Epigenetics, ethics and the evolution of science

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by Shea Robison (@EpigeneticsGuy)

In this post and others, the historical, cultural and philosophical underpinnings of the scientific assumptions of genetics are discussed. That the assumptions of genetics are contingent on these historical and philosophical precedents likely seems trivial to laypersons and nonsensical to geneticists. Genetics, as a product of modern science, exhibits the patina of scientific objectivity which appears to negate these historical and cultural influences. However, as described encyclopedically by Daston and Galison (2007)[1], this conception of scientific objectivity is itself a product of the same historical and cultural processes which have shaped our contemporary politics and ethics, and is therefore just as contingent as our politics and our ethics.

Daston and Galison do not address the field of genetics in their analysis of the history of scientific objectivity, but both the modern concept of genes and the practice of genetics are prototypical examples of the epistemology and the ethos of modern scientific objectivity identified by Daston and Galison. How genes and genetics embody this concept of objectivity and its accompanying ethos also demonstrates how the scientific challenges of epigenetics to genetics are at once also challenges to the prevailing ethics of contemporary society.

The self and scientific objectivity

What lends contemporary science much of its current power is its objectivity. Objective in this sense is the ostensible removal—as much as possible—of any traces of the subjective individual(s) conducting the science: The less the presence of the individual scientist is a factor in the results of a scientific experiment, the more faithfully does the experiment reflect nature, the more replicable it is by other scientists, and so on.

In their discussion of the evolution of this concept of objectivity which so epitomizes contemporary science, Daston and Galison draw the critical connection between what constitutes knowledge with what is the accompanying conception of self: “The answer to the question ‘Why objectivity?’ lies precisely in the history of the scientific self to be eliminated”[2]. In other words, change the composition of this subjective self conducting a science, and the resulting scientific objectivity also changes.

This objectivity is also based upon a specific concept of the self. Just as this modern self has not always been the prevailing concept of self, as discussed here and here, the accompanying concept of scientific objectivity has likewise not always been the prevailing concept of scientific objectivity. In particular, Daston and Galison contrast the Enlightenment views of the self with the conceptions of the self that began to form after the transformative work of Immanuel Kant, and how these changes in conceptions of the self manifested as changes in the accepted definitions of scientific objectivity.

Daston and Galison use Kant as a reference point for these changes, and as an authoritative mouthpiece, but they are also careful to allow that whether Kant was actually the source of these changes in the conceptualization of the self is immaterial for their purposes; what is important is that Kant was at the very least a “precocious philosophical witness” of these changes “that reverberated with seismic intensity in every domain of nineteenth-century intellectual life, from science to literature”[3].

These differences in the assumptions about what constitutes the self and how this self gains knowledge (e.g., science) have direct relevance to how genes have been defined and conceptualized, and what are the implicit ethical commitments of genetics, and therefore why epigenetics presents the scientific and ethical and political challenges that it does.

The Enlightenment self

Citing various primary sources, Daston and Galison detail how during the Enlightenment era the self became perceived as a network or “skein of threads”[4] centered on an origin “as a spiderweb is centered on the spider”[5]. Personal identity was similarly conceived as “fragile as cobweb, guaranteed only by memory and the continuity of consciousness”[6]. As such, “the self was imagined as permeable, sometimes too permeable, to its milieu, a self characterized by receptivity rather than assertive dynamism”[7]. The “sovereignty of reason” located at the origin of this network—which is the seat of the self—“was always under threat from within (the vagaries of imagination and the uprisings of the branches of the network) and without (the barrage of sensations registered by the receptive network)”[8]. This Enlightenment concept of a passive, impressionable, malleable self is perhaps best embodied through John Locke’s concept of persons born as tablua rasa, or “white paper, void of all characters, without any ideas” upon which sensations from the external world are impressed[9].

Science and ethical behavior in the Enlightenment

The scientific ethos of this concept of self and its accompanying epistemology can be characterized as such: Because of the inherent passivity of this Enlightenment self as a permeable and impressionable network of competing influences, achieving truth-to-nature for Enlightenment scientists “required that they actively select, sift and synthesize the sensations that constantly flooded the too-receptive mind”[10]. Allowing oneself to be overwhelmed and carried away by the cacophony of sensations of the natural world as it is was “to be at best confused”[11], which is a moral and intellectual failure. In other words, the objectivity of the Enlightenment scientist was to interpose their selves—understood as the reasonable centers of the web of impressions which constitute their consciousness—into the flow of sensations from their external worlds

The ethics which were produced from this concept of self can be characterized as such: Good character—like good science—was seen as first overcoming this natural passivity by acting upon the world rather than being acted upon by the world[12]. Good character was also manifest in the balancing of this constant influx of stimulus from all the branches of this network by the reasonable center[13]. In contrast, bad character—like bad science—resulted from either an excess of passivity and impressionability or from this reasonable center being overwhelmed by all these other influences, manifest as supine acquiescence  to authority, or a surfeit of imagination, or other flights of irrationality[14].

