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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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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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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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· . . . . . · ~.,
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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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.
Representative terms from entire chapter:
biographical memoirs