The history of developmental biology is interwoven with debates as to whether mechanistic explanations of development are possible or whether alternative explanatory principles or even vital forces need to be assumed. In particular, the demonstrated ability of embryonic cells to tune their developmental fate precisely to their relative position and the overall size of the embryo was once thought to be inexplicable in mechanistic terms. Taking a causal perspective, this Element examines to what extent and how developmental biology, having turned (...) molecular about four decades ago, has been able to meet the vitalist challenge. It focuses not only on the nature of explanations but also on the usefulness of causal knowledge – including the knowledge of classical experimental embryology – for further scientific discovery. It also shows how this causal perspective allows us to understand the nature and significance of some key concepts, including organizer, signal and morphogen. This title is also available as Open Access on Cambridge Core. (shrink)

I examine to what extent accounts of mechanisms based on formal interventionist theories of causality can adequately represent biological mechanisms with complex dynamics. Using a differential equation model for a circadian clock mechanism as an example, I first show that there exists an iterative solution that can be interpreted as a structural causal model. Thus, in principle, it is possible to integrate causal difference-making information with dynamical information. However, the differential equation model itself lacks the right modularity properties for a (...) full integration. A formal mechanistic model will therefore have to leave out either noncausal or causal explanatory relations. (shrink)

Philosophers use historical case studies to support wide-ranging claims about science. This practice is often criticized as problematic. In this paper we suggest that the function of case studies can be understood and justified by analogy to a well-established practice in biology: the investigation of model organisms. We argue that inferences based on case studies are no more problematic than inferences from model organisms to larger classes of organisms in biology. We demonstrate our view in detail by reference to a (...) case study with a long history: Semmelweis’s discovery of the cause of childbed fever. (shrink)

Laws in the special sciences are usually regarded to be non-universal. A theory of laws in the special sciences faces two challenges. (I) According to Lange's dilemma, laws in the special sciences are either false or trivially true. (II) They have to meet the ?requirement of relevance?, which is a way to require the non-accidentality of special science laws. I argue that both challenges can be met if one distinguishes four dimensions of (non-) universality. The upshot is that I argue (...) for the following explication of special science laws: L is a special science law just if (1) L is a system law, (2) L is quasi-Newtonian, and (3) L is minimally invariant. (shrink)

The basic idea behind the Competitive Exclusion Principle is that species that have similar or identical niches cannot stably coexist in the same place for long periods of time when their common resources are limiting. A more exact definition of the CEP states that, in equilibrium, n number of sympatric species competing for a common set of limiting resources cannot stably coexist indefinitely on fewer than n number of resources. The magnitude or intensity of competition between species is proportional to (...) the degree of overlap in their niches or use of resources. Species that stably coexist do so because there are “important differences” in their niches or use of common limiting resources. (shrink)

The import of Hodgkin and Huxley’s classic model of the action potential has been hotly debated in recent years, with particular controversy surrounding claims by prominent proponents of mechanistic explanation. For these authors, the Hodgkin-Huxley model is an excellent predictive tool but ultimately lacks causal/explanatory import. What is more, they claim that this is how Hodgkin and Huxley themselves saw the model. I argue that these claims rest on a problematic reading of the work. Hodgkin and Huxley’s model is both (...) causal and, in an important sense, explanatory. (shrink)

