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JOHN ROBERT RAPER October3? 1911-May21, 1974 BY KENNETH B. RAPER PRIOR TO HIS UNTIMELY DEATH in 1974, John R. Raper was recognized as the foremost investigator of sex- uality in the fungi. Beginning at the University of North Car- olina in Chapel Hill ant! continuing at Harvard University, he gained wicle recognition while still a graduate student for his imaginative researches on the hormonal control of sex- uality in species of Achlya, a common genus of aquatic fungi. Further pioneering studies followed while he was a National Research Fellow at the California Institute of Technology anct as a member of the botany staffat Inctiana University; dur- ing WorIcl War II he was recruited as a radiation biologist for the Manhattan Project in Chicago and Oak Ridge. At the University of Chicago after the war, he further refiner! and extendecl his observations on the induction and regulation of sexual interactions in aquatic fungi by diffusible hormones that are produced in a sequential and invariant pattern. Then as if sexuality in Achlya were too straightforward he turned his attention to the more complicated phenomena in the higher fungi ant! chose as the primary object for ex- perimentation a small woo(l-rotting basicliomycete, Schizo- phyllum commune. Following his return to Harvard in 1954, and for twenty years thereafter, John, his wife CarIene, and his students ant! associates were cledicatect to unclerstancTing 347

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348 BIOGRAPHICAL MEMOIRS and revealing the bewildering intricacies of this complex sys- tem. They were singularly successful in this task, which led eventually to Johns authoritative book, Genetics of Sexuality in High Fungi. John R. Raper, the eighth child and seventh son of Wil- liam Franklin and Julia Crouse Raper, was born on a farm near Welcome, North Carolina, on October 3, 1911. As the youngest in the family, he received special attention and con- sideration. But more was involved than his tender age: John was someone special, not just in our immediate household but in the community at large. He was handsome, he was smart, and he was talented. Whether he enjoyed performing before church and public school audiences, or whether he just accepted the role because it was expected of him, one cannot say. But perform he did, reciting in a singularly clear voice and singing with a tonal quality seldom heard in the rural community where we lived. John was a lover of good music, a taste he acquired quite early and one that sustained him throughout his life. He grew up in a Moravian community where brass choirs were as much a part of special church services as were the "love- feasts," featuring coffee and hot cross buns. Although the latter delighted all of us, the music must have held a special fascination for John, for he was playing the trumpet almost by the time he could hold a horn to his lips. His proficiency with the instrument increased steadily: by the time he was a student at the university in Chapel Hilland first trumpet in the newly organized North Carolina Symphony he was seriously considering music as a profession. Fortunately, he chose biology not that he would have been a poor musician, but fortunately because otherwise he could not have made the important discoveries and contributions that marked his . career in science.

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JOHN ROBERT RAPER 349 Concerning his chilc~hooct and adolescence, John has writ- ten in part: My father had suffered a massive heart attack before my birth and I never knew him as a robust man. The hard work on the farm and largely the direction of its operations were thus left to my older brothers, each of whom in turn went away to high school (there being none locally), then college, and soon thereafter developed careers elsewhere. In the matter of education and independent development there was every possible en- couragement from home save appreciable financial assistance. Born dur- ing the Civil War, my father was unable to secure the education and train- ing he desired, and feeling that he had been trapped by circumstances, did what he could to insure that his children obtain the education he had been denied. Mother's attitude was somewhat different. Having grown up on a highly productive farm, her greatest wish was that one of her sons would take over the farm and operate it efficiently. No one of the seven accepted the challenge. Most of our social activities revolved around the public school and three churches, Friedberg and Enterprise, Moravian, and NIt. Olivet, Methodist, among which the family membership oscillated. Services were attended by the whole family, occasionally at all three on the same Sunday! Our home was bone dry and dancing and card playing were not encour- aged. As the youngest member of this rather strict paternalistic family, I was always the object of much affection if not frivolity. My first four years of school were spent in a 3-room school with no pretensions beyond the elementary level. At this time, a new, consolidated school (for which my father was a leading organizer and donor of the building site) was opened, and my further education through high school was received there. Aside from two years in the fifth grade for reasons never learned secondary school was reasonably uneventful, and I grad- uated from high school in 2nd or 3rd place in a class of 11. I was not good in sports or in other extra-curricular activities, such as dramatics, for which some rudimentary opportunities were provided. I was reasonably profi- cient with the trumpet, but the school, with neither musical instruction nor performing groups, provided no outlet for musical expression. English grammar and literature were my preferred subjects, and French was en- joyable. Science instruction was very rudimentary and generated only mild interest.

