A Brief History of Epigenetics: C.H. Waddington


by Shea Robison (@EpigeneticsGuy)

Read updated posts on Waddington here, with additional details, as excerpts from my forthcoming book :

C.H. Waddington: Genesis of the Original Epigeneticist

More About Waddington: Socialism, Science, and Epigenetics

(See also A Brief History of Epigenetics: Jean-Baptiste Lamarck)

In the history of contemporary Western evolutionary theory, the first use of the term epigenetics is generally attributed to Conrad Waddington in an article published in 1942. In this article Waddington used epigenetics as the name for the study of the causal mechanisms through which genes bring about their phenotypic effects and which necessarily involve adaptive interaction with the environment—although it bears mentioning that the ‘epigenetics’ of Waddington was rooted exclusively in embryology and development, and is not quite the epigenetics of today with its emphasis on non-genetic inheritance, but is better conceived of as “developmental genetics” (Gilbert 2012). Years before this 1942 article Waddington had proposed the appearance of particular organs as the product of the interactions of the genotype and what he called the “epigenotype” with the external environment (Waddington 1939). He then subsequently developed this idea of an epigenotype into the more general notion of epigenetics.

For Waddington, the postulation of the epigenotype as a distinct biological layer and the recognition of epigenetics as distinct processes in biological development were necessary steps for the progress of genetics. According to Waddington himself, Waddington’s original project in promoting the explicit recognition of this biological layer between the genome and the environment was not to challenge but rather to extend the conventional understanding of genetics through a more sophisticated approach which bridges the gap between the genotype and the developing phenotype via the epigenome (Waddington 1940; See also Gilbert 2012; Jablonka & Lamb 2012). Important evidence which could contribute to a more complete understanding of biological development was being overlooked because, per Waddington, the nexus of the epigenotype between the genome and the environment was being so maligned by the prevailing gene-environment dichotomy.

Science and ideology

However, at this point, it may be useful to mention something about Waddington’s politics. Although it would probably be imprecise to label Waddington a Marxist, per se, it is clear that many of his closest associates were unabashed Marxists, and that if Waddington himself was not a card-carrying Marxist he had strong ideological tendencies in that direction (Peterson 2010). Waddington’s ideological inclinations and his openness to a more holistic and emergentist biology were not unique. Val Dusek, in his depiction of the emergence of the anti-mechanist and anti-reductionist biology in the late 19th and early 20th century, identifies many of the prominent scientists involved in research in this area—such as J.B.S. Haldane, a prominent figure in the mathematical theory of population genetics, and J.D. Bernal, a pioneer in X-ray crystallography—as “self-proclaimed Marxists,” and discusses a couple of the ways that their ideological inclinations were manifest in their scientific work (1999, 21-22). Swann and Aprahamian also detail a number of ways in which the dialectical materialism of Marx and Engels fit the assumptions of the experimental work of these scientists including Waddington (1999, xvi-xix). These connections between politics and science were not lost on the scientists themselves, as Waddington observed that “a scientist’s metaphysical beliefs are not mere epiphenomena, but have a definite and ascertainable influence on the work he produces” (2009, 72).

Bearing this background in mind, Waddington highlighted the philosophical roots of the dogmatic limitations of the Modern Synthesis sixty years ago, and discussed the implications of the challenges introduced by epigenetics to these basic ontological commitments. In particular, Waddington described the insuperable wall between genes and their environment as evidence of the “exaggerated atomism” of modern genetics which is the “gravest defect” not just of modern genetics but of modern science as a whole (1953, 188). This assumption of atomism in modern genetics is also reflective of similar assumptions of the atomistic individual which is a central pillar of modern liberalism, as expounded most notably by John Locke and Thomas Hobbes (Barbour 2006; Den Uyl & Rasmussen 2006; Hurtgen 2002; Taylor 1985).

The problem with such atomisms and their resulting dualisms—in this case the alleged isolation of genes from their environments—is that they are at best exaggerated and oversimplified, as demonstrated by the recent recognition of the importance of epigenetics in evolution and biological development. As writes Waddington, this logic of dichotomization commits modern evolutionary theorists and geneticists to the idea that:

All living things, man included, had been brought into being by the collocation of two entirely independent factors: on the one hand the occurrence of mutations whose nature was totally unconnected with any ambient circumstances, and on the other hand a sieving process in which the environment merely selected from organisms which were offered to it ready made as units of being…each [factor] having its character in its own right, which come together with as little essential inter-relation as a sieve and a shovelful of pebbles thrown on to it. (1953)

According to Waddington, this dichotomization of environment and organism which is so fundamental to modern genetics and the Modern Synthesis constitutes a breach with reality “as complete as the Cartesian dualism of mind and matter” (1953).

Mind, matter and genes

The Cartesian dualism of mind as substantially distinct from the matter it observes and manipulates is the ontological and epistemological basis of modern science—manifest primarily as the subject-object distinction which guides modern scientific investigation. As described in encyclopedic detail by P.F.M. Fontaine (1986), some form of dualism has been the implicit if not explicit assumption of most philosophies and sciences and religions throughout Western history going back to Plato and Aristotle. The philosopher Alfred Whitehead locates the pervasiveness of this “facile vice of bifurcation” as the inevitable result of the commonsense—but incorrect—perception of objects ‘out there’ as obviously distinct from the entity ‘in here’ that is perceiving those objects ([1920] 1964).

