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KATHERINE ESAU April 3, 1 898-June 4, 1997 BY RAY F . EVE RT KATHERINE ESAU, WORLD renowned botanist, recipient of the National Mecial of Science, author of six textbooks, en c! teacher par excellence, cliec! Tune 4, 1997, at her home in Santa Barbara, California. She was ninety-nine years young. Her work on plant structure coverer! seven-plus clecacles en c! lee! to much of the current research on plant function. Throughout her career, Esau continues! research on phIo- em both in relation to the effects of phioem-limitec! viruses on plant structure en c! clevelopment en c! to the unique struc- ture of the sieve tube as a conduit for foocI. She clemon- stratec! an exceptional ability for attacking basic problems en c! she set new stanciarcis of excellence for the investiga- tion of anatomical problems in the plant sciences. Esau was born on April 3, IS9S, in the city of YekaterinosTav, now caller! Dnepropetrovsk, in the Ukraine. She liver! there until the enc! of 191S, when she en c! her family fleck to Germany cluring the Bolshevik Revolution. Her family was Mennonite, clescenciants of the German Mennonites that Catherine the Great invites! to Russia to promote agricul- ture on the Ukrainian steppes. Naturally suspicious of any- one from the outside, the Ukrainians ostracizer! the Men- nonites, who liver! in colonies, clevelopec! very successful 91

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92 BIOGRAPHICAL MEMOIRS farms, schoolec! their chiTciren, en c! practicer! their religion. To break clown this wall of distrust, the Ukrainian authori- ties asker! the Mennonites to sent! some of their chilciren to Russian schools. Esau's uncle en c! father were the first from their colony to stucly in Russian schools. Her Uncle Jacob stucliec! medicine. Her father {ohan trainee! as a mechani- cal engineer en c! became the city engineer of YekaterinosTav. Builcling the city's waterworks, a streetcar line, en c! large city buildings, including several schools, he worked very hare! to make the city more liveable. Consequently, he was well likes! by the citizenry, who electec! him mayor. An in- novation of his was to borrow money from France for the buiTcling projects. With the turmoil of the Bolshevik Revo- lution en c! being a city leacler, it was a only matter of time before the Bolsheviks market! him for execution. The revo- lutionary government remover! him from his post en c! kept him uncler constant surveillance. The Esau home was searcher! for banner! goocis, en c! valuable items were taken. With the advent of Woric! War I, the German Army acI- vancec! en c! succeeclec! in occupying the Ukraine. Most of the peaceful population welcomer! this turn of events, be- cause it saver! them from Bolshevik occupation en c! from invasion by the unorganized! bancis that were massacring people en c! destroying property. John Esau was reinstatec! as mayor. As the war wounc! clown, the German officers warner! that the Esau family wouIc! be in great cianger after the army left en c! acivisec! them to flee with them to Ger- many. This was heartbreaking for the oicler Esaus, because their roots were in Yekaterinoslav, where their chilciren Katherine en c! Paul en c! Mrs. Esau (the former Margrethe Toews) were born. The Esaus, along with many other people, followed the German advice. Paul was an administrator on a ship in the Black Sea and was unable to go with them on the journey, but he joined them soon after they arrived in

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KATHERINE ESAU 93 Berlin. They traveler! thirc! class in a train with wooden benches, together with the officers, the injured, en c! other refugees. The journey to Berlin laster! two weeks rather than the usual two clays because of various clifficulties en c! obstacles put in their way by the revolutionary government in the cities through which they passed. The clay after the Esaus left YekaterinosTav, posters appearec! in the city pro- cIaiming that the new city managers were looking for her father, whom they characterizec! as a member of the coun- terrevolutionary bourgeoisie en c! an enemy of the country. During the turbulent last years the family spent in Rus- sia, Esau's father gave money to a family frienc! to deposit in a Swiss bank in case the Esaus neeclec! to leave the coun- try in a hurry. The frienc! prover! worthy of their friencI- ship, en c! the money was there when the family reacher! Berlin. Later when the frienc! neeclec! money to clevelop a patent on an oil-well part, Esau's father stakes! him. That investment proviclec! the means for the Esaus to live com- fortably for the rest of their lives. When the Esaus fleck Russia, Katherine Esau hac! com pletec! her first year of study at the Golitsin Women's Agri- cultural College in Moscow. Fortunately, she hac! asker! for a transcript of her course work en c! gracles at the enc! of the term, en c! upon the family's arrival in Berlin, she registered in the Berlin Lanc~wirtschaftTiche HochschuTe (Agricultural College of Berlin), where she resumes! her studies, this time in German, not Russian. Her brother Paul also macle the transition en c! stucliec! chemistry. As part of her studies, she spent two semesters in Hohenheim near Stuttgart, where she enrollee! in various agricultural courses. After two more semesters in Berlin en c! a final examination, she receiver! the title "Landwirtschaftlehrerin." Following additional stud- ies, she passer! a Zusatzprufung in plant brawling given by the famous geneticist Erwin Bauer, who urger! her to re