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350 BIOGRAPHICAL MEMOIRS About this time, cows became very prominent in my life. With the successive defection of my older brothers, there was no longer the labor force necessary to continue intensive tobacco farming, and a herd of mis- cellaneous cows became a major source of farm revenue. All through high school, usually with the help of a hired man, the care, feeding and milking of the cows was a constant preoccupation. Otherwise, I read avidly and widely and a reasonably good library at Winston Salem, some 10 miles distant, was routinely visited about once a week. It was there also that I discovered serious music via the record department of a furniture store in which an uncle worked; all available cash for some years went into classical recordings. Poorly prepared, and under considerable stress, my first year at the University of North Carolina was quite difficult, and it was made more so by the necessity of working 30-40 hours per week. In the spring quarter, however, my first science course, Introductory Botany under Professor John N. Couch, kindled an intense interest such as I had not previously known. In the second year, more botany and introductory zoology sus- tained and heightened this interest, and the offer of a teaching assistant- ship in the Department of Botany for the following year was recognized as an opportunity to indulge more fully my newfound interest. Aside from a passing flurry of musical activity (see above), there has been no subse- quent significant distraction from the pursuit of scientific interests. Major influences throughout this period in determining the selection of a partic- ular field of specialization were the enthusiasm and dedication of three teachers with whom I early worked: John N. Couch and William C. Coker of the University of North Carolina and William H. Weston of Harvard. Kenneth Raper's influence was also considerable during my student days. ~ohn's first contribution to science (1936) was an attempt to determine the pattern of sexual interaction between self- sterile strains of Achlya bisexualis, a heterothallic water moicI, previously clescribecl by Coker. Approximately 500 new col- lections mostly from streams anct ponds near Chapel Hill- were examined; of these 500, 32 were self-sterile. Of the lat- ter, 27 were identifiect as A. bisexualis anc! could be ciassifiect as female All, male (7), anct hermaphroctitic-female (12~. The last were of special interest because in the presence of strong males they behaved as true females.

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JOHN ROBERT RAPER 351 In the autumn of 1936, John transferred to Harvard Uni- versity anti, as the recipient of an Austin Teaching Fellow- ship, resumed his researches on A. bisexualis under the guicl- ance of Professor Weston. About a year later he isolated from the Charles River several cultures of a new anct even more interesting Achlya that he describec! as A. ambisexualis. This specific name was chosen because some of the isolates pos- sessec! both male and female potential ancT could behave as either sex, clepencling on the stronger sexual character of a paired mate. For this ant] other reasons, increased attention was subsequently given to the latter species. It soon became clear that the mating process consisted of a number of welI- defined steps or stages, that these occurred in an orderly and invariant sequence, anct that each was associated with a vi- sually evident developmental change in the paired culture. Additionally, because the stages appeared reciprocally in the two plants (thalli), and with these generally separated by some distance, it was reasonable to surmise that the successive changes observed in one plant were induced by cliffusible substances, or hormones, produced by the other. Subsequent researches involving selected cultures and interspecific crosses amply confirn~ecl this supposition; they also providecl the first unequivocal proof of hormonal, or pheromonal (al- ternate term), control of sexual reproduction in lower plants. In fact, if one wished to demonstrate "courtship" in plants, no better example exists than that of the interacting male and female thalli of the water mold Achlya which John so carefully described and so beautifully illustratecl some forty years ago. The sexual process in Achlya was shown to proceed in this manner. When strong male and female strains were im- plantec! at some distance in an agar plate or when they were grown on halvecl hemp seed floating in water- the terminal areas of the male hyphae began to branch profusely as the

