John D. Biggers (1923–2018)

John D. Biggers, one of the true pioneers in the field of reproductive biology, passed away at his home in Lexington, Massachusetts on April 7, 2018. His scientific career spanned seven decades, with more than 175 peer-reviewed papers and many reviews and book chapters published beginning in 1951 and continuing through 2016. John was born in Reading, England, in 1923. He graduated from the Royal Veterinary College and obtained his Ph.D. from the University of London. He then moved to the University of Sydney in Australia to take up a post in Veterinary Physiology where he investigated estrogen signaling and developed estrogen assays in collaboration with Peter Claringbold in the department headed by Clifford Emmens. This period helped establish John’s lifelong passion for statistical and biometric analysis. In 1955, John D. Biggers returned to England, taking a faculty position at the Royal Veterinary College at the University of London, where he embarked on studies of tissue culture, initially focusing on organ culture of bones. The development of media for long-term growth of embryonic bones in vitro (Heyner & Biggers, 1958) established one of the research themes that drove his entire career—the development and optimization of chemically defined culture media. Anne McLaren was also in London by this time, and their complementary interests led to their historic collaboration. Their 1958 Nature paper, “Successful Development and Birth of Mice Cultivated In Vitro as Early Embryos,” provided the first demonstration that embryos that had developed in culture could produce normal offspring (McLaren & Biggers, 1958). This was one of the most important advances for experimental embryology and the eventual development of assisted reproduction technologies. In 1959, John moved to the US, where he held faculty positions at the University of Pennsylvania and The Johns Hopkins University before being appointed as the Professor of Physiology at Harvard Medical School, in 1972. From 1960s through early 1980s, he published extensively on the metabolism of preimplantation embryos and regulation of oocyte maturation, with trainees and collaborators who included Ralph Brinster, Wes Whitten, David Whittingham, Dale Benos, and many other well-known reproductive biologists. During this time, he also established methodologies for analysis of the preimplantation embryo and egg microenvironment in a long-lived collaboration with the biophysicist Claude Lechene and with Henry Leese. John was a founding member and second president of the Society for the Study of Reproduction (SSR), as well as the Editor of their journal Biology of Reproduction. He received the SSR’s highest award, the Carl G. Hartman Award, in 1987. The mid-1980s saw the establishment of the National Cooperative Program on Non-Human In Vitro Fertilization and Preimplantation Development under the leadership of Richard J (Dick) Tasca at the US National Institutes of Health. It was within this program, informally called the “Culture Club,” that the first media capable of supporting complete preimplantation embryo development from fertilized eggs to blastocysts were developed. Biggers’ KSOM embryo culture medium (Lawitts & Biggers, 1993) that was developed in an extensive series of investigations with his postdoctoral fellow Joel Lawitts, which used sophisticated statistical optimization techniques to drive medium formulation, is widely used today and is the basis for successful human clinical IVF culture media. The importance of the Culture Club, particularly the contributions of the Biggers laboratory, to the success of Human Assisted Reproductive Technologies (ART) cannot be overstated. John D. Biggers was also very much interested in the ethics of ART and wrote and lectured extensively on this subject. He was called to testify before the US Senate about IVF and embryo transfer and appointed by President Carter as Chief Scientific Advisor on the Ethics Board of the US Department of Health, Education and Welfare. John remained active in research until the end of his life. Most recently, he had an active collaboration with Mehmet Toner to develop methods of long-term storage of mouse sperm and had just completed a book chapter with his collaborator Catherine Racowsky. Only 2 years ago, he and his coauthor Carol Kountz authored a book on Walter Heape, an underappreciated pioneer in the field of reproduction who performed the first successful embryo transfer at the end of the 19th century (Biggers & Kountz, 2016). John traveled extensively including all seven continents, often with his wife, late Betsey Williams, a former faculty member in the Anatomy Department at Harvard. He was an excellent cook and a passionate bird watcher and photographer. He will be remembered as a valued collaborator, a superb mentor who trained many top researchers, and a talented scientist whose work includes some of the most important advances made in the field of reproduction (Figure 1). John D. Biggers and Anne McLaren with one of the mice from their landmark 1958 paper (McLaren & Biggers, 1958), which first reported that embryos that had been cultured in vitro before being transferred to a recipient female could give rise to a live offspring. Photo from John D. Biggers (Biggers, 2001)

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