American J o u r n a l of Medical Genetics 401-25 (1991)
Historical Study: Johann Gregor Mendel 1822-1884
F’ranz Weiling
Bonn-Lengsdorf, Germany
~
The life and personality of Johann Gregor
Mendel (1822-1884), the founder of scientific
genetics, are reviewed against the contempor-
ary background of his times. At the end are
weighed the benefits for Mendel (as charged
by Sir Ronald Fisher) to have documented his
results on hand of falsified data.
Mendel was born into a humble farm family
in the “Kuhlandchen”, then a predominantly
German area of Northern Moravia. On the
basis of great gifts Mendel was able to begin
higher studies; however, he found himself in
serious financial difficulties because of his
father’s accident and incapacitation. His
hardships engendered illness which threat-
ened continuation and completion of his
studies until he was afforded the chance of
absolving successfully theological studies as
an Augustinian monk in the famous chapter
of St. Thomas in Altbrunn (Stare Brno).
Psychosomatic indisposition made Mendel
unfit for practical pastoral duties. Thus, he
was directed to teach but without appropriate
state certification; an attempt to pass such an
examination failed. At that point he was sent
to the University of Vienna for a 2-year course
of studies, with emphasis on physics and bot-
any, to prepare him for the exam. His scien-
tific and methodologic training enabled him
to plan studies of the laws of inheritance,
which had begun to interest him already dur-
ing his theology training, and to choose the
appropriate experimental plant.
In 1865, after 12 years of systematic investi-
gations on peas, he presented his results in
the famous paper “Versuche uber Pflanzen-
hybriden.” Three years after his return from
Vienna he failed to attain his teaching certi-
fication a second time. Only by virtue of his
exceptional qualifications did he continue to
function as a Supplementary Professor of
Physics and Natural History in the two lowest
Received for publication September 12, 1989; revision received
October 1, 1990.
Address reprint requests to Universitats Professor Dr. rer. nat.
Dr. hum. lett. Franz Weiling, Zur Marterkapelle 65, D-5300 Bonn-
Lengsdorf, Germany.
0 1991 Wiley-Liss, Inc.
classes of a secondary school. In 1868 he was
elected Abbot of his chapter, and freed from
teaching duties, was able to pursue his many
scientific interests with greater efficiency.
This included meteorology, the measurement
of ground water levels, further hybridization
in plants (a.0. involving the hawk week Hier-
acium up to about 1873),vegetable and fruit
tree horticulture, apiculture, and agriculture
in general. This involved Mendel’s active par-
ticipation in many organizations interested in
advancing these fields at a time when appro-
priate research institutes did not exist in
Brunn. Some of the positions he took in his
capacity of Abbot had severe repercussions
and further taxed Mendel’s already over-
stressed system. The worst of these was a 10-
year confrontation with the government
about the taxation of the monastery. Attempts
to conciliate (a.0. by appointing Mendel first
as Vice-Director, then as Director of the Mora-
vian Mortgage Bank) did not alter Mendel’s
position.
The recently-dated genetic note-sheet
shows that Mendel was preoccupied with the
interpretation of the results of his Pisum ex-
periments till the end of his life.
KEY WORDS: Mendel-life and personality,
history, heredity and plant
breeding experiments, Pisum,
genetics, origins; apiculture,
meteorology, dating the ge-
netic note-sheet, Mendel-a
forger of data?
INTRODUCTION
Modern genetics has attracted the interest of nu-
merous investigators, as well as the fascination of many
lay persons, in part due to the complexity of its science,
possibilities of its application, and its methodologicchal-
lenges. The story of the discovery of the fundamental
laws of genetics over 120 years ago by the Augustinian
monk Johann Gregor Mendel is still of interest and
importance to science and to mankind in general.
Following the “rediscovery” of the laws presented by
Mendel in 1865 (and published in the following year as
2 Weiling

