Biological Complexity and Integrative Pluralism (Cambridge Studies in Philosophy and Biology)

Reductionism in Biology
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Cite this Email this Add to favourites Print this page. You must be logged in to Tag Records. Broken link? Introduction 2. Constitutive Complexity 2. Compositional Complexity and the Superorganism Metaphor 3. Dynamic Complexity 3. The Evolution of Division of Labor 4. Evolved Diversity 4. Competing Units of Selection? A Case of Symbiosis 4.

The Units of Behavior in Evolutionary Explanations 4. On Biological Functions 5. Laws more Laws 5. Critics of reductionism do not argue that each higher level type corresponds to several lower level tokens which is trivially true or that molecular biology can study token phenomena only. Rather, the point is that a higher level type corresponds to several lower level types , and for each of these lower level kinds a distinct molecular account obtains.

Thus, a unified as opposed to disjunctive causal explanation of the higher level phenomenon appears impossible. In some cases, disjunctive explanations at a lower level seem to explain better than unified explanations at higher levels. Elliott Sober extends this line of argument: an explanation in terms of higher level kinds can be more unified or general by encompassing heterogeneous lower level kinds, but this same heterogeneity exhibits important differences in scientifically interesting properties.

Only the lower level science can account for such differences and therefore, in this sense, offers a deeper explanation than the higher level science.

As a result, explanations are not better or worse along a one-dimensional scale, but there are several epistemic virtues an explanation can possess—generality as well as depth. Sometimes a lower level explanation is better relative to one epistemic property, while a higher level explanation is preferable relative to another epistemic quality, such as unification.

And unification itself can be explicated differently in terms of functional or structural features of biological systems see, e. It is natural for contemporary accounts of explanatory reduction to focus on mechanisms and the study of model organisms. One reply to the multiple realization objection from this perspective explicitly argues for token-token reduction, discussing how the causal description of a token mechanism explains Delehanty However, this stratagem ignores the key point of contention because token-token reduction is uncontroversial.

It is a relatively trivial claim that follows from physicalism. The focus on token phenomena or individual organisms fails to address an important epistemological feature of scientific knowledge: some scientific explanations pertain to a single organism only, but in general explanations cover types of phenomena.

In many biological contexts, such as developmental biology, the intellectual aim is to explain phenomena instantiated in many organisms; and the use of model organisms is predicated on this aim. Explaining development—as understood by many developmental biologists—means to have an account of ontogeny exhibited by individual organisms belonging to a larger group e. As the same morphological structure may develop in different species by means of different processes and based on the action of different genes, multiple realization obtains and is relevant for such explanations Laubichler and Wagner Moreover, even if the aim is to explain a feature of a single organism—the situation considered above by Weber —it does not follow that an explanation should exclusively appeal to the factors on the lowest level.

Levels of Organization in Biology

An explanation should only include factors that are explanatorily relevant; an irrelevant factor is one whose omission from or modification in the explanation does not prevent the explanandum from following Strevens This also holds for mechanistic explanation, which should cite those features as components of a mechanism whose modification would make a causal difference to the explanandum phenomenon Craver For instance, chromosomes are relevant parts of the mechanisms needed to explain Mendelian patterns of inheritance Darden , but this particular explanation will go through regardless of what the lower level, molecular composition of the chromosomes is e.

In the context of developmental biology, even during the development of a single organism, a structure may be present across time and have a stable developmental-functional role, yet its underlying molecular and cellular basis may change Brigandt , Kirschner Entities above the molecular level can be more robust so that when some developmental processes change or break down these higher level entities are not modified or removed.

Knocking out a gene need not have any impact on ontogeny due to genetic redundancy, while eliminating a particular cell or group of cells may dramatically interfere with normal development Brigandt , Mitchell A higher level causal connection can be more salient in that the regular operation of the mechanism strongly depends on the presence of this kind of causal relation. This is not to deny that such a higher level cause is realized by molecular entities and their interactions i.