The post-Kantian self

Distinct from this passive and permeable network self of the Enlightenment which is shaped by its environment, Kant  proposed a concept of self as an autonomous center which imposes order onto the world around it both by vetting all perceptions “like callers at the door”[15] and by projecting its assumptions outward onto the world.

Kant began from the proposition that sensations do not of themselves cohere into the perception of objects, much less into concepts. If order is observed, this awareness of order among the objects and sensations ‘out there’ therefore necessarily presupposes an ordering consciousness ‘in here’ that precedes perception[16]. Likewise, if this consciousness is imposing an order, it must also be ordered itself “as a necessary precondition for fusing raw sensations into coherent experience”[17]. Kant then asserts the necessity for this ordering and ordered consciousness to also be a consistent synchronic and temporal “unity of consciousness” across the many sensations and experiences being experienced[18]. Thus emerges the concept of the self as a unified and consistent fundamental entity which expresses itself through a faculty Kant calls the will.

Science and ethical behavior after Kant

This new concept of self has moral and scientific and political implications. In particular, as Kant writes, “freedom in the practical sense, is the independence of the will of coercion by sensuous impulses”[19]. Rational morality, according to Kant, is the subjection of this autonomous will to objective moral laws—objective meaning “free from all influence of contingent grounds”[20]. However, for this will to be actually free and therefore actually moral, the will cannot be simply subject to the law “but subject in such a way that it must be conceived also as itself prescribing the law”[21]. Obedience to these non-contingent universal a priori moral laws, regardless of incentives or benefits, is the root of Kant’s deontological moral philosophy. In other words, in contrast to the diffused and permeable self of the Enlightenment, Daston and Galison describe this emerging Kantian moral self as “monolithic and tightly organized around the will [and] posited as free and autonomous (literally ‘giving the law to itself’)”[22].

The scientific ethos associated with this concept of an active and autonomous self (and its accompanying epistemology) can be distinguished from the Enlightenment ethos as such: There is an objective world ‘out there,’ but perception of this world is always mediated through the senses and this ordering consciousness, and therefore this world is never knowable as it is in itself. Because this post-Kantian self “was viewed as overactive and prone to impose its preconceptions and pet hypotheses on the data,” the appropriately objective scientific approach is now one of “self-denying passivity”[23]. In other words, “the only way for the active self to attain the desired receptivity to nature was to turn its domineering will inward—to practice self-discipline, self-restraint, self-abnegation, self-annihilation, and a multitude of other techniques of self-imposed selflessness”[24]. This is the dispassionate scientific objectivity we recognize today.

Ethical Isomorphisms?

As described above, each of these different conceptions of the self also have their own unique epistemology and ethos. These two concepts of self are also each conceptually isomorphic with either genes or the epigenome. The significance of these conceptual isomorphisms is that they demonstrate some of the unique ethical commitments of genetics and epigenetics which are often obscured as scientific differences.

For example, the monolithic, tightly organized, active and autonomous post-Kantian self is a very apt description of the genes. Within the Modern Synthesis genes are conceived of as “information, blueprints, books, recipes, programs, instructions, and further as active causal agents as that which is responsible for putting the information to use as the program that runs itself”[25]. These genes as active causal agents—as exemplified most prominently by Richard Dawkins’ popularization of “selfish” genes, for which organisms are merely vehicles[26]—are identified as both the focus and the impetus of evolution[27]. Further, as expressed by the “central dogma of molecular biology”[28], information necessarily only flows from the gene outwards and not from the environment back into the gene such that genes are functionally isolated and therefore independent from their immediate environments. Thus, genes are post-Kantian as they are also tightly organized, unitary, active and autonomous agents which are likewise the authors of their own laws.

The Enlightenment self as an open, permeable, embedded network of influences with a coherent center but also constantly in flux is a very apt description of the contemporary epigenome. Epigenetic mechanisms are described in contemporary sources as “nongenetic cellular memory, which records developmental and environmental cues”[29] through responses to signals which “come from inside the cell, from neighboring cells, or from the outside world (environment)”[30]. Notably, the changes in phenotypes (i.e., appearances, behaviors, health outcomes, etc.) in reaction to the environment are produced as the result of “several converging and reinforcing signals” that are fundamental for the this ability of both cells and the epigenome as a whole to “‘remember’ past events, such as changes in the external environment or developmental cues”[31]. These scientific descriptions of cellular epigenetics and of the epigenome as a whole sound very much like the “cobweb” self of the Enlightenment “characterized by receptivity rather than assertive dynamism”[32], constantly bombarded by impulses from within and without, whose continuity is guaranteed “only by memory”[33].

