International Journal of Epidemiology, 2016, Vol. 45, No. 1 habits that may have been acquired. It ought to assign limits to reasonable doubt. We should learn that the number of times in which an acquired habit was inherited to any sensible degree, after some specified number of generations, did certainly not exceed so and so, per cent. We might also learn that it certainly did exceed some lesser number. We cannot prove any negative and therefore cannot hope to prove that acquired habits are never inherited, but we may perhaps find no case of their being inherited in any appreciable degree, even after many generations. This seems to have been already done in the case of many artificial mutilations. But as the loss of a limb belongs to a different order of events to the loss of an instinct, it would be dangerous to conclude that what has been found to occur in the one case would necessarily occur in the other. It therefore appears to me that the possibility of experiments along the lines I have suggested, on creatures reared from eggs and away from their parents, are well worth considering. It is only through the hard compulsion of unquestion- 15 able facts, that we are likely to forego the wholesome moral doctrine that the self-denials and toils of the parent will earn for his children a happier nature than he had himself been gifted with. That they will be congenitally more disposed than he was to deny themselves harmful pleasures, and to work generously for the good of others. It must be recollected that we are no longer justified in idly holding this comfortable belief, and in preaching about it as if it were unquestionably true. So far from this being the case the balance of probability seems strongly against its truth. But the question is not yet solved. The possibility of a child inheriting even the simplest aptitudes that the parent has acquired against the grain of his nature, can only be determined by a direct appeal to suitable experiments. The object of this memoir is to obtain the opinion of experts as to the most feasible experiments for the purpose. International Journal of Epidemiology, 2016, 15–20 Commentary: Before doi: 10.1093/ije/dyw004 Weismann and germplasm there was Galton and eugenics: the biological and political meaning of the inheritance of acquired characteristics in the late 19th century Ruth Schwartz Cowan Philadelphia, PA 19104, USA E-mail: [email protected] Accepted 11 January 2016 Francis Galton was a Victorian gentleman of means, more a polymath than a disciplined man of science. He was born in 1822 into a distinguished and affluent Birmingham family. His father, Samuel Tertius Galton, was a banker; his mother, Violetta Darwin Galton, was one of the many daughters of Erasmus Darwin, which made her an aunt (and Francis therefore a cousin) to Charles Darwin. Francis Galton’s paternal and maternal grandfathers had been Quakers and founding members of the famous Lunar Society of Birmingham, an association of men who were interested in what we would now call science and technology. According to family accounts, Francis was a very bright and inquisitive boy—but he disliked school and was an in- different student. As a young man he studied medicine for a few years and then, over some parental objections, went to Cambridge; he left 3 years later with only an ordinary degree, having done poorly in all of his examinations. In the autumn of 1844 shortly after receiving his degree, his father died, leaving an enormous inheritance. After that Galton had no need of gainful employment; until his death, 76 years later, he was free to spend all of his time and a good deal of his money on his personal interests, almost all of them (following in the footsteps of his family) either scientific or mathematical. Over the course of his long life, Galton wrote and published more than 300 papers and 9 full-length books, almost all of which had some C The Author 2016; all rights reserved. Published by Oxford University Press on behalf of the International Epidemiological Association V 16 International Journal of Epidemiology, 2016, Vol. 45, No. 1 observational and statistical content. Historians of the various sciences credit him with having made significant contributions to the study of African geography, anthropology, anthropometry, psychology, statistics, meteorology, criminology—and, of particular concern here— genetics, which in Galton’s day was called the study of heredity. In 1889, when Galton delivered the essay (published here in full for the first time) as a talk at the annual meeting of the British Association for the Advancement of Science, heredity was being investigated by two kinds of biologists: naturalists and cytologists. The naturalists were interested for two reasons; one was practical (how can we breed better domesticated animals and plants?) and the other was theoretical (evolution by natural selection requires an explanation—which was missing in those years—of how adaptive traits are passed from one generation of organisms to the next). The cytologists also had two reasons; they suspected that the cellular and nuclear structures that they were discovering