REVIEW Rebel Without Applause: , Developmental and the Sex-Determination Controversy Roxanne Khamsi ’02 With the recent mapping of the human single instruments, but less than the productive genome, scientists have dramatically increased machinery of a laboratory or research field” their ability to explain inheritance patterns. Yet, (Kohler, 2001). Indeed, Goldschmidt’s choice of even so, developmental genetics remains relative- Lymantria as the organism for his studies on ly elusive. The discrepancy between these two genic balance during the early twentieth century approaches to genetics is not a new divide in the may have influenced the historical reception of research community. Richard Goldschmidt is his work. one scientist in the past who attempted to bridge Those who have sought to revive a dis- this divide.As a German scientist (fed on the cussion regarding his science, including C. Lynne holistic notion of Entwicklungsmechanik, which Littlefield, Peter Bryant, and Stephen emphasizes a physiological approach to embryol- Gay Gould, emphasize the questions he raised as ogy), he felt compelled to relate action to paramount to the conclusions he reached. In the problem of transmission. Consequently, he their eyes, he did the discipline of genetics a serv- confronted scientists of his time with several ice by highlighting the shortcomings of the stat- controversial developmental theories. In his ic genomic model. Goldschmidt’s rebellious keynote speech at the 1951 Cold Spring Harbor attack of the conventional concept of the gene Symposium, Goldschmidt upset many attendees may have indeed induced a healthy debate by proposing that the constituted among geneticists, but it undoubtedly cast a stub- the unit of function and that there was no such born shadow over his authority with respect to thing as a gene.His suggestion has become a the theory of genic balance. stock joke for historians. Nonetheless, the obscured significance of his work proves more NO FOREIGNER TO SCIENCE interesting than the humor of his inaccuracies. Many questions surrounding Richard Goldschmidt was born on April Goldschmidt’s career deserve the attention of 12, 1878, in -am-Main, , into a history: Why did his Drosophila work of the well-respected family (Stern, 1980). This social 1930s and 1940s go widely unnoticed? (James and cultural elite to which he belonged owed Crow calls Goldschmidt’s article on chromosome their status more to education than maps in the fly “particularly foolish” and later (Dietrich, 1995). Indeed, Goldschmidt himself refers to a genetic theory of sex determination demonstrated a passion for learning. By the age proposed by Calvin Bridges—rather than the of seventeen, he could read French, English, émigré German (Crow, 2000). Robert Kohler Italian, Latin and Greek and attempted to edu- fails to properly crown Goldschmidt as a “Lord of cate himself about the works of difficult philoso- the Fly” in his well-known book.) What made his phers (Stern, 1980). He enrolled in the experiments on Lymantria so controversial and Heidelberg University as a medical student and later forgotten? To answer these questions we left for Munich two years later, after passing his must understand Goldschmidt as a scientist geo- examinations. Abandoning his medical training, graphically, ideologically and historically dis- Goldschmidt became a student of Richard placed. We are also obliged to take into account Hertwig, and by the age of 21, he had completed the material culture of his experimental practice, his first scientific paper. After returning to described by Kohler as “something more than Heidelberg to write his thesis and giving a year of his life to the compulsory training period in the was not ignored by his contemporaries. German army, he joined Hertwig’s staff in Gilbert states that Goldschmidt strove for Munich, where he stayed through the early 1910s a noble life and considered himself “a self-aware (Stern, 1980). king in the scientific world ofinterbellum According to Scott Gilbert, Goldschmidt Germany” (Gilbert, 1988). The scientist’s contro- battled anti-Semitism throughout his scientific versial character embodied the stereotypical career. Because of his Jewish background, he German professor of his time. According to encountered difficulties while researching under Gould,he was ‘arrogant,haughty,yet invariably Hertwig. Gilbert writes, “Goldschmidt made the kind and even courtly” (Gould, 1982). But most of his reputation while working as Richard Goldschmidt’s life changed drastically in the half Hertwig’s assistant, and he could never hope to decade preceding World War II. In 1936, he was get a tenured position in the university even forced to flee Nazi Germany, leaving behind the though he essentially ran Hertwig’s laboratory Kaiser-Wilhelm Institute where he had been a and taught Hertwig’s courses” (Gilbert, 1988).Yet director for 23 years (Dietrich, 1996). Goldschmidt triumphed over the anti-Semitic Goldschmidt suffered a blow to his authoritative sentiment