Conclusion

The actual genealogical connections between this Enlightenment self and the contemporary epigenome and the post-Kantian self and the gene have only yet been loosely traced. This comparison, though, suggests that as epigenetics and conventional genetics each mirror one of these different historical and philosophical conceptions of self, they each also reflect the accompanying ethical commitments of these different conceptions of self as well. Ultimately, while it remains to be seen to what extent there is a direct connection between the more open and fluid self of the Enlightenment and the contemporary epigenome, or between the fixed and atomistic post-Kantian self and the scientific definitions of genes, this foray into the history of science reveals some of the substantial differences in ethics produced by a concept of atomistic and autonomous essences versus diffuse and permeable essences.

As I discuss in more detail in this post and this postI contend that much of the current resistance to epigenetics, which is usually framed in scientific terms, is actually more a function of these unseen fundamental ethical incompatibilities than about the science. In reality, there are likely far more science-based reasons for the inclusion of epigenetics within genetics than for its exclusion, which actually makes this exclusion that much more of a puzzle, at least on scientific grounds.

Instead of fundamental scientific differences, I propose that the real challenges from epigenetics stem from the interconnectedness and openness it asserts, which run so counter to the basic ethical commitments of contemporary society. By way of comparison, the widespread acceptance and assimilation of conventional genetics likewise suggests that it shares the same basic ethical commitments as contemporary society, which ethical congruence both facilitates this acceptance but also camouflages these ethical commitments of genetics, thereby reinforcing its appearance of objective scientific value-neutrality. 

This post is therefore an effort to further establish more of the history and the parameters of these  fundamental ethical differences. Subsequent posts will further develop the unique ethics and ethical challenges from epigenetics. For even more detail, you can also read my paper-length treatment of the political implications of these fundamental ethical incompatibilities between genetics and epigenetics which I presented at the Association of Politics and the Life Sciences annual conference at Emory University in October 2014 here.

I am curious to hear what you think so far. Are these conceptual isomorphisms indicative of fundamental relationships? Or are they just coincidences? Leave your comments below and I will respond.

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.

[1] Lorraine Daston and Peter Galison. (2007). Objectivity. New York: Zone Books.

[2] Daston and Galison (2007): 197.

[3] Daston and Galison (2007): 205-6.

[4] Daston and Galison (2007): 200.

[5] Daston and Galison (2007): 200.

[6] Daston and Galison (2007): 201.

[7] Daston and Galison (2007): 225.

[8] Daston and Galison (2007): 201.

[9] John Locke. 1995. Essay Concerning Human Understanding. Amherst, NY: Prometheus, Book II Chapter 1 p. 59.

[10] Daston and Galison (2007): 203.

[11] Daston and Galison (2007): 203.

[12] Daston and Galison (2007): 209.

[13] Daston and Galison (2007): 203.

[14] Daston and Galison (2007): 200; 225.

[15] Daston and Galison (2007): 201.

[16] Immanuel Kant (1781). Critique of Pure Reason. J.M. Meiklejohn (trans.)  Buffalo, NY: Prometheus Books, 1990: 77.

[17] Daston and Galison (2007): 201.

[18] Kant (1781): 79; Kant (1781): 77-8.

[19] Kant (1781): 300.

[20] Immanuel Kant (1785). Foundations of the Metaphysics of Morals. Lewis White Beck (trans.). Upper Saddle River, NJ: Prentice-Hall, Inc., 1997: 43.

[21] Kant (1785): 48.

[22] Daston and Galison (2007): 210.

[23] Daston and Galison (2007): 203.

[24] Daston and Galison (2007): 203.

[25] Moss, L. (2003) What Genes Can’t Do, Cambridge: MIT Press.

[26] Dawkins, R. (1989) The Selfish Gene, New York: Oxford University Press.

[27] Wilson, R. (2005) Genes and the Agents of Life: The Individual in the Fragile Sciences. New York: Cambridge University Press.

[28] Crick, F. “Central Dogma of Molecular Biology” Nature 227 (1970): 561-563.

[29] Guy RiddihoughLaura M. Zahn (2010). “What Is Epigenetics?” Science 330(6004), 611.

[30] http://learn.genetics.utah.edu/content/epigenetics/epi_learns/

[31] Roberto Bonasio Shengjiang Tu Danny Reinberg (2010). “Molecular Signals of Epigenetic States.” Science 330(6004), 612-616.

[32] Daston and Galison (2007): 225.

[33] Daston and Galison (2007): 201.

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A Brief History of Epigenetics: Jean-Baptiste Lamarck

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by Shea Robison (@EpigeneticsGuy)

(See also A Brief History of Epigenetics: C.H. Waddington)

See updated posts on Lamarck here, as excerpts from my forthcoming book:

Lamarck’s Actual Lamarckism (or How Contemporary Epigenetics is not Lamarckian)

Lamarckism and the Biology of Discontent in the 1800s

The Unfortunate Legacy of Jean-Baptiste Lamarck

Jean-Baptiste Lamarck

Epigenetics is just now emerging into the scientific and public awareness seemingly out of nowhere. The results from epigenetics raise a number of significant science-based doubts about many of the fundamental tenets of modern genetics. These challenges to the orthodoxy of genetics spark a significant amount of skepticism about—and outright antagonism towards—epigenetics. However, this antipathy towards epigenetics is nothing new. Many of the ideas and concepts revealed by contemporary epigenetics have been around for centuries, and this resistance to epigenetics goes back just as far.

