Genetics and Epigenetics Come Home from the War

Excerpt from my forthcoming book Epigenetics and Public Policy The Tangled Web of Science and Politics to be released February 2018 by Praeger

As discussed before, prior to World War II there were substantial geographical differences in the approach to and the understanding of genetics: There was the more European emphasis on embryology and on the processes of biological development which focused on the environment of the genes, in contrast to the emerging American focus on the genes as ultimately controlling development and heredity. These were the two main currents of biology that Waddington, with his feet planted firmly in both streams, attempted to combine through his postulation of the epigenotype and epigenetics. Eventually, though, the American-led emphasis on molecular genetics carried the day to become the sine qua non of biology in the West, relegating embryology and development to secondary status.

This outcome is usually portrayed as the product of the inevitable and impartial progress of science, but the material effects of the Second World War and its aftermath on the particular trajectory of the science of genetics are rarely considered. When these factors are taken into account, the inevitability or incontestability of this increasingly reductive focus of genetic research—including the decades-long exclusion of epigenetic mechanisms—are brought into question.

Other voices, other rooms

For example, the development of the science of genetics in France after World War II initially followed a substantially different path than in America.[1] Although the French genetics research program eventually merged into the international mainstream of molecular genetics by the mid-1960s—in part for the reasons to be discussed in a subsequent section—this initial development of a distinct and yet still fruitful focus provides a counter-balance to the conventional story of the inevitability of the gene-centric focus of molecular genetics as we now know it.

Genetics in France, as lead by the Russo-French geneticist Boris Ephrussi, was much more focused on the combination of embryology and genetics. Ephrussi, who was appointed to the first chair of genetics in France at the University of Paris after WW II, had, like Waddington, been initially trained in embryology but had also studied genetics in America under T.H. Morgan. After the war, Ephrussi discovered the non-Mendelian inheritance of deep physiological changes in cells, and other evidence of the significance of the cytoplasm in heredity.[2] Given these empirical results, Ephrussi pushed for the cytoplasm to be a focus of French genetics specifically against the American preoccupation with the nuclear gene.[3] Ephrussi  famously expressed his dissatisfaction with the distinctly Americanized position that heredity was exclusively controlled by the genes writing that “we cannot determine the truth of a hypothesis by counting the number of people who believe it.”[4]

Regardless of the efforts of Ephrussi and others to maintain a distinct concentration for genetics research in France, they were ultimately unable to resist the rising wave of the focus on molecular genetics emanating from America—coincident with the solidification of U.S. geopolitical hegemony.

Follow the money

In this context, especially given the dramatic ascension to superpower status of the United States following World War II, the distribution of the funding for scientific research in the postwar bipolar world constitutes a significant and often overlooked factor in the development of genetics and the virtual exclusion of epigenetics.

Until the first World War, most scientific projects around the world were funded by wealthy patrons, private benefactors, or industry, with only modest support from governments. Government support for science increased somewhat through the 1800s but never constituted much of an influence. After the first World War, government funding of science increased but was still not a significant amount, and even private sources of funding support for science had dwindled (for example, in 1931, total grants from American foundations amounted to just over $50 million, by 1934 it was $34 million, and by 1940 it was only around $40 million [5]).

With World War II, though, all of this changed dramatically—especially in the United States. Vannevar Bush, an engineer and vice-president at MIT, with a one page proposal and a fifteen minute meeting with President Roosevelt in June of 1940, was able to secure the funding for the National Defense Research Committee (NDRC) to coordinate scientific research on “the problems underlying the development, production, and use of mechanisms and devices of warfare.”[6] The NDRC was then superseded a year later by the Office of Scientific Research and Development (OSRD), also overseen by Bush, which by 1946 was allocated in excess of $536 million from the Emergency Fund of the President for projects of all kinds, including the Manhattan project to develop the first atomic weapons.[7]

In the waning days of WW II, Bush submitted a report to President Roosevelt called “Science: The Endless Frontier” in which Bush proposed the continued funding of science by the government at wartime levels, but without the shackles of military utility.[8] In particular, Bush identified what he called “basic research,” or “research in the purest realms of science” without concern for direct application, as the proper focus of the government funding of science. “Scientific progress on a broad front,” Bush wrote, “results from the free play of free intellects, working on subjects of their own choice, in the manner dictated by their curiosity for exploration of the unknown.”[9] However, Bush also explicitly painted this scientific superiority in the light of maintaining national security, as the best defense against aggression. Congress eventually agreed with Bush, and created the National Science Foundation according to his recommendation.

As detailed extensively by Daniel Greenberg, Bush’s insistence on federal patronage for the definition and advancement of scientific knowledge in the United States was a dramatic departure from previous funding practices, which eventually came to characterize federal science policy after World War II—bringing with it substantial political and ethical concerns.[10] The scale of the government funding of science only escalated after the onset of the Cold War, quickly becoming the new norm as universities competed for this funding to fuel “the steepest expansion of higher education in American history (if not the whole world).”[11] For example, by 1953 the federal funding in the U.S. for ‘basic’ research alone was over $256 million, and federal research contracts constituted more than 90% of the annual operating budget of MIT.

The sheer magnitude of the funding available for science in the U.S. at this historical moment after World War II and at the beginning of the Cold War is especially extreme when compared with the situation in Europe where, for example, as part of the Marshall Plan the U.S. was in the process of spending $12 billion ($120 billion in current value) to rebuild the infrastructures and economies of Europe. In other words, at this crucial historical moment in the development of the science of genetics, substantial financial resources for scientific work were readily available to those involved in promoting a distinctly molecular and atomistic focus for genetics. In contrast, those who were developing alternatives to this molecular focus in Europe not only had to conduct their work within demolished infrastructures being rebuilt with substantial material support from the U.S., but also had to appeal to external sources primarily from the U.S.—where molecular genetics was the emerging consensus—for much of the funding for their scientific work.[12]

The road not taken

Again, in the context of the development of the orthodox science of genetics, which practically excluded epigenetics for so long, the question of who had access to money and resources for research and who did not is very much a live issue. While the gene-centric focus of molecular genetics is now often perceived as the obvious and inevitable victor over other potential alternatives, these financial factors, combined with the geographical, political, and ideological factors discussed before, instead describe a drastically lopsided playing field.

All this is not to say that mainstream genetics is therefore invalid, to be replaced by epigenetics (if anything, I hope this history has demonstrated just how inseparable are genetics and contemporary epigenetics). Rather, this is to suggest that the ascendance of the molecular emphasis of genetics that developed from out of this historical moment—including the decades-long omission of epigenetics—was contingent on many other factors beyond purely scientific considerations which influenced the science and the research of this time. Had it not been for this particular convergence of factors, the science of genetics which resulted after World War II may have been significantly different, even potentially incorporating epigenetic mechanisms into its basic theoretical frameworks sixty years or more before the recent explosion of interest in epigenetics. If epigenetics had been incorporated into the orthodoxy of genetics at this earlier time, as it very well could have been given other circumstances, then not only would it not be as controversial as it is now, but we would also already be sixty years beyond the advances in our understanding of gene function which we are just now gaining from the recent work being done in epigenetics.

 

[1] Burian, R. M., Gayon, J., & Zallen, D. (1988). The singular fate of genetics in the history of French biology, 1900–1940. Journal of the History of Biology21(3), 357-402.

Burian, R. M., & Gayon, J. (1999). The French school of genetics: From physiological and population genetics to regulatory molecular genetics. Annual Review of Genetics33(1), 313-349.

Gayon, J., & Burian, R. M. (2004). Timeline: National traditions and the emergence of genetics: the French example. Nature reviews. Genetics5(2), 150.

[2] Ephrussi B. (1953). Nucleo-cytoplasmic relations in micro-organisms: their bearing on cell heredity and differentiation. Oxford.

[3] Sapp, Jan. (1986). Inside the Cell: Genetic Methodology and the Case of the Cytoplasm. In The politics and rhetoric of scientific method: Historical studies (Vol. 4), Schuster, J. and Yeo, R.R. eds. Springer Science & Business Media.