“Versuche uber Pflanzenhybriden”) [Mendel, 1865a,bl, two dcughters [Weiling,1983~1. His birthplace Heinzen-
confirmation was brought in all areas of plant, animal, dorf (Czech: HynEice) in the Kuhlandchen, was located
and human life. However, hardly any of Johann Gregor in the border area of that small part of Silesia which
Mendel’s discoveries have raised so many questions; remained with Austria after the Silesian wars (1740-
their elucidation took decades while the appreciation of 1742, 1744, 1756-1763) fought between King F’re-
the discoverer Mendel fluctuated between highest rec- derick I1 of Russia and Empress Maria Theresia of
ognition and virtually complete oblivion or misinterpre- Austria. Thus, in practically all later official documents,
tation. Mendel was referred to as a Silesian (Sil.: Silesius).
Thus we may pose the following questions: 1)Exactly Ecclesiastically, Heinzendorf belonged to the Moravian
parish of Gross-Petersdorf (6s: Vrazne) located about a
how did this monastic priest arrive at this biological core
question and its elegant solution; 2) how is it possible to
half hour away. Its parish priest, Johann A.E. Schreiber
explain that his discovery and knowledge about his per- (1769-18501, had a special interest in natural history
son remained unknown for such a long time that it took and was known as an expert fruit grower; he had a
decades of research to arrive at a half-way satisfactory lasting influence on the boy Mendel’s development
answer; and 3) how was it possible for the founder of [Weiling, 1984el. It was Schreiber, more than anyone
modern statistics, Sir Ronald Aylmer Fisher (1890- else, who recognized the gifts of the young Johannes and
1962),to conclude in 1936 that “the data of most, if notpersuaded his parents to release their only son for
all, of his experiments have been falsified, so as to agree
higher education in spite of their constrained financial
closely with Mendel’s expectations” [Fisher, 19361, situation. Initially Mendel attended the village school
whereas the interpretation of his experiments is ac- in Heinzendorf. At the age of 11 (1833) he transferred
cepted as valid. During the past 20 years this statement into the third grade of the Piarist Central School in
caused Mendel to be widely accused of falsifying his dataLeipnik (Cs: Lipnik) and in the following year he was
[Weiling, 19851 (Fig. 1). sent to the Gymnasium in Troppau (Opava) where he
The following is an attempt to offer an answer corre- completed all 6 years successfully.
sponding as closely as possible to our present knowledge. At that time the Austrian school system provided in
the public gymnasia four grammar classes followed by
MENDEL’S LIFE two classes in the humanities always taught by the same
Johannes (his birthname) Mendel was born on July 20 grammar and humanities instructor over the entire
or 22,1822, the only son of peasant farmers who also had 6-year cycle. This teacher taught all subjects (Latin,

Fig. 1. The “Rogues’gallery” of Wade [Wade, 19831 including Johann Gregor Mendel.

