Abstract
In Chapter 6, I develop an ontic account of explanatory reduction in biology by analyzing paradigmatic and important examples of reductive (and non-reductive) explanations and discussions about reductionism from biological practice. I start with briefly specifying two concepts that occupy center stage in my account: the concept of a biological part (or of a part-whole relation) and the concept of levels of organization. On the basis of these conceptual clarifications I then answer the central question of my book, namely what are the characteristics that determine whether a biological explanation is reductive or not. The main result of my analysis of biological practice will be that reductive explanations in biology possess three features (two of which are necessary conditions, one of which is only a typical feature that most reductive explanations exhibit): they display a lower-level character, focus on factors that are internal to the biological object of interest, and describe the biological parts of this object only as parts in isolation.
“A prevalent… stance in biology is… reductionism, which predicates the study of biological systems at the lowest possible level with the objective of uncovering molecular and biochemical causes.” (Ana M. Soto and Carlos Sonnenschein 2010, 364)
“These examples support the importance of being able to think holistically, to look outwardly from the boundaries of the phenomenon under study and thereby place it in a more inclusive context.” (William Z. Lidicker 1988, 280)
“[G]ene sequencing and other techniques will soon have isolated all the cell’s individual parts and spelled out their isolated functions. Now, it is time to move beyond reductionism. […] Now we need to know how all these things are integrated.” (Robert F. Service 1999, 81)
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Notes
- 1.
Part-whole relations in general are analyzed in mereology (e.g., Simons 1987).
- 2.
In this book, I employ a very broad notion of environment, similar to Brandon’s “external environment” (1990, 47–49) but which is not restricted to the environment of organisms (since the category of biological objects comprises much more than just organisms).
- 3.
Craver makes a similar point with respect to levels (2007a, 187f).
- 4.
I use the term ‘line’ metaphorically. Since objects are extended in at least a three-dimensional space the boundary of Y is, of course, a surface, not a line.
- 5.
At least if one rejects four-dimensionalism (see e.g., Sider 2001).
- 6.
In what follows, I focus on processes and do not talk about occurrents in general for reasons of simplicity and because processes are of particular importance in the biological realm (Dupré 2012).
- 7.
However, if one endorses an etiological theory of function (rather than, for instance, a causal-role theory) the class of characteristic behaviors will be larger than the class of realizations of functions (cf. Kitcher 1993). I thus prefer not to speak about functions but rather about behaviors or processes in general.
- 8.
- 9.
Note that this holistic aspect does not render my account of biological parthood circular since one can identify a biological object Y and demarcate Y from its context by identifying its characteristic behaviors (and by identifying its natural boundary) without, at the same time, knowing what the biological parts of Y are.
- 10.
The conditions of spatial inclusion and of relevance are not completely detached from each other as, for instance, the existence of a natural boundary requires the existence of certain causal cut-offs at that boundary.
- 11.
In reference to Simon’s classic paper on “The Architecture of Complexity” (1962), Wimsatt states that “levels of organization are a deep, non-arbitrary, and extremely important feature of the ontological architecture of our natural world” (2007, 203). For a rejection of this view see Ladyman and Ross 2007, 55–57.
- 12.
Hence, I understand levels of organization to be levels of nature, not levels of science (as e.g., Oppenheim and Putnam 1958 did). That is, levels relate entities in the world such as objects, properties, or processes, not epistemic entities such as theories, explanations, or scientific disciplines.
- 13.
Wimsatt characterizes a level of organization as a “local maximum of predictability and regularity” (1976b, 238).
- 14.
I agree with Craver on the point that levels always are, as he frames it, “levels of behaving components” (2007a, 190) and not only levels of objects. In my terms, since levels are determined by part-whole relations, which cannot be identified by considering only objects (processes must be taken into account as well), not only are objects located on levels but objects that are involved in certain processes or that behave in a certain way.
- 15.
There might be additional reasons not to locate all of Y’s parts on the same level of organization (see the second point).
- 16.
This does not mean that these biologists accept the simple and, admittedly, unconvincing view that everything in nature is organized into a few, monolithic levels. An intermediate position is possible that rejects the radical view that levels are defined only in local explanatory contexts and conceives levels to be more global (but not too global) things.
- 17.
Since I want my account to apply to biology in general and not to be restricted to certain biological disciplines, the strategy Sarkar (1998), and Hüttemann and Love (2011) chose, namely to focus on the analysis of examples from only a few biological fields (e.g., genetics, molecular biology or developmental biology), is not a viable option for me.
