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Motivations, Practice, and Conflict in the History of Nuclear Transplantation, 1925-1970

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Motivations, Practice, and Conflict in the History of Nuclear Transplantation, 1925-1970 A 'Fantastical' Experiment: Motivations, Practice, and Conflict in the History of Nuclear Transplantation, 1925-1970 A DISSERTATION SUBMITTED TO THE FACULTY OF THE GRADUATE SCHOOL OF THE UNIVERSITY OF MINNESOTA BY Nathan Paul Crowe IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY Mark Borrello, Adviser December, 2011 © Nathan Paul Crowe 2011 All Rights Reserved Acknowledgements Though completing a dissertation is often seen as a solitary venture, the truth is rather different. Without the help of family, friends, colleagues, librarians, and archivists, I would never have been able to finish my first year in graduate school, let alone my thesis. During the course of my research I came in contact with many people who were invaluable to my eventual success. First and foremost are those librarians, archivists, and collections specialists who helped guide me through a sea of records. I am indebted to all of those who helped me in the process: in particular, I would like to thank Janice Goldblum and Daniel Barbiero at the National Academy of Sciences, Lee Hiltzik at the Rockefeller Archives Center, and National Cancer Institute Librarian Judy Grosberg. At the National Institutes of Health, librarian Barbara Harkins, information specialist Richard Mandel, and historian David Cantor were all extremely useful in helping me find what I needed. I also can't imagine anyone being more accommodating than librarian Beth Lewis and her staff at the Fox Chase Cancer Center Library. I was able to complete my research with the financial support of several institutions. My thanks go out to the Rockefeller Archive Center for their generous help. The University of Minnesota Consortium on Law and Values in Health, Environment, and the Life Sciences provided the bulk of my funding for research trips and materials. I'm indebted to their support, and I want to particularly thank Audrey Boyle, who helped me manage the piles of receipts and last-minute changes in plans that so often come up during the length of a grant. Out of all my funding, perhaps the most important was the Dissertation and Development Fellowship provided by the University of Minnesota i Graduate School. I cannot imagine how I could have researched and written my dissertation without the help of this year-long fellowship. I hope that the administration continues to support graduate students (particularly in the humanities) during their last years; it can mean a world of difference. Also, institutional support can come in forms other than money. Thank you to Barbara Eastwold, who helped me successfully navigate through the bureaucracy of University of Minnesota. You deserve to be in the acknowledgements of every student who comes out of our program. Intellectually, I am indebted to a number of people. Primarily, my adviser, Mark Borrello, supplied me with insightful comments and critiques throughout the process, encouraging me when necessary and always reining me in when I got intellectually wild. I'm thankful for his guidance not just in this dissertation, but also throughout graduate school in general. Of course, Sally Gregory Kohlstedt, as my first director of graduate studies and later my department head, was influential in my becoming the professional historian I am today. If only more academics were as hard-working and as caring, higher education would enjoy an impeccable reputation. I would also like to thank the Minnesota Center for the Philosophy of Science and members of the Biological Interest Group, particularly professors Alan Love and Ken Waters (both of whom also served as dissertation committee members) and Mike Travisano. Being a part of their weekly intellectual grappling opened my eyes to new ways of understanding the world and taught me to think (and write) more clearly and at a higher level. Also, this dissertation would have been much harder to write without the insights of those who experienced it. Particularly, I would like to thank Robert McKinnell, whose ii vivid tales, both during our formal interview but also during many lunches, helped make the past become so visible. Likewise, I would like to thank Marie Di Berardino for her cooperation. I could not do her historical contributions justice in this dissertation, but promise to make her story more prominent in the future. One of the best parts of graduate school is the comrades-in-arms that you gain along the way. Special thanks to my original classmate, Maggie Hofius. She was always around to celebrate the good times and to help get through the tough ones. My fellow students from DAWGS (our Dissertation And Writing Group), Frazier Benya and Christine Manganaro, were both instrumental in helping me not only finish my dissertation, but also in keeping my sanity. I should also thank Rachel Mason Dentinger and Gina Rumore for all the great mentorship over the years. Of course, my