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Opportunities for Women In Early Genetics

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  See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/5950334 Opportunities for women in early genetics  Article   in  Nature Reviews Genetics · December 2007 DOI: 10.1038/nrg2200 · Source: PubMed CITATIONS 6 READS 17 1 author: Marsha L RichmondWayne State University 32   PUBLICATIONS   170   CITATIONS   SEE PROFILE All content following this page was uploaded by Marsha L Richmond on 13 January 2017. The user has requested enhancement of the downloaded file. All in-text references underlined in blue are added to the srcinal documentand are linked to publications on ResearchGate, letting you access and read them immediately.  The rediscovery in 1900 of Gregor Mendel’s seminal 1866 paper on hybridization in peas ushered in a new approach to heredity that soon spawned the birth of genetics. The epis-temic aspects of the founding and early years of this new discipline have been well surveyed by historians. We now have a good under-standing of how rapidly biologists recognized the potential breakthroughs that were offered by Mendel’s laws and flocked to the study of heredity, variation and evolution, which in 1906 the British biologist William Bateson named ‘genetics’. The new methodology that was used by geneticists — controlled hybrid crosses followed by statistical analysis of the appearance of alternative characters among the progeny — gave biologists a powerful new tool by which to explore the mechanism of heredity. Within the space of only a few years, confirmation and extension of Mendel’s results led to the general acceptance of the  validity of genetics. Thomas Hunt Morgan’s 1910 discovery of the ‘white-eye’ mutation in Drosophila melanogaster   marked a crucial turning point, culminating in the 1915 publication of The Mechanism of Mendelian Heredity   by Morgan, Sturtevant, Muller and Bridges, which, combining Mendelism with the chromosome theory of heredity, ushered in the ‘new genetics’. This work heralded the remarkable advances of the classical period of genetics (1900–1935), which revolutionized not only studies of heredity, but also most areas of the life sciences 1–4 . Women in early genetics Although this brief sketch of the early his-tory of genetics is broadly accurate, it fails to appreciate a cardinal feature that is less well recognized — namely, that women were well represented among the field’s initial workers. Indeed, genetics was one of the earliest emerging disciplines in twentieth-century biology to benefit from the contributions of women. This knowl-edge is newly won. None of the leading historical surveys of genetics mentions that a significant number of the early contributors to genetics were women. However, recent scholarship has begun to trace the roles of women in advancing early Mendelian research and classical genetics 5–9 . One interesting finding is the close connection between the opening of higher education to women in the last three decades of the nine-teenth century and the entry of women in greater numbers into the scientific workforce at the turn of the century. Importance of higher education of women.   Emerging as a new field of study at precisely the time that Western women were graduat-ing from universities in increasing numbers, genetics highlights general trends pertaining to women in science. After 1900, middle-class women demanded greater accessibility to the work force. Many were especially well poised to pursue advanced research in sci-ence, having gained entry to university study in the sciences in the United Kingdom and the United States from the 1860s. By 1900, a significant number of women in these coun-tries had received excellent undergraduate and even graduate training in biology at leading universities, and they sought oppor-tunities to pursue advanced research and a career in science 10–13 . Research opportunities were opening worldwide, as economic development promoted the establishment of university laboratories, as well as private and governmental establishments and industrial research complexes 14–23 . These developments created unprecedented opportunities for women to enter science as a career 22 . As the President of the University of Chicago stated in 1901, “The women now being graduated, with the Doctor’s degree, from our strongest institutions, are, in almost every particular, as able and as strong as the men. If oppor-tunity were offered, these women would show that they possess the qualifications demanded.” 23  An international trend.  