Explanation in biology has long been characterized as being very different from explanation in other scientific disciplines, very much so from explanation in physics. One of the reasons was the existence in biology of explanation types that were unheard of in the physical sciences: teleological explanations (e.g. Hull 1974), evolutionary explanations (e.g. Mayr 1988), or even functional explanations (e.g. Neander 1991). More recently, and owing much to the rise of molecular biology, biological explanations (...) have been depicted as mechanisms (e.g; Machamer, Darden and Craver 2000). The aim of this volume is to shed some new light on the diversity of explanation types in biology. What are the different types of explanation that occur in biology? Are these types of explanation specific to particular sub-disciplines of biology, or to particular types of problems across biology? How do they relate to each another? Do they compete with one another for answering the same questions? Or do they complement each other, providing insights to different questions? What are the reasons for such diversity? Can this diversity be overcome by a broader unifying model of explanation or is it more profound and irreducible? Why? This volume aims at making sense of this diversity of types of explanations that are found in biology, of their relationship with one another. After all, explanation in biology may prove not only different from explanation in the physical sciences, but also much more diverse than originally anticipated. (shrink)

This essay analyzes and develops recent views about explanation in biology. Philosophers of biology have parted with the received deductive-nomological model of scientific explanation primarily by attempting to capture actual biological theorizing and practice. This includes an endorsement of different kinds of explanation (e.g., mathematical and causal-mechanistic), a joint study of discovery and explanation, and an abandonment of models of theory reduction in favor of accounts of explanatory reduction. Of particular current interest are philosophical (...) accounts of complex explanations that appeal to different levels of organismal organization and use contributions from different biological disciplines. The essay lays out one model that views explanatory integration across different disciplines as being structured by scientific problems. I emphasize the philosophical need to take the explanatory aims pursued by different groups of scientists into account, as explanatory aims determine whether different explanations are competing or complementary and govern the dynamics of scientific practice, including interdisciplinary research. I distinguish different kinds of pluralism that philosophers have endorsed in the context of explanation in biology, and draw several implications for science education, especially the need to teach science as an interdisciplinary and dynamic practice guided by scientific problems and explanatory aims. (shrink)

This paper is concerned with the problem of giving a correct analysis of function statements as they are used in biology. Examples of such statements are (1) The function of the myelin sheath is to insulate the nerve fiber and (2) The function of chlorophyll is to enable photosynthesis to take place. After criticizing analyses of such statements developed by Braithwaite, Nagel and Hempel an analysis is presented by the author. Finally the question of whether function statements are explanations (...) is discussed briefly. (shrink)

This paper evaluates Kuipers' account of functional explanation in biology in view of an example of such an explanation taken from real biology. The example is the explanation of why electric fishes swim backwards (Lannoo and Lannoo 1993). Kuipers' account depicts the answer to a request for functional explanation as consisting only of statements that articulate a certain kind of consequence. It is argued that such an account fails to do justice to the main (...) insight provided by the example explanation, namely the insight into why backwards swimming is needed by fishes that locate their food by means of an electric radar. The paper sketches an improved account that does justice to this kind of insight. It is argued that this account is consistent with and complementary to Kuipers' insight that function attributions are established by means of a process of hypothetico-deductive reasoning guided by a heuristic principle. (shrink)

Among philosophers of science, there is now a widespread agreement that the DN model of explanation is poorly equipped to account for explanations in biology. Rather than identifying laws, so the consensus goes, researchers explain biological capacities by constructing a model of the underlying mechanism.We think that the dichotomy between DN explanations and mechanistic explanations is misleading. In this article, we argue that there are cases in which biological capacities are explained without constructing a model of the underlying (...) mechanism. Although these explanations do not conform to Hempel’s DN model, they do invoke more or less stable generalisations. Because they invoke generalisations and have the form of an argument, we call them inferential explanations. We support this claim by considering two examples of explanations of biological capacities: pigeon navigation and photoperiodism. Next, we will argue that these non-mechanistic explanations are crucial to biology in three ways: sometimes, they are the only thing we have, they are heuristically useful, and they provide genuine understanding and so are interesting in their own right.In the last sections we discuss the relation between types of explanations and types of experiments and situate our views within some relevant debates on explanatory power and explanatory virtues. (shrink)