[4] Ephrussi (1953); This line by Ephrussi was actually a paraphrase of an earlier comment by the philosopher of science J. H. Woodger—a close friend of Waddington and also a member of the Theoretical Biology Club at Cambridge—who wrote elsewhere that “Admittedly, some hypotheses have become so well established that no one doubts them. But this does not mean that they are known to be true. We cannot determine the truth of a hypothesis by counting the number of people who believe it, and a hypothesis does not cease to be a hypothesis when a lot of people believe it.” [Woodger, J. H. (1948). “Observations on the present state of embryology”. Symposium of the Society for Experimental Biology. 2 (Growth in Relation to Differentiation and Morphogenesis].

[5] Neal, H.A., Smith, T.L. and McCormick, J.B., 2008. Beyond Sputnik: US science policy in the 21st century. Ann Arbor, Michigan: University of Michigan Press.

[6] James Phinney Baxter III, Scientists Against Time (Boston: Little, Brown & Co., 1946), p. 14; draft of order attached to undated, unsigned memorandum in OSRD Box 212.

[7] Stewart, Irvin (1948). Organizing Scientific Research for War: The Administrative History of the Office of Scientific Research and Development. Boston: Little, Brown and Company

[8] Bush, V., 1945. Science: The Endless Frontier: a Report to the President on a Program for Postwar Scientific Research, July 1945. United States Government Printing Office.

[9] Bush (1945).

[10] Greenberg, Daniel S. (2001). Science, Money, and Politics: Political Triumph and Ethical Erosion. Chicago: University of Chicago Press.

[11] Kaiser, David. (2011). The Search for Clean Cash. Nature 472 (7341), pp. 30–31.

[12] Strasser, B. (2003). The transformation of the biological sciences in post‐war Europe. EMBO reports4(6), 540-543.

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More About Waddington: Socialism, Science, and Epigenetics

Excerpt from my forthcoming book Epigenetics and Public Policy The Tangled Web of Science and Politics to be released February 2018 by Praeger

The connections between the deep-seeded philosophical inclinations of C.H. Waddington and his eventual ‘discovery’ of epigenetics have been detailed elsewhere. Given these connections between his philosophy and his scientific work, it should be little surprise that there are similar connections between his politics and his science. These connections not only provide additional context for his scientific work, they also help to explain the icy reception of epigenetics when he first proposed it in the 1940s, given the nature of Waddington’s political inclinations and the geopolitical circumstances of the time. These connections in turn help to explain why epigenetics has only recently emerged within the last couple of decades, sixty years since Waddington initially proposed epigenetics as an intermediary layer between genes and the environment.

Science and socialism

Gary Werskey, in his extensive work on the “scientific socialists” of the 1930s,[1] describes the intertwined lives and careers of five prominent British scientists—J.D. Bernal, J.B.S. Haldane, Lancelot Hogben, Hyman Levy, and Joseph Needham—who openly professed both a socialist politics and a socialist conception of science, and four of whom were formal members of the Communist Party of Great Britain. Although Waddington is not one of Werskey’s subjects, he worked with Haldane and published a paper with him. He was close lifelong friends with Bernal, a pioneer in X-ray crystallography which played such an important role in the discovery of the double helix of DNA, and Needham, a specialist in embryology and morphogenesis as well as a respected sinologist, through an informal ‘club’ the three had founded while they were in school together in Cambridge in 1931.

Werskey concludes that the politics of these men did not directly influence their science, primarily because they worked in the mainstream of the science of their time. I suggest instead that while these men did make substantial contributions to the mainstream science of their time, there were also identifiable influences of their politics on their scientific work, and that these political influences are found in their common approach to biology. This convergence of politics, ideology, and biology—especially given how the geopolitical history of the world was to soon unfold—is also pertinent to the development of Waddington’s conception of epigenetics in the 1940s, and to the lack of acceptance of epigenetics until relatively recent. In the same way, this convergence of biology and ideology is equally pertinent to the development of the science of genetics as we now know it, which until the last decade or so more or less excluded epigenetics from serious consideration.

Waddington and the Theoretical Biology Club

This ‘club’ that Bernal, Needham, and Waddington (along with the philosopher of biology Joseph Woodger and the mathematician and biochemical theorist Dorothy Wrinch) formed while at Cambridge was called the Theoretical Biology Club. As the name denotes, the club was organized around discussions of both the philosophy and the science of biology that was just then emerging at that time. In particular, the primary theme or topic of this club was the discussion of the concept of organicism in biology. Organicism, which is related to the philosophy of organism of Whitehead described before, is the idea that wholes are greater than the sum of their parts, that “the properties of each part are dependent upon the context of the part within the whole in which they operate,” and that wholes exert some regulative control over their parts.[2]

Coincidentally or not, organicism applied to biology was championed by J.B.S. Haldane’s father, J.S. Haldane, an internationally respected physiologist also working at Cambridge at this time.[3] As a working natural scientist, the elder Haldane conceived of organicism through his work on the regulative processes of the body, and respiration in particular, in response to changes in the environment.[4] Haldane the father also saw organicism in biology as a much-needed middle way between the unscientific vitalism of Lamarckism and the overly reductionist and dualistic conception of biology which was then emerging in concert with genetics.[5] At the same time, as historian Peter Bowler observes, the organicism of the elder Haldane also coincided with his support for a social philosophy which advocated a significant role for the state in coordinating the actions of individuals to secure the greater good, in contrast to the prevailing liberal philosophy of extreme individualism which Haldane saw as leading to selfishness and expressive of the more avaricious aspects of humankind.[6]

Likewise, in addition to organicism in biology, another major topic of conversation of the Theoretical Biology Club was their shared socialist and Marxist beliefs, which they saw as inextricably linked with their views of biology. These ideological beliefs were not just sophomoric exuberances, though, but were deeply held sentiments which were maintained by all into their subsequent work as well-regarded scientists. For example, Brenda Swann and Francis Aprahamian detail a number of ways in which the dialectical materialism of Marx and Engels fit the assumptions of the experimental work of these men as mature scientists, including Waddington.[7] In particular, Swann and Aprahamian identify Marxism’s historical perspective and its concern with transformation over time, as well as its “vision of the totality of the phenomena in nature that allowed both for its unity and its limitless diversity…that was not at the same time mechanically reductionist” as its likely appeal to these “adventurous young talents…that refused to accept disciplinary limitations and boundaries.”[8]

J.B.S. Haldane

For example, J.B.S. Haldane—who was not a member of the Theoretical Biology Club, but was well acquainted with all who were—was a prominent figure in the emerging mathematical theories of population genetics, and was also a card-carrying Marxist.

At the time, the mathematical formalisms of population genetics treated genes as entirely independent units, with the assumption that most traits were rigidly determined by genetic inheritance. This emphasis on genes was also coupled with the belief that genetic change only happened rarely, via random mutations, and that for changes in genes to distribute through a population required geological time scales (from the background assumptions of uniformitarianism and gradualism carried over from geology, as described elsewhere). This strict emphasis on genes as atomistic sources of control of traits and as insulated from their environments seems to share little in common with either an organicist approach to biology or a collectivist ideology like Marxism, thus precluding a connection between Haldane’s politics and his scientific work in population genetics.

However, in this context it is especially interesting that one of the main emphases of Haldane’s mathematical work was to show that selection coefficients could be larger than other population geneticists generally assumed, which allowed for a much more rapid evolution than was imagined possible by population geneticists before Haldane.[9]  In particular, Haldane’s work attributed much more of a link to the environment than was allowed by most other population geneticists. For example, in his paper on the famous case of the moths of Manchester,[10] Haldane demonstrated mathematically how the evolution from disproportionate numbers of speckled moths to disproportionate numbers of black moths within 50 years was plausible given the changes in environmental conditions around the industrializing city of Manchester, which was a practical impossibility given the then-accepted rates of genetic variation.[11] This link between environmental change and a commensurately rapid change in the biological constitution of organisms was in a way a confirmation of sorts of the Marxian assertion of the connection between humans and their environments, particularly in the context of industrialization, as described by Friedrich Engels, even though it was also well within the mainstream science of the time.