- 18.
I admit that the choice of paradigmatic and important examples of reductive explanations in biology sometimes might be affected by a certain pre-concept of reduction one (explicitly or implicitly) endorses – for example, that reduction leads to simplification, that reduction is closely related to mechanisms and part-whole explanation, that reduction involves showing that something is nothing over and above something else, etc. It seems to me that this is not bad – as long as the influence of this pre-concept is not too strong, as long as this influence is revisable in the light of important insights about what scientists treat as reductive explanation, that is, as long as this influence does not result in imposing an ill-fitting view of reduction on biological practice.
- 19.
Also in biology itself molecular biology is frequently characterized as the “triumph of reductionism” (Fang and Casadevall 2011, 1402).
- 20.
This thesis should not be confused with the view of reduction that is known as “layer-cake reduction” (e.g. Rosenberg 2006, 28; Waters 2008; Steel 2004, 60) and which can be traced back to Oppenheim’s and Putnam’s layer-cake model of the unity of science (1958). My account of reduction is neither concerned with theory reduction (recall Chap. 3, Sect. 3.1), nor does it accept Oppenheim’s and Putnam’s concept of level.
- 21.
- 22.
Note that the direction of a reductive explanation runs in the opposite direction than the reductive method decomposition. Biological objects or systems are decomposed into their parts by looking down one or more levels, whereas the behavior of an object is explained by looking upward.
- 23.
Mikkelson uses this characterization of reductive explanation to offer a quite moderate view of the reductionist position: “The reductionist considers ‘upward’ causal and explanatory relationships to be more important than ‘downward’ causation and explanation.” According to his view, only a “strict reductionist… believes that downward explanations are illegitimate, and should therefore be expunged altogether from science.” (2004, 120f)
- 24.
In a different paper Soto and Sonnenschein explain this difference as follows: “In this view, causes act from the bottom-up. Contrary to reductionism, organicism considers both bottom-up and top-down causation.” (2010, 364)
- 25.
Already Wimsatt has recognized the importance of questions like these. He states that “the issue between scientists who are reductionists and holists is not over the in principle possibility of an analysis in lower-level terms but on the complexity and scope of the properties and analyses required. The more holistically inclined scientists usually argue that higher-order relational properties of the lower-level entities are required, and the reductionists argue that a given simple, lower-level model… is adequate.” (1980, 146)
- 26.
For more information about limb development see Gilbert 2006, Chapter 16.
- 27.
Since this sounds very eliminative it should be stressed that Rosenberg seeks to defend a non-eliminative version of explanatory reductionism in biology. However, in Chap. 4, Sect. 1 I have already pointed out that it remains unclear what exactly this non-eliminative character amounts to. On the one hand, Rosenberg claims that reductionism tolerates “terms like cell as acceptable expressions in biological description” and “accepts the reality of cells and their causal roles”. On the other hand, he denies that “there are distinct causal properties of the items such terms name that are not open to identification in molecular terms” (2006, 84). Other statements, too, suggest that Rosenberg assigns only a very minor role to functional biology’s concepts (such as ‘cell’). In his view, they may play an important role for identifying the phenomena to be explained, but they are not part of the explanantia (see Chap. 4, Sect. 1.1.2).
- 28.
One should be aware of the fact that not all reductive explanations in developmental biology must bottom out at the molecular level. For instance, gastrulation can be described as a process in which the whole blastula is involved. Gastrulation of the blastula is explained by appealing to certain regions of the blastula and to the tree germ layers. These explanatory relevant factors are biological parts of the blastula and located on a lower level than it, but they are not molecules.
- 29.
Van Regenmortel explains this as follows: “the specificity of a complex biological activity does not arise from the specificity of the individual molecules that are involved, as these components frequently function in many different processes. […] It is the particular cellular compartment and environment… that allow a gene product to have a unique effect. Biological specificity results from the way in which these components assemble and function together.” (2004a, 1016; see also van Regenmortel 1998)
- 30.
This is compatible with the view that there might exist an additional feature of reductive explanation, according to which at least in some reductive explanations structural properties are regarded as being more important than functional ones. This view can be found, for instance, in Wimsatt’s work (e.g. 2006a, 468) and it is implicitly present in Sarkar’s analysis of reductive explanation in genetics and molecular biology (e.g. 2005, 9f). However, I do not endorse this view because it is not supported by the biological literature.
