The comprehension of living organisms in all their complexity poses a major challenge to the biological sciences. Recently, systems biology has been proposed as a new candidate in the development of such a comprehension. The main objective of this paper is to address what systems biology is and how it is practised. To this end, the basic tools of a systems biological approach are explored and illustrated. In addition, it is questioned whether systems biology ‘revolutionizes’ molecular biology and ‘transcends’ its (...) assumed reductionism. The strength of this claim appears to depend on how molecular and systems biology are characterised and on how reductionism is interpreted. Doing credit to molecular biology and to methodological reductionism, it is argued that the distinction between molecular and systems biology is gradual rather than sharp. As such, the classical challenge in biology to manage, interpret and integrate biological data into functional wholes is further intensified by systems biology’s use of modelling and bioinformatics, and by its scale enlargement. (shrink)

Ever since David Hume, empiricists have barred powers and capacities from nature. In this book Cartwright argues that capacities are essential in our scientific world, and, contrary to empiricist orthodoxy, that they can meet sufficiently strict demands for testability. Econometrics is one discipline where probabilities are used to measure causal capacities, and the technology of modern physics provides several examples of testing capacities (such as lasers). Cartwright concludes by applying the lessons of the book about capacities and probabilities to the (...) explanation of the role of causality in quantum mechanics. (shrink)

We will show that there is a strong form of emergence in cell biology. Beginning with C.D. Broad’s classic discussion of emergence, we distinguish two conditions sufficient for emergence. Emergence in biology must be compatible with the thought that all explanations of systemic properties are mechanistic explanations and with their sufficiency. Explanations of systemic properties are always in terms of the properties of the parts within the system. Nonetheless, systemic properties can still be emergent. If the properties of the components (...) within the system cannot be predicted, even in principle, from the behavior of the system’s parts within simpler wholes then there also will be systemic properties which cannot be predicted, even in principle, on basis of the behavior of these parts. We show in an explicit case study drawn from molecular cell physiology that biochemical networks display this kind of emergence, even though they deploy only mechanistic explanations. This illustrates emergence and its place in nature. (shrink)

We will show that there is a strong form of emergence in cell biology. Beginning with C.D. Broad's classic discussion of emergence, we distinguish two conditions sufficient for emergence. Emergence in biology must be compatible with the thought that all explanations of systemic properties are mechanistic explanations and with their sufficiency. Explanations of systemic properties are always in terms of the properties of the parts within the system. Nonetheless, systemic properties can still be emergent. If the properties of the components (...) within the system cannot be predicted, even in principle, from the behavior of the system's parts within simpler wholes then there also will be systemic properties which cannot be predicted, even in principle, on basis of the behavior of these parts. We show in an explicit case study drawn from molecular cell physiology that biochemical networks display this kind of emergence, even though they deploy only mechanistic explanations. This illustrates emergence and its place in nature. (shrink)

Explaining the complex dynamics exhibited in many biological mechanisms requires extending the recent philosophical treatment of mechanisms that emphasizes sequences of operations. To understand how nonsequentially organized mechanisms will behave, scientists often advance what we call dynamic mechanistic explanations. These begin with a decomposition of the mechanism into component parts and operations, using a variety of laboratory-based strategies. Crucially, the mechanism is then recomposed by means of computational models in which variables or terms in differential equations correspond to properties of (...) its parts and operations. We provide two illustrations drawn from research on circadian rhythms. Once biologists identified some of the components of the molecular mechanism thought to be responsible for circadian rhythms, computational models were used to determine whether the proposed mechanisms could generate sustained oscillations. Modeling has become even more important as researchers have recognized that the oscillations generated in individual neurons are synchronized within networks; we describe models being employed to assess how different possible network architectures could produce the observed synchronized activity. (shrink)

This article argues that the basic account of mechanism and mechanistic explanation, involving sequential execution of qualitatively characterized operations, is itself insufficient to explain biological phenomena such as the capacity of living organisms to maintain themselves as systems distinct from their environment. This capacity depends on cyclic organization, including positive and negative feedback loops, which can generate complex dynamics. Understanding cyclically organized mechanisms with complex dynamics requires coordinating research directed at decomposing mechanisms into parts and operations with research using computational (...) models to recompose mechanisms and determine their dynamic behavior. This coordinated endeavor yields dynamic mechanistic explanations. (shrink)

