It is a great honor to be able to convene this presentation ceremony of the 2002 International Prize for Biology in the presence of Their Majesties the Emperor and Empress and with the attendance of many eminent persons and distinguished guests from both Japan and abroad.

The International Prize for Biology was established in 1985 to celebrate the 60th anniversary of the reign of the Emperor Showa and to commemorate his long-held interest in scientific research in the field of biology. Since its inception, the purpose of this prize has been to recognize and to promote important advances in the study of biology.

This year marks the 18th award presentation. Having come thus far, I wish to take this opportunity to express my gratitude to all who are gathered here and to the many others whose valuable cooperation and support have made it possible to sustain this effort over so many years.

Dr. Masatoshi Nei, the winner of this year�fs Prize, developed statistical methods, and by applying them to molecular-level data, he devised a theoretical basis for explaining many experimental observations that had previous defied interpretation. Among them are the evolutional relationship among diverse species, the times of species divergence from common ancestors, and the location of gene regions in which natural selection operates. These achievements laid the theoretical foundation of the current field of molecular evolutionary biology, and have been of enormous contribution to scientific progress in biology.

Dr. Nei is most highly deserving of this prize. On behalf of the Committee, I would like to both congratulate him and extend him our deepest esteem for his most valuable work as a scientist and a researcher.

At this time, I would like to thank the members of the Selection Committee who bore the weighty responsibility of making this nomination. I would also like to take this opportunity to express my appreciation to the members of the universities, research centers, academic associations and other organizations, both in Japan and abroad, who have favored us with their recommendations of candidates for this year's prize.

As I explained at the beginning, having been awarded eighteen times now the International Prize for Biology has earned a degree of acclaim suitable to commemorating Emperor Showa�fs long devotion to biological research. The Committee on the International Prize for Biology has committed itself to making every effort to sustain the stature of this international Prize.

In closing, may I ask you for your continued encouragement and support for the work of the Prize Committee.

On behalf of the Selection Committee of the 2002 International Prize for Biology, I am pleased to report on this year's selection process.

The Selection Committee consisted of 19 members, including myself. Four of our members were highly authoritative foreign researchers, who had been specially commissioned to serve on the committee.

This year, the applicable area of the prize was stipulated as "Biology of Evolution." The committee distributed a total of 1,787 recommendation forms to Japanese and foreign universities, research centers, academic associations, individual researchers, and international academic organizations involved in this field of biology. In response, the committee received a total of 51 recommendations. As there was some overlapping, the actual number of individuals recommended was 37. They reside in 14 countries spread throughout the world.

The Selection Committee met a total of four times and very carefully reviewed all the candidates. Ultimately, the committee decided to recommend Dr. Masatoshi Nei of the United States of America as the awardee of the 2002 International Prize for Biology.

Dr. Nei was born in Miyazaki Prefecture in 1931. He graduated from Miyazaki University in 1953. After which, he went on to graduate school at Kyoto University, where he became an Assistant Professor. He was, then, employed as a Geneticist at the National Institute of Radiological Sciences in Chiba. In the meantime, he received his Ph.D. from Kyoto University in 1959. In 1969, he moved to the United States, where he became an Associate Professor and a full Professor at Brown University, and then a Professor at University of Texas at Houston. In 1990, he became a Distinguished Professor of Biology at Pennsylvania State University and the Director of its Institute of Molecular Evolutionary Genetics, where he continues to serve today. In 1985, he acquired US citizenship.

Dr. Nei carried out original work through which he devised innovative statistical techniques for conducting molecular-level studies of genetic diversity in populations and evolutionary relationships among species. He also developed methods for estimating the times of divergence of different species and detecting the gene regions in which natural selection takes place. Accordingly, Dr. Masatoshi Nei has contributed immensely to laying the theoretical foundations of the modern field of molecular evolutionary biology.

One of the best-known statistical methods developed by Dr. Nei defines, from molecular data on DNA and protein molecules, the degree of genetic difference between populations as �ggenetic distance.�h This method, named �gNei�fs Genetic Distance�h made it possible to estimate from molecular data the origins of populations and the times of their divergence from common ancestors, and is still in frequent use around the world. Applying the method to human populations, Dr. Nei obtained the first evidence pointing to the African origins of modern humans.

