Alfred D

Alfred D. Hershey

Born: 4 December 1908, Owosso, MI, USA

Died: 22 May 1997, Syosset, NY, USA

Affiliation at the time of the award:Carnegie Institution of Washington, Long Island, New York, NY, USA

Prize motivation: "for their discoveries concerning the replication mechanism and the genetic structure of

viruses"

Alfred Day Hershey was born on December 4th, 1908, in Owosso, Michigan. He studied at the Michigan State College, where he obtained B.S. in 1930, and Ph.D. in 1934. In 1967 he got an honorary D.Sc. at the University of Chicago.

From 1934 till 1950 he was engaged in teaching and research, at the Department of Bacteriology, Washington University School of Medicine. In 1950 he became a Staff Member, at the Department of Genetics, Carnegie Institution of Washington, Cold Spring Harbor, New York; in 1962 he was appointed Director of the Genetics Research Unit of the same institution.

Alfred Hershey married Harriet Davidson in 1945, they have one son, Peter.

Alfred Hershey is a Member of the American Society for Microbiology, the National Academy of Sciences, and the American Academy of Arts and Sciences. Hershey is Recipient of the Kimber Genetics Award of the National Academy of Sciences, 1965. Michigan State University honored him with an M.D.h.c. in 1970.

Robert Koch

Born: 11 December 1843, Clausthal (now Clausthal-Zellerfeld), Germany

Died: 27 May 1910, Baden-Baden, Germany

Affiliation at the time of the award:Institute for Infectious Diseases, Berlin, Germany

Prize motivation: "for his investigations and discoveries in relation to tuberculosis"

Robert Koch was born on December 11, 1843, at Clausthal in the Upper Harz Mountains. The son of a mining

engineer, he astounded his parents at the age of five by telling them that he had, with the aid of the newspapers,

taught himself to read, a feat which foreshadowed the intelligence and methodical persistence which were to be

so characteristic of him in later life. He attended the local high school («Gymnasium») and there showed an

interest in biology and, like his father, a strong urge to travel.

In 1862 Koch went to the University of Göttingen to study medicine. Here the Professor of Anatomy was Jacob

Henle and Koch was, no doubt, influenced by Henle's view, published in 1840, that infectious diseases were

caused by living, parasitic organisms. After taking his M.D. degree in 1866, Koch went to Berlin for six months of

chemical study and there came under the influence of Virchow. In 1867 he settled, after a period as Assistant in

the General Hospital at Hamburg, in general practice, first at Langenhagen and soon after, in 1869, at Rackwitz,

in the Province of Posen. Here he passed his District Medical Officer's Examination. In 1870 he volunteered for

service in the Franco-Prussian war and from 1872 to 1880 he was District Medical Officer for Wollstein. It was

here that he carried out the epoch-making researches which placed him at one step in the front rank of scientific

workers.

Anthrax was, at that time, prevalent among the farm animals in the Wollstein district and Koch, although he had

no scientific equipment and was cut off entirely from libraries and contact with other scientific workers, embarked,

in spite of the demands made on him by his busy practice, on a study of this disease. His laboratory was the 4-

roomed flat that was his home, and his equipment, apart from the microscope given to him by his wife, he

provided for himself. Earlier the anthrax bacillus had been discovered by Pollender, Rayer and Davaine, and

Koch set himself to prove scientifically that this bacillus is, in fact, the cause of the disease. He inoculated mice,

by means of home-made slivers of wood, with anthrax bacilli taken from the spleens of farm animals that had

died of anthrax, and found that these mice were all killed by the bacilli, whereas mice inoculated at the same time

with blood from the spleens of healthy animals did not suffer from the disease. This confirmed the work of others

who had shown that the disease can be transmitted by means of the blood of animals suffering from anthrax.

But this did not satisfy Koch. He also wanted to know whether anthrax bacilli that had never been in contact with

any kind of animal could cause the disease. To solve this problem he obtained pure cultures of the bacilli by

growing them on the aqueous humour of the ox's eye. By studying, drawing and photographing these cultures,

Koch recorded the multiplication of the bacilli and noted that, when conditions are unfavourable to them, they

produce inside themselves rounded spores which can resist adverse conditions, especially lack of oxygen and

that, when suitable conditions of life are restored, the spores give rise to bacilli again. Koch grew the bacilli for

several generations in these pure cultures and showed that, although they had had no contact with any kind of

animal, they could still cause anthrax.