When philosophers discuss the topic of explanation, they usually have in mind the following question: given the beliefs one has and some proposition that one wishes to explain, which subset of the beliefs constitutes an explanation of the target proposition? That is, the philosophical ‘problem of explanation’ typically has bracketed the issue of how one obtains the beliefs; they are taken as given. The problem of explanation has been the problem of understanding the relation ‘x explains (...) y’. Since Hempel did so much to canonize this way of thinking about explanation, it deserves to be called ‘Hempel's problem’. (shrink)

When philosophers discuss the topic of explanation, they usually have in mind the following question: given the beliefs one has and some proposition that one wishes to explain, which subset of the beliefs constitutes an explanation of the target proposition? That is, the philosophical ‘problem of explanation’ typically has bracketed the issue of how one obtains the beliefs; they are taken as given. The problem of explanation has been the problem of understanding the relation ‘x explains (...) y’. Since Hempel did so much to canonize this way of thinking about explanation, it deserves to be called ‘Hempel's problem’. (shrink)

One of biology's fundamental aims is to generate understanding of the living world around—and within—us. In this chapter, I aim to provide a relatively nonpartisan discussion of the nature of explanation in biology, grounded in widely shared philosophical views about scientific explanation. But this discussion also reflects what I think is important for philosophers and biologists alike to appreciate about successful scientific explanations, so some points will be controversial, at least among philosophers. I make three main (...) points: (1) causal relationships and broad patterns have often been granted importance to scientific explanations, and they are in fact both important; (2) some explanations in biology cite the components of or processes in systems that account for the systems’ features, whereas other explanations feature large-scale or structural causes that influence a system; and (3) there can be multiple different explanations of a given biological phenomenon, explanations that respond to different research aims and can thus be compatible with one another even when they may seem to disagree. (shrink)

Until recently, the notions of function and multiple realization were supposed to save the autonomy of psychological explanations. Furthermore, the concept of supervenience presumably allows both dependence of mind on brain and non-reducibility of mind to brain, reconciling materialism with an independent explanatory role for mental and functional concepts and explanations. Eliminativism is often seen as the main or only alternative to such autonomy. It gladly accepts abandoning or thoroughly reconstructing the psychological level, and considers reduction if successful as equivalent (...) with elimination. In comparison with the philosophy of mind, the philosophy of biology has developed more subtle and complex ideas about functions, laws, and reductive explanation than the stark dichotomy of autonomy or elimination. It has been argued that biology is a patchwork of local laws, each with different explanatory interests and more or less limited scope. This points to a pluralistic, domain-specific and multi-level view of explanations in biology. Explanatory pluralism has been proposed as an alternative to eliminativism on the one hand and methodological dualism on the other hand. It holds that theories at different levels of description, like psychology and neuroscience, can co-evolve, and mutually influence each other, without the higher-level theory being replaced by, or reduced to, the lower-level one. Such ideas seem to tally with the pluralistic character of biological explanation. In biological psychology, explanatory pluralism would lead us to expect many local and non-reductive interactions between biological, neurophysiological, psychological and evolutionary explanations of mind and behavior. This idea is illustrated by an example from behavioral genetics, where genetics, physiology and psychology constitute distinct but interrelated levels of explanation. Accounting for such a complex patchwork of related explanations seems to require a more sophisticated and precise way of looking at levels than the existing ideas on (reductive and non-reductive) explanation in the philosophy of mind. (shrink)

