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New Genetics and Society

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Epigenetics for the social sciences: justice,embodiment, and inheritance in the postgenomicage

Maurizio Meloni

To cite this article: Maurizio Meloni (2015) Epigenetics for the social sciences: justice,embodiment, and inheritance in the postgenomic age, New Genetics and Society, 34:2,125-151, DOI: 10.1080/14636778.2015.1034850

To link to this article: http://dx.doi.org/10.1080/14636778.2015.1034850

Published online: 12 May 2015.

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Epigenetics for the social sciences: justice, embodiment,and inheritance in the postgenomic age

Maurizio Meloni∗

Institute for Advanced Study, School of Social Science, Princeton, USA

(Received 11 May 2014; final version received 10 December 2014)

In this paper, I firstly situate the current rise of interest in epigenetics in thebroader history of attempts to go “beyond the gene” in twentieth-centurybiology. In the second part, after a summary of the main differences betweenepigenetic and genetic mutations, I consider what kind of implications the suigeneris features of epigenetic mutations may have for the social sciences. Ifocus in particular on two sites of investigation: (a) the blurring of theboundaries between natural and social inequalities in theories of justice andtheir possible implications for public policy and public health and (b) adeepening of the notion that the constitution of the body is deeply dependenton its material and socially shaped surroundings (“embodied constructivism”).In conclusion, I advance some cautionary reflections on some of the (knownand unprecedented) problems that the circulation of epigenetics in widersociety may present.

Keywords: epigenetics; natural/social inequalities; public health implications

Introduction

Epigenetics represents one of the key terms with which to grasp the profile of thenew biological landscape that has taken shape in the so-called postgenomic age(so-called because it is conventionally used to define the period dating after thecompletion of the Human Genome Project in 2003). Of this postgenomic age,with its tensions and promises, hypes and controversies, epigenetics is a perfectincarnation and an excellent theoretical spyglass through which to see the changingthought-style (and possibly ethos) of the biosciences in this early twenty-firstcentury.

Epigenetics was originally conceived by embryologist Conrad H. Waddington(1905–1975; see Slack 2002) as the investigation of the unfolding of geneticmaterial into a final phenotype (1942; see Holliday 1990, 2006). Today, it isincreasingly understood in a molecular sense, as “the study of changes in genefunction that are mitotically and/or meiotically heritable and that do not entail a

New Genetics and Society, 2015Vol. 34, No. 2, 125–151, http://dx.doi.org/10.1080/14636778.2015.1034850

∗Emails: meloni@ias.edu, m.meloni@exeter.ac.uk

# 2015 Taylor & Francis

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change in the sequence of DNA” (Armstrong 2014). In the context of a postge-nomic proliferation of – omics studies (genome, microbiome, transcriptome, expo-some, etc.) – epigenetics’ contribution is the study of the epigenome, the set of thepotentially “heritable changes in gene expressions that occur in the absence ofchanges to the DNA sequence itself” (Dolinoy and Jirtle 2008).

In this article, I have a threefold goal. First, I situate the last decade’sexplosion of interest in epigenetics within a broader perspective that takes inthe tensions and persisting challenges to notions of hard-heredity in the historyof biology, and genetics in particular. Second, I investigate some of the possibleimplications of epigenetics for social science and wider society. I will look inparticular at two ossified conceptual dichotomies in political and social theorythat are likely to be challenged by the rise of epigenetics: (a) the boundarybetween natural and social inequalities in theories of justice and (b) the opposi-tion between biomedical and social constructionist views of the body. Finally, Iwill address some of the counterintuitive implications that claims of a replace-ment of hard-heredity (heritable material impervious to direct environmentalinfluences) with an epigenetic or soft view of inheritance (heredity is malleable,modulated by direct environmental variations) may have on public policy andpublic health.

The context: from the crisis of the gene to the “reactive genome”

The quest for mechanisms of nongenetic inheritance is far from new in the historyof biology. Repeated attempts to go “beyond the gene” (Sonneborn 1949; Sapp1987) and search for extrachromosomal forms of inheritance (Harwood 1993)have characterized twentieth-century biology more than textbook histories ofbiology allows to think. Far from being an ideological return to Lysenko, as polemi-cally claimed (Maderspacher 2010), the current rise of interest in the mechanismsof epigenetic inheritance can be better seen as the last symptom of a longer termcrisis in the notion of the gene with important implications for modern evolutionarytheory and a remaking of the notion of heredity.

Postgenomic genome

A crisis in the notion of the gene is nothing new. The gene has always been aconcept in “tension” (Falk 2003), situated in a complex history of experimental,technological, and conceptual settings (Beurton et al. 2003; Rheinberger andMuller-Wille 2010). However, what has happened after the completion of theHuman Genome Project has made this crisis even more conspicuous.

To paraphrase Portin (2002, 274), our current postgenomic understanding hasoutgrown conventional knowledge of the gene. In a postgenomic context, whereit is becoming harder to maintain a realist and particulate notion of the gene(Barnes and Dupre 2008), what is increasingly brought to light is the reactivity

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of the new postgenomic genome to environmental signals (Keller 2012, 2014; seealso Gilbert 2003; Griffiths and Stotz 2013). The genome is best described today asa “vast reactive system” (Keller 2012), a mechanism “for regulating the productionof specific proteins in response to the constantly changing signals it receives fromits environment” (Keller 2014, 2427).

This shift in focus from the centrality of DNA to its broader regulatory (cellularand environmental) context may open the way to a different view of evolution andinheritance. Whereas the “transmission genetics” model (Mameli 2005; Amundson2005) saw heredity merely as the simple passage of DNA from one generation toanother (see also: Muller-Wille and Rheinberger 2012), we are today increasinglyencouraged to look at the fact that “heredity involves more than DNA”, and thatvariations arising during development may be inherited (Jablonka and Lamb2008). The same so-called central dogma of molecular biology, which states theunidirectional nature of the flow of information from DNA to protein (Crick1970), is increasingly challenged on the ground that information can also go inthe reverse direction (Dupre 2012). It is in this turbulent intellectual context thatthe current wave of interest in epigenetics has to be situated (Jablonka and Raz2009).

