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ONE HOW EPIGENETICS DEALS WITH BIOLOGICAL COMPLEXITY One can best feel in dealing with living things how primitive physics still is. ALBERT EINSTEIN, 1947 The word explanation occurs so continuously, and has so important a place in philosophy, that a little time spent in fixing the meaning of it will be profitably employed. JOHN STUART MILL, 1843 EXPLAINING COMPLEX LIVING SYSTEMS UNTIL RELATIVELY RECENTLY, JOHN Stuart Mill’s call for dealing with scientific explanation was addressed by philosophers of science as well as scientists in a rather monistic way. Many were convinced that a phenomenon is explained only if some law covers it as a special case. Accordingly, philosophers usually focused on phenomena in the exact sciences or held 14 ◂ HOW EPIGENETICS DEALS WITH BIOLOGICAL COMPLEXITY the view that all scientific explanations can be reduced to a small number of laws or theories at a physical level. This view of scientific explanation has imposed an overstated criterion of how certain explanations do, in fact, explain, which has led some to describe biologists’ scientific program as a technological application of basic laws of nature, similar to that of, for example, engineering (Smart 1959). However, due in no small part to the increasing success of molecular biologists in “decoding” the genome and the importance of their findings for humans, as well as the emergence of the field of philosophy of biology in the 1960s to 1970s, this view has gradually changed. 1 Two consequences have to be considered in this context. First, both the applicability of the so-called covering-law account of explanation in biology and the existence of biological laws have been called into question (see chap. 3). Second, new accounts of scientific explanations that fit the unique explanatory challenges in biology and the special sciences more generally have been (re)developed. In philosophy of biology, these new explanatory accounts draw on a long tradition of conceptualizing relationships in and between living systems (between structures of various size, from DNA to organisms, and of different strength), as well as analyzing the role with which they figure into explanation. For example, by revising the Aristotelian concept of teleology (Aristotle 1984, 195a23–24, 198b8–9), Ernst Mayr (1961) developed an idea of final cause that he considered to be explanatorily relevant exclusively for evolutionary biologists. In addition, it has been debated how the seemingly goal-directedness of living beings can be addressed properly by functional explanations (Lehman 1965; Wimsatt 1972). More recently, philosophers of science have focused on clarifying the nature of causal dependency relations at higher levels of organization. These dependencies are usually cited as explanantia in the special sciences in general and in biology in particular. Causal relations have been grounded in complex regularities, as argued by John Leslie Mackie (1974), or in counterfactual dependencies described in the form “if event c had not occurred, event e would not have occurred” (Lewis 1973, 2000). In addition, they have been also understood as relations of probabilistic relevance (Pearl 2000; Williamson 2009) and as relations involving manipulability (Woodward 2003). The latter of these has been highly influential in recent philosophy of science. This approach is exemplified by the so-called interventionist account of James Woodward, which integrates counterfactual and probabi- ▸ 15 HOW EPIGENETICS DEALS WITH BIOLOGICAL COMPLEXITY listic theories with the view that causes are (probabilistically) dependent on an intervention and are thus exploitable for purposes of control. In particular , interventionism has been invoked in philosophy of biology to specify causal relations and how they figure in causal explanation in a variety of research fields, ranging from molecular genetics to ecology (see chap. 3). Moreover, this theory has been used extensively to make sense of how biological mechanisms constitute a phenomenon and how mechanistic explanation functions in biology (see chap. 4). These more recent accounts of causal and mechanistic explanation have been in large part developed to address problems that arise due to the complexity of living systems investigated in biology. For example, models of these systems often include events with low probabilities. This means that there are dependencies in the living world—between genes, cells, organisms , and their environment—that only hold occasionally or even rarely. Are these dependencies causal? In order to answer such a question, we first have to understand what constitutes a living system’s complexity. Complexity has been defined in various ways (see S. Lloyd 2001; Emmeche 1997; Holland 1998, 225–31; Mitchell 2003, 4–7; and Northrop 2010, 2–4). With respect to the issue at...


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