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REBECCA CRAIGHILL LANCEFIELD January 5, 1895-March 3, 1981 BY MACLYN McCARTY REBECCA CRAIGHILL LANCEFIELD was born onJanu- ary 5, 1895, in Fort Wadsworth, New York, where her father, Col. William E. Craighill, was stationed as an officer in the U.S. Army Engineer Corps. As a member of an Army family, she livecl in many different communities cluring her early years. After graduating from Wellesley College, how- ever, and spending one year teaching in a girls school in Ver- mont, she returned to New York City. Except for a year's sojourn at the University of Oregon, she spent the remainder of her life there. Her first move toward a career in science apparently came at Wellesley. Stimulated by her roommate's course in zoology, she dropped her notion of majoring in French and English and concentrated her efforts on biology. By the time she err in 1 91 fi she was easer to herein Graduate training. C~ ~ ~ ~ ~ ~ ~ ~ ~ c~ c~ c~ ~ But she was forced to compromise: funds were short because of the death of her father, and her mother needed her help in supporting her five sisters. She saved enough from her earnings as a teacher cluring the following year to enable her to accept a scholarship with graduate tuition at Teachers' Col- lege of Columbia University. Fortunately, although this schol- arship (established by the Daughters of the Cincinnati for daughters of Army anc! Navy officers) specified Teachers' 227

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228 BIOGRAPHICAL MEMOIRS College, it was not necessary for her to take her courses there. Thus she spent the year in Hans Zinsser's Department of Bacteriology at the College of Physicians and Surgeons of Columbia University. Here she was able to broaden substan- tially her knowledge and experience in the branch of biology that interested her most. There were a series of notable events in Rebecca Craig- hill's life in the spring of 1918. She receiver! her master's degree from Columbia University and shortly thereafter was married to Donalc] Lancefield, a fellow gracluate student at Columbia who was in the famous Department of Genetics under T. H. Morgan. Even more significant from the point of view of her future career in research, her application for a position at the Rockefeller Institute for Medical Research was accepted. That June she became a technical assistant to 0. T. Avery and A. R. Dochez. The timing of her arrival at the Rockefeller Hospital was of considerable importance in shaping the course of her life's work. Until late in 19 ~ 7, Avery and Dochez had concentrated their efforts on studies of the pneumococcus. At that time, however, they traveled to Texas as consultants to the Surgeon General of the Army to investigate an outbreak of serious streptococcal infections that tract been superimposed on a measles epidemic in a number of military installations there. Returning to New York with a collection of streptococcal strains that tract been isolates] cluring the visit, they set about trying to determine whether, as in pneumococcus, there were separate and distinct types of streptococci involved in the ep- idemic rather than a single unvarying pathogen. Their ap- proach was to use the serological procedures that hacl prover! successful in delineating pneumococcal types: the agglutin- ation reaction anc! the protection of mice with specific anti- sera. Their progress in these efforts was reported at a confer-

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REBECCA CRAIGHILL LANCEFIELD 229 ence on Streptococcus hemolyticus that was held at the Princeton laboratories of the Rockefeller Institute on June I, LOIS. The discussions at the conference dealt with various aspects of the problem of streptococcal infections, but much attention was focused on the recently isolated strains. Avery prefaced his comments with this statement: "It is rather difficult from a stucly of the strains that we have isolated to tell whether they are all alike, whether they constitute one or several types." Dochez later enlarged on this point, describing the difficulties they had encountered with agglutination reactions as well as with mouse protection experiments. He conclucled: "Up to now, however, we have been unable to obtain immune serum which affords any considerable degree of protection for white mice against experimental infection. We are still work- ing along this line and it is possible that the proper combi- nation of immune serum and test animal may be obtained." It was to assist in this effort that Rebecca Lancefield was brought into the laboratory soon after. Anct although she had had no real opportunity before to display her special talents, it was clearly a case of bringing the right person to the right place at the right time. Within a year they had together iden- tifiecl four distinct serological typesas cletermined both by agglutination and mouse protectionthat server! to classify 70 percent of the 125 strains stucliecl. The paper describing these results was submitter! for publication on June I, 1919, one year to the clay after the conference. There can be little oubt that Rebecca Lancefielcl's native talent for solving this type of problem, perhaps accelerated in its development under the tutelage of two established masters, was a prime factor in the success of these studies. That she contributed much more than simply technical help was tacitly acknowI- eciged by the inclusion of her name as a coauthor of the paper, a type of recognition selclom accorded to technical assistants in those clays. It was a major publication, running