The Hodgkin–Huxley (HH) model of the action potential is a theoretical pillar of modern neurobiology. In a number of recent publications, Carl Craver ([2006], [2007], [2008]) has argued that the model is explanatorily deficient because it does not reveal enough about underlying molecular mechanisms. I offer an alternative picture of the HH model, according to which it deliberately abstracts from molecular specifics. By doing so, the model explains whole-cell behaviour as the product of a mass of underlying low-level events. The (...) issue goes beyond cellular neurobiology, for the strategy of abstraction exhibited in the HH case is found in a range of biological contexts. I discuss why it has been largely neglected by advocates of the mechanist approach to explanation. 1 Introduction2 A Primer on the HH Model2.1 The basic qualitative picture2.2 The quantitative model3 Interlude: What Did Hodgkin and Huxley Think?4 Craver’s View4.1 Mechanistic explanation4.2 Sketches4.3 Craver's view: The HH model as a mechanism sketch5 An Alternative View of the HH Model5.1 Another look at the equations5.2 The discrete-gating picture5.3 The road paved by Hodgkin and Huxley5.4 Summary and comparison to Craver6 Conclusion: The HH Model and Mechanistic Explanation6.1 Sketches and abstractions6.2 Why has aggregative abstraction been overlooked? (shrink)

Today, mechanisms and mechanistic explanation are very popular in philosophy of science and are deemed a welcome alternative to laws of nature and deductive‐nomological explanation. Starting from Mitchell's pragmatic notion of laws, I cast doubt on their status as a genuine alternative. I argue that (1) all complex‐systems mechanisms ontologically must rely on stable regularities, while (2) the reverse need not hold. Analogously, (3) models of mechanisms must incorporate pragmatic laws, while (4) such laws themselves need not always refer to (...) underlying mechanisms. Finally, I show that Mitchell's account is more encompassing than the mechanistic account *Received August 2008; revised January 2010. †To contact the author, please write to: Centre for Logic and Philosophy of Science, Ghent University, Blandijnberg 2, B‐9000 Belgium; e‐mail: [email protected]. (shrink)

I examine the warrants we have in light of the empirical successes of a kind of model I call ‘ hybrid models ’, a kind that includes climate models among its members. I argue that these warrants ’ strengths depend on inferential virtues that are not just explanatory virtues, contrary to what would be the case if inference to the best explanation provided the warrants. I also argue that the warrants in question, unlike those IBE provides, guide inferences only to (...) model implications about which there is real uncertainty. My conclusion provides criteria of adequacy for epistemologies of climate and other hybrid models. (shrink)

Recently, it has been provocatively claimed that dynamical modeling approaches signal the emergence of a new explanatory framework distinct from that of mechanistic explanation. This paper rejects this proposal and argues that dynamical explanations are fully compatible with, even naturally construed as, instances of mechanistic explanations. Specifically, it is argued that the mathematical framework of dynamics provides a powerful descriptive scheme for revealing temporal features of activities in mechanisms and plays an explanatory role to the extent it is deployed for (...) this purpose. It is also suggested that more attention should be paid to the distinctive methodological contributions of the dynamical framework including its usefulness as a heuristic for mechanism discovery and hypothesis generation in contemporary neuroscience and biology. (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)

Traditionally, a scientific model is thought to provide a good scientific explanation to the extent that it satisfies certain scientific goals that are thought to be constitutive of explanation. Problems arise when we realize that individual scientific models cannot simultaneously satisfy all the scientific goals typically associated with explanation. A given model’s ability to satisfy some goals must always come at the expense of satisfying others. This has resulted in philosophical disputes regarding which of these goals are in fact necessary (...) for explanation, and as such which types of models can and cannot provide explanations. Explanatory monists argue that one goal will be explanatory in all contexts, while explanatory pluralists argue that the goal will vary based on pragmatic considerations. In this paper, I argue that such debates are misguided, and that both monists and pluralists are incorrect. Instead of any goal being given explanatory priority over others in a given context, the different goals are all deeply dependent on one another for their explanatory power. Any model that sacrifices some explanatory goals to attain others will always necessarily undermine its own explanatory power in the process. And so when forced to choose between individual scientific models, there can be no explanatory victors. Given that no model can satisfy all the goals typically associated with explanation, no one model in isolation can provide a good scientific explanation. Instead we must appeal to collections of models. Collections of models provide an explanation when they satisfy the web of interconnected goals that justify the explanatory power of one another. (shrink)