Similar trends were occurring in other countries. The first female students entered Dutch universities in the early 1870s, and by 1900 almost 5% of the student population was female, half of whom studied mathematics or the natural sciences. In Germany, although universities were not widely open to female students until after 1900, a number of women were able to receive an education in the sciences and were eager to pursue advanced research 6,24,25 . Russian women participated in the work of the two leading centres for genetics: Nikolai Kol’tsov’s Institute of Experimental Biology in Moscow and Nikolai Vavilov’s group in St Petersburg (known at the time as Leningrad). It was not accidental, then, that a significant number of well qualified women were eager to join the various research groups that began to emerge in genetics after 1900. Women were important to new fields Biologists undertaking Mendelian studies welcomed the opportunity to gain qualified workers to expand their experimental efforts in a field that had not yet gained full scientific authority, and hence was not widely attractive to young male research TIMELINE Opportunities for women in early genetics Marsha L. Richmond  Abstract | Although women have long been engaged in science, their participation in large numbers was limited until they gained access to higher education in the last decades of the nineteenth century. The rediscovery of Mendel’s work in 1900 coincided with the availability of a well trained female scientific workforce, and women entered the new field in significant numbers. Exploring their activities reveals much about the early development of the field that soon revolutionized biology, and about the role of gender in the social organization of science. NATURE REVIEWS |   GENETICS  VOLUME 8 |  NOVEMBER 2007 |   897 PERSPECTIVES  ©    2007   Nature Publishing Group    students seeking to advance their future career prospects (this phenomenon is discussed by sociologists in the context of queuing theory, with the expectation of finding women in fields that are not yet  viewed as legitimate 12,26 ).Often, those who accepted female workers were sympathetic supporters of the ‘woman question’ and served as helpful mentors. This was an important prerequisite for working in science at a time when women were not generally admitted to scientific societies and therefore had limited access to established organs of publication 27 . They also generally lacked access to the the institutional infrastructure that sup-ported extended work in science, including studentships, grants, fellowships and estab-lished career tracks in university teaching 28 . The new field of genetics, which was not yet well institutionalized, offered these highly trained women the means to participate in and contribute to an area of study that held great promise for advancing biology. ‘Invisible technicians’ Barriers to the participation of women.  Between 1900 and 1935, many women seized the opportunity, and the excitement, that was offered by Mendelian genetics, and helped to advance knowledge in heredity (see below). Nonetheless, women faced several visible and invisible barriers that restricted their full participation in the scientific community, as well as their ability to pursue extended careers in science 12,29–32 . As women began to enter the scientific workforce in greater numbers, a system developed that restricted them to certain roles and job titles, even after they gained advanced certification and a master’s or doctoral degree. They fitted into a system that rendered them the modern counter-parts of the ‘invisible technician’, described by the science historian Steven Shapin as assisting the men who pursued chemical and physical studies in the seventeenth century  33 .   Women were present, for example, in Morgan’s laboratory at Columbia University, New York, USA, just as they were at other leading centres of early genetics, including the Cold Spring Harbor Station for Experimental Evolution, USA, which was established in 1904 and directed by Charles Benedict Davenport, and the John Innes Horticultural Institute, London, UK, which was headed by William Bateson when he left Cambridge in 1910. However, the job titles they held were most often those of ‘assistant’, ‘technician’, ‘stock-keeper’ or even sometimes unpaid working wife 7,34–36 .   Michael Dietrich and Brandi Tambasco suggest that women who worked in early Drosophila genetics experienced ‘hierarchical segregation’. This suggestion was based on a demographic survey of the period from 1934 to 1970, which stated that “…while the majority of men work-ing in Drosophila  genetics were Faculty, the majority of women were Technical Assistants.” 36 This trend seems to have been established in earlier decades.It should not be assumed, however, that women holding such job titles were simply support staff. As Maria Rentetzi has shown, women who worked in early radioactivity physics were “…not merely assistants and members of the laboratory support staff, setting up experiments and performing tedious preparatory tasks for their male colleagues…” but active participants in laboratory life, making “…steady contribu-tions to radioactivity research” and being “…as scientifically productive as their male counterparts, publishing not only in the institute’s journal but in other periodicals as well.” 37,25  The same is true of the women who worked in the Manhattan Project in the 1940s. Although Margaret Rossiter, in her classic text on women in American science, could count only 11 female ‘scientists’ work-ing on this project, there were, it turns out, more than 300 women who were classified as technicians. Of these, many “…held bach-elor’s degrees, some even in the appropriate scientific fields, but as was the practice in that era, women were given lesser job titles and far lower pay than men with similar backgrounds.” 