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.

A biochemical pathway is a sequence of chemical reactions in a biological organism. Such pathways specify mechanisms that explain how cells carry out their major functions by means of molecules and reactions that produce regular changes. Many diseases can be explained by defects in pathways, and new treatments often involve finding drugs that correct those defects. This paper presents explanation schemas and treatment strategies that characterize how thinking about pathways contributes to biomedical discovery. It discusses the significance of pathways for (...) understanding the nature of diseases, explanations, and theories. (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)

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)

The dominant neuroscientific theory of spatial memory is, like many theories in neuroscience, a multilevel description of a mechanism. The theory links the activities of molecules, cells, brain regions, and whole organisms into an integrated sketch of an explanation for the ability of organisms to navigate novel environments. Here I develop a taxonomy of interlevel experimental strategies for integrating the levels in such multilevel mechanisms. These experimental strategies include activation strategies, interference strategies, and additive strategies. These strategies are mutually reinforcing, (...) providing a kind of interlevel and intratheoretic robustness that has not previously been recognized. (shrink)

The dominant neuroscientific theory of spatial memory is, like many theories in neuroscience, a multilevel description of a mechanism. The theory links the activities of molecules, cells, brain regions, and whole organisms into an integrated sketch of an explanation for the ability of organisms to navigate novel environments. Here I develop a taxonomy of interlevel experimental strategies for integrating the levels in such multilevel mechanisms. These experimental strategies include activation strategies, interference strategies, and additive strategies. These strategies are mutually reinforcing, (...) providing a kind of interlevel and intratheoretic robustness that has not previously been recognized. (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.

Explanations in the life sciences frequently involve presenting a model of the mechanism taken to be responsible for a given phenomenon. Such explanations depart in numerous ways from nomological explanations commonly presented in philosophy of science. This paper focuses on three sorts of differences. First, scientists who develop mechanistic explanations are not limited to linguistic representations and logical inference; they frequently employ diagrams to characterize mechanisms and simulations to reason about them. Thus, the epistemic resources for presenting mechanistic explanations are (...) considerably richer than those suggested by a nomological framework. Second, the fact that mechanisms involve organized systems of component parts and operations provides direction to both the discovery and testing of mechanistic explanations. Finally, models of mechanisms are developed for specific exemplars and are not represented in terms of universally quantified statements. Generalization involves investigating both the similarity of new exemplars to those already studied and the variations between them. (shrink)

Philosophers of science increasingly believe that much of science is concerned with understanding the mechanisms responsible for the production of natural phenomena. An adequate understanding of scientific research requires an account of how scientists develop and test models of mechanisms. This paper offers a general account of the nature of mechanical models, discussing the representational relationship that holds between mechanisms and their models as well as the techniques that can be used to test and refine such models. The analysis is (...) supported by study of two competing models of a mechanism of speech perception. (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)

The philosophical conception of mechanistic explanation is grounded on a limited number of canonical examples. These examples provide an overly narrow view of contemporary scientific practice, because they do not reflect the extent to which the heuristic strategies and descriptive practices that contribute to mechanistic explanation have evolved beyond the well-known methods of decomposition, localization, and pictorial representation. Recent examples from evolutionary robotics and network approaches to biology and neuroscience demonstrate the increasingly important role played by computer simulations and mathematical (...) representations in the epistemic practices of mechanism discovery and mechanism description. These examples also indicate that the scope of mechanistic explanation must be re-examined: With new and increasingly powerful methods of discovery and description comes the possibility of describing mechanisms far more complex than traditionally assumed. (shrink)

Recently philosophers of science have begun to pay more attention to the use of highly idealized mathematical models in scientific theorizing. An important example of this kind of highly idealized modeling is the widespread use of optimality models within evolutionary biology. One way to understand the explanations provided by these models is as a censored causal explanation: an explanation that omits certain causal factors in order to focus on a modular subset of the causal processes that led to the explanandum. (...) In this paper, I first argue that the censored causal model approach fails to establish a permanent explanatory role for optimality models in biology and mischaracterizes the explanatory virtues of biological optimality modeling. In addition, I argue that many biological optimality explanations cannot be characterized as censored causal explanations. In response, I propose an alternative approach that analyzes optimality models’ reliance on synchronically representing a system’s constraints and tradeoffs as well as their employment of various kinds of idealization in order to provide equilibrium explanations. (shrink)