This paper develops an account of explanation in biology which does not involve appeal to laws of nature, at least as traditionally conceived. Explanatory generalizations in biology must satisfy a requirement that I call invariance, but need not satisfy most of the other standard criteria for lawfulness. Once this point is recognized, there is little motivation for regarding such generalizations as laws of nature. Some of the differences between invariance and the related notions of stability and resiliency, (...) due respectively to Sandra Mitchell and Brian Skyrms, are explored. (shrink)

Besides mechanistic explanations of phenomena, which have been seriously investigated in the last decade, biology and ecology also include explanations that pinpoint specific mathematical properties as explanatory of the explanandum under focus. Among these structural explanations, one finds topological explanations, and recent science pervasively relies on them. This reliance is especially due to the necessity to model large sets of data with no practical possibility to track the proper activities of all the numerous entities. The paper first defines topological (...) explanations and then explains why topological explanations and mechanisms are different in principle. Then it shows that they are pervasive both in the study of networks—whose importance has been increasingly acknowledged at each level of the biological hierarchy—and in contexts where the notion of selective neutrality is crucial; this allows me to capture the difference between mechanisms and topological explanations in terms of practical modelling practices. The rest of the paper investigates how in practice mechanisms and topologies are combined. They may be articulated in theoretical structures and explanatory strategies, first through a relation of constraint, second in interlevel theories, or they may condition each other. Finally, I explore how a particular model can integrate mechanistic informations, by focusing on the recent practice of merging networks in ecology and its consequences upon multiscale modelling. (shrink)

A common view of how-possibly explanations in biology treats them as explanatorily incomplete. In addition to this interpretation of how-possibly explanation, I argue that there is another interpretation, one which features what I term “explanatory strategies.” This strategy-centered interpretation of how-possibly explanation centers on there being a different explanatory context within which how-possibly explanations are offered. I contend that, in conditions where this strategy context is recognized, how-possibly explanations can be understood as complete explanations. I defend this (...) alternative interpretation by analyzing the explanatory value of simple physical models the nineteenth century developmental biologist Wilhelm His constructed for animal development. (shrink)

During the war, I worked in aircraft design. About a year after D-day, an exhibition was arranged at Farnborough of the mass of German equipment that had been captured, including the doodlebug and the V2 rocket. I and a friend spent a fascinating two days wandering round the exhibits. The questions that kept arising were ‘Why did they make it like that?’, or, equivalently ‘I wonder what that is for?’ We were particularly puzzled by a gyroscope in the control system (...) of the V2. One's first assumption was that the gyroscope maintained the rocket on its course, but, instead of being connected to the steering vanes, it was connected to the fuel supply to the rocket. Ultimately my friend remembered that the rate of precession of a gyro depends on acceleration, and saw that the Germans had used this fact to design an ingenious device for switching off the engines when the required velocity had been reached. (shrink)

The inapplicability of variations on theory reduction in the context of genetics and their irrelevance to ongoing research has led to an anti-reductionist consensus in philosophy of biology. One response to this situation is to focus on forms of reductive explanation that better correspond to actual scientific reasoning (e.g. part–whole relations). Working from this perspective, we explore three different aspects (intrinsicality, fundamentality, and temporality) that arise from distinct facets of reductive explanation: composition and causation. Concentrating on these (...) aspects generates new forms of reductive explanation and conditions for their success or failure in biology and other sciences. This analysis is illustrated using the case of protein folding in molecular biology, which demonstrates its applicability and relevance, as well as illuminating the complexity of reductive reasoning in a specific biological context. (shrink)

Biologists in many different fields of research give how-possibly explanations of the phenomena they study. Although such explanations lack empirical support, and might be regarded by some as unscientific, they play an important heuristic role in biology by helping biologists develop theories and concepts and suggesting new areas of research. How-possibly explanations serve as a useful framework for conducting research in the absence of adequate empiri cal data, and they can even become how-actually explanations if they gain enough empirical (...) support. (shrink)

