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JOHN GORDON TORREY February 22, 1921–January 7, 1993 BY LEWIS FELDMAN AND ALISON BERRY J cipled man, and a groundbreaking plant scientist. Allprin- forthright, honest, highly OHN GORDON TORREY WAS A who associated with John, or “JGT” as he was called by his graduate students, valued greatly his opinions and wise counsel. He was born in Philadelphia on February 22, 1921, the third of four children, and the second son of Edward and Elsie (Gordon) Torrey. He died January 7, 1993, in Greenfield, Massachusetts. Torrey graduated from Williams College (Williamstown, Massachusetts) in 1942 and shortly thereafter enlisted as an officer in the U.S. Army, serving for the duration of World War II in the Medical Adminis- trative Corps in both the United States and Europe. As noted by Kenneth Thimann, John Torrey’s Ph.D. disserta- tion supervisor at Harvard, Torrey “came to Harvard while still in uniform and was the first graduate student to come into the Biology Department after World War II.” In 1947 while still a graduate student, he was awarded a traveling fellowship allowing him to spend a year (1948-1949) at Cam- bridge University in the Botany School. In 1949 in England he married Noreen Lea-Wilson whom he had met during his earlier military service in the United Kingdom. Also in 1949 when he returned to Harvard, Torrey submitted a the- sis titled “Studies on the Physiology of Lateral Root Forma- 345

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346 BIOGRAPHICAL MEMOIRS tion and Root Growth,” for which he was awarded a Ph.D. in biology in 1950. Thereafter roots would be at the center of his long and highly influential research career. Torrey received assistant professor offers from Berkeley, Santa Barbara, and McGill. Thimann lobbied for McGill, but Torrey instead chose Berkeley and joined the Depart- ment of Botany in late 1949. There he became part of a young cadre of new faculty, including Leonard Machlis and Johannes Proskauer, who together rejuvenated the depart- ment and expanded its role from that of service to agricul- ture to include the broader discipline of plant science. While at Berkeley, Torrey began to elaborate his research on roots, focusing mainly on their growth and development. His dissertation adviser, Professor Thimann, had played a central role in characterizing the structure and function of the plant growth regulator, auxin (indole-3-acetic acid), and Torrey built on this knowledge to provide definitive evi- dence for the involvement of auxin in lateral root initiation and outgrowth. Based on his previous tissue culture experi- ence at Harvard with Professor Ralph Wetmore, Torrey in- corporated sterile culture techniques into his studies of roots, allowing him to manipulate root growth and development. From these efforts came seminal papers that characterized the controls of patterning in roots, with special emphasis on vascular patterning. Torrey was able to show that pat- terning was regulated by the root apical meristem. His re- search papers and the reviews arising from this work serve as the foundation for much of contemporary experimental root biology. While at Berkeley, Torrey taught, with Leonard Machlis, a plant physiology course that included a laboratory. As no suitable laboratory manual was then available, he and Machlis wrote Plants in Action; a Laboratory Manual of Plant Physi- ology, which not only became the standard text for plant

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347 JOHN GORDON TORREY physiology laboratory classes throughout the United States but also served as a valuable laboratory reference. In 1960 and now an associate professor at Berkeley, Torrey accepted an invitation to return to Harvard as a professor in the Department of Biology. There he expanded his stud- ies of roots and began to focus on an aspect of root devel- opment that had earlier attracted his attention, namely, the fixation of atmospheric nitrogen (nitrogen fixation) in rootborne structures known as nodules. Torrey believed that his earlier work on lateral root development could provide a context for further discoveries of this scientifically inter- esting and economically important process. His initial stud- ies of this phenomenon were with members of the pea fam- ily (legumes), in which nitrogen fixation occurs as a consequence of the association between the root and a bac- terium belonging to the genus Rhizobium. Torrey was in- terested in the beginning stages of nodule initiation, and focused much of his early research on understanding the reprogramming of the root cortical cells allowing them to develop into nodules. During the early years following his return to Harvard, Torrey authored (in 1967) Development in Flowering Plants, in which he pointed the way to the challenges ahead for plant developmental biologists. In the early 1970s Torrey moved his research activities to the Harvard Forest, in Petersham, Massachusetts, about 60 miles west of Cambridge. Coincident with this move was a redirection and refocusing of his research to include ni- trogen fixation in root nodules of perennial, nonlegume plants, with initial emphasis on the genus Comptonia (Sweet Fern). This plant grew abundantly in and around the Harvard Forest, inhabiting open woodlands and clearings. Torrey and his group showed that the organism causing nodule formation in Comptonia and responsible for nitrogen fixa- tion was neither a fungus, as was once believed, nor a mem-