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)

Motile bacteria respond to environmental cues to move to more favorable locations. The components of the chemotaxis signal transduction systems that mediate these responses are highly conserved among prokaryotes including both eubacterial and archael species. The best‐studied system is that found in Escherichia coli. Attractant and repellant chemicals are sensed through their interactions with transmembrane chemoreceptor proteins that are localized in multimeric assemblies at one or both cell poles together with a histidine protein kinase, CheA, an SH3‐like adaptor protein, CheW, (...) and a phosphoprotein phosphatase, CheZ. These multimeric protein assemblies act to control the level of phosphorylation of a response regulator, CheY, which dictates flagellar motion. Bacterial chemotaxis is one of the most‐understood signal transduction systems, and many biochemical and structural details of this system have been elucidated. This is an exciting field of study because the depth of knowledge now allows the detailed molecular mechanisms of transmembrane signaling and signal processing to be investigated. BioEssays 28:9–22, 2006. © 2005 Wiley Periodicals, Inc. (shrink)

The comprehension of living organisms in all their complexity poses a major challenge to the biological sciences. Recently, systems biology has been proposed as a new candidate in the development of such a comprehension. The main objective of this paper is to address what systems biology is and how it is practised. To this end, the basic tools of a systems biological approach are explored and illustrated. In addition, it is questioned whether systems biology ‘revolutionizes’ molecular biology and ‘transcends’ its (...) assumed reductionism. The strength of this claim appears to depend on how molecular and systems biology are characterised and on how reductionism is interpreted. Doing credit to molecular biology and to methodological reductionism, it is argued that the distinction between molecular and systems biology is gradual rather than sharp. As such, the classical challenge in biology to manage, interpret and integrate biological data into functional wholes is further intensified by systems biology’s use of modelling and bioinformatics, and by its scale enlargement. (shrink)

Even though the evidence‐based medicine movement (EBM) labels mechanisms a low quality form of evidence, consideration of the mechanisms on which medicine relies, and the distinct roles that mechanisms might play in clinical practice, offers a number of insights into EBM itself. In this paper, I examine the connections between EBM and mechanisms from several angles. I diagnose what went wrong in two examples where mechanistic reasoning failed to generate accurate predictions for how a dysfunctional mechanism would respond to intervention. (...) I then use these examples to explain why we should expect this kind of mechanistic reasoning to fail in systematic ways, by situating these failures in terms of evolved complexity of the causal system(s) in question. I argue that there is still a different role in which mechanisms continue to figure as evidence in EBM: namely, in guiding the application of population‐level recommendations to individual patients. Thus, even though the evidence‐based movement rejects one role in which mechanistic reasoning serves as evidence, there are other evidentiary roles for mechanistic reasoning. This renders plausible the claims of some critics of evidencebased medicine who point to the ineliminable role of clinical experience. Clearly specifying the ways in which mechanisms and mechanistic reasoning can be involved in clinical practice frames the discussion about EBM and clinical experience in more fruitful terms. (shrink)

Models are of central importance in many scientific contexts. The centrality of models such as inflationary models in cosmology, general-circulation models of the global climate, the double-helix model of DNA, evolutionary models in biology, agent-based models in the social sciences, and general-equilibrium models of markets in their respective domains is a case in point (the Other Internet Resources section at the end of this entry contains links to online resources that discuss these models). Scientists spend significant amounts of time building, (...) testing, comparing, and revising models, and much journal space is dedicated to interpreting and discussing the implications of models. In short, models are one of the principal instruments of modern science. (shrink)

This paper defends a counterfactual account of explanation, according to which successful explanation requires tracing patterns of counterfactual dependence of a special sort, involving what I call active counterfactuals. Explanations having this feature must appeal to generalizations that are invariant--stable under certain sorts of changes. These ideas are illustrated by examples drawn from physics and econometrics.