- 31.
With regard to the latter question see, for instance, Sarkar 2005, 117–143.
- 32.
“[G]ene sequencing and other techniques will soon have isolated all the cell’s individual parts and spelled out their isolated functions. Now, it is time to move beyond reductionism. […] Now we need to know how all these things are integrated.” (Service 1999, 81)
- 33.
I am aware that this way of talking about the one molecular level is exposed to criticism. The already discussed example of photosynthesis shows that the notion of molecules encompasses objects that are of very different kind and size (e.g. complexes of macromolecules, ions, and subatomic particles) and that partly compose each other (e.g. electrons are parts of photosystems). But the fact that the molecular level (in a wide sense) can be characterized in a more fine-grained manner by dividing it into the level of complexes of macromolecules, the level of macromolecules, the level of molecules (in a narrow sense), the level of atoms and so on does not imply that the way of talking about the one molecular level (which can be found frequently in biological practice) is illegitimate.
- 34.
This notion of a reductive explanation in ecology is underpinned by the following statements of ecologists: “reductionism in ecology… see[s] ‘true causes’ arising at only one level […]. … for reductionists, organisms are the only real objects, while the higher levels of organization are the surface of the truly important events…” (Korfiatis and Stamou 1999, 388) And: “Reductionism… sees the individual species, or ultimately the individuals…, as the only ‘real’ objects while higher levels are again descriptions of convenience without causal reality.” (Levins and Lewontin 1980, 51)
- 35.
The version of reductionism that corresponds to reductive explanations of this kind is also known as “methodological individualism” (Sarkar 2009, Sect. 5; Dupré 1993, 107–120). In the social sciences methodological individualism prescribes explaining social phenomena by appealing to the behavior of individuals, ignoring or simplifying influences from the environment and from higher levels as the societal level.
- 36.
The New Mechanists formulates this point as follows: “[T]he components that are accepted as relatively fundamental or taken to be unproblematic for the purposes of a give scientist, research group, or field… [i.e., the level, on which] the explanation comes to an end… is relative. […] [And] what is considered as the bottom out level may change” (Machamer et al. 2000, 13f).
- 37.
- 38.
Wimsatt describes the reductive method that corresponds to this kind of reductive explanation as follows: “Assume that all descriptions and processes are to be referred to entities at a given level” (2006a, 468).
- 39.
The controversial assumption, that the appearance of a particular phenotypic trait of an organism is exclusively determined by its genes, is called genetic determinism. This thesis is nowadays rejected by almost everyone (Byerly 2003).
- 40.
This revolutionist stance also finds expression in the titles of their papers, for example “Somatic Mutation Theory of Carcinogenesis: Why It Should Be Dropped and Replaced” (Sonnenschein and Soto 2000).
- 41.
The stance that underlies SMT can be condensed to the following slogan: “A gene is broken, fix the broken gene and cure disease” (Joyner and Pedersen 2011, 1018).
- 42.
The proponents of SMT more and more take into account also non-genetic factors, such as communication signals between cells and other extra-cellular factors (Hahn and Weinberg 2002; Malaterre 2007, 59). However, they do not regard changes at the tissue level as being important, as proponents of TOFT claim.
- 43.
- 44.
Genetic explanations can also be characterized as single-factor explanations (see Sect. 2.4.3). This is particularly true for genetic explanations that appeal to single genes or single kinds of genes.
- 45.
Muscle fibers originate during development from the fusion of several undifferentiated immature cells, called myoblasts, into long, cylindrical, multi-nucleated cells. Muscle fibers are composed of actin and myosin myofibrils repeated as a sarcomere, the basic functional unit of the muscle fiber. The term ‘muscle’ refers to multiple bundles of muscle fibers held together by connective tissue.
- 46.
One may wonder why this type of reductive explanation is discussed in this section, that is, under the label of the lower-level character of reductive explanations, and not, for instance, in the context of the internal character of reductive explanations (Sect. 3). I admit that one could allocate this subtype differently. But I think that it suits this section well because the single factor is chosen among the lower-level factors.
- 47.
Examples are genetic explanations that refer to the one gene that is taken to cause a particular phenotypic trait. Explanations of this kind are often reported by such statements as: “The gene for x (e.g. aggression, obesity, crib death, intelligence, etc.) has been found.”