A common reductionist assumption is that macro-scale behaviors can be described "bottom-up" if only sufficient details about lower-scale processes are available. The view that an "ideal" or "fundamental" physics would be sufficient to explain all macro-scale phenomena has been met with criticism from philosophers of biology. Specifically, scholars have pointed to the impossibility of deducing biological explanations from physical ones, and to the irreducible nature of distinctively biological processes such as gene regulation and evolution. This paper takes a step back (...) in asking whether bottom-up modeling is feasible even when modeling simple physical systems across scales. By comparing examples of multi-scale modeling in physics and biology, we argue that the “tyranny of scales” problem presents a challenge to reductive explanations in both physics and biology. The problem refers to the scale-dependency of physical and biological behaviors that forces researchers to combine different models relying on different scale-specific mathematical strategies and boundary conditions. Analyzing the ways in which different models are combined in multi-scale modeling also has implications for the relation between physics and biology. Contrary to the assumption that physical science approaches provide reductive explanations in biology, we exemplify how inputs from physics often reveal the importance of macro-scale models and explanations. We illustrate this through an examination of the role of biomechanics modeling in developmental biology. In such contexts, the relation between models at different scales and from different disciplines is neither reductive nor completely autonomous, but interdependent. (shrink)

The goal of this paper is to present and defend an empiricist, neo-Hempelian account of scientific explanation as ampliative, specialized embedding. The proposal aims to preserve what I take to be the core of Hempel’s empiricist account, by weakening it in some respects and strengthening it in others, introducing two new conditions that solve most of Hempel’s problems without abandoning his empiricist strictures. According to this proposal, to explain a phenomenon is to make it expectable by introducing new conceptual/ontological machinery (...) and using special, and non-ad hoc, non-accidental regularities. The new conditions are elaborated making essential use of two central structuralist ideas, namely T-theoreticity and specialization. I first introduce and qualify the project, then present the new account in detail and assess it vis-à-vis its rivals, and finally discuss some possible objections, concluding that the account fares better than its monistic rivals and well enough to qualify as a promising neo-Hempelian account. Even for those unpersuaded by its monistic goals, it has the merit of calling attention to two new necessary conditions not explicitly emphasized thus far and showing how they serve to answer many of the criticisms addressed against Hempel’s account. (shrink)

Completeness is an important but misunderstood norm of explanation. It has recently been argued that mechanistic accounts of scientific explanation are committed to the thesis that models are complete only if they describe everything about a mechanism and, as a corollary, that incomplete models are always improved by adding more details. If so, mechanistic accounts are at odds with the obvious and important role of abstraction in scientific modelling. We respond to this characterization of the mechanist’s views about abstraction and (...) articulate norms of completeness for mechanistic explanations that have no such unwanted implications. _1_ Introduction _2_ A Balancing Act: When Do Details Matter? _3_ The Norms of Causal Explanation _4_ The Norms of Constitutive Explanation _5_ Salmon-Completeness _6_ From More Details to More Relevant Details _7_ Non-explanatory Virtues of Abstraction _8_ From Explanatory Models to Explanatory Knowledge _9_ Mechanistic Completeness Reconsidered _10_ Conclusion. (shrink)

The most expressive account of explanations in neuroscience is currently the causal-mechanistic model formulated by Carl Craver. According to him, explanations in neuroscience describe mechanisms, in other words, it points out how parts organize themselves and interact to engender the phenomenon. Furthermore, neuroscience is unified as scientists from different areas that compose it work together to develop mechanisms. This model was extensively discussed in the last years and several criticisms were raised towards it. Still, it remains as the soundest model (...) for explanations in neuroscience nowadays. This paper is presented as a review of this model, as well as the critiques worked out against it and finishes with a brief consideration of the problem of explanation in neuroscience. (shrink)