Using molecular data, Dr. Nei also constructed a mathematical theory for elucidating phylogenetic relationships. He and his colleagues developed a technique of inferring molecular phylogenies, known as the �gneighbor-joining method,�h which has become the most widely used method of constructing phylogenetic trees.

Furthermore, Dr. Nei was first to clarify the theoretical relationships of gene trees and species trees, thus providing a theoretical basis for explaining many experimental observations that previously could not be elucidated.

As we reviewed Dr. Nei's outstanding achievements, which have stimulated and propelled the advancement of scientific research in this field, we were convinced that his acceptance of the International Prize for Biology would add to the Prizes luminescence and prestige throughout the world.

In making our selection, the major criteria used by the Committee were the originality of the candidate's research and its international significance and contribution to advancing progress in the selected field of biology. We found Dr. Nei's work to more than amply satisfy each and every one of these criteria; and on this basis, we judged him to be the most highly suited candidate to receive this year's International Prize for Biology.

The Committee on the International Prize for Biology accepted our committee's recommendation of Dr. Nei and has bestowed upon him the 2002 International Prize for Biology.

With this, I conclude my report on the Selection Committee.

Thank you.

Address by His Majesty the Emperor

I should like to express my heartfelt congratulations to Dr. Masatoshi Nei on being awarded the 18th International Prize for Biology. This year, the Prize is awarded in the field of �gBiology of Evolution.�h Through his long years of research, Dr. Nei has conducted molecular-level studies on genetic diversity in populations and evolutionary relationships among species, developing his own innovative statistical methods, such as techniques for accurately estimating the time of divergence of different species and detecting the gene regions in which natural selection takes place. These advances have laid the theoretical foundation of contemporary �gmolecular evolutionary biology.�h

Among the statistical methodologies that Dr. Nei has devised is his world-recognized technique of estimating the degree of genetic difference between populations from protein and DNA data by defining the �ggenetic distance.�h This technique, called �gNei�fs genetic distance,�h is both widely used and highly regarded by researchers worldwide. Furthermore, Dr. Nei was the first to clarify the theoretical relationship between gene trees and species trees. He and his collaborator also developed the �gneighbor-joining method�h for inferring molecular phylogenies. This has become the most widely used method in the world for constructing phylogenetic trees.

I myself have long studied the gobioid fishes from a morphological point of view, and I co-authored and published a paper with a number of colleagues two years ago on an analysis of these fishes conducted from a molecular phylogenetic perspective. In this study, we applied the neighbor-joining method, and we obtained interesting findings on the morphogenetic relationships of these fishes. I was deeply impressed with the value of Dr. Nei�fs methods in understanding the affinities and phylogenies of organisms.

The products of Dr. Nei�fs research are widely used by researchers and contribute immensely to the advancement of biological sciences. With my sincerest hope that his research will continue to yield much fruit, I should like to reiterate my heartiest congratulations to Dr. Masatoshi Nei.

Distinguished guests, ladies, and gentlemen:

I should like to offer my congratulations on the occasion of this award ceremony for the 18th International Prize for Biology, which is taking place here today in the presence of Their Majesties the Emperor and Empress.

The International Prize for Biology was established in honor of the 60th year of Emperor Showa�fs reign and to commemorate his long years of biological research. Over the past eighteen years, the prize has earned high acclaim throughout the world as a prestigious award. I would like to express my congratulations to Dr. Masatoshi Nei who is being presented with the prize this year.

I understand that Dr. Nei has contributed immensely to all the biological sciences by laying the theoretical foundations of current molecular evolutionary biology through his original R&D on innovative statistical methods, including a method for estimating the times of species divergence from common ancestors. I should like to express my deepest respect to Dr. Nei for his achievements.

The Government of Japan is now energetically promoting scientific research and development under a national policy to make Japan into �ga nation richly endowed in scientific and technological creativity�h at the apex of global excellence. Underpinning this effort can be none other than a steady accumulation of scientific research achievements in biology and other fields of scientific inquiry. The promotion of creative and pioneering scientific research is indispensable to opening up new realms of possibility for humankind and creating a foundation for the further advancement of society. To this end, I encourage the scientists present today to persevere in your work and ask for everyone�fs understanding of and cooperation in your endeavor to expand the frontiers of human knowledge.