The results of this painstaking work were demonstrated by Koch to Ferdinand Cohn, Professor of Botany at the

University of Breslau, who called a meeting of his colleagues to witness this demonstration, among whom was

Professor Cohnheim, Professor of Pathological Anatomy. Both Cohn and Cohnheim were deeply impressed by

Koch's work and when Cohn, in 1876, published Koch's work in the botanical journal of which he was the editor,

Koch immediately became famous. He continued, nevertheless, to work at Wollstein for a further four years and

during this period he improved his methods of fixing, staining and photographing bacteria and did further

important work on the study of diseases caused by bacterial infections of wounds, publishing his results in 1878.

In this work he provided, as he had done with anthrax, a practical and scientific basis for the control of these

infections.

Koch was still, however, without adequate quarters or conditions for his work and it was not until 1880, when he

was appointed a member of the «Reichs-Gesundheitsamt» (Imperial Health Bureau) in Berlin, that he was

provided, first with a narrow, inadequate room, and later with a better laboratory, in which he could work with

Loeffler, Gaffky and others, as his assistants. Here Koch continued to refine the bacteriological methods he had

used in Wollstein. He invented new methods - «Reinkulturen» - of cultivating pure cultures of bacteria on solid

media such as potato, and on agar kept in the special kind of flat dish invented by his colleague Petri, which is

still in common use. He also developed new methods of staining bacteria which made them more easily visible

and helped to identify them. The result of all this work was the introduction of methods by which pathogenic

bacteria could be simply and easily obtained in pure culture, free from other organisms and by which they could

be detected and identified. Koch also laid down the conditions, known as Koch's postulates, which must be

satisfied before it can be accepted that particular bacteria cause particular diseases.

Some two years after his arrival in Berlin Koch discovered the tubercle bacillus and also a method of growing it in

pure culture. In 1882 he published his classical work on this bacillus. He was still busy with work on tuberculosis

when he was sent, in 1883, to Egypt as Leader of the German Cholera Commission, to investigate an outbreak

of cholera in that country. Here he discovered the vibrio that causes cholera and brought back pure cultures of it

to Germany. He also studied cholera in India.

On the basis of his knowledge of the biology and mode of distribution of the cholera vibrio, Koch formulated rules

for the control of epidemics of cholera which were approved by the Great Powers in Dresden in 1893 and formed

the basis of the methods of control which are still used today. His work on cholera, for which a Prize of 100,000

German Marks was awarded to him, also had an important influence on plans for the conservation of water

supplies.

In 1885 Koch was appointed Professor of Hygiene in the University of Berlin and Director of the newly

established Institute of Hygiene in the University there. In 1890 he was appointed Surgeon General (Generalarzt)

Class I and Freeman of the City of Berlin. In 1891 he became an Honorary Professor of the Medical Faculty of

Berlin and Director of the new Institute for Infectious Diseases, where he was fortunate to have among his

colleagues, such men asEhrlich, von Behring and Kitasato, who themselves made great discoveries.

During this period Koch returned to his work on tuberculosis. He sought to arrest the disease by means of a

preparation, which he called tuberculin, made from cultures of tubercle bacilli. He made two preparations of this

kind called the old and the new tuberculin respectively, and his first communication on the old tuberculin aroused

considerable controversy. Unfortunately, the healing power that Koch claimed for this preparation was greatly

exaggerated and, because hopes raised by it were not fulfilled, opinion went against it and against Koch. The

new tuberculin was announced by Koch in 1896 and the curative value of this also was disappointing; but it led,

nevertheless, to the discovery of substances of diagnostic value. While this work on tuberculin was going on, his

colleagues at the Institute for Infectious Diseases, von Behring, Ehrlich and Kitasato, carried out and published

their epoch-making work on the immunology of diphtheria (see the biographies of Ehrlich and von Behring).

In 1896 Koch went to South Africa to study the origin of rinderpest and although he did not identify the cause of

this disease, he succeeded in limiting the outbreak of it by injection into healthy farm-stock of bile taken from the

gall bladders of infected animals. Then followed work in India and Africa on malaria, blackwater fever, surra of

cattle and horses and plague, and the publication of his observations on these diseases in 1898. Soon after his

return to Germany he was sent to Italy and the tropics where he confirmed the work of Sir Ronald Ross in

malaria and did useful work on the aetiology of the different forms of malaria and their control with quinine.

