Vernon M. Ingram THE WILLIAM ALLAN MEMORIAL AWARD Presented at the annual meeting of the American Society of Human Genetics Toronto December 1, 1967

CITATION The contributions for which we honor Vernon Ingram represent a landmark in the history of molecular . Using an ingenious technique, he showed in 1956 that the difference between normal and sickle was caused by a single substitution. This work, followed by his demonstration of a similar mechanism for and hemo- globin E, established clearly the nature of mutations in structural proteins and laid the groundwork for our concepts of gene action on the molecular level. 287 288 THE WILLIAM ALLAN MEMORIAL AWARD Vernon Ingram's investigations played a key role in the full integration of human genetics into the mainstream of modern genetic research. For the first time in the history of genetics, a phenomenon of fundamental significance for all forms of life was first demonstrated in man. Human genetics had come of age and became respectable to our more basically inclined colleagues. Vernon Ingram, as a scientist, represents a model for our students. Some investigators are analytical, while others are synthesizers and paint a broad sweep. Some are lone workers, others perform better in a team. Most biologists work in one field-within the framework of Ingram's interests, either as biochemists, as hematologists, as geneticists, or as evolutionists. Vernon Ingram has performed with versatility and distinction in all of these roles. As an analytical chemist, he developed the "fingerprinting" method of peptide separation, which he used to test theories of mutations. Against a broad background of genetic biology and his own findings in , he elaborated the now classic scheme of globin chain evolu- tion-a highly pertinent model for the evolution of other proteins. Always eager to see the broad implications of his work, he popularized the now generally accepted concept of the a and i thalassemias-diseases caused by genetically determined suppression of globin chain synthesis. Ever seeing fundamental problems, he has in recent years addressed himself to the experimental study of control mechanisms affecting hemoglobin synthesis. Few scientists have the enviable record of having stimulated large groups of investigators all over the world with both concepts and methods. Every scientist working in biochemical genetics, molecular evolution, hematology, and physical anthropology is in Vernon Ingram's intellectual debt. Vernon Ingram was educated as an organic chemist and obtained his Ph.D. at Birkbeck College in London. After fellowships in biochemistry at Rockefeller Institute and Yale Uni- versity in this country, he joined the Unit of Molecular Biology at the in Cambridge, England. It is here that the exciting work on hemoglobin was done. The in- tellectual atmosphere at the Cavendish Laboratory of Cambridge and particularly at its Molecular Biology Unit was particularly propitious for the discoveries he was to make. Here the double helix of DNA and its genetic significance had been worked out by Crick and Watson and the structure of myoglobin and hemoglobin was being elucidated by Kendrew and Perutz. Vernon Ingram was not content to test his ideas in discussions with Crick and Perutz in Cambridge. Our previous William Allan medalist, Jim Neel, also played a role as godfather in the development of Ingram's concepts, as did Herman Lehmann in London and later Corrado Baglioni, Park Gerald, Salvadore Luria, and Cyrus Levinthal in Boston. In 1958, Great Britain lost an outstanding scientist, and England's loss became our gain. A professor of biochemistry at Massachusetts Institute of Technology, Vernon has become a friend and adviser to many workers who are interested in hemoglobin and its genetics. His book on the genetics and evolution of hemoglobin has had a wide impact on biologists of all persuasions. A more recent book on The Biosynthesis of Macromolecules shows his great abili- ties as a teacher and points to his recent interests in the structure and function of transfer RNA. Always intent on developing the human implications of his work, Ingram devotes some of his time to consultation and lecturing in the Department of Medicine at , in New York. He has served on the Hematology Study Section of the National Institutes of Health. He has been a Harvey Society lecturer and a Jessup lecturer at Columbia University. Always eager to expand his horizons, he is spending this year on a sabbatical leave as a Guggenheim fellow at University College in London to learn tissue culture tech- niques. The American Society of Human Genetics is proud and delighted to present the William Allan Memorial Award to Vernon Ingram for work which has brought distinction not only to the recipient, but to human genetics as a discipline. (Arno G. Motulsky)