Many biologists and philosophers have worried that importing models of reasoning from the physical sciences obscures our understanding of reasoning in the life sciences. In this paper we discuss one example that partially validates this concern: part-whole reductive explanations. Biology and physics tend to incorporate different models of temporality in part-whole reductive explanations. This results from differential emphases on compositional and causal facets of reductive explanations, which have not been distinguished reliably in prior philosophical analyses. Keeping these two facets (...) distinct facilitates the identifi cation of two further aspects of reductive explanation: intrinsicality and fundamentality. Our account provides resources for discriminating between different types of reductive explanation and suggests a new approach to comprehending similarities and differences in the explanatory reasoning found in biology and physics. (shrink)

The notion of how-possibly explanations emerged with William Dray in response to Carl Hempel’s influential deductive-nomological model of scientific explanation. Dray’s aim was to distinguish explanations of states of affairs that might occur, in contrast to the aim of D-N explanations working to establish that states of affairs must actually occur. More recently, interest in how-possibly explanations has been particularly keen among philosophers of biology. One of the concerns philosophers of biology have focused on is whether how-possibly (...) explanations are “complete” explanations. Dray considered how-possibly explanations to be complete explanations, but the view that they are incomplete explanations became prominent with Robert Brandon’s articulation of successful adaptation explanation in evolutionary biology. According to Brandon, how-possibly adaptation explanations are incomplete, since they lack the requisite empirical evidence. Moreover, Brandon’s characterization of how-possibly explanation can be seen as having set the stage for thinking broadly about how-possibly explanations in biology. Echoing Brandon’s account for adaptation explanations, for example, David Resnik contends that how-possibly explanations are inherently incomplete, and yet they serve an important heuristic role throughout biology by setting out an agenda for future research. (shrink)

It seems that the above account of explanation-strategy in the area of temperature adaptation underscores many of the points made earlier. First, it discloses the fruitful interaction of classical, evolutionary, and molecular approaches. Secondly, it indicates that biological characterizations are not rival accounts to chemical ones. Thirdly, it stresses the importance of the DNA sequence order in chemical explanations of biological organisms.One feature which this area does not seem to reveal, which genetics does, is the development of a biological (...) (that is, non-chemical) theory which would explain temperature adaptation in nonchemical terms. What this shows is that one should not always expect systematization at the biological level, in the sense of being able to refer many phenomena to a small number of basic principles. The future development of laws and theories in biological terms in this area is, of course, not precluded by the fact that none have yet been discovered. The case does indicate that physical and chemical language and theories are extremely useful in biological investigations, and the physicochemical approach may well be the best approach in this area. In any event, ultimate reduction of the biological to the chemical seems to be a foregone non-controversial conclusion here as elsewhere. (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.

In opposition to the etiological conception of biological functions, this paper attempts to show that explanations by natural selection, far from justifying functional attributions, presuppose them, and that these attributions may be understood by appealing to a particular specification of the systemic conception of function, that is, the biological conception of function. This paper argues that the etiological conception of function is based on two fundamental errors: confusing the concept of function with the concept of adaptation, and confusing selectional explanations (...) with functional analyses. This explains the inappropriate demand that those analyses include an etiological explanatory content and make possible the distinction between adaptations and exaptations. (shrink)

Is Neo-Aristotelian hylomorphism compatible mechanistic science? In this essay I forge a rapprochement between Neo-Aristotelian hylomorphism and the "new mechanist philosophy" in biology, neuroscience, and psychology by drawing attention to their shared commitments concerning multilevel organization, mechanisms, and teleology. Significantly, the new mechanists endorse organization realism (a touchstone of hylomorphism). Similarly, Neo-Aristotelian hylomorphism is committed to the reality of mechanisms or causal powers that produce, underlie, or maintain the behavior of (i) phenomena that are constituted through the (ii) spatial, (...) temporal, and active organization of their (iii) component entities and (iv) component activities (the four hallmarks of the new mechanist philosophy). In the course of the essay I address potential disagreements between these two positions pertaining to emergence, downward causation, and teleology. I conclude that Neo-Aristotelian hylomorphism should not been seen as fundamentally opposed to mechanisms; rather, hylomorphism provides a rigorous ontological framework that complements the insights of the new mechanist philosophy of biology, neuroscience and psychology. (shrink)

