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HARRY ALFRED BORTHWICK January 7~1898_May 21,1974 BY STERLING B. HENDRICKS HARRY BORTHWICK was born in Otsego, Minnesota, a small village about thirty miles from Minneapolis. His mother, Frances, was the aunt of Hubert Humphrey, who became Vice- President of the United States, and the sister of Harry B. Humphrey, a leading phytopathologist. It was the latter who influenced Harry to enter the School of Agriculture of the University of Minnesota in 1917. When his parents moved to San Jose, California, in 1919, Harry transferred to Stanford University, where he majored in botany. After receiving a 13A. degree in 1921, he continued in graduate school with research interest in plant morphology, leading in 1924 to an M.S. degree. Harry became a research assistant in the Division of Botany of the Agriculture School of the University of California at Davis in 1922, shortly before his marriage to Myrtis Hall. At Davis, Harry was first an assistant to and later a close associate of E. C. Robbins, a botany teacher and author of a well-known textbook in that field, who was engaged in research on crop plants. Harry continued working toward a doctorate at Stanford, devoting his attention mainly to the reproduction and development of both higher and lower plants. The position at Davis also required attention to basic and applied aspects of vegetable crop plants. These endeavors fashioned the pattern of his later scientific efforts. The main themes were: A wide 105

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106 BIOGRAPHICAL MEMOIRS knowledge of the functioning of plants of all types, with close attention to minute details of form and development and con- cern for ways to turn the more basic work to practical use in agriculture. Harry and Katherine Esau divided the work at Davis on the development of the vegetable crops of California" asparagus, beans, sugar beets, and carrots, among others. One of the first undertakings at Davis was the study of the development of lettuce from ovum fertilization to seed matu- rity. This classic work, often referred to over the course of the last fifty years, also served as a background for later work on seed germination. In the early 1930s, Harry worked on thresher injury to beans and on carrot seed development. The latter study was undertaken with L. T. Emsweller, who left Davis in 1935 to take charge of work in floriculture at the U.S. Depart- ment of Agriculture station in Beltsville, Maryland. Shortly after Emsweller went to Beltsville, the Congress passed what was known as the Bankhead-Jones Act, providing for research in depth in several aspects of agriculture. One of the most fundamental discoveries in biology had been made in 1920 in the Department of Agriculture by H. A. Allard and W. W. Garner. This was photoperiodism, or dependence of plants and animals on the length of day. In 1936 a decision was made to establish under the Bankhead-Jones Act a small group to look further into the nature of photoperiodism and its signifi- cance in agriculture. Because the photoperiodic response in plants partly regulates flowering, it was thought that progress might best result from attention to the morphological aspects involved in plants changing from vegetative to reproductive growth and to the underlying physiology. Emsweller recom- mended Harry Borthwick to undertake the work. Marion W. Parker, who was then teaching plant physiology at the nearby University of Maryland, was invited to join the effort. Borthwick and Parker foresaw that an understanding of photoperiodism would probably require growing plants under

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HARRY ALFRED BORTHWICK 107 closely controlled environmental conditions in order not to confound responses to light with temperature and other changes. Such controlled conditions, which are now commonplace, had not previously been obtained except in a very limited way at the Boyce-Thompson Institute. They decided that a.c. carbon arc lamps, developed for treatment of tuberculosis, were the only available type giving the required high light intensities. Growth rooms were constructed! with such lamps as light sources, but the plants grown in them were poorly developed compared to those from the field. Because the rare-earth loaded carbons gave radiation that was relatively more intense in the blue parts of the spectrum than in the red, Borthwick and Parker experimented with supplementary radiation from incan- descent filament lamps to enhance the red. The resulting growth of the selected plants was entirely satisfactory and of low vari- ability. The high requirement for red light anticipated what later became known as the high energy reaction for plant growth, the exact nature of which is still much debated. On H. A. Allard's recommendation, a soybean and cocklebur variety sensitive to light and requiring short days for flowering was selected. Barley par. Win tex was chosen as a long-day plant. The findings by others that the leaf was the receptor organ for the effective light and that transport of the stimulus to the terminal of the plant required phloem continuity were soon verified. A major discovery at this juncture was the effectiveness of short irradiations near the middle of Tong nights in prevent- ing flowering response. In the 1940s, the tendency of those interested in control of flowering was to attempt detection of florigen, a hypothetical hormone. Parker and Borthwick con- sidered this a poor approach, offering little opportunity for examination by experiment. They were more inclined to con- duct quantitative studies of the involvement of light in the flowering process. This would require measurement of action spectra. Wanting advice on methods and instrumentation for