While there are justifiable scientific reservations about many of the unorthodox claims from epigenetics, the underlying model of my overall project is that much of this resistance both in the past and in the present comes from other than scientific reasons. In reality, there are likely far more science-based reasons for the inclusion of epigenetics within genetics than for its exclusion, which actually makes this exclusion that much more of a puzzle. Explaining this puzzle is a main focus of mine, and requires delving into this convoluted and colorful common history between epigenetics and genetics.This is where Jean-Baptise Lamarck (1744-1829) enters the picture.

The first extensively developed theory for how adaptive characteristics acquired during the life of an organism can be passed on to future generations was formulated by Lamarck, and is often referred to as Lamarckism.

The classic example of Lamarckian inheritance is that of giraffe necks elongating as a result of giraffes having to stretch to reach available forage located higher and higher in the trees. This elongation from the stretching of the necks of giraffes in one generation was then passed on to their offspring in the next generation. Giraffes in these subsequent generations were then able to reach even higher into the trees, which raised the height of available forage, thereby requiring even further stretching which was then passed on to subsequent generations, and so on, thus resulting in the very long necks of the giraffe species[1].

However, it bears mentioning that this example was not a central aspect of Lamarck’s theoretical exposition of the mechanisms of inheritance. As described by Ron Roizen, Lamarck’s theoretical treatise regarding inheritance, Philosophy Zoologique, “runs fully 405 pages in its English translation yet it contains only two sentences specifically about giraffes”[2].Thus, the historical and scientific significance of this specific example as a characterization of Lamarck’s thought appears to be greatly exaggerated.

Also, in judging the validity of Lamarck’s theories, there is the issue of what was known and knowable in Lamarck’s time. As Hugh Samuel Roger Elliott writes in his introduction to Philosophie Zoologique, at the time of Lamarck “many of the known facts of evolution might be accounted for either by use-inheritance or by natural selection. If it is true that acquired characteristic are hereditary, then the giraffe might well have developed his neck through that agency. The hypothesis fits the facts.” However, Elliot goes on to also note that “but so also does the hypothesis of special creation,” and also that “similarly, again, natural selection is equally satisfactory as an a priori hypothesis.” In other words, for Lamarck and those of his time, “the facts are covered by at least three different and mutually exclusive hypotheses,” and without the benefit of a posteriori experiments, for example, Lamarck opted for the a priori validity of use-inheritance. What was required next, Elliott observes, was to verify empirically the mechanisms for such inheritances, which neither Lamarck nor anyone else of his time pursued. Still, concludes Elliot, given the specific circumstances of Lamarck “few indeed are the people who are competent to judge of the correct use of deduction in difficult biological inquiries”[3].

Given the march of history, Lamarck’s choice now appears clearly as the incorrect choice. There are a couple of reasons for this, in particular Lamarck’s a priori method of reasoning combined with his philosophical orientation–neither of which are or should be considered indicative of contemporary epigenetics. As Michael Ghiselin writes:

Lamarck’s approach to evolution was that of a metaphysician rather than a natural scientist. It invoked a mystical assumption (the notion that organisms sought “perfection” and tended to become increasingly complex and man-like) which could not be treated scientifically and could not be supported or contravened by evidence. For that very reason, Lamarck’s construct was not a proper theory and was not at all comparable to the theory that Darwin would later present in On the Origin of Species. Darwin’s concept was a well articulated body of scientific thought that could be, and was, tested by recourse to facts. Lamarck’s was not[4].

In other words, making scientific comparisons between Lamarck and Darwin is like making a scientific comparison between…Deepak Chopra and Stephen Jay Gould, which is as fair to Chopra as it is to Lamarck, and as equally unfair to contemporary epigenetics.

Now, while Ghiselin’s point is not to defend Lamarck’s theories of inheritance per se, but rather to demonstrate that “the Lamarck presented in schoolbooks, however, is a fiction — an imaginary figure who has been fashioned from hearsay and wrong guesses, and who has been replicated in countless books by successive teams of plagiarists”[5], my point is to show to what extent the juxtapositions of Lamarckism against Darwinism as a demonstration of the invalidity of non-selectionist theories of evolution and development are little more than crude straw man arguments. As ‘Lamarckism’ is so often used as an epithet against contemporary epigenetics, as a shorthand means to discount the validity of its claims, I hope the preceding shows just how much Lamarckism as such does not apply to contemporary findings like epigenetic inheritance, and therefore should not be used to try and invalidate such science-based claims.