It was during these later years of his life that Koch came to the conclusion that the bacilli that caused human and

bovine tuberculosis are not identical and his statement of this view at the International Medical Congress on

Tuberculosis in London in 1901 caused much controversy and opposition; but it is now known that Koch's view

was the right one. His work on typhus led to the idea, then a new one, that this disease is transmitted much more

often from man to man than from drinking water and this led to new control measures.

In December, 1904, Koch was sent to German East Africa to study East Coast fever of cattle and he made

important observations, not only on this disease, but also on pathogenic species

of Babesia and Trypanosoma and on tickborne spirochaetosis, continuing his work on these organisms when he

returned home.

Koch was the recipient of many prizes and medals, honorary doctorates of the Universities of Heidelberg and

Bologna, honorary citizenships of Berlin, Wollstein and his native Clausthal, and honorary memberships of

learned societies and academies in Berlin, Vienna, Posen, Perugia, Naples and New York. He was awarded the

German Order of the Crown, the Grand Cross of the German Order of the Red Eagle (the first time this high

distinction was awarded a medical man), and Orders from Russia and Turkey. Long after his death, he was

posthumously honoured by memorials and in other ways in several countries.

In 1905 he was awarded the Nobel Prize for Physiology or Medicine. In 1906, he returned to Central Africa to

work on the control of human trypanosomiasis, and there he reported that atoxyl is as effective against this

disease as quinine is against malaria. Thereafter Koch continued his experimental work on bacteriology and

serology.

In 1866 Koch married Emmy Fraats. She bore him his only child, Gertrud (b. 1865), who became the wife of Dr.

E. Pfuhl. In 1893 Koch married Hedwig Freiberg.

Dr. Koch died on May 27, 1910, in Baden-Baden.

Robert W. Holley

Born: 28 January 1922, Urbana, IL, USA

Died: 11 February 1993, Los Gatos, CA, USA

Affiliation at the time of the award:Cornell University, Ithaca, NY, USA

Prize motivation: "for their interpretation of the genetic code and its function in protein synthesis"

Robert W. Holley was born in Urbana, Illinois, on January 28th, 1922, one of four sons of Charles and Viola Holley. His parents were both educators. He attended public schools in Illinois, California and Idaho, and graduated from Urbana High School in 1938. He studied chemistry at the University of Illinois and received his B. A. degree in 1942. Graduate work was at Cornell University, where the Ph.D. degree in organic chemistry, with Professor Alfred T. Blomquist, was awarded in 1947. Graduate work was interrupted during the war. He spent two years, 1944-1946, with Professor Vincent du Vigneaud at Cornell University Medical College, where he participated in the first chemical synthesis of penicillin.

After completing the Ph. D. degree, Holley spent 1947-1948 as an American Chemical Society Postdoctoral Fellow with Professor Carl M. Stevens at Washington State University. He then returned to Cornell University as Assistant Professor of Organic Chemistry at the Geneva Experiment Station in 1948. He was Associate Professor there from 1950-1957. During a sabbatical year, 1955-1956, he was a Guggenheim Memorial Fellow in the Division of Biology at the California Institute of Technology. In 1958, he returned to Ithaca, New York, as a Research Chemist at the U. S. Plant, Soil and Nutrition Laboratory, a U. S. Department of Agriculture Laboratory on the Cornell University campus. He had an appointment in the University throughout this period and became Professor of Biochemistry in 1962. He rejoined the faculty of Cornell University full time in 1964 as Professor of Biochemistry and Molecular Biology, and was Chairman of the Department from 1965 to 1966. The following year, 1966-1967, was spent at the Salk Institute for Biological Studies and the Scripps Clinic and Research Foundation in La Jolla, California, as a National Science Foundation Postdoctoral Fellow. In 1968, though maintaining an affiliation with Cornell University, he joined the permanent staff of the Salk Institute, where he is a Resident Fellow and an American Cancer Society Professor of Molecular Biology. He is also an Adjunct Professor at the University of California at San Diego.

Holley's training as a chemist did not alter his basic interest in living things. This interest has influenced his choice of research, which began with the organic chemistry of natural products. There followed a gradual drift toward more biological subjects, with work on amino acids and peptides, and eventually work on the biosynthesis of proteins. During the latter, the alanine transfer RNA was discovered. The following 10 years were spent working with this RNA, first concentrating on the isolation of the RNA, and then working on the determination of the structure of the RNA. The nucleotide sequence was completed at the end of 1964. It was for this work that the Nobel Prize was awarded. More recently, his work has been concerned with factors that control cell division in mammalian cells.