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108 BIOGRAPHICAL MEMOIRS such an undertaking, they sought me out as one who might be interested. We soon began an informal cooperation that would endure for the next twenty-five years and unfold a whole new area of knowledge about plants. Action spectra for short-day plants were obtained in 1945. These indicated the presence of a blue pigment related to phycocyamin of the blue-green algae as the receptor for light. Long-day plants were found to have the same action spectra but with the opposite flowering response. With this finding of uni- versality in control of flowering, attention was turned to other responses to light. One of these was etiolation of plants growing in darkness. When examined, with cooperation of F. W. Went, who was visiting Beltsville, it, too, had the action spectrum for flowering control. Many students of plant growth at the time were little inclined to believe such findings, but some came to Beltsville on their own volition to observe. Harry's awareness, open mindedness, and courtesy encouraged an informality of approach and devotion to finding the meaning of things. The most basic finding, however, came from close at hand in an unexpected way. For more than a century, many seeds were known to require light for germination. Eben and Vivian Toole, who worked with seeds in a laboratory adjacent to the room where the action spectra were measured, proposed exam- ining the promotion of lettuce seed germination. The flowering- action spectrum again was found. But, most important, the potentiated action of red light, which required a day for dis- play, was found to be immediately reversible by a short expo- sure to far-red radiation. This indicated that the photoresponsive pigment was photoreversible and thus had at least two forms, only one of which was biologically effective. When attention was returned to flowering and etiolation, their potentiations were also seen to be photoreversible. The action spectra, more- over, were closely the same for all responses. Harry had throughout these years paid close attention to

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HARRY ALFRED BORTHWICK 109 the agricultural implications of the findings. These were chiefly applicable to the control of weeds and, through plant breeding, to many crops such as soybeans. With H. M. Cathey, who had succeeded Emsweller in floricultural work at Beltsville, Harry studied the control of flowering of poinsettia and chrysanthe- mum, which came into wide use in flower production. With A. A. Piringer and R. I. Downs, he studied the effect of light on woody perennials (trees and shrubs). These studies clearly emphasized the high energy action of far-red radiation, which had first been sensed in the development of lighting for growth rooms. The main response was control of the dormancy of buds by moderate periods of radiation. This is one of the many aspects of photoperiodism. The years were fruitful ones, with rewards wherever attention turned. The reversibility of photoresponsiveness was the keystone to progress about photoperiodism in the molecular sense. Through its use the product of molecular absorbancies of the receptive pigment and the quantum efficiency for conversion could be measured after the method used by Otto Warburg in work o cytochrome oxidase. In this way the pigment was established as deeply colored and present only in minute amounts in both albino and green plants long before it was seen. It, moreover, was probably a protein. Although the work on photoperiodism was steadfastly sup- ported for fifty years in the U.S. Department of Agriculture, the many pressures of limited funds militated against enlarging the effort when promising paths of investigation arose. Harry, foreseeing the possibility that great rewards might result from biochemical applications, used his limited funds to induce H. W. Siegelman to join the group. In a few years it became obvious that the receptive pigment could only be detected in viva by its photoreversibility. But, would reversibility work in vitro to serve for assay in an attempted isolation? Again, nearby cooperation was at hand. Karl Norris, an