with their increasingly powerful microscopes (and staining techniques) might provide that missing explanation—and might also hold the key to the linked mysteries of reproduction and embryological development. Galton, however, was neither a naturalist nor a cytologist; he was interested in heredity for an entirely different reason, a reason which he had first explored in print in 1865.1 ‘Hereditary talent and character’ was Galton’s first foray into the study of heredity, and its very first paragraph holds the key to understanding both the nature and the depth of his interest in the subject: The power of man over animal life, in producing whatever varieties of form he pleases, is enormously great. It would seem as though the physical structure of future generations was almost as plastic as clay, under the control of the breeder’s will. It is my desire to show more pointedly than—as far as I am aware—has been attempted before, that mental qualities are equally under control. (p.157)1 Galton was, as this paragraph makes clear, interested in the inheritance of human mental qualities for philanthropic reasons; he hoped it would prove possible to improve the human race by controlling its breeding patterns—just as if humans were domestic animals, like horses, dogs and sheep, and as if behavioural traits were akin to physical characteristics. To put the matter another way, Galton’s argument rests on two analogies, which he carefully expounds at the beginning of the article: first, that mental traits can be inherited in the same way as physical traits; second, that heredity in man is no different from heredity in animals. If this argument and these analogies sound familiar to 21stcentury ears, it is because they are the foundations for both a scientific discipline and a social movement, each of which still exists (although considerably altered) in our own day. The scientific discipline is now called human genetics; the social movement was then, and still is, called eugenics. Over the course of his long career, Galton never gave up his conviction that the best hope for social reform rested with better knowledge of human heredity and better application of its laws. Galton himself coined the term ‘eugenics’—combining the Greek roots for ‘good’ and ‘ancestry’—in 1883 (pp. 24–25).2 His eugenic convictions, first articulated in 1865, underlay all his subsequent studies of heredity. About one-quarter of ‘Hereditary talent and character’ is devoted to developing the empirical evidence in favour of Galton’s thesis, ‘ . . . that talent and peculiarities of character are found in the children, when they have existed in either of the parents, to an extent beyond all question greater than in the children of ordinary persons’. (p.158)1 To do this, he attempts a statistical analysis of several biographical dictionaries. He finds in one such volume that out of the 605 individuals listed, 102 were relatives of someone else on the list, with a frequency of 1 in 6. Doing the same analysis on another biographical dictionary of eminent men, he finds a frequency of 1 in 11: in a listing of painters, 1 in 6; for musicians, 1 in 10; for scientists, 1 in 12; among Lord Chancellors (the law being ‘by far the most open to fair competition of all the professions’) he finds a frequency of 1 in 3 (p.161).1 The conclusion, he argues, is ineluctable: ‘Everywhere is the enormous power of hereditary influence forced on our attention’. (p.163)1 Not one whit of additional evidence is provided, however; the remaining two-thirds of Galton’s article is concerned with the practical implications of this conclusion (can young people be persuaded to find intellectual superiority more romantically attractive than physical beauty? Can the government be persuaded to give large cash wedding presents to intellectually superior couples?). If modern readers find the evidence unconvincing, so too did many readers in his own day (Cowan, pp.10–20, 60–65; Gillham, pp.169–172).3,4 In subsequent researches, Galton improved his investigatory methods but never found any reason to alter his original conclusion about the enormous power of hereditary influence. Furthermore, he retained his scepticism, already announced in ‘Hereditary talent and character’ about the idea (which was then a part of both biological and political orthodoxy) that characters acquired during the lifetime of an individual (characters both physical and mental) could be inherited. ‘Can we hand anything down to our children’, he asked at the beginning of the second part of the International Journal of Epidemiology, 2016, Vol. 45, No. 1 article, ‘that we have fairly won by own independent exertions?’ (p.321)1 Probably not, he answers; the sons of soldiers, for example, do not learn to drill any faster than the sons of other men, nor are the sons of sailors noticeably more accustomed to the sea than the sons of those who are land bound. ‘Or are we’, he goes on to ask, ‘no more than passive transmitters of a nature we have received, and which we have no power to modify?’ (p.322)1 Probably, he continues, making a most unorthodox (for his day) suggestion, we should ‘ . . . consider our own embryos to have sprung immediately from those embryos whence our parents were developed, and these from the embryos of their parents, and so on forever’. If this suggestion sounds uncannily like August Weismann’s concept of the continuity of germplasm, that is because it is, with one crucial caveat; Weismann came to his conclusion (in the 1880s) after decades of careful biological research, but Galton came to his because it followed, almost logically, from his political convictions. Liberalism was the dominant political orthodoxy in England in the middle years of the 19th century. Victorian liberals believed that men (and they meant men as a sex, not as a generic term for humans) were basically the same at the time of their birth and were only made different from each other by the effects of their environments and educations; many liberal reform efforts were, therefore, focused on relieving poverty and improving access to education. Victorian conservatives, on the other hand, took the opposite view, holding that men were inherently different from each other at birth, a difference which could not be overcome by environment and education; this view made them disinclined to believe that society could be, or even needed to be, reformed. Galton was a lifelong conservative. ‘It is in the most unqualified manner’, as he wrote in his book Hereditary Genius, ‘that I object to pretensions of natural equality’. (p.56)5 As a result, he tended to be sceptical of the biological notion that underlay liberal reform efforts, namely that we can pass anything biological down to our children that we can acquire during our lifetimes. Since the last decades of the 18th century, that biological notion had been an essential component of the scientific discussion about whether species had evolved over time. A prominent French naturalist, George Louis Leclerc, Comte du Buffon (1707-88) had started the argument by declaring that the fossil record (just then being, literally, unearthed) could be understood as the degeneration of the original God-created species as a result of the inherited effects of climate and diet. A generation later, a German comparative anatomist, Johann Friedrich Blumenbach (1752–1840) had asserted that the first human beings were light-skinned and that dark-skinned humans were, therefore, a degenerate form that had arisen when humans 17 migrated to places in which they suffered over-exposure to the sun. Erasmus Darwin, who was of course the common grandfather of Charles Darwin and Francis Galton, used the idea that acquired characters could be inherited to buttress his more progressive concept of evolution in Zoonomia; or the Laws of Organic Life, which was published in 1794. Subsequently, Buffon’s famous disciple Jean Baptiste Lamarck, whose major work Philosophie Zoologique appeared in 1809, followed Erasmus Darwin in turning his mentor’s vision on its head, seeing evolution as progressive rather than retrogressive, going from simplicity to complexity while retaining the same proposed mechanism, a ‘life force’ which allowed organisms to pass on traits that they had acquired (in response to changing environmental circumstances) during their lifetimes.6–9 Charles Darwin, perhaps the most observationally acute naturalist of the 19th century, also believed in the inheritance of acquired characters. In 1865, when Galton wrote ’Hereditary talent and character’, he could not have known that (since Darwin did not mention it in The Origin of Species, 1859) but he certainly learned it three years later, when Darwin published his dense two-volume tome The Variations in Animals and Plants Under Domestication. The first volume of The Variations contains huge amounts of information about the numerous varieties of domestic animals and plants, information that Darwin had spent several decades culling from the writings of animal and plant breeders; there are chapters on dogs and cats, horses, pigeons, fruits, ornamental trees, cereals and many other domesticated organisms. The second volume opens with several chapters in which Darwin tries to discern patterns of inheritance in all the data he had presented in the first. Some of these patterns did not involve the inheritance of acquired characters (hybridization for example) but a very large number of them did (‘use and disuse’, for example). In the penultimate chapter of the book, furthermore, Darwin proposes a physiological mechanism that might explain all the patterns that he had discovered; he called it ‘the provisional hypothesis of pangenesis’.6 Darwin speculates that every organism may contain a large number of very small heredity-particles (he called them ‘gemmules’) which are continuously produced by all the organs of the body and which continuously circulate throughout the body. Eventually, he concludes, the gemmules (or some portion of them) must collect in the reproductive organs, so that they can be passed on to the next generation. Quite explicitly, pangenesis allowed for and explained how acquired characters could be inherited. Galton had a very strong reaction to The Variations; his personal copy is filled with marginal notes.7 