officemate, Jacob Steere-Williams deserves my infinite gratitude. Not only did he bear much of the intellectual drama as the wall off of which I bounced my ideas, but he also helped make life enjoyable along the way. Thanks for all the beer, bourbon, baseball, basketball, board games, and general brotherhood. And to all the other graduate students in the program, past and present, thanks for making the University of Minnesota the best place to spend five years. Finally, I never would have made it half as far without my family. My parents, Paul and Lisa, and my brother Colin gave me more support and encouragement than anyone could ask for. They never seemed to doubt that I would finish. That goes double for my Aunt Maria, whose emphatic assumption that I would succeed was infectious. I would also like to thank my parents-in-law, Art and Sue Horowitz, for their constant enthusiasm and thought-provoking questions. iii Last but not least, my wife Stephanie. A "thank you" can never be enough for everything you did for me during graduate school, all the unpleasantness that you had to deal with, all the papers that you edited, and all the dishes that often went unwashed because of my work. I look forward to having the rest of my life to show you how much I appreciate everything you did for me and everything about you. iv For my grandmother, Thea. v Abstract In 1952, Robert Briggs and Thomas King published a paper announcing the development of a new technique, nuclear transplantation, which could have profound consequences in the study of developmental biology. Forty-four years later, in 1996, researchers in Scotland used a variation of nuclear transplantation to produce a cloned sheep. The sheep was named Dolly and became a cultural, scientific, and controversial symbol for biology's successes and promises. Since then, the historical relevance of nuclear transplantation has always been its connection to the successful cloning of Dolly. I argue in this dissertation, however, that the history of nuclear transplantation before Dolly offers valuable insights into the history of developmental biology, genetics, cancer research, and bioethics. As essentially a biography of the technique, my narrative weaves together these often distinct historiographical traditions, showing the intricate institutional and intellectual connections between them. Though the first successful nuclear transplantation in vertebrates occurred in the early 1950s, this dissertation traces back the relevant historical origins to the early 1920s with the development of the cancer research center in which Briggs and his colleagues eventually worked out nuclear transplantation. In subsequent chapters this dissertation follows the development of the technique and the successes and controversies that it encountered in the 1950s related to the work of John Gurdon. From there, I show how nuclear transplantation moved from strictly a laboratory discussion to a cultural phenomenon related to human cloning in the 1960s when Nobel Laureate Joshua Lederberg helped co-opt nuclear transplantation to fuel democratic discussion over the direction of biological research. vi Table of Contents Introduction 1 Chapter One: A Research Program Takes Shape: The Founding and 15 Development of Lankenau Hospital Research Institute, 1925-1942 Chapter Two: Embodying Conflict: Robert Briggs and the Potential of 62 Nuclear Transplantation Chapter Three: The Proliferation and Interpretation of Nuclear Transplantation 118 Chapter Four: Going Public: Nuclear Transplantation's Transition to a 178 Bioethical Challenge Conclusion 237 Bibliography 243 vii Introduction On July 5, 1996, a sheep named Dolly was born. Unlike other lambs born that day, Dolly was special. She was a clone–a genetic duplicate of a six-year-old sheep already in existence. Dolly was not the first cloned animal, nor was she even the first cloned mammal. Rather, Dolly's subsequent fame derived from the type of cell used to create her. Instead of an embryo or an early developing cell – one that still held the potential to become any part of the body – Dolly came from the nucleus of a fully specialized cell using the technique of nuclear transplantation.1 Her creators used the nucleus of a mammary cell from one sheep to direct development of an unfertilized egg cell from another sheep that had had its nucleus removed, thus making a duplicate organism from the donor cell.2 The circumstances of her birth gained her both scientific and cultural fame. The birth of Dolly was culturally important because it supposedly removed all scientific barriers to human cloning. New York Times science writer Gina Kolata's 1998 book, Clone: The Road to Dolly and the Path Ahead, was indicative of the type of coverage that the event received in the press, beginning with the first news of the birth that Kolata herself broke for the New York Times.3 "When the time comes to write the history of our age, this quiet birth, the creation of a little lamb, will stand out," Kolata 1 The phrases "nuclear transplantation" and "nuclear transfer" are interchangeable.
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