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94 BIOGRAPHICAL MEMOIRS turn to Russia, saying that the country neeclec! her. Fortu- nately for the worIc! en c! the advancement of science, she clic! not hem! his acivice. From Berlin, Esau went to a large estate in northern Ger- many that houses! a moclel seecI-breecling station for wheat. She joiner! the workers there in the fielcis en c! barns cloing various chores. The son of the farm's owner was very smit- ten with Katherine, but he conic! not persuacle her to marry him. She returnee! to Berlin, where a teaching assistantship awaited, but by then the Esaus hac! cleciclec! to settle in the Uniter! States, en c! preparations were underway to clo so. The political situation in Germany was deteriorating en c! John Esau felt his family wouic! be safer in the Uniter! States. The Esaus left for America in micI-October of 1922. Brother Paul stayer! behinc! to finish his last year of studies in chem- istry. The Esaus crosser! the continent by train en c! the ocean by boat, like so many other immigrants, they enterer! the Uniter! States at Ellis Island. Their destination was ReecIley, California, where there was a large Mennonite community. Esau's father wantec! to buy a farm for her to manage, but she convincer! him that she neeclec! more working knowI- ecige of California agriculture en c! American-styTe manage- ment. At first, she worker! as a house cleaner en c! chilcicare worker in Fresno, California, all the while perfecting her English en c! learning American customs. When Esau felt comfortable with the language en c! Ameri- can practices, she took a job with Sloan Seec! Company in Oxnard, California. She later mover! to the Spreckles Sugar Company in the Salinas Valley, where she brat! strains of sugar beets for resistance to the virus causing curly-top clis- ease. At that time, she began to consicler continuing her education. It was serendipity that Professor Wilfred Robbins of the University of California, Davis, campus made a visit to the company, en c! Esau was asker! to show him her re

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KATHERINE ESAU 95 search project. She inquirer! about the possibility of stucly en c! later an invitation was extenclec! to her to clo graduate work atDavis. Esau arriver! in Davis in the fall of 1927, registering as a graduate student in the College of Agriculture for the 1928 spring semester. (The Davis campus clic! not aware! a Ph.D. at that time, so the degree was awarclec! from Berkeley.) She intenclec! to clevelop a sugar beet that was resistant to curly-top virus. However, that wouIc! have requires! releas- ing the beet leafl,opper into the university fielcis to infect the sugar beets. This was opposer! by other plant research- ers en c! growers, en c! she was toIc! that it was incompatible with other crop research going on at Davis. Accordingly, she changer! the direction of her research to the stucly of the transmission of curly-top virus en c! its effect on the sugar beet phioem, directing her research from applied to the more basic stucly of plant anatomy as it relates to the disease. Esau receiver! her Ph.D. in 1932 from Berkeley. She re- mainec! at Davis as an instructor, later becoming professor of botany. Esau left Davis in 1963, close to her official re- tirement ciate, to join her Tong-time research collaborator Vernon I. CheacIle at the University of California, Santa Barbara, where he was chancellor. She remainec! actively engages! in research for twenty-four more years! Esau con- siclerec! the years in Santa Barbara her most productive en c! fulfilling. She hac! been introclucec! to electron microscopy just before leaving Davis, en c! she was interested in applying this new too! to her anatomical research. She colIaboratec! and published with many people during this period. Today the electron microscope facility bears her name. During her tenure at Davis, Esau stucliec! both cliseasec! en c! healthy plants, inclucling celery, tobacco, carrot, en c! pear. Her work with CheacIle on the comparative structure