Enter epigenetics

Bases and mechanisms

To understand epigenetics, it is a good idea to start from the biochemical bases ofthe process, namely the fact that, in eukaryotic cells, DNA is tightly wrapped intochromatin and that modifications of the chromatin structure can affect DNAexpression (Meaney and Szyf 2005; Feil and Fraga 2012).

DNA methylation (the most recognized mechanism of epigenetic mutations, andstudied since the late 1960s, see Holliday 2006, see however Daxinger and White-law 2012) is one such change by which the attachment of a methyl group to a DNAbase results in chromatin de-activation and inhibition of gene transcription. Methyl-ation works therefore as a sort of “physical barrier to transcription factors” (Gluck-man et al. 2011) and is regulated by nutritional and environmental factors,especially during phases of early development (Dolinoy and Jirtle 2008).

Further epigenetic mechanisms include histone modification and regulation bynon-coding RNA, although they are considered less stable than methylationpatterns.

There are several excellent examples of very visible phenotypic changes in theanimal kingdom driven by changes in methylation patterns as a result of differentenvironmental exposures and nutritional inputs. For instance, in honeybees (Apismellifera), genetically identical larvae following differential feeding (royal jellyversus less rich food) produce alternative adult phenotypic (sterile worker andfertile queen: Kucharski et al. 2008).

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In experimental contexts, the most well-known case of nutritional control ofalternative phenotypic outcomes is probably the switching on and off of theagouti gene in mice that is obtained by feeding a rich in methyl-donor diet to preg-nant agouti mice. Exposing the pregnant mouse to lack of methyl-rich donor dietresults into hypomethylation (and enhanced expression) of the promoter of theagouti gene. This brings to highly visible phenotypic changes because offspringare no longer slim and brown, but yellow, fat, and prone to diabetes (Waterlandand Jirtle 2003).

If we move from nutritional to behavioral exposures, the most well-known caseis Meaney’s group study on how high versus low licking and grooming behaviorsin rats alter the methylation patterns of the promoter of the glucocorticoid receptorin pups (Weaver et al. 2004; see an updated review in Lutz and Turecki 2014).

The most hotly debated issue in epigenetics concern probably the stability andevolutionary significance of transgenerational epigenetic inheritance (Richards2006; Jablonka and Raz 2009; Daxinger and Whitelaw 2010, 2012; Grossniklauset al. 2013; Lim and Brunet 2013; Heard and Martienssen 2014). To clarifywhat is at stake here, it is important to distinguish two alternative, but often con-fused, aspects that go under the label of epigenetic transgenerational effects: (a)germ line inheritance (where the epigenetic signature is not entirely cleared ingametogenesis and can be transmitted through the germ line: Chong and Whitelaw2004; see Anway et al. 2005) and (b) non-germ line, experience-dependent epige-netic inheritance (where the epigenetic signature is re-established in each succes-sive generation by the reoccurrence of the “behaviour or environment thatinduces the mark”, also known as “niche recreation”: Gluckman et al. 2011) asin the case of the Meaney’s group studies above mentioned.

The possibility of transgenerational epigenetics phenomena in humans remainsvery controversial. There are very well-known epidemiological studies on thetransgenerational effects of chronic disease in individuals prenatally exposed tofamine during the Dutch Hongerwinter (Hunger Winter) in 1944–1945 duringGerman occupation (Heijmans et al. 2008; Painter et al. 2008), as well as studieson the link between ancestors’ access to food and longevity drawn from the histori-cal record of the Overkalix cohorts in Northern Sweden (Bygren, Kaati, andEdvinsson 2001).1 These studies establish connections between malnutrition inutero and in early life, and persisting metabolic disorders (up to six decades laterthat is including the second generation, Painter et al. 2008) via changes in thehuman epigenome.

The debate, however, on the mechanisms, significance, and magnitude of thesetransgenerational2 effects remains very much open, given the fact that the erasureof methylation marks between generations is what is in principle to be expected inmammalian development (although not a universal phenomenon in other organisms,and not complete even in mammals, see Richards 2006, and Lim and Brunet 2013, on“unconventional modes of inheritance”).3 Cautiousness is certainly necessary, asscientists remind us that there is still a “long way to go” to fully understand “the

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involvement of epigenetics in environmentally triggered phenotypes and diseases”(Feil and Fraga 2012, 107; see also Heard and Martienssen 2014).

The uniqueness of epigenetic mutations

Scientific controversies and hype undoubtedly surround epigenetic research(Meloni and Testa 2014). Speculation on such a scientifically unstable field maybe seen as premature. However, recognition of exaggerated claims and controver-sial issues is not a sufficient reason to shy away from the potential of epigeneticresearch, especially when understood in sophisticated postgenomic frameworks.My argument is that what we already know about epigenetics and the new reactiveview of the genome that is emerging from postgenomics offer us enough concep-tual elements to think in a different way the separation of the human world into a“biological” and a “social” domain that was characteristics of the century of thegene (and a reflex of its hard-hereditarian framework). Before assessing some ofthe distinctive implications of epigenetics for the social sciences, it is importantto recap the uniqueness displayed by epigenetic mechanisms when compared togenetic ones.

As the following table indicates (elaborating on Jablonka and Lamb 1995; Randoand Verstrepen 2007; Loi, Del Savio, and Stupka 2013; Landecker and Panofsky2013), the crucial differences between epigenetic and genetic mutations can besummed up in seven dimensions:

Epigenetic mutations Genetic mutations

(1) Sensitivity to the environment in short socialtime span

(1) Unresponsive to direct environmental signals(with exceptions of accidental effects, such asmutations due to exposure to X-rays, nuclearradiations, etc.)