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230 BIOGRAPHICAL MEMOIRS to some thirty-four pages and replete with tables that docu- mented the findings in great detail. In addition to being the first account of specific types among the hemolytic strepto- cocci, it also represented a record of the first encounter be- tween these microorganisms and the investigator who was destined over the next five decades to become the master of their diversity. At this point, however, the work with these streptococcal strains ended temporarily. The war was over and with it the Army support for the studies. Dochez went to Johns Hop- kins; Avery returned to his first love, the pneumococcus; and Lancefield moved back to Columbia where she worked as a research assistant on problems of Drosophila genetics. Never- theless, the streptococcal strains were not all simply dis- carded. Some of them remain today in the Lancefield colIec- tion as reference type strains of group A streptococci, still identified by the same letter and number designation that was assigned on their isolation in ~ 9 ~ 7. In 1922, after her year at the University of Oregon, dur- ing which she and her husband Donald both taught, I.ance- field came back to the Rockefeller Hospital for good. Mrs. L. (as she came to be affectionately known to her colleagues) was now associated with the rheumatic fever service of Dr. Homer Swift rather than with the pneumonia service. She was also enrolled again as a graduate student at Columbia, and most of the laboratory work for her Ph.D. thesis was carried out at Rockefeller on a problem concerned with the so-called "green" or viridans streptococci. These streptococci were erroneously suspected of having something to do with rheumatic fever; her studies, published in her thesis and in two papers in the Journal of Experimental Medicine, helped to dispel this notion. The viridans streptococci are an extraor- dinarily heterogeneous and protean group of microorga- nisms. It must have been of some relief to her and of con-

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REBECCA CRAIGHILL LANCEFIELD 231 siclerable importance to science that she was able before too long to resume her stucties of hemolytic streptococci. It is important to realize that the relationship of hemolytic streptococci to human disease was not well characterized in the perioc! immecliately following WorIct War I. They were looked upon as important primarily as secondary invaders in such situations as puerperal fever, wound infections, and pneumonia that followed measles or influenza, as in the Army camp epidemics. The great prevalence of primary streptococcal sore throat clid not appear to be clearly recog- nizecl, ant! the key role of streptococci in scarlet fever was yet to be discoverecI. There was even less of a clue with regard to their implication in the pathogenesis of rheumatic fever and gIomerulonephritis. Thus Lancefielct's early studies were initiated before the present picture of streptococcal disease had been formulatecI. The results that she obtained hacT much to do with originating anct crystallizing these concepts ant! with providing a basis for understanding the clinical and epidemiological patterns of disease caused by these orga- nisms. Although Lancefield was no longer directly associated with Avery, their laboratories were in close proximity, ant! she continued to look to him for advice ant] counsel in the de- velopment of her research. As a great admirer of his scientific insights and approaches, she was well prepared to bring to her studies of the streptococci the same points of view that he tract usect so successfully in the case of pneumococci. Con- sequently, she consiclered the laborious and cletailed serolog- ical analysis of the large family of streptococci as being pri- marily an essential means to a more significant enct: that of determining the chemical nature and biological significance of the antigenic substances responsible for the serological re- actions. The systematic classification that emerged from her serological grouping and typing of streptococci was not in

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232 BIOGRAPHICAL MEMOIRS her minct the ultimate goal of her research. Rather, it was a neecled step in identifying the most significant antigens and determining their role in the ctisease-producing capacity of . . the microorganisms. In the micI-1920s she succeecled in obtaining two antigens in soluble form from hemolytic streptococci: one that was type specific and responsible for the distinction between the strains from the epidemic in 1918 and another that was spe- cies specific anct present in all of the human strains that she examined. She soon encountered a surprising result in at- tempting to determine the nature of the type-specific anti- gen. Avery and Hei~lelberger had earlier establishecl that the type-specific antigens of the pneumococcus were polysac- charicles present in the capsule of the organism; subse- quently, other pathogenic bacteria had been founct to be similarly equippec! with capsular polysaccharicles that deter- minect type specificity. Lancefielct thus anticipated a similar situation in streptococci, but after careful studies was forced to conclucle that this was not the case. Her soluble, type- specific antigen of streptococcus was clearly a protein, which she later clesignated as M-protein on the basis of the associa- tion of the antigen with the matt colony form of the organism when grown on an agar medium. The M-protein appeared to serve essentially the same function in determining the vir- ulence of hemolytic streptococci that the capsular polysac- charicle diet in the pneumococcus. Her soluble species-specific antigen did, however, prove to be carbohydrate in nature and was designatecl the C- carbohycTrate. (The continuing close relationship with the Avery laboratory is illustratect by the fact that when Avery anct his colleagues shortly thereafter found an analogous species-specific carbohydrate in pneumococcus, it was also callecI C-carbohydrate or C-substance.) The great impor- tance of the streptococcal C-carbohydrate, however, proved

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REBECCA CRAIGHILL LANCEFIELD 233 to be in the sorting out of the many different varieties of hemolytic streptococci that exist in nature. As she received more strains from numerous sources, it became apparent to Lancefield that her species-specific antigen was really group specific. It was common to strains isolated from strep throat ant] certain other human diseases, but a group of strains from bovine mastitis tract a quite different group-specific car- bohyctrate and those from horses with strangles still a third. A continuation of this process establishect that there are sev- eral distinct serological groups of hemolytic streptococci in nature. Their differentiation provect of great importance in the study of streptococcal disease. Lancefielc! designatect the human strains that tract been the object of her initial studies as group A and assigned let- ters of the alphabet to the others in sequence. Group A strep- tococci are responsible for most of the serious streptococcal infections of man, anc! it is infection with this group of or- ganisms that leads to the poststreptococcal sequelae, rheu- matic fever and gIomerutonephritis. But the other groups of streptococci, regarcIless of their normal habitat, also occur in man and may be associated with disease. Group B strepto- cocci, for example, which were initially encountered in cattle, are not uncommon in man and today are receiving much attention as the cause of septicemia anct meningitis of the newborn. Lancefielct carried out extensive studies of group B streptococci that laict the necessary groundwork for the present efforts to clear with this pediatric problem. In con- trast to the situation in group A with its M-protein, she found that the type-specific antigens of group B streptococci are capsular polysaccharides, fully analogous to the pneumococ- cal polysaccharides. In working out the interrelationships be- tween the several prevalent types of group B streptococci, she showed that specific antibodies to the capsular polysac- charide were highly protective against experimental infec-

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234 BIOGRAPHICAL MEMOIRS tions. Many years later she returned to work on group B streptococci and initiates! studies on their complex biochem- ical and antigenic structure that continue to be pursuccT in numerous laboratories throughout the world. In company with many other experimentalists, Rebecca Lancefield's enthusiasm for working at the laboratory bench clicl not extent! to the painful process of writing up the work for publication. She worked for nearly four years on the he- molytic streptococci without publishing any of her findings, but she quickly remectiect the situation with a flurry of seven papers, all appearing in the Journal of Experimental Medicine in 1928. These papers includes! the first description of her M-protein ant] C-carbohydrate, with details of their chemical anc! immunological properties, and some information on their relationship to the bacterial cell. Her continuing work built on this base of new knowledge and lecl to the ctifferen- tiation of serological groups of streptococci and clelineation of the biological significance of the type-specific M-protein. An interesting episode in the further sorting out of strep- tococcal diversity relates to Lancefield's exchanges with Fred Griffith, the noted British microbiologist. Griffith, after his famous work on the discovery of the transformation of pneu- mococcal types, hac! turned to studies of hemolytic strepto- cocci. His technical approach differed significantly from that of Lancefield: he dependecl primarily on slide agglutination for serological differentiation of his strains, and she used a precipitin technique that depended on the property of her soluble antigens to give visible precipitates when mixed with antisera. Both workers usecI extensive adsorption of their antisera with heterologous strains to eliminate cross- reactions. Griffith examined a large number of human strains by his procedure and published his first extensive cle- scription of types of streptococci early in 1935.

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REBECCA CRAIGHILL LANCEFIELD 235 On January 22, 1935, LancefielcT wrote to Griffith request- ing a reprint of his paper, and she included the following comment: "l have just react your paper in the current Journal of Hygiene with the greatest interest. ~ shouIcl not have sup- posec! it possible to classify the majority of strains of S. py- ogenes into so small a number of types as 27. It certainly makes a much more workable situation in this group if one can clo that." Her interest had obviously been captured, and two months later she wrote requesting his cultures and samples of his antisera "to compare the types that ~ have encountered with yours." This began a long series of ex- changes of strains, sera, and ciata that was prematurely ter- minatect by Griffith's tragic death in the London Blitz in ~ 940. The two workers had great respect for one another even though they did not see eye to eye on methodology and were never converted to each other's approaches. There was much in common between the types clefinec! by the two different techniques, and Lancefielct arloptect the numbers that Grif- fith had assigned to his types in order to achieve uniformity. In a few cases discrepancies arose because Griffith was not grouping his strains on the basis of C-carbohyctrate, and strains that slid not belong to group A were inclu(lecI among his types. Another source of discrepancy lecI Lancefield to the discovery of a seconc! surface protein antigen of group A streptococci, which she designatecl as T-antigen. T-antigen couIc] take part in slide agglutination and thus be cletectect by Griffith, but it was not present in the soluble M-protein extracts. Subsequently, LancefielcI ant! her colleagues were able to show that T-antigen unlike M-protein has no re- lation to virulence and, further, that the same or closely re- latecl T-antigen may be present on cli~erent M types. In the end, M types became the standarc! classification for bacteri- ological, clinical, and epiclemiological stuclies, even though T

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236 BIOGRAPHICAL MEMOIRS typing by slide agglutination remains an adjunct technique applicable to a number of situations in which M-protein is absent or difficult to detect. As it turned out, the doubts that Lancefield expressed to Griffith about being able "to classify the majority of strains of S. pyogenes into so small a number of types as 27" proved to be well founded. The total number of recognized types has been added to by laboratories all over the world and is now well over sixty. She herself had little interest in the busi- ness of identifying new types, preferring to devote her ener- gies to the biological properties of the organism and their bearing on disease-producing capacity. The most dramatic illustration of the fruits of this approach is the unfolding of the story of the central role of M-protein in streptococcal infections. This surface antigen not only determines the type specificity of the numerous strains of group A streptococci but also serves to protect the organism from host defenses. When M-protein is present, the white blood cells appear to be unable to engulf and destroy the organisms; in the pres- ence of specific antibody, however, this protective effect of M- protein is neutralized and the white cells can do their job. These facts led to the concept that immunity to streptococcal disease is primarily type specific and that recovery from in- fection with one type does little to provide protection against the numerous other types of group A organisms. This served to explain why repeated strep throats were so common in childhood and why rheumatic fever is a notoriously recurrent disease. Thus her work on this antigen provided the basis for a better understanding of the epidemiology of the disease and a more rational approach to its control. The work that she and her colleagues pursued during World War IT continued with the sorting out of the various antigens, especially the relationships between M- and T- antigens. At the same time she supervised the large-scale pro-

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REBECCA CRAIGHILL LANCEFIELD 237 auction of the grouping and typing sera that were provided to the military services for the first intensive studies of the epiclemiology of streptococcal disease using the powerful tools that she tract clevelopect. In the postwar years she re- sumec] her efforts to purify anc! characterize the properties of the important antigens. She carried out extensive studies of representative M- and T-antigens, a new surface protein that she designated as it-antigen, and the polysaccharide an- tigens of group B streptococci. In addition, in an illuminating study of the persistence of type-specific antibodies in man following group A streptococcal infections, she showed that lasting immunity to the M-antigen is commonly encountered. Over the course of her work a vast number of streptococ- cal strains were sent to her; most of these are still preserved in the Tyophilized state in her collection of some thousands of different strains. They were sent to her for identification, for a confirmation of identity, or because of some special fea- ture of the situation in which they were obtained. They all received attention and analysis, resulting in a few clozen vol- umes of loose-leaf notebooks, in sturdy harcI-cover binders, in which the data on each strain are recorcled. Much of this information is written in her own hand, anct it took some experience to be able to decipher her notes. But with per- sistence one couicI usually learn what he wanted to know about the strain in question. An equally large set of note- books dealing with her research projects also exists, and these are even more Circuit to decipher. (On occasion she even had trouble herself when trying to review experiments car- riec! out two or three clecacles earlier.) Rebecca Lancefield's devotion to her streptococcal studies was just as clurable and persistent as the type-specific anti- bodies that she had describecI, and she maintained her lab- oratory activity until a few months before her death. In June 1979, sixty years after her arrival at the Rockefeller Hospital

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238 BIOGRAPHICAL MEMOIRS to work with Avery anct Dochez, she was still coming in reg- ularly, driving her own car back and forth from Douglaston, Queens, as she had since before the war. Although the an- noying infirmities of age began to make it impossible for her to maintain her customary schedule, she clict not abandon the effort until Thanksgiving Day, 1980, when she fell at home anct broke her hip. She never regained full mobility, ancT she cried on March 3, 1981. Many of her colleagues fee! that there was an inexplicable delay in general recognition and appreciation of her great scientific contributions. There is certainly some truth in this, but it must also be notect that among microbiologists she had long ago attainer! international stature as the outstanding au- thority on streptococci. Both the national and international organizations devotee] to streptococcal problems have re- namec! their groups "The Lancefield Society," the former while she was still active. As further evidence of her recog- nition within the general field, she was elected president of the Society of American Bacteriologists in 1943 and of the American Association of Immunologists in ~ 96 ~ . Even though they may have been somewhat clelaye(1 in arriving, a number of other honors also came to her. She receiver! the T. Duckett {ones Award of the Helen Hay Whitney Foun- ciation in 1960, the American Heart Association Achieve- ment Award in 1964, and the Medal of the New York Acacl- emy of Medicine in 1973. Rockefeller University recognized her contributions anct long service to the institution with an honorary D.Sc. in 1973; her alma mater Wellesley College followed suit with a similar honor on the occasion of the six- tieth anniversary of her graduation in 1976. She was electect to the National Academy of Sciences in 1970. Regrettably, this came too late for the Lancefiel(ls to enjoy the fellowship of the annual meetings with their long- time friends, the A. H. Sturtevants. "Sturt" and his wife,

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REBECCA CRAIGHIEE EANCEFIEED 239 Phoebe, had regalecl them for years with tales of the Acact- emy meetings cluring their sharer! summer holidays at Woods Hole. Rebecca attendee] a few meetings, but T am sure that she missed the special flavor that might have been contrib- utecl by the presence of her friends. She always spent the summer at Woods Hole, a place that was second in her heart only to her laboratory. For the most part she dial not engage in laboratory work or writing there, the stay being reserved for renewal ant! recreation. In fact, a major aim was to escape the hot, humid weather of a New York City summer, which she detested; and in any event, throughout her early decades at Rockefeller it was impractical to try to do bacteriological or immunological work where neither the laboratories nor the animal quarters were air conctitionecI. She found Woods Hole ideal for relaxation, tennis, anc! especially swimming, an activity that she pursued to her final summer. The description of Lancefield's scientific contributions gives an incomplete picture of her life in the laboratory. As single-mindecl as she was in the pursuit of her research goals, she coup always finct time to provide advice and assistance to other workers, both within and outside the laboratory. A visitor with an interest in streptococcal problems would leave with a thorough indoctrination anct with most of his ques- tions answeredas well as with a collection of cultures of reference streptococcal strains anct samples of the relevant antisera. Streptococcal strains and antisera, together with di- rections for their use, were freely supplied to laboratories all over the worIcI. The younger associates and postdoctoral fel- lows in our group found that she was not only ready to help whenever neecled but that she expected to participate fully in all of the activities of the laboratory, including the parties and informal get-togethers. The pre-Thanksgiving eggnog party that she initiated is still carrier! on today, using her recipe.

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240 BIOGRAPHICAL MEMOIRS Since ~ became the heact of the rheumatic fever service after Dr. Swift's retirement in ~ 946, my own clirect association with Rebecca Lancefielcl extendect over more than half of her career at Rockefeller. Many of her major contributions tract been completec! anct the groundwork already laid for others by this time, but ~ tract ample opportunity to observe her working methods at first hand and to collaborate with her in more than one research project. Out of this came some in- sight into the qualities that were responsible for her success as an investigator. Because of her intuitive recognition of the great complexity of hemolytic streptococci, she was fully aware of the inherent danger of drawing premature conclu- sions from limiter] data. Accorctingly, she could never be sat- isfiec! with the results obtained with one or two strains exhib- iting a given characteristic after analysis with one or two antisera. It was always necessary to examine all available strains with each of many antisera, a procedure that greatly increased the burden of the analysis because of the diversity of the organisms and the heterogeneity of the antibody re- sponse of different rabbits to the multiple antigens involvecI. Such careful investigations, however, prevented her from drawing misleacling and oversimplistic conclusions, and her meticulous approach is responsible, ~ believe, for the great durability and reproclucibility of her publishecl finctings. Rebecca Lancefield never clevelopecl very much sympathy for the modern feminist's point of view on women in science. She was not enthusiastic about honors that recognized her as the "first woman" to clo this or that and preferred those that came without reference to her sex. She had no illusions about the difficulties of having both a scientific career anct a family, but she felt that with determination and hard work it was possible without special treatment. In the case of her own small family, her efforts to provide a rewarding home life along with her scientific pursuits were notably successful,

