John Robert Raper

John Robert Raper deposited specimens in NCU

(3 October 1911 – 21 May 1974)

The University of North Carolina at Chapel Hill Herbarium (NCU) has cataloged about thirty vascular plant specimens collected by John Robert Raper.  His most frequent collecting locations included Chapel Hill and the Great Smoky Mountains National Park, and orchids seem to have been his particular interest.  Although John Raper was a mycologist, we have found few fungal specimens collected by him in our collection.  Several specimens have simply “Raper” as the collector, and as both John and his brother, Kenneth Bryan Raper, attended UNC-CH and deposited specimens in NCU, it is difficult to attribute those collections accurately.  NCU’s mycological collection is cataloged and available to the public at and our vascular plants are cataloged at .

Other herbaria curating  John Raper’s fungal specimen include the New York Botanical Garden (NY; most collected in the Boston area), the US National Fungus Collection (BPI; most collected from southern Florida), and the University of Tennessee (TENN; most collected in eastern Tennessee, especially the Great Smoky Mountains National Park).

John Robert Raper was born in Welcome, Davidson County, North Carolina on 3 October 1911 to Julia Salina (Crouse) Raper and William Franklin Rapera farmer.  He was the youngest of eight children, four of whom attended the University of North Carolina at Chapel Hill — Arthur F. (class of 1924, Ph.D. 1931), Howard D. (class of 1927), and Kenneth B. (class of 1929).

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 encouragement save appreciable financial assistance. Born during the Civil War, my father was unable to secure the education and training 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 [sons] accepted the challenge.1

John Raper entered the University of North Carolina at Chapel Hill in 1929 and was known as “Red” due to his hair color.2  According to his brother, Kenneth, “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 Hill – and 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.” 1

Poorly [academically] prepared, and under considerable stress, my first year at the University of North Carolina [at Chapel Hill] was quite difficulty, and 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 sustained and heightened this interest, and the offer of a teaching assistantship in the Department of Botany for the following year was recognized as an opportunity to indulge more fully my newfound interest. 1

John Raper earned an A.B. in Botany in 1933.  Classmate Dudley Williams wrote of Raper, “He was one of the heroes of the depression era – working his way through [undergraduate school] at the library and cringing when not allowed to take his finals because his tuition was not fully paid even by then.”3

He completed an M.A. at UNC-Chapel Hill under the direction of Drs. William Chambers Coker and John Nathaniel Couch in 1936 with his thesis, “Heterothallism and sterility in Achlya and observations on the cytology of Achlya bisexualis,” portions of which were published in the Journal of the Elisha Mitchell Scientific Society (52:  274-289.  1936.)  As part of his thesis work, John Raper made about 500 new collections of Achlya bisexualis, a water mold described by William Chambers Coker & John Nathaniel Couch, from ponds and streams near Chapel Hill.1

In 1936 John Raper married fellow graduate student Ruth Scholz (M.A. University of North Carolina at Chapel Hill, 1934) , and together they had one son, William. Their marriage lasted until 1948. 1, 2

Raper moved to Cambridge Massachusetts and completed another M.A. as well as a Ph.D. (1939) at Harvard University under the direction of Professor William H. Weston, continuing his studies on Achlya.  He was a post-doctoral fellow at the California Institute of Technology in the laboratory of Professor A. J. Haagen-Smit.  1

From 1943-1946, he “got mixed up with the atom-bomb works at Chicago and Oak Ridge,” studying the biological effects of radiation.2  “Investigations for the project centered on the effects of irradiating laboratory animals with beta and gamma rays, such tests commonly representing total body exposure to beta irradiation…it was observed that when different animals were subjected to sublethal doses of total surface beta irradiation, some species (rats and mice) showed a significantly higher incidence of tumor formation than did others (rabbits and guinea pigs)…Out of it also came a man chastened by the experience and frightened by the implications of what he had learned.”1

During my first year at the University of Chicago after the war years as a radiobiologist, work was continued on [hormones] 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 floor 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… 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 situation on the other, I could only urge the completion of the task and enjoin her not to cry over spilt hormone… Perhaps my failure to erupt into the violent display of temper… convinced her that I might be human after all.  In any event, a couple of years later [1949] we were married, but over the years I’ve come to appreciate the monicker [sic] of “Spilly” bestowed on her by her family at a very early age.1