of his contemporaries when he position in Germany when he immigrated in the became a division director of the Kaiser-Wilhelm 1930s. While he had enjoyed a celebrated status Institute for Biology in 1913. as one of the most imposing figures of German Between the first and second World intellectual life during the pre-Hitler days, he Wars, Goldschmidt established himself at the became “one of the large number of professors forefront of science. His experiments on sex whose prestige was limited” when he came to the determination and differentiation in the gypsy (Stern, 1980). According to Stern, moth Lymantria dispar led him to conclude that Goldschmidt’s status was questioned by many of sex determination was controlled by interactions his colleagues at this time. He writes in his biog- between specific male and female factors—and raphy of his mentor, “The attacks against their that the potency of these factors varied between opinions, which he had waged for so long and races. Without the help of sex-linked , their critical attitude to his own pronouncements he did his best to interpret the chromosomal did not provide a background for ready accept- constitution of the individuals during the course ance except where the person, not the scientist of his investigations (Littlefield, 1980). Based was involved” (Stern, 1980). Goldschmidt irritat- upon the results of his experiments, he produced ed researchers when he assaulted the theory of the concept of genic balance,the time-law of the gene in “Morgan’s country.”During this time, intersexuality and theories about heterochro- he continued to reject the corpuscular picture of matin and sex determination. The first of these the gene and maintained that regions of specific ideas proposes that competitive interactions function along the chromosome formed a single, between sex-determining factors on different long molecule (Caspari, 1980). Most geneticists, (rather than individual chromo- however, had accepted a model in which the somes alone) determine sex (Littlefield, 1980) location of specific on specific chromo- Thus the theory of genic balance, with its empha- somes.They pointed to studies ofradiation- sis on complex systems, provides a representative induced mutations as evidence to support the example of the holistic approach that grounded model of corpuscular genes. Goldschmidt’s science. And though his theory’s In “An Evaluation of Goldschmidt’s Work flaws were later revealed, the model was initially after Twenty Years,” Ernst Caspari reasons that well received. Littlefield and Bryant state that, because Goldschmidt’s views ran counter to “The theory of genic balance as outlined by those of the majority, he found himself scientifi- Goldschmidt met with general approval from cally isolated and his work widely disregarded other workers studying lepidopteran intersex- when he came to the United States (Caspari, es…” (Littlefield, 1980). His Lymantria research 1980). Goldschmidt, who argued that “the whole

33 conception of the gene” was obsolete, found him- and 1940s (Caspari, 1980). self increasingly marginalized by the predomi- Instead of attributing the idea of genic nant idea of a static genome (Comfort, 2000). balance to Goldschmidt, Lords of the Fly gives According to Gilbert, credit to Bridges. “When Richard Goldschmidt By 1938, Goldschmidt had already alien- laid claim to his theory of sex determination,” ated himself from the majority of geneti- writes Kohler,“Bridges refused to argue.”The cists with a series of increasingly serious word “his” in this sentence offers problematic breaks with the genetic “orthodoxy” of readings. I understand “his” to be synonymous to the Morgan School…[He] disagreed with “Bridges’s” because of its context within the para- the simple chromosomal genetics of sex graph (the previous seven sentences discuss the determination espoused by Morgan, scholarship student at Columbia rather than the Bridges, and Sturtevant, preferring German outsider).In elevating the significance instead the physiological approach of the of Bridges’s work, Kohler creates a self-serving German school that he helped lead presentation of the theory of genic balance of sex (Gilbert, 1988). determination. (He did, after all, write a book about Drosophila and not Lymantria.) Ascribing When Goldschmidt announced that “The genic balance to American researchers, he theory of the gene is—dead!” (Stern, 1980) most aggrandizes the importance of Morgan’s fly geneticists actually believed that the concept was group at the expense of historical accuracy. alive and well. His keynote speech at the 1951 Kohler is not alone in his historical treat- Cold Spring Harbor Symposium struck a sour ment (or, rather, mistreatment) of Goldschmidt’s note with scientists studying heredity. research on genic balance. In his essay celebrat- Goldschmidt aggressively defended his own sci- ing the diamond anniversary of the publication entific views, but his intellectual independence Genetics, James Crow applauds