But if a higher level process is less sensitive to disturbance or intervention than certain molecular causes, the more powerful mechanistic explanation can be the one that appeals to more robust causal connections. This is one reason why some biological sciences preferentially invoke higher level entities and processes in the face of multiple realization, explaining in terms of higher level natural kinds and causes. In summary, the level s of organization a successful explanation addresses often depends on the particular explanandum. If the aim is to explain a type of phenomenon, multiple realization issues many-one may arise. In the explanation of a token phenomenon, the explanation should include lower level factors only to the extent that they are explanatorily relevant for that particular explanandum. Sarkar's account of explanatory reduction draws attention to the requirement of representation: natural phenomena must be symbolized, embodied, pictured, or designated through media such as equations, scale miniatures, or abstract diagrams.

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Every reductive explanation in science involves a representation of the systems or domains to be related by reduction. Note that this is not the question of how theory structure is represented; see, e.

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Almost all of the discussion surrounding one-many Section 4. Sarkar distinguishes three representational criteria for a reductive explanation: i fundamentalism: the explanation of the phenomena relies entirely on features that pertain to a more fundamental realm; ii abstract hierarchy: the system is represented as a hierarchy, where the lower levels are regarded as the more fundamental; and, iii spatial hierarchy: the abstract hierarchy is rendered physical by the requirement that the entities on the lower level are spatial parts of the entities at the higher levels see Section 3.

Whether or not there are one-many or many-one relations between different hierarchical levels turns on the nature of the hierarchy invoked and how it is characterized. The decomposition of a system is not a univocal enterprise and, depending on the principles utilized, can generate both competing and complementary sets of part representations from the same system Kauffman , Wimsatt , Craver Certain governing principles are often visible, such as functional versus structural decompositions Bechtel and Richardson , Winther Therefore, prior to determinations of whether reductive explanations succeed or fail, questions of representational choice and adequacy need to be explicitly addressed.

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These representation issues are not idle with respect to differing explanations, both past and present, because the same line of argument about the non-independence of organismal parts constituted reasons for treating the organism from a non-reductionist standpoint Aristotle, Parts of Animals II.

Individuation of parts and their explanatory relationship to wholes is critically dependent on the principles utilized in representing them Love Wimsatt , , , has claimed that reduction and emergence are compatible within the context of explaining organizational complexity in living systems, going so far as to argue that reductionist methodology can identify emergence when specific conditions of aggregativity are delineated.

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In most discussions of epistemic reduction, both theoretical and explanatory, no explicit distinction has been drawn between constitutional or spatial relations arrangements and causal or temporal relations dynamics; but see Mitchell One source of the focus on spatial relations is the objection to reductionism from structural organization Polanyi ; cf.

Delehanty , Frost-Arnold , Schaffner , ch. Kellert , Nagel presciently recognized differences between spatial and temporal reasoning in his discussion of teleological explanation in part because he was reading the literature on explaining ontogeny by theoretical biologists; see Section 2. What this distinction misses is the possibility of diachronic aspects of part-whole or interlevel relations. Scientific explanations commonly invoke dynamic causal processes involving entities on several levels of organization Craver and Bechtel This is one of the core reasons why development is a persisting biological topic for reductionism.

2. Philosophical Accounts of Levels of Organization

During ontogeny there are causal interactions over time among parts and activities both homogeneous and heterogeneous to bring about new parts and activities both homogeneous and heterogeneous , eventually generating an integrated adult whole organism. Although temporal or process hierarchies have not received the same attention as spatial hierarchies such as in Wimsatt's aggregativity criteria , their importance is seen in explanations that invoke higher level features e.

For example, a study of the developmental origin of aortic arch asymmetry involves a mixture of reductionist and non-reductionist representations according to the criterion of temporality Yashiro et al. The explanation depicts several events in a sequential process. First, fundamental level properties gene expression change non-fundamental level properties arterial structure , which alters blood flow dynamics thereby fulfilling the temporality condition. These new dynamics induce a subsequent change in gene expression thereby violating the temporality condition.

This altered gene expression then produces the non-fundamental property of interest, aortic arch asymmetry thereby fulfilling the temporality condition. Thus, depending on how time is represented and which temporal partitions are utilized, an explanation may be reductive, non-reductive, or both.