Greek, geography, history, and mathematics) for 18
hours per week; 2 hours per week were devoted to mathe-
matics [Timp, 19681. The prefect at that time, who also
functioned as religion in$ructor, was an Augustinian
Father of the Altbriinn (Cs: Stare Brno) chapter [Weil-
ing, 1984al.
At that time natural history was not a didactic subject
in the school system. However, Mendel had the oppor-
tunity to regularly visit the Museum of Natural History
of the school (today Schlesisches Landesmuseum) (cs:
Slezske Muzeum Opava) which had been founded in
1822. There he was, presumably, of great assistance to
its director and co-founder (also grammar teacher) Pro-
fessor Faustin Ens (1782-1858), in caring for and deve-
loping the collections [Weiling, 1983al.
The support which the parents were able to offer their
son was restricted to regular “care packages.” However,
these stopped when Mendel’s father became incapaci-
tated after a severe accident and finally had to turn the
running of his property over to his son-in-law. In his
Autobiography [Mendel,18501Mendel wrote that “after
several disasters following in rapid succession. . . his
parents became completely unable to pay for the neces-
sary expenses of [his] studies” (Fig. 2).
Mendel tried to stave off his direst needs by offering
remedial tutoring for which he had been approved
through a special examination after a private course.
Nevertheless, his constant deprivations led to a severe
illness which forced him t o rest in his parents’ home
from Pentecost till September in 1839 [Iltis, 19241.
Mendel’s miserable existence did not improve after
completion of his gymnasium studies in the middle of
1840and matriculation into the 2-year course of philoso-
Fig. 2. Final portion of Mendel’s handwritten account of his life [Mendel, 18501submitted by him to the
Examination Commission of Vienna together with other documents.
Mendel 3
phy at the Philosophical Institute of the University of
Olmutz (Olomouc). This course of studies included
philosophy, Latin and Greek philology, mathematics,
physics, religion, as well as the facultative fields of natu-
ral history and history. Greek, physics, and history were
taught in the third and fourth semesters. As a stranger
in town Mendel found no opportunity of supporting him-
self through remedial instruction [Mendel, 18501. He
became ill during the final exams of the first semester,
after having passed the examinations in mathematics
and Latin philology with highest grades (Fig. 3). Thus,
Mendel lost not only credit for the entire year, but also
the free time granted to him till then for his studies. The
spring and summer of 1841were apparently spent recu-
perating in his parent’s home, according to his Auto-
biography, even “a full year.” It was only when his youn-
ger sister Theresia offered him a portion of her
inheritance that he was able to renew his studies in 1842
[Weiling, 19881, now as a paying student. “Using all of
his strength he succeeded this time t o complete his two
years of study” [Iltis, 1924, Mendel, 18501. He attained
the highest grade in all courses (e = eminens) except for
theoretical and practical philosophy, in which he got the
second-best grade (1=prima classis). Thus, it appears
that the practical courses, especially mathematics and
physics, had a much greater interest for Mendel than a
more speculative field such as philosophy, a conclusion
supported by the qualifiers used to describe his efforts
(“by application”) (Fig. 3).
In this connection it is important to note that of the 20
or so hours of classes attended per week in Olmutz, 7 and
8, respectively, were devoted to mathematics in the first
year and t o physics in the second both were areas of
4 Weiling

Fig. 3. Excerpts of the records of studies of the Philosophical Institute of the University of Olmutz with
the entries pertaining to Johann Mendel. A the year 1841;B: the years 1842 and 1843 (State Archive
Olomouc, Czechoslovakia). During the earlier attempt Mendel had the number 49,during his subsequent
course (as repeater) the number 42.

special importance to him in his later scientific career.
Mendel’s mathematics instructor was Professor Johann
F’ux (1785-1848) [Weiling, 1984fl (Fig. 4) who was
widely known for his textbooks; his physics instructor
was Professor Friedrich F’ranz (1796-1860) [Weiling,
1983b1, a Prqmonstratensian and later Abbot of Neure-
isch (Nova Ris6). Before his appointment in Olmiitz
(1842), F’ranz had taught physics for 19 years a t the
Philosophical Institute in Briinn, and during that time
he had lived in the Altbriinn monastery of the Augus-
tinians [Weiling, 1982al. Franz gave Mendel a solid
character reference, in fact referred to him “as almost
the best” in physics [Iltis, 19241.On his recommendation
Mendel was accepted as a novice in the Altbriinn chap-
ter after completion of his philosophical studies (1843).
This freed him at once of his struggle for existence and
“courage and strength returned with that level of com-
fort desirable for all studies” [Mendel, 18501. In his Au-
tobiography Mendel wrote that his condition decided on
his change of status. This was not meant to refer to the
priestly state, as has sometimes been assumed, but
rather to the monastic state, i.e., membership in an
order. In the contract (1842) in which Mendel’s father
turned over his home and farm to his son-in-law,he also
makes provision for his son Johann in case he were “to
follow his wishes and enter the priestly state” and in
case he would be “prevented through a disease” from
“attaining the priestly state’’ [Iltis, 19241.