Still, for a while, Lamarckism competed with Darwinism (Darwin himself actually proposed Lamarck-esque mechanisms for the transmission of acquired traits which Darwin called “pangenesis”[6]). However, this Lamarckian “soft inheritance”[7] is now generally dismissed as a quaint pre-Mendelian theory of evolution in lieu of the Modern Synthesis which locates the source of adaptation exclusively in the random mutations of genes. Again, though, even if actual Lamarckism is seriously flawed (which it likely is), this actually has little significance for contemporary epigenetics. However, that Lamarckism as such has been so unilaterally and uncritically and unfairly applied as an invective against epigenetics is, I think, of substantial significance.

The scientific rationale for the rejection of Lamarckian inheritance is largely provided by experiments such as August Weissmann’s tests of whether mutilations of parents (e.g., cutting off the tails of rats) could be passed on to their offspring, and the transplantation of guinea pig ovaries by Castle and Phillips in 1911, which appeared to verify empirically that adaptations of such characteristics were not inheritable[8]. Even though these experimental protocols did not accurately reflect the mechanisms of inheritance as theorized by Lamarck, the results of these experiments were taken as definitive disproof of the possibility of the inheritance of acquired characteristics. (In other words, in this case as well misinterpretations of Lamarck’s theories are used as evidence of the validity of gene-focused selection-based theories which isolate our biological essences from our environments, and the invalidity of alternative explanations which closely link these essences with our environments.) From these experiments was postulated Weismann’s Barrier, a tissue barrier between those cells involved in forming the body of an organism (somatic) and those cells involved in sexual reproduction (germinal). This barrier allegedly protects the germ cells from any type of environmentally-induced change, which therefore prevents Lamarckian inheritance.

Of particular interest at this point, though, is Hugh Samuel Roger Elliott’s observation that, as of 1914, Weissmann’s postulation of this barrier against use-inheritance was as pure and as dangerous a deduction as was Lamarck’s a priori support for use-inheritance:

But while we remain in our present ignorance as to the causes of development, and of the extraordinary and specific influence which the pituitary, thyroid, reproductive and other glands exert upon remote parts of the body, and indeed of many other remarkable correlations existing between apparently disconnected parts, our empirical knowledge is surely far too slight to offer any kind of firm basis for a large reaching deduction like that of Weissmann[9].

In other words, for all the experiments of Weismann, the results did not of themselves justify the blanket denial of use-inheritance as such, but still needed further verification as to the mechanisms involved; all that Weismann was able to show was that the mutilation of parents was not inherited by the offspring. Subsequent research has supported the presence of Weissman’s Barrier, but the actual mechanisms of this Barrier are even just now being identified[10]. The recency of these verifications suggests that other mechanisms which operate outside of Weissman’s Barrier may yet be discovered, and there is suggestive evidence that these other epigenetic mechanisms are also being found[11]. In the comments to this post, Abhay Sharma has provided links to recent research that demonstrates the mechanisms of the transmission of information from the soma to germline in plants and mammals.

I suggest that there are underlying ethical and political reasons for the acceptance of the provisional findings of Weissman and the others which fit the emerging paradigm of isolated essences. These early choices as to which line of research to pursue necessarily influenced the subsequent development of the theory. Further discussion of these underlying motivations will be conducted in subsequent posts, but the specific historical trajectory of the development of the Modern Synthesis and of conventional genetics, with the exclusion of epigenetic inheritance, is an important point to keep in mind.

In the 1950s the theoretical importance of this isolation of our biological essences from our environments is revealed again when Francis Crick and James Watson asserted the “central dogma of molecular biology” that DNA is also insulated from the vagaries of the environment[12]. According to the central dogma, “the chemical language of genetic information in the form of DNA sequences can be directly copied into complementary nucleic acid base sequences termed RNA which in turn can then be translated into a protein sequence of amino acids, a quite different chemical language”[13].  This chemical language is noncommutative, though, which means that “genetic information never flows in reverse from a sequence string of amino acids into a complementary sequence of DNA or RNA bases”[14]. In other words, DNA is not influenced by its environments, which means there is no adaptation of DNA, and that genetic variation only happens randomly. Thus, barring very rare and random—i.e., accidental and uncontrollable—genetic mutations, each person is born with a pristine normal copy of the human genome.

This assumed isolation of our biological essences—although arrived at through scientific processes—is a central tenet of the modern theory of genetics. However, this insulation of our essences from our environments also has significant, though often overlooked, ethical implications, especially in light of this blanket and facile (and now highly suspect) dismissal of all non-selectionist theories through comparison with Lamarckism. The motivations behind these comparisons with Lamarckism are one way history, philosophy and ethics enter the ostensibly exclusive scientific domain of genetics. The elaboration of these ethical implications will be the focus of subsequent posts, such as Epigenetics and the Extended Synthesis and The Genetics of the Ethics of the Science of Genetics.