As a reader of ‘Hereditary talent and character’ might have easily guessed, he was dubious about the inheritance of acquired 18 characters which was, as he had pointed out, logically incompatible with his conviction that both physical and intellectual traits were innate. Where, for example, Darwin had written (Vol II, p.24) about cases of a child born with an injury similar to one that a parent had acquired, Galton wrote: ‘He might have had an infirm organ to begin with or . . . his wife, or his wife’s family must have’. On the other hand, he was enthusiastic about the notion of gemmules, because they could be mathematized; in Hereditary Genius, which Galton published just a year later, he was explicit on this point. ‘The theory of Pangenesis’, he wrote, ‘brings all the influences that bear on heredity into a form that is appropriate for the grasp of mathematical analysis’. (p.426)5 Until the publication of Hereditary Genius, Darwin and Galton had not had much of a personal relationship despite their family ties. But Darwin was delighted by Galton’s appreciation of pangenesis and so, when Galton wrote to him in December 1869 regarding some experiments that might test the theory, Darwin responded enthusiastically. Galton proposed to transfuse blood between rabbits of different breeds in order to see whether the traits of the donor rabbits (particularly coat colour) would appear in the offspring of the recipients. Darwin put Galton in touch with rabbit breeders and, for the next year, insisted on being kept apprised (by letter) of the progress of the experiments. To Galton’s delight—and Darwin’s embarrassment—the results were negative; out of 124 offspring in 21 litters born by transfused rabbit does, not a single ‘mongrel’ had appeared. Galton presented a full report of his findings to the Royal Society in March, 1871.12 (The correspondence between Darwin and Galton about the rabbit experiments can be found in Pearson’s Life, Letters and Labours of Sir Francis Galton, Vol. 2.13) Darwin reacted to Galton’s report uncharacteristically; he behaved badly. Galton found himself publicly admonished for having unwittingly misinterpreted his cousin’s theory. ‘In my chapter of Pangenesis in my ‘Variations of Animals and Plants under Domestication’, Darwin wrote in a letter to Nature, ‘I have not said one word about the blood, or about any fluid proper to any circulating system’. (p.502)14 Darwin was nit-picking. What he had actually said was ‘the gemmules in each organism must be thoroughly diffused; nor does this seem improbable considering their minuteness, and the steady circulation of fluids throughout the body’,—and in all the correspondence that he had had with Galton, he never expressed any doubts about the experimental efficacy of transfusing blood between breeds of rabbits.10 Galton’s reply to Darwin was a model of British forbearance. ‘I do not much complain’, he replied, a week later, ‘of having been sent on a false quest by ambiguous language, for I know how conscientious Mr. Darwin is in all he writes, International Journal of Epidemiology, 2016, Vol. 45, No. 1 how difficult it is to put thoughts into accurate speech, and again, how words have conveyed false impressions on the simplest matters from the earliest times’.15 But Galton never forgot his annoyance; this explains the first sentence of his B.A.A.S. speech (‘Feasible experiments have yet to be designed that shall be recognised by experts as satisfactory tests of the hereditary transmission of acquired habits’) as well as the fact that he waited until Darwin had died (1882) to discuss the matter again publicly. Despite the failure of the rabbit experiments, Galton never gave up his conviction that some sort of hereditary particle existed: his apologetic letter to Nature ends with the words, ‘Vive Pangenesis’.15 Indeed, between 1872 and 1876 he published two papers in which he developed a variant of the theory (which he called ‘stirp’). Galton understood stirp to be the entirety of all the hereditary particles in the body, but— unlike Darwin—he thought that, because of the large number of particles, the laws of heredity were most likely probabilistic.16,17 Over the course of the next decade, Galton pursued this hypothesis by undertaking experimental studies of the inheritance of height in ornamental sweet pea plants (the same plant, ironically, and one of the same traits that Mendel had studied) and a series of controlled observations on the heights of children compared with the heights of their parents. His analysis of those experiments and observations led him to the discovery of what we now call regression to the population mean, one of this major contributions to the science of statistics.18 Stirp theory did not, however, permit the inheritance of acquired characters. The text of Blood Relationship makes it very clear that Galton still held to his notion that every embryo is the direct, physical descendant of another embryo. ‘The span of the true hereditary link’, he wrote, ‘connects . . . not the parent with the offspring, but the primary elements of the two [stirp that is passed from one generation to the next] such