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110 BIOGRAPHICAL MEMOIRS engineer working in the field of agricultural marketing, had developed spectroscopic methods for measuring the quality of apples. The method involved measuring minute absorbancies at two frequencies in optically dense media. With Norris and his associate, Warren Butler, the method, when applied re- versibly, immediately indicated the presence of the pigment in etiolated maize tissue, both before and after grinding. With the assay perfected, isolation of the pigment was finally achieved. With the pigment, by then known as phytochrome, identi- fied, Harry again looked toward finding some clue to its actions. He knowingly left the inviting molecular approaches to others of suitable interest and training. One unexpected clue to the method of action was quick display of response, rather than the long delayed ones that had previously been studied. This was demonstrated by J. C. Fondeville, who had come from France to work temporarily in Beltsville. He had been studying the diurnal movement of mimosa leaflets as an example of photo- periodism or biological rhythm, a favorite topic of P. Bert in Bordeaux more than a century earlier and of Charles Darwin and his son, Francis, in their Power of Movement in Plants. It was soon established that the leaflet closure upon placing mimosa plants in darkness could be prevented by changing the phytochrome from the far-red to the red-absorbing form. The response was displayed within ten minutes and was photore- versible. This established the role of phytochrome in the control of turgor with most likely action on a controlling membrane. Others soon turned their attention to this inviting approach and are now elaborating it in many promising directions. Harry by then (1969) had retired, fully aware that in this long and often lonely journey others now stood on the threshold of opportunity to look even deeper and perhaps in the end to find out more about differentiation in flowering, which is still elusive. In 1972, in his last publication, Borthwick looked back over

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HARRY ALFRED BORTHWICK 111 thirty-five years of research on photoperiodism in plants. A1- though fully aware of the measurement of time by plants, he had never accepted what others have called "the biological clock," or endogenous rhythm. These were meaningless words to himwords that clid more to obscure than to enlighten, and little to advance, experimentation. Also, he had rejected "flori- gen," the postulate flowering hormone, as more of the imagina- tion than of fact. Instead, the course he had charted into the core of photoperiodism consisted of a sound mixing of biologi- cal understanding and physical experimentation. Honors came to Harry Borthwick in his later years. He was elected to the National Academy of Sciences in 1961. He was President of the American Society of Plant Physiologists for a term and the recipient of its highest honor, the biennial Stephen Hales Award, and of a life membership. He received the Hob- litzelle Award for distinguished service to agriculture, the Joachem-Hafiz aware! from Switzerland, and the Distinguished Service Award from the U.S. Department of Agriculture. His greatest pleasure and deepest recognition, however, came from his many associates in research. They knew and honored him for his unselfish dedication to a central effort.

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112 KEY TO ABBREVIA TIONS BIOGRAPHICAL MEMOIRS BIBLIOGRAPHY Am. i. Bot. = American Journal of Botany Am. Nat. = American Naturalist Ann. Bot. = Annali di Botanica Annul Rev. Plant Physiol. = Annual Review of Plant Physiology Bot. Gaz. = Botanical Gazette Florists' Rev. = Florists' Review Plant Physiol. = Plant Physiology Proc. Am. Soc. Hortic. Sci. = Proceedings of the American Society for Hor- ticultural Science Proc. Int. Seed Test. Assoc. = Proceedings of the International Seed Test- . . . . 1ng Assoclatlon Proc. Natl. Acad. Sci. USA = Proceedings of the National Academy of Sci- ences of the United States of America Proc. Plant Propag. Soc. = Proceedings of the Plant Propagators Society U.S. Dep. Agric. Misc. Publ. = U.S. Department of Agriculture Miscella- neous Publication 1925 Factors influencing the rate of germination of the seed of Asparagus officinalis. California Agricultural Experiment Station Technical Paper, no. 18. With W. W. Robbins. Development of the seed of Asparagus offi- cinalis. Bot. Gaz., 80:426-38. 1928 With W. W. Robbins. Lettuce seed and its germination. Hilgardia, 3:275-305. 1931 Development of the macrogametophyte and embryo of Daucus carota. Bot. Gaz., 92:23-44. 1932 Thresher injury in baby lima beans. Journal of Agricultural Re- search, 44:503-10. Carrot seed germination. Proc. Am. Soc. Hortic. Sci., 28:310-14. 1933 With S. L. Emsweller. Carrot breeding experiments. Proc. Am. Soc. Hortic. Sci., 30:531-33.