i b c r
Irit@tuetiP nab aiebctc %leebra.
--#wwwwM-
SImQe, 1839.
;Drub u n b ‘ p a p t t r @ o n 2ldlIol6 e r a r n i b f .
Fig. 4. The textbook of Arithmetic and Basic Algebra from the
series on pure mathematics by Johann Fux, Olmutz, 1839.
After completion of his novitiate, Mendel absolved his
theological studies between 1844 and 1848 at the Epis-
copal Seminary in Briinn. On August 6, 1847 he was
ordained a priest. During the course of his studies he
also completed a l-year (1845-1846) agricultural cur-
riculum (Fig. 5) at the Philosophical Institute in Briinn
under Professor Franz Diebl (1770-1859). At the same
time he attended a semester course on fruit growing and
viniculture (Fig. 6). At the beginning of the 19th cen-
tury (however,not at Mendel‘s time anymore) these ad-
ditional studies were required of theology students,
since the Austrian government intended the rural
priesthood to effectively teach and pass on their agri-
cultural knowledge to the local population. These
studies were also in the best interest of the Altbriinn
monastery with its own agricultural domains, and thus
of its Abbot, Franz Cyril1 Napp (1792-1867) [Weiling,
1968, 1971bl (Fig. 7).
After a brilliant performance in his professional ex-
aminations of the theological Faculty of the University
Mendel 5
of Vienna, Napp was appointed Professor of old-testa-
mentary hermeneutics and ancient Oriental languages
(including Arabic, Syriac, and Chaldean) at the Theo-
logical Institute in Briinn. In 1824 Napp has been elec-
ted Abbot and quickly mastered the administrative1
economic duties of the monastery. He was a member of
the Executive Committee of the Moravian-Silesian Ag-
ricultural Society; Director of the Pomological Society
(as of 1827),the predecessor of the later section for fruit,
horticulture, and viniculture of the Agriculture Society;
and, as of 1849, Vice-Director,and in 1865 the General
Director of the Agricultural Society. As such, Napp
manifested not only profound technical knowledge and
interests, but also proved himself to be a skillful judge of
people and a splendid organizer. And as the Director of
the State Gymnasia of Moravia and Silesia he was for
many years the supervisor of higher education until the
reorganization of the Austrian school system after the
revolution of 1848.
Not only Napp, but also other members of the chapter
held public office. Thus, the chapter had to provide the
professors of philosophy and mathematics at the Philo-
sophical Institute in Briinn and to keep appropriate
candidates ready for these positions at all times [Weil-
ing, 1982al. Even for the chair of Natural History and
Agronomy the monastery was prepared to provide a
trained candidate after the vacancy of 1825-1828
[Weiling, 19761.
It is t o be expected that the theology student Gregor
Mendel was in constant contact with these obligations of
the chapter, particularly since he himself additionally
completed the entire agriculture curriculum then avail-
able. In his Autobiography he emphasized that during
his theological studies he had dealt with botanical and
natural-historical problems in the framework of the
available opportunities and that, as a consequence, his
liking for natural history increased greatly.
With great likelihood Mendel was also confronted
during these years with questions of heredity, i.e., the
question of how, according to which rules, above all
valuable traits are transmitted from generation to gen-
eration, and how crossing of sorts and types could com-
bine and increase valuable traits. Astonishing results of
some sheep breeders in the Briinn area had regularly
occasioned detailed discussion of these problems in the
1820s and 1830s at the meetings of the Agricultural
Society; even at the beginning of the 1840s they were a
subject at an international scientific meeting at which
Napp participated substantially in the discussion [Weil-
ing, 19881. Botanically, the problem of plant improve-
ment through crossing, especially in fruit growing, was
of special interest. But also the efforts in collecting ex-
tensive assortments of different types, e.g., in sheep
breeding (wool improvement) and in viniculture (Fig. 8),
pointed in this direction and may have been ofinterest to
the young Mendel.
Soon after completion of his theological training the
young Mendel turned out to be little fit for practical
pastoral duties. According to a report by Napp to the
Bishop of Briinn “at the bedside of the ill and suffering
[Mendell was overcome with a paralyzing shyness, and
he himself then became dangerously ill” [Iltis, 19241.
6 Weiling

Fig. 5. “Report card” concerning the outstandingly successful performance by Mendel during the
second semester of the study of agriculture at the Episcopal Philosophical Institute of Briinn. The certifi-
cate is signed not only by the instructor Professor Diebl, but additionally by the certifying commissioners of
the Bishop and of the Moravian Silesian Agricultural Society.