as they existed in the newly impregnated ova’. (p.400)16 Just a few months before Galton gave his B.A.A.S lecture, August Weismann wrote to Galton acknowledging that Galton had expressed the central idea of continuity of germplasm earlier than he himself had done. After thanking Galton for having sent a copy of his most recent book, Natural Inheritance, in addition to some of his earlier papers on heredity, Weismann expressed his regret about not having previously read Galton’s A Theory of Heredity because ‘ . . . you have exposed in your paper an idea which is in one essential point nearly allied to my theory of the continuity of germplasm’. After explaining that an English translation of his essays was about to appear,19 Weismann went on to remark that, ‘I shall profit from the next occasion which offers itself to me to give a more extensive International Journal of Epidemiology, 2016, Vol. 45, No. 1 account of your views and to point out the differences between our views’.20 Weismann does not elaborate on what he thought those differences might be, but subsequent commentators have repeatedly done so.21,22 Weismann, one of the ablest of the German founders of cytology, is famous for having given the inheritance of acquired characters its final coup de grace by cutting off the tails of newborn laboratory mice and observing their offspring for several generations without once finding a newborn tail-less mouse. But Weismann only did that experiment after first convincing himself, through his cytological and embryological studies, that the physical substance responsible for generational continuity was located in the nucleus of the cell (he called it ‘germplasm’ as opposed to ‘protoplasm’). Galton never located stirp in the body except to say that it eventually collected in the reproductive organs; not being a naturalist, let alone a cytologist, Galton knew nothing about the existence of cells, let alone about the contents of their nuclei. Weismann’s version, as a result, had biological meaning in a way that Galton’s did not. It led to the discovery of the chromosomes and to the understanding of both mitosis and meiosis—and, eventually, helped naturalists to understand the significance of Mendel’s laws. Galton’s concept of continuity was not historically sterile; it laid the foundation for what we now call biostatistics. Weismann’s concept, however, laid the foundation for what we now call genetics. Galton’s concept also laid the foundation for what became, early in the 20th century, the eugenics movement. Galton had never given up his conviction, first expressed in 1865 and embedded in both the title and the content of his BAAS speech a quarter-century later, that human talent and character were determined more by nature than by nurture, and in his old age he was happy to lend his support to the social movement that some of his disciples were creating.(The pairing of nature and nurture appears in Shakespeare’s The Tempest (Act 4, Scene 1) but its modern usage is ascribed to Galton in the title of his book23). The title of the B.A.A.S paper reminds us that Galton was more interested in mental habits than in physical traits. The text also reminds us which organism he cared most about; it refers, repeatedly, to ‘parents’ and ‘children’, even though the animals on which he suggests experimenting are chickens, moths and fish. Galton focused on these organisms because they can be bred relatively easily and kept under controlled environmental conditions; he already knew— having undertaken a pioneering study of twins in the same years that he was developing his stirp theory—that human beings were far from the ideal experimental animals for this purpose. 24 Galton was a eugenicist even before he coined the term—and his passionate commitment to the idea that bet- 19 ter breeding would produce a better human being underlay all of his effort to study heredity and all of his efforts to deny the likelihood of the inheritance of acquired traits— as this paper aptly demonstrates. A note about the persons mentioned in Galton’s paper August Weismann (1834-914) has already been identified, but it is worth noting that Galton spelled his name wrongly here and continued to make the same mistake in subsequent publications. Mr Welldon is undoubtedly W. F. R. Weldon (1860–1906) who started his career as an invertebrate morphologist, but who developed an interest in the statistical study of variation in the 1890s, becoming a disciple of Galton’s and a collaborator with Karl Pearson. In the first decade of the 20th century, when the English Galtonians waged a fierce battle of words with the English Mendelians, Weldon and Pearson were the leaders of the Galtonian camp. Frank Merrifield was an entomologist living in Brighton, England, in the 1890s. He undertook some experiments with moths at Galton’s behest, the first of which are reported in ‘Practical Methods and Enquiries as to the Method of Breeding Selenia illustraria for the Purpose of Obtaining Data for Mr. Galton’, Trans. Entomol. Soc 1887;I:19-14. It has proved impossible to identify Miss Pridhom. Mobius is not the mathematician August Ferdinand Möbius (1790-1868) but rather the zoologist Karl August Möbius (1825-1908). The work in which he reported his experiment is a 20-page pamphlet, Die Bewegungen der Thiere, und ihr psychischer Horizont [The movement of animals and their psychological horizon] (Kiel, 1873). Charles Darwin gave a full account of the experiment in Chapter 3 of The Descent of Man, and Selection Related to Sex (1871)—which is undoubtedly where Galton encountered it, as he did not read German. Funding The research on which this essay is based was done forty years ago, in 1966-1967, when the author was a pre-doctoral fellow, under an NIH Training Grant on the history of genetics. The principal investigator was William Coleman, PhD. (deceased, 1988) then a faculty member at The Johns Hopkins University, where the author was then a graduate student. References 1. Galton F. Hereditary talent and character. MacMillan’s Magazine 1865;12:157–66, 318–27. 20 2. Galton F. Inquiries into Human Faculty and Its Development. London: Macmillan,1883. 3. Cowan RS. Sir Francis Galton and the Study of Heredity in the Nineteenth Century. New York, NY: Garland Press, 1985. 4. Gillham NW. A Life of Sir Francis Galton: From African Exploration to the Birth of Eugenics. Oxford, UK: Oxford University Press, 2001. 5. Galton F. Hereditary Genius. London: Macmillan, 1869. 6. Zirkle C. The Inheritance of Acquired Characters and the Provisional Hypothesis of Pangenesis. The American Naturalist 1935;69:417–45. 7. Green JC. The Death of Adam: Evolution and its Impact on Western Thought. New York, NY: Mentor, 1961. 8. Burkhardt RW. The Spirit of System: Lamarck and Evolutionary Biology. Cambridge, MA: Harvard University Press, 1977. 9. Jordanova LJ. Lamarck. Oxford, UK: Oxford University Press, 1984. 10. Darwin C. The Variations of Animals and Plants under Domestication. 2 vols. London: John Murray, 1868. 11. Located in Galton Papers, Archives, University College London, London UK. 12. Galton F. Experiments in pangenesis by breeding from rabbits of a pure variety, into whose circulation blood taken from other varieties had previously been transfused. Proc R Soc 1870-71; 19:404. International Journal of Epidemiology, 2016, Vol. 45, No. 1 13. Pearson K. Life, Letters and Labours of Sir Francis Galton. Vol. 2. Cambridge, UK: Cambridge University Press, 1914-30. 14. Darwin C. Pangenesis. Nature 27 April 1871:502–03. 15. Galton F. Pangenesis. Nature 4 May 1871:5–6. 16. Galton F. On blood relationship. Proc R. Soc 1872;21:394–401. 17. Galton F. A theory of heredity. J R Anthropol Inst 1876;5: 329–48. 18. Cowan RS. Francis Galton’s statistical ideas: the influence of eugenics. Isis 1972;63:509–28. 19. Weismann A. Essays upon Heredity and Kindred Biological Problems. Oxford, UK: Clarendon, 1889. 20. Weismann to Galton, 23 February 1889. In: Pearson K. Life, Letters and Labours of Sir Francis Galton. III. Cambridge, UK: Cambridge University Press, 1914-1930. 21. Churchill FB. August Weismann, Development, Heredity and Evolution. Cambridge, MA: Harvard University Press, 2015. 22. Schwartz Cowan R. Sir Francis Galton and the continuity of germplasm: a biological idea with political roots. In: Proceedings of the 12th International Congress of the History of Science, vol. 8. Paris: Albert Blanchard, 1970. 23. Galton F. English Men of Science: Their Nature and Nurture. London: Macmillan, 1874. 24. Galton F. The history of twins as a criterion of the relative powers of nature and nurture. Fraser’s Magazine 1875;12: 566–76. Reprinted in Int J Epidemiol 2012;41:905–11. International Journal of Epidemiology, 2016, 20–23 Commentary: Inheritance of doi: 10.1093/ije/dyw031 acquired characteristics: eliminating alternatives in the search for mechanisms. Commentary on Galton F: Feasible experiments on the possibility of transmitting acquired habits by means of inheritance BG Dias Emory University School of Medicine, Department of Psychiatry and Behavioral Sciences, Yerkes National Primate Research Center, Atlanta GA 30329, USA. E-mail: [email protected] Accepted 11 January 2016 Increasing scientific evidence suggests a role for the inheritance of physiological and behavioural traits across generations.1–17 By way of clarification, the inheritance being talked about refers to traits that have first been acquired by an ancestral generation as a consequence of an environmental trigger and then subsequently inherited by the descendant generations. This is in contrast to congenital traits that are inherited from the ancestral lineage. Several broad conceptual questions dominate critiques about the inheritance of characters acquired by ancestral populations and ought to be addressed if such inheritance is to be proven unequivocally.18–20 Among these: (i) is the acquisition of the trait in question a case of true biological inheritance or a consequence of information transmitted across generations via social means? and (ii) over how many generations should this acquisition be observed to C The Author 2016; all rights reserved. Published by Oxford University Press on behalf of the International Epidemiological Association V
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