1. Introduction

This is a good example of how ontological and epistemic reduction can yield strikingly different answers. A biologist using a higher level feature at an earlier time to explain a lower level feature at a later time a non-reductionist explanation by the criterion of temporality is not denying that this higher level feature is composed of lower level features e. Functional proteins are folded structures composed of amino acid components linked together into a linear chain. If we ask whether the folded protein is mereologically composed of its amino acid parts given current representations in molecular biology, then the affirmative answer seems to favor explanatory reduction with respect to the spatial hierarchy.

But if we ask whether the linear amino acid chain folds into a functional protein a causal process with a temporal dimension purely as consequence of its linked amino acid parts, then the answer is less clear. Empirical studies have emphasized the necessary role of other folded proteins to assist in the proper folding of newly generated linear amino acid chains Frydman That the linked amino acid components alone are insufficient causally, even if they are sufficient constitutionally, allows for a more explicit appreciation of the significance of temporality and dynamics Mitchell , especially because the relations of interest concern only molecular biological phenomena as opposed to higher levels of organization, such as cells or anatomy.

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A final representational feature is intrinsicality, i. In the protein folding case it appears that the failure of temporally indexed explanatory reduction involves causal powers of something extrinsic to an instance of the process under scrutiny i. This line of thought relates to the context-dependency objection Section 4.

Once we incorporate distinctions regarding different types of reduction ontological, epistemological, and methodological , the different interpretations of these types e. The multiplicity and heterogeneity of biological subdisciplines only reinforces this argument and suggests to some that we should move beyond reductionism entirely.

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Much of the past discussion about reduction presupposes a unitary view of the scientific enterprise that involves a close relation between natural kinds, laws, counterfactual dependence, explanation, and confirmation Godfrey-Smith One of the ubiquitous features of modern biology that seems prima facie incompatible with many methodological and epistemological reductionist theses is the proliferation and flourishing of diverse biological subdisciplines, molecular and otherwise.

Lindley Darden and Nancy Maull were at the forefront of this discussion with the concept of an inter-field theory Darden and Maull , Maull For example, the advent of the chromosome theory of inheritance in the s bridged the previously unrelated fields of Mendelian genetics which studied phenotypic patterns of inheritance across generations and cytology which dealt with the material contents of cells.

This interfield theory effected a unification of these two fields but Mendelian genetics and cytology were not reduced to each other, nor did the interfield theory reduce both fields. More recent work has articulated interfield relations without relying on the notion of a theory Bechtel , Burian , Grantham a, b, Mitchell This parallels the trend in models of explanatory reduction of moving away from theories as the only epistemic units of interest Section 3.

In this context, Alan Love , uses the notion of a problem agenda to argue that some scientific problems not only motivate interdisciplinary research but also provide structure for the requisite intellectual coordination see also Brigandt , , Brigandt and Love Problem agendas consist of numerous component questions that stand in systematic relations and are tied to associated standards of explanatory adequacy.

These standards imply what epistemic resources and different disciplines are needed to address the problem. The relations among component questions of the problem agenda i.

2. Historical Background: Philosophical and Scientific

Building on this perspective, Ingo Brigandt suggests that there is no single, linear hierarchy among scientific fields e. These philosophical accounts align with general research on interdisciplinarity, which has identified the presence and articulation of a complex question as a prerequisite Repko , Szostak , Thus, instead of unification being a regulative ideal with biologists seeking as much unity as possible in their explanations of the complex natural world e.

William Bechtel's , studies of cell biology document how the formation of new subdisciplines generates both integration among approaches and some disintegration due to new specializations. Bechtel's attention to the institutional underpinnings of biology also meshes with sociological analyses of disciplinary organization Gerson Many philosophers view experimental biology as fundamentally concerned with the discovery of mechanisms Darden and Craver , Tabery , where mechanisms are understood as akin though not equivalent to machines with interconnected, organized parts operating to produce regular or expected outcomes.

Conceiving of biological explanation in terms of mechanisms provides a philosophical framework that closely tracks biological practice and does not rely on the notion of a theory.