Life-science phenomena are often explained by specifying the mechanisms that bring them about. The new mechanistic philosophers have done much to substantiate this claim and to provide us with a better understanding of what mechanisms are and how they explain. Although there is disagreement among current mechanists on various issues, they share a common core position and a seeming commitment to some form of scientific realism. But is such a commitment necessary? Is it the best way to go about mechanistic (...) explanation? In this article, we propose an alternative antirealist account that also fits explanatory practice in the life sciences. We pay special attention to mechanistic models, i.e. scientific models that involve a mechanism, and to the role of coherence considerations in building such models. To illustrate our points, we consider the mechanism for the action potential. 1 Introduction2 Some Core Features of Mechanistic Explanation3 Scientific Realism and Mechanistic Explanation4 Antirealist Mechanistic Explanation: The Case of the Action Potential5 Some Outstanding Issues for the Antirealist Mechanist6 Two Problems for the Realist Mechanist7 Conclusions. (shrink)

In a recent paper, Kaplan (Synthese 183:339–373, 2011) takes up the task of extending Craver’s (Explaining the brain, 2007) mechanistic account of explanation in neuroscience to the new territory of computational neuroscience. He presents the model to mechanism mapping (3M) criterion as a condition for a model’s explanatory adequacy. This mechanistic approach is intended to replace earlier accounts which posited a level of computational analysis conceived as distinct and autonomous from underlying mechanistic details. In this paper I discuss work in (...) computational neuroscience that creates difficulties for the mechanist project. Carandini and Heeger (Nat Rev Neurosci 13:51–62, 2012) propose that many neural response properties can be understood in terms of canonical neural computations. These are “standard computational modules that apply the same fundamental operations in a variety of contexts.” Importantly, these computations can have numerous biophysical realisations, and so straightforward examination of the mechanisms underlying these computations carries little explanatory weight. Through a comparison between this modelling approach and minimal models in other branches of science, I argue that computational neuroscience frequently employs a distinct explanatory style, namely, efficient coding explanation. Such explanations cannot be assimilated into the mechanistic framework but do bear interesting similarities with evolutionary and optimality explanations elsewhere in biology. (shrink)

New mechanical philosophy posits that explanations in the life sciences involve the decomposition of a system into its entities and their respective activities and organization that are responsible for the explanandum phenomenon. This mechanistic account of explanation has proven problematic in its application to mathematical models, leading the mechanists to suggest different ways of aligning abstract models with the mechanist program. Initially, the discussion centered on whether the Hodgkin-Huxley model is explanatory. Network models provided another complication, as they apply to (...) a wide number of materially diverse systems. In this article, we examine the various attempts to integrate abstract models within the mechanist program, also presenting a further challenge: the Heimburg-Jackson model, which was introduced as an alternative to the Hodgkin-Huxley model. We argue that although the notion of abstraction as the omission of irrelevant mechanistic details appears to give a mechanistic solution for accommodating abstract models, this notion does not suit models whose epistemic strategy is not decompositional. As a result, the mechanist has to choose whether to dilute the mechanistic approach nearly beyond recognition or to claim that many, if not most, abstract theoretical models do not deliver mechanistic explanations, or qualify as explanatory at all. (shrink)

It is a widespread assumption in philosophy of science that data is what is explained by theory—that data itself is not explanatory. I draw on instances of representational and explanatory practice from mammalian chronobiology to suggest that this assumption is unsustainable. In many instances, biologists employ representations of data in explanatory ways that are not reducible to constraints on or evidence for representations of mechanisms. Data graphs are used to exemplify relationships between quantities in the mechanism, and often these representations (...) are necessary for explaining particular aspects of the phenomena under study. I argue that this kind of representation is distinct from representing laws or generalizations, and its primary purpose is to convey particular types or patterns of quantitative relationships. The benefit of the analysis is two-fold. First, it provides a more accurate account of explanatory practice in broadly mechanistic analysis in biology. Second, it suggests that there is not an explanatory “fundamental” type of representation in biology. Rather, the practice of explanation consists in the construction of different types of representations and their employment for distinct explanatory purposes. (shrink)