Holley is a member of the National Academy of Sciences, the American Academy of Arts and Sciences, the American Association for the Advancement of Science, The American Society of Biological Chemists and the American Chemical Society. He received the Albert Lasker Award in Basic Medical Research in 1965, the Distinguished Service Award of the U. S. Department of Agriculture in 1965, and the U. S. Steel Foundation Award in Molecular Biology of the National Academy of Sciences in 1967.

Holley was married to Ann Dworkin in 1945. They have one son, Frederick. Mrs. Holley's professional interests are concerned with the teaching of mathematics. The three of them especially enjoy the ocean and the mountains.

Born: 9 January 1922, Raipur, India

Affiliation at the time of the award:University of Wisconsin, Madison, WI, USA

Prize motivation: "for their interpretation of the genetic code and its function in protein synthesis"

Har Gobind Khorana was born of Hindu parents in Raipur, a little village in Punjab, which is now part of eastern Pakistan. The correct date of his birth is not known; that shown in documents is January 9th, 1922. He is the youngest of a family of one daughter and four sons. His father was a «patwari», a village agricultural taxation clerk in the British Indian system of government. Although poor, his father was dedicated to educating his children and they were practically the only literate family in the village inhabited by about 100 people.

Har Gobind Khorana attended D.A.V. High School in Multan (now West Punjab); Ratan Lal, one of his teachers, influenced him greatly during that period. Later, he studied at the Punjab University in Lahore where he obtained an M. Sc. degree. Mahan Singh, a great teacher and accurate experimentalist, was his supervisor.

Khorana lived in India until 1945, when the award of a Government of India Fellowship made it possible for him to go to England and he studied for a Ph. D. degree at the University of Liverpool. Roger J. S. Beer supervised his research, and, in addition, looked after him diligently. It was the introduction of Khorana to Western civilization and culture.

Khorana spent a postdoctoral year (1948-1949) at the Eidgenössische Technische Hochschule in Zurich with Professor Vladimir Prelog. The association with Professor Prelog molded immeasurably his thought and philosophy towards science, work, and effort.

After a brief period in India in the fall of 1949, Khorana returned to England where he obtained a fellowship to work with Dr. (now Professor) G. W. Kenner and Professor (now Lord) A. R. Todd. He stayed in Cambridge from

1950 till 1952. Again, this stay proved to be of decisive value to Khorana. Interest in both proteins and nucleic acids took root at that time.

A job offer in 1952 from Dr. Gordon M. Shrum of British Columbia (now Chancellor of Simon Fraser University, British Columbia) took him to Vancouver. The British Columbia Research Council offered at that time very little by way of facilities, but there was «all the freedom in the world», to use Dr. Shrum's words, to do what the researcher liked to do. During the following years, with Dr. Shrum's inspiration and encouragement and frequent help and scientific counsel from Dr. Jack Campbell (now Head of the Department of Microbiology at the University of British Columbia), a group began to work in the field of biologically interesting phosphate esters and nucleic acids. Among the many devoted and loyal colleagues of this period, there should, in particular, be mention of Dr. Gordon M. Tener (now a Professor in the Biochemistry Department of the University of British Columbia), who contributed much to the spiritual and intellectual well-being of the group.

In 1960 Khorana moved to the Institute for Enzyme Research at the University of Wisconsin. He became a naturalized citizen of the United States. As of the fall of 1970 Khorana has been Alfred P. Sloan Professor of Biology and Chemistry at the Massachusetts Institute of Technology.

Har Gobind Khorana was married in 1952 to Esther Elizabeth Sibler, who is of Swiss origin. Esther brought a consistent sense of purpose into his life at a time when, after six years' absence from the country of his birth, Khorana felt out of place everywhere and at home nowhere. They have three children: Julia Elizabeth (born May 4th, 1953), Emily Anne (born October 18th, 1954), and Dave Roy (born July 26th, 1958).

