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GEORGE DAVIS SNELL December ~ 9, ~ 903-fune 6, ~ 996 BY N. AVRION MITCHISON GENETICIST GEORGE SNEER iS known principally for his part in the discovery of H2, the major histocompatibility complex (MHC) of the mouse en cl the first known MHC. For this he shared the 1980 Nobel Prize in physiology or medicine. He was electec! to the National Academy of Sci- ences in 1970. Most of his life was spent at Bar Harbor, Maine, where he workocl in the Jackson Laboratory. George was prouc! of his New Englanc! roots, moral en c! intellectual. His life was passed in the northeast, apart from brief spells in Texas and the Midwest. He was born in BraciforcI, Massachusetts, ant! at the age of ~ 9 went to Dartmouth College, where he obtained his B.S, degree in biology in 1926. He went on to Harvard University, where he obtainer! his D.Sc. four years later at the Bussey Institu- tion. During his last year he served as an instructor back at Dartmouth, and in the following year served again as an instructor at Brown University. He then obtainer! a National Research Council Fellowship to work at the University of Texas in the laboratory of H. I. Muller (1931-33) en cl re- turnec! there 20 years later to spenc! a sabbatical year reacI- ing up on ethics, as mentioned below. He movecl to Wash- ington University in St Louis as an assistant professor 253
254 B I O G RA P H I C A L EMOIRS (1933-34~. In 1935 at the age of 32 he joiner! the Jackson Laboratory, then clirectecl by its founder Clarence Cook Little, where he remained until retirement in ~ 973. His cleath follower! a year after the Toss of his belovec! wife, Rhocia. They hacl three sons, who became respectively a manager of a ciata processing center, a manufacturer of hi- f; Toucispeakers, en c! an architect, all in New EnglancI. SacITy, one son suffered an untimely cleath in 1984. In an autobiographical note written in 1989 George writes. My paternal grandfather, my father, and a brother all held patents; now a son has one. None were big money makers, but in each case at least one had commercial value. I would thus assume that insofar as I have an incli- nation to invent, this came from my father's side of the family. My mother was . . . a natural planner, a faculty which showed in her carefully designed and tended garden. Gathering and arranging facts are, I think, important antecedents to scientific creativity, and insofar as I have been effective in coping with these antecedents, I think my debt is mostly to my mother. As a boy, aside from enjoying science and mathematics in school and read- ing an occasional book on science at home, I don't think I showed any unusual scientific bent. My family spent the summer months in South Woodstock, Vermont, which was then primarily a farming community. Every farmer had a rifle for hunting.... I remember trying to devise a mechanism for a repeating rifle that would be different from the two I was familiar with. This never got beyond the thinking stage and I doubt if it had a design that would work, but it was an activity that I enjoyed. In our year-round house in Brookline, Massachusetts, one of my friends and I had a rainy day activ- ity telling what we called change-around stories that certainly required some imagination. The idea was to get the hero of the story into the worst possible predicament and then leave it to the other storyteller to extricate him. It was not until I studied genetics with Professor John Gerould at Dartmouth College that I became sufficiently involved in any branch of science to think of making it a career. Even then, it was not until I gradu- ated that I finally decided, with the encouragement of Professor Gerould, to enter a graduate school.