Back cover: This book develops a philosophical account that reveals the major characteristics that make an explanation in the life sciences reductive and distinguish them from non-reductive explanations. Understanding what reductive explanations are enables one to assess the conditions under which reductive explanations are adequate and thus enhances debates about explanatory reductionism. The account of reductive explanation presented in this book has three major characteristics. First, it emerges from a critical reconstruction of the explanatory practice of the life (...) sciences itself. Second, the account is monistic since it specifies one set of criteria that apply to explanations in the life sciences in general. Finally, the account is ontic in that it traces the reductivity of an explanation back to certain relations that exist between objects in the world (such as part-whole relations and level relations), rather than to the logical relations between sentences. Beginning with a disclosure of the meta-philosophical assumptions that underlie the author’s analysis of reductive explanation, the book leads into the debate about reduction(ism) in the philosophy of biology and continues with a discussion on the two perspectives on explanatory reduction that have been proposed in the philosophy of biology so far. The author scrutinizes how the issue of reduction becomes entangled with explanation and analyzes two concepts, the concept of a biological part and the concept of a level of organization. The results of these five chapters constitute the ground on which the author bases her final chapter, developing her ontic account of reductive explanation. (shrink)

This paper attempts to elucidate three characteristics of causal relationships that are important in biological contexts. Stability has to do with whether a causal relationship continues to hold under changes in background conditions. Proportionality has to do with whether changes in the state of the cause “line up” in the right way with changes in the state of the effect and with whether the cause and effect are characterized in a way that contains irrelevant detail. Specificity is connected both to (...) David Lewis’ notion of “influence” and also with the extent to which a causal relation approximates to the ideal of one cause–one effect. Interrelations among these notions and their possible biological significance are also discussed. (shrink)

Biology and history are often viewed as closely related disciplines, with biology informed by history, especially in its task of charting our evolutionary past. Maximizing the opportunities for cross-fertilization in these two fields requires an accurate reckoning of their commonalities and differences-precisely what this volume sets out to achieve. Specially commissioned essays by a team of recognized international researchers cover the full panoply of topics in these fields and include notable contributions on the correlativity of evolutionary and historical (...) explanations, applying to history the latest causal-mechanical approach in the philosophy of biology, and the question of generalized laws that might pertain across the two subjects. The collection opens with a vital interrogation of general issues on explanation that apart from potentially fruitful areas of interaction (could the etiology of the causal-mechanical perspective in biology account for the historical trajectory of the Roman Empire?) this volume also seeks to chart relative certainties distinguishing explanations in biology and history. It also assesses techniques such as the use of probabilities in biological reconstruction, deployed to overcome the inevitable gaps in physical evidence on early evolution. Methodologies such as causal graphs and semantic explanation receive in-depth analysis. Contributions from a host of prominent and widely read philosophers ensure that this new volume has the stature of a major addition to the literature. (shrink)

It is argued that once biological systems reach a certain level of complexity, mechanistic explanations provide an inadequate account of many relevant phenomena. In this article, I evaluate such claims with respect to a representative programme in systems biological research: the study of regulatory networks within single-celled organisms. I argue that these networks are amenable to mechanistic philosophy without need to appeal to some alternate form of explanation. In particular, I claim that we can understand the mathematical modelling techniques (...) of systems biologists as part of a broader practice of constructing and evaluating mechanism schemas. This argument is elaborated by considering the case of bacterial chemotactic networks, where some research has been interpreted as explaining phenomena by means of abstract design principles. (shrink)