I sincerely hope that the International Prize for Biology will contribute to the further advancement of scientific research in Japan and throughout the world. In closing, I should like to wish you all the best of health.

Thank you.

Distinguished guests, ladies, and gentlemen:

I am truly delighted that the awards ceremony for the 18th International Prize for Biology is taking place here today, in the presence of Their Majesties the Emperor and Empress.

The International Prize for Biology was established in honor of the 60th year of Emperor Showa�fs reign, in order to commemorate his long years of biological research and to promote research in the field of biology. Every year, this honor is bestowed on a scientist who has achieved outstanding accomplishments in biological research and has made great contributions to the advancement of science throughout the world.

I understand that Dr. Masatoshi Nei, who has just been presented with this prize, performed research over many years on the genetic diversity of populations and evolutionary relationships among organisms on a molecular level, and that he has laid the theoretical foundation of current molecular biology. His superb work, which has contributed tremendously to all the biological sciences, is indeed most deserving of recognition by the International Prize for Biology-which is awarded for highly original, vanguard research. This is because Dr. Nei�fs research has provided the theoretical infrastructure for today�fs molecular evolutionary biology, a field on the razor edge of which outstanding scientists are compering fiercely, throughout the world. I am delighted to express my wholehearted congratulations to Dr. Nei, and to wish him every success in his future research.

As intellectual and creative work carried out in search of truth, scientific research has intrinsic cultural value and is critical in opening up entirely new frontiers of inquiry and achieving breakthroughs that can change the complexion of future society.

For this reason, the Ministry of Education, Culture, Sports, Science and Technology is working to make Japan �ga nation richly endowed in scientific and technological creativity.�h In this pursuit, the Ministry places particular emphasis on the life sciences as a priority field. We are actively promoting basic research that is both pioneering and cutting edge, and are advancing R&D that yields richly creative results at the world�fs highest level of excellence. I would like to urge all of the distinguished researchers gathered here today to redouble their efforts so as to advance further scientific research around the world.

In closing, I should like to express my respect and gratitude to the members of the International Prize for Biology Committee and to all the other individuals who have worked so hard in the administration of this celebrated award.

Thank you.

Your Majesties, Ladies, and Gentlemen:

It is a great honor to receive the 2002 International Prize for Biology, which is for the field of evolutionary biology. I understand that His Majesty the Emperor is an enthusiastic biologist and has published many papers on the taxonomy and evolution of Gobioid fishes including a paper on molecular phylogeny. This makes me especially happy to receive this year�fs award as an evolutionary biologist.

When I was a college student, I read a number of books on evolution. However, I was not particularly interested in this subject, because evolutionary biology at that time was highly speculative and it was difficult to prove or disprove various arguments presented for explaining evolution. I was then interested in population genetics, in which short-term evolution was studied by using mathematical theory and experimental verification. I was particularly interested in developing mathematical theory by which one could predict short-term genetic change of biological populations.

It was in the 1960�fs and the 1970�fs that various molecular techniques were introduced in the study of evolution. Before this time, the only way to identify homologous genes between different individuals was to conduct hybridization experiments, but in principle this could not be done between different species. Therefore, most experimental studies of population genetics or evolution were confined to individuals within species. Introduction of the new molecular techniques radically changed this situation. It removed the species barrier and made it possible to study evolutionary change of genes between distantly related organisms even between bacteria and humans. This opened up the new field of molecular evolutionary biology. One of the pioneers who developed various mathematical theories of molecular evolution was my mentor, Professor Motoo Kimura, who was the recipient of the 1988 International Prize for Biology.

Although the conceptual framework of molecular evolutionary biology was laid out largely by Professor Kimura and others, it was still necessary to develop statistical methods by which one can infer the evolutionary histories of populations and species and test various evolutionary theories with empirical data. I initiated this type of work around 1970 and tried to make evolutionary biology a testable science rather than a speculative one. I continued this work for over 30 years and applied the statistical methods developed to find some general principles of evolution.