The science and ideology of Waddington

While it would be imprecise to label Waddington a Marxist per se (e.g., given that he once asserted that the organicist philosophy of Whitehead had actually superseded Marx’s dialectical materialism “with a fuller view of nature”[12]), it is clear that many of his closest associates were unabashed Marxists, and that Waddington had ideological inclinations which leaned in that direction as well.[13] Given the connections between social reform movements and open biologies described elsewhere, and given Waddington’s emphasis on the interactions of genes with their environments, it is perhaps not surprising that Waddington also advocated for socialist politics.

That said, these connections between ideological inclinations towards socialism and organicism in science during this era were not unique to Waddington and his close group of friends. Val Dusek, in his own account of the emergence of the anti-mechanistic, anti-reductionist biology and physics around this time, identifies many of the prominent scientists who embraced this more holistic view and who also proclaimed themselves as Marxists, and discusses the ways in which their ideological inclinations were manifest in their scientific work.[14]

Notably, the significance of these connections between politics and science were not lost on the scientists themselves. In his later life Waddington himself remarked on what he called the practical consequences of metaphysical beliefs on scientists’ work, observing from his own experience that “a scientist’s metaphysical beliefs are not mere epiphenomena, but have a definite and ascertainable influence on the work he produces.”[15]

This open acknowledgement of the connection between ideological beliefs and scientific work makes many scientists today uncomfortable, as if it should discredit the work of overt socialists and Marxists like Waddington, Haldane, and Bernal because of the level of the influence of their political beliefs on their work. However, given the caliber of these scientists and of their scientific work—some of which now constitutes the bedrock of contemporary genetics—this assumption that the influence of ideology automatically invalidates scientific work does not hold water.

Likewise, to assume that extra-scientific beliefs like political ideology only influenced the work of these few socialist scientists working in 1930s and 1940s who openly acknowledged this connection, while the work of other non-socialist scientists who did not acknowledge such a connection is somehow exempt from such influences is not supported by the political and scientific history of the science of genetics. As discussed in previous chapters, political ideology has been a pervasive influence on the development of the science of biology throughout its history, including on many of those innovations which are now accepted as its core orthodoxy. As this history shows again and again, claims about the ideological neutrality of scientific programs are usually at best unintentionally myopic, or at worst hubristic—and are themselves likely the manifestations of a particular ideology which is taken as self-evidently true. Again, this influence of ideology does not necessarily render such scientific claims unscientific—otherwise there would have been no development of science over the course of this history—but it does mean that ideologies must also be a consideration in the evaluation of scientific claims.

As such, in the final section of this book on the public policy implications of epigenetics, these connections between ideologies and science and policies will be seen to still be live concerns in regards to contemporary epigenetics—although now in our contemporary political contexts and not necessarily as a contest of socialism contra capitalism and democracy, as in the geopolitical context of World War II and the Cold War, which is the subject of the next chapter.

[1] Werskey, G., 1978. The Visible College: The Collective Biography of British Scientific Socialists of 1930s. New York: Holt, Rinehart, and Winston.

[2] Gilbert, S. F., & Sarkar, S. (2000). Embracing complexity: organicism for the 21st century. Developmental dynamics219(1), 1-9.

Bedau, M.A. and Cleland, C.E., 2010. The nature of life: classical and contemporary perspectives from philosophy and science. Cambridge University Press, p. 95.

[3] Peterson, E. (2010). Finding mind, form, organism, and person in a reductionist age: The challenge of Gregory Bateson and CH Waddington to biological and anthropological orthodoxy, 1924–1980 (Doctoral dissertation). Retrieved from CurateND. (https://curate.nd.edu/downloads/js956d5968k), pp. 39-41.

[4] Haldane the elder is also famous for his penchant for conducting his respiratory experiments upon himself, and for inventing the first gas mask from his firsthand observations of poison gas attacks during World War I.

[5] Ibid.

[6] Bowler, P.J., 2010. Reconciling science and religion: The debate in early-twentieth-century Britain. University of Chicago Press, p. 169.

[7] Swann, B., & Aprahamian, F. (Eds.). (1999). JD Bernal: a life in science and politics. Verso, pp. xvi-xviii.

[8] Ibid., p. xviii.

[9] Crow, J. F. (1987). Population genetics history: a personal view. Annual review of genetics21(1), pp. 5-7.

[10] Haldane JBS. A mathematical theory of natural and artificial selection. Trans Cambridge Philos Soc. 1924;23:19–41.

[11] Larson, E. J. (2004). Evolution: the remarkable history of a scientific theory (Vol. 17). Random House Digital, Inc., pp. 218-224.

[12] Gilbert, S. F. (1991). Induction and the origins of developmental genetics. In A conceptual history of modern embryology (pp. 181-206). Springer US.

[13] Bowler, P. J. (2010). Reconciling science and religion: The debate in early-twentieth-century Britain. University of- Chicago Press, pp. 174-175

Waddington, C. H. (1942). Science and Ethics: An Essay. George Allen And Unwin Ltd.; London.

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

[15] Waddington, C.H. 2009. The Practical Consequences of Metaphysical Beliefs on a Biologist’s Work: an Autobiographical Note. In C.H. Waddington ed. Sketching Theoretical Biology: Toward a Theoretical Biology (Vol. 2). Transaction Publishers.

A Tale of Two Fields: Epigenetics and Biology Between the Wars

Excerpt from my forthcoming book Epigenetics and Public Policy The Tangled Web of Science and Politics to be released February 2018 by Praeger

One of the roots of the decades-long delay in the acceptance of epigenetics is found in the divergence of emphasis in biology between the emerging science of genetics and the more established field of embryology in the 1930s and 1940s. As will be discussed, this divergence had material, methodological, and geographical aspects. My contention is that this divergence also had substantial political and ideological aspects which were to become even more apparent in the onset of the Second World War, as these ostensible methodological differences also mirrored these impending political fractures.

This schism between embryology and genetics also complicates the conventional picture of the development of science as a logical, inevitable progression, and of the differences between fields as merely the result of a functional division of labor. Instead, at least in this case, this divergence between embryology and genetics appears to have occurred both for scientific reasons, but also as “a struggle for power and authority.”[1]

These dynamics are not exclusive to genetics but seem to be characteristic of science itself. For example, the sociologist Pierre Bordieu[2] and the historian Steven L. Goldman,[3] in their own analyses of the history of science, identify how what often becomes accepted as science is not obviously and necessarily more valid than what is deemed unscientific; instead, legitimate science is often determined just as much by “those who manage to impose the definition of science [as] having, being and doing what they have, are or do,”[4] which outcomes then become justified by assumption after the fact.

In other words, often there are no clear natural distinctions between different possible interpretations of the same natural phenomena, or no objective ways that nature can be ‘carved at its joints,’ and so what come to be the defining assumptions of a science have to be decided by other means. For example, in this case of the schism between genetics and embryology, it is fair to say that neither side was ‘wrong’ as such about the phenomena in question. What was primarily different was in the focus of each, or where the locus of causation was being placed. A legitimate science of biology built upon the work being done in embryology at this time, with genetics as a subsidiary component, is as conceivable as the genetics with embryology as a subsidiary component which actually did develop. In fact, the recent emergence of contemporary epigenetics, in which both these emphases are combined, indicates what such a science would have looked like. The actual result of this schism in biology, though, was a science of biology with the gene as the primary—and practically exclusive—cause.

All this is not to suggest that the science of genetics as it has developed since this time is therefore somehow not legitimate science. It has been an undoubtedly successful scientific enterprise by any standard. This is rather to say that the gene-centric focus of molecular biology is not the only direction a legitimate science of biology could have taken at this particular juncture, and that there were other equally viable alternative routes. That genetics developed as it did, though, is due to both scientific and political factors, although again this observation should not be construed as a condemnation of genetics.