Epigenetics as a mechanism for flexible anddynamic responses to the solicitations from achanging environment

Directed variation as a consequence of a“specific environmental agent inducingspecific and predictable heritable changes”(Jablonka and Lamb 1995)

Random mutations: “independent of selectivepressure” (Rando and Verstrepen 2007); onlydue “to the imperfections of the copy-systemor to non-directed effects of environmentalfactors” (Jablonka and Lamb 1995)

The randomness of epigenetic variations isquestioned (Jablonka and Lamb 2005; Randoand Verstrepen 2007). Non-random, althoughnot necessarily adaptive responses (Jablonkaand Lamb 1995)

(2) Potential reversibility through practice,lifestyle or therapy (e.g. pharmacologicalintervention) and other post hoc interventions(Szyf 2009)

(2) Not affected by lifestyle in “short” social time,only over a long evolutionary timescale (e.g.gene-culture coevolution: lactase persistencein specific human populations with increasednutrition from dairy)

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(3) Early programming (3) Insensitive to in utero or post-natal events(with the exceptions of events indicated in 1)

In utero or perinatal events set stably epigeneticmarks

No link between early-life events and geneticsequences

Embedding of early-life experiences and early-life environment in the genome via epigeneticmechanisms (Champagne and Curley 2009;Szyf and Bick 2013)

Genome-wide effects of epigenetic responses toearly-life adversity (McGowan et al. 2011)

(4) Short-term adaptive flexibility (4) Long-term adaptive flexibilityStable enough to allow intergenerational

transmissibility over a certain (limited)number of generations

Inherited features acting only over longevolutionary timescale/countless generations

Many ecological challenges are too transient tobe “dealt with effectively by changes in genefrequencies”, but “too chronic to be efficientlybuffered by homeostasis or allostasis”: hence,importance of an epigenetic adjustment tovariable environmental and behavioralexperiences (Kuzawa and Bragg 2012; seeJablonka and Raz 2009)

Thousands of years before fixing a change in thegenetic pool

(5) Tissue and cell specificity (differentepigenetic marks depending on different cells)

(5) One body one genome (although this isincreasingly challenged: chimerism,mosaicism, see Dupre 2012, Chap. 7)One body many epigenomes (Wade 2009)

(6) Time dependency: epigenetic marks willchange depending on the time a sample istaken

(6) One-time DNA sample invariant for life

Age-related changes No change of DNA during life(7) Unit of inheritance: broad (7) Unit of inheritance: narrowThe whole cellular architecture, or gene’s

phenotype (Jablonka and Lamb 1995)including DNA, chromatin structure, etc.

The nucleotide sequence

It is building upon this specificity and singularity of epigenetic phenomena that itis possible to anticipate some of the possible reconfigurations of the relationshipbetween the biological and the social in a postgenomic scenario.

A significant body of scholarship has already started to outline the implicationsof these anticipated differences between the specific features of this nongeneticpathway of inheritance vis-a-vis a genetic one. Many of the points in the tablehave already received a significant level of attention in social science investigation.Issues of extended personal, social, and legal responsibility arise from points 1–2(Rothstein, Cai, and Marchant 2009; Dupras, Ravitsky, and Williams-Jones 2014;Hedlund 2012). Increasing attention and intervention on the maternal body are aneffect of point 3 which clearly situates mothers and their behaviors at the center ofepigenetic attention (Richardson, forthcoming; Richardson et al. 2014), often inter-laced with racial themes (Mansfield 2012). Risk of classifying sub-populations with

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different epigenetic marks because of the reproduction/perpetuation of hard socialconditions (Katz 2013 ; Meloni 2014; Meloni and Testa 2014) is a consequence ofpoint 4. And the issue of a new class of sensitive information in need of privacyprotection arises from point 5 (Rothstein, Cai, and Marchant 2009).

In the second part of this article, I will therefore direct my attention to two otherpossible implications of epigenetic research for social and political science, whichhave received relatively less attention so far. In conclusion, I will raise a prudentwarning about some unforeseen implications that the circulation of epigeneticsin wider society may present.

Epigenetics and political theory: blurring the boundaries between naturaland social inequalities

How much is genetic luck really due to luck?

The first area in which I would like to measure the social implications of epigeneticsresearch may seem speculative, but has important implications in the way notionsof inequalities and theories of justice are constructed.

Differences between “natural” and “non-natural” features of human experienceare amongst the most deeply rooted assumptions in Western metaphysics. Thisdichotomy has been visible since Antiquity in both medical thinking (Galen) andtheories of justice (Plato, Aristotle). In both fields, the separation between“natural” and “not natural” plays the same powerful function to distinguishingwhat is within human control and what is outside it.

In medical thinking, categories like “naturals” and “non-naturals” were stillinfluential until early modernity to mark the boundaries between non-mutableand modifiable aspects of human life: temperament and constitution for the first,nutrition, and climate for the latter (Beltran 2002; Muller-Wille and Rheinberger2012).

In socio-political thought, we see at stake a similar dichotomy (Beteille 1983;Lewens 2010). In Plato and Aristotle, notions of natural inequalities perform akey function in sustaining a stratified view of society. From its classical origin,the notion of natural inequality represents what lies outside the domain of justiceand human intervention. If an inequality is rooted in nature, there is less (or no)reason to intervene. In various forms, such a way of thinking has persisted untilthe twentieth century, with very few exceptions. Theologically oriented traditionsin political theory that have built the conception of nature upon notions of deityhave further performed this function (Daston and Vidal 2007).

What is particularly striking is that this idea of a state of nature as radically priorto, and uncontaminated by, social relationships has found a novel legitimation inmodernity through the incorporation of biological arguments into public affairs.Talks of innate, natural or congenital differences between races, classes, orgenders stemming from biological thinking have re-legitimized ideas of naturalinequalities in politics (Beteille 1983).

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In particular, political theorists in the twentieth century have referred explicitly togenetics to provide an example of something that is separated or prior to the pol-itical domain. Genetics has been often taken as a perfect case of a diverse distri-bution of talents (the so-called genetic lottery) on which justice has no reason tointervene or, at the very least, morality rather than justice can be invoked tojustify some sort of rectification. In large areas of political theory, with the excep-tion probably of some egalitarian or consequentialist thinkers,4 genetic disadvan-tage is seen as a natural asset whose unfortunate effects can be mitigated orneutralized in a decent society but on which there is no obligation to intervene.