Raper continued to teach and do research at the University of Chicago, then accepted a faculty position in the Biology Department of Harvard University in 1954.  John Raper was elected to the National Academy of Sciences in 1964.  In 1974 he was Chair of the Biology Department at Harvard University.1

John Robert Raper died at age 62 in Cambridge, Massachusetts, survived by his wife (and research partner), Carlene Marie, their two children, Jonathan and Linda Carlene, and by William, his son from his first marriage.1


Raper, John R. (1936)  Heterothallism and sterility in Achlya and observations on the cytology of Achlya bisexualis.  J. Elisha Mitchell Sci. Soc. 52:  274-289.
— (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:  321-322.
— (1939)  Sexual hormones in Achlya.  I.  Indicative evidence for a hormonal coordinating mechanism.  Am. J. Bot. 26:  639-650.
— (1940)  Sexuality in Achlya ambisexualisMycologia, 32:710-727.
— (1940)  Sexual hormones in Achlya. II. Distance reactions, conclusive evidence for a hormonal coordinating mechanism.  Am. J. Bot. 27:  162-173.
— (1942)  Sexual hormones in Achlya. III. Hormone A and the initial male reaction. Am. J. Bot., 29:  159-166.
Raper, John R. and A. J. Haagen-Smit.  (1942)  . Sexual hormones in Achlya. IV. Properties of hormone A of Achlya bisexualis. J. Biol. Chem., 143:311-312.
Raper, John R. (1942)  Sexual hormones in Achlya. V. Hormone A’, a male-secreted augmenter or activator of hormone A.  Proc. Nat. Acad. Sci. USA 28:  509-516.
— (1947)  Effects of total surface beta irradiation. Radiology, 49:314-324.
— (1950)  Beta rays: Biological effects. In: Medical Physics, vol. 2, ed. Otto Glasser, pp. 66-71. Chicago, 111.: The Year Book Publishers.
— (1950)  Sexual hormones in Achlya. VI. The hormones of the A-complex.  Proc. Nat. Acad. Sci. USA 36:  524-533.
— (1950)  Sexual hormones in Achlya. VII. The hormonal mechanism in homothallic species. Bot. Gaz., 112:1-24.
— (1951)  Sexual hormones in Achlya. Am. Sci., 39:110-112.
— (1951)  Chemical regulation of sexual processes in fungi. In: Plant Growth Substances, ed. F. Skoog, pp. 301—13. Madison: University of Wisconsin Press.
Raper, John R., R. E. Zirkle and K. K. Barnes (1951)  Techniques of external irradiation with beta rays.  Nat. Nuclear Energy Ser. Div, IV 22E:  1-41.
Raper, John R., J. E. Wirth and K. K. Barnes (1951)  Gross effects of beta irradiation on restricted surface of rabbits. Natl. Nuclear Energy Ser. Div. IV-22E:42-61.
Raper, John R., R.E. Zirkle and K. K. Barnes (1951)  Comparative lethal effects of external beta irradiation. Natl. Nuclear Energy Ser. Div., IV-22E:62-76.
Raper, John R. and K.K. Barnes (1951)  Gross effects of total-surface beta irradiation.  Natl. Nuclear Energy Ser. Div., IV-22E:77-109.
— (1951)  Rate of recovery from total-surface beta irradiation.  Natl. Nuclear Energy Ser. Div., IV-22E: 110-20.
— (1951)  Additivity of lethal effects of external beta and gamma irradiation (I). Natl. Nuclear Energy Ser. Div., IV- 22E:121-29.
— (1951)  Additivity of lethal effects of external beta and gamma irradiation (II). Natl. Nuclear Energy Ser. Div., IV-22E: 130-36.
Raper, John R. and R.S. Snider (1951)  Histopathological effects of single doses of total-surface beta radiation on mice.  Nat. Nuclear Energy Ser. Div., IV-22E: 152-78.
Raper, John R. and K.K. Barnes (1951)  Effects of external irradiation with beta rays on the peripheral blood of rabbits. Natl. Nuclear Energy Ser. Div., IV-22E: 179-84.