Therefore he was freed up for teaching and in Septem-
ber 1849 he was appointed adjunct instructor at the
Gymnasium in Znaim (Znojmo)where he taught Latin,
Greek, German, and mathematics during the winter
semester [Kfiienecky, 1965; Czihak, 19841. However,
since he lacked the required state qualifications to
teach, at the urging of the Director of the School after
this semester he applied to the appropriate agency in
Vienna to take the examination in Natural History for
all grades, but in physics only for the lower grades.
Mendel did not pass this examination; however, the noti-
fication, dated October 17, 1850, was not forwarded to
him till August 1851 [Weiling, 19861. At the advice of
Professor Andreas von Baumgartner (1793-1865) (Fig.

t. f. ')lroiqTor bcr P~nbmirtl)id)~ft6kf)reunb aUjemcintn %JtUrl
8ef4iete Jn Err pl)ilofop$ifd)en EtlpnPdlt iu Brdiin unb Binit0
glirte mef)rrrer IunEloirt~f@JftlidW@efCuid)aften 3nns
unb %u6lanbe&
-a-
--- - ----=-==zT;;&-- _ _ __
SrCitut 1644,
Bcbrudt bei %ubolp$ %ol)rer'4 f d Bittior.
Fig. 6. Professor Diebl's textbook of arboriculture, especially per-
taining to fruit trees, viniculture, and forestry, Briinn, 1844.
9) who had been the examiner in physics but in the
meantime had been appointed the Austrian Minister of
Commerce, Mendel enrolled as a special student for
further studies a t the University of Vienna.
During the reorganization of higher studies in Aus-
tria after the revolution of 1848 the philosophical insti-
tutes had been dissolved; they existed only in locations
where there was also a seminary since theological train-
ing had t o be preceded by attendance a t a philosophical
institute. The natural-historical subjects of these insti-
tutions were transferred to the gymnasia whose curricu-
lum was thereby expanded by the 7th and 8th grades. In
order to meet the need for appropriately trained instruc-
Mendel 7
Fig. 7. Prelate Franz Cyril1 Napp in contemporary clerical garb.
tors, chairs were created or reorganized, primarily
within the philosophical faculty of the University of
Vienna, in order to offer required courses in physics,
botany, zoology, and chemistry [Weiling, 19861.
Thus, in Vienna a new Physics Institute was created
under the direction of Professor Christian Doppler
(1803-1853) (Fig. 10); it began instruction late in the
winter semester 1851-1852 in a recently completed
and newly equipped institute building. We are partic-
ularly well-informed on the operations of this institute
because it was based on regulations which had been
formulated by the Ministry on the basis of a draft re-
quested by Doppler. Accordingly, the maximally three
semester-long training courses in physics consisted of 2
hours per day of active classroom instruction, with the
entire rest of the day devoted to individual demonstra-
tions or research projects. In this connection each of the
candidates, after completing the work and experiments
required for school instruction, was to perform an indi-
vidually assigned project. These projects were to include
the appropriate library research, the experimental de-
sign and protocol, as well as a final report under contin-
uous supervision and eventual evaluation and correc-
tion by the Director of the Institute. In addition, these
research papers were to be submitted to the Ministry at
the end of academic year together with the Director's
progress report [Weiling, 19861.
Mendel began his studies in Vienna late, 4 weeks after
the beginning of the semester, however, in time for the
8 Weiling

Fig. 8. List of the type of grapes grown close to the chapter of St. Thomas, Altbriinn [Mittheilungen,
1868, No. 21, May 1830, pp 166-1671. During the first half of the 19th century there were similar
collections of grapes in two other places in Brunn.

opening of the Physics Institute. As a special student
(ausserordentlicher Eleve) he attended exclusively the
Physics Institute during the first semester and part of
the interval between semesters. It may have helped
Mendel that, like Mendel, the Assistant a t the Institute,
Franz Pekarek, who held this position over the next
several years, had also been active as an adjunct instruc-
tor during the winter semester 184911850 at the Gym-
nasium in Znaim.
Only during his second semester did Mendel take
additional courses: plant morphology and systematics in
botany under Eduard Fenzl (1808-1879) (Fig. 91, and
systematics and lab in zoology under Rudolf Kner
(1810-1869). However, there were some scheduling
conflicts with physics. During the third semester it was
not possible for Mendel t o attend the lectures announced
by Doppler who became ill and died shortly afterwards
(1853). Doppler was immediately replaced by Professor
Andreas von Ettingshausen (1796-1878) (Fig. 9) under
whose direction the practical projects at the Physics
Institute remained considerably restricted during the
first few years. During this semester Mendel took most
of his non-physical courses: under F'ranz Unger (1800-
1870) (Fig. 9)anatomy and physiology of plants as well
as the first lab course at the University of Vienna in
botanical microscopy; under Josef Redtenbacher (1810-
1870) (Fig. 9) general and pharmaceutical chemistry as
well as methods of chemical analysis. In addition he
 Mendel 9