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96 . . . . . ~., BIOGRAPHICAL MEMOIRS of the secondary phioem of clicotyTeclons in the 1950s pro- viclec! valuable information regarding the evolutionary spe- cialization of the phIoem tissue in relation to function. In 1953 her classic Plant Anatomy-known woric~wicle as the bible of plant anatomy-was publishecI. This was follower! by Anatomy of Seed Plants in 1960. Both of these books have been publisher! in several languages, inclucling Russian, en c! have extenclec! her influence on the quality of instruction of plant anatomy into classrooms all over the worIcI. The clevelopmental aspects of her studies maturer! into Vascular Differentiation in Plants (1965) en c! her interest in virus-plant host relations into Plants, Viruses and Insects (1961) en c! Vi- ruses in Plant Hosts ~ ~ 968) . In ~ 969 Gebrucler Borntraeger published The Phloem. In it Esau reviewed the structure and clevelopment of phIoem beginning with the earliest records of the tissue. She redrew many of the oic! illustrations from the original articles en c! books. Her mastery of languages, inclucling French, Spanish, English, Russian, German, en c! Portuguese, allowed! her to prepare a thorough review of the very early en c! important German, Russian, en c! French articles. Esau was a superb teacher, in part because she genuinely likes! students. She never failer! to reply to a note or letter from a student, -as- a ' otter~ng encouragement anct praise. Even in her early nineties she answerer! any correspondence she receiver! from a student. She once sail! to a friend, "Don't they know I'm retired?" Her course in plant anatomy was exceptional. A gift for story telling, total command of and enthusiasm for the sub- ject matter, en c! a clelightful sense of humor macle her a truly outstanding teacher. On one occasion when she be- gan a lecture humorously with "Once upon a time," a graduate student quipped, "Aha, another of Esau's fables!" Her abili- ties as a teacher en c! researcher were recognizec! by fellow

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KATHERINE ESAU 97 staff members when in 1946 while still an associate profes- sor she was selectee! to give the Faculty Research Lecture at Davis. Although she servec! as major professor to only fifteen cloctoral students, there are numerous botanists, inclucling many who have never met her but have stucliec! her papers en c! books, who consicler themselves her students. She in- stillec! in her students an appreciation of precision en c! rigor that go into truly excellent studies of plant structure en c! clevelopment. In every aspect of her work she set new stan- ciarcis of excellence. Esau servec! as president of the Botanical Society of America in 1951, en c! in 1956 was one of the original recipients of the Merit Awarc! of that society at its fiftieth anniversary meeting. The Certificate of Merit react: "Katherine Esau, plant anatomist en c! histologist, for her numerous contribu- tions on tissue clevelopment of vascular plants en c! in par- ticular for her outstanding studies on the structure, clevel- opment, en c! evolution of phioem." Katherine Esau was the personification of excellence en c! integrity. Despite her numerous successes en c! many hon- ors, she remainec! moclest en c! close to her Mennonite roots. In 1959, when questionec! about her election to the Na- tional Academy of Sciences, she commentec! "I never wor- riec! about being a woman. It never occurrec! to me that that was an important thing. I always thought that women conic! clo just as well as men.... My surprise at being electec! to the National Academy of Sciences was not be- cause I was a woman, but because I clicin't think that I hac! clone enough to be electecI." Some of her other honors inclucle honorary degrees from Mills College, OaklancI, CaTi- fornia (1962) en c! the University of California (1966) en c! election to the American Academy of Arts en c! Sciences,

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98 BIOGRAPHICAL MEMOIRS the American Philosophical Society, en c! the Swoclish Royal Academy of Science. In 1989, she was awarclec! the National Mecial of Science. The citation accompanying the mecial react: "In recognition of her clistinguishec! service to the American community of plant biologists, en c! for the excellence of her pioneering research, both basic en c! appliecI, on plant structure en c! clevelopment, which has spanner! more than six clecacles, for her superlative performance as an educator, in the cIass- room en c! through her books, for the encouragement en c! inspiration she has given a legion of young aspiring plant biologists, and for providing a special role model for women in science."