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348 BIOGRAPHICAL MEMOIRS ber of the bacterial group called Rhizobia, which carries out nitrogen fixation in root nodules of the pea family. Instead, the microsymbiont in Comptonia root nodules and those of related plant hosts, belonged to the genus Frankia, bacteria in the Actinomycete group, which are evolution- arily distant from the Rhizobia. A significant accomplish- ment stemming from these efforts was the discovery of how to grow Frankia in culture, outside the root nodule envi- ronment. These landmark discoveries formed the basis of the most productive period in Torrey’s research career, in which more than 70 papers and one coedited volume (Ap- plications of Continuous and Steady-State Methods to Root Biology) were published between 1978 and 1991, detailing various aspects of the development of the association be- tween omptonia roots and Frankia. In 1965 while still in Cambridge, Torrey assumed a ma- jor administrative role as the fifth director of the Maria Moors Cabot Foundation for Botanical Research. Using fair, balanced judgment he was instrumental in directing foun- dation funds to support a much needed updating of botani- cal facilities at Harvard, including the expansion of the University Herbaria and the establishment of the Controlled Environment Facility at the Harvard Forest. He also en- sured that foundation funds were made available to junior faculty, and that the granting of these funds would involve a minimum of paperwork. Later, in 1984, he continued his administrative duties through his appointment as the Charles Bullard Professor of Forestry and director of the Harvard Forest. He main- tained these positions until 1990. As director he had great impact on activities at the forest, from initiating freshmen seminars (in those days a novelty at universities) to spear- heading a consortium of scientists from several institutions that eventually led to the awarding of a large National Sci-

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349 JOHN GORDON TORREY ence Foundation grant supporting the establishment of a Long Term Ecological Research site at the forest. The legacy of his efforts and his perspectives are today reflected in the expanded and varied ecological research activities at the forest. Although located in Petersham, Torrey continued to teach classes, supervise graduate students, and meet with visiting scholars in Cambridge. He also established an association with the University of Massachusetts in nearby Amherst, and there offered his mentoring skills and served on doctoral dissertation theses. He was a strong supporter of women in science and encouraged his female associates, both graduate students and visitors, to aim high. The legacy of this mentorship is today evident in the many successful academic careers of women who worked with John Torrey. “In a tough academic world,” one former female graduate advisee noted, “he was truly a hero.” John Torrey was a recipient of many awards and ap- pointments, including a Guggenheim Fellowship (1965-1966) and a Fulbright Senior Research Scholar Fellowship (1984). He was a member of the American Academy of Arts and Sciences, the Botanical Society of America, and the Society for Developmental Biology (president, 1963), among other organizations. He was elected to the National Academy of Sciences in 1981. Torrey was known to his many associates worldwide for his interests in etchings, particularly those from Scotland, England, and New England, from the late 19th and early 20th centuries. In his retirement he had intended to ex- plore “the interaction, interplay and influence of the group of British etchers on the Americans and vice versa.” When- ever he traveled he would reserve some time to visit local antique and art establishments. In these adventures he was

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350 BIOGRAPHICAL MEMOIRS usually, but not always, successful in acquiring a new etch- ing or watercolor. John Torrey was occasionally confused with the noted 19th-century American botanist of the same name, founder of the Torrey Botanical Club. Although there was no rela- tion, he was mildly bemused by the confusion occasioned by the coincidence of both their names and occupations. His wry sense of humor was not often seen but could be noted occasionally, such as when Torrey drove past a grave- yard and chuckled at the sign reading “One Way.” Reflecting on John Torrey, his Harvard colleagues rec- ollected that he was “outspoken and, in his controlled way, passionate about what he thought was right and what he thought was wrong.” On one occasion when a Harvard col- league off-handedly informed Torrey in his role as director of the Cabot Foundation that he (the colleague) would be spending funds differently from what was originally bud- geted, Torrey noted firmly, “No, I don’t think you should spend that money in a way that differed from your original intentions.” John and his wife, Noreen (Norah), were married for 43 years and had five daughters: Jennifer, Joanna, Susan, Sa- rah, and Carolyn. John Torrey was a man of high ethics both in his sci- ence and in the way he lived his life. He treated everyone equally, with respect, dignity, and honesty, and he expected no less from others. At his passing his daughter Joanna penned, “We lay him to full rest, sorry that he is not here with us, hat in hand, to point up into the trees, or down to the earth, sharing his faith in that which grows.”

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351 JOHN GORDON TORREY Some information used in preparing this remembrance was obtained from the Memorial Minute of the Harvard University Faculty of Arts and Sciences, appearing in the November 7, 1996, edition of the Harvard Gazette.

