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In the early days of molecular biology, it was an evangelical movement. Most people were against us. Most of the biochemists didn’t understand the nature of the problems that we thought were interesting and important. They had a completely different set of attitudes. SYDNEY BRENNER, 1988 There is not the slightest evidence that the findings of epigenetics will dispel the main ideas of neo-Darwinism. JERRY COYNE, 2011 I now groan audibly when a journalist (usually from continental Europe, where they spend too much time learning philosophy rather than science) asks me the now inevitable “what about epigenetics?” question. . . . I am heartily sick of the “epigenetics ” bandwagon and almost look forward to the next one, whatever it turns out to be. RICHARD DAWKINS, 2011 TWO CHALLENGES OF EPIGENETICS IN LIGHT OF THE EXTENDED EVOLUTIONARY SYNTHESIS 42 ◂ CHALLENGES OF EPIGENETICS NOT EVERYONE HAS WELCOMED the recent rise of epigenetics. In fact, some evolutionary biologists doubt whether epigeneticists should have a say in evolutionary biology. This reluctance may be due in part to sensational reporting of new epigenetic research results and simplistic public debates about “Lamarck’s return” and “Darwin’s death.” Besides these issues, however, there are more serious challenges currently hindering theoretical integration of epigenetics. In this chapter, I discuss two kinds of tensions that arise due to the explananda and explanantia chosen by epigeneticists and due to the methodological strategies chosen to investigate the relationships between these explanantia and explananda. First, I give an overview on some challenges the developmentally oriented view of epigenetics poses for the field of evolutionary biology or, more specifically, for orthodox neo-Darwinian explanation. This includes the issues of whether (and how) molecular epigenetic explanations that are based on highly artificial experimental setups and that focus on developmental proximate causes rather than on ultimate causes can address evolutionary explananda. The latter problem in particular becomes apparent in the recent debate about a so-called “extended evolutionary synthesis.” Second, in addition to the tensions between epigenetics and other disciplines —above all evolutionary biology—we also find intradisciplinary tensions concerning how to understand the field of epigenetics conceptually or methodologically, as well as how (and what) epigeneticists should explain. These latter issues display epigenetics’ status as a young and developing research field. Its inner tension and heterogeneity are reconstructed by means of a historical analysis of approaches using Waddington’s epigenetic landscape images as a methodological tool to mathematically model the role of epigenetic determinants in cell differentiation and reprogramming. TOWARD AN EXTENDED EVOLUTIONARY SYNTHESIS Since the 1990s, an increasing number of epigenetic factors and processes have been revealed. Due to their importance for research in both development and heredity they have often been juxtaposed with the somewhat antiquated concept of inheritance of acquired characteristics. As described in the previous chapter, this Lamarckian claim has recently been accom- ▸ 43 CHALLENGES OF EPIGENETICS panied by a call for an “extended evolutionary synthesis” or even a “postDarwinian synthesis” that includes not only epigenetics (West-Eberhard 2003; Kutschera and Niklas 2004; Jablonka and Lamb 2005, 2007, 2010; Jablonka 2006; Pigliucci 2007, 2009; Cabej 2008; Gissis and Jablonka 2011; Mesoudi et al. 2013; Laland et al. 2014, 2015) but also related fields such as evo-devo (Johnson and Porter 2001; Müller 2007; S. Gilbert and Epel 2009; Kirschner and Gerhart 2010; Laubichler 2010; Müller 2010; Newman 2010; Laland et al. 2014, 2015) and niche construction theory (Odling-Smee et al. 2003; Laland et al 2009, 2014, 2015; Danchin et al. 2011; Piersma and van Gils 2011; Mesoudi et al. 2013). 1 This novel extended theory is thought to be based on a “development first view” on evolution, which emphasizes the evolutionary significance of developmental responsiveness. In addition, it should help to overcome gene-centrism and relegate genes to the role of “followers” in evolutionary processes (see, e.g., Pigliucci and Müller 2010a, 14). The leading role is now allocated to developmental interactions, their inheritance, and the mobilization of phenotypic traits through the plastic responses of environmentally sensitive and adaptive developmental systems. Accordingly, genes are mainly responsible for routinization and progressive fixation of traits in evolution. The various developmentally oriented approaches calling for an expansion of evolutionary theory have been challenged in at least three ways. First, particularly orthodox-minded population geneticists have doubted whether novel concepts, like evolvability and robustness in evo-devo, actually offer anything substantially new compared to the conceptual framework of the modern synthesis (Lynch 2007; Coyne quoted in both Pennisi 2008 and...


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