Marshall W. Nirenberg

Born: 10 April 1927, New York, NY, USA

Died: 15 January 2010, New York, NY, USA

Affiliation at the time of the award:National Institutes of Health, Bethesda, MD, USA

Prize motivation: "for their interpretation of the genetic code and its function in protein synthesis"

Marshall Warren Nirenberg was born in New York City on April 10th, 1927, the son of Harry and Minerva Nirenberg. The family moved to Orlando, Florida in 1939. He early developed an interest in biology. In 1948 he received a B. Sc. degree, and in 1952, a M. Sc. degree in Zoology from the University of Florida at Gainesville. His dissertation for the Master's thesis was an ecological and taxonomic study of caddis flies (Trichoptera).

During this period he became interested in biochemistry. He continued studies in this field at the University of Michigan, Ann Arbor, and in 1957 received the Ph. D. degree from the Department of Biological Chemistry. Nirenberg's thesis, performed under the guidance of Dr. James Hogg, was a study of a permease for hexose transport in ascites tumor cells.

From 1957 to 1959 he obtained postdoctoral training with DeWitt Stetten Jr., and with William Jakoby at the National Institutes of Health as a fellow of the American Cancer society. During the next year he held a Public Health Service Fellowship and in 1960 became a research biochemist in the Section of Metabolic Enzymes, headed by Dr. Gordon Tompkins, at the National Institutes of Health.

In 1959 he began to study the steps that relate DNA, RNA and protein. These investigations led to the demonstration with H. Matthaei that messenger RNA is required for protein synthesis and that synthetic messenger RNA preparations can be used to decipher various aspects of the genetic code.

In 1962 he became head of the Section of Biochemical Genetics at the National Institutes of Health.

Nirenberg holds honorary degrees from the University of Michigan, Yale University, University of Chicago, University of Windsor (Ontario) and Harvard University. Other honours include: The Molecular Biology Award, National Academy of Sciences, 1962; Paul Lewis Award in Enzyme Chemistry, American Chemical Society, 1964; The National Medal of Science, 1965; The Research Corporation Award, 1966; the Hildebrand Award, 1966; the Gairdner Foundation Award of Merit, 1967; The Prix Charles Leopold Meyer, French Academy of Sciences, 1967; the Joseph Priestly Award, 1968; and the Franklin Medal, 1968. The Louisa Gross Horwitz Prize, Columbia University, and the Lasker Award were shared with H. G. Khorana in 1968. He is a member of the American Academy of Arts and Sciences and the National Academy of Sciences.

He was married in 1961 to Perola Zaltzman, a chemist from the University of Brazil, Rio de Janeiro. She is now a biochemist at the National Institutes of Health.

Susumu Tonegawa

Born: 6 September 1939, Nagoya, Japan

Affiliation at the time of the award:Massachusetts Institute of Technology (MIT), Cambridge, MA, USA

Prize motivation: "for his discovery of the genetic principle for generation of antibody diversity"

I was born in Nagoya, Japan on September 6th, 1939, the second of three sons. I have also a younger sister. My father was an engineer working for a textile company that had several factories scattered in rural towns in the southern part of Japan. The company policy made it necessary for my father to move from one factory to another every few years. I and my brothers and sister spent most of our childhood in these small provincial towns, enjoying the space and freedom of the countryside. As my elder brother and I reached adolescence, however, my parents decided to send us to Tokyo so that we could receive a better education.

I commuted to the prestigious Hibiya high school from my Uncle's home in Tokyo. During the high school years I developed an interest in chemistry, so upon graduation, I chose to take an entrance examination for the Department of Chemistry of the University of Kyoto, the old capital of Japan. After having failed once, I was admitted to this University in 1959. This happened to be one year before the first ten-year term of the defence treaty between Japan and the United States expired and the governments of both countries were preparing for a second ten-year term.

The nation was deeply divided between the pragmatic pro-American conservatives and the idealistic anti-military leftists. Being the home of the most radical leftist student groups, classes at Kyoto University were often cancelled due to frequent political discussions and demonstrations on the streets. I was only a passive participant, withdrawn from the turmoil, but could not help having a feeling of defeat shared with many of my classmates when the treaty was finally extended for the next ten-year term. I believe that this experience might have been a major factor in making me give up the original goal of becoming a chemical engineer to pursue the

academic life.