GEORGE DAVIS SNELL 255 George also mentions his love of ball games, from chiTcI- hoocl on. Later his colleagues remember him playing volley ball with enthusiasm en cl he was very goocI. My own memory of George is of the warm welcome he gave me for a very happy year spent in his laboratory, in the excellent company of Nathan Kaliss, Sheila Counce, en cl Gustave Hoecker. George himself was away in Texas writing his ethics book for much of the time, but a rich moment in my life was at the end of the year when he whiskocl me off into the awesome presence of Little. I vastly appreciates! the liberal encouragement that they both gave (but still with a touch of caution on their part about referring to "antigens"), en c! was cluly impresser! when George later en- couragecl me to publish the work on my own. Personally George was gentle en cl considerate, but at the same time intellectually stalwart, determined, and cre- ative. Neither flamboyant nor self-assertive, he never built a school or in his formative years published many multi-au- thor papers, en c! he fount! little neec! for technical innova- tion. He worked within a tradition of classical Mendelian genetics that flourished through most of his lifetime en cl still connects tociay with molecular genetics. The Jackson Laboratory with its magnificent mouse facility suited George perfectly, en cl he proviclecl exactly the foresight en cl drive that it neeclecI. He was not a goof! speaker, so the relative isolation there must have been a benefit. In fact, George clefinecl the Jackson phenotype: Stick to your knitting for as Tong as it takes en c! let the breecling of mice set your pace. This is well illustratecl by his relationship with cellular im- munology. George was already working in transplantation at the end of World War II. He realized that immunology wouIcl burgeon en cl that his work on the MHC wouIcl help it to clo so. He initiated work on immunological enhance- ment en c! reviewer! clevelopments in immunology on sev-
256 B I O G RA P H I C A L EMOIRS eral occasions. Yet he never allowed! himself to be clivertec! from his commitment to genetics. Stories grew around this friencIly en cl unassuming man. Following the 1947 blaze in the Acaclia National Forest that clestroyocl much of the Jackson facility he restarted his re- search from the remnants that he helpecl rescue. In the furor after the Nobel Prize the Jackson receptionist cleniec! knowledge of him, and the reporters were told by his neigh- bors that yes, they had been expecting him to get a prize- for his vegetables. The stock from his prize chives is still hanclecl clown among the Jackson geneticists, en cl his veg- etable patch can still be seen on AtIan tic Avenue. {an Klein cites the mice that bear the label "/Sn" as his living monu- ment. On the acivice of GerouIcI, George went to graduate school at Harvarc! uncler the guidance of William Castle, a pioneer of mammalian genetics. George usecl to say that Castle as- signecl him to work with mice because he himself clicin't like their smell. The mice of the time were clomesticatecI, but clicl not belong to clefinecl laboratory strains of the moclern kind. To start with, George workocl on linkage in mice of the "fancy," using mutations collected! by amateur brawlers, such as short-ear, dwarf, ringed hair, hairless, en cl nakocI. By 1996 (his last en cl posthumous paper) he hacl stucliecl a total of 26 such visible mutations. This represented a major contribution to formal genetics, whose task it was to estab- lish the linkage groups of selected species such as the mouse. He clelightec! in the molecular characterization of these genes that began in the last years of his life. Certainly the visible mutations provecl very useful later when he came to map his immunological genes. In the 1930s George clevelopecl an interest in the new fielcl of physiological genetics. The control of growth in- trigued him, as it did others at the time, including Little. In
GEORGE DAVIS SNELL ~ .~ 257 retrospect one is amazes! at the temerity of the biologists of that era. In Oxford the young Peter Meciawar, whose icleas about immunology were later to converge with those of George, had begun his research career by studying the growth of embryos. George colIaboratecl briefly with Douglas Fal- coner of Edinburgh University, later a great authority on the genetics of mouse growth. Tociay the quantitative ge- netics of growth is still regarclecl as a formiciably clifficult subject. While at Harvard, en c! as was the custom of Harvarc! biologists, George spent summers working at Woocis Hole. There he joined Phineas Whiting, an earlier student of Castle's, in studying the genetics of the parasitic wasp Habrobracon. This species is the prototype example of haplo- diploidy (i.e., haploid mares, diploid females), end his 1932 en c! 1935 papers are clevotec! to this subject, in particular to the role of male parthenogenesis in the evolution of the social hymenoptera. The topic was to emerge again later, when Hamilton identified the relative genetic proximity of sisters in haplo-cliploicl species as a key to the evolution of altruism. George discusses the point in his 1988 book on ethics, en c! one wonclers what part this wasp playact in form- ing his abiding interest in the evolution of social behavior. Might he have become a sociobiologist hacl the right iclea struck him in time? For his postcloctoral work he joined the laboratory of H. I. Muller at the University of Texas. Muller hacl cliscoverecl that racliation incluces mutations in Drosophila. In a series of papers between 1933 en cl 1946 George proved that same effect couIcl be obtained in the mouse, as a representative species of mammal. The most striking effect of racliation, he found, was to produce transIocations en cl other chromo- somal abnormalities, which often reclucecl fertility. His careful analysis helped establish that these effects resulted from
