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Man’s mind cannot grasp the causes of events in their completeness , but the desire to find those causes is implanted in man’s soul. LEO TOLSTOY, 1868–69 Manipulation of the variables in a model, or simulation, is intended to provide answers to the critical question, “What would happen if . . .” CONRAD HAL WADDINGTON, 1977 SO FAR WE HAVE seen that epigenetics will have to travel a rocky road to be integrated into mainstream biology. The difficulties on this path, however , exist not just for historical reasons, such as the methodological traditions addressed in the previous chapter. As this chapter and the next show, a more detailed philosophical investigation brings to light a number of additional features in epigenetics that set it apart from the way many scientists understand molecular and cell biology. These features and their associated challenges refer to the concept of scientific explanation. THREE CAUSAL EXPLANATION ▸ 91 CAUSAL EXPLANATION Since the mid-1990s, the nature of biological explanation has been the focus of various philosophical analyses. In particular, philosophical accounts of explanation have turned out to be inadequate for explaining biological phenomena, and the existence and explanatory role of laws in biology have been questioned. As a reaction to this period of uncertainty, a very influential position has been developed. This position, adopted by many philosophers of science, draws heavily on James Woodward’s interventionist account of explanation (Woodward 1997, 2003, 2004, 2007a; Woodward and Hitchcock 2003). It serves as a blueprint for more detailed accounts of both causal explanation and mechanistic explanation in biology (see, e.g., Waters 2007; and Craver 2007). While the former kind of explanation relates biological events, the latter relates causal properties of biological entities, usually across different levels of organization. The former traces diachronous causal relations and the latter, synchronous constitutive relations. This chapter focuses on causal explanation in epigenetics before turning to constitutive explanation. I show here that the interventionist account elucidates and justifies the way that epigeneticists in molecular biology infer causality in complex living systems. Therefore, experimental work on epigenetic inheritance phenomena , as well as the comprehensive methodological framework that rests upon manipulation of inducing environmental variables and multifactorial experimentation, is reviewed. In addition, I discuss whether the explanatory practices of nonmanipulative field studies in evolutionary epigenetics , described in the previous chapter, fit the interventionist schema. Then, against the popular belief that interventionist explanation is in urgent need of supplementary information gathered from lower levels (so-called “mechanistic detail”) that addresses why a certain causal dependency relation exists, I argue that tracing interventionist causality forms an exhaustive explanatory approach in biology. This argument has two components. First, more fundamental factors such as genes do not have a unique ontic or epistemic status, and this means that it is not necessary to list them as a mechanistic detail in every explanans addressing biological complexity. In other words, genes are considered to be causal factors that often have a mere potential, non-actual difference-making effect on phenotypic traits. Second, the populations in which higher-level nongenetic factors are identified as actual difference-making causes are distinct from the actual populations in which genes make a difference. These arguments imply the autonomy of 92 ◂ CAUSAL EXPLANATION higher-level epigenetic explanation, which requires little or no mechanistic detail in molecular and cell biology. 1 THE INTERVENTIONIST ACCOUNT OF CAUSAL EXPLANATION The idea that causal relationships are somehow exploitable for purposes of control and manipulation has intuitive appeal—especially, but not exclusively , to biologists who are experimentally acting upon phenomenato -be-explained. Francis Bacon ([1620] 1994) famously advocated this idea in his Novum Organon. According to Bacon, we can only uncover and explain the hidden causal structures of nature by interacting with them. In other words, we have to manipulate putative causes in order to reveal them. This view of causal explanation was subsequently developed further in the twentieth century by philosophers as well as scientists (Collingwood 1940; von Wright 1971; P. Holland 1986; Menzies and Price 1993; Freedman 1997; Pearl 2000). It is commonly called the manipulationist or interventionist account of causal explanation. The discussion of causal explanation in epigenetics presented below focuses mainly on James Woodward’s contribution to the interventionist account. In contrast to the other notions of manipulation, Woodward refers to a special kind of manipulation relevant to causal explanation, which he calls interventions. Interventions are not based on any kind of human agency, which is often thought to be necessary for them to be carried out. 2...


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