Bricks in their walls

The fields of genetics and embryology had begun to diverge noticeably from each other by the mid-1920s, and this split became definitive by the 1940s—not that embryology disappeared, but that it was relegated or subordinated to genetics. According to the Nobel-prize winning geneticist T.H. Morgan, “an embryologist who inadvertently founded the gene theory,”[5] in his 1934 book Embryology and Genetics, this split was ostensibly along the lines of focus: Genetics and geneticists focused on the transmission of hereditary traits, while embryology and embryologists focused on the expression of those traits.[6]

However, while this may now seem an inevitable division, why such a split was deemed necessary in the first place, and why it should be along the lines of genetics versus embryology, is not so straightforward. This ambiguity is one indication that extra-scientific factors also played a role in this schism, which in turn influenced the timing of the rejection and the eventual begrudging acceptance of epigenetics, which takes more of an embryological approach. For this reason, the underlying social and political currents of this split are of particular relevance to the focus of this book on the policy implications of epigenetics to be discussed in the final section, which are a product of the novelty of epigenetic explanations in the accepted science of genetics.

Notably, this split was facilitated in large part by the work and the influence of Morgan himself, who set the stage for a distinctly American genetics in the 1920s—which would go on to become the prevailing conception of genetics and biology after World War II coincident with the zenith of American global hegemony. Morgan accomplished this division primarily by promoting the nuclear envelope, or the membrane which surrounds the genetic material in cells, as the primary conceptual and disciplinary boundary between genetics and embryology: what occurred within the envelope was the domain of genetics and what occurred outside was the domain of embryology. However, as the historian of biology Jan Sapp observes, the priority Morgan gave to this membrane, and to the disciplinary distinctions which resulted, were not “an intrinsic logical necessity of scientific thought,” but rather “depended directly on both the technical capacity and the institutional power of the discipline within which they were produced.” [7]

In particular, Morgan’s physiological distinction also highlighted an important geographical difference as well. Most biologists in Europe at this time did not recognize the priority of the physical boundary of the nuclear envelope as asserted by Morgan, but rather considered it to be just one component of the physiology of genes and the cell and the organism. Peter Bowler in his own history of evolutionary thought also notes this key geographical and cultural difference, observing that in Europe “genetics developed in a much less dogmatic form,” with much more interaction between these different subfields, and much more openness even to “non-Darwinian mechanisms of evolution” than in America in particular.[8]

The European approach to biology, in particular as championed by German scientists, asserted that the cytoplasm—or the non-genetic material within a cell not including the nucleus—played an important role in gene function, especially in providing the building material for the chromosomes and the genes themselves. In this way the external environment and the properties of the cells were seen to have a substantial influence on the functions of the genes within the cells. As such, both development and heredity were implicated in this more holistic focus, such that biologists in Europe did not have to confine themselves to studying either inheritance or development, as evidenced by the career of Waddington described elsewhere.

Morgan himself, as an embryologist, was declaring as late as 1910 that “We have come to look upon the problem of heredity as identical to the problem of development,”[9] which was well within this more European and German approach to genes and development. However, by 1926 Morgan the geneticist was asserting that cell composition and structure could be ignored in relation to the genes, such that the explanations of both inheritance and development could be found exclusively within the genes.[10]

The ability of this gene-centered faction of biology to resist absorption by the more established field of embryology constituted what the historian Scott Gilbert identifies as the “last chapter” in the emergence of a particularly American biology, which was to go on to become the prevailing gene-centered conception of biology more generally following World War II. “When had American biology finished ‘emerging’?” Gilbert asks, “I suspect that stage was reached when it had successfully resisted the last attempts to integrate it into European-dominated traditions of inquiry.”[11]

Thus, beyond the legitimate scientific rationales for this emphasis on the nuclear envelope which in part fomented the schism between embryology and genetics, some of the impetus behind this division of fields also stemmed from geneticists asserting their disciplinary independence from embryology, and from the desire of the American school of genetics in particular to assert its independence from the “European-dominated traditions of inquiry” which prioritized embryology and development over a near-exclusive focus on the genes.[12]

Again, per the guiding model of this project, that the sides in these disciplinary quarrels in biology coincided with the eventual sides taken in the Second World War, and that genetics ultimately emerged as the hegemonic victor in biology at the same time the U.S. emerged as the global hegemon after the war, is not merely coincidental. One of the effects of this distinctly Americanized focus on genes which emerged after the Second World War, and which was further solidified during the Cold War, was the antipathy towards epigenetic explanations like those proposed by C.H. Waddington in the 1940s which integrated genetics with embryology. This longstanding antipathy towards epigenenetics resulting in part from this disciplinary divergence of genetics and embryology from before the war helps to explain the timing of the recent (re)emergence of contemporary epigenetics from within genetics. This long delay in the reintegration of epigenetic explanations into genetics in turn helps to explain the political challenges now presented by epigenetics, which are the focus of this book: If epigenetics had been incorporated into the edifice of modern genetics as it was being constructed through the 1940s-1960s—as it very well could have been, given other social and political circumstances—then epigenetics would not present the conceptual and interpretive issues that it does now.

This intra-disciplinary contest between genetics and embryology was just one field of battle in this clash of science, politics, and ideologies during the interwar years. There were many other arenas in which developments in science mirrored the conflicts between political ideologies leading up to both the Second World War and the subsequent Cold War. Describing the circumstances of this convergence of science and ideology on a global scale from before World War Two through the Cold War, and how they pertain to epigenetics, will be the focus of subsequent chapters.

[1] Sapp, J. (1983). The struggle for authority in the field of heredity, 1900–1932: New perspectives on the rise of genetics. Journal of the History of Biology16(3), 311-342.

[2] Bourdieu, P. (1999). The specificity of the scientific field and the social conditions of the progress of reason. Sociology of Science 14(6): 19-47.

[3] Goldman, S.L., 2006. Science wars: What scientists know and how they know it. Teaching Company.

[4] Bourdieu 1999, p. 24.

[5] Gilbert, S.F. (1991). Cellular politics: Ernest Everett Just, Richard B. Goldschmidt, and the attempt to reconcile embryology and genetics. In Rainger R, Benson KR, Maienschein J. (eds) The American Development of Biology. Philadelphia: University of Pennsylvania Press. Available at http://10e.devbio.com/article.php?id=26

[6] Morgan, Thomas Hunt. (1934). Embryology and Genetics. New York, NY: Columbia University Press.

[7] Sapp 1983.

[8] Bowler, Peter J. (2003). Evolution: the History of an Idea (3rd ed.). California: University of California Press.

[9] T. H. Morgan, “Chromosomes and Heredity,” American Naturalist, 1910, 44: 449-496.

[10] T. H. Morgan, “Genetics and the Physiology of Development,” Am. Nat., 1926, 60:459-515.

[11] Ibid.

[12] Gilbert, S.F. (1991). Cellular politics: Ernest Everett Just, Richard B. Goldschmidt, and the attempt to reconcile embryology and genetics. In Rainger R, Benson KR, Maienschein J. (eds) The American Development of Biology. Philadelphia: University of Pennsylvania Press, p. 311. (Available at http://10e.devbio.com/article.php?id=26)

Alfred Russel Wallace, Ideology, and Evolution

Excerpt from my forthcoming book Epigenetics and Public Policy The Tangled Web of Science and Politics to be released February 2018 by Praeger

One specific and particularly intriguing example of these differences in the content of the natural science of the 1800s due to social class and ideological inclination is the contrast in the description of natural selection by Alfred Russel Wallace with natural selection as described by Darwin.

Although largely unknown by most people outside of biology today, within biology Wallace is rightfully credited as both an independent co-discoverer of the theory of natural selection with Darwin and as providing the critical impetus for Darwin to finally publish his theory. That natural selection as described by Darwin has since assumed priority over that of Wallace is generally chalked up to minor but important theoretical differences and Darwin’s more complete elaboration of the concept in his book On the Origin of Species. I suggest, as do others,[1] that there are important differences between the two conceptions of natural selection. Further, I suggest that these theoretical differences are related to the social and political factors mentioned before, and that these social and political factors contributed as much to the eventual triumph of the Darwinian version as any claims of its enhanced correspondence with empirical facts.