Molecular biologists and biochemists often use diagrams to present hypotheses. Analysis of diagrams shows that their content can be expressed with linguistic representations. Why do biologists use visual representations instead? One reason is simple comprehensibility: some diagrams present information which is readily understood from the diagram format, but which would not be comprehensible if the same information was expressed linguistically. But often diagrams are used even when concise, comprehensible linguistic alternatives are available. I explain this phenomenon by showing why diagrammatic (...) representation is especially well suited for a particular kind of explanation common in molecular biology and biochemistry: namely, functional analysis, in which a capacity of the system is explained in terms of capacities of its component parts. (shrink)

In biology a great variety of models can be distinguished: there is a gradation scale from the more realistic to the more idealistic ones. The place of a model on this scale depends on the role of the fundamental ideas, apriorisms and empirisms, which inspire the direction of thought. The relation between reality, models and explanatory theory is worked out. The interplay between model and ideas makes it understandable why in biology several kinds of explanation are possible.

Relations between physics and biology have been always difficult. One reason is that physical approaches to the phenomena of life have frequently been conceived by their authors as alternatives to biological explanations. My argument is that molecular descriptions and explanations have been pushed so far that they have reached their limits: these limits constitute a favourable niche in which physical explanations can develop. I will focus on the field of molecular and cell biology and give many examples of (...) these recent physical studies made possible by the precision of molecular observations. The nature of these niches is probably diverse. I consider that it is too early to have a global view of the interactions between biological and physical explanations, and to organize them into different categories. Such interactions are not new within the life sciences: the history of biology reveals a complex, permanently moving landscape of interactions between biological and physical explanations. (shrink)

It is widely assumed that mechanistic explanations are causal explanations. Many prominent new mechanists endorse interventionism as the correct analysis of explanatory causal models in biology and other fields. This article argues that interventionism is not entirely satisfactory in this regard. A case study of Jacob and Monod’s operon model shows that at least some important mechanistic explanations in biology present significant contrasts with the interventionist account. This result motivates a more inclusive approach to mechanistic explanation, allowing (...) for noncausal aspects. (shrink)

This article presents a challenge that those philosophers who deny the causal interpretation of explanations provided by population genetics might have to address. Indeed, some philosophers, known as statisticalists, claim that the concept of natural selection is statistical in character and cannot be construed in causal terms. On the contrary, other philosophers, known as causalists, argue against the statistical view and support the causal interpretation of natural selection. The problem I am concerned with here arises for the statisticalists because the (...) debate on the nature of natural selection intersects the debate on whether mathematical explanations of empirical facts are genuine scientific explanations. I argue that if the explanations provided by population genetics are regarded by the statisticalists as non-causal explanations of that kind, then statisticalism risks being incompatible with a naturalist stance. The statisticalist faces a dilemma: either she maintains statisticalism but has to renounce naturalism; or she maintains naturalism but has to content herself with an account of the explanations provided by population genetics that she deems unsatisfactory. This challenge is relevant to the statisticalists because many of them see themselves as naturalists. (shrink)

Taxonomists in biology have traditionally been concerned to delimit and classify actual (or previously actual) biological forms orkinds. But not all useful classification schemes are of actualisedforms. This paper focuses on the need to delimit and classifynon‐actual forms when offering contrastive explanations in evolutionary biology. Such a classification scheme sorts actual and nonfactual forms according to their modal status. Such a sorting has been offered by theoretical morphologists, but these efforts havepaid insufficient attention to the metaphysics of modality. (...) Contemporary approaches to the metaphysics and epistemology of modalityare also found wanting. The paper ends by arguing that the needed intellectual resources are to be found in the imaginative use of Aristotle. (shrink)

This paper: (1) gives a schema of the logical structure of functional explanation in biology; (2) shows that it falls under the covering law model of explanation by proving that the explanandum follows from the explanans; and (3) supports the claim that it captures the logical structure underlying the biological usage by analyzing in detail two cases from biology.