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KATHERINE ESAU SELECTED BIBLIOGRAPHY 1930 99 Studies of the breeding of sugar beets for resistance to curly top. Hilgardia 4:417-41. 1933 Pathologic changes in the anatomy of leaves of the sugar beet, Beta vulgaris L., affected by curly top. Phytopathology 23:679-712. 1935 Ontogeny of the phloem in sugar beets affected by the curly-top disease. Am. f. Bot. 22:149-63. 1943 Origin and development of primary vascular tissues in seed plants. Bot. Rev. 9:125-206. 1944 Apomixis in guayule. Proc. Natl. Acad. Sci. U. S. A. 30:352-55. 1948 Some anatomical aspects of plant virus disease problems. II. Bot. Rev. 14:413-49. 1950 Development and structure of the phloem tissue. II. Bot. Rev. 16:67- 114. 1953 Plant Anatomy. New York: Wiley. 1954 Primary vascular differentiation in plants. Biol. Rev. 29:46-86. 1956 An anatomist's view of virus diseases. Am. f. Bot. 43:739-48.

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00 BIOGRAPHICAL MEMOIRS 1957 Phloem degeneration in Gramineae affected by the barley yellow- dwarf virus. Am. I. Bot. 44:245-51. 1960 Anatomy of Seed Plants. New York: Wiley. 1961 Plants, Viruses, and Insects. Cambridge, Mass.: Harvard University Press. 1965 With R. H. Gill. Observations on cytokinesis. Planta 67:168-81. Plant Anatomy, 2nd ed. New York: Wiley. Vascular Differentiation in Plants. New York: Holt, Rinehart & Win- ston. 1967 Anatomy of plant virus i infections. Annul Rev. Phytopathol. 5:45-76. 1968 Viruses in Plant Hosts: Form, Distribution, and Pathologic Effects. The 1968 John Charles Walker Lectures, with a foreword by G. S. Pound. Madison: University of Wisconsin Press. 1969 With R. H. Gill. Tobacco mosaic virus in dividing cells of Nicotiana. Virology 38:464-72. The Phloem: Handbuch der Pflanzenanatomie. Band V. Tell 2, Histologic. Berlin-Stuttgart: Gebruder Borntraeger. 1971 With R. H. Gill. Aggregation of endoplasmic reticulum and its rela- tion to the nucleus in a differentiating sieve element. 7. Ultrastruct. Res. 34:144-58. 1972 With L. L. Hoefert. Development of infection with beet western yellows virus in the sugar beet. Virology 48:724-38.

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KATHERINE ESAU 101 With L. L. Hoefert. Ultrastructure of sugar beet leaves infected with beet western yellows virus. 7. Ultrastruct. Res. 40:556-71. With R. H. Gill. Nucleus and endoplasmic reticulum in differentiat- ing protophloem of Nicotiana tabacum. f. Ultrastruct. Res. 41:160- 75. 1976 With A. C. Magyarosy and V. Breazeale. Studies of the mycoplasma- like organism (MLO) in spinach leaf affected by the aster yellow disease. Protoplasma 90:189-203. 1977 Anatomy of Seed Plants, 2nd ed. New York: Wiley. 1981 With L. L. Hoefert. Beet yellow stunt virus in the phloem of Sonchus oleraceus L. 7. Ultrastruct. Res. 75: 326-38. 1982 With T. Thorsch. Nuclear crystalloids in sieve elements of species of Echium (Boraginaceae). 7. Cell. Sci. 54:149-60. 1985 With T. Thorsch. Sieve plate pores and plasmodesmata, the commu- nication channels of the symplast: Ultrastructural aspects and developmental relations. Am. f. Bot. 72: 1641 -53. 1991 With R. H. Gill. Distribution of vacuoles and some other organelles in dividing cells. Bot. Gaz. 152:397-407.