Why has this fundamental misconstrual of Lamarck played such a significant role in the dismissal of epigenetics, if not at least in part for political and/or ethical reasons? I am curious to hear what you think. Leave your comments below and I will respond.

For further discussion of the convergence of politics and science around Lamarck’s time, see also The History of Epigenetics and the Science of Social Progress. For discussion of the politics and science of epigenetics in the 20th century, read A Brief History of Epigenetics: C.H. Waddington,  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.

[1]Moore, J. N. 1970. Biology: a search for order in complexity. Grand Rapids: Zondervan Pub. House.

[2] Roizen, Ron. 1971. “The Argument of  Philosophy Zoologique.” http://www.roizen.com/ron/Lamarck.htm.

[3] Lamarck, Jean-Baptiste. [1809] 2011. Zoological Philosophy: An Exposition with Regard to the Natural History of Animals. Cambridge: Cambridge University Press, xlii.

[4] Ghiselin, M. 1994. “The Imaginary Lamarck:A Look at Bogus “History” in Schoolbooks.” The Textbook Letter 5(4). http://www.textbookleague.org/54marck.htm.

[5] Ghiselin, M. 1994. “The Imaginary Lamarck:A Look at Bogus “History” in Schoolbooks.” The Textbook Letter 5(4). http://www.textbookleague.org/54marck.htm.

[6] Geison, G. L. (1969). “Darwin and heredity: The evolution of his hypothesis of pangenesis”. J Hist Med Allied Sci XXIV (4): 375–411.

[7] Mayr, Ernst. 1980. “Prologue: Some Thoughts on the History of the Evolutionary Synthesis.” In The Evolutionary synthesis: perspectives on the unification of biology, eds. William B. Provine and Ernst Mayr. Cambridge: Harvard University Press.

[8] E.J. Steele. 1999. Lamarck’s Signature : How Retrogenes Are Changing Darwin’s Natural Selection Paradigm. Basic Books.

[9] Lamarck, Jean-Baptiste. [1809] 2011. Zoological Philosophy: An Exposition with Regard to the Natural History of Animals. Cambridge: Cambridge University Press, xliv.

[10] Solana, J. (2013). Closing the circle of germline and stem cells: the Primordial Stem Cell hypothesis. EvoDevo4(1), 1-17.

[11] Sharma, A. (2013). Transgenerational epigenetic inheritance: focus on soma to germline information transfer. Progress in biophysics and molecular biology,113(3), 439-446.

[12] Francis Crick. 1970. “Central Dogma of Molecular Biology.” Nature 227(5258):561-3.

[13] E.J. Steele. 2000. “The Evidence for Lamarck.” Quadrant 364(44): 47-56.

[14] E.J. Steele. 2000. “The Evidence for Lamarck.” Quadrant 364(44): 47-56.

A Brief History of Epigenetics: C.H. Waddington

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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.

Epigenetics and the Extended Synthesis

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by Shea Robison (@EpigeneticsGuy)

Epigenetics is just one aspect of what is being called the Extended Synthesis of evolutionary biology[1], in distinction to what is referred to as the Modern Synthesis of evolutionary biology. This Modern Synthesis is the fusion of Mendelian genetics with Darwinian evolution as interpreted through the theory and practice of population (i.e., statistical) genetics[2]. This integration of ideas is called the Modern Synthesis because until the synthesizing work of Theodosius Dobzhansky and others during the 1920s, 1930s and 1940s, Mendelian genetics and Darwinian evolution were actually competing and seemingly irreconcilable paradigms[3].

The major tenets of the Modern Synthesis can be summarized as 1) that populations contain genetic variation which arises by random (i.e., not adaptively directed) mutation and recombination; 2) that populations evolve by changes in gene frequency through random genetic drift, gene flow, and especially natural selection; 3) and that most individual phenotypic effects are very slight, so that most phenotypic changes are very gradual[4].

The most salient aspects of the Modern Synthesis in the context of this post are first, the emphasis on populations and not on individuals in describing variation; second, the emphasis on randomness as the source of variation; third, the lack of adaptiveness (i.e., lack of responsiveness to the immediate environment) which results from this assumption of randomness as the source of variation; and, finally, that phenotypic variation is very slight and very gradual. Epigenetics poses significant challenges to each of these basic. assumptions

While each of these tenets was derived through scientific methods, for all their scientific pretenses each of these assumptions have a long non-scientific history which predates their incorporation as tenets of the Modern Synthesis. These tenets and their attendant assumptions constitute a very specific ethical framework which allows certain interpretations of behaviors and outcomes and excludes others. However, this framework and these ethical implications of these scientific assumptions are often not recognized as such. What is this underlying ethical framework of genetics and what are the politics which result from this framework will be discussed, but the main point is that these ‘scientific’ assumptions are actually the doorway through which ethics and politics steal into the henhouse of science.