Bridges for his cost him his popularity. article on genic balance and leaves out any men- tion of Goldschmidt’s claim to the concept. The frequency, phenotype and synapsis and segrega- WHERE IS RICHARD tion patterns of nondisjuctional types “led GOLDSCHMIDT? Bridges to his genic balance theory of sex deter- In his book, Lords of the Fly,Robert mination” (Crow, 2000). Like Kohler, Crow ideal- Kohler examines the network of scientists under izes Bridges’s persona. The small-town orphan who engineered the use of turned bad-boy scientist died unexpectedly in Drosophila as an essential laboratory instrument 1938 at age 49—making him appear in many in genetic research. Kohler briefly mentions texts as the James Dean of genetics. His magnan- Goldschmidt’s work in several chapters ofthe imous and seemingly humble character, as pre- text. He contrasts the German scientist’s holistic sented by the two aforementioned authors, con- biological approach with the “narrowly reduc- trasts sharply with the haughty comportment of tionist perspective of genetics” produced by the scientific “Bosses” who reigned in the labora- Morgan’s fly group (Kohler, 1994). More interest- tories during the early twentieth century. ingly and perhaps more memorably, he uses Furthermore, Crow highlights the reconciliatory Goldschmidt’s claim to intellectual priority on letter Bridges wrote to Muller and remarks that the topic of genic balance to illustrate the toler- the founder of the Drosophila Information ance and generosity of Calvin Bridges, who, Service was underpaid throughout his life.When according to Kohler, “never held a Crow does speak of Goldschmidt’s articles, he grudge”(Kohler, 1994). In limiting his mention of uses them merely to stress that earlier works in Goldschmidt’s research, Kohler continues the Genetics were not always in English (implicitly tradition of ignoring the Drosophila work that inviting the reader to exclaim, “Look how far the foreign scientist produced in the late 1930s we’ve come!”). He ridicules the German scien- tist’s second paper in the publication,which gled story of authorship looming behind genic argued against chromosome maps in Drosophila balance. Dunn initially states, “[T]he most spec- (Crow, 2000). Crow omits Goldschmidt from the tacular application of the genic balance theo- scientific narrative behind genic balance while ry…was attained in Bridges’ (1922) interpreta- casting a glowing spotlight on Bridges, a tion of sex types in Drosophila” (Dunn, 1965). He Drosophilist like himself. then proceeds to acknowledge that the ideas There is, of course, evidence of important involved in this theory had been previously pro- contributions made by Calvin Bridges to the the- posed by Goldschmidt. Dunn reasons that ory of genic balance. In 1921, using the genetic Goldschmidt failed to garner recognition to the manipulations made possible with Drosophila, same extent as Drosophilists working on the Bridges analyzed the effect of obtained triploid problem because he produced gypsy moth inter- (2X3A) intersexes by crossing triploid (3X3A) sexes within the normal diploid genome and did females to diploid (XY2A) males (Littlefield & not identify or associate genes leading in the Bryant, 1980). The relatively advanced analyses male or female direction with specific chromo- in the fly labs enabled him to find that sex was somes (and was therefore unable to directly determined by a quantitative relationship measure the relative strengths ofhis factors). between the number of X chromosomes and the Dunn writes, number of sets of autosomes. Reluctant to accept Thus, although Goldschmidt was Goldschmidt’s idea that the alleles of just one or undoubtedly the originator of the bal- two genes controlled sex, Bridges concluded that ance theory of sex, the convincing nature there were many sex-modifying genes in the ofthe proofs provided by Bridges and genome.In this fly model,he outlined that others working on Drosophila prevailed female modifiers dominated the X chromosome in giving that work greater influence while male modifiers governed autosomes. Each upon the subsequent development of the organism’s sex was determined by the ratio genic balance theory (Dunn, 1965). between these two types of modifiers; intersexu- als arose as a result of abnormal ratios between Caspari offers a more recent and celebratory per- female and male sex-modifying genes.According spective on Goldschmidt’s claim to this theory. to Littleton and Bryant, He presents a broad narrative of Goldschmidt’s Bridges pointed out that although place in the , emphasizing that Goldschmidt had occasionally found the reception of the scientist’s work changed with with Lymantria that the strength of male the in the scope of the discipline factors became altered and had attributed between the 1930s and 1940s (including “the rise it to a mutational event, recombination of biochemical genetics, microbial genetics and between several closely linked sex-modi- finally molecular genetics”). His essay reveals fying genes could have given the same that during the 1950s, when new complexities result (Littlefield & Bryant, 1980). challenged earlier models of the gene, geneticists once again turned to Goldschmidt’s research and Bridges’s research expanded and challenged that ideas (Caspari, 1980). of Goldschmidt. Furthermore, the organism he Caspari portrays a positive turn in the chose for his study had a wide appeal in the popularity of Goldschmidt’s science; however, American scientific community. Although histo- recent texts (such as those of Kohler and Crow) rians such as Kohler and Crow have good reason continue the tradition of mocking and neglect- to sing the praises of Bridges’s studies, they nev- ing the German émigré. Surveying the table of ertheless deceive their readers by playing down contents and indexes of these books, a careful Goldschmidt’s own contributions to the theory reader begins to wonder,“Where is Richard of genic balance. Goldschmidt?” When authors attribute the theo- Both Dunn and Caspari address the tan- ry of genic balance to Bridges alone, they demon-

35 strate their bias in favor of American scientists flies in genetic research. In fact, Lords of the Fly working under Morgan as well as their prejudice specifically focuses on the dramatic success of against a foreign geneticist who went so far as to Morgan’s researchers in using Drosophila in a attack the classical concept of the gene. revolutionary system of genetic exploration. The new mutants and new genetic knowledge fueled THE MATERIAL BASIS FOR FAME more of the same in an autocatalytic process. In If anti-Semitism, American patriotism Kohler’s words, “Drosophila became,in effect,a and “Drosophile” forces have marginalized biological breeder reactor, creating more materi- Goldschmidt’s science, his choice of Lymantria as al for new breeding experiments than was con- the organism for his studies also detracted from sumed in the process” (Kohler, 1994). Morgan’s the permanence of his fame.While he conducted network of labs became a “hegemonic” establish- his experiments, collecting samples from around ment and many bright minds were eager to join the world, the Fly Group managed to develop the fly bandwagon. Goldschmidt’s preferred revolutionary analytical technologies and give organism for his sex-determination research researchers, such as Bridges, a scientific edge. (Lymantria) therefore placed him on a path that Littlefield and Bryant assert that Goldschmidt diverged from the overwhelmingly dominant Fly had chosen an organism that “is by no means Group. ideal for genetic manipulations…” (Littlefield & Goldschmidt did more than just hold off on Bryant, 1980). In contrast, Bridges had a wealth becoming a Drosophilist; he maintained a holis- of new techniques to examine his Drosophila. tic approach to the study of heredity that ran Moreover, his use of flies in his investigations contrary to the pragmatic style of American concerning genic balance came with a built-in geneticist (Dietrich, 1996). According to Gilbert, audience that only grew with time. This audience Goldschmidt sought “to place American genetics existed because Morgan had institutionalized a into the German type of developmental physiol- system of etiquette that enabled the ogy”(Gilbert, 1988). This challenge presented Drosophilists to “make such progress in genetics great difficulty at the time. Employing Wilhelm that by 1920 they dominated the field” (Dietrich, Johansenn’s distinction between genotype and 1996). The prevalent use of Drosophila in genet- phenotype,“Morgan had separated the transmis- ics today remains as evidence of the organism’s sion of hereditary traits (genetics) from the popularity as a scientific tool. expression of those traits (embryology)” In selecting Lymantria for his research, (Gilbert, 1988). Using Drosophila to churn out Goldschmidt adopted what Robert Kohler would dazzling discoveries, the Fly Group made this call a non-standard “system ofproduction.” question of gene transmission more exciting Kohler urges historians to consider organisms than that ofgene action.The used in the laboratory, specifically those that Entwicklungsmechanik perspective Goldschmidt were altered for the purposes of the experiment, maintained cost him the esteem of scientists in as constructed artifacts (Kohler, 2001). America for many years. Although he came to “Standard” animals, produced by genera- the United States and devoted himself to analyz- tions of inbreeding are unambiguous ing the spontaneous mutations in Drosophila examples of engineered from 1939 to 1945 (Dietrich, 1996), Goldschmidt instruments…Organisms and instru- refused to surrender his complex view of heredi- ments come to embody libraries of ty. Thus, even though he adopted the material of knowledge and routine practices. That, Morgan’s labs, he remained at odds with many by definition, is how they become sys- geneticists of his time and failed to break into the tems of production (Kohler, 2001). limelight in history books.

The practice of using Lymantria was not popu- larized to anywhere near the extent as the use of THE REBEL’S REDEMPTION could “fit their parts into the yet imaginary struc- Those who seek to resurrect discourse ture” (Stern, 1980). The German Boss was, in about Goldschmidt’s work bring attention to the Stern’s opinion, a designer of frameworks of the cruel treatment he has received. In the introduc- future (Stern, 1980). tion to Goldschmidt’s book, The Material Basis of Additionally, Goldschmidt’s holistic per- Evolution, Gould writes that the scientist “suf- spective has gained newfound appreciation fered the worst fate of all: to be ridiculed and among historians. Littlefield and Bryant con- unread” (Gould, 1982). In science courses, stu- clude that his proposal that genes control rates of dents laugh at Goldschmidt’s ideas dutifully to reaction “shed one of the first rays of light into prove that they are not guilty of either ignorance the still-very-dark black box that connects the or heresy (Gould, 1982). His heretical denial of genotype with the phenotype” (Littlefield & the conventional gene, however, provides histori- Bryant, 1980). These commemorative statements ans with more than comedy. It presents one pos- give Goldschmidt the applause that the majority sible explanation for the neglect of his theories. of “Drosophile” historians do not offer in their In the revisionists’ story, most geneticists were texts. not ready to confront the challenging models he championed. Goldschmidt’s own audacity cast a THE EVOLUTION OF CELEBRITY shadow over his insight. Just as Kohler and Crow If we evaluate Goldschmidt’s refutation of depict Bridges as a social rebel censured for his the classical concept of the gene out of context, it promiscuous behavior, Allen and Dietrich por- becomes easy to ridicule. This presentist tenden- tray Goldschmidt as a scientific rebel outcast for cy to find humor in his mistaken models serves his rejection of the classical concept of the gene. no constructive purpose. Instead, it merely Seen in the context of his time, Goldschmidt diverts our attention from his authority on ideas acquires a new allure of someone who possessed such as genic balance. Kohler and Crow both err the courage to stand apart from the crowd to in de-emphasizing Goldschmidt’s theoretical promote what he believed. contributions. We cannot write him into the Building upon this interpretation of story of the Fly Group, but we must not careless- Goldschmidt as an intellectual rebel, Allen and ly disregard him either. Comfort suggest that he should be remembered It would incorrect to conclude that histo- for the kinds of questions that he posed if not for rians have reached a definitive agreement over his actual research. His contrarian views— the importance of Goldschmidt’s role in genetics. including his struggle against the notion of a One need only compare Kohler’s discrediting static genome—pushed the discipline of genetics presentation of Goldschmidt’s relationship to the forward. According to Comfort, “By playing the theory of genic balance with the earlier essay by devil’s advocate, Goldschmidt forced geneticists Littlefield and Bryant to see that the debate con- to reconsider their assumptions. For all his tinues. Dietrich has recently written on the crotchetiness, Goldschmidt was and is an impor- exceptional significance of the German scientist’s tant figure in genetics” (Comfort, 2000). work. He writes, “Goldschmidt’s work stood Similarly, Allen writes, “Goldschmidt may often apart from typical Mendelian accounts by pro- have been wrong, but his ideas had imagination viding a quantitative interpretations of sex deter- and fire” (Allen, 1980). He adds that,“Our under- mination” (Dietrich, 1996). Thus, our views on standing of,and approach to,genetic processes Goldschmidt’s importance continuously evolve. today bears the mark of the kinds of questions His place in history is by no means static. which Goldschmidt persisted in asking, often irreverently, throughout his entire career” (Allen, 1980). Stern likens Goldschmidt’s thinking to the scaffolding of a building on which other scien- tists were able to reach the spaces where they

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