Raper, John R. and J.E. Wirth (1951)  Reactions of human skin to single doses of beta rays. Natl. Nuclear Energy Ser. Div., IV-22E: 193-99.
Raper, John R., P.S. Henshaw and R.S. Snider (1951)  Delayed effects of single exposures to external beta rays. Natl. Nuclear Energy Ser. Div., IV-22E:200-211.
— (1951)  Effects of periodic total-surface beta irradiation. Natl. Nuclear Energy Ser. Div., IV-22E:212-26.
Raper, John R. (1952)  Chemical regulation of sexual processes in the Thallophytes. Bot. Rev., 18:447-545.
Raper, John R. (1953)  Tetrapolar sexuality. Q. Rev. Biol., 28:233-59.
Raper, John R. and J.P. San Antonio (1954)  Heterokaryotic mutagenesis in Hymmenomycetes I. Heterokaryosis in Schizophyllum commune. Am. J. Bot. 41:  69-86.
Raper, John R. (1954)  Life cycles, sexuality, and sexual mechanisms in the fungi. In: Sex in Microorganisms, pp. 42—81. Washington, D.C.: American Association 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.
— (1955)  Heterokaryosis and sexuality in fungi. Trans. N.Y. Acad. Sci. (II), 17:627-35.
Raper, John R., P. G. Miles and H. Lund (1956)  The identification of indigo as a pigment produced by a mutant culture of Schizophyllum commune.  Arch. BiochemBiophys., 62:1—5.
Raper, John R. and P. G. Miles (1956)  Recovery of the component strains from dikaryotic mycelia.  Mycologia  48:484—94.
Raper, John R. (1957)  Hormones and sexuality in lower plants. Symp. Soc. Exp. Biol., 3-65.
Raper, John R., G.S. Krongelb and M.G. Baxter (1958)  The number and distribution of incompatibility factors in Schizophyllum. Am. Nat 92:  221-232.
Raper, John R., M.G. Baxter and R. B. Middleton (1958)  The genetic structure of the incompatibility factors in Schizophyllum commune. Proc. Natl. Acad. Sci. USA 44:889-900.
Raper, John R. and P.G. Miles (1958)  The genetics of Schizophyllum commune. Genetics 43:530-46.
Raper, John R., J.P. San Antonio and P.G. Miles (1958)  The expression of mutations in common-A heterokaryons of Schizophyllum commune. Z. Vererbungsl., 89:540-58.
Raper, John R. and G.S. Krongelb (1958)  Genetic and environmental aspects of fruiting in Schizophyllum commune Fr. Mycologia, 50:707-740.
Raper, John R. and P.J. Snider (1958)  Nuclear migration in the Basidiomycete Schizophyllum commune. Am. J. Bot., 45:538—46.
Raper, John R. (1959)  Sexual versatility and evolutionary processes in the fungi. Mycologia  51:107-25.
— (1959)  Schizophyllum umbrinum Berkeley in culture. Mycologia, 51:474—7.
— (1960)  The control of sex in fungi. Am. J. Bot., 47:794-808
Raper, John R., M.G. Baxter and A. H. Ellingboe (1960)  The genetic structure of the incompatibility factors of Schizophyllum commune: The A-factor.  Proc. Natl. Acad. Sci. USA, 46:833-42.
Raper, John R. and Y. Parag (1960)  Genetic recombination in a common-beta cross of Schizophyllum commune. Nature, 188:765—6.
Raper, John R. (1960)  Tetrapolarity in Schizophyllum fasciatumMycologia, 52:334-36.
Raper, John R. and S. Dick (1961)  Origin of expressed mutations in Schizophyllum commune.  Nature 189:  81-82.
Raper, John R. and K. Esser (1961)  Antigenic differences due to the incompatibility factors in Schizophyllum commune. Z. Vererbungsl., 92:439-4.
Raper, John R. (1961)  Incompabilitat bei den Basidiomyceten Schizophyllum communeBertGes., 74:326-28.
— (1961)  Parasexual phenomena in Basidiomycetes. In: Recent Advances in Botany, pp. 379-83. Toronto: University of Toronto Press.