The bottom line argument in much political philosophy seems to be, as Lewenswrites in an insightful article that when, as in the case of genetics, “inequality isnobody’s fault, it is not the concern of justice to correct it” (2010, 265). As Bucha-nan and colleagues have remarked, conventional thought in political theory is that:

nature, or the natural, is often thought to be not only that which is given but also thatwhich must be accepted as beyond human control. In that sense, to say that some-thing is due to nature is to relegate that to the realm of fortune or misfortune,rather than justice or injustice. ( . . . ) It is not surprising, then, that to a large extenttraditional thinking about justice has associated natural disadvantages with misfor-tune rather than injustice, since there was little or nothing that could be done toprevent them. (2000, 83, my italics)

Tom Nagel’s classical article on Nature and Justice is very representative of thisway of thinking for which society has to be seen as less “accountable for thoseinequalities in whose generation nature plays a central role” (1997, 305; Lewens2010). It is not by chance that Nagel picks as a key example a degenerativeillness produced by a defective gene to make the argument for the differentweight that should be assigned to social and natural inequalities, that is, to makea distinction between injustice and bad luck (or “natural unfairness”). WhatNagel hints at are spheres of human life that “have nothing to do with justice,and that are not mandatory in the same way” (1997, 303): genetics would be oneof these cases.

Nagel may represent here a particularly conspicuous example of this social/natural distinction. However, he is not the only one to take this distinction as sig-nificant. In Rawls for instance, such a dichotomy between the natural and social isalso taken for granted. As he famously wrote in A Theory of Justice, “The naturaldistribution is neither just nor unjust” (Rawls 1971, 87). His semi-consequentialismimplies that “natural primary goods, such as ‘health and vigour, intelligence andimagination’, fall outside the master patterns by which feasible alternative basicstructures are assessed” (Pogge 1989, 64; Lewens 2010). More recently, inanother very influential article, Anderson (1999, 331) claims that the “distributionof natural assets” has to be seen out of a “democratic equality” framework. Only theresponse of institutions to this distribution, she believes, is politically relevant, butnatural assets are not as such socialized. And many other examples could be given.

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What shall we do with the idea of natural inequalities in an epigenetic age?

Over the last decades, philosophers of biology have convincingly illustrated theconceptual confusion of the innate/acquired distinction (Griffiths 2002). In thelight of a developmentalist view of biology and genetics, it is easy for them toargue against any neat distinction between the “effects of natural differencesfrom effects of social differences” (Lewens 2010, 271).

The first argument I want to make here, expanding in particular on an importantarticle by Loi, Del Savio, and Stupka (2013), is that epigenetics makes the inappro-priateness of the natural/social divide for political theory highlighted by Lewenseven more flagrant. Epigenetics provides “a chain of connections” (Loi, DelSavio, and Stupka 2013, 143) between what we used to think of separately as“social” and “natural” inequalities, radically cutting across the distinctionbetween these two domains. From what we have said in the first part of thearticle, epigenetics seems to point to the existence of a double social opening inthe architecture of the gene. Upstream, genetic expression is regulated by the incor-poration of certain environmental exposures; downstream, genetic expression canbe reversed via social and pharmacological intervention (Szyf 2009). Thisdouble hole in the structure of the gene calls radically into question the leak-proof separation of the “genetics” from the “social” and, accordingly, the equationof genetics with a natural lottery.

Identifying genetics with a lottery implies understanding it as operating blindlyand immune from the distortions of social structures. The notion of a lottery impliesthe idea of a game of chance (Buchanan et al. 2000, 83), a matter of mere misfor-tune upon which no call for responsibility can be made. In abstract, being born witha faulty gene (Nagel’s case) may be thought of as just poor chance.

The idea of a genetic lottery fits well with the twentieth-century view of heredityas a mere transmission of DNA sequence. However, things become more compli-cated in epigenetic scenarios when we include in heredity not only DNA sequencebut also how these sequences are expressed. If we take seriously, as epigeneticistsclaim, the possibility that genetic expression may incorporate our early life experi-ences or even the experiences of our ancestors, the picture of the relationshipbetween the social and the natural, misfortune and choice, appears to change sig-nificantly. How much of a role can be given to “poor chance” in cases like theDutch Hunger Winter, in which biological disadvantage of two generations isseen as a direct consequence of the social hardship suffered in 1944? And inother scenarios where smoking, alcohol, styles of nurturing, or various psychoso-cial factors seem to directly affect the well-being of present and future generations?

The implications of these studies for public health are clearly far-reaching. Novelinteresting studies inspired by a developmentalist and epigenetic approach havealready started to elaborate a model by which social conditions and material con-texts experienced by past generations may become biologically embedded andsomatically transmitted in specific social groups. In these novel models, social

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factors such as nutrition, health care, financial and social capital, education andpublic policies in general are embodied and transmitted transgenerationally, thusperpetuating disadvantaged situations (see the “maternal fitness model”, Wells2007, 2010; and the “intergenerational phenotypic inertia model”, for instance:Kuzawa and Sweet 2009).

These studies highlight that neat boundaries between what is “given in nature”,and therefore not subject to the demands of justice (for instance genetic malfunc-tioning), and what is instead “socially transmitted” (the long-term effects ofunjust social structures), appear very precarious to maintain. To the extent thatthe social operates via the biological and vice versa, chance and choice tendincreasingly to overlap.

This is why epigenetics has the potential to represent an interesting problemati-zation for theories of justice (Loi, Del Savio, and Stupka 2013). Where should apolitical demand for justice stop if these epigenetic findings will be further con-firmed? And where are the boundaries between personal and collective responsibil-ity in a context where social factors seem so massively engaged in producingaspects of our own individual biology?

Of course, it is not my goal here to adjudicate between different approaches totheories of justice. Different political philosophers will look in different ways atthe potential implications of epigenetics. Epigenetics will probably not be a signifi-cant change for the two opposite positions of radical libertarians and luck egalitar-ians (justice as equality of fortune). For the first, no matter the social or naturalorigins of an inequality, everyone will be entitled to her/his own lottery ticketreceived at birth. For the latter, arbitrary (unchosen) original conditions, be themnatural or social, have to be corrected, so the “faulty gene” of traditional geneticexplanation would be no different from an abnormal methylation pattern causedby (or, associated with) “unjust” social structure.5

However, leaving these two positions aside, for the remaining arc of socio-pol-itical theories, taking epigenetics seriously will have important implications,although the nature of those implications may be less unidirectional than manyhave noticed so far.

Loading the genetic dices?

To reiterate the argument: if genetics is a lottery, from an epigenetic point of view,“the dice are loaded in the womb” (Gardner 2012). Arguments a la Nagel about adefective gene’s effects being out of the scope of social justice will be assessedcarefully. At least a second look will be required to adjudicate whether there is asense in which previous unjust social situations may have had an influence inabnormal genetic functioning.