In this paper, we compare the mechanisms of protein synthesis and natural selection. We identify three core elements of mechanistic explanation: functional individuation, hierarchical nestedness or decomposition, and organization. These are now well understood elements of mechanistic explanation in fields such as protein synthesis, and widely accepted in the mechanisms literature. But Skipper and Millstein have argued that natural selection is neither decomposable nor organized. This would mean that much of the current mechanisms literature does not apply to the mechanism (...) of natural selection.We take each element of mechanistic explanation in turn. Having appreciated the importance of functional individuation, we show how decomposition and organization should be better understood in these terms. We thereby show that mechanistic explanation by protein synthesis and natural selection are more closely analogous than they appear—both possess all three of these core elements of a mechanism widely recognized in the mechanisms literature. (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)

The optimality approach to modeling natural selection has been criticized by many biologists and philosophers of biology. For instance, Lewontin (1979) argues that the optimality approach is a shortcut that will be replaced by models incorporating genetic information, if and when such models become available. In contrast, I think that optimality models have a permanent role in evolutionary study. I base my argument for this claim on what I think it takes to best explain an event. In certain contexts, optimality (...) and game-theoretic models best explain some central types of evolutionary phenomena. ‡Thanks to Michael Friedman, Helen Longino, Michael Weisberg, and especially Elliott Sober for comments on earlier drafts of this paper. †To contact the author, please write to: Department of Philosophy, Stanford University, Stanford, CA 94305-2155; e-mail: [email protected]. (shrink)

Due to the variation, contingency and complexity of living systems, biology is often taken to be a science without fundamental theories, laws or general principles. I revisit this question in light of the quest for design principles in systems biology and show that different views can be reconciled if we distinguish between different types of generality. The philosophical literature has primarily focused on generality of specific models or explanations, or on the heuristic role of abstraction. This paper takes a different (...) approach in emphasizing a theory-constituting role of general principles. Design principles signify general dependency-relations between structures and functions, given a set of formally defined constraints. I contend that design principles increase our understanding of living systems by relating specific models to general types. The categorization of types is based on a delineation of the scope of biological possibilities, which serves to identify and define the generic features of classes of systems. To characterize the basis for general principles through generic abstraction and reasoning about possibility spaces, I coin the term constraint-based generality. I show that constraint-based generality is distinct from other types of generality in biology, and argue that general principles play a unifying role that does not entail theory reduction. (shrink)

In many domains of biology, explanation takes the form of characterizing the mechanism responsible for a particular phenomenon in a specific biological system. How are such explanations generalized? One important strategy assumes conservation of mechanisms through evolutionary descent. But conservation is seldom complete. In the case discussed, the central mechanism for circadian rhythms in animals was first identified in Drosophila and then extended to mammals. Scientists' working assumption that the clock mechanisms would be conserved both yielded important generalizations and served (...) as a heuristic for discovery, especially when significant differences between the insect and mammalian mechanism were identified. †To contact the author, please write to: Department of Philosophy and Interdisciplinary Programs in Science Studies and Cognitive Science, 0119, University of California, San Diego, La Jolla, CA 92093‐0119; e‐mail: [email protected]. (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)

This paper is concerned with reasonings that purport to explain why certain organisms have certain traits by showing that their actual design is better than contrasting designs. Biologists call such reasonings 'functional explanations'. To avoid confusion with other uses of that phrase, I call them 'design explanations'. This paper discusses the structure of design explanations and how they contribute to scientific understanding. Design explanations are contrastive and often compare real organisms to hypothetical organisms that cannot possibly exist. They are not (...) causal but appeal to functional dependencies between an organism's different traits. These explanations point out that because an organism has certain traits, it cannot be alive if the trait to be explained were replaced by a specified alternative. They can be understood from a mechanistic point of view as revealing the constraints on what mechanisms can be alive. (shrink)