For example, in contrast to the privileged station Darwin enjoyed in Victorian society, Wallace emerged from a working class background, was self-educated in Mechanic’s Institutes and ‘Halls of Science,’ which were set up for adult education by utopian Owenite societies,[2] and was also a committed socialist.[3] To wit, while Darwin emphasized selection at the level of individual organisms and the competitive aspect of evolution, which reflected the prevailing beliefs of his station as a proper Victorian, Wallace’s theory of natural selection emphasized environmental pressures on varieties or groups instead of just individuals, and cooperation instead of competition, both of which also reflected his strong socialist inclinations.[4] Thus, while there are striking similarities between the two versions of natural selection, there are some substantial differences as well, most of which break along the class and ideological lines described before. That the Darwinian version of selection eventually became the predominant interpretation is, according to the guiding model of this project, not merely a coincidence but also a function of its congruence with these prevailing social and political factors.

On the other hand

At the same time, though, all of this is not to say that there is always an exclusive one-to-one correspondence between an ideology and a specific understanding of biology—or that a belief in socialism necessarily requires an adherence to a Lamarckian understanding of biology, as was prevalent during the 1800s. For example, one noteworthy exception to the pattern of relationships described before—and one which merits much more scrutiny and commentary than I can give here—is that Wallace himself, contrary to expectations given his  socialist ideology, explicitly and consistently disavowed Lamarckian inheritance in his conception of natural selection to a greater degree than even Darwin.[5] Where Darwin explicitly included a role for Lamarckian use-disuse inheritance in evolution,[6] and later even proposed his hypothesis of pangenesis as a mechanism for such inheritance, Wallace was much more definitive in asserting that this kind of inheritance was unnecessary in natural selection.

Notably, as described in more detail elsewhere, the version of Darwinism which ultimately formed the foundation of modern genetics was ‘Neo-Darwinism,’ which is basically Darwinian natural selection scrubbed of any equivocation on the inheritance of acquired traits—which is actually more akin to Wallaceism. This theoretical difference suggests that relatively few modifications to Wallaceism would have been required to be congruent with genetics, as were required of Darwinism, such that the Modern Synthesis of genetics with natural selection could perhaps have emerged decades earlier if Wallace’s version of natural selection had been selected as the explanation of choice for natural selection. If Wallace’s version had been selected, though, given the differences just cited, the resulting theory of genetics would have been substantially different in many ways than the contemporary version of genetics as we now know it—including, perhaps, in the even earlier acceptance of epigenetics. This is an intriguing counterfactual which would be worth more exploration. That Wallace’s version of natural selection was not selected by the prevailing science of the time, though, also seems due at least in part to these social and political differences mentioned before, in addition to these differences in the scientific content.

Likewise, as will be shown in the next chapter, in the decades after Darwin many advocates of what would now be considered a distinctly conservative laissez faire ideology invoked a version of Lamarckism to justify economic and social policies, while progressives—particularly in America—actually turned to a reinterpreted Darwinian biology and the emerging science of genetics to explain and justify their politics and policies. This opposition between these Lamarckian ‘conservatives’ and these Darwinian ‘progressives’ constituted one of the important fundamental ideological oppositions of this era, and would shape the politics of the 20th century not only in America but on a global scale.

Per the guiding model of this project, what is consistent throughout all these examples is that in each case these interpretations of biology are made congruent with the ideology. In other words, even though these conservatives in the early 1900s were using Lamarckism while the progressives were using Darwinism—the exact opposite as in the 1800s—in each case the biology was interpreted in line with the dictates of the ideology, and vice versa. As such, these seeming counterexamples actually provide support for this notion of the necessary connection between social and political ideology and biological science. This ultimate congruence of biology with ideology is a particularly important point for the discussion of epigenetics and public policy in the final section of this book.

[1] Kutschera, U. (19 December 2003). “A comparative analysis of the Darwin–Wallace papers and the development of the concept of natural selection”. Theory in Biosciences122 (4): 343–59.

Glickman, S. E. (2009). Charles Darwin, Alfred Russel Wallace and the Evolution/Creation of the Human Mind. Gayana (Concepción)73, 32-41.

Gross, C. (2010). Alfred Russell Wallace and the evolution of the human mind. The Neuroscientist,16(5), 496-507.

Ruse, M. (2013). Charles Robert Darwin and Alfred Russel Wallace: their dispute over the units of selection. Theory in Biosciences132(4), 215-224.

[2] Harrison, J. (2009). Robert Owen and the Owenites in Britain and America: the quest for the new moral world. Taylor & Francis, p. 189.

[3] Green, J. (2012). Alfred Russel Wallace: Socialist and co-founder of evolutionary theoryLondon Progressive Journal. Retrieved 15 August 2017, from http://londonprogressivejournal.com/article/view/1049/alfred-russel-wallace-socialist-and-cofounder-of-evolutionary-theory.

Cervantez, S. R. (2016). Facts Are Stubborn Things: The Foundation of Alfred Russel Wallace’s Theories, 1823-1848 (Master’s thesis). Retrieved from http://digitalcommons.lsu.edu/gradschool_theses/1900.

[4] Jones, G. (2002). Alfred Russel Wallace, Robert Owen and the theory of natural selection. The British Journal for the History of Science35(01), 73-96.

[5] Wallace, A. R. (1889). Lamarck versus Weismann. Nature40(1043), 619-620.

Stack, D. (2003), The First Darwinian Left: Socialism and Darwinism, 1859-1914, London: New Clarion, p. 29.

[6] “Curiously few evolutionists have noted that, in addition to natural selection, Darwin admits use and disuse as an important evolutionary mechanism. In this he is perfectly clear. For instance,…on page 137 he says that the reduced size of the eyes in moles and other burrowing mammals is “probably due to gradual reduction from disuse, but aided perhaps by natural selection.” In the case of cave animals, when speaking of the loss of eyes he says, “I attribute their loss wholly to disuse” (p. 137). On page 455 he begins unequivocally, “At whatever period of life disuse or selection reduces an organ…” The importance he gives to use or disuse is indicated by the frequency with which he invokes this agent of evolution in the Origin. I find references on pages 11, 43, 134, 135, 136, 137, 447, 454, 455, 472, 479, and 480.” (Mayr, E. (1964/1859). “Introduction.” In Charles Darwin. On the Origin of Species: a Facsimile of the First Edition. Harvard University Press.)

Lamarckism and the Biology of Discontent in the 1800s

Excerpt from my forthcoming book Epigenetics and Public Policy The Tangled Web of Science and Politics to be released February 2018 by Praeger

Just how significant and real were the threats being confronted by the defenders of the status quo at this time? As described by John Bellamy Foster, England in the mid-1800s was “a seething cauldron of discontent” due to the sudden social, political and economic changes which accompanied rapid industrialization.[1]

Throughout England during this time there were strikes, demonstrations and riots against work conditions and wage disparities, and especially against the enforcement of the Poor Laws, passed in 1834. For example, the phrase ‘reading the Riot Act’ can be traced through this era as during many of these demonstrations the Riot Act was literally read aloud to advise demonstrators that they were assembling unlawfully and that deadly force would be applied if they did not disperse.

The Chartist movement in particular, so-called for the People’s Charter published in 1838, was a major force in this social unrest.[2] The Chartist movement focused primarily on reforms of the political system in England which favored the working classes, and was able to marshal the support of millions of people from all around the country, but especially from the newly industrialized areas. In 1839, 1842, and 1848, Chartist petitions with millions of signatures each were presented to the House of Commons, each of which were summarily dismissed without a hearing. As can be imagined, this refusal of the formal political institutions of the state to even hear the grievances of such a significant proportion of the population only increased the tension and the rancor.