More importantly, because of its supposed protection from the distortion ofsocial structures, the notion of a genetic lottery still possesses a democratic andegalitarian allure in the eyes of many political philosophers. This argument has

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a long tradition in biology (firstly in Weismann 1891), and was remade in the lasttwo decades by Francis Fukuyama, when he claimed that far from being “inher-ently unfair” “because it condemns certain people to lesser intelligence, or badlooks, or disabilities of one sort or another”, the genetic lottery is actually:

profoundly egalitarian, since everyone, regardless of social class, race or ethnicity,has to play in it. The wealthiest man can and often does have a good-for-nothingson ( . . . ) the genetic lottery guarantees that the son or daughter of a rich and success-ful parent will not necessarily inherit the talent and abilities that created conditionsconducive to the parents’ success. (2002, 156)6

My argument is that this democratic allure of genetics will be easily lost if epi-genetics prevails. Epigenetic findings undermine Fukuyama’s view that the geneticlottery is inherently impartial and blind. In an epigenetic view, the genetic lottery(genetic transmission + genetic expression) is always influenced by the social dis-parities of the past, from individual behaviors (smoking, etc.) to social processes(war, poverty, injustice, etc.).

If genes have their own biological memory, the genetic lottery loses its originalveil of ignorance. It brings the mark of a certain social inertia, is no longer impar-tial, and we cannot fully count on it as a fair referee distributing randomly naturalassets. Such a denaturalization of the genetic lottery has two profound implications.

On one side, it may be used for broadening the scope of theories of justice. Intheir seminal book on genetics and justice, Buchanan and colleagues anticipateda colonization of the natural by notions of justice from the viewpoint of newgenetic technologies making possible a change of human nature (2000). In a differ-ent sense, epigenetics may bring a similar colonization of the natural via socialjustice. Loi, Del Savio, and Stupka (2013) have argued that a deconstruction ofthe natural/social border implies that luck-egalitarian proposals in which thereare no moral differences between unchosen natural and social disadvantages, andRawlsian approaches that are limited to social inequalities, will increasinglycome to overlap. To the extent that for instance class inequalities are now seento operate via biological endowments, the Rawlsian principle for a fair equalityof opportunity can be broadened to include biological assets (Loi, Del Savio,and Stupka 2013).

On the other side, however, this translation of epigenetic themes in public policyand public health debates may be less unidirectional (expanding justice) than thisliberal agenda might like. Here is where some knowledge of the history of biopo-litics may help. What has not to be forgotten is that the public health implications ofsoft-inheritance are full of problematic and often counterintuitive aspects.Lamarckism in public policy, of which today epigenetics would be an actualization,has always been an ambiguous force (Bowler 1984; Meloni, forthcoming). It can beused to promote social reform because, obviously, investing massively on the socialis not a wasted effort if good habits can be passed on across generations (as “left-Lamarckians” like Paul Kammerer highlighted in the 1920s).

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However, the inheritance of acquired characters is a double-edged sword: alsobad habits can become bad biology, and the scars of past exposures and traumascan give rise to ideas of specific groups being “too damaged7” to be rescued. Inheri-tance in sum can be “poisoned”, as a recent headline in The Economist (2013) hassignificantly claimed, commenting on new epigenetics studies.8 If one looks at thedebate in Europe and America until the 1910s, when soft-hereditarian ideas werestill used in public policy and public health debates, this notion of a poisoned her-edity (or a “racial poison”) was widespread and not always employed for progress-ive goals. For instance, the notion that alcoholism or life in the slums couldirremediably damage not only the germ-plasm of exposed people but also that oftheir offspring was quite common at the time. In sum, the sins of the fatherswould leave the new generation in a situation of biological inferiority from thevery beginning. Upon this poisoned heredity, a racism or classism complementaryto but different from its more famous hard-hereditarian version was easily promoted(Meloni, forthcoming).

Several neo-Darwinians and Mendelians, from Wallace to Haldane and Huxley,have highlighted over time these problematic aspects of Lamarckian inheritance.However, the most significant use of this anti-Lamarckian argument in socialpolicy to my knowledge was made in the 1920s by Soviet geneticist and eugenicistYuri Filipchenko when he claimed, in Loren Graham’s words, that if Lamarckismnot Mendelism was true:

all socially or physically deprived groups, races, and classes of people-such as theproletariat and peasantry and the nonwhite races-would have inherited the debilitat-ing effects of having lived for centuries under deprived conditions. Far from promis-ing rapid social reform, the inheritance of acquired characteristics would mean thatthe upper classes are not only socially and economically advantaged, but geneticallyprivileged as well, a result of centuries of living in a beneficial environment. Thus theproletariat in Soviet Russia would never be capable of running the state; it was geneti-cally lamed by the inheritance of the effects of its poverty. (1977, my italics; see alsoBabkov 2013)

As I will say in my critical final coda, Filipchenko’s argument may still have itssignificance today.

The rise of the epigenetic body: how social structures get under the skin

A second contribution of epigenetics to a reformulation of the relationship betweenthe biological and the social is in the area of theories of embodiment. Since the1990s, we have witnessed in social theory (Shilling 2003; Williams, Birke, andBendelow 2003) and social epidemiology (Krieger 2001a, 2001b, 2004, 2011;Krieger and Davey Smith 2004) to an increasing recourse to the notion of “embo-diment” to undermine the sterile opposition between biomedical models (endogen-ous causes of disease) on one side, and social constructionist views (body as theeffect of language/power structures), on the other.

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As I will highlight next, also in theories of embodiment, epigenetics can signifi-cantly contribute to undermine established dichotomies between the “natural” andthe “social”.

Embodiment before embodiment

As social epidemiologists are well aware of, there is a long and oft-forgotten history(Waitzkin 2005) of how social conditions are literally inscribed into human bodiesand socioeconomic disparities are replicated in the bodily structures of peopleaffected by these differences (Krieger 2005, 2011). The writings of the Frenchdoctor and economist Rene Villerme, of cell-biologist Rudolf Virchow, as wellas Friedrich Engels’ classical work on the destructive impact of industrializationon the physiological and moral life of the working class (1844), can be read as nine-teenth-century anticipation of an embodiment paradigm (see extended references inKrieger and Davey Smith 2004; Waitzkin 2005).