A prominent approach to scientific explanation and modeling claims that for a model to provide an explanation it must accurately represent at least some of the actual causes in the event's causal history. In this paper, I argue that many optimality explanations present a serious challenge to this causal approach. I contend that many optimality models provide highly idealized equilibrium explanations that do not accurately represent the causes of their target system. Furthermore, in many contexts, it is in virtue of (...) their independence of causes that optimality models are able to provide a better explanation than competing causal models. Consequently, our account of explanation and modeling must expand beyond the causal approach. (shrink)

Discovery proceeds in stages of construction, evaluation, and revision. Each of these stages is constrained by what is known or conjectured about what is being discovered. A new characterization of mechanism aids in specifying what is to be discovered when a mechanism is sought. Guidance in discovering mechanisms may be provided by the reasoning strategies of schema instantiation, modular subassembly, and forward/backward chaining. Examples are found in mechanisms in molecular biology, biochemistry, immunology, and evolutionary biology.

Discovery proceeds in stages of construction, evaluation, and revision. Each of these stages is constrained by what is known or conjectured about what is being discovered. A new characterization of mechanism aids in specifying what is to be discovered when a mechanism is sought. Guidance in discovering mechanisms may be provided by the reasoning strategies of schema instantiation, modular subassembly, and forward/backward chaining. Examples are found in mechanisms in molecular biology, biochemistry, immunology, and evolutionary biology.

Proponents of mechanistic explanation all acknowledge the importance of organization. But they have also tended to emphasize specificity with respect to parts and operations in mechanisms. We argue that in understanding one important mode of organization—patterns of causal connectivity—a successful explanatory strategy abstracts from the specifics of the mechanism and invokes tools such as those of graph theory to explain how mechanisms with a particular mode of connectivity will behave. We discuss the connection between organization, abstraction, and mechanistic explanation and (...) illustrate our claims by looking at an example from recent research on so-called network motifs. (shrink)

Design thinking in general, and optimality modeling in particular, have traditionally been associated with adaptationism—a research agenda that gives pride of place to natural selection in shaping biological characters. Our goal is to evaluate the role of design thinking in non-evolutionary analyses. Specifically, we focus on research into abstract design principles that underpin the functional organization of extant organisms. Drawing on case studies from engineering-inspired approaches in biology we show how optimality analysis, and other design-related methods, play a specific methodological (...) role that is tangential to the study of adaptation. To account for the role of these reasoning strategies in contemporary biology, we therefore suggest a reevaluation of the connection between design thinking and adaptationism. (shrink)

In the 1950s and 1960s, an interfield interaction between molecular biologists and biochemists integrated important discoveries about the mechanism of protein synthesis. This extended discovery episode reveals two general reasoning strategies for eliminating gaps in descriptions of the productive continuity of mechanisms: schema instantiation and forward chaining/backtracking. Schema instantiation involves filling roles in an overall framework for the mechanism. Forward chaining and backtracking eliminate gaps using knowledge about types of entities and their activities. Attention to mechanisms highlights salient features of (...) this historical episode while providing general reasoning strategies for mechanism discovery. (shrink)

With In Search of Mechanisms, Carl F. Craver and Lindley Darden offer both a descriptive and an instructional account of how biologists discover mechanisms. Drawing on examples from across the life sciences and through the centuries, Craver and Darden compile an impressive toolbox of strategies that biologists have used and will use again to reveal the mechanisms that produce, underlie, or maintain the phenomena characteristic of living things. They discuss the questions that figure in the search for mechanisms, characterizing the (...) experimental, observational, and conceptual considerations used to answer them, all the while providing examples from the history of biology to highlight the kinds of evidence and reasoning strategies employed to assess mechanisms. At a deeper level, Craver and Darden pose a systematic view of what biology is, of how biology makes progress, of how biological discoveries are and might be made, and of why knowledge of biological mechanisms is important for the future of the human species. (shrink)