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352 BIOGRAPHICAL MEMOIRS thalli approached each other. (The branches Lantheridial] were quite thin, highly ramified, anct at this first stage ran- clomly ctistributect.) In the second stage, several hours later, the female hyphae nearest the activated male began to swell terminally or to produce short, lateral, club-shapecI branches that in a few hours enIargecl to form gIobose structures (oo- gonial initials). In the thirct stage the antheridial hyphae grew towarc! the oogonial initials and on reaching their surfaces became appressed against them. The fourth stage was again expressed by the antheri(lial hyphae. Soon after these ap- pliecl themselves against the oogonial wails, protoplasm ac- cumulatec! in their tips anc! transverse walls were laict down to clelimit the male gametangia, or antherictia. The fifth stage followocl soon thereafter and was marked by the appearance of crosswalIs that delimited the swollen termini of the female hyphae and their club-shaped branches. The spherical struc- tures thus formed were the oogonia, or female gametangia. Whereas elapsed time varied appreciably clepencling on cul- tural and environmental conditions, the entire process could be completecl in thirty to thirty-six hours on agar or in ap- preciably shorter periods when plants were cultivated in water. In either case the sequence was the same, and the in- tervals between stages were proportional. The reciprocating responses, and the markedly shorter time necessary for stage one to be expresse(1 in water, strongly indicated that the for- mation of the anthericlial branches was clepenclent upon some substances procluced by the vegetative female plant; this in turn suggested that the female initiated the entire sex- ual process. These suppositions were then confirmed in clif- ferent ways. In one experiment male and female plants were separated by permeable membranes, either in the form of tubes or as sheets laid under the agar for some distance anti then bent upwarc! at 90 degrees. When the plants arrived at positions

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JOHN ROBERT RAPER 353 on opposite sides of the membranes, the male plant began to produce antheridial hyphae; six to eight hours later, oogonial initials were seen to emerge on the female plant, and as these matured the antheridial hyphae grew toward them and spread outward on the membrane nearest the oogonial ini- tials. In another experiment, male and female plants were grown separately on halved hemp seeds in petri dishes. When these reached maturity, the water was drained off and fresh water was added and allowed to remain for twenty-four hours. Liquid from these vessels was then drawn through Seitz filters and added to plants of the opposite sex. Anther- idial hyphae appeared on the male plants at seven hours and were very abundant at twelve hours. The response of female plants was less rapid but nonetheless positive. A perfusion experiment was the most dramatic of all. It was accomplished with a series of four connected micro-aquaria through which water flowed at a constant rate. The experiment was de- scribed in this way: In the first cell were placed two vegetatively mature female plants of Achlya amb?sexualis; in the second, two male plants of that species; in the third, two females; and in the fourth, a single male. Beginning about 5-6 hours after the introduction of the plants, a few antheridial hyphae were formed on the male plants in cells 2 and 4, but no reaction was given by the female plants in cell 3. Accordingly this experiment was repeated, but two female plants of A. bisexuals were placed in the first cell, since, as previously found, the male of A. ambisexual?s reacts more strongly to this than to female plants of the same species; the plants in the remaining three cells were selected as be- fore. Approximately 3 hours after the introduction of the plants, the male in cell 2 was seen to be reacting strongly. Two hours later the male plant in cell 4 was reacting vigorously. Twelve to fourteen hours after the initia- tion of the female reaction in cell 2, oogonial initials began to appear scattered over the entire female mycelium in cell 3. Following the begin- ning of the male reaction, directional growth of antheridial hyphae in the vicinity of the siphon tip in cell 4 began to take place, and at the end of another day this directional growth was fairly pronounced.

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354 BIOGRAPHICAL MEMOIRS From these and other studies John concluclecT that four major hormones, alternately pro(lucect by the female anct male plants, were responsible for initiating anct regulating the sexual process in Achlya. These were clesignated hor- mones A, B. C, anct D; they were characterized as follows: hormone A, proclucec! by the mycelium of the female plant, incluces the formation of antheridial hyphae on the male; hormone B. producer! by the antherictial hyphae of the male plant, induces the formation of oogonial initials on the fe- male; hormone C, produced by the oogonial initials, attracts antheridial hyphae to themselves and promotes delimitation of antheridia; and hormone D, producecl by the antheridia, causes clelimitation of the oogonia and subsequent differen- tiation of oospheres. For technical reasons, fertilization and maturation of oospheres couIcT not be followed. Upon receiving his doctorate from Harvard, John was awarded a National Research Fellowship. He subsequently went to work with Professor A. I. Haagen-Smit at the Cali- fornia Institute of Technology, his avowed purpose being to isolate anal, if possible, chemically characterize hormone A. In this he was partially successful. Much was learned about the properties of hormone A: "a final fraction, weighing 0.0002 g and still impure, contained 37 percent of the initial hormone-A activity of 1,440 liters of filtrate from female plants and incluced antheridial hyphal formation when tested in a dilution of I/10,000,000,000,000." For several reasons- economic anc! otherwise the work could not be continued at that time, but a very small amount of hormone-A concen- trate was retained for future study. Not until a quarter of a century later was hormone A finally isolated and character- izect by Trevor McMorris and Alma Barksdale at the New York Botanical Garden. It was found to be a stero] and re- named antheri(lao1t, the first steroid hormone found in plants. It is of more than passing interest that in the course of their work they reexamined a concentrate of hormone A that John