Philosophers of science increasingly recognize the importance of idealization: the intentional introduction of distortion into scientific theories. Yet this recognition has not yielded consensus about the nature of idealization. e literature of the past thirty years contains disparate characterizations and justifications, but little evidence of convergence towards a common position.

Mechanistic explanation has an impressive track record of advancing our understanding of complex, hierarchically organized physical systems, particularly biological and neural systems. But not every complex system can be understood mechanistically. Psychological capacities are often understood by providing cognitive models of the systems that underlie them. I argue that these models, while superficially similar to mechanistic models, in fact have a substantially more complex relation to the real underlying system. They are typically constructed using a range of techniques for abstracting (...) the functional properties of the system, which may not coincide with its mechanistic organization. I describe these techniques and show that despite being non-mechanistic, these cognitive models can satisfy the normative constraints on good explanations. (shrink)

This paper explores whether natural selection, a putative evolutionary mechanism, and a main one at that, can be characterized on either of the two dominant conceptions of mechanism, due to Glennan and the team of Machamer, Darden, and Craver, that constitute the “new mechanistic philosophy.” The results of the analysis are that neither of the dominant conceptions of mechanism adequately captures natural selection. Nevertheless, the new mechanistic philosophy possesses the resources for an understanding of natural selection under the rubric.

Several articles have recently appeared arguing that there really are no viable alternatives to mechanistic explanation in the biological sciences (Kaplan and Bechtel; Kaplan and Craver). We argue that mechanistic explanation is defined by localization and decomposition. We argue further that systems neuroscience contains explanations that violate both localization and decomposition. We conclude that the mechanistic model of explanation needs to either stretch to now include explanations wherein localization or decomposition fail or acknowledge that there are counterexamples to mechanistic explanation (...) in the biological sciences. (shrink)

Chemists take mechanisms to be an important way of explaining chemical change. I examine the usefulness of the mechanism approach in the recent philosophical literature in explicating the explanatory use of mechanisms by organic chemists. I argue that chemists consider a mechanism to be explanatory because it accounts for the “dynamic process of bringing about” (Tabery 2004 , 10) chemical change. For chemists, mechanisms are causal explanations based on interventions that show “how some possibilities depend on others” (Woodward 2003 , (...) 375). Only possibilities are achievable because chemists face a number of challenges when they explain by means of a mechanism. †To contact the author, please write to: Department of Philosophy, Smith College, Northampton, MA 01063; e‐mail: [email protected]. (shrink)

This book on the philosophy of science argues for an empiricism, opposed to the tradition of David Hume, in which singular rather than general causal claims are primary; causal laws express facts about singular causes whereas the general causal claims of science are ascriptions of capacities or causal powers, capacities to make things happen. Taking science as measurement, Cartwright argues that capacities are necessary for science and that these can be measured, provided suitable conditions are met. There are case studies (...) from both econometrics and quantum mechanics. (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.

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)

In this paper I offer an analysis of causation based upon a theory of mechanisms-complex systems whose internal parts interact to produce a system's external behavior. I argue that all but the fundamental laws of physics can be explained by reference to mechanisms. Mechanisms provide an epistemologically unproblematic way to explain the necessity which is often taken to distinguish laws from other generalizations. This account of necessity leads to a theory of causation according to which events are causally related when (...) there is a mechanism that connects them. I present reasons why the lack of an account of fundamental physical causation does not undermine the mechanical account. (shrink)

This textbook is aimed at newcomers to nonlinear dynamics and chaos, especially students taking a first course in the subject. The presentation stresses analytical methods, concrete examples and geometric intuition. The theory is developed systematically, starting with first-order differential equations and their bifurcations, followed by phase plane analysis, limit cycles and their bifurcations, and culminating with the Lorenz equations, chaos, iterated maps, period doubling, renormalization, fractals, and strange attractors.A unique feature of the book is its emphasis on applications. These include (...) mechanical vibrations, lasers, biological rhythms, superconducting circuits, insect outbreaks, chemical oscillators, genetic control systems, chaotic waterwheels, and even a technique for using chaos to send secret messages. In each case, the scientific background is explained at an elementary level and closely integrated with the mathematical theory.Richly illustrated, and with many exercises and worked examples, this book is ideal for an introductory course at the junior/senior or first-year graduate level. It is also ideal for the scientist who has not had formal instruction in nonlinear dynamics, but who now desires to begin informal study. The prerequisites are multivariable calculus and introductory physics. (shrink)