We outline a framework of multilevel neurocognitive mechanisms that incorporates representation and computation. We argue that paradigmatic explanations in cognitive neuroscience fit this framework and thus that cognitive neuroscience constitutes a revolutionary break from traditional cognitive science. Whereas traditional cognitive scientific explanations were supposed to be distinct and autonomous from mechanistic explanations, neurocognitive explanations aim to be mechanistic through and through. Neurocognitive explanations aim to integrate computational and representational functions and structures across multiple levels of organization in order to explain (...) cognition. To a large extent, practicing cognitive neuroscientists have already accepted this shift, but philosophical theory has not fully acknowledged and appreciated its significance. As a result, the explanatory framework underlying cognitive neuroscience has remained largely implicit. We explicate this framework and demonstrate its contrast with previous approaches. (shrink)

Mechanistic models in molecular systems biology are generally mathematical models of the action of networks of biochemical reactions, involving metabolism, signal transduction, and/or gene expression. They can be either simulated numerically or analyzed analytically. Systems biology integrates quantitative molecular data acquisition with mathematical models to design new experiments, discriminate between alternative mechanisms and explain the molecular basis of cellular properties. At the heart of this approach are mechanistic models of molecular networks. We focus on the articulation and development of mechanistic (...) models, identifying five constraints which guide the articulation of models in molecular systems biology. These constraints are not independent of one another, with the result that modeling becomes an iterative process. We illustrate the use of these constraints in the modeling of the mechanism for bistability in the lac operon. (shrink)

In this article, two issues regarding mechanisms are discussed. The first concerns the relationships between “mechanism description” and “mechanism explanation.” It is proposed that it is rather plausible to think of them as two distinct epistemic acts. The second deals with the different molecular biology explanatory contexts, and it is shown that some of them require physics and its laws.

Explanations of biological phenomena such as cell division, protein synthesis or circadian rhythms commonly take the form of models of the responsible mechanisms. Recently philosophers of science have attempted to analyze this practice, presenting mechanisms as organized collections of parts performing operations that together produce the phenomenon. But in some cases what researchers seek to explain is not a general phenomenon, but a specific feature of a more fine-grained phenomenon. In some of these cases, it is not the model of (...) the mechanism that performs the explanatory work. I consider a case in which the investigator offered an abstract representation of a fine-grained phenomenon to show why in had the feature in question. I consider a second case in which a researcher abstracted from the mechanism to identify a design principle that explains why the functioning mechanism exhibits a specific feature. (shrink)

A survey of models in immunology is conducted and distinct kinds of models are characterized based on whether models are material or conceptual, the distinctiveness of their epistemic purpose, and the criteria for evaluating the goodness of a model relative to its intended purpose. I argue that the diversity of models in interdisciplinary fields such as immunology reflects the fact that information about the phenomena of interest is gathered from different sources using multiple methods of investigation. To each model is (...) attached a description specifying how information about a phenomenon of interest has been acquired, highlighting points of commonality and difference between the methodological and epistemic histories of the information encapsulated in different models. These points of commonality and difference allow investigators to integrate findings from different models into more comprehensive explanatory accounts, as well as to troubleshoot anomalies and faulty accounts by going back to the original building blocks. (shrink)

In this field guide, I distinguish five separate senses with which the term ‘mechanism’ is used in contemporary philosophy of science. Many of these senses have overlapping areas of application but involve distinct philosophical claims and characterize the target mechanisms in relevantly different ways. This field guide will clarify the key features of each sense and introduce some main debates, distinguishing those that transpire within a given sense from those that are best understood as concerning two distinct senses. The ‘new (...) mechanisms’ sense is the primary sense from which other senses will be distinguished. In part II of this field guide, I consider three further senses of the term that are ontologically ‘flat’ or at least not explicitly hierarchical in character: equations in structural equation models of causation, causal-physical processes, and information-theoretic constraints on states available to systems. After characterizing each sense, I clarify its ontological commitments, its methodological implications, how it figures in explanations, its implications for reduction, and the key manners in which it differs from other senses of mechanism. I conclude that there is no substantive core meaning shared by all senses, and that debates in contemporary philosophy of science can benefit from clarification regarding precisely which sense of mechanism is at stake. (shrink)