The paradox, however, is that in the twentieth century, such references to the wayclass differences are manifest in the visual cues of people (“class physiognomies”:Laurell, quoted in Krieger 2005), have been rather marginalized in social epidemio-logical studies (Shaw, Tunstall, and Davey Smith 2003). Such discourse was seenas too resonant with darker biologistic arguments from Lombrosian psychiatry toeugenics.

In the last two decades, however, something has started to change again in thisdomain. From neuroscience to social epidemiology, it seems possible again toinvoke the physiological and the somatic level in social explanations, withoutfalling back into classical deterministic arguments. Rather, what is emphasized inthis new wave of studies is the continuous and plastic interchange of the bodywith its material surroundings.

The new embodiment

Since the 1990s, cumulative evidence of the impact of social contexts on the humanbody has become increasingly available. Data from biomedical and psychoneur-oimmunological research have highlighted the dependence of neuroendocrineand immune functions on social structures and social and psychological processes(Cacioppo 1994; Cohen and Herbert 1996), and emphasized how social character-istics and structures of neighborhood and communities (Diez Roux, Jacobs, andKiefe 2002; Sampson 2003) have a profound impact on well-being and cardiovas-cular health.

Social relationships, social integration, support, and social networks have beenincreasingly linked to longevity, and seen as “intrinsically beneficial” to physicalhealth (Seeman 1996; Cohen 2004; Brown et al. 2009; Cohen and Janicki-Deverts 2009; Kok and Fredrickson 2010). On the other side, social isolationand rejection, loneliness, and disruption of social networks are increasingly seen

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as major causes for morbidity and mortality (Cacioppo et al. 2002, 2006; Cacioppoand Patrick 2008; Cacioppo, Hawkley, and Thisted 2010).

Moving from the individual body to the “body economic”, there is now anincreasing number of studies focusing on the way in which austerity policiesand inequalities are literally “killing” factors (Stuckler and Basu 2013; Therborn2014). These recent works constitute the apex of a longer tradition of socialresearch on the way in which inequality, discrimination, and racism literallyharm health (Clark et al. 1999; Krieger 2001a, 2001b, 2011, 2004). Probablythe most significant and investigated construct in this area, for its implicationsfor public health, is socioeconomic status (SES). Since the early 1990s, studieson SES discovered a strong link between low social status and higher mortalityrates (Marmot et al. 1991). Detailed studies exist today connecting SES toasthma (Chen et al. 2006), changes in immune responses (Chen et al. 2003;Dowd and Aiello 2009), increased cardiovascular disease risk factors (particularlyamongst non-Hispanic Blacks: Sharma et al. 2004), abnormal cortisol secretion inyoung children (Lupien et al. 2000, 2001), dysregulation of the stress axis andresponse (Evans and Kim 2007). In general, the pathways by which SES isseen to drive disease include systemic inflammation, cellular processes, andgenomic pathways (Chen and Miller 2013; see also Wolfe, Evans, and Seeman2012).

Finally, in the last decade, neuroscience has gained the lion’s share in this litera-ture on the embodiment of social influences and the effects of SES in particular. Atthe core of the recent wave of neuroscientific studies is the growing appreciation forthe plasticity of the brain, including the adult brain (Rosenzweig and Barnes 2003;Rubin 2009; McEwen and Gianaros 2010). The recognition of plasticity, how thebrain constantly responds to and is literally shaped by environmental contexts,makes possible to establish increasing connections between social settings andbrain morphology and functions.

Several neuroscience-based studies have addressed directly how SES impactsnot only cognitive and emotional development, but also the volumes of the hippo-campus and the amygdala, memory, and certain aspects of executive function,including cognitive control (respectively: Noble, Norman, and Farah 2005; Farahet al. 2006; Noble, McCandliss, and Farah 2007; Hackman and Farah 2009;Hackman, Farah, and Meaney 2010; Noble et al. 2012). Two recent fMRIstudies have reported SES differences in the function of certain language-support-ing brain regions, namely the left fusiform (Noble et al. 2006) and left inferiorfrontal gyrus (Raizada et al. 2008).

Epigenetic research plays a twofold function in this context. First, it provides akey missing link to explain the molecular pathways by which transient environ-mental factors can leave marks or be inscribed on the biological body. The oftenelusive role of social factors (poverty, inequality, stress, etc.) in affecting body,brain, and behavior is likely to find a less ethereal mechanism in epigeneticmutations (Champagne 2008; Miller 2010; Vineis, Stringhini, and Porta 2014).

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Obviously, if the trend toward transgenerational epigenetic inheritance consoli-dates, the mechanism will involve the exposed as well as unexposed generations.

Second, epigenetics represents the conceptual climax of a more profoundappreciation of the many material flows connecting the wider environment andthe body (Niewohner 2015). I will conclude this section looking at the potentialpresent in epigenetics to move toward a more ecosocial view of embodiment(Krieger 2011), beyond superficial interactionism.

The epigenetic body: embodied constructivism beyond the genetic body

Several works have recently emphasized the potential in the epigenetic discourse ofundermining the still influential idea of a body autonomous and insulated from itsmaterial surroundings. Especially when understood in a broader ecological sense,epigenetics becomes a perfect epitome for the extreme porosity between the milieuand the body (Niewohner 2011; Guthman and Mansfield 2013; Landecker andPanofsky 2013).

Meaney’s experiments on the transmission of inadequate mothering in rats areillustrative of this dynamic for which social events become regulative of the organ-ism physiology. Certain behavioral exposures (low licking/grooming) shapechanges in methylation patterns which modify genetic expression and neurochemi-cal responses in the brain. This modifies the physiology of the pups that, once adult,will reproduce the inducing behavior and consequently modify their environmentalniche, thus shaping the biological life of the forthcoming generations.

In epigenetics, therefore, the dictum that “bodies express ecology” (Piersma andvan Gils 2011) seems to find a stronger confirmation. We are well beyond, asNiewohner insightfully writes, Western views of the individual as separated fromthe external world by their “impenetrable” skin. An “altogether different degreeof entanglement between body and context” is brought to light (Niewohner2011, 12).