- 48.
Bechtel and Richardson put a lot of effort in depicting these “psychological constraints” for developing explanations (see e.g., 2010, 234–243).
- 49.
Exceptions are for instance homeostatic systems, which possess the capacity of self-regulation, i.e., of sustaining the internal conditions under a certain range of variations in the environmental conditions. The most common kind of homeostasis is the regulation of the body temperature that can be found in endothermic animals like birds and mammals (see Reece et al. 2011, 906–914).
- 50.
Consider the following reductionistic heuristics: “(3) Interface determinism: […] black-world perspectivalism – all that matters about the environment is what comes in across system boundaries… […] (5) Modelling localization: look for an intra-systemic mechanism to explain a systemic property rather than an inter-systemic one. […] (6) Contextual simplification:… simplify environment before simplifying system. […] (7) Generalization:… focus on generalizing or elaborating the internal structure, at the cost of ignoring generalizations or elaborations of the environmental structure.” (2006a, 468f)
- 51.
The term ‘intrinsic’ might have the advantage (compared to ‘internal’) that it can also be spelled out in a non-spatial way and thus captures also more rare types of reductive explanations (e.g. structural explanations). However, I stick to the term ‘internal’ because my view of part-whole relations is closely linked to the idea that parts are spatially included in their wholes. Moreover, the term ‘intrinsic’ has the drawback that it confounds the internal character with the third characteristic of reductive explanations, which is introduced in Sect. 4.
- 52.
These “steps” are distinguished for heuristic reasons. In reality, they do not occur in such a neat, clearly successive fashion as suggested here (see e.g. Dobson 2003).
- 53.
This thesis is sometimes referred to as (one reading of) the “linear sequence hypothesis” (Hüttemann and Love 2011, 17), in short “LSH”.
- 54.
Another example of a lower-level explanation that does not also exhibit an internal character is the explanation of how the PI 3-kinase/Akt pathway promotes cell survival (see Sect. 3.3). This explanation also describes how an extracellular survival signal binds to a particular receptor of the cell. This factor is external to the cell, but at the same time it is a lower-level factor since it belongs to the kind of macromolecules, to which also most of the cell’s parts belong.
- 55.
I only found two statements of biologists that are relevant to these questions. But even they provide only limited insights. For instance, Greenspan characterizes reductionism (in molecular genetics) as “the neat view of biological systems made up of dedicated components… and in which particular starting conditions give rise to uniquely predictable responses” (2001, 386; my emphasis). Furthermore, Levins and Lewontin assume that with respect to ecology “reductionism takes the form of regarding each species as a separate element existing in an environment that consists of the physical world and of other species. The interaction of the species and its environment is unidirectional: the species experiences, reacts to, and evolves in response to its environment. The reciprocal phenomenon, the reaction and evolution of the environment in response to the species, is put aside.” (1980, 49)
- 56.
Machamer et al. speak about start or set-up conditions, too (2000, 11). There is, however, an important difference between our positions: they regard these conditions to be components of a certain mechanism, whereas I think that (in most cases) it is more appropriate to interpret them as parts of the context of a biological object Y, not as biological parts of Y itself.
- 57.
RTK is a transmembrane receptor protein. As such it is a part of the cell membrane and, hence, an internal factor.
- 58.
In accordance with my thesis, Wimsatt identifies a certain reductionistic heuristic, which he calls “black-world perspectivalism”. He characterizes this reductionistic strategy as the assumption that “all that matters about the environment is what comes in across system boundaries” and that “the order of study is from a system with its input-output relations to its subsystems with theirs, and so on.” (Wimsatt 2006a, 468)
- 59.
It is not the case that no philosopher has paid attention to this issue so far. For instance, in the chapter on “Mechanisms and its Alternatives” in his book “The Mind and its Place in Nature” (1925) Broad makes the following claim: “It is clear that in no case could the behavior of a whole composed of certain constituents be predicted merely from a knowledge of the properties of these constituents, taken separately” (1925, 63; my emphasis). In his paper on “The Watson-Crick model and Reductionism” (1969) Schaffner addresses the same issue: “given an organism composed out of chemical constituents, the present behavior of that organism is a function of the constituents as they are characterisable in isolation plus the… causal inter-structure of the chemical constituents” (1969, 346; my emphasis).
- 60.