258 B I O G RA P H I C A L EMOIRS chromosome entanglements former! at meiotic pairing that interfere to a variable extent with chromosome segrega- tion. The 1946 paper shows George scrupulously citing the work of a student of his who hac! been ciraftec! for military service, as well as that of his competitor Peter Hertwig, who hacl continual to publish papers on mouse genetics in Ber- lin until ~ 942! George's pioneering work on transiocation in mammals pointed in three future directions. Mis-segre- gation of rearranged chromosomes was founcl to unclerlie the weirs! phenomena clisplayoc! by the t-alleles at H2. Ra- cliation-inclucecl chromosomal rearrangements proviclecl fun- ciamental insight into the life span of human T cells. An cl postwar studies of the genetic consequences of the atomic explosions hinged largely on chromosomal rearrangements. The advent of nuclear weapons make his 1937 paper on the genetic effects of neutrons seem remarkably prescient. In- cleecl it was fortunate for immunology that George came to feel that racliation genetics hacl reached the point of climin- ishing returns, as otherwise he might have been sucker! into the post-1945 resurgence of the subject. In 1941 the first edition of Biology of the Laboratory Mouse appeared, edited by George and largely written by staff of the Jackson Laboratory. It became the stanciarcl work on the subject, along with the second eclition publishecl in 1966 that container! no less that nine chapters coauthored! by George. George's entry into immunogenetics, in 1943, came in the form of a stucly of sperm iso-agglutinins (i.e., strain- specific antibodies macle in one mouse strain are able to agglutinate sperm of another strain). This approach was natural to him, as he hac! Tong been interested! in breecling mice, en cl similar antibodies hacl long been known against reel bloocl A cells. In his previous work he hacl encountered mate sterility inclucec! by racliation en c! hac! stucliec! several
GEORGE DAVIS SNELL 259 of the aspects of reproduction relevant to running a large mouse colony. In retrospect it is worth noting that these anti-sperm antibodies have since become one of the few human immune responses that show clear-cut regulation by the major histocompatibility complex. In the same year George publishecl a joint paper with A. M. Clouciman that market! his clebut in tumor transplanta- tion, a fielcl of research that he came to dominate en cl in which he macle his great discovery of the major histocom- patibility complex. ClouUman had Tong been working with Little at the Jackson Laboratory. In ~914 Little (Science 40:904-906) proposal a genetic theory of tumor transplan- tation postulating that the susceptibility to a tumor trans- plant was cleterminecl by several dominant genes. An cl he en cl Tyzzer J. MecI. Res. 33~1916~:393-453) went on to esti- mate the number of these factors by challenging an F2 population with grandparental tumor. Thanks to Little's foresight Jackson Laboratory proceeclecl to collect en cl in- breec! numerous strains of mice en c! has ever since server! as the worIcl center for the distribution of mouse strains. Using the collection, George formulatecl the "funciamental rules of transplantation": that tumors could be transplanted freely within an inbred strain, into its F] hybrids with other strains, and into a Mendelian proportion of an F2. They were however rejectee! by other strains, as were tumors that originated in F] hybrids en cl were transplantecl into one of the parental strains. Inbred mouse strains, they concluclecI, clifferec! by only a few rejection-inclucing genes. Opinion grew that these rules must reflect an immune response to antigens expressed by the tumors, as well as by normal transplanter! tissue. From the early years of the twen- tieth century it was known that transplanted tumors were often rejected, en cl that this might represent a response to something specific to cancer a possibility that was to prove
260 B I O G RA P H I C A L EMOIRS an enduring hope of cancer research. W. Wogiom in his influential 1929 review rejected that view en cl arguccl in- steacl that tumors en cl normal tissue share a similar ability to elicit immunity. Later l. B. S. Haitians visitec! Little en c! brought back to Lonclon on the pet fleck of the liner Mauritania some of the new inbred strains. In his 1933 lec- ture to the Royal Institution he suggester! that each of Little en cl Clouciman's rejection genes might be "responsible for the manufacture of a particular antigen, as in the case of the rec! corpuscles." He encourages! his young colleague Peter Gorer to search for such antigens. Gorer workocl in parallel with Irwin en cl Coles, who hacl coiner! the term "immunogenetics" to describe their work on antigens of avian reel cells. Gorer raisecl rabbit antisera to reel bloocl cells of mice, which upon absorption clistin- guishec! two antigens present in different strains. He then joined George in demonstrating that his antigen II segre- gated in F2 mice together with the gene fused (Fu), which George hac! fount! to be linker! to transplant rejection. On this basis the gene uncoiling the antigen was namecl H2 (H for histocompatibility en cl 2 for antigen II) en cl represents the first sighting of what later came to be caller! the major histocompatibility complex. It is worth noting that their 1947 paper wisely refrains from claiming that the antigen expresser! on rec! blooc! cell antigen causer! the rejection, or that antibodies of the iso-agglutinin type were respon- sible. For another five years at least, George continued to refer to histocompatibility factors rather than antigens. Af- ter all, 20 years earlier WogIom had cited evidence that antibodies clicl not mecliate transplant rejection. George at this point made the wise decision to split the effort of his laboratory. To Nathan Kaliss he left the prob- lem of characterizing the histocompatibility factors, while his own group concentrated on the genetics. The isolation
GEORGE DAVIS SNELL 26 work hac! originally begun in collaboration with Clouciman. They sought simply to preserve tumors by freezing. Next they cliscoverecl that the new trick of freeze cirying couIcl be user! to preserve the immunizing material in tumors, al- though to their surprise this material often prolongecl (en- hancecI) rather than shortened the survival of subsequent transplants. KaTiss took up the problem with only limiter! success. Although the conditions uncler which enhancement takes place were clefinecI, little progress was macle with char- acterizing its mechanism. In 1960 Henry Winn shower! that lymphocytes couIcl transfer the effect. In retrospect the ef- fect joins other assorted clown-regulatory phenomena such as the transfusion effect (suppressing rejection of transplanter! tissue by prior transfusion of clonor bloocI) en cl the activity of various regulatory T cells. A clevelopment of this work sees that rare occurrence, George abroad. During the "Prague spring" of 1968 he vis- itecl Prague to collect the Menclel mecial, the first interna- tional recognition of his contributions to immunogenetics. He received a warm welcome from Milan Hasek, whose group he recognized for its scientific excellence, its stalwart clevo- tion to science through clifficult times, en c! the extent to which it shared his interest in immunogenetics. After the sacl encling of the "spring" Hilgert and Dement joined George's laboratory for a while. Hilgert attempted to carry on the work of Kaliss but with little success: The fielcl was waiting for better biochemical methods. Meanwhile, the genetics of the MHC, the work for which George received the Nobel Prize, steamed ahead on an ex- panding scale. The work depended on two simple but inge- nious procedures. One was to type existing inbrec! lines at H2 by means of the linkage with the Fused gene (as de- scribecl in 1951~. The second was to make new congenic lines (originally termec! "isogenic resistant," abbreviates! to
262 B I O G RA P H I C A L EMOIRS JR), in which various histocompatibility alleles were back- crossecl onto the same background. The H2 alleles were later termed "haplotypes" after the composite nature of this genetic region was cliscoverecI. Each step of the backcross- ing required 2 generations, en cl George cleciclecl that up to 20 generations were neeclecl to establish a new line. Over a clecacle this prodigious task proceeclec! steacliTy. By 1953 a total of 102 typings had yielded 9 H2 alleles, by 1958 the number rose to 12 alleles, en cl by 1969 to IS, encompassing all the main laboratory strains. With his colleague Graff, George later showocl that a congenic line wouIcl also reject skin grafts from its pair. In the meantime DonaTc! Shreffler en c! Jan Klein began to employ antibodies to explore H2 en cl clivicle it into its components within George's congenic lines. In this quest they sought en c! fount! recombination between ens! mark- ers of H2 en cl began to construct the map of the H2 region that figures in moclern textbooks. The term "major histo- compatibility complex (MHC)" was coined to describe this series of closely linkocl genes. In parallel Hugh McDevitt cliscoverecl that genes of the MHC hacl the unexpected func- tion of controlling the level of the immune response. In collaboration with George these two approaches converged in 1972 to map Ire] (immune response gene one, now iclen- tifiec! as H2A en c! H2E) within H2. With the advent of mono- clonal antibodies en cl later of DNA sequencing the map- ping proceeclecl rapicIly, curtailecl only by Shreffler's untimely cleath in the micist of his work on the C4 complement (Ss, SIp) locus. Ian McKenzie from Australia contributed to this effort during his sojourn in George's lab, in collaboration with the main serological work concluctec! there by Peter Demant and Marianna Cherry. The prodigious polymorphism of H2 required explana- tion, since obviously it clic! not exist merely to bother trans-
GEORGE DAVIS SNELL 263 plant surgeons. George in his 1981 "Future" paper rightly iclentifiecl regulation of the immune response as the func- tion of the MHC. He saw resistance to viral infection as the main driving force in its evolution, en c! its polymorphism as sustained by the wicler range of reactivity enjoyocl by het- erozygotes. Both these views are now generally accepted. Not all congenic pairs clifferec! from one another at H2, as jucigecl by the linkage test with the Fusecl gene. Differ- ences at the remaining "minor" H loci resulted in a weaker en c! more variable rejection that often requires! pre-immu- nization to prevent tumor growth. Goocl quantitation of the difference was obtained by Winn's transfer test. Con- secutive numbers were assignee! to these minor loci (e.g., H1, H3, Hip. By this en cl similar methods some 60 minor H loci have now been mapped. The frequency of single nucle- oticle polymorphisms in the genome suggests that there may be many more. The H3 complex is of particular interest, as shown in George's 1964 en c! 1967 papers. As Roopenian en c! Simpson write, "Snell and his collaborators' masterful exploitation of the fortuitous linkage of H3 to agouti en cl other visually detectable linker! loci . . . prover! that there were a mini- mum of two H genes within the H3 segment." Since then minor transplantation antigens have provecl valuable tools for probing the working of the immune sys- tem, notably in delineating the role of T-cell subsets en cl in studies of energy en cl other forms of peripheral tolerance. They are consiclerec! likely to have a therapeutic future, as contributing to the so-called graft-versus-leukemia effect af- ter bone marrow transplantation. As George's retirement approached, Cherry and McKenzie contributed to the discovery of non-H2 antigens on the surface of mouse lymphocytes, clefinecl by antibodies. The unfortunate name "cluster o If differentiation" (CD) is now
264 B I O G RA P H I C A L EMOIRS given to these molecules, which turn out to have important functions, as George preclictecl in his 1981 "Future" paper. Snell hacl an abicling worry about the contradiction be- tween evolution en c! ethics. He relates that this first struck him while teaching genetics en cl evolution at Washington University in 1933-34. He founcl the genetics easy, but the survival of the fittest clic! not seem compatible with his New Englancl upbringing. In 1953-54 he took a sabbatical to react further at the University of Texas in Austin en cl at Dartmouth College, leacling eventually to his book Search for a Ratio- na] Ethic in 1987. This is an extensive survey of human evolution from a genetic en cl anthropological standpoint, which argues that the origins of ethical behavior can be tracecl to particular periods en cl structures of human soci- ety. The book is hare! going. The mea cusps, surely clue to human genetics as practical in the twentieth century, is missing (couIcl George have been unaware of the ricliculous views about the genetics of human merit expresser! by his one-time collaborator R. A. Fisher, a grand oIcl man of ge- netics?) . Mussolini gets favorable mention, for what we wouIcl now call anti-terrorist activity (against the Mafia in Sicily) but not Hitler or Stalin. The 01d Testament and the Koran receive attention but not the Israeli-Arab conflict. A sen- sible discussion of kinship in ethology gets mixed up with some far-fetched sociobiology. From this book the author emerges as a true scholar, expert in biology but confused by ethics en c! the social sciences en c! quite unconcerned! with urgent problems of the day. George's discovery en cl characterization of the MHC is of funciamental importance to immunology en c! medicine. It enablecl the MHC to be subcliviclecl into sets of genes of different type. It allowocl the normal function of these genes to be cleterminecI, en c! lee! eventually to the structural en c!
GEORGE DAVIS SNELL 265 molecular characterization of the proteins that they encocle. It prepared the ground for HLA (the MHC of man), which rapicIly gainecl importance in organ en cl bone marrow trans- plantation en c! has saver! many lives. Tociay it is also impor- tant in predicting disease susceptibility en cl in the elusion of pepticle vaccines. Well clicl it merit its Nobel Prize. a Science moves on. Immunogenetics, in man en c! mouse, is now a sub-specialty of molecular genetics en cl genomics. The laborious methods of immunogenetics in the past are being replaced by DNA sequencing. Worldwide the bone marrow transplantation groups are engaged in clecicling whether sequencing HLA-cIass genes is worthwhile in prac- tice. The oic! transplantation tests en c! serology now seem oIcl hat. Nevertheless it was those oIcler methods that laicl the foundations on which we now builcI, en cl in viva trans- plantation tests will remain the endpoint for clinical trans- plantation. I AM GRATEFUL TO Jan Mein, Elizabeth Simpson, Derry Roopenian, Will Silvers, and Henry Metzger for help in preparing this memoir. The library of Jackson Laboratory has an extensive Snell archive, a useful Snell reprint collection, and a full Snell bibliography.
266 B I O G RA P H I C A L S E L E C T E D EMOIRS B I B L I O G RAP H Y 1928 A crossover between the genes for short-ear and density in the house mouse. Proc. Natl. A cad. Sci. U. S. A. 14:926-28. 1932 The role of male parthenogenesis in the evolution of the social hymenoptera. Am. Nat. 66:381-84. 1935 The induction by X-rays of hereditary changes in mice. Genetics 20:545-67. 1939 The induction by irradiation with neutrons of hereditary changes in mice. Proc. Natl. Acad. Sci. U. S. A. 25:11-14. 1941 Ed. Biology of the Laboratory Mouse. New York: McGraw-Hill. 1944 Antigenic differences between the sperm of different inbred strains of mice. Science 100:272-73. 1946 An analysis of translocations in the mouse. Genetics 31:157-80. 1948 With R. A. Fisher. A twelfth linkage group of the house mouse. Heredity 2:271-73. With P. A. Gorer and S. Lyman. Studies on the genetic and anti- genic basis of tumour transplantation: Linkage between a histo- compatibility gene and "Fused" in mice. Proc. R. Soc. Lond. Biol. 135 :449-505. 1951 With N. Kaliss. The effects of injections of lyophilized normal and
GEORGE DAVIS SNELL 267 neoplastic mouse tissues on the growth of tumor homoiotransplants in mice. Cancer Res. 11:122-26. With G. F. Higgins. Alleles at the histocompatibility-2 locus in the mouse as determined by tumor transplantation. Genetics 36:306- 10. 1954 The enhancing effect (or actively acquired tolerance) and the his- tocompatibility-2 locus in the mouse. 7. Natl. Cancer Inst. 15:665- 77. 1956 With S. Counce, P. Smith, and R. Barth. Strong and weak histocom- patibility gene differences in mice and their role in the rejection of homografts tumors and skin. Ann. Surg. 144:198-204. 1958 Histocompatibility genes of the mouse. I. Demonstration of weak Histocompatibility differences by immunization and controlled tumor dosage. 7. Natl. Cancer Inst. 20:787-824. Histocompatibility genes of the mouse. II. Production and analysis of isogenic resistant lines. 7. Natl. Cancer Inst. 21:843-77. 1960 With H. J. Winn, J. H. Stimpfling, and S. J. Parker. Depression by antibody of the immune response to homografts and its role in immunological enhancement. 7. Exp. Med. 112:293-314. 1961 With H. T. Winn and A. A. Kandutsch. A quantitative study of cellu- lar immunity. J. Immunol. 87:1-17. 1965 With H. P. Bunker. Histocompatibility genes of mice. V. Five new Histocompatibility loci identified by congenic resistant lines on a C57BL/10 background. Transplantation 3:235-52. 1966 With T. H. Stimpfling. Genetics of tissue transplantation. In Biology
268 BIOGRAPHICAL MEMOIRS of the Laboratory Mouse, ed. E. L. Green, pp. 457-91. New York: McGraw-Hill. 1969 With D. C. Shreffler. The distribution of thirteen H-2 alloantigenic specificities among the products of eighteen H-2 alleles. Trans- plantation 8: 435-50. 1972 With H. O. McDevitt, B. D. Deak, D. C. Shreffler, T. Klein, and T. H. Stimpfling. Genetic control of the immune response. Mapping of the Ir-1 locus. 7. Exp. Med. 135:1259-78. 1973 With I. F. McKenzie. Comparative immunogenicity and enhanceability of individual H-2K and H-2D specificities of the murine histo- compatibility-2 complex. J. Exp. Med. 138:259-77. With M. Cherry and P. Demant. H-2: Its structure and similarity to ML-A. Transplant Rev. 15: 3-25 1976 With T. Dausset and S. Nathenson. Histocompatibility. New York: Academic Press. 1980 The future of immunogenetics. Prog. Clin. Biol. Res. 45:241-72.