38  (see also REFS 12,39 ) The question thus becomes, were the women working in genetics at such leading centres as Cold Spring Harbor and the John Innes Institute (as well as in university settings) merely technicians, or were they actually doing work on a par with men who held the position of investigator?The system of hierarchical segrega-tion resulted in many of the women who contributed to early genetics remaining invisible in the historiography of genetics. Despite publishing papers and advancing the programmes of their respective research groups, few were able to pursue a long-term academic career. This was especially true after genetics gained increasing legitimacy following the 1915 publication of The  Mechanism of Mendelian Heredity  . Few women, even those who received their doctorates under Morgan, were able to sur-mount the existing barriers and continue to work in genetics 12 . Many women geneticists therefore became ‘silent scientists’ — those who did not publish beyond their disserta-tion 40 . Some women moved to other fields, and those who married generally left science altogether, which was the common practice among working women before the 1930s. Even those who succeeded in making a Box 1 | Edith Rebecca (Becky) Saunders (1865–1945) in her garden allotment The daughter of a hotel keeper in Brighton, England, Saunders was educated at Handsworth Ladies’ College. She entered Newnham College, Cambridge, in 1884, taking Part I (class II) of the Natural Sciences Tripos in 1887, and Part II (class I) in 1888. She continued into postgraduate research as a Bathurst student, 1888–1889, and served as a demonstrator in botany at the Balfour Biological Laboratory for Women, becoming director in 1899. She held a College Research Fellowship, 1906–1909, and was the director of studies in natural science at Newnham College, 1918–1925, and Girton College, 1904–1914. She was a fellow of the Royal Horticulture Society, receiving the Banksian Medal in 1906, and was one of the first women to be elected as a fellow of the Linnean Society of London (1905). She served as president of the botanical section of the British Association for the Advancement of Science in 1920 and of the Genetical Society from 1936 to 1938. She died in 1945 from injuries suffered in a bicycle accident. Image courtesy of Newnham College. PERSPECTIVES 898 |  NOVEMBER 2007 |  VOLUME 8 www.nature.com/reviews/genetics  ©    2007   Nature Publishing Group    career in genetics, however, generally remained invisible owing to the tendency for historians to focus on those who made notable (‘heroic’) contributions to the field, most of whom were men 41 . Addressing women’s participation is an important aspect of current work in the his-tory of genetics, and not simply as a quest to ‘right’ the historical record. For example, as Shapin notes, “…in the now-vast academic literature in the history and sociology of sci-ence there still does not exist a single study systematically documenting and interpreting technicians’ work, past or present.” 33  Because women were most often classified as technicians, their work has generally been overlooked, and hence they might have been actively and unfairly rendered invisible 41 . Several issues are at play here. The first is whether women holding the title of techni-cian were well trained in advanced science and made srcinal contributions, and could therefore be considered scientists. Second, even if they were simply technicians, does their work, as Shapin notes, not merit scholarly examination as being a necessary component of the scientific enterprise? Third, we currently know little about the actual organization of early genetics labora-tories, including the parcelling out of work and credit. As sociologists emphasize, science is a social endeavour in which relationships are important 23,26,29–32 . Focusing on the work of women reveals important aspects of the sociology of science, including the part that gender has played in research groups and laboratory practices in science 7 . Collecting and analysing data on women in genetics therefore promises to revise our understanding of the practice, organization, functioning and ‘moral economy’ of the laboratory, and thereby shed further light on crucial aspects of the rise of modern experimental science 42–44 . Bateson’s group as an illustration Our knowledge about women’s participation in genetics has expanded in recent years, with historians of women in science provid-ing important information about the career paths of numerous individuals 5–9,45–52 . In tracing trends, however, it is often helpful to identify collective cases. One of the most prominent examples of women working Table 1 | Women geneticists at Cambridge University, 1900–1910 NameDate of birth and deathCollege affiliationResearch Interests Darwin, Nora (Mrs Alan Barlow)1885–1989NoneTrimorphic forms (primroses, Lythrum and Oxalis )Durham, Florence Margaret1869–1948Girton Coat colour in mice and canariesKillby, Hilda Nanette Blanche Praeger1877–1962NewnhamHeredity in goats, rabbits and peasMarryat, Dorothea Charlotte Edith (Mrs J. J. Lister)1880–1928NewnhamEye colour and sex in canaries; variation in Mirabilis jalapa Saunders, Edith Rebecca (Becky)1865–1945Newnham  Biscutella laevigata , Datura , Matthiola  and  Atropa Sollas, Igerna Brünhild Johnson (Hilda)1877–1965NewnhamCoat colour in guinea pigs and wing coloration in