Caution, Overload: the Troubled Past of Genetic Load
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| PERSPECTIVES Caution, Overload: The Troubled Past of Genetic Load Amir Teicher1 Department of History, Tel Aviv University, Israel 6997801 ORCID ID: 0000-0002-6890-2073 (A.T.) KEYWORDS genetic load; history; eugenics; psychiatry; radiation; population genetics REOCCUPATION with “genetic load” seems to be on the Dowbiggin 1985, p. 191). In mid nineteenth-century France, Prise again, fueled by the new opportunities that full hu- the notion that heredity, and in particular morbid heredity, man genome sequencing provides (e.g., Simons et al. 2014; was the underlying cause of a variety of nervous and mental Lynch 2016a; Stewart et al. 2017; Verduijn et al. 2017). The disorders, was already becoming widespread. Even though fear of the ultimately devastating effect that the accumula- there was no precise understanding of the mechanism gov- tion of deleterious mutations would have on the future of erning hereditary transmission at the time, the fact of herita- humanity was articulated nearly 70 years ago by Hermann bility itself seemed undeniable, and apparently could explain J. Muller in a 1950 seminal paper, Our Load of Mutations. why, in certain families, social, medical, and psychological Four decades later, in his 50 Years of Genetic Load: An Odyssey, deviations were so abundant. These ideas received their Bruce Wallace (1991) recounted how the concept of genetic clearest expression in Bénédict Augustin Morel’s Treatise on load had evolved in the twentieth century, and traced its the Physical, Intellectual and Moral Degeneracy of the Human origins further back to J. B. S. Haldane’s 1937, The Effect of Race (Morel 1857). For Morel, degeneracy was a process of Variation on Fitness. This paper by Haldane, wrote Wallace, progressive mental, physical, and moral deterioration from “marks the origin of the genetic load concept.” Both points of one generation to the next, whose initial impetus may have departure, however—Muller (1950) as well as Haldane been external (poor nutrition, urban living conditions, ner- (1937)—furnish a history of the concept “genetic load” that vous strain from industrial work, and alcohol) but whose in- utterly ignores its eugenic roots, manifested most clearly in fluence was inherited and whose ultimate result was the the concept’s German predecessor: erbliche Belastung.To production of imbeciles and the annihilation of entire familial properly assess the meaning and complexities of the concept stocks. of genetic load, as well as its potential social implications, it is The same idea was also developed by other contemporary vital that we acknowledge these scientific and cultural ori- thinkers, in and outside of France (reviewed in Chamberlin gins, some of whose trajectories are still pertinent today. and Gilman 1985; Pick 1989). In Italy, for example, Cesare Lombroso related “born” criminal behavior to degeneracy; impressed by Lombroso’s writings, and under the impression Origins of the Concept of Hereditary Load that, “degenerates are not always criminals, prostitutes, an- “[T]oday we possess numerous examples, not only of trans- archists and pronounced lunatics; they are often authors and mission but as well of hereditary accumulation of morbid artists,” the Hungarian-born doctor and social critic Max predisposition [to mental anomalies].” These are the words Nordau characterized all fin-de-siècle “decadent” cultural of L. F. E. Renaudin, Director of the mental asylum at fashions as expressions of moral and social degeneracy. Maréville, in his 1854 Medical Psychological Studies on Men- Nordau defined degeneracy as “a morbid deviation from an tal Alienation (Renaudin 1854, p. 33, translation following original type. This deviation, even if, at the outset, it was ever so slight, contained transmissible elements of such a nature Copyright © 2018 by the Genetics Society of America that anyone bearing in him the germs becomes more and more doi: https://doi.org/10.1534/genetics.118.301093 incapable of fulfilling his functions in the world; and mental Manuscript received May 1, 2018; accepted for publication May 22, 2018. fi 1Address for correspondence: Department of History, Tel Aviv University, Tel Aviv, progress, already checked in his own person, nds itself Israel 6997801. E-mail: [email protected] menaced also in his descendants.” (Nordau 1895, p. vii, 16). Genetics, Vol. 210, 747–755 November 2018 747 The concept of degeneration was also popular in the reasoning to the psychiatric domain indicated that most men- Anglo-Saxon world (e.g., Talbot 1898). In America, studies on tal diseases were recessive and that, even when dominant “social degenerate” families (the Jukes, the Ishmaels, the factors were involved, the more severe mental aberrations