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352 BIOGRAPHICAL MEMOIRS SELECTED BIBLIOGRAPHY 1950 The induction of lateral roots by indoleacetic acid and root decapi- tation. Am. J. Bot. 37:257-264. 1951 Cambial formation in isolated pea roots following decapitation. Am. J. Bot. 38:596-604. 1957 Auxin control of vascular pattern formation in regenerating pea root meristems grown in vitro. Am. J. Bot. 44:859-870. 1958 Endogenous bud and root formation by isolated roots of convolvu- lus grown in vitro. Plant Physiol. 33:258-263. 1959 A chemical inhibitor of auxin-induced lateral root initiation in roots of Pisum. Physiol. Plantarum 12:873-887. 1972 On the initiation of organization in the root apex. In The Dynamics of Meristem Cell Populations, eds. M. W. Miller and C. C. Kuehnert, pp.1-10. New York: Plenum Press. 1975 With D. T. Clarkson, eds. The Development and Function of Roots. London: Academic Press. 1976 With L. J. Feldman. The isolation and culture in vitro of the quies- cent center of Zea mays. Am. J. Bot. 63:345-355.

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353 JOHN GORDON TORREY 1978 With D. Callaham. Determinate development of nodule roots in actinomycete-induced root nodules of Myrica gale. Canad. J. Bot. 56: 1357-1364. 1979 With W. Newcomb, D. Callaham, and R. L. Petersen. Morphogen- esis and fine structure of the actinomycetous endophyte of nitro- gen-fixing root nodules of Comptonia peregrina. Bot. Gaz. 140:S22- S34. 1980 With D. Baker and W. Newcomb. Characterization of an ineffective actinorhizal microsymbiont, Frankia sp. EuIl (Actinomycetales). Canad. J. Microbiol. 26:1072-1089. 1981 With J. D. Tjepkema, G. L. Turner, F. J. Bergersen, and A. H. Gibson. Dinitrogen fixation by cultures of Frankia sp. CpI1 demonstrated by 15N2 incorporation. Plant Physiol. 68:983-984. 1985 With D. J. Marvel, G. Kuldau, A. Hirsch, E. Richards, and F. M. Ausubel. Conservation of nodulation genes between Rhizobium meliloti and a slow-growing Rhizobium strain that nodulates a nonlegume host. Proc. Natl. Acad. Sci. 82:5841-5845. 1986 With M. F. Lopez and P. Young. A comparison of carbon source utilization for growth and nitrogenase activity in two Frankia iso- lates. Canad. J. Microbiol. 32: 353-358. With Z. Zhang and M. A. Murry. Culture conditions influencing growth and nitrogen fixation in Frankia sp. HFPCcI3 isolated from Casuarina. Plant Soil 91:3-15.

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354 BIOGRAPHICAL MEMOIRS 1987 With D. J. Marvel and F. M. Ausubel. Rhizobium symbiotic genes required for nodulation of legume and nonlegume hosts. Proc. Natl. Acad. Sci. 84:1319-1323. Endophyte sporulation in root nodules of actinorhizal plants. Physiol. Plantarum 70:279-288. 1989 With S. Racette. The isolation, culture and infectivity of a Frankia strain from Gymnostoma papuanum (Casuarinaceae). Plant Soil 118:165-170. With S. Racette. Root nodule initiation in Gymnostoma (Casuarinaceae) and Shepherdia (Elaeagnaceae) induced by Frankia strain HFPGp11. Canad. J. Bot. 67: 2873-2879. With L. J. Winship, eds. Applications of Continuous and Steady- State Methods to Root Biology. Dordrecht: Kluwer Academic. With S. S. Tzeany. Spore germination and the life cycle of Frankia in vitro. Canad. J. Microbiol. 35:801-806. 1990 With S. Burleigh. Effectiveness of different Frankia cell types as inocula for the actinorhizal plant Casuarina. Appl. Environ. Microbiol. 56:2565-2567. With S. R. Mansour and A. Dewedar. Isolation, culture, and behav- ior of Frankia strain HFPCg14 from root nodules of Casuarina glauca. Bot. Gaz. 151:490-496. With W. Newcomb, S. Jackson, and S. Racette. Ultrastructure of infected cells in the actinorhizal root nodules of Gymnostoma papuanum (Casuarinaceae) prepared by high pressure freezing and chemical fixation. Protoplasma 157:172-181. 1991 With S. R. Mansour. Frankia spores of strain HFPCg14 as inoculum for seedlings of Casuarina glauca. Canad. J. Bot. 69:1251-1256.

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355 JOHN GORDON TORREY