The Extended Synthesis in one sense, as indicated by the name, is properly understood as the extension of the Modern Synthesis and not its replacement. As explained by Massimo Pigliucci:

Let me again be clear on a fundamental point underlying this whole discussion: one can reasonably argue that none of this contradicts any tenet of the [Modern Synthesis], although it seems to me at least reasonable to concede that the new concepts and empirical findings…may eventually force a shift of emphasis away from the population genetic-centered view of evolution that characterizes the [Modern Synthesis][5].

In this sense at least the Extended Synthesis and its constituent parts are not fundamental challenges to genetics. In particular, according to Pigliucci, this shift of emphasis merely involves the incorporation into conventional genetics and developmental biology of concepts such as evolvability, phenotypic plasticity, epigenetics, complexity and the nonlinearity of adaptation in high-dimension adaptive landscapes[6].

However, this point of view does not consider the ethical implications of these extensions of conventional genetics. At the level of science, it is probable that these ideas can eventually be incorporated into the theoretical structure of genetics without too much difficulty, but this does not consider the possible ethical and political issues that might be involved. In fact, there are likely far more science-based reasons for their inclusion than for their exclusion, which actually makes the exclusion of fields like epigenetics from genetics that much more of a puzzle: If there are so many science-based reasons for the inclusion of epigenetics within genetics, why has it been so maligned for so long? A plausible answer is that there are even stronger non-scientific reasons for this exclusion of epigenetics, and my working theory in this post is that these non-scientific reasons are a function of key differences in the ‘hidden’ ethical commitments of epigenetics and conventional genetics. Thus, again the focus of this post is on first revealing the unseen ethical commitments behind conventional genetics, then elaborating the ethical commitments of epigenetics, and analyzing the political implications of these new ethical challenges presented by the (re)emergence of epigenetics.

Also, at this point I do not know if the other components of the Extended Synthesis share the same background ethical commitments as epigenetics and therefore constitute a coherent ethical whole as does the Modern Synthesis, or if the ethical commitments of these different components are unique from each other, or much else about the ethical landscape of the Extended Synthesis. My assumption is that the underlying ethics of these other components of the Extended Synthesis will at the least be more similar to those of epigenetics than of conventional genetics, but this is work that will have to wait.

I am curious to hear what you think. Leave your comments below and I will respond.

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.

[1] Pigliucci, M. & Muller, G. (2010). Evolution: The Extended Synthesis. Cambridge, Mass.: MIT Press.

[2] Mayr, E. (1980). The Evolutionary synthesis: perspectives on the unification of biology. Cambridge, Mass.: Harvard University Press.

[3] Burian, R. (2005). The Epistemology of Development, Evolution, and Genetics. Cambridge: Cambridge University Press. See also Bowler, P. J. (1983). The Eclipse of Darwinism. Baltimore: The Johns Hopkins University Press; Bowler, P. J. (1988). The Non-Darwinian Revolution. Baltimore, MD: Johns Hopkins University Press; Bowler, P. J. (1989). The Mendelian Revolution. Baltimore, MD: Johns Hopkins University Press; Depew, D. J., and B. H. Weber (1995). Darwinism Evolving : Systems Dynamics and the Genealogy of Natural Selection. Cambridge, MA: MIT Press; Gayon, J. (1998). Darwinism’s Struggle for Survival: Heredity and the Hypothesis of Natural Selection. Translated by M. Cobb. Cambridge: Cambridge University Press; Provine, W. B. (1971). The Origins of Theoretical Population Genetics. Chicago: Chicago University Press.

[4] Futuyama, Douglas (1986). Evolutionary Biology. Sunderland, MA: Sinauer: 12.

[5] Pigliucci, Massimo (2007). “Do We Need an Extended Evolutionary Synthesis?” Evolution 61(12): 2743-2749.

[6] Pigliucci, Massimo (2007). “Do We Need an Extended Evolutionary Synthesis?” Evolution 61(12): 2743-2749.

The Genetics of the Ethics of the Science of Genetics

Me2

by Shea Robison (@EpigeneticsGuy)

To begin, the basic assumptions of the science of genetics are not unique to contemporary genetics but are actually the products of a very specific intellectual and cultural history. This history goes back all the way to the metaphysics of Aristotle as filtered through the Medieval theology of St. Thomas Aquinas and the natural and political philosophy of John Locke which had such a significant influence on the Founding Fathers of the United States of America. From this common intellectual history were born both the philosophical concept of equally created persons as the ethical focus of modern politics as well as the scientific description of genes as pristine copies of our common human essence.

That the assumptions of genetics are contingent on these historical and philosophical precedents likely seems trivial to laypersons and nonsensical to geneticists. Genetics, as a product of modern science, exhibits the patina of scientific objectivity which appears to negate these historical and cultural influences. However, one of the unanticipated benefits of this recent (re)emergence of epigenetics is how it exposes these hidden ethical assumptions of genetics—which are themselves reflections of the prevailing ethics of our time—against which epigenetics proposes its own unique ethics. These novel ethical and political implications of epigenetics are in many ways prior to and perhaps even more significant than its scientific implications.