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JOHN ROBERT RAPER 355 hac! sealed in a vial in 1943; upon assay, they found that little of its activity hac! been lost. More recently, McMorris and coworkers have resolvecT hormone B into three steroidal com- poun(ls, which have been designated oogoniol-l, oogoniol-2, and oogoniol-3. Work on Achlya continucct when John was at Incliana Uni- versity; it was resumed when he returned to Chicago after the war. There at the university important events transpired, and about one of these he tells this story: During my first year at the University of Chicago after the war years as a radiobiologist, work was continued on the hormonal action of hor- mone A and the physiology of antheridial induction in Achlya. For this work, there was available a pitifully small supply of hormone A of high purity and standardized activity, and this vial of standard was dear to my heart it being used only in critical experiments and then only in 0.01 ml portions. Imagine my horror upon returning from a lecture to find my assistant, a fair-haired, first-year graduate student (Carlene Allen), on her hands and knees in the middle of the laboratory sucking up this precious liquid with a tiny pipette. She had dropped the bottle, which had broken, and had intuitively gone about the rational business of recovering what she could of the hormonal solution with the equipment at hand. In a mixture of shock at the obvious carelessness on the one hand and my admiration of her initiative in making the best of a totally unnecessary and bad situ- ation on the other, I could only urge the completion of the task and enjoin her not to cry over spilt hormone. There was, of course, no possibility of precise comparison of the activity of the recovered hormone with the orig- inal, and it may well be that the quantitative aspects of the work with Achlya underwent a slight discontinuity as a result of the accident. Forgiveness, however, was apparently not too difficult. Perhaps my failure to erupt into the violent display of temper that had been suggested in earlier and far less serious situations convinced her that I might be human after all. In any event, a couple of years later we were married, but over the years I've come to appreciate the monicker of "Spilly" bestowed on her by her family at a very early age. Following their marriage, CarIene continued to work with John in the laboratory anct soon became a full partner in his

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356 BIOGRAPHICAL MEMOIRS researches. In the years after his cleath, she obtainer! her cloctorate from Harvarc! University and developed an inde- pendent research career centered upon the genetics of higher fungi. In the early 1950s, John's attention began to shift increas- ingly to a quite ctifferent area of experimental mycology: the analysis of tetrapolar sexuality in the Basidiomycetes. Hans Kniep, H. R. R. Buller, anc! others had outlinect the broact picture of the genetic control system and the developmental sequence from spore to spore. Adclitionally, Haig Papazian, one of John's stuclents, had expancled this work and discov- erect several unusual features, including the appearance of rare, new, mating types, presumably as a result of recombi- nation; the existence of hemi-compatible heterokaryons; anc! the frequent occurrence of morphological mutations in cer- tain of the heterokaryons. Intrigued by these discoveries and impelled by his deep interest in the sexuality of all fungi, John spent the remainder of his professional career probing every facet of the biology of the Basidiomycetes, particularly Schizophy/Zum commune. Among the problems he adctressect were the analysis of the genetic fine structure of the incom- patibility system, the biochemical mechanism of incompati- bility in the Basidiomycetes, the genetics of fruiting, the mu- tational dissection of the morphogenetic sequences of heterokaryosis, anti the physiological consequences of com- patible and incompatible mycelial interactions. His efforts and those of an ever-expancling group of students and as- sociates raised S. commune to prominence as the best- understood representative of the Basicliomycetes; his labo- ratory, then at Harvarc! University, became a mecca for re- search in experimental mycology of the higher fungi. Some highlights of the SchizophyIlum research conclucted there are briefly noted below. John clemonstratecl in great detail the immensity of the