Raper, John R. and A. H. Ellingoe (1962)  Somatic recombination in Schizophyllum commune.  Genetics, 47:85-98.
— (1962)  The Buller phenomenon in Schizophyllum communeInternuclear selection in compatible dikaryotic-homokaryotic matings. Am. J. Bot., 49:545-49.
Raper, John R. and M.T. Oettinger (1962)  Anomalous segregation of incompatibility factors in Schizophyllum commune. Rev. Biol. (Lisbon), 3:205—221.
— (1963)  Patterns of sexuality in fungi. Mycologia, 55:79—92.
Raper, John R. and E.A. Hyatt.  (1963)  Modified press for disruption of microorganisms. J. Bacteriol., 85:712-13.
Raper, John R. (1963)  Device for isolation of spores. J. Bacteriol., 86:342—44.
Raper, John R. and G.N. Bastis (1963)  Heterothallism and sexuality in Ascobolus stercorarius.  Am. J. Bot. 50:  880-891.
Raper, John R. and K. Esser (1964)  The fungi. In: The Cell, ed. J. Brachet and A. E. Mirsky, vol. 6, pp. 139—244. New York: Academic Press.
Raper, John R. and C.A. Raper (1964)  Mutations affecting heterokaryosis in Schizophyllum commune.  Am. J. Bot., 51:503—12.
Raper, John R., D.H. Boyd and C.A. Raper (1965) Primary and secondary mutations at the incompatibility loci in Schizophyllum. Proc. Nat. Acad. Sci. USA, 53:1324-32.
Raper, John R. and P.J. Snider (1965)  Nuclear ratios and complementation in common-A heterokaryons of Schizophyllum commune. Am. J. Bot., 52:547=552.
Raper, John R. (1965)  Introduction. In: Incompatibility in Fungi, ed. K. Esser and J. R. Raper, pp. 1-6. Berlin: Springer-Verlag.
Raper, John R. and J.T. Mullins (1965)  Heterothallism in biflagellate aquatic fungi: Preliminary  analysis. Science, 150:1174-75.
Raper, John R. (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.
Raper, John R. and Y. Koltin (1966) Schizophyllum commune: New mutations in the B incompatibility factor.  Science  154: 510-511.
Raper, John R. (1966)  Genetics of Sexuality in Higher Fungi. New York: Ronald Press.
Raper, John R. and C.A. Raper (1966)  Mutations modifying sexual morphogenesis in Schizophyllum. Genetics, 54:1151—68.
Raper, John R. (1967)  The role of specific secretions in the induction and development of sexual organs and in the determination of sexual affinity. In: Handbuch der pflanzen Physiologie, ed. H. F. Linskins, vol. 18, pp. 214-34. Heidelberg, Berlin, and New York: Springer-Verlag.
Raper, John R., Y. Koltin and G. Simchen.  (1967)  Genetic structure of the incompatibility factors of Schizophyllum: The B factor. Proc. Nat. Acad. Sci. USA, 57:55-62.
Raper, John R. and Y. Koltin (1967)  The genetic structure of the incompatibility factors of Schizophyllum commune: Three functionally distinct classes of beta factors.  Proc. Natl. Acad. Sci. USA, 58:1220-26.
— (1967)  The genetic structure of the incompatibility factors of Schizophyllum commune: The resolution of class III factors.  Mol. Gen. Genet., 100:275-82.
Raper, John R. and M. Raudaskowski (1968)  Secondary mutations at the B beta locus of Schizophyllum.  Heredity 23:  109-117.
Raper, John R. and C. A. Raper (1968)  Genetic regulation of sexual morphogenesis in Schizophyllum commune. J. Elisha Mitchell Sci. Soc, 84:267—273.
Raper, John R. (1968) On the evolution of fungi. In: The Fungi (III), ed. G. C. Ainsworth and A. S. Sussman, pp. 677—93. New York: Academic Press.
Raper, John R. and Y. Koltin.  (1968)  Dikaryosis: Genetic determination in Schizophyllum.  Science, 160:85-86.
Raper, John R. (1968)  Steroid sexual hormones in a water mould. Proc. Sect. Sci. Isr. d. Sci. Humanit., 11:1—8.