Much to the alarm of the political establishment, and to the consternation of the English religious establishment of the early 1800s, most of the agitators of this time favored the materialistic (i.e., godless) natural philosophies coming out of revolutionary France, and Lamarckism in particular. In contrast to the hierarchical but increasingly individualistic and competitive view of nature that was promoted by mainstream science in Britain—coincident with the emergence of industrialization, capitalism, and modern liberal politics in British society—these other groups envisioned societies organized around more collectivist and symbiotic principles, and Lamarckism provided a biological explanation for how such societies could be realized out of the present state of affairs.[3] Buoyed by these reinterpretations—or misinterpretations—of Lamarckism, by asserting that at a basic level organisms respond rapidly to their environments, these groups advocated for changes in the environment in the form of fundamental reconfigurations of society to achieve their goals of “egalitarianism, female emancipation, [and] secularization” in the progressive development from barbarism to civilization.[4]

In other words, while the clerical naturalists and gentrified scientists of this era saw a natural—and therefore a social—world that was set and ordered by divine command, millions of others more exposed to the vicissitudes of the recent societal upheavals instead saw a social—and therefore a natural—world that can and did change, sometimes precipitously, and without the providential oversight of Deity. That each side gravitated towards an understanding of biology which mirrored their lived experience is not only understandable but in many ways inevitable.

As described by Margaret Anne Loose in her analysis of the Chartist literature of the era, “Lamarck’s 1809 hypothesis that offspring acquired traits based on the associations and environments of their forebears was favored by many working-class thinkers (perhaps because it allowed one to hope that s/he could alter the future).”[5] In other words, if things were not good now, Lamarckism—at least as interpreted by these reformist groups—provided a reasonable explanation for how they could be made better. In contrast, the mainstream science of the time instead allowed only the possibility of incrementally small changes over geological time scales (keeping in mind that Lamarck himself also actually employed these same ‘conservative’ uniformitarian and gradualist principles). These Lamarckesque ideas about adaptation and inheritance also had the benefit of seeming common sense, as this was how the natural world appeared to work to most people of this time. Thus, this more explicitly socialist version of Lamarckism seemed to confirm to many what they already knew about the world, and provided reasons to imagine the possibility of a future fundamentally different than the present.

These hopes for a better future via Lamarckian adaptations were predicated on alterations of current environments as sweeping social and political changes. As such, Lamarckism became a fundamental aspect of many of the rationalist (i.e., secular) progressive reform movements of the mid-1800s in which “a belief in the perfectability of humankind and the self-organizing power of matter according to natural laws [was] joined to a faith in the environment as a determinant of form and character.”[6] These interpretations of Lamarckian biology supported the expectation that “through the appropriate social and material environment, humanity’s spiritual qualities could be molded as a prelude to political change.”[7]

…For example, Friedrich Engels, the cofounder of Marxism with Karl Marx, explicitly incorporated a Lamarckian understanding of evolution into his formulation of communism and the labor theory of history by suggesting that the physical adaptations to work played a crucial part in the biological transition from ape to man, such that “in a sense, we have to say that labour created man himself.”[8] From these origins, Engels traces the subsequent technological developments relative to labor which have continued to shape man, culminating in the creation of the steam engine, which instrument “more than any other was to revolutionise social relations throughout the world.”[9] Engels continues:

By long and often cruel experience, and by collecting and analysing historical material, we are gradually learning to get a clear view of the indirect, more remote social effects of our production activity, and so are afforded an opportunity to control and regulate these effects as well. This regulation, however, requires something more than mere knowledge. It requires a complete revolution in our hitherto existing mode of production, and simultaneously a revolution in our whole contemporary social order.[10]

Against these direct challenges to the status quo, the geologist Charles Lyell—who would go on to have such an influence on Charles Darwin via his three volume Principles of Geology—along with many others in the mainstream scientific establishment in Britain undertook concerted action against Lamarckism and any other such notions of the malleability of essential forms, with the primary intention to demonstrate “that morals were not the better part of brute instinct” and particularly “to prove that man was no transformed ape.”[11] Part of this rearguard action by scientists such as Lyell was to intentionally restructure geology and paleontology “along safe non-progressionist lines…to preserve man’s unique status in creation.” These intentions were so explicit that, according to the historian of science Adrian Desmond, without a doubt “Lyell’s biology and geology were inextricably related in Principles of Geology and his ideology affected his science as a whole.”[12] This was the openly ideological nature of the geology which had such an overwhelming influence on Charles Darwin in the eventual formulation of his epoch-marking theory of biological evolution, which in turn came to eventually constitute some of the basic assumptions of contemporary genetics—including its entrenched dispositions against the responsiveness to the environment and inheritance via epigenetics.

As such, people on both sides of this social cleavage in 19th century Britain identified the ideas of biological responsiveness to the immediate environment and the inheritance of those adaptations with calls for fundamental reforms of the prevailing social order. Further, as will be discussed in more detail later, Engel’s combination of history and economics with a Lamarckian biology directly influenced the subsequent development of Leninism, Stalinism and Maoism, which would go on to play such a significant role in the social and political history of the 20th century in antagonism with the liberalism of the West. As will be shown in later chapters, these political and ideological antagonisms, with their roots in these disputes over biological theories, also undoubtedly influenced the reception to epigenetics in the West, with its biological openness to the immediate environment and the inheritance of those adaptations.

However, what cannot be forgotten, per the previous descriptions of Lamarck’s actual theories, is how similar his theory of evolution actually was to Darwin’s eventual theory of evolution, particularly in reference to the minute internal variations and the geological time scales required for change to occur. Thus, the Lamarckism utilized by these reformist groups downplayed or ignored these other more uniformitarian aspects of Lamarck’s Lamarckism, and instead emphasized those aspects which fit with their desire to describe human nature as malleable and history as progressive. Regardless, Lamarckism did at least provide them with a platform in natural science to explain and justify their social and political agendas.

This history thus suggests that contemporary epigenetics may also have affinities with specific contemporary political ideologies, and will likely also be put to similar political uses which may or may not accurately reflect the underlying science. Being aware of this likelihood for the ideological support of, or opposition to, the results from epigenetics may be of use to both scientists and policymakers (and those concerned with public policy) as epigenetics become more of a factor in policy and politics.

[1] Foster, John Bellamy. 2000. Marx’s ecology materialism and nature. New York: Monthly Review Press, p. 179.

[2] Thompson, D. (1984). The Chartists: popular politics in the Industrial Revolution. Pantheon.

[3] Harrison, J. F. C.  (1969). Quest for the New Moral World: Robert Owen and the Owenites in Britain and America. New York.

[4] Lenoir & Ross 1996, p. 376

[5] Loose, M. (2010). Literary Form and Social Reform: The Politics of Chartist Literature (Doctoral dissertation). Retrieved from ProQuest Dissertation Express (UMI No. 3225641).

[6] Burkhardt, R. W. (1995). The Spirit of System: Lamarck and Evolutionary Biology: Now with” Lamarck in 1995″. Harvard University Press, p. 59.

Burkhart, R. (2011).  Lamarck, Cuvier and Darwin on Animal Behavior. In Gissis, S., & Jablonka, E. Eds. Transformations of Lamarckism: From subtle fluids to molecular biology. MIT Press, p. 40.

Haig, D. (2011). Lamarck Ascending! Philosophy & Theory in Biology3:e204.

Laurent, J., & Nightingale, J. (2001). Darwinism and evolutionary economics. Edward Elgar Publishing, p. 128.

[7] Lenoir & Ross 1996, p. 375.

[8] Engels, F. (1876/2015). The part played by labour in the transition from ape to man. In Dialectics of Nature: Explanation about Dialectical Materialism. CreateSpace, p. 171.

[9] Ibid., p. 191.

[10] Ibid.

[11] Desmond (1985), p. 25.

[12] Ibid.

C.H. Waddington: Genesis of the Original Epigeneticist

Excerpt from my forthcoming book Epigenetics and Public Policy The Tangled Web of Science and Politics to be released February 2018 by Praeger

Conrad Hal Waddington, who initially proposed epigenetics in 1939, is one of the more eclectic and interesting personalities in the natural science of his time, not for any flamboyance or peculiar affectation on his part, but for the range of his interests and work, which beyond biology also included an especial interest in the arts and philosophy. For example, although Waddington was a widely respected laboratory experimentalist, writer, and lecturer in embryology and genetics, holding prestigious research and university positions throughout his career, he accomplished all this without being academically credentialed in genetics or even biology. This diverse background, including Waddington’s unorthodox politics, will be shown to have a significant influence in his ‘discovery’ of epigenetics, and also provides clues as to why epigenetics was as ignored it was until fairly recently.