Because “organism and environment are not two separate things,” Waddington proposed his integrative and more interactive epigenetic theory of biological development as a means of “healing” this unwarranted and ultimately unscientific separation of ourselves from our environments (1953). Likewise, a main goal of my project is to challenge and to unravel this fundamental dichotomization as a means to identify and potentially reconcile the politically relevant narratives of genetics and epigenetics, to begin to discuss how epigenetics could make its impact on public policy.

Waddington was not alone in seeing the opportunities from the integration of genetics with epigenetics. He and others had been able to marshal significant empirical evidence in demonstration of the role of the epigenotype in phenotypic plasticity (Kirpichnikov 1947; Snyder 1950) and even in non-genetic inheritance (Ephrussi 1958; Mitchell & Mitchell 1952; Russell 1942; Waddington 1942) in the 1940s and 1950s. Regardless of this work, though, epigenetics was in Waddington’s words still “so completely rejected by the rest of the scientific world that it is hardly considered to be worthy of discussion” (1953), an attitude as shown before which was carried on well into the 1990s (Haig 2012).

This almost blanket rejection of the evidence in favor of epigenetics in the development of the Modern Synthesis was for Waddington not the inevitable and justifiable outcome of scientific progress. Rather this rejection was proof of the “extremist” nature of the Modern Synthesists and their practically exclusive focus on genetic adaptation to the neglect of “the doctrines emerging from other fields of modern biology” which could be combined with genetics to produce significantly different (and ostensibly more comprehensive) conclusions (1953). The political and ethical underpinnings of both this rejection of epigenetics and of epigenetics itself in the mid-20th century will be discussed in more detail in other posts. Suffice to say, this antagonistic attitude towards epigenetics delayed for over sixty years the significant advances being reported almost daily from research in epigenetics across a wide swath of domains, the human costs of which are incalculable.

I am curious to hear your thoughts about this historical connection between science qua epigenetics and politics. Leave your comments below and I will respond. For discussion of the political and scientific antecedents of Waddington, see A Brief History of Epigenetics: Jean-Baptiste Lamarck and The History of Epigenetics and the Science of Social Progress. For further discussion of the politics and science of epigenetics around the time of Waddington, read Epigenetics and the Dustbin of History and Epigenetics and the geopolitical history of the 20th century.

Also, if you find these thoughts I’ve shared interesting and worthwhile, Like this post, Reblog it, or Tweet about it using the buttons below.

Barbour, R. (2005). Bacon, Atomism, and Imposture: The True and the Useful in History, Myth, and Theory. Francis Bacon and the Refiguring of Early Modern Thought: Essays to Commemorate the Advancement of Learning (1605–2005), 17-44.

Den Uyl, Douglas J. and Douglas B. Rasmussen. 2006. The Myth of Atomism. The Review of Metaphysics. 59(4), 841-868.

Dusek, V. (1999). The holistic inspirations of physics: The underground history of electromagnetic theory. Rutgers University Press.

Ephrussi B (1958). The cytoplasm and somatic cell variation. J Cell Comp Physiol 52 Suppl 1:35-53.

Fontaine, P. F. M. (1986). The Light and the Dark: A Cultural History of Dualism. Vol. 1-21. JC Geiben.

Gilbert, S. F. (2012). Commentary: ‘The Epigenotype’ by C.H. Waddington. International Journal of Epidemiology, 41(1), 20-23.

Haig D. (2012). The epidemiology of epigenetics. International of Journal of Epidemiology 41:13–16.

Hurtgen, J. R. (2002). The divided mind of American liberalism. Lexington Books. Taylor, Charles. 1985. “Atomism,” in Philosophy and the Human Sciences: Philosophical Papers 2. Cambridge: Cambridge University Press.

Jablonka, Eva and Marion Lamb. 2010. “Transgenerational Epigenetic Inheritance.” In Evolution, the Extended Synthesis, eds. Massimo Pigliucci and Gerd Müller. Cambridge, Mass.: MIT Press, 150-152.

Mitchell M.B., Mitchell H.K. (1952). A case of “maternal” inheritance in Neurospora crassa. Proc. Natl. Acad. Sci. U.S.A. 38 (5): 442–9.

Peterson, Erik L. 2010. “Finding Mind, Form, Organism, and Person in a Reductionist Age: The Challenge of Gregory Bateson and C. H. Waddington to Biological and Anthropological Orthodoxy, 1924–1980.” Ph.D. diss. University of Notre Dame.

Russell, E. (1942). The Inheritance of Tumors in Drosophila Melano-Gaster, with Special Reference to an Isogenic Strain of St Sr Tumor 36AL. Genetics 27(622).

Swann, B., & Aprahamian, F. (Eds.). (1999). JD Bernal: a life in science and politics. Verso.

Taylor, Charles. 1985. “Atomism,” in Philosophy and the Human Sciences: Philosophical Papers 2. Cambridge: Cambridge University Press.

Waddington C.H. (1939). An introduction to modern genetics. Macmillan, New York.

Waddington C.H. (1940). Organisers and Genes. Cambridge: Cambridge University Press

Waddington C.H. (1942). The epigenotype. Endeavour 1: 18.

Waddington, C. H. (1942). Canalization of Development and the Inheritance of Acquired Characters. Nature 150 (3811), 563-565.

Waddington C.H. (1953). The strategy of the genes. London: George Allen & Unwin, 188.

Waddington, C. H. (Ed.). (2009). Sketching Theoretical Biology: Toward a Theoretical Biology(Vol. 2). Transaction Publishers.

Whitehead, Alfred North. [1920] 1964. The concept of nature. Cambridge: Cambridge University Press, 188.

3 thoughts on “A Brief History of Epigenetics: C.H. Waddington

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