Fig. 9. Group portrait of the members of the mathematical-historical class of the Academy of Sciences
in Vienna (about 1855)[Meister, 1947, plate 19, with five of Mendel’s teachers A.v. Baumgartner (second
sitting from left), A.v. Ettingshausen (first sitting from left), E. Fenzl (fifth standing from left), F. Unger
(first standing from right), J. Fkdtenbacher (fourth standing from left).

attended two 1-hour lectures on paleontology as well as a
1-hour lecture on logarithmic-trigonometric tables.
During his fourth and last semester Mendel took von
Ettingshausen’s course on introduction to and use of
physical instruments and higher mathematical physics,
and Redtenbacher’s course on organic chemistry. In ad-
dition, he established connection, perhaps as early as
the second semester, with the zoological division of the
Court Natural History Collection and its director Vin-
zenz Kollar (1797-1860). This resulted in a paper
which Mendel presented on July 21,1853, a few weeks
before his final return to Brunn, before the Zoological-
Botanical Society of Vienna [Mendel, 1853; Weiling,
19861.
Thus, Mendel’s studies in Vienna were primarily de-
voted to physics, having worked completely or exten-
sively at least 1full year in the Physics Institute under
Doppler. Doppler died early, but here Mendel may have
acquired his later so successfully applied scientific
method, especially since it can be assumed that during
the first year of the newly created Institute Doppler
proceeded strictly according t o the rules of the Institute,
largely due to his own initiative. In addition, Doppler
was far more practically oriented, and didactical in his
mode of work, than his successor von Ettingshausen.
Since the rules of the Physics Institute contain detailed
instructions to study the literature and since it can be
assumed that Mendel personally acquired at the latest
during his stay in .Vienna C. F’r. von Gartner’s
(1772-1850) book: “Uber die Bastarderzeugung i m
PfZanzenreich” (concerning the production of hybrids in
the plant kingdom, 1849; Figs. 11, 12); it can be con-
cluded that under the influence of that book Mendel
read the paper he later quoted by J. G. Kolreuter
(1733-1806) [Kolreuter, 1761-17661, W. Herbert
(1778-1847) [Herbert, 18371,and H. Lecoq (1802-1871)
[Lecoq, 18451, probably accessible only in Vienna. We
can therefore assume that Mendel made the detailed
plans for his later Pisum experiments in Vienna, and
began them, i.e. obtained and tested the various types
and sorts of seeds, immediately after his return to
Brunn.
However, upon return t o Brunn (18531, Mendel did
not, as might have been expected,present himself for the
final exam, but only 3 years later when difficulties arose
in the as yet incomplete Oberrealschule in Brunn where
he taught without having taken the exam. Mendel did
not pass this exam either. According to Kfiienecky
[19631 a t the beginning of the written exam on May 5,
1856in Vienna, Mendel became so ill that he was unable
to write and gave up. It is known that Mendel was in
poor health while preparing for the exam presumably
through overexertion. Now he became so ill that, in spite
of difficult travel conditions, his father and uncle rushed
over 120 km from Heinzendorf to support him
[KEiienecky, 19651. Thereafter Mendel did not attempt
to repeat the exam, and the Director of the Ober-
realschule, Josef Auspitz (1812-1889) was evidently
able, on the basis of Mendel’s exceptional pedagogic per-
formance and in the face of the official rules, to retain
him as a professor at the school (Fig. 13).
Mendel informed the Munich botanist Carl Nageli
(1817-1891) on April 18, 1867 that he had performed
the actual experiments of his Pisum work from 1856
until 1863, inclusively [Correns, 19051. In contrast to
Fisher 119361 (who later corrected himself) [Fisher,
19551and in agreement with Orel [Orel, 1971al it may
be assumed that the first crosses were performed in
10 Weiling