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JOHN ROBERT RAPER 361 and his wife were especially fond of classical music, ant] he equipped his home with excellent facilities for playing re- corcis ant] tapes, of which he tract a large ancT varied collec- tion. He enjoyed good food, and he was adept in preparing special dishes that on occasion were served to house guests with obvious joy and pricle. Thanksgiving at the John Rapers' was a warm tradition well remembered by several generations of students who couIcin't make it home for the holiday. A bountiful dinner was always preceded by a walk around Tho- reau's Walden Ponct; it was followed by listening to music before an open fire. John was a man of many talents who enjoyed life thor- oughly anct who enriched the lives of all the people who knew him commonly serene, sometimes impulsive, occasionally quixotic, but never dull. In recognition of his outstand- ing accomplishments, John received many honors. He was awarded a Guggenheim fellowship and a Fulbright scholar- ship to carry on researches in Germany in 1960 anct 1961, and he received the Aware] of Merit of the Botanical Society of America in 1969. He served as vice-presiclent and presi- clent of the Mycological Society of America, and he was a fellow and secretary of the American Acaclemy of Arts and Sciences. (As the holder of that office, he signed the letter of felicitation sent by that Academy on the occasion of the cen- tennial of the National Academy of Sciences in 1963.) He was elected to the National Academy the following year. Active in university affairs, he was nearing the end of a four-year term as chairman of the Department of Biological Sciences of Harvard University at the time of his cleath. John died on May 2l, 1974, after a brief illness. Quite fittingly a memorial service was held in Harvarct's Memorial Chapel that consisted primarily of choral music by J. S. Bach, Montevercli, ant! Vittoria. He is survived by his wife and coworker, CarIene; his son

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362 BIOGRAPHICAL MEMOIRS Jonathan, a clevelopmental neurobiologist anct recipient of an appointment as research scientist at the Max Planck In- stitut fur Virusforschung in Tubingen, Germany; his claugh- ter Lincia CarIene, a professional quilt artist; and, by a pre- vious marriage, his son William, a high school teacher. THE WRITER WISHES to express his appreciation to his brother John for having deposited with the Academy comments and rem- iniscences concerning his childhood and early adult life; to his wife, Dr. Carlene Raper, for his portrait and for her counsel and suggestions; and to Dr. T. I. Leonard for summarizing {ohn's stud- ies of sexuality in the higher Basidiomycetes.

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JOHN ROBERT RAPER B I B LI OGRAPHY 1936 363 Heterothallism and sterility in Achlya and observations on the cy- tology of Achlya bisexualis. ]. Elisha Mitchell Sci. Soc., 52:274- 89. 1937 A method of freeing fungi from bacterial contamination. Science, 85:342. 1939 Role of hormones in the sexual reaction of heterothallic Achlyas. Science, 89:321-22. Sexual hormones in Achlya. I. Indicative evidence for a hormonal coordinating mechanism. Am. i. Bot., 26:639-50. 1940 Sexuality in Achlya ambisexualis. Mycologia, 32:710-27. Sexual hormones in Achlya. II. Distance reactions, conclusive evi- dence for a hormonal coordinating mechanism. Am. J. Bot., 27: 162-73. 1942 Sexual hormones in Achlya. III. Hormone A and the initial male reaction. Am. I. Bot., 29:159-66. With A. I. Haagen-Smit. Sexual hormones in Achlya. IV. Properties of hormone A of Achlya bisexualis. ]. Biol. Chem., 143:311-20. Sexual hormones in Achlya. V. Hormone A', a male-secreted aug- menter or activator of hormone A. Proc. Natl. Acad. Sci. USA, 28:509-16. 1947 Effects of total surface beta irradiation. Radiology, 49:314-24. 1950 Beta rays: Biological eEects. In: Medical Physics, vol. 2, ed. Otto Glasser, pp. 66-71. Chicago, Ill.: The Year Book Publishers.