During these years, I was very fortunate to have many students and collaborators, who joined in my efforts to understand evolution at the molecular level. I should also mention that the statistical study of molecular evolution was initiated not only by my group but also by other investigators, some of whom are attending this ceremony. I would like to take this opportunity to thank all of them who participated in our joint venture to make evolutionary biology a quantitative and hypothesis-testing discipline.

I have been a man of curiosity from the time of childhood. I feel very happy when I find some explanation even for a small mystery. This is especially so when I discover a simple rule underlying a seemingly complicated phenomenon. In 1960 and 1961, I visited the United States for the first time for over one year. At that time Japan was poor, and it was important to improve the economic status. By contrast, the United States was rich and allowed scientists to do both basic and applied research. In fact, original research was strongly encouraged. This environment inflamed my interest in doing research driven by curiosity, and in 1969 I moved to the United States to do basic research.

Japan is now a rich country, and I understand both basic and applied research is encouraged. I also hear that the universities and research institutes in Japan are now under reorganization to enhance original research and accountability to the public. With this new environment, I am confident that in the near future Japan will play an important role in the international scientific community as well as the world economy.

I am most grateful for the Selection Committee of the International Prize for Biology for recognizing the importance of molecular study of evolution and for selecting me as the recipient of this prestigious award. I humbly accept this award as a representative of the many scientists who have contributed to the progress of molecular evolutionary biology during the last 30 years.

I am very grateful for all the nice words that have been said about me. I also would like to thank all of you who have come here to attend the award-presentation ceremony. I finally would like to thank my wife, Nobuko, and my children, Keitaro and Maromi, for their patience and understanding while I devoted a large portion of my life to scientific research. Without their support and affection, I could not have accomplished any significant work.

The Committee on the International Prize for Biology awards the 2002 International Prize for Biology in the field of "Biology of Evolution" to Dr. Masatoshi Nei, Pennsylvania State University, USA     On September 17, 2002, the Committee on the International Prize for Biology (chaired by Dr. Saburo Nagakura, President of the Japan Academy) decided, based on the recommendation of the Prize�fs Selection Committee, to present the 2002 International Prize for Biology to Dr. Masatoshi Nei, Professor, Pennsylvania State University, USA. The field of specialization for the 2002 Prize is �gBiology of Evolution.�h   Process of Selection   The Selection Committee, composed of a chairman (Dr. Kunio Iwatsuki, Professor, The University of the Air) and 18 other members, including four foreign members, functioned as a sub-committee of the Committee on the International Prize for Biology. It sent out invitations for candidate nominations to academic institutions/associations and individual experts both at home and abroad in the subject field of specialization. Altogether these comprised 1,787 mailing addressees. Fifty-one nominations were received by the deadline. After adjustment for overlapping nominations, there were nominations for 37 individuals who reside in 14 countries spread throughout the world. Upon careful deliberation of the nominees at its first through fourth meetings, the Selection Committee decided to recommend Dr. Nei as the nominee for the 2002 Prize. Thereafter, the Committee on the International Prize for Biology formally endorsed the Selection Committee�fs recommendation.   Achievements Recognized by the Award   Dr. Masatoshi Nei has contributed immensely to laying the theoretical foundations of current molecular evolutionary biology. The award recognizes these distinguished contributions, which arise from his original work in developing many innovative statistical techniques for the study of genetic diversity in populations and evolutionary relationships among species at the molecular level, including methods for estimating the times of divergence of different species and detecting gene regions in which natural selection is operating. In a career spanning more than forty years, working with many collaborators, Dr. Nei has developed various statistical methods to determine the molecular mechanisms of biological diversity and evolution; he has also elaborated the underlying theory and applied these methods to the analysis of actual data. One of the best-known statistical methods developed by Dr. Nei defines the degree of genetic difference between populations and estimates this �ggenetic distance�h from molecular data on DNA and protein molecules. This measure makes it possible to estimate the origins of populations and the times of their divergence from common ancestors utilizing molecular data; known as �gNei�fs genetic distance,�h it is still in frequent use around the world. Dr. Nei himself applied it to human populations and obtained the first evidence pointing to the African origins of modern humans. In his research on mathematical modeling of DNA evolution, Dr. Nei devised several statistical methods for estimating the number of synonymous and nonsynonymous nucleotide substitutions. By applying these methods to the major histocompatibility complex (MHC) genes, which are involved in immune responses such as the rejection of transplanted organs, he then demonstrated that they make it possible to identify gene regions in which natural selection operates strongly. These methods continue to be used by a large number of investigators today. Dr. Nei further discovered that the puzzling diversity of the MHC genes in human populations is due to a special pattern of DNA evolution that occurs when genes protect their hosts from invaders such as viruses and bacteria. Dr. Nei has also constructed a mathematical theory of the phylogenetic relationships of genes using molecular data. With one of his graduate students, he developed a technique of inferring molecular phylogenies known as �gthe neighbor-joining method,�h which has become the most widely used method of constructing phylogenetic trees. Through these achievements, which laid the foundations for the theoretical framework of current molecular evolutionary biology, Dr. Nei has contributed enormously to the development not only of his own field, but of every branch of biology that is concerned with diversity and evolution.   Ceremony and Commemorative Symposium   The presentation ceremony for the 2002 International Prize will be convened on Monday, December 2, 2002 at the Japan Academy, Tokyo; and a commemorative international symposium will be held on December 3-4 at Plaza HEISEI in Tokyo. Committee on the International Prize for Biology Japan Society for the Promotion of Science 6 Ichiban-cho, Chiyoda-ku, Tokyo 102-8471 Telephone: +81-3-3263-1722, Facsimile: +81-3-3221-2470

Date of Birth:	January 2, 1931
Nationality:	USA
Position:	Evan Pugh Professor of Biology, Pennsylvania State University, University Park
Address:	Institute of Molecular Evolutionary Genetics Pennsylvania State University 328 Mueller Lab University Park, PA 16802 USA
		�@
Education and Career:
	1953	B.S. from Miyazaki University, Miyazaki, Japan
	1959	Ph.D. from Kyoto University, Japan
	1958-1962	Assistant Professor, Kyoto University, Japan
	1960-1961	Rockefeller Fellow, University of California and North Carolina State University
	1962-1969	Geneticist, National Institute of Radiological Sciences, Chiba, Japan
	1969-1972	Associate Professor to Professor of Biology, Brown University, USA
	1972-1990	Professor of Population Genetics, University of Texas at Houston
	1990-1994	Distinguished Professor of Biology, Pennsylvania State University, University Park
	1990-present	Director, Institute of Molecular Evolutionary Genetic, Pennsylvania State University, University Park
	1994-present	Evan Pugh Professor of Biology, Pennsylvania State University, University Park
		�@
Awards and Distinctions:
	1977	Japan Society of Human Genetics Award
	1989	Honorary Member, Genetics Society of Japan
	1990	Fellow, American Academy of Arts and Sciences
	1990	Kihara Prize, Genetics Society of Japan
	1993	Fellow, American Association for the Advancement of Science
	1996	Honorary Member, Japan Society of Human Genetics
	1997	Member, National Academy of Sciences, USA
	2000	Honorary Member, Japan Society for Histocompatibility and Immunogenetics

Dr. Masatoshi Nei is one of the main architects who laid the theoretical foundations of current molecular evolutionary biology. By developing various statistical methods and applying them to molecular data, he pioneered new ways of studying the genetic diversity of populations, evolutionary relationships among organisms, the times of species divergence from common ancestors, the location of gene regions in which natural selection is operating, and related areas. The methods he introduced have made it possible to obtain quantitative estimates of various parameters of evolutionary importance that could not have been measured experimentally. Through these achievements, Dr. Nei has not only made the latest findings at the molecular level available to evolutionary biologists, but has contributed greatly to the birth of molecular evolutionary biology and its establishment as a positive science in which hypotheses can be verified quantitatively, rather than being discussed solely on a conceptual level.