Fig. 11. Carl Friedrich von Gartner [Stubbe, 1965, p 1021.

netic study on differences in flowering time) intended for

publication were concluded in 1863 to obtain “space and
time for the cultivation of other experimental plants”
[Correns, 19051. At that time infestation with the pea
beetle Bruchuspisi was so heavy that beginning in 1864
substantial damage was recorded and in 1865 barely
20-25% of harvested peas were uninvolved. He men-
tions in the same letter that in the vicinity of Brunn the
cultivation of peas had to cease altogether.
The final analysis of his experiments and the prepara-
tion of the manuscript probably occurred in 1864. Men-
Fig. 10. Bust of Christian Doppler in the courtyard of the Univer- del reported on his work in detail t o the Naturforschende
sity of Vienna. Verein of Brunn (founded in 1861) a t the sessions of
February 8, and March 8, 1865. Mendel’s paper was
printed in the volume of Proceedings for the year 1865;
however, it appeared only toward the end of 1866 (Fig.
1856. This would involve at least the 118 crosses per- 15).The reason for this delay was the war of 1866 [Weil-
formed on 25 pea plants, with the traits smooth and ing, 1970~1, which visited on Brunn a temporary occupa-
wrinkled seeds respective green and yellow seed color, tion by Russian troops, food shortages, and a serious
during the year of his disastrous second board examina- cholera epidemic with some 3,000 deaths (1,000 citizens
tion. In the two preceding years he had subjected 34 and more than 2,000 Prussians) [Mendel, 18663. Accord-
“more or less different” types of peas to a test of purity, ing to the information on its yellow covers, this volume
and selected 22 types for his subsequent experiments appeared in an edition of 500 copies of which 115 went to
[Mendel, 1865a,bl. scientific institutes or libraries [Weiling, 1969133.There
In order to appreciate his work during the years of his were 40 reprints of Mendel’s work; the whereabouts of 13
Pisum experiments it must be mentioned that by no of these is presently known [Weiling, 1984~1.
means all experiments entered his later publication and Already 6 months after its appearance Mendel’s work
that since 1854 he also worked at the Brunn Ober- was cited in the May 14, 1867 issue of the well-known
realschule teaching the two lowest classes of Naturlehre botanical journal Flora in a “repertoire of the periodic
(Physics) and natural history with, depending on the botanical literature.” In addition, Mendel’s work is
number of parallel classes, 18 and 27 hours per week of quoted both in the register of authors as well as in the
instruction and 60 to 120 students per class [Programme index of subjects (key words: Bastarde and Hybridittit) of
of the Oberrealschule Brunn] (Fig. 14). the volume of this journal [Weiling, 1969133.By 1870one
According t o Mendel’s own information (second letter other publication, by 1880 another four, by 1890 a fur-
to Professor-Nageli) all experiments (except for one ge- ther two, and one other by 1900 in the German, Swedish,
 Mendel 11