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364 BIOGRAPHICAL MEMOIRS Sexual hormones in Achlya. VI. The hormones of the A-complex. Proc. Natl. Acad. Sci. USA, 36:524-33. Sexual hormones in Achlya. VII. The hormonal mechanism in homothallic species. Bot. Gaz., 112: 1-24. 1951 Sexual hormones in Achlya. Am. Sci., 39: 110-20. Chemical regulation of sexual processes in fungi. In: Plant Growth Substances, ed. F. Skoog, pp. 301-13. Madison: University of Wisconsin Press. With R. E. Zirkle and K. K. Barnes. Techniques of external irra- diation with beta rays. Natl. Nuclear Energy Ser. Div., IV- 22E: 1 - 41. With I. E. Wirth and K. K. Barnes. Gross effects of beta irradiation on restricted surface of rabbits. Natl. Nuclear Energy Ser. Div., IV-22E:42 - 61. With R. E. Zirkle and K. K. Barnes. Comparative lethal effects of external beta irradiation. Natl. Nuclear Energy Ser. Div., IV- 22E:62-76. With K. K. Barnes. Gross effects of total-surface beta irradiation. Natl. Nuclear Energy Ser. Div., IV-22E:77-109. With K. K. Barnes. Rate of recovery from total-surface beta irra- diation. Natl. Nuclear Energy Ser. Div., IV-22E: 110-20. With K. K. Barnes. Additivity of lethal effects of external beta and gamma irradiation (I). Natl. Nuclear Energy Ser. Div., IV- 22E: 121-29. With K. K. Barnes. Additivity of lethal effects of external beta and gamma irradiation (II). Natl. Nuclear Energy Ser. Div., IV- 22E: 130-36. With R. S. Snider. Histopathological effects of single doses of total- surface beta radiation on mice. Natl. Nuclear Energy Ser. Div., IV-22E: 152 - 78. With K. K. Barnes. Effects of external irradiation with beta rays on the peripheral blood of rabbits. Natl. Nuclear Energy Ser. Div., IV-22E: 179-84. With I. E. Wirth. Reactions of human skin to single doses of beta rays. Natl. Nuclear Energy Ser. Div., IV-22E:193-99. With P. S. Henshaw and R. S. Snider. Delayed effects of single

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JOHN ROBERT RAPER 365 exposures to external beta rays. Natl. Nuclear Energy Ser. Div., IV-22E:200 - 11. With P. S. Henshaw and R. S. Snider. Effects of periodic total- surface beta irradiation. Natl. Nuclear Energy Ser. Div., IV- 22E:212-26. 1952 Chemical regulation of sexual processes in the Thallophytes. Bot. Rev., 18:447-545. 1953 Tetrapolar sexuality. Q. Rev. Biol., 28:233-59. 1954 With J. P. San Antonio. Heterokaryotic mutagenesis in Hymmen- omycetes. I. Heterokaryosis in Schizophyllum commune. Am. I. Bot., 41:69-86. Life cycles, sexuality, and sexual mechanisms in the fungi. In: Sex in Microorganisms, pp. 42 - 81. Washington, D.C.: American As- sociation for the Advancement of Science. 1955 Some problems of specificity in the sexuality of plants. In: Biological Specificity and Growth, ed. E. G. Butler, pp. 119-40. Princeton: Princeton University Press. Heterokaryosis and sexuality in fungi. Trans. N.Y. Acad. Sci. (II), 17:627-35. 1956 With P. G. Miles and H. Lund. The identification of indigo as a pigment produced by a mutant culture of Schizophyllum com- mune. Arch. Biochem. Biophys., 62:1-5. With P. G. Miles. Recovery of the component strains from dikar- yotic mycelia. Mycologia, 48 :484-94. 1957 Hormones and sexuality in lower plants. Symp. Soc. Exp. Biol., 11: 143-65.