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REBECCA CRAIGHILL LANCEFIELD 241 even though there must have been problems at times in adapting. She commuted by car from Douglaston, Long Is- lanct, for over forty years, which by itself was something of a triumph, considering bad weather, gasoline shortages, anct the like. Donalct Lancefield survived Rebecca by only a few months. Their daughter, Jane Hersey, clid not follow her par- ents into a career in biology ant! received her education in the classics. She has not managed to avoid science altogether, however; for some time, she served as a book review editor for The American Scientist. She ant] her husband, George Her- sey, have two sons, Donald ant! fames. M U C H O F T H E M A T E R ~ A ~ on which I drew for this memoir came from my own files. I am indebted, however, to the Rockefeller Uni- versity Archives for the opportunity to reread some of Rebecca Lancefield's correspondence and for access to the annual reports to the Board of Scientific Directors of the Rockefeller Institute, which were helpful in piecing together the early history of her work.

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242 BIOGRAPHICAL MEMOIRS SELECTED BIBLIOGRAPHY 1919 With O. T. Avery and A. R. Dochez. Studies on the biology of streptococcus. I. Antigenic relationship between strains of streptococcus haemolyticus. I. Exp. Med., 30:179-213. 1921 With C. W. Metz. Non-disjunction and the chromosome relation- ship of Drosophila willistoni. Proc. Natl. Acad. Sci. USA, 7:225 29. 1922 With C. W. Metz. The sex-linked group of mutant characters in Drosophila willastoni. Am. Nat., 36:211 - 41. 1924 . Antigenic relationships of the nucleo-proteins from the Gram- positive cocci. Proc. Soc. Exp. Biol. Med., 22: 109-11. 1925 The immunological relationships of streptococcus viridans and certain of its chemical fractions. I. Serological reactions ob- tained with antibacterial sera. I. Exp. Med., 42:377-95. The immunological relationships of streptococcus viridans and certain of its chemical fractions. II. Serological reactions ob- tained with antinucleoprotein sera. i. Exp. Med., 42:397-412. 1928 The antigenic complex of Streptococcus haemolyticus. I. Demonstra- tion of a type-specific substance in extracts of Streptococcus hae- molyticus. J. Exp. Med., 47:91-103; II. Chemical and immuno- logical properties of the protein fractions, 469-80; III. Chemical and immunological properties of the species-specific substance, 481-91; IV. Anaphylaxis with two non-type specific fractions, 843-55; V. Anaphylaxis with the type-specific sub- stance, 857-75. With E. W. Todd. Variants of hemolytic streptococci; their relation

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REBECCA CRAIGHILL LANCEFIELD 243 to type-specific substance, virulence, and toxin. }. Exp. Med., 48:751-67. With E. W. Todd. Antigenic differences between matt hemolytic streptococci and their glossy variants. I. Exp. Med., 48:769-90. 1933 A serological differentiation of human and other groups of he- molytic streptococci. I. Exp. Med., 57:57 1-95. 1934 A serological differentiation of specific types of bovine hemolytic streptococci. I. Exp. Med., 59:441-58. Loss of the properties of hemolysin and pigment formation with- out change in immunological specificity in a strain of Streptococ- cus haemolyticus. ]. Exp. Med., 59:459-69. 1935 With K. Goodner and H. F. Swift. The serological classification of hemolytic streptococci in relation to epidemiological problems. Am. I. Med. Sci., 190:445-53. With R. Hare. The serological differentiation of pathogenic and nonpathogenic strains of hemolytic streptococci from partu- rient women. I. Exp. Med., 61:335-49. 1938 Two serological types of group B hemolytic streptococci with re- lated, but not identical, type-specific substances. J. Exp. Med., 67:25-40. A micro precipitin-technic for classifying hemolytic streptococci, and improved methods for producing antisera. Proc. Soc. Exp. Biol. Med., 38:473-78. 1939 With G. K. Hirst. Antigenic properties of the type-specific sub- stance derived from group A hemolytic streptococci. I. Exp. Med., 69:425-45.

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244 BIOGRAPHICAL MEMOIRS 1940 Type-specific antigens, M and T. of matt and glossy variants of group A hemolytic streptococci. J. Exp. Med., 71:521-37. The significance of M and T antigens in the cross reactions be- tween certain types of group A hemolytic streptococci. i. Exp. Med., 71:539-50. 1941 Specific relationship of cell composition to biological activity of he- molytic streptococci. Harvey Lect., 36:251-90. 1943 With H. F. Swift and A. T. Wilson. Typing group A hemolytic strep- tococci by M precipitin reactions in capillary pipettes. I. Exp. Med., 78:127-33. Studies on the antigenic composition of group A hemolytic strep- tococci. I. Effects of proteolytic enzymes on streptococcal cells. J. Exp. Med., 78:465-76. 1944 With W. A. Stewart. Studies on the antigenic composition of group A hemolytic streptococci. II. The occurrence of strains of a given type containing M but no T antigen. J. Exp. Med.,79:79- 88. With R. F. Watson. Studies on the antigenic composition of group A hemolytic streptococci. III. Types with serologically identical M but distinct T antigens: Types 10 and 12. J. Exp. Med., 79:89-98. With W. A. Stewart, A. T. Wilson, and H. F. Swift. Studies on the antigenic composition of group A hemolytic streptococci. IV. Related T but distinct M antigens in types 15, 17, 19, 30, and in types 4, 24, 26, 28, 29, 46. Identification by slide agglutina- tion. J. Exp. Med., 79:99 -114. 1946 With V. P. Dole. The properties of T antigens extracted from group A hemolytic streptococci. i. Exp. Med., 84:449-70`

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REBECCA CRAIGHILL LANCEFIELD 245 1952 With G. E. Perlmann. Preparation and properties of type-specific M antigen isolated from a group A, type 1, hemolytic strepto- coccus. I. Exp. Med., 96:72-82. With G. E. Perlmann. Preparation and properties of a protein (R antigen) occurring in streptococci of group A, type 28, and in certain streptococci of other serological groups. J. Exp. Med., 96:83-97. 1954 Cellular constituents of group A streptococci concerned in anti- genicity and virulence. In: Streptococcal Infections, ed. M. Mc- Carty, pp. 3-18. New York: Columbia University Press. 1955 With M. McCarty. Variation in the group specific carbohydrate of group A streptococci. I. Immunochemical studies on the car- bohydrates of various strains. I. Exp. Med., 102: 11-28. 1957 Differentiation of group A streptococci with a common R antigen into three serological types, with special reference to the bac- tericidal test. J. Exp. Med., 106:525-44. 1958 Occurrence of R antigen specific for group A type 3 streptococci. J. Exp. Med., 108:329-41. 1959 Persistence of type-specific antibodies in man following with group A streptococci. l. Exp. Med., 110:271-92. 1960 . ,~ . Infection With E. W. Hook and R. R. Wagner. An epizootic in Swiss mice caused by a group A streptococcus, newly designated type 50. Am. J. Hyg., 72:111-19.

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246 BIOGRAPHICAL MEMOIRS 1962 Current knowledge of type-specific M antigens of group A strep- tococci. I. Immunol., 89: 307-13. 1964 With E. H. Freimer. Type-specific polysaccharide antigens of group B streptococci. J. Hyg., 64:191-202. 1971 With I. Rotta, R. M. Krause, W. Everly, and H. Lackland. New approaches for the laboratory recognition of M types of group A streptococci. i. Exp. Med., 134: 1298-315. 1975 With M. McCarty and W. Everly. Multiple mouse-protective anti- bodies directed against group B streptococci. Special reference to antibodies effective against protein antigens. J. Exp. Med., 142: 165-79. 1977 With S. D. Elliott and M. McCarty. Teichoic acids of group D strep- tococci with special reference to strains from pig meningitis (Streptococcus suds). J. Exp. Med., 145:490-99. 1979 With J. Y. Tai and E. C. Gotschlich. Isolation of type-specific poly- saccharide antigen from group B type Ib streptococci. J. Exp. Med., 149:58-66.

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