality thesis, and show why some alternative approaches are nonstarters. (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)

Are learning processes selection processes? This paper takes a slightly modified version of the account of selection presented in Hull et al. (Behav Brain Sci 24:511–527, 2001) and asks whether it applies to learning processes. The answer is that although some learning processes are selectional, many are not. This has consequences for teleological theories of mental content. According to these theories, mental states have content in virtue of having proper functions, and they have proper functions in virtue of being the (...) products of selection processes. For some mental states, it is plausible that the relevant selection process is natural selection, but there are many for which it is not plausible. One response to this (due to David Papineau) is to suggest that the learning processes by which we acquire non-innate mental states are selection processes and can therefore confer proper functions on mental states. This paper considers two ways in which this response could be elaborated, and argues that neither of them succeed: the teleosemanticist cannot rely on the claim that learning processes are selection processes in order to justify the attribution of proper functions to beliefs. (shrink)

Sober and Lewontin's critique of genic selectionism is based upon the principle that a unit of selection should make a context‐independent contribution to fitness. Critics have effectively shown that this principle is flawed. In this paper I show that the context independence principle is an instance of a more general principle for characterizing causes,called the contextual unanimity principle. I argue that this latter principle, while widely accepted, is erroneous. What is needed is to replace the approach to causality characterized by (...) the contextual unanimity criterion with an approach based on the concept of causal mechanism. After sketching such an approach, I show how it can be used to shed light on the units of selection problem. (shrink)

Selection type theories solve adaptation problems. Natural selection, clonal selection for antibody production, and selective theories of higher brain function are examples. An abstract characterization of typical selection processes is generated by analyzing and extending previous work on the nature of natural selection. Once constructed, this abstraction provides a useful tool for analyzing the nature of other selection theories and may be of use in new instances of theory construction. This suggests the potential fruitfulness of research to find other theory (...) types and construct their abstractions. (shrink)

The ultimate source of explanation in biology is the principle of natural selection. Natural selection means differential reproduction of genes and gene combinations. It is a mechanistic process which accounts for the existence in living organisms of end-directed structures and processes. It is argued that teleological explanations in biology are not only acceptable but indeed indispensable. There are at least three categories of biological phenomena where teleological explanations are appropriate.

Was human nature designed by natural selection in the Pleistocene epoch? The dominant view in evolutionary psychology holds that it was -- that our psychological adaptations were designed tens of thousands of years ago to solve problems faced by our hunter-gatherer ancestors. In this provocative and lively book, David Buller examines in detail the major claims of evolutionary psychology -- the paradigm popularized by Steven Pinker in The Blank Slate and by David Buss in The Evolution of Desire -- and (...) rejects them all. This does not mean that we cannot apply evolutionary theory to human psychology, says Buller, but that the conventional wisdom in evolutionary psychology is misguided.Evolutionary psychology employs a kind of reverse engineering to explain the evolved design of the mind, figuring out the adaptive problems our ancestors faced and then inferring the psychological adaptations that evolved to solve them. In the carefully argued central chapters of Adapting Minds, Buller scrutinizes several of evolutionary psychology's most highly publicized "discoveries," including "discriminative parental solicitude". Drawing on a wide range of empirical research, including his own large-scale study of child abuse, he shows that none is actually supported by the evidence.Buller argues that our minds are not adapted to the Pleistocene, but, like the immune system, are continually adapting, over both evolutionary time and individual lifetimes. We must move beyond the reigning orthodoxy of evolutionary psychology to reach an accurate understanding of how human psychology is influenced by evolution. When we do, Buller claims, we will abandon not only the quest for human nature but the very idea of human nature itself. (shrink)

I argue that there are at least four different ways in which the term ‘function’ is used in connection with the study of living organisms, namely: function as activity, function as biological role, function as biological advantage, and function as selected effect. Notion refers to what an item does by itself; refers to the contribution of an item or activity to a complex activity or capacity of an organism; refers to the value for the organism of an item having a (...) certain character rather than another; refers to the way in which a trait acquired and has maintained its current share in the population. The recognition of a separate notion of function as biological advantage solves the problem of the indeterminate reference situation that has been raised against a counterfactual analysis of function, and emphasizes the importance of counterfactual comparison in the explanatory practice of organismal biology. This reveals a neglected problem in the philosophy of biology, namely that of accounting for the insights provided by counterfactual comparison. (shrink)

An analysis of two heuristic strategies for the development of mechanistic models, illustrated with historical examples from the life sciences. In Discovering Complexity, William Bechtel and Robert Richardson examine two heuristics that guided the development of mechanistic models in the life sciences: decomposition and localization. Drawing on historical cases from disciplines including cell biology, cognitive neuroscience, and genetics, they identify a number of "choice points" that life scientists confront in developing mechanistic explanations and show how different choices result in divergent (...) explanatory models. Describing decomposition as the attempt to differentiate functional and structural components of a system and localization as the assignment of responsibility for specific functions to specific structures, Bechtel and Richardson examine the usefulness of these heuristics as well as their fallibility—the sometimes false assumption underlying them that nature is significantly decomposable and hierarchically organized. When Discovering Complexity was originally published in 1993, few philosophers of science perceived the centrality of seeking mechanisms to explain phenomena in biology, relying instead on the model of nomological explanation advanced by the logical positivists (a model Bechtel and Richardson found to be utterly inapplicable to the examples from the life sciences in their study). Since then, mechanism and mechanistic explanation have become widely discussed. In a substantive new introduction to this MIT Press edition of their book, Bechtel and Richardson examine both philosophical and scientific developments in research on mechanistic models since 1993. (shrink)

A formalism is introduced to represent the connective organization of an evolving neuronal network and the effects of environment on this organization by stabilization or degeneration of labile synapses associated with functioning. Learning, or the acquisition of an associative property, is related to a characteristic variability of the connective organization: the interaction of the environment with the genetic program is printed as a particular pattern of such organization through neuronal functioning. An application of the theory to the development of the (...) neuromuscular junction is proposed and the basic selective aspect of learning emphasized. (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)

This paper examines the relationship between a family of concepts involving reliable correlation, and a family of concepts involving adaptation and biological function, as these concepts are used in the naturalistic semantic theory of Dretske's "Explaining Behavior." I argue that Dretske's attempt to marry correlation and function to produce representation fails, though aspects of his failure point the way forward to a better theory.

The occasion for this critical review is the publication of a second, revised edition of David M Balme's translation, with introduction and notes, of Aristotle's "De Partimus Animalium I" and "De Generatione Animalium I" in the Clarendon Aristotle series. The second edition includes an appendix by Allan Gotthelf with a selective bibliography and a guide to the philosophical literature occasioned by Balme's notes. This critical review discusses Balme's unique style of translation and commentary, analyzing in detail a number of key (...) passages in both. (shrink)

There are two general approaches to characterising biological functions. One originates with Cummins. According to this approach, the function of a part of a system is just its causal contribution to some specified activity of the system. Call this the ‘C-function’ concept. The other approach ties the function of a trait to some aspect of its evolutionary significance. Call this the ‘E-function’ concept. According to the latter view, a trait's function is determined by the forces of natural selection. The C-function (...) and E-function concepts are clearly quite different, but there is an important relation between them which heretofore has gone unnoticed. The purpose of this paper is to outline that relation.This is not the first paper to discuss the relation of C-function and E-function. Previous attempts all follow either one of two strategies. The first proposes that the two concepts are ‘unified.’ The other proposes that they are radically distinct and apply to wholly different fields within biology. (shrink)

Authors frequently refer to gene-based selection in biological evolution, the reaction of the immune system to antigens, and operant learning as exemplifying selection processes in the same sense of this term. However, as obvious as this claim may seem on the surface, setting out an account of “selection” that is general enough to incorporate all three of these processes without becoming so general as to be vacuous is far from easy. In this target article, we set out such a general (...) account of selection to see how well it accommodates these very different sorts of selection. The three fundamental elements of this account are replication, variation, and environmental interaction. For selection to occur, these three processes must be related in a very specific way. In particular, replication must alternate with environmental interaction so that any changes that occur in replication are passed on differentially because of environmental interaction. One of the main differences among the three sorts of selection that we investigate concerns the role of organisms. In traditional biological evolution, organisms play a central role with respect to environmental interaction. Although environmental interaction can occur at other levels of the organizational hierarchy, organisms are the primary focus of environmental interaction. In the functioning of the immune system, organisms function as containers. The interactions that result in selection of antibodies during a lifetime are between entities (antibodies and antigens) contained within the organism. Resulting changes in the immune system of one organism are not passed on to later organisms. Nor are changes in operant behavior resulting from behavioral selection passed on to later organisms. But operant behavior is not contained in the organism because most of the interactions that lead to differential replication include parts of the world outside the organism. Changes in the organism's nervous system are the effects of those interactions. The role of genes also varies in these three systems. Biological evolution is gene-based (i.e., genes are the primary replicators). Genes play very different roles in operant behavior and the immune system. However, in all three systems, iteration is central. All three selection processes are also incredibly wasteful and inefficient. They can generate complexity and novelty primarily because they are so wasteful and inefficient. Key Words: evolution; immunology; interaction; operant behavior; operant learning; replication; selection; variation. (shrink)

Much of our work with enriched experience and training in animals supports the Quartz & Sejnowski (Q&S) thesis that environmental information can interact with pre-existing neural structures to produce new synapses and neural structure. However, substantial data as well as an evolutionary perspective indicate that multiple information-capture systems exist: some are constructivist, some are selectionist, and some may be tightly constrained.