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366 BIOGRAPHICAL MEMOIRS 1958 With G. S. Krongelb and M. G. Baxter. The number and distri- bution of incompatibility factors in Schizophyllum. Am. Nat., 92:221-32. With M. G. Baxter and R. B. Middleton. The genetic structure of the incompatibility factors in Schizophyllum commune. Proc. Natl. Acad. Sci. USA, 44:889 - 900. With P. G. Miles. The genetics of Schizophyllum commune. Genetics, 43:530-46. With I. P. San Antonio and P. G. Miles. The expression of mutations in common-A heterokaryons of Schizophyllum commune. Z. Ver- erbungsl., 89:540-58. With G. S. Krongelb. Genetic and environmental aspects of fruit- ing in Schizophyllum commune Fr. Mycologia, 50:707-40. With P. I. Snider. Nuclear migration in the Basidiomycete Schizo- phyllum commune. Am. I. Bot., 45:538 - 46. 1959 Sexual versatility and evolutionary processes in the fungi. Mycol- ogia, 51:107-25. Schizophyllum umbrinum Berkeley in culture. Mycologia,51 :474-76. 1960 The control of sex in fungi. Am. I. Bot., 47:794-808. With M. G. Baxter and A. H. Ellingboe. The genetic structure of the incompatibility factors of Schizophyllum commune: The A-fac- tor. Proc. Natl. Acad. Sci. USA, 46:833-42. With Y. Parag. Genetic recombination in a common-B cross of Schi- zophyllum commune. Nature, 188:765 - 66. Tetrapolarity in Schizophyllumfasciatum. Mycologia, 52:334 - 36. 1961 With S. Dick. Origin of expressed mutations in Schizophyllum com- mune. Nature, 189:81-82. With K. Esser. Antigenic differences due to the incompatibility fac- tors in Schizophyllum commune. Z. Vererbungsl., 92:439-44. Incompabilitat bei den Basidiomyceten Schizophyllum commune. Ber. Dtsch. Bot. Ges., 74:326-28.

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JOHN ROBERT RAPER 367 Parasexual phenomena in Basidiomycetes. In: Recent Advances in Botany, pp. 379 - 83. Toronto: University of Toronto Press. 1962 With A. H. Ellingboe. Somatic recombination in Schizophyllum com- mune. Genetics, 47:85-98. With A. H. Ellingboe. The Butler phenomenon in Schizophyllum commune: Internuclear selection in compatible dikaryotic-homo- karyotic matings. Am. i. Bot., 49:545-49. With M. T. Oettinger. Anomalous segregation of incompatibility factors in Schizophyllum commune. Rev. Biol. (Lisbon), 3:205 - 21. 1963 Patterns of sexuality in fungi. Mycologia, 55:79-92. With E. A. Hyatt. Modified press for disruption of microorganisms. I. Bacteriol., 85:712-13. Device for isolation of spores. I. Bacteriol., 86:342-44. With G. N. Bistis. Heterothallism and sexuality in Ascobolus stercor- arius. Am. I. Bot., 50:880-91. 1964 With K. Esser. The fungi. In: The Cell, ed. i. Brachet and A. E. Mirsky, vol. 6, pp. 139-244. New York: Academic Press. With C. A. Raper. Mutations affecting heterokaryosis in Schizophyl- lum commune. Am. I. Bot., 51 :503 - 12. 1965 With D. H. Boyd and C. A. Raper. Primary and secondary muta- tions at the incompatibility loci in Schizophyllum. Proc. Natl. Acad. Sci. USA, 53:1324-32. With P. I. Snider. Nuclear ratios and complementation in common- A heterokaryons of Schizophyllum commune. Am. J. Bot., 52:547- 52. introduction. In: Incompatibility in Fungi, ed. K. Esser and I. R. Raper, pp. 1 - 6. Berlin: Springer-Verlag. With I. T. Mullins. Heterothallism in biflagellate aquatic fungi:.Pre- liminary genetic analysis. Science, 150: 1174-75.

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368 BIOGRAPHICAL MEMOIRS 1966 Life cycles, basic patterns of sexuality and sexual mechanisms. In: The Fungi (II>, ed. G. C. Ainsworth and A. S. Sussman, pp.473- 511. New York: Academic Press. With Y. Koltin. Schizophyllum commune: New mutations in the B in- compatibility factor. Science, 154:510-11. Genetics of Sexuality in Higher Fungi. New York: Ronald Press. With C. A. Raper. Mutations modifying sexual morphogenesis in Schizophyllum. Genetics, 54: 1151-68. 1967 The role of specific secretions in the induction and development of sexual organs and in the determination of sexual affinity. In: Handbuch der ppanzen Physiologie, ed. H. F. Linskins, vol. 18, pp. 214-34. Heidelberg, Berlin, and New York: Springer-Verlag. With Y. Koltin and G. Simchen. Genetic structure of the incom- patibility factors of Schizophyllum: The B factor. Proc. Natl. Acad. Sci. USA, 57:55-62. With Y. Koltin. The genetic structure of the incompatibility factors of Schizophyllum commune: Three functionally distinct classes of B factors. Proc. Natl. Acad. Sci. USA, 58:1220-26. With Y. Koltin. The genetic structure of the incompatibility factors of Schizophyllum commune: The resolution of class III factors. Mol. Gen. Genet., 100:275-82. 1968 With M. Raudaskowski. Secondary mutations at the B,`3 locus of Schi- zophyllum. Heredity, 23: 109-17. . . With c. A. Kaper. Genetic regulation of sexual morphogenesis in Schizophyllum commune. ],. Elisha Mitchell Sci. Soc., 84:267-73. On the evolution of fungi. In: The Fungi (IIIJ, ed. G. C. Ainsworth and A. S. Sussman, pp. 677-93. New York: Academic Press. With Y. Koltin. Dikaryosis: Genetic determination in Schizophyllum. Science, 160:85-86. Steroid sexual hormones in a water mould. Proc. Sect. Sci. Isr. Acad. Sci. Humanit., 11: 1-8.