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HARRY ALFRED BORTHWICK 1934 113 With R. Bainere Thresher and other mechanical injury to seed, beans of the lima type. California Agricultural Experimental Station Bulletin, no. 580. 1935 With S. L. Emsweller and P. C. Burrell. Studies on the inheritance of carotene in carrots. Proc. Am. Soc. Hortic. Sci., 33:508-11. 1936 Retarded germination in the seed of Hypericum perforatum. Bot. Gaz., 98:270-82. 1937 With S. L. Emsweller. Effects of short periods of low temperatures on flower production in stock. Proc. Am. Soc. Hortic. Sci., 35: 755-57. Histological and microchemical studies of the reactions of tomato plants to indolacetic acid. Bot. Gaz., 98:491-519. 1938 With M. W. Parker. Influence of photoperiods upon the differentia- tion of meristems and the blossoming of Biloxi soybeans. Bot. Gaz., 99:825-39. With M. W. Parker. Effectiveness of photoperiodic treatments of plants of different ages. Bot. Gaz., 100: 245~9. With M. W. Parker. Photoperiodic perception in Biloxi soybeans. Bot. Gaz., 100:274-87. 1939 With M. W. Parker. Effect of photoperiod on development and metabolism of the Biloxi soybean. Bot. Gaz., 100:651-89. With M. W. Parker. Effect of variation in temperature during the photoperiod induction upon initiation of flower primordia in Biloxi soybeans. Bo t. Gaz., 1 0 1:1 45-67. With M. W. Parker. Photoperiodic responses of several varieties of soybeans. Bot. Gaz., 101:341-65.

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114 BIOGRAPHICAL MEMOIRS 1940 With M. W. Parker. Floral initiation in Biloxi soybeans as influ- enced by age and position of leaf receiving photoperiodic treat- ment. Bot. Gaz., 101: 906-17. With M. W. Parker. Floral initiation in Biloxi soybeans as influ- enced by photosynthetic activity during the induction period. Bot. Gaz., 102: 256-68. 1941 With M. W. Parker and P. H. Heinze. Influence of localized low temperature on Biloxi soybeans during photoperiodic induction. Bot. Gaz., 102: 792-800. With M. W. Parker and P. H. Heinze. Effect of photoperiod and temperature on development of barley. Bot. Gaz., 103:326~1. 1942 With P. H. Heinze and M. W. :Parker. Floral initiation in Biloxi soybeans as influenced by grafting. Bot. Gaz., 103:518-30. With M. W. Parker. Day length and crop yields. U.S. Dep. Agric. Misc. Publ. no. 507. 1943 With M. W. Parker. Influence of temperature on photoperiodic reactions in leaf blades of Biloxi soybeans. Bot. Gaz., 104:612-19. With M. W. Parker and N. l. Scully. Effects of photoperiod and temperature on growth and development of Kok-saghyz. Bot. Gaz., 105:100-107. 1945 With M. W. Parker, S. B. Hendricks, and N. J. Scully. Action spec- trum for the photoperiodic control of floral initiation in Biloxi soybeans. Science, 102:152-55. With N. J. Scully and M. W. Parker. Interaction of nitrogen nutri- tion and photoperiod as expressed in bulbing and flowerstalk development on onion. Bot. Gaz., 107:52-61. With N. J. Scully and M. W. Parker. Relationship of photoperiod and nitrogen nutrition to initiation of flower primordia in soy- bean varieties. Bot. Gaz., 107:218-31.