I became fascinated by the then blossoming science of molecular biology when in my senior year I happened to read the papers by François Jacob and Jacques Monod on the operon theory. I decided to pursue graduate study in molecular biology and was accepted by Professor Itaru Watanabe's laboratory at the Institute for Virus Research at the University of Kyoto, one of a few laboratories in Japan where U.S.-trained molecular biologists were actively engaged in research. However, only two months after I started my work in his laboratory, Professor Watanabe called me into his office and suggested that I carry out my graduate study in the United States. He explained how inadequate the graduate training programs in molecular biology laboratories were in Japan, including his own, and offered to help in my application to some major universities in the United States, if I would seriously consider studying abroad.

At that time, it was a common career development for a Japanese molecular biologist to go to the United States for a few years of postdoctoral study after obtaining the Ph.D. in Japan. I already had a vague wish to follow that pattern. Professor Watanabe's advice to enroll in an American graduate school therefore came to me as a bit of a surprise, but I was excited by the idea and accepted his help immediately. I cannot thank Professor Watanabe enough for this critical suggestion in the early phase of my scientific career.

With the additional help of Dr. Takashi Yura, then an assistant professor in Watanabe's laboratory, I was admitted to the graduate school of the Department of Biology of the University of California at San Diego that had recently been established by Professor David Bonner in La Jolla, the beautiful southern Californian town near the Mexican border.

At UCSD I studied in the laboratory of Professor Masaki Hayashi, carrying out a thesis project on the transcriptional control of phage lambda and received my Ph.D. in molecular biology in 1968. I remained in Professor Hayashi's laboratory as a postdoctoral fellow working on the morphogenesis of a phage, ØX174, until early 1969. Then I moved, also as a postdoctoral fellow, across the street to the laboratory of Dr. Renato Dulbecco at the Salk Institute.

Like many others, I believed that the golden age of prokaryotic molecular biology was coming to an end and that the great excitement would be in higher organisms. However, the complexity of high organisms was baffling and the necessary tools seemed hopelessly insufficient. Small tumor viruses like polyoma and simian virus 40, the biological material primarily dealt with in Dulbecco's laboratory, seemed to offer a bridge for the gap between prokaryotes and eukaryotes. Indeed Dulbecco's laboratory was filled with first-class postdoctoral fellows from around the world, who were trained in prokaryotic molecular biology and who came there intending to expand their research into eukaryotic molecular biology.

My project was to define the transcripts of SV40 during lytic infection and in transformed cells. Since this was the pre-restriction enzyme and pre-recombinant DNA age, the information I could obtain was very limited. However, being a member of the best laboratory in the field I glimpsed the excitement of the cutting edge of scientific research. Furthermore, I very much enjoyed the free and stimulating atmosphere of the laboratory. Unfortunately, as an awardee of a Fulbright travel grant, my U.S. visa was to expire by the end of 1970 and I had to leave the country for at least two years before I was eligible for another U.S. visa.

I had two or three job possibilities outside of the United States, but none were particularly interesting. In the autumn of 1970, only a few months before my visa was to expire I received a letter from Renato Dulbecco who was travelling in Europe. Renato mentioned the newly established Basel Institute for Immunology in Basel, Switzerland, and suggested that the time might be ripe for a molecular biologist to tackle immunological problems. I had very little knowledge of immunology, but decided to take Dr. Dulbecco's advice and sent an application letter to the Director of the Institute, Professor Niels Kaj Jerne, who offered me a two-year contract.

In the winter of 1971, I thus found myself surrounded by immunologists in this small town located in the middle of Europe. I must admit that the first year in the Institute was not easy for me. I had a continuing interest in work on SV40, but I was also keenly aware that I would not be able to take much advantage of the circumstances if I isolated myself by pursuing that subject. I therefore decided to study immunology in the hope of finding an interesting project.

An immunologist, Dr. Ita Askonas, and a geneticist, Charles Steinberg, were very helpful to me on my entering

the new field. By the end of 1971, I was introduced to the great debate on the genetic origins of antibody diversity. I felt from the beginning that I could contribute to resolving this question by applying the recently invented techniques of molecular biology, namely, restriction enzymes and recombinant DNA. Initially I worked only with my skillful technicians, Monica Shöld and Rita Schuller, but was soon joined by Drs. Nobumichi Hozumi, Minoru Hirama, and Christine Brack. Later, as my research group expanded, I had the good fortune to work with many capable postdoctoral fellows and devoted technical assistants. In addition, Charles Steinberg was a very important collaborator and consultant, particularly in the initial phase of the research.