In other words, in posing the significant challenges to the science of genetics that it does, epigenetics also poses equally significant ethical and political challenges. The big difference, though, is that while these scientific challenges are readily apparent, these ethical and political dimensions are not—primarily because these ethical and political implications are often masked by the obvious scientific differences.

While modern scientific methods on their own—in a vacuum, as it were—would eventually achieve an appropriately scientific resolution to the current discrepancies between genetics and epigenetics, science is not ahistorical and is not exempt from the prevailing politics and ethics of its time. Because the influence of these non-scientific biases are often not recognized as such, scientific positions often function as unconscious proxies for underlying ideologies, and ideological differences cannot be resolved through scientific methods but are rather contestable as politics.

That the scientific and ethical and political are all knit from the same cloth suggests two things: First, given the widespread acceptance of genetics, that the scientific assumptions of genetics dovetail well enough with the basic assumptions of our prevailing ethics and politics that there are no irreconcilable differences between them. Second, that much of the scientific resistance against epigenetics stems from the ethical exceptionalism of epigenetics[1]. In such circumstances, for the methods of science to work to their intended effect scientific claims must be untangled from the underlying ethics as much as possible so that scientific problems can be treated scientifically and ethical claims can be dealt with politically. Thus, an important component in the science of epigenetics should be to untangle the scientific from the ethical, and this post is one such effort.

Even in the ‘hard’ domains of the natural and life sciences much of the contest over the validity of epigenetics is a process of narrative formation, or an ongoing contest over how epigenetics is talked about and perceived both within and outside of its scientific contexts[2]. This is a critically important distinction to make because instead of being simply true or false, narratives are—like the epigenome itself—malleable and highly susceptible to internal and external influences at critical moments in their development.

While scientific content is obviously a key component in the formation of science-based narratives, so are the ethical and political aspects (e.g., the controversy around vaccinations and autism, or the ongoing global warming/climate change debate). For example, the coevolution of epigenetics and genetics during the Cold War was significantly influenced by the prevailing political ideologies of the time[3], the effects of which still reverberate today. However, the narratives of epigenetics during the Cold War are not the narrative of epigenetics today, and for any number of reasons what are the narratives of epigenetics today will likely not be the narratives of epigenetics tomorrow.

Awareness of the malleability of narratives and of how this process of narrative formation works could be invaluable for practitioners and proponents of epigenetics in influencing or nudging the emerging narrative of epigenetics towards more conducive and politically acceptable forms. This post is one part of a greater project to flesh out the political and ethical implications of epigenetics. My working theory in this project is that many of the scientific criticisms of epigenetics are ultimately political and ethical in origin, and that these scientific disagreements will not be resolved until these underlying ethical complications from epigenetics are at least acknowledged and addressed. Elaborating these ethical challenges from epigenetics through an analysis of fundamental concepts in political philosophy is the primary purpose of this post.

Even though my primary domain in this project is largely philosophical, my goal is very practical: To provide practitioners and proponents of epigenetics with the tools they need to better understand the role of some of these additional non-scientific factors in the acceptance of or resistance to epigenetics.

I am curious to hear what you think. Leave your comments below and I will respond.

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.

[1] Mark Rothstein (2013). “Epigenetic Exceptionalism.” Journal of Law, Medicine and Ethics 41(3): 733-736.

[2] Bruner, Jerome. 1991. “The Narrative Construction of Reality.” Critical Inquiry 18 (1): 1–21; Roe, Emery. 1994. Narrative Policy Analysis: Theory and Practice. Durham, CT: Duke University Press; Jones, Michael D. and Mark K. McBeth. 2010. A Narrative Policy Framework: Clear Enough to Be Wrong? The Policy Studies Journal, 38(2): 329-353.

[3] Sapp, J. (1994). Symbiogenesis in Russia. In Evolution by association a history of symbiosis. New York: Oxford University Press; Gaissinovitch, A. E. (1980). The origins of Soviet genetics and the struggle with Lamarckism, 1922-1929. Journal of the History of Biology, 13(1),1-51; DeJong-Lambert, William. 2007. “The Cold War Politics of Genetic Research.” Presented at The7th Annual Havighurst Center for Russian & Post-Soviet Studies International Young Researchers Conference, Miami, OH; Graham, L.R. (2004). Science in Russia and the Soviet Union. A Short History. Series: Cambridge Studies in the History of Science. Cambridge University Press; Wrinch, P. N. (1951). Science and politics in the USSR: the genetics debate. World Politics, 3(04), 486-519; Adams, M. B. (1991). “Through the looking glass: The evolution of Soviet Darwinism.” In New Perspectives on Evolution, edited by L. Warren and H. Kropowski, New York: Wiley-Liss, 37-63.