A variety of scientific disciplines have set as their task explaining mental activities, recognizing that in some way these activities depend upon our brain. But, until recently, the opportunities to conduct experiments directly on our brains were limited. As a result, research efforts were split between disciplines such as cognitive psychology, linguistics, and artificial intelligence that investigated behavior, while disciplines such as neuroanatomy, neurophysiology, and genetics experimented on the brains of non-human animals. In recent decades these disciplines integrated, and with (...) the advent of techniques for imaging activity in human brains, the term cognitive neuroscience has been applied to the integrated investigations of mind and brain. This book is a philosophical examination of how these disciplines continue in the mission of explaining our mental capacities. (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)

Carl F. Craver investigates what we are doing when we use neuroscience to explain what's going on in the brain. When does an explanation succeed and when does it fail? Craver offers explicit standards for successful explanation of the workings of the brain, on the basis of a systematic view about what neuroscientific explanations are.

Carl Craver investigates what we are doing when we sue neuroscience to explain what's going on in the brain.

Evolutionary developmental biology (evo-devo) is considered a ‘mechanistic science,’ in that it causally explains morphological evolution in terms of changes in developmental mechanisms. Evo-devo is also an interdisciplinary and integrative approach, as its explanations use contributions from many fields and pertain to different levels of organismal organization. Philosophical accounts of mechanistic explanation are currently highly prominent, and have been particularly able to capture the integrative nature of multifield and multilevel explanations. However, I argue that evo-devo demonstrates the need for a (...) broadened philosophical conception of mechanisms and mechanistic explanation. Mechanistic explanation (in terms of the qualitative interactions of the structural parts of a whole) has been developed as an alternative to the traditional idea of explanation as derivation from laws or quantitative principles. Against the picture promoted by Carl Craver, that mathematical models describe but usually do not explain, my discussion of cases from the strand of evo-devo which is concerned with developmental processes points to qualitative phenomena where quantitative mathematical models are an indispensable part of the explanation. While philosophical accounts have focused on the actual organization and operation of mechanisms, properties of developmental mechanisms that are about how a mechanism reacts to modifications are of major evolutionary significance, including robustness, phenotypic plasticity, and modularity. A philosophical conception of mechanisms is needed that takes into account quantitative changes, transient entities and the generation of novel types of entities, feedback loops and complex interaction networks, emergent properties, and, in particular, functional-dynamical aspects of mechanisms, including functional (as opposed to structural) organization and distributed, system-wide phenomena. I conclude with general remarks on philosophical accounts of explanation. (shrink)

In recent debates mechanisms are often discussed in the context of ‘complex systems’ which are understood as having a complicated compositional structure. I want to draw the attention to another, radically different kind of complex system, in fact one that many scientists regard as the only genuine kind of complex system. Instead of being compositionally complex these systems rather exhibit highly non-trivial dynamical patterns on the basis of structurally simple arrangements of large numbers of non-linearly interacting constituents. The characteristic dynamical (...) patterns in what I call “dynamically complex systems” arise from the interaction of the system’s parts largely irrespective of many properties of these parts. Dynamically complex systems can exhibit surprising statistical characteristics, the robustness of which calls for an explanation in terms of underlying generating mechanisms. However, I want to argue, dynamically complex systems are not sufficiently covered by the available conceptions of mechanisms. I will explore how the notion of a mechanism has to be modified to accommodate this case. Moreover, I will show under which conditions the widespread, if not inflationary talk about mechanisms in complex systems stretches the notion of mechanisms beyond its reasonable limits and is no longer legitimate. (shrink)

1. The Naturalistic Turn in Philosophy of Science 2. The Framework of Mechanistic Explanation: Parts, Operations, and Organization 3. Representing and Reasoning About Mechanisms 4. Mental Mechanisms: Mechanisms that Process Information 5. Discovering Mental Mechanisms 6 . Summary.