Given the importance of geology in the unfolding of the science of evolution detailed over the past few chapters, perhaps the first most conspicuous biographical fact about Waddington is that he earned his bachelor’s degree at Cambridge with a specialization in geology in 1926. He then went to graduate school also at Cambridge to study paleontology, focusing on ammonites, an extinct group of marine mollusks, which again mirrors significant developments in the early history of the theory of evolution. In these details, at least, Waddington reflected the origins and development of evolutionary thought from the previous century.

As a graduate student, Waddington actually held two studentships: one in geology and another in philosophy,[1] neither of which seem to have much of a link to the geneticist and epigeneticist that Waddington would become. However, Waddington’s interests in philosophy can be shown to have direct connections with his eventual ‘discovery’ of epigenetics. In particular, Waddington’s philosophy of choice was the process philosophy of the mathematician and philosopher Alfred North Whitehead, and especially Whitehead’s “philosophy of organism.” According to this philosophy, the world is composed not of material objects with their own independent existences, but rather of deeply interdependent processes and events.[2] The impact on Waddington of process philosophy and Whitehead’s book Science and the Modern World as an undergraduate appears to have been so profound that it impelled his exit from geology and his entrance into the world of biology and genetics,[3] which culminated in his postulation of epigenetics.

Waddington and the philosophy of process

Process philosophy is famously distinct from the prevailing reductionist and analytic paradigm of Western philosophy and science since Plato, which identifies objects as independent entities which can be broken down into their constituent parts until the simplest components are revealed, which will then explain the entire object. Instead, in process philosophy, an object is just “an ingredient” in the character of some event, only having effects via its interactions with other objects in these “events.”[4] For Whitehead, the provisional successes of this reductionist approach in science rendered what was a useful heuristic into an unquestionable worldview—but one which reasoned in a circle by taking its stipulated assumptions about independent and reducible objects as evidence of their existence.[5] Instead, Whitehead proposed that all existing things were the organic product of the ongoing interactions of many different processes, existing only as “events,” or as the persistence of the product of these interactions.[6]

Process philosophy was a lifelong inspiration for Waddington in perceiving the organic world as composed not of individual and independent entities but as the product of ongoing interconnected processes. In the last year of his life, Waddington describes the enduring effects of this philosophy on his scientific work, remarking that this early exposure to Whitehead had “totally inoculated me against the present epidemic intellectual disease, which causes people to argue that the reality of anything is proportional to the precision with which it can be defined in molecular or atomic terms.”[7]

As a signal of what was to come, in the paper which won him his philosophy scholarship in 1929, Waddington focused on the implications for biology of the process philosophy of Whitehead. In this paper, Waddington suggests that if all things actually are the product of interacting processes, instead of independent entities, then the “scientific explanation of the process of evolution, as that it is brought about by natural selection acting on gene mutations,” would still not qualify as a complete general explanation of evolution until it even genes are conceived as events, or as the ongoing product of these other processes.[8]

After only three years of graduate school, Waddington left Cambridge without having completed his planned doctoral thesis in geology to pursue his burgeoning interests in biology. Through a close friend (the equally eclectic and iconoclastic anthropologist Gregory Bateson, son of the geneticist William Bateson who actually coined the term ‘genetics’ in 1905), Waddington was given the opportunity to work with the eminent horticulturist and geneticist Edith Saunders. Often referred to as The Mother of British Plant Genetics, Saunders is most known for her significant role in the rediscovery of Mendelian heredity. Waddington worked with Saunders on a longstanding problem in the genetics of maladaptive recessive traits in plants, writing a paper on his treatment of the problem which was published in the prestigious Journal of Genetics in 1929.[9] In 1931, Waddington published a highly technical statistics-based paper on inbreeding and genetic linkage in the equally prestigious journal Genetics,[10] which he wrote in collaboration with the respected population geneticist J.B.S. Haldane.[11]

The move from genetics to embryology (and back)

Even with these early successes, though, Waddington was dissatisfied with the emphasis on reduction that permeated the study of genetics. Through another close friend Waddington secured a job at the Strangeways lab at Cambridge to study embryology, which he felt better reflected his inclinations towards process and development. Within a year, Waddington was able to complete difficult experimental work on the development mechanics of embryos in vitro that the lab founders had been unable to figure out, and to publish the results in Nature.[12] Waddington was then able to parlay these successes into a six-month stint at one of the most prestigious laboratories of embryology in the world in Berlin (the lab of Hans Spemann and Otto Mangold). Upon his return to Cambridge, Waddington was offered a position teaching experimental zoology and a fellowship with the Medical Research Council. Also, with another of his closest friends (the biochemist and polymath Joseph Needham), Waddington was able to build up one of the most respected laboratories of embryology in Europe on a shoestring budget.[13]

In his laboratory work, Waddington focused for the most part on the ‘organizer,’ which is the cluster of cells in developing embryos which induces the development of the central nervous system.[14] The discovery of the organizer and its basic mechanisms in 1921 by Ph.D. student Hilde Mangold and her advisor Hans Spemann merited a Nobel Prize for Spemann in 1935, but not Mangold who died from severe burns suffered in a kitchen explosion in her apartment in 1924.[15] Through this lab work Waddington repeatedly observed firsthand how the processes of development depended on specific configurations of the results of prior developments, and how differences in these prior conditions altered the outcomes of development. These observations would be pivotal in his subsequent postulation of epigenetics.

However, during this period Waddington also spent time at the Drosophilia (fruit fly) labs in Caltech and at the labs in Cold Spring Harbor. These labs were the epicenters of the emergence of molecular genetics which established the conception of genes as atomistic units. At these labs Waddington worked with some of the foremost researchers in genetics, such as Theodosius Dobzhansky and Alfred Sturtevant, just as they were making the discoveries which would contribute so much to the gene-centric focus of the Modern Synthesis. From this work, Waddington published numerous papers on the genetic control of wing development in normal and mutant strains of drosophila in journals such as The Proceedings of the National Academy of Science of the U.S.A, the Journal of Genetics, and Nature.

After World War II, during which Waddington contributed to the Allied war effort via operational research with the Royal Air Force, Waddington accepted an offer to be the chair of animal genetics at the University of Edinburgh where he remained until his death in 1975. At the time, this was one of only three chairs of genetics in the United Kingdom.

Thus, although Waddington was at heart and in practice an embryologist, and had not earned a degree in genetics, he was deeply involved in the development of the genetics of his time, and was respected enough to be considered one of the top scientists in the field. As will be discussed elsewhere, though, Waddington’s identification as an embryologist was in many ways as politically charged as his controversial political associations, to be discussed next. This discussion will demonstrate the intimate connections between Waddington’s background, his scientific work, and his politics, culminating in his postulation of epigenetics.

[1] The majority of these biographical details are gathered from the excellent and highly detailed dissertation of Erik L. Peterson, 2010, Finding mind, form, organism, and person in a reductionist age: The challenge of Gregory Bateson and CH Waddington to biological and anthropological orthodoxy, 1924–1980. University of Notre Dame.

[2] Whitehead, A.N. (1929/2010). Process and Reality. New York: Simon and Schuster, p. 73.

[3] Peterson, E.L., 2011. The excluded philosophy of evo-devo? Revisiting CH Waddington’s failed attempt to embed Alfred North Whitehead’s” organicism” in evolutionary biology. History and philosophy of the life sciences, pp.301-320.

[4] Whitehead, A.N. (1920/2004). The Concept of Nature. Mineola, NY: Dover Publications, Inc., p. 143.

[5] Whitehead, A. N. (1925/2011). Science and the modern world. Cambridge University Press, pp. 50-51.

[6] Whitehead (1925/2011), pp. 151-152.

[7] C. H. Waddington, “Fifty Years On,” Nature 258 (1975): 20–21.