What is important to emphasize here is that this openness of the epigenetic bodyto the world is a significant rupture with the mainstream lesson of twentieth-centurybiology, especially genetics (Meloni and Testa 2014). The “genetic body” was builton a segregation of the somatic from the genotypic level (Griesemer 2002). First,through the Weismannian separation of the soma from the germ line (impervious-ness of the hereditary material). Second, this passivity of the somatic level waspowerfully reinforced by the so-called central dogma of molecular biologywhich stated that no information can travel back from protein to DNA.

Relegated to the passive end of the genetic chain of information (Gudding 1996),the agency, situatedness, and reality of the body were elided in twentieth-centurygenetics (Keller 2001; Robert 2004). Not by chance, as social epidemiologistsand historians of medicine have noted (Krieger 2011; Rosenberg 2012), the main-stream biomedical approach, with its search for inward causes, ended up viewingthe body as a passive biophysical abstraction with no serious consideration of its

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embedding in local surroundings, nor appreciation of it being marked by materialand local contexts (Lock 1993, 2001, 2013; Lock and Nguyen 2010).

Against this marginalization of the body in twentieth-century genetics and bio-medicine, epigenetics, especially when theorized in sophisticated conceptual fra-meworks, can be seen as a decisive advance toward what can here be called“embodied constructivism”, a position that tries to overcome the shortcomings ofboth reductionistic biomedical theories of the body and disembodied social con-structionist views.

Although used sometimes in phenomenological contexts, the term “embodiedconstructivism” is here named after the “constructivist interactionism” of develop-mental systems theory (DST). In the spirit of DST, it is meant as an invitation tothink in terms of non-dichotomous, jointly determined, and reciprocally contingentbiosocial factors when explaining human development and social life (Gray 1992;Oyama 2000; Griffiths and Gray 2005). Expanding on the “parity thesis” of DST(i.e. equality between the role of DNA and non-DNA elements in explaining devel-opment), embodied constructivism defines a non-hierarchical and relational ontol-ogy in which social structures can be seen as the sources, as well as the effects, ofbiological factors. There is a genetic component to social processes, as there is asocial shaping and driving of genetic expression (Landecker and Panofsky2013). Biological realities always emerge socially situated and “interwoven withmeanings” and culture is always in human biology (Goodman 2013).

A good example of what I mean by embodied constructivism is the recent workby Clarence Gravlee on the material pathways through which “race becomesbiology”. The article explores the double biosocial movement by which racismand social inequalities (as a sociocultural categories) are embedded and materia-lized in the “the biology of racialized groups and individuals”; while at the sametime embodied inequalities (the biological level) perpetuate and reinforce “a racia-lized understanding of human biology” (Gravlee 2009, 54).

Epigenetics can fruitfully complement this line of thought: the body bears theinscriptions of its socially and materially situated milieu, and the milieu is consti-tuted by the socially modulated present and past biographies of the body (nutri-tional, metabolic, behavioral, toxicological, psychosocial, cultural, etc.).

Not only is genetic expression socially modulated (that is influenced by powerstructures in society), but it is also the source of novel environments that will shapein their turn the “socially modulated biologies of further generations of organisms.The causal arrows go both ways, and the ontology of the gene as content and theenvironment as context ceases to make sense” (Landecker and Panofsky 2013, 351).

Critical coda: soft-inheritance and its quandary

I have mainly focused in my article on the theoretical potential of epigenetics tobring together biology and the social sciences after the acrimonious conflicts ofthe twentieth century, conflicts partly determined by taking a narrow view of the

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biological and equating heredity with transmission genetics. In the light of this, itwould be tempting to say that, in the novel postgenomic scenario, we finally havethe opportunity to speak of the biological in the social sciences with less anxietyabout biomedical reductionism or the return of eugenics. Although tempting, itwould be more careful to postpone this claim. The reason for this cautiousnesscomes, in my view, from two main lines of argument.

First, there are too many uncertainties not only about the biological significanceof the epigenetics revolution but also about which stakeholders are going to prevailin the epigenetic arena. It is far from obvious that gene-centrism will be effectivelyovercome rather than reinforced by epigenetic findings (Lock 2005, 2012; Richard-son, forthcoming; Waggoner and Uller 2015). The epigenetics revolution has beencompared to the Gramscian passive revolution (Meloni and Testa 2014): a revolu-tion that is less a radical break and more a limping compromise with existing forces.The analogy seems valid especially in the sense that underneath the surface ofhyped claims about a rupture with gene-centrism, the prevailing tendency is touse epigenetic marks as a sort of “fifth letter” of the genetic code and a tonic forthe unkept promises of various genetic projects.

A second, different but likewise important line of critical reflection originatesfrom opposite reasons. It is the possibility that a new epigenetic approach willprevail in wider society as a distinctive way to understand the relationshipbetween human beings and their biology. It is possible (much more so than threedecades ago) that hard-heredity will no longer be considered the only game intown. This idea has been at the center of my article: with the advent of epigenetics,a paradigm is emerging that is very different from previous articulations of geneticsand society. If we look at the two main implications of epigenetics for the socialsciences as I have described them: (1) the denaturalization of the genetic lotteryand (2) the deepening of the notion of embodiment – what seems to emerge ascommon point is the erasure of any residual distance between the biological andthe social in the ontology of human beings.

To reiterate the previous findings: present social contexts get under the skin anddeeply impregnate our biology; various past environmental events shape ourgenetic lottery, at least at the level of its expression; local social events becomeembedded in human biology potentially contributing to differences and variationamongst human groups. The bottom line is that a demarcation between the biologi-cal and the social becomes increasingly untenable.

However, this blurring of the line between the biological and the social mayassume several contradictory guises in society, with unpredictable effects uponmoral reasoning. What it is often too easily forgotten is that the separation of thebiological from the social established by hard-heredity played a complex concep-tual and political role in the early twentieth century. In favoring an emancipation ofthe sociocultural from its biological bases, hard-heredity became (somehow para-doxically) a helpful construct for cultural anthropologists eager to get rid of the“vitiated mixture” (Kroeber 1915) between the heredity and civilization,

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symbolized by the Lamarckian trends in social sciences (Stocking 1968). It is unde-niable that Kroeber, one of the fathers of cultural anthropology, saw in hard-her-edity his best ally to get rid of racialist discourses that were part and parcel ofneo-Lamarckian social science (Stocking 1968; Kronfeldner 2009). Of course,there was an awful lot of racism also amongst Mendelians (as Boas well knew),but Kroeber thought that by taking advantage of the separation of heredity fromculture, this latter (aka the Superorganic) could become a space that is muchmore malleable by human effort.