One of the best-known statistical methods developed by Dr. Nei is concerned with defining the degree of genetic difference between populations and estimating this �ggenetic distance�h from protein and DNA data. Dr. Nei�fs method, which he proposed in 1972, uses polymorphism data (individual differences in proteins and DNA) to estimate the origins of human and other populations, the times at which they diverged from common ancestral forms, and so on. Under the name �gNei�fs genetic distance,�h it has been adopted worldwide by most researchers in related fields and is still frequently used today.
In 1974, Dr. Nei applied his new measure to human populations and showed that the genetic variation among Europeans, Asians, and Africans is only about 10 percent of the average variation within the three major human ethnic groups; he also estimated that Africans diverged from Europeans and Asians about 100,000 years ago, and Europeans and Asians diverged from each other about 50,000 years ago. This was the first evidence pointing to the African origins of modern humans.

Focusing at an early stage on DNA sequence data, from the late 1970s Dr. Nei initiated mathematical modeling of DNA evolution. In the mid-1980s, he worked on statistical methods for estimating the number of nonsynonymous substitutions (nucleotide substitutions that result in amino acid replacements) and synonymous substitutions (those that do not result in amino acid replacements), and proposed that the ratio between the two be used to identify gene regions in which natural selection is operating strongly. These methods are now being utilized by a large number of investigators.

During the 1980s, Dr. Nei began to construct a mathematical theory of the phylogenetic relationships of genes, using molecular data such as DNA and protein sequences. He and his collaborators were the first to clarify the theoretical relationships of gene trees and species trees, thus providing a theoretical basis for explaining many experimental observations that had previously defied interpretation. With one of his graduate students, Dr. Nei also developed a technique of inferring molecular phylogenies known as the �gneighbor-joining method,�h which has become the most widely used method of constructing phylogenetic trees.

In 1988, Dr. Nei turned his attention to the major histocompatibility complex (MHC) genes, which are involved in immune responses such as rejection of organ transplants, and showed that the puzzling variability of these genes in human populations is due to a special pattern of evolutionary change of DNA that occurs when MHC genes protect the host from invaders such as viruses and bacteria. This evolutionary pattern has since been shown to hold true not only for humans but for vertebrates in general.

Dr. Nei also studied �gmultigene families,�h in which multiple copies of member genes are present in the genome, as is generally the case for the immune system genes. He discovered that the member genes typically evolve according to a pattern called �gbirth-and-death evolution,�h in which, after gene duplication, the member genes evolve more or less independently of one another, with some staying in the genome for a long time and others disappearing.


These achievements laid the foundation on which rests the theoretical framework of current molecular evolutionary biology. Thus, Dr. Nei has made an enormous contribution to the development of his chosen field of inquiry.

Nei, M. 1969. Gene duplication and nucleotide substitution in evolution. Nature 221:40-42.
Nei, M. 1972. Genetic distance between populations. American Naturalist 106:283-292.
Nei, M. 1973. Analysis of gene diversity in subdivided populations. Proceedings of the National Academy of Sciences USA 70:3321-3323.
Nei, M. 1975. Molecular Population Genetics and Evolution. North-Holland, Amsterdam and New York.
Nei, M. 1978. Estimation of average heterozygosity and genetic distance from a small number of individuals. Genetics 89:583-590.
Nei, and Li, W. H. 1979. Mathematical model for studying genetic variation in terms of restriction endonucleases. Proceedings of the National Academy of Sciences USA 76:5269-5273.
Nei, M. and Roychoudhury, A. K. 1982. Genetic relationship and evolution of human races.�@Evolutionary Biology 14:1-59.
Saitou, N. and Nei, M. 1987. The neighbor-joining method: a new method for reconstructing�@phylogenetic trees. Molecular Biology and Evolution 4:406-425.
Nei, M. 1987. Molecular Evolutionary Genetics. Columbia University Press, New York.
Hughes, A. L. and Nei, M. 1988. Pattern of nucleotide substitution at major histocompatibility complex class I loci reveals overdominant selection. Nature 335:167-170.
Nei, M. Gu., X., and Sitnikova, T. 1997. Evolution by the birth-and-death process in multigene�@families of the vertebrate immune system. Proceedings of the National Academy of Sciences USA 94:7799-7806.
Nei, M. and Kumar, S. 2000. Molecular Evolution and Phylogenetics. Oxford University Press, Oxford.