mothsWheldale, Muriel (the Right Honourable Mrs Huia Onslow)1880–1932NewnhamFlower colour in  Antirrhinum  and plant pigmentation in generalBox 2 | (Emma) Nora Darwin Barlow (1885–1989) at the John Innes Horticultural Institute, 1913 Nora Darwin (daughter of Horace Darwin and granddaughter of Charles Darwin) studied botany at Cambridge and was a student of William Bateson’s 1906 course on ‘variation and heredity’, which he first named ‘genetics’ the same year (her notebook with lecture notes is in the Cambridge University Library). She later recalled this experience : ‘‘My first introduction to the whole subject [of genetics] … was when William Bateson was giving what we called his Bible Class, in a remote lecture room, in the back of one of the colleges. It was outside the ordinary curriculum. It was a five or six o’clock lecture. And there he introduced a small set of people into the elements of the new Genetics. Mendelism was just coming in … He was a brilliant lecturer and, of course, he had an entirely new view of ordinary heredity … It was very inspiring indeed.” 55   Following her marriage to Alan Barlow in 1911 and even after the birth of her six children (1912–1921), Nora Barlow continued to study the genetics of trimorphic species, visiting the John Innes Institute each summer until 1926 to examine the flowers that were grown for her there, and publishing the results of her work. She was among the founders of the Genetical Society in 1919 and attended its meetings regularly. Her son, Professor Horace Barlow (Department of Physiology, Development and Neuroscience, Cambridge University) recalls that when he was a child his mother did “…controlled pollinations of flowers that spent their time in muslin bags. She also went to meetings of the Genetics Society, and kept up with genetical friends such as R. A. Fisher — in fact we had at least one dog that was the result of one of his genetical experiments. But I don’t think any of this resulted in any publications on genetics” (personal communication). At her memorial service in 1989, Alex Pankhurst recognized her scientific interests: “Nora studied genetics at Cambridge, and remained fascinated by the subject. From early on she tried her hand at hybridizing various flowers, including aquilegias — her experiments indicated by little muslin bags over the flower heads. On one occasion, however, she unwisely showed her children how to break off an aquilegias spur and suck out the honey. Thereafter quite a few of her experiments were tampered with.” 56 Today, Nora Barlow is best remembered for her pioneering editing of her grandfather’s scientific and autobiographical works. Reproduced with permission from REF. 56     (1992) Cambridge Desktop Bureau. PERSPECTIVES NATURE REVIEWS |   GENETICS  VOLUME 8 |  NOVEMBER 2007 |   899  ©    2007   Nature Publishing Group    communally in early genetics comes from Bateson’s group at Cambridge University, UK. In the 1880s, Bateson rejected evolu-tionary morphology as a prime means of studying evolution, focusing instead on ‘saltatory’ variations — cases involving the sudden appearance of new kinds — as important in the production of new species. In 1895, seeking to discover the laws that govern inheritance, he began “systematic experiments in breeding” in collaboration with Edith Rebecca Saunders (1865–1945), a botany instructor at Newnham College, Cambridge, and the director of the Balfour Laboratory for Women 9,45   (BOX 1) . In 1899, Saunders published the results of breeding experiments in Biscutella laevigata,  in which she found evidence that crosses between hairy-leaved and smooth-leaved plants did not produce blended offspring, thus supporting Bateson’s belief in ‘discon-tinuous’ heredity. However, neither she nor Bateson could identify any consistent patterns in the crosses that they made. It was therefore propitious that in May 1900 Bateson learnt about Gregor Mendel’s work and found in it the laws of heredity that he and Saunders were seeking. They became immediate converts to what soon became known as Mendelism.Revising their experimental setup to reflect the kind of controlled hybrid crosses that Mendel required, Bateson and Saunders published the preliminary results of their work in 1902 in the first of five Reports to the Evolution Committee of the Royal Society  . However, they fully realized that the participation of many more workers was required to establish the validity of Mendel’s laws, especially in the face of the severe attack that was launched by the biometricians. Although Bateson had gained the assistance of several horticulturalists and amateur breeders, he needed university researchers to convince the scientific community of the legitimacy and importance of Mendel’s approach to heredity. However, he was unable to attract the students of the School of Morphology, largely because its head, Adam Sedgwick, was skeptical of both Bateson’s unorthodox  views and his methodology  9 . It was thus extremely fortuitous that Bateson and Saunders were able to recruit early workers from among the women who studied the life sciences at Cambridge. In 1901, Bateson offered for the first time his course of lectures on heredity and  variation, in which he described Mendel’s work. Aided by Saunders’s contacts with life science students from