Raper, John R. and C.-S. Wang (1969)  Protein specificity and sexual morphogenesis in Schizophyllum commune. J. Bacteriol., 99:291—9.
Raper, John R. and T. J. Leonard (1969)  Schizophyllum commune: Gene controlling haploid fruiting.  Science 165:  190.
Raper, John R. (1970) Chemical ecology among lower plants. In: Perspectives in Chemical Ecology, ed. E. Sondheimer, pp. 21—43. New York: Academic Press.
Raper, John R. and A. S. Flexer (1970)  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. J. G. Knight, pp. 401-32. New York:  Cambridge University Press.
Raper, John R. and J.H. Perkins (1970)  Morphogenesis in Schizophyllum commune. III.  A mutation that blocks initiation of fruiting.  Mol. Gen. Genet. 106:  151-154.
Raper, John R. and C.-S. Wang (1970)  Isozyme patterns and sexual morphogenesis in Schizophyllum. Proc. Natl. Acad. Sci. USA, 66:882-89.
Raper, John R. (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.
Raper, John R. and A.S. Flexer (1971)  Mating systems and evolution of Basidiomycetes.  In: Evolution in the Higher Basidiomycetes, ed. R. H. Petersen,, pp. 149-167.  Knoxville:  University of Tennessee.
Raper, John R. and R. M. Hoffman (1971)  Genetic restriction of energy conservation in Schizophyllum. Science, 171:418—19.
— (1972)  Lowered respiratory response to adenosine diphosphate of mitochondria isolated from a mutant-B strain of Schizophyllum commune. J. Bacteriol., 110:780-81.
Raper, John R. and C. A. Raper (1921)  Life cycle and prospects for interstrain breeding of Agaricus bisporus. Mushroom Sci., 8:1—9
Raper, John R., C. A. Raper and R. E. Miller (1972)  Genetic analysis of the life cycle of Agaricus bisporusMycologia, 64:1088—117.
Raper, John R. and C. A. Raper (1973)  Incompatibility factors: Regulatory genes for sexual morphogenesis in higher fungi. In: Basic Mechanisms in Plant Morphogenesis, pp. 19-39. Brookhaven Symposia in Biology no. 25. Upton, N.Y,: Brookhaven National Laboratory.
— (1973)  Mutational analysis of a regulatory gene for morphogenesis in Schizophyllum. Proc. Natl. Acad. Sci. USA  70:1427-31.
Raper, John R. and A. S. Flexer (1974)  Heterothallism in Basidiomycetes. In: Mycology Guidebook, ed. R. B. Stevens, pp. 524-39. Seattle: University of Washington Press.
Raper, John R. and R.M. Hoffman (1974)  Schizophyllum. In: Handbook of Genetics, vol. 1, ed. R. C. King, pp. 597-626. New York: Plenum Press.
— (1974)  Genetic impairment of energy conservation n development of Schizophyllum: Efficient mitochondria in energy-starved cells. J. Gen. Microbiol., 82:67—75.
Raper, John R. and R. C. Ullrich (1974)  Number and distribution of bipolar incompatibility factors in Sistotrema brinkmannii. Am. Nat., 108:507-18.
— (1975)  Primary homothallism—relation to heterothallism in the regulation of sexual morphogenesis in Sisotrema.  Genetics 90:  311-312.


  1. Raper, Kenneth B. (1987)  John Robert Raper (1911-1974).  Biographical Memoir.  National Academy of Sciences, Washington, D.C.
    2.  1958 John R. Raper Dope Sheet ’33 Yackety Yack Revisited.  University of North Carolina at Chapel Hill Alumni Records.
    3.  Williams, Dudley (1974)   Memorandum for Alumni Records.  University of North Carolina at Chapel Hill.

Information compiled by Carol Ann McCormick, September, 2013 from documents generously provided by Meredith Tozzer, University of North Carolina at Chapel Hill Alumni Records.  Updated December, 2019 by C. A. McCormick