[8] Waddington 1929 [unpublished], p. 66 in Peterson, 2011.

[9] Waddington, C. H. (1929). Pollen germination in stocks and the possibility of applying a lethal factor hypothesis to the interpretation of their breeding. Journal of Genetics, 21(2), 193-206.

[10] Haldane, J. B. S., & Waddington, C. H. (1931). Inbreeding and linkage. Genetics, 16(4), 357.

[11] Waddington’s relationship with Haldane will come up again in regards to their politics, but it is important to note at this point that Waddington later recalled his experience with Haldane and the “thin gruel of mathematical formalism” as a rather joyless endeavor, and said of Haldane that “I never knew him well, and I don’t actually think very much of all the ‘mathematical theory of evolution’ stuff which he started [because] it is all ultimately based on the quite falacious [sic] notion that selection coefficients belong to genes, whereas actually they belong to phenotypes,” which for Waddington was a distinction with “profound consequences.”

[12] Waddington, C. H. (1930). Developmental mechanics of chicken and duck embryos. Nature, 125, 924-925.

[13] Peterson 2011, p. 307.

[14] Philbrick, S., O’Neil, E. (2012, January 12) Spemann-Mangold Organizer. Embryo Project Encyclopedia . ISSN: 1940-5030 http://embryo.asu.edu/handle/10776/2330.

[15] Doty, M. (2011, May 9) Hilde Mangold (1898-1924). Embryo Project Encyclopedia. ISSN: 1940-5030 http://embryo.asu.edu/handle/10776/1743.

The Unfortunate Legacy of Jean-Baptiste Lamarck

Excerpt from my forthcoming book Epigenetics and Public Policy The Tangled Web of Science and Politics to be released February 2018 by Praeger

Although the ultimate goal of this book is to explain or predict the policy implications of epigenetics, which probably seems to not require much attention to the history of epigenetics, this history is actually necessary to understand a number of aspects of contemporary epigenetics and its political implications. For example, this history is absolutely necessary to understand why epigenetics has only begun to emerge within the last couple decades, especially given that epigenetics as we now know it—minus the suggestions of transgenerational inheritance—was first proposed around eighty years ago.

In this context, a common tactic used to downplay or denigrate epigenetics is to refer to epigenetics as ‘Lamarckian.’ This is a reference to the almost universally discredited theory of evolution of Jean-Baptiste Lamarck, which is contrasted with the Darwinian theory of evolution by natural selection. The twist of all this, though, is that this version of history is a fundamental misconstrual of both Lamarckism and epigenetics. Why Lamarckism was misconstrued in this specific way, and how Lamarckism came to be anathema to mainstream genetics, and why epigenetics came to be misidentified as Lamarckian are all questions which are answered by this history.

Lamarck, evolution, and world history

The first comprehensive theory of organic evolution was formulated by Jean-Baptiste Lamarck (1744-1829), and first publicly presented by him at the Museum of Natural History in Paris in May of 1802, more than 50 years before Darwin presented his theory of evolution by natural selection. Although Lamarck is mostly remembered now, if at all, as a footnote in the history of evolutionary theory, it was Lamarck’s theory of evolution as the inheritance of acquired characteristics which was utilized by the progressive social reformers of the 1800s, such as Marx, Engels, and many others, in their struggles against the defenders of the status quo. At the same time, Lamarckism is also the theoretical framework which was invoked by Herbert Spencer and the other so-called Social ‘Darwinians’ at the end of the 19th century, and which was ostensibly the basis for the state-sanctioned science of the Soviet Union in the mid-20th century.

In other words, Jean-Baptiste Lamarck has had as much an impact on the political history of the modern world as any biologist, perhaps save Darwin—but, notably, much of this impact is based on misunderstandings and reinterpretations of Lamarck’s actual theory of evolution. Likewise, contemporary epigenetics is often compared with Lamarckism in a way which paints both in an unfavorable light; but, as I will also show, these comparisons are also again usually based on misunderstandings of Lamarck’s actual theory of evolution, or misunderstandings of contemporary epigenetics, or both.

The unfortunate legacy of Lamarck

First, although Lamarck is remembered most for his description of biological change over time via the inheritance of adaptive characteristics acquired during the life of an organism, as a member of the French Academy of Sciences and a professor of botany and zoology at the Museum of History in Paris his scientific interests and pursuits were much broader than just biological inheritance. In fact, according to the historian of science Jean Gayon,[1] heredity as such was actually only of minor interest to Lamarck in his scientific work.

As described in detail by historian Richard Burkhardt in a recent article published in the journal Genetics,[2] the history of science is often marked by three almost perverse kinds of remembrances: When an event or discovery that seems significant in retrospect is barely noticed at the time,[3] or when an event or discovery is trumpeted as significant at the time but disappears from history, or when someone is remembered for something that is not what he or she would have considered to be his or her most significant achievement. Lamarck is an unfortunate example of the last of these kinds of perverse remembrances.

As Burkhardt explains, in contrast to the common perception of Lamarck, he did not claim as his own the notion that acquired characteristics could be inherited. “While it is true that Lamarck endorsed the idea of the inheritance of acquired characters and made use of it in his evolutionary theorizing,” Burkhart observes, “neither Lamarck nor his contemporaries treated this as Lamarck’s ‘signature’ idea.”[4] That inheritance of this kind was an accepted explanation of the time helps also to explain why Lamarck did not feel the need to confirm his theory for his audience through a vast assemblage of supporting facts or by experimentation. Rather, Lamarck, like most of his contemporaries, treated this idea as the well-known alternative theory it was—even though it was counter to the more generally accepted theory of the time of species as fixed.

For example, in the introduction to his multi-volume Histoire naturelle des animaux sans vertèbres (Natural History of the Invertebrates) published in 1815, Lamarck described the idea that individuals of one generation inherit the biological organization acquired by their parents during their lives as a “law of nature” which is “so much attested by the facts, that there is no observer who has been unable to convince himself of its reality” (Lamarck 1815, p. 200). Burkhardt also describes how the eminent naturalist Charles-Georges LeRoy (1723–1789) and the political philosopher Marquis de Condorcet (1743–1794) also invoked this inheritance of acquired characters in regard to the perfectibility of both animals and humans as examples of the company Lamarck kept in upholding such a theory. Likewise, the historian of science Pietro Corsi, in a meticulous analysis of French scientific thought,[5] places Lamarck squarely amidst the scientific debates of the time, and not as the ridiculous, possibly insane outcast on the fringes of science as he is so often portrayed in conventional accounts.

In other words, and contrary to how Lamarck is usually described, he was neither on the lunatic fringe of the science of his time with his theory of the inheritance of acquired characteristics, nor was he even particularly concerned with the inheritance with which he is now so indelibly associated. Instead, it seems, Lamarck was using a common alternative account of biological change as one piece of his more comprehensive systematic theory of development and change.

As such, a basic understanding of Lamarck’s actual theories of evolution and inheritance is important in understanding not only how biological inheritance and interaction with the environment was understood and used politically in the past, but also why contemporary epigenetics is not Lamarckism, which will help to reveal the actual ‘hidden’ political content of contemporary references to epigenetics as Lamarckism.

[1] Gayon, J. (2006). Hérédité des caractères acquis, pp. 105–163 in Lamarck, Philosophe de la Nature, edited by P. Corsi, J. Gayon, G. Gohau, and S. Tirard. Presses Universitaires de France, Paris.

[2] Burkhardt, R. W. (2013). Lamarck, evolution, and the inheritance of acquired characters. Genetics194(4), 793-805.

[3] Burkhardt gives the example of Thomas Bell, the president of the Linnean Society of London in 1858, who in his annual review of the Society’s meetings for that year concluded that nothing revolutionary had been brought up in their meetings of that year, even though Bell was actually presiding over the Society meeting of July 1, 1858 in which a paper by Charles Darwin and Alfred Russel Wallace expressing their views on natural selection was read and discussed.

[4] Ibid., p.794.

[5] Corsi, P. (1988). The Age of Lamarck: Evolutionary Theories in France, 1790–1830. University of California Press, Berkeley.