Kallikaks) and investigation into degenerate heredity by such were caused by recessive variants (Davenport 1908; Rüdin prominent scientists as Charles B. Davenport and Henry H. 1911; Lenz 1912, p. 597). When it came to hereditary load, Goddard, ostensibly furnished the links between crime, however, the concept of recessivity also enabled, at least in poverty, alcoholism, and mental deviancy, and helped frame theory, the narrowing down of the idea of familial burden to social problems as inherently biological ones (see discussions only those members of a family who really carried harmful by Rafter 1988 and Carlson 2001). To counter the looming hereditary factors. “Every individual, in whose family circle... degeneration of human germplasm, some form of counter- mental illness appeared, was termed ‘belastet,’” recalled regeneration was needed. The British polymath Francis psychiatrist Ernst Wittermann in 1913, referring to pre- Galton, Charles Darwin’s half-cousin, proposed the term “eu- Mendelian times. He explained, however, that Mendelian the- genics” to define the means and methods to be mobilized for ory supported a distinction between homozygous individuals, taking control over human evolution and for redirecting it who were free from a certain recessive disease (DD), and toward a better path, whether by checking the reproduction heterozygous carriers (DR). As long as Belastung continued of those deemed less fit, or by boosting the propagation of to be a familial concept, Wittermann (1913) argued, the two those who were of higher quality. Throughout this period, types remained indistinguishable, despite the fact that only when French, American, and British scholars wanted to ad- the second type of individuals (DR) were truly belastet. dress the nature of degeneracy or hereditary processes, they The mid 1910s saw the establishment of a novel kind of spoke quite generally of hereditary transmission, hereditary analysis of hereditary relations in German psychiatry. The particles (Darwin’s “gemmules”) or even accumulating he- limitations of simple Mendelian schemes were already reditary influences or “taint.” Disregarding religious- and becoming apparent, and psychiatrist Ernst Rüdin, greatly economic-related contexts, they did not yet use in any sys- assisted by Wilhelm Weinberg (better known today for the tematic fashion the term hereditary “load” or “burden.” Hardy-Weinberg equilibrium), developed innovative meth- Their German-speaking counterparts, however, did. The ods to assess the hereditary risk that the mentally ill posed term Belastung in German is translated as strain, load, or to their relatives (Weinberg 1912, 1913a,b, 1929; Rüdin burden. When accompanied by the adjective, erblich (hered- 1916; see also Just 1920; Crow 1999). The new statistical itary) this term was often used to refer to mentally ill indi- toolbox, which quickly acquired international recognition viduals in one’s family circle or community. Throughout the and some of whose results are considered robust to this nineteenth century, German psychiatrists made repeated at- day, was officially referred to under the heading, “empirical tempts to assess the relative significance of heredity in the hereditary prognosis” (Rüdin 1933), but was often labeled causation of mental abnormalities (see Gausemeier 2015). In simply as “statistics of load” (Belastungsstatistik). Different 1895, for example, an assistant at the Burghölzli psychiatric scholars published the results of their studies on the “load” clinic in Zurich by the name of Jenny Koller turned “heredi- of the “average population” with respect to various mental tary load” into her direct object of inquiry. Koller (1895) not disorders, and evaluated the chances that children, siblings, only offered a detailed analysis of the frequencies of different or even nephews or nieces of the mentally ill would mani- mental disorders in the families of 370 asylum patients, but fest mental deviances (Kattentidt 1926; Schulz 1927, 1931; also compared these results to a control group of (reportedly) Luxenburger 1928; Brugger 1929; Panse 1935). healthy individuals; this was a true methodological novelty at As in the case of the American eugenics movement, the the time. Ten years later, another scholar from the same in- underlying agenda informing these works was that families stitute, Otto Diem, published an additional massive study on with members displaying mental illnesses—and not only the the degree of Belastung of both mentally ill and healthy indi- mentally ill themselves—needed to be socially isolated and viduals. Diem (1905) stated with dissatisfaction that, “[i]t their reproduction stemmed. The calculation of morbidity has evolved into an actual dogma that hereditary load is risk for nephews and nieces of a disease sufferer (Schulz the most important