I admit that the unbiased character of my account might be threatened a bit by my strategy to take into account also the way how biologists dispute about reductionism. It might be argued that their notion of reductive explanation is affected by their wish to defend or to criticize reductionism, too. In fact, this may be applicable to some of their statements. However, I think that I managed to identify these cases and to treat them with special care. This will become apparent in the following sections.
- 61.
- 62.
- 63.
- 64.
The add-on ‘dynamically’ is superfluous since any interaction is temporally extended, involves changes and, as such, is dynamic and not static.
- 65.
Wimsatt describes this assumption as the reductionistic strategy to assume that “the results of studies done with parts studied under different… conditions are context-independent, and thus still valid when put together to give an explanation of the behavior of the whole”. He refers to this heuristic as “Articulation-of-Parts (AP) coherence” (2006a, 470).
- 66.
By contrast, whether an object exhibits a relational (or extrinsic) property depends on the object’s relationships to other things.
- 67.
To be exact, interactions are not relational properties themselves, but rather “occasions on which a change in a [frequently relational] property of one part brings about a change in a [frequently relational] property of another part” (Glennan 2002, 344).
- 68.
The forth condition reveals a close connection between the assumption that biological objects are aggregative systems and the assumption that the organization of and interactions between biological parts can be represented as neat, linear causal chains.
- 69.
Philosophers, too, agree with me on this point. For instance, Wimsatt and Sarkar write that in most biological explanations “much of the explanatory weight is borne by the organization of… parts into… the higher-level system”. Hence, they emphasize that “it is not appropriate for interlevel reduction to be tarred with the ontologically corrosive reputation of aggregativity.” (2006, 702)
- 70.
This underestimation of complexity is a part of what Dennett calls “greedy reductionism” (1995, 82).
- 71.
Bechtel and Richardson (2010) place special emphasis on Wimsatt’s last condition of aggregativity since they are primarily concerned with exploring variations in organizational structure.
- 72.
Similarly, Wimsatt admits that “[i]t is rare indeed that all of these conditions are met” (1997, 375).
- 73.
- 74.
Nagel has already emphasized this point: “What is distinctive of such systems [i.e., of organic wholes]…is that their parts do not act, and do not possess characteristics, independently of one another.” (1952, 26)
- 75.
This high degree of integration can be traced back to the fact that the component subsystems have evolved together (Levins 1970, 77).
- 76.
Bechtel and Richardson primarily discuss “connectionist systems” as examples for non-decomposable systems (2010, 199–229).
- 77.
However, I do not insist on this thesis. Biological practice exhibits a sometimes surprising diversity. Perhaps in some biological field there exists a kind of reductive explanation I have overlooked, but which is crucial to that field. Future work on this issue will show whether my list is complete or whether it must be supplemented by other characteristics.
- 78.
Alternatively, it might be that the complex entanglement of system and environment results in a failure of explanation altogether because the system cannot be separated from its environment and, thus, the phenomenon to be explained cannot be identified.
- 79.
- 80.
See for instance Boyle, “The Excellency of Theology” (1966), 69–71, 77f and Descartes, “Principles” (1991 [1644]), Part III, Section 46. This traditional idea can still be found in the statements of some biologists. For instance: “Reductionism seeks to explain the wide variety of natural phenomena on the basis of the behavior of a limited number of simple constituents” (Grizzi and Chiriva-Internati 2006, 5; my emphasis).
- 81.
- 82.
The New Mechanists claim that “set-up conditions”, “enabling conditions”, or even background conditions, which often include external factors, should be regarded as “parts of the mechanism” (Machamer et al. 2000, 11).
- 83.
These relations can but need not be relations of ontological reduction. For instance, relations between levels as well as internal-external relations trace back to constitutional relations between a system and its parts. These can but need not be characterized as instances of ontological reduction (recall Chap. 3, Sect. 2.1.1).
- 84.
This, however, does not imply that the world completely determines the reductivity of an explanation. As I point out in Kaiser (forthcoming a), pragmatic factors might have an influence on how the line between a biological object or system and its environment is drawn and on which objects and processes are identified as biological parts. In this way pragmatic factors might also affect the reductive character of an explanation.
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Kaiser, M.I. (2015). The Ontic Account of Explanatory Reduction. In: Reductive Explanation in the Biological Sciences. History, Philosophy and Theory of the Life Sciences. Springer, Cham. https://doi.org/10.1007/978-3-319-25310-7_6
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