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HARRY ALFRED BORTHWICK 115 1946 With M. W. Parker, S. B. Hendricks, and N. J. Scully. Action spec- tra for photoperiodic control of floral initiation in short-day plants. Bot. Gaz., 108:1-26. Photoperiodic response as a factor in choice of plants for testing soil deficiencies. Soil Science, 62:99-107. 1947 Day length and flowering. In: Yearbook of Agriculture, pp. 273-83. Washington, D.C.: U.S. Gout. Print. Off. 1948 With M. W. Parker and S. B. Hendricks. Wave length dependence and the nature of photoperiodism. Lotsya, 1: 71-78. With S. B. Hendricks and M. W. Parker. Action spectrum for photo- periodic control of floral initiation of a long-day plant, Wintex barley (Hordeum vulgare). Bot. Gaz., 110:103-18. 1949 With M. W. Parker. Photoperiodic responses of gladiolus. Gladiolus Magazine, 13: 26-31, 36-41. With M. W. Parker, S. B. Hendricks, and F. W. Went. Spectral sensitivities for leaf and stem growth of etiolated pea seedlings and their similarity to action spectra for photoperiodism. Am. J. Bot., 36:194-204. With M. W. Parker. Growth and composition of Biloxi soybeans grown in a controlled environment with radiation from different carbon-arc sources. Plant Physiol., 24: 345-58. 1950 With M. W. Parker. A modified circuit for slimline fluorescent lamps for plant chambers. Plant Physiol., 25:86-91. With M. W. Parker. Influence of light on plant growth. Annul Rev. Plant Physiol., 1:43-58. With M. W. Parker and S. B. Hendricks. Action spectrum for the photoperiodic control of floral initiation of the long-day plant Hyoscyamus niger. Bot. Gaz., 111: 242-52.

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116 BIOGRAPHICAL MEMOIRS With M. W. Parker and S. B. Hendricks. Recent developments in the control of flowering by photoperiod. Am. Nat., 84:117-34. With M. W. Parker and Laura E. Rappleye. Photoperiodic responses of poinsettia. Florists' Exchange. With M. W. Parker and N. W. Stuart. Tulips under light. Florists' Rev., 107:31-32. 1951 With M. W. Parker and Laura E. Rappleye. Photoperiodic responses of azaleas. Florists' Rev., 108:29-30. With S. B. Hendricks and M. W. Parker. Action spectrum for inhi- bition of stem growth in dark-grown seedlings of albino and non-albino barley (Hordeum vulgarej. Bot. Gaz., 1 1 3: 95-105. With M. W. Parker. Photoperiodic responses of soybean varieties. Soybean Digest, 11:26-30. 1952 With M. W. Parker, S. B. Hendricks, and C. E. Jenner. Photoperi- odic responses of plants and animals. Nature, 169:242. With M. W. Parker. Photoperiodism. In: Grolier Society Encyclo- pedia Yearbook ("The Story of Our Time"), pp. 310-12. New York: Grolier Society, Inc. With S. B. Hendricks, M. W. Parker, E. H. Toole, and V. K. Toole. A reversible photoreaction controlling seed germination. Proc. Natl. Acad. Sci. USA, 38:662-66. With S. B. Hendricks and M. W. Parker. The reaction controlling floral initiation. Proc. Natl. Acad. Sci. USA, 38:929-34. 1953 With M. W. Parker. Light in relation to flowering and vegetative development. Report of the 1 3th International Horticultural Congress of the Royal Horticultural Society, London, pp. 801- 10. London: Royal Horticultural Society. Photoperiodismthe dark secret; how nights and light affect plant growth. Electricity on the Farm Magazine, 26:11-13. With E. H. Toole, S. B. Hendricks, and V. K. Toole. Physiological studies of the effects of light and temperature on seed germina- tion. Proc. Int.. Seed Test. Assoc., 18:267-76.

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HARRY ALFRED BORTHWICK 1954 117 With S. :13. Hendricks, E. H. Toole, and V. K. Toole. Action of light on lettuce seed germination. Bot. Gaz., 1 15: 205-25. With G. M. Darrow. Fasciation in the strawberry; its inheritance and the relationship of photoperiodism. Journal of Heredity, 45:298-304. With C. E. Hagen and S. B. Hendricks. Oxygen consumption of lettuce seed in relation to photocontrol of germination. Bot. Gaz., 115:360-64. With N. i. Scully. Photoperiodic responses of hemp. Bot. Gaz., 116: 14-29. 1955 Some effects of light on plants. Annual Report of Vegetable Growers Association of America, pp. 71-77. Light and plant propagation. Proc. Plant Propag. Soc., pp. 63-70. With A. A. Piringer. Photoperiodic responses of coffee. Turrialba, 5:72-77. Daylength and crop production. 1955 Report Joint Committee on Grassland Farming, pp. 4-8. With E. H. Toole' V. K. Toole, and S. B. Hendricks. Photocontrol of Lepidium seed germination. Plant Physiol., 30:15-21. 1956 - With S. B. Hendricks and M. W. Parker. Photoperiodism. In: Radi- ation Biology III, ed. by A. Hollaender, pp. 479-517. New York: McGraw-Hill Book Co. With E. H. Toole, S. B. Hendricks, and V. K. Toole. Physiology of seed germination. Annul Rev. Plant Physiol., 7:299-324. With S. B. Hendricks and R. J. Downs. Pigment conversion in the formative responseses of plants to radiation. Proc. Natl. Acad. Sci. USA, 42:19-26. With R. l. Downs. Effect of photoperiod upon the vegetative growth of Weigela forida var. Variegata. Proc. Am. Soc. Hortic. Sci., -68:518-21. With A. A. Piringer, R. J. Downs, and S. B. Hendricks. A reversible photoreaction controlling photoperiodic response, seed germina- tion, and other phenomena. In: Congres international de bota-

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118 BIOGRAPHICAL MEMOIRS nique. Rapports et communications parvenus avant le congres, aux sections 11 et 12, pp. 321-23. (Proceedings of the 8th Inter- national Congress of Botany, Paris) Paris: Pierre Andrd. With S. B. Hendricks. Time dependencies in photoperiodism. In: Congres international de botanique. Rapports et communica- tions parvenus avant le congres, aux sections 11 et 12, pp. 323- 24. (Proceedings of the 8th International Congress of Botany, Paris) Paris: Pierre Andre. With S. B. Hendricks. Photoperiodism in plants. In: Proceedings of the 1st International Photobiological Congress, Amsterdam, pp. 23-25. Wageningen, Netherlands: H. Veenman & Zonen. With R. J. Downs. Effects of photoperiod on growth of trees. Bot. Gaz., 117:310-26. Light and some plant responses. Proc. Plant Propag. Soc., 5:63-72. Light studies and plant reaction. American Peony Society Bulletin, 141:11-14. With S. B. Hendricks. Photoresponsive growth. In: Aspects of Syn- thesis and Order in Growth, ed. by Dorothea Rudnick, pp. 149- 69. Princeton: Princeton Univ. Press. With E. H. Toole, V. K. Toole, and S. B. Hendricks. Interaction of temperature and light in germination of seeds. Plant Physiol., 30:473-78. With N. W. Stuart and A. A. Piringer. Photoperiodic responses of hydrangeas. In: Report of the 14th International Horticultural Congress, vol. 1, pp. 337-41. Held in the Netherlands. Wagenin- gen, Netherlands: H. Veenman & Zonen. 1957 With E. H. Toole, V. K. Toole, and S. B. Hendricks. Effect of tem- perature on germination of light-sensitive seeds. Proc. Int. Seed Test. Assoc., 22: 196-204. With V. K. Toole, E. H. Toole, and S. B. Hendricks. Physiology of seed dormancy. Proc. Int. Seed Test. Assoc., 22:205-19. With E. H. Toole and V. K. Toole. Growth and production of snap beans stored under favorable and unfavorable conditions. Proc. Int. Seed Test. Assoc., 22:418-22. With R. l. Downs and S. B. Hendricks. Photoreversible control of elongation of pinto beans and other plants under normal condi- tions of growth. Bot. Gaz., 118: 199-208.

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HARRY ALFRED BORTHWICK 1 l9 Light effects on tree growth and seed germination. Ohio Journal of Science, 57: 357-64. With H. M. Cathey. Photoreversibility of floral initiation in chry- santhemum. Bot. Gaz., 1 19: 71-76. 1958 With A. A. Piringer and R. T. Downs. Effect of photoperiod on Rawolfia. Am. J. Bot., 45: 323-26. With R. T. Downs and A. A. Piringer. Comparison of incandescent and fluorescent lamps for lengthening photoperiods. Proc. Am. Soc. Hortic. Sci., 71:568-78. 1959 With R. J. Downs, A. A. Piringer, and G. A. Wiebe. Effect of photo- period and kind of supplemental light on growth and reproduc- tion of several varieties of wheat and barley. Bot. Gaz., 120:170- 77. With S. B. Hendricks. Photocontrol of plant development by the simultaneous excitations of two interconvertible pigments. Proc. Natl. Acad. Sci. USA, 45:344-- 49. With S. B. Hendricks. Photocontrol of plant development by the simultaneous excitations of two interconvertible pigments. II. Theory and control of anthocyanin synthesis. Bot. Gaz., 120: 187-93. Photoperiodic control of flowering. In: Photoperiodism and Related Phenomena in Plants and Animals, ed. by Robert B. Withrow, pp. 275-87. AAAS Publication, no. 55. With S. B. Hendricks, E. H. Toole, and V. K. Toole. Photocontrol of plant development by the simultaneous excitations of two interconvertible pigments. III. Control of seed germination and axis elongation. Bot. Gaz., 121: 1-8. 1960 With S. Nakayama and S. B. Hendricks. Failure of photoreversible control of flowering in Pharbitis nil. Bot. Gaz., 121:237~3. With S. B. Hendricks. Photoperiodism in plants. Science, 132:1223- 28. With H. W. Johnson and R. C. Leffel. Effects of photoperiod and time of planting on rates of development of the soybean in vari- ous stages of the life cycle. Bot. Gaz., 122:77-95.

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120 BIOGRAPHICAL MEMOIRS 1961 With V. K. Toole, E. H. Toole, S. B. Hendricks, and A. G. Snow, fir. Responses of seeds of Pin us virginiana to light. Plant Physiol., 36:285-90. With S. Nakayama and S. B. Hendricks. Failure of reversibility of the photoreaction controlling plant growth. In: Progress in Pho- tobiology. Proceed ings of the 3d International Congress on Photobiology, pp. 394-98. Amsterdam, Netherlands: Elsevier Publishing Co. With R. l. Downs and A. A. Piringer. Light and plants. U.S. Dep. Agric. Misc. Publ. no. 879, p. 26. With H. M. Cathey and W. A. Bailey. Cyclic lightingto reduce cost of timing chrysanthemum flowering. Florists' Rev., 29:21-22, 72-75, 9~95. With A. A. Piringer. Effects of photoperiod and kind of supple- mental light on growth, flowering, and stem fasciating of celosia. Am. i. Bot., 48:588-92. With A. A. Piringer and R. I. Downs. Effects of photoperiod and kind of supplemental light on the growth of three species of citrus and Poncirus trifoliata. Proc. Am. Soc. Hortic. Sci., 77: 202-10. With S. B. Hendricks. Effects of radiation on growth and develop- ment. In: Handbuch der Ppanzenthysiologie, ed. by W. Ruh- land, vol. 16, pp. 299-330. Berlin: Springer-Verlag. With H. M. Cathey. Cyclic lighting for controlling flowering of chrysanthemums. Proc. Am. Soc. Hortic. Sci., 78:545-52. 1962 With V. K. Toole, E. H. Toole, and A. G. Snow, Jr. Responses of seeds of Pinus tueda and P. strobes to light. Plant Physiol., 37: 228-33. With H. M. Cathey. Role of phytochrome in control of flowering of chrysanthemum. Bot. Gaz., 123: 155-62. 1963 With A. A. Piringer and R. l. Downs. Photocontrol of growth and flowering of caryopteris. Am. I. Bot., 50: 86-90. With S. B. Hendricks. Control of plant growth by light. In: En-