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JOHN ROBERT RAPER 1969 369 With C.-S. Wang. Protein specificity and sexual morphogenesis in Schizophyllum commune. ]. Bacteriol., 99:291-97. With T. I. Leonard. Schizophyllum commune: Gene controlling hap- loid fruiting. Science, 165: 190. 1970 Chemical ecology among lower plants. In: Perspectives in Chemical Ecology, ed. E. Sondheimer, pp. 21-43. New York: Academic Press. With A. S. Flexer. The road to diploidy with emphasis on a detour. In: Organization and Control in Prokaryotic and Eukaryotic Cells, ed. H. P. Charles and B. C. I. G. Knight, pp. 401-32. New York: Cambridge University Press. With I. H. Perkins. Morphogenesis in Schizophyllum commune. III. A mutation that blocks initiation of fruiting. Mol. Gen. Genet., 106: 151-54. With C.-S. Wang. Isozyme patterns and sexual morphogenesis in Schizophyllum. Proc. Natl. Acad. Sci. USA, 66:882-89. 1971 Growth and reproduction of fungi. In: Plant Physiology, A Treatise, ed. F. C. Steward, vol. 6, pp. 167-230. New York and London: Academic Press. With A. S. Flexer. Mating systems and evolution of Basidiomycetes. In: Evolution in the Higher Basidiomycetes, ed. R. H. Petersen, pp. 149 - 67. Knoxville: University of Tennessee. With R. M. Hoffman. Genetic restriction of energy conservation in Schizophyllum. Science, 171.418-19. 1972 With R. M. Hoffman. Lowered respiratory response to adenosine diphosphate of mitochondria isolated from a mutant-B strain of Schizophyllum commune. ]. Bacteriol., 110: 780-81. With C. A. Raper. Life cycle and prospects for interstrain breeding of Agaricus bisporus. Mushroom Sci., 8: 1-9. With C. A. Raper and R. E. Miller. 1972. Genetic analysis of the life cycle of Agaricus bisporus. Mycologia, 64:1088-117.

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370 BIOGRAPHICAL MEMOIRS 1973 With C. A. Raper. Incompatibility factors: Regulatory genes for sexual morphogenesis in higher fungi. In: Basic Mechanisms in Plant Morphogenesis, pp. 19-39. Brookhaven Symposia in Biol- ogy no. 25. Upton, N.Y.: Brookhaven National Laboratory. With C. A. Raper. Mutational analysis of a regulatory gene for morphogenesis in Schizophyllum. Proc. Natl. Acad. Sci. USA, 70: 1427-31. 1974 With A. S. Flexer. Heterothallism in Basidiomycetes. In: Mycology Guidebook, ed. R. B. Stevens, pp. 524-39. Seattle: University of Washington Press. With R. M. Hoffman. Schizophyllum. In: Handbook of Genetics, vol. 1, ed. R. C. King, pp. 597-626. New York: Plenum Press. With R. M. Hoffman. Genetic impairment of energy conservation in development of Schizophyllum: Efficient mitochondria in energy-starved cells. i. Gen. Microbiol., 82:67-75. With R. C. Ullrich. Number and distribution of bipolar incompat- ibility factors in Sistotrema brinkmannii. Am. Nat., 108:507 - 18. 1975 With R. C. Ullrich. Primary homothallism- relation to hetero- thallism in the regulation of sexual morphogenesis in Sistotrema. Genetics, 80:311-21.

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