Looking back, the research progressed with amazing speed from 1974 to 1981, the year I left Basel. We all worked hard and had had a great deal of fun. Our work resolved the long held debate on the genetic origin of antibody diversity. It turned out that this diversity is generated by somatic recombination of the inherited gene segments and by somatic mutation. To our very good fortune, Director Niels Jerne was quick to understand the importance of our approach and became a staunch supporter of the research in its early phase.

In the beginning of the 1980's I began to feel that the great mystery of antibody diversity had been solved, at least in its outlines. I thought that it might be good to change my environment to launch into a new project. I also recalled that I had initially come to Switzwerland with the intention of staying for two years and then returning to the United States. Fortunately, I received a few offers from the United States and decided in 1981 to take a professorship at the Center for Cancer Research at M.I.T. Professor Salvador E. Luria, Director of the Cancer Center, was extremely helpful, not only in bringing me to M.I.T., but also in providing me with a beautiful laboratory.

The research projects on which I had decided concerned two major problems. One was to investigate the role of somatic rearrangement in the activation of the rearranged antibody gene, and the second was to extend the research in Basel to "the other half" of the immune system, namely, to the antigen receptor of T cells. Fortunately, we could contribute to the understanding of both problems by discovering a tissue-specific transcriptional enhancer in the immunoglobulin heavy chain gene and by identifying, cloning, and sequencing genes coding for the polypeptide subunits of the T cell receptor. A particularly intriguing development made during the latter study was the identification of a gene that led to the discovery of a new T cell receptor, gamma delta. While the function of the T cells bearing this receptor is currently unknown, data accumulated during the past year in our laboratory as well as many other laboratories suggest that these T cells may be involved in an entirely new aspect of immunity.

When I look back on my scientific career to-date, I am amazed at my good fortune. At every major turn, I met scientists who were not only at the very top in their own fields, but who also gave me insightful advice and generous help. I am most grateful to Professors Itaru Watanabe, Renato Dulbecco, Niels Kaj Jerne, Charles Steinberg, and Salvadore Luria. I also wish to extend my unending gratitude to many colleagues and technical assistants.

My parents were firm believers that education is the best asset that parents can give to their children. I am deeply grateful to them for their outstanding support of my study and professional career. I am extremely grateful to my wife, Mayumi, whom I married in September 1985 for her devotion, interest, encouragement and criticism. I also wish to express my sincere thanks to my first wife, Kyoko, for her limitless devotion during my days in La Jolla and Basel.

I have been fortunate enough to receive many professional honors which include: The Cloetta Prize of Foundation Professor Dr. Max Cloetta, Switzerland (1978), Warren Triennial Prize of the Massachusetts General Hospital, U.S.A. (1980), Genetics Grand Prize of Genetics Promotion Foundation, Japan (1981), Avery Landsteiner Prize of the Gesselshat für Immunologie, West Germany (1981), Asahi Prize of Asahi-Shimbun (Asahi Press), Tokyo, Japan (1982), Louisa Gross Horwitz Prize of Columbia University, New York, U.S.A. (1982), The V.D. Mattia Award of the Roch Institute of Molecular Biology, Nutley, U.S.A. (1983), Gairdner Foundation International Awards of the Gairdner Foundation, Toronto, Canada (1983), Person of Cultural Merit "Bunkakorosha" of the Japanese Government (1983), Order of Culture "Bunkakunsho" from the Emperor of Japan (1984), Bristol-Myers Award for Distinguished Achievement in Cancer Research (1986), Robert Koch Prize of the Robert Koch Foundation, West Germany (1986). Albert and Mary Lasker Award, New York City (1987) and NOBEL PRIZE in Physiology or Medicine, Stockholm, Sweden (1987).

Gerald M. Edelman

Born: 1 July 1929, New York, NY, USA

Affiliation at the time of the award:Rockefeller University, New York, NY, USA

Prize motivation: "for their discoveries concerning the chemical structure of antibodies"

Dr. Gerald M. Edelman was born on July 1, 1929 in New York City to Edward Edelman and Anna Freedman Edelman. His father is a practicing physician in New York.

After his education in New York public schools, Edelman attended Ursinus College in Pennsylvania and received the B.S. degree, magna cum laude, in 1950. He then attended the Medical School of the University of Pennsylvania where he received the M.D. degree in 1954. In the succeeding year, he was a Medical House Officer at the Massachusetts General Hospital. He became a Captain in the U.S. Army Medical Corps in 1955 and practiced general medicine at a Station Hospital connected with the American Hospital in Paris, France. In 1957, he joined the Rockefeller Institute as a graduate fellow in the laboratory of Dr. Henry G. Kunkel.

After receiving the Ph.D. degree in 1960, he remained at the Rockefeller Institute as Assistant Dean of Graduate Studies and started work in his own laboratory. In 1963, he became Associate Dean of Graduate Studies, a position from which he retired in 1966. From that time to the present, he has been a Professor of the Rockefeller University.

Edelman is a member of the National Academy of Sciences, the American Academy of Arts and Sciences, the

American Society of Biological Chemists and the American Association of Immunologists, as well as a number of other scientific societies. He was a member of the Biophysics and Biophysical Chemistry Study Section of the National Institutes of Health from 1964 to 1967. Presently, he is an Associate of the Neurosciences Research Program at Massachusetts Institute of Technology, a member of the Board of Governors of the Weizmann Institute of Science and a member of the Advisory Board of the Basel Institute for Immunology.

He has given the Carter-Wallace Lectures at Princeton University in 1965, the National Institutes of Health Biophysics and Bioorganic Chemistry Lectureship at Cornell University in 1971, and delivered the Darwin Centennial Lectures at the Rockefeller University in 1971. In 1972, he was the first Felton Bequest Visiting Professor at the Walter and Eliza Hall Institute for Medical Research in Melbourne, Australia.

Edelman has received the Spencer Morris Award of the University of Pennsylvania in 1954, the Eli Lilly Award in Biological Chemistry given by the American Chemical Society in 1965, and the Annual Alumni Award of Ursinus College in 1969. In addition to his studies of antibody structure, his research interests have included the application of fluorescence spectroscopy and fluorescent probes to the study of proteins and the development of new methods of fractionation of both molecules and cells. His present research interests include work on the primary and three-dimensional structures of proteins, experiments on the structure and function of plant mitogens and studies of the cell surface.

In 1950, Edelman married Maxine M. Morrison. They have two sons, Eric and David and one daughter, Judith.

Rodney R. Porter

Born: 8 October 1917, Newton-le-Willows, United Kingdom

Died: 6 September 1985, Winchester, United Kingdom

Affiliation at the time of the award:University of Oxford, Oxford, United Kingdom

Prize motivation: "for their discoveries concerning the chemical structure of antibodies"

Rodney Robert Porter born 8 October 1917 at Newton-le-Willows, Lancashire, England.

He was educated at the Ashton-in-Makerfield Grammar School taking his Hons.B.Sc. (Biochemistry) in 1939 at the University of Liverpool and his Ph.D. at the University of Cambridge in 1948.

After one year's postdoctoral work at Cambridge, Professor Porter joined the scientific staff of the National Institute of Medical Research in 1949 and was there until 1960 when he joined St. Mary's Hospital Medical School, London University as the first Pfizer Professor of Immunology.

In 1967, he was appointed Whitley Professor of Biochemistry in the University of Oxford and Fellow of Trinity College, Oxford.

Amongst his awards are those of:

Fellow of the Royal Society, 1964Gairdner Foundation Award of Merit, 1966Ciba Medal (Biochemical Society), 1967Karl Landsteiner Memorial Award from the American Association of Blood Banks, 1968National Academy of Sciences, U.S.A., Foreign Member 1972

He took his Ph.D. at Cambridge under the supervision of Dr. F. Sanger investigating protein chemistry. In 1948 Professor Porter started to investigate the structure of antibodies, but on moving to Mill Hill he worked on methods of protein fractionation collaborating with Dr. A. J. P. Martin. The particular interest was in chromatographic methods of fractionation.

He returned to the study of the chemical structure of antibodies leading to the finding of the three fragments produced by splitting with papain in 1958-59. He continued this work at St. Mary's Hospital Medical School and put forward the peptide chain structure of antibodies in 1962.

Since moving to Oxford he has been concerned with the structure of antibody combining site, of the genetic markers of immunoglobulins and recently in the chemical structure of some of the early complement components.

During the war years 1940-46 Professor Porter was in the army serving with the R.A., R.E., and R.A.S.C., finishing with the rank of Major. He was with the First Army in 1942 in the invasion of Algeria and with the 8th Army during the invasion of Silicy and then Italy. He remained with the Central Mediterranean Forces in Italy,

Austria, Greece and Crete until January 1946.