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HARRY ALFRED BORTHWICK 121 vironmer~tal Control of Plant Growth, pp. 233-63. New York: Academic Press, Inc. With M. J. Kasperbauer and H. M. Cathey. Cyclic lighting for pro- motion of flowering of sweetclover, Melilotus alba Desr. Crop Science, 3:230-32. With M. i. Kasperbauer and S. B. Hendricks. Inhibition of flower- ing of Chenopodium rubrum by prolonged far-red radiation. Bot. Gaz., 124:444-51. Photochemical aspects of plant photoperiodicity. In: Photophysiol- ogy, ed. by E. Geise, vol. 1, pp. 305-31. New York: Academic Press, Inc. 1964 With M. l. Kasperbauer and S. B. Hendricks. Reversion of phyto- chrome 730 (Pfr) to P 660 (Pr) assayed by flowering in Cheno- pod ium rubrum. Bot. Gaz., 125: 75-80. Phytochrome action and its time displays. Am. Nat., 95:347-55. With A. L. Mancinelli. Photocontrol of germination and phyto- chrome reaction in dark-germinating seeds of Lactuca saliva L. Ann. Bot., 28: 9-24. With E. H. Toole, and V. K. Toole. Phytochrome control of Pau- lownia seed germination. Israel journal of Botany, 13: 122-33. 1965 With L. T. Evans and S. B. Hendricks. The role of light in suppress- ing hypocotyl elongation in lettuce and petunia. Planta (Ber- lin), 64:201-18. With B. G. Cumming and S. B. Hendricks. Rhythmic flowering re- sponses and phytochrome changes in a selection o Chenopodium rubrum. Canadian Journal of Botany, 43: 825-53. With S. B. Hendricks. The physiological functions of phytochrome. In: Biochemistry of Plant P'gments, ed. by T. W. Goodwin, pp. 405-36. New York: Academic Press, Inc. With L. T. Evans and S. B. Hendricks. Inflorescence initiation in Lolium temulentum L. VII. The spectral dependence of induc- tion. Australian Journal of Biological Sciences, 18:745-62. Light effects with particular reference to seed germination. Proc. Int. Seed Test. Assoc., 30:15-27. With H. C. Lane, H. M. Cathey, and L. T. Evans. The dependence

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122 BIOGRAPHICAL MEMOIRS of flowering in several long-day plants on the spectral composi- tion of light extending the photoperiod. Am. J. Bot., 52:1006-14. 1966 With A. L. Mancinelli and S. B. Hendricks. Phytochrome action in tomato-seed germination. Bot. Gaz.j 127: 1-5. With J. C. Fondeville and S. B. Hendricks. Leaflet movement of Mimosa pud ica L. indicative of phytochrome action. Planta (Berlin), 69:357-64. With i. C. Fondeville, M. l. Schneider, and S. B. Hendricks. Photo- control of Mimosa pudica L. leaf movement. Planta (Berlin), 75:228-38. 1967 With S. B. Hendricks. The function of phytochrome in regulation of plant growth. Proc. Natl. Acad. Sci. USA, 58:2125-30. With M. l. Schneider and S. B. Hendricks. Effects of radiation on flowering of Hyoscyamus niger. Am. J. Bot., 54:1241-49. 1968 With V. K. Toole. The photoreaction controlling seed germination in Eragrostis curoula. Plant and Cell Physiology, 9:125-36. With S. B. Hendricks and V. K. Toole. Opposing actions of light in seed germination of Poa pratensis and Amaranthus arenicola. Plant Physiol., 42: 2023-28. With V. K. Toole. Light responses of Eragostis seeds. Proc. Int. Seed Test. Assoc., 33: 2-16. 1969 With S. B. Hendricks, M. l. Schneider, R. B. Taylorson, and V. K. Toole. The high-energy reaction controlling plant responses and development. Proc. Natl. Acad. Sci. USA, 64:479-86. lD71 With V. K. Toole. Effect of light, temperature, and their interactions on germination of seeds of Kentucky bluegrass (Poa pratensis L). T. Am. Soc. Hortic. Sci., 96: 301~. 1972 History of phytochrome: In: Phytochrome, ed. by K. Mitrakos and W. Shropshire, ir., pp. 3~4. New York: Academic Press, Inc.

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