on cell division and the behavior of the easily stainable

bodies in the nucleus (chromosomes)which appeared to
have something to do with the genetic mechanism. The
British genius, Sir Francis Galton (1822-19111, who
worked extensively with the heredity of human faculties
and character traits and developed his own model of
heredity, apparently was unacquainted with Mendel’s
work.
Following the strong recommendation of the influen-
tial Munich botanist Carl Nageli, Mendel was induced
to begin detailed crosses in Hieracium. However,
through behavior which is peculiar to this plant
(apogamy) and which was recognized and worked out
only much later, the reproductive mechanism functions
in a totally deviant manner. New and completely differ-
ent challenges which Mendel had to meet as of 1868 did
not allow him the required time for further systematic
genetic investigations. Nevertheless, the Hieracium ex-
periments, presented by Mendel before the Natur-
forschende Verein in Brunn on June 9, 1869, are re-
markable [Mendel, 18691. Hieracium was a difficult
genetic subject because of the delicate construction of
the individual blossoms; however, the fact that with only
one exception all known artificially produced Hieraciurn
hybrids derive from these experiments of Mendel attest
to his extraordinary experimental skill. From five of six
successful hybrids Mendel sent living material includ-
ing the parental stock to Nageli in Munich. It was propa-
gated in the Munich botanical garden and then was
incorporated into the subsequent systematic investiga-
tions of Nageli and his coworker Albert Peter (1853-
1937) [Peter, 1884,1885; Nageli and Peter, 18851.Peter
included specimens of 11 original forms and 17 hybrid
from Mendel’s stock in his four volume herbarium of the
Hieraciae [Peter, 1885, 18861 (Fig. 16). Of this work, I
was able to track down 26 admittedly in part incomplete
specimens in herbaria of 13 different European coun-
tries [Weiling, 1969al.
On March 31,1868 Mendel was elected the successor
of the deceased Abbot Franz Cyril1Napp of the Altbrunn
Fig. 12. Mendel’s personal copy of the book by Gartner. It contains chapter. This brought to an end not only his work as
numerous annotations and notes in Mendel’s handwriting, especially
on the front fly leaves. adjunct Professor in the Oberrealschule, but also, after a
short phase-out period, the complete cessation of his
work in genetics. Administrative and public service
obligations predominated, especially in the executive
committee of the Moravian-Silesian Agricultural
Russian, and Anglo-Americanliterature had cited Men- Society.
del’s Pisum work [Weiling, 1971al. A particularly thor- In 1874, a tax law was enacted which obligated mon-
ough discussion is in the 1874 St. Petersburg disserta- asteries and convents to substantial contributions to the
tion of Johannes Schmalhausen, a descendant of a Religious fund from which the State drew means to
family originally from Bremen [Weiling, 1966, 1974, support the church. Mendel’s protest grew into a pro-
unpublished manuscript]. The formerly frequently tracted and wearisome confrontation. Since he viewed
voiced opinion that Mendel’s work was unknown before the law as unjust, Mendel, as the only prelate in the
1900 and thus had fallen into oblivion is untenable on Austrian empire, resisted it, in the face of the ensuring
the basis of these data. confiscation of a substantial portion of the monastery
However, the problem of the genetic process itself and income. After Mendel’s death, his successor compro-
its laws lost importance in Brunn because of the increas- mised; however, he insisted, successfully, on return of
ing cotton industry and simultaneous reduction of the the overpayments. At the same time the chapter was
importance of sheep wool. On the other hand, work in relieved of any further payments of this tax, since the
cytology, especially by Walther Flemming (1843-1905) true income of the monastery was below the level stipu-
a.o., August Weismann (1834-19141, and Eduard lated by the law.
Strasburger (1844-1912), yielded detailed knowledge Early in the morning of Epiphany (January 6,1884)
12 Weiling

Fig. 13. A previously little known (incomplete) group portrait of members of the Altbrunn chapter of
St. Thomas from a time between 1861-1864 (University Library Brno, Czechoslovakia). This is a different
grouping than that in Iltis [1924,plate 41 involving the same persons and taken at the same occasion.Ofthe
persons portrayed here, the following were engaged as teachers: Mendel holding a fuchsia blossom; Fr.
Thomas Bratranek (standing third from left), for 30 years Professor ofGerman Literature at the University
of Krakau, today Krakow, Poland: Fr. Antonin Alt (standing, second from right): elementary mathematics,
at times director of the Gymnasium of Troppau (Opava, Czechoslovakia); Fr. Benedikt Fogler (standing,
first from the left), instructor of German, Italian, and French at the Oberrealschule in Brunn; Fr. Paul
KfiikowskJi (standing second from left), composer, director of the Thurn-Foundation for Musical Education
(conservatory) at the Altbrunn chapter (later attended by the famous Moravian composer LeoB JanaEek,
1854-19281, as well as choir director of the Cathedral of Olmutz (Olomouc).

Mendel died of chronic renal disease and heart failure His failure during the first board examination must
(Bright’s disease and dropsy according to the death cer- also be regarded as a blow of fate. He had to fail, because
tificate) [Kfifenecky, 1965; Weiling, 1984dl. without appropriate specialty training, he had to sub-
ject himself t o much higher standards of examination
MENDEL’S PERSONALITY which had been improved during the reorganization of
Mendel was a multifaceted personality whose life was higher education following the 1848 revolution.
marked not only by remