DUJS 2002S V. Ii

REVIEW Rebel Without Applause: Richard Goldschmidt, Developmental Genetics 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, Curt Stern and Stephen emphasizes a physiological approach to embryol- Gay Gould, emphasize the questions he raised as ogy), he felt compelled to relate gene 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 chromosome 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 Frankfurt-am-Main, Germany, 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 heredity 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 United States (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 mutations, 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, chromosomes (rather than individual chromo- however, had accepted a model in which the somes alone) determine sex (Littlefield, 1980) location of specific genes 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)

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