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)

Over the past three decades, philosophy of science has grown increasingly “local.” Concerns have switched from general features of scientific practice to concepts, issues, and puzzles specific to particular disciplines. Philosophy of neuroscience is a natural result. This emerging area was also spurred by remarkable recent growth in the neurosciences. Cognitive and computational neuroscience continues to encroach upon issues traditionally addressed within the humanities, including the nature of consciousness, action, knowledge, and normativity. Empirical discoveries about brain structure and function suggest (...) ways that “naturalistic” programs might develop in detail, beyond the abstract philosophical considerations in their favor. -/- The literature distinguishes “philosophy of neuroscience” and “neurophilosophy.” The former concerns foundational issues within the neurosciences. The latter concerns application of neuroscientific concepts to traditional philosophical questions. Exploring various concepts of representation employed in neuroscientific theories is an example of the former. Examining implications of neurological syndromes for the concept of a unified self is an example of the latter. In this entry, we will assume this distinction and discuss examples of both. (shrink)

Proponents of mechanistic explanation have argued that dynamical models are mere phenomenal models, in that they describe rather than explain the scientific phenomena produced by complex systems. I argue instead that dynamical models can, in fact, be explanatory. Using an example from neuroscientific research on epilepsy, I show that dynamical models can meet the explanatory demands met by mechanistic models, and as such occupy their own unique place within the space of explanatory scientific models.

This paper employs an interventionist framework to elucidate some issues having to do with explanation in neurobiology and with the differences between mechanistic and non-mechanistic explanations.

Craver’s (2007) account of explanation in neurobiology offers one of the most sophisticated explications of the mechanism concept. This paper argues that despite groundbreaking advances in understanding mechanistic explanation, serious challenges remain. The first goal of this paper is to address the notorious problem of explanatory relevance concerning mechanistic explanation. I argue that Craver underestimates the importance of pragmatic constraints on the individuation of mechanisms, and that his suggestion for a solution of the explanatory relevance problem is therefore insufficient on (...) several counts. My second goal is to develop an alternative that explicitly incorporates both pragmatic and ontic aspects of mechanism individuation. (shrink)

There are two traditions of thinking about idealization offering almost opposite views on their functioning and epistemic status. While one tradition views idealizations as epistemic deficiencies, the other one highlights the epistemic benefits of idealization. Both of these, however, identify idealization with misrepresentation. In this article, we instead approach idealization from the artifactual perspective, comparing it to the distortion-to-reality accounts of idealization, and exemplifying it through the case of the Hodgkin and Huxley model of nerve impulse. From the artifactual perspective, (...) the epistemic benefits and deficiencies introduced by idealization frequently come in a package due to the way idealization draws together different resources in model construction. Accordingly, idealization tends to be holistic in that it is not often easily attributable to just some specific parts of the model. Instead, the idealizing process tightly embeds theoretical concepts and formal tools into the construction of a model. (shrink)

I examine the adequacy of the causal graph-structural equations approach to causation for modeling biological mechanisms. I focus in particular on mechanisms with complex dynamics such as the PER biological clock mechanism in Drosophila. I show that a quantitative model of this mechanism that uses coupled differential equations – the well-known Goldbeter model – cannot be adequately represented in the standard causal graph framework, even though this framework does permit causal cycles. The reason is that the model contains dynamical information (...) about the mechanism that concerns causal properties but that does not correspond to variables that could be subject to independent interventions. Thus, a representation of the mechanisms as a causal structural model necessarily suppresses causally relevant information. (shrink)