the Cambridge colleges Newnham and Girton at the Balfour Laboratory, the two soon attracted to the cause a “group of eager students”, all of whom were initially women who were reading for Parts 1 and 2 of the Natural Sciences Tripos (TABLE 1). These included:   Igerna Brünhild Johnson Sollas (1877–1965), daughter of the Oxford professor of geology, who investigated the inheritance of coat colour in guinea pigs; Florence Margaret Durham (1869–1948), Bateson’s sister in law and future UK Medical Research Council investigator, who studied coat colour in mice and coloration in canaries; Muriel Wheldale, later Onslow (1880–1932), who worked on the genetics of flower colour in  Antirrhinum; and Nora Darwin, later Lady Barlow (1885–1989) (BOX 2) , who, following in the footsteps of her famous grandfather, crossed trimorphic forms, such as plants of the genus Oxalis . Other early workers included Hilda Nanette Blanche Praeger Killby (1877–1962), who investigated heredity in goats and assisted both Bateson and Saunders; and Dorothea Charlotte Edith Marryat, later Lister (1880–1928), who studied variation in  Mirabilis jalapa . This small band of dedicated followers later expanded to include men as Mendelism gained credibility (BOX 3) , with Reginald Crundall Punnett (1875–1967) joining Bateson in 1904 and several more thereafter. By 1906, the group was recognized as the leading ‘school of genetics’ and, by 1910, had collectively produced convincing evidence to establish the validity and value of Mendelism in probing the secrets of heredity  9,45 .In 1910, Bateson left Cambridge to direct the newly founded John Innes Horticultural Institute in Merton outside London, disap-pointed by the university’s failure to fully support genetic research. Many of the men in his group continued to work in the field (although this number was diminished and dispersed following the First World War); however, the fate of the women illustrates well several of the previously discussed points about the difficulties that women faced. Several women (Durham, Killby and Darwin) followed Bateson to the John Innes Institute and continued to work for a time in plant genetics, but as invisible assistants with no formal job title (and apparently no salary). Durham left the John Innes Institute during the war to work at the Medical Research Committee (later Council), thereby securing a successful research position thanks to the opportu-nity offered by governmental support for medicine, from which she retired in 1930. Sollas gave up genetics to take charge of her father’s household in Oxford (passing along her guinea pigs and her interest in genetics to the young J. B. S. Haldane and his sister, Naomi). So too did Marryat on her marriage to the Cambridge zoologist Box 3 | The Biometrical–Mendelism controversy Even before Bateson’s conversion to Mendelism in 1900, he clashed with the views of his former friend and Cambridge colleague W. F. R. (Frank) Weldon. Weldon upheld neo-Darwinian principles of gradualistic evolution, believing that natural selection operated on minute, fortuitous variations in organisms. Bateson, on the other hand, viewed speciation as the product of abrupt changes in form, and sought evidence for this process through the study of variation and its causes. Interestingly, both Bateson and Weldon found support for their respective beliefs in the work of Francis Galton, each emphasizing different facets of Galton’s law of ancestry and his idea of regression to the mean. Weldon believed that heredity was determined by the input of traits that were derived from an individual’s parents, grandparents and other distant ancestors, in decreasing proportion. Seeking empirical evidence to support his contrasting view of discontinuous heredity, Bateson (with the assistance of Becky Saunders) began experimental crosses in the mid-1890s between varieties that exhibited alternative characters. They were thus well prepared to appreciate the significance of Mendel’s law of segregation when it became publicized in the spring of 1900. The era of Mendelism was effectively launched with the 1902 publication of their first Report to the Evolution Committee of the Royal Society  . Weldon, however, launched a vociferous challenge to Mendelism in the pages of his new journal Biometrika , supported by his colleague and fellow biometrician Karl Pearson. They argued, in part, that by focusing solely on the parental generation, Mendelians omitted the important component of ancestrian heredity, and that their results were simply a ‘special case’ of the broader laws of inheritance. Bateson, recognizing that the biometrical challenge could effectively squelch both the acceptance of Mendel’s principles and the workforce needed to establish Mendelian genetics, responded with equal vitriol in his 1902 book, Mendel’s Principles of Heredity: A Defense . This controversy, which effectively ended with Weldon’s death in 1906, has been well studied by historians, sociologists and philosophers alike, seeking to identify and analyse the major factors that contribute to the generation and resolution of scientific disputes 57–60 . PERSPECTIVES 900 |  NOVEMBER 2007 |  VOLUME 8 www.nature.com/reviews/genetics  ©    2007   Nature Publishing Group  
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