If we are heading therefore toward a potentially neo-Lamarckian situation wheresocial experiences can become part of our biological memories, we need to con-sider carefully what this may imply for the social sciences. Leaving aside ontologi-cal questions about the robustness of the category of the “social” (and therefore of“social sciences”) in a situation of continuous osmosis between “the biological”and “the sociocultural”, some more directly political questions remain to beanswered for the epigenetic scenario.

For instance: How will the plethora of epigenetic studies showing how varioustraumas travel across generations be translated into public policy? How completewill the identification be between specific groups and the pathogenic environmentsto which they have been historically exposed? Will this become the basis for repro-ducing and consolidating structural differences in society (class, gender, and race)?What if epigenetic discourses on different levels of methylation between socialgroups are used to underwrite new discourses on “the biological inferiority ofthe poor” (Katz 2013) or on an incapacity of the poor to seriously take care of them-selves (Mansfield 2012)? Will the embedding of the body in its own socioeconomiccontext become so profound as to be seen as an unbreakable chain? Or will thenovel emphasis on the reversibility of epigenetic signatures always keep openthe possibility of change even for the most “damaged” social groups?

It is likely that the answer will be somewhere in between this two. As EvaJablonka has pointed out (personal communication), a critique a la Filipchenkoon the permanent scars left by historical oppression on the proletariat is based onthe idea that with time epigenetic effects become more difficult to reverse. Thisdoes not need to be, however, as the reversibility of epigenetic signatures candepend on different factors, such as the persistence of certain environmental con-ditions, etc. Epigeneticists seem in sum more optimistic today about the easy rever-sibility of epigenetic marks. Scientifically, this may be a satisfying answer, but thepolitical conditions in which the costs of reversing bad epigenetic signatures willtake place are not for scientists to decide. We too often take for granted the natur-alness of our human rights framework in which we expect epigenetics to help pro-moting better schools for children, or more serious cures for traumatized people.

However, what the last century has taught us is that when the broader politicalconditions deteriorate, even good scientific ideas such as Mendelian inheritancecan easily be turned into ammunitions for race or class discrimination. Therefore,in principle, the relatively easy reversibility of epigenetic marks should be a

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reassuring answer against fears of a new biological racism or classism under epige-netic guises. However, how costly for society this reversibility will be is not in thehands of scientists. Cost–benefit considerations on the availability of publicresources to invest on such policies, or political considerations on the opportunityto do so is not a scientific but a political decision.

These are the set of problems to which this article has no answer, but nonethelessneed to be raised to set the perimeter of the specific ethos of our postgenomic world.Certainly many of these social sciences translations appear speculative at present,given the uncertain status of epigenetic research and, particularly, transgenerationalinheritance, as said above.

However, we must also not forget that much of the science behind eugenics (theunit of character hypothesis for instance) was, to say the least, precarious and con-troversial at the time (and in the end wrong), but this did not stop a direct translationof these scientific arguments into harsh policy measures, under certain existing con-ditions of “law and sentiment”, as Galton would have said. My mixed conclusion isthat, as usual in the history of how biological arguments are imported into thepublic sphere, there is no one-to-one relationship between scientific theories andsocial values (Meloni, forthcoming): much will depend on the broader socio-politi-cal context in which science circulates.

Epigenetics has lots of potential to rethink the relationship between the biologi-cal and the social world. However, even the best conceptual framework may beopen to the most unpredictable and sometimes unfortunate socio-politicaloutcome. This is why social and political scientists need to be prepared to themany nuances of the novel scenario.

Acknowledgments

Thanks to Eva Jablonka, Danielle Allen, Didier Fassin, and Alexandre Guerrero for helpful commentsand bibliographic suggestions on the text. Of course, all the responsibility for the claims made isexclusively mine. Thanks as usual to Andrew Turner for his help with the English language in thetext. Thanks also to John Dupre and the Egenis Centre, College of Social Sciences & InternationalStudies, Exeter University (UK) for offering me an Honorary Senior Lectureship that has be veryhelpful in carrying out this research.

Disclosure statementNo potential conflict of interest was reported by the author.

FundingI acknowledge the contribution of a Marie Curie ERG grant, FP7-PEOPLE-2010-RG (research titled“The Seductive Power of the Neurosciences: An Intellectual Genealogy”), of a CAS/University ofNottingham pilot grant on epigenetics, and of a Institute for Advanced Study (School of SocialScience), Princeton, NJ, annual membership funding.

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Notes

1. See a subsequent work by the same group on the transgenerational effects (on the male line) ofsmoking during puberty in Pembrey et al. (2006).

2. Often, a technical distinction is introduced between inter-generational (or parental) andtransgenerational effects, with the former shorter and limited to two generations, and thesecond spanning over multiple generations (see Grossniklaus et al., 2013; Heard andMartienssen, 2014).

3. A recent interesting update on epigenetic inheritance is Dias and Ressler (2014) study on thetransgenerational effects of olfactory sensitivity in mice. See for a comment Szyf (2014).

4. It is the case of so-called luck egalitarians for which any inequality of fortune, that isbeyond one’s control, no matter whether natural or social, needs to be corrected in society(for instance in authors like Cohen and Dworkin; see for a critique Anderson, 1999). Asimilar disregard for the difference between natural and social in the cause of unfair situationsapplies in principle also to a full-fledged consequentialist position, for which what matters isthe nature of the outcome.

5. Although the key “moral” distinction in luck egalitarianism, which is between option luck andbrute luck, that is between choices and circumstances (Anderson, 1999; Tan 2012) may alsobe in trouble in an epigenetic view.

6. The core of Fukuyama’s argument was against the idea of the genetic supermarket, i.e. replacingthe blind genetic lottery with consumerist choices of engineered reproduction that wouldeventually “increase the disparity between the top and bottom of the social hierarchy” (2002,157).

7. See the title “Babies born into poverty are damaged forever before birth” in the newspaper TheScotsman as a comment to an epigenetic study on social exposure in the city of Glasgow, UK(McLaughlin 2012).

8. A study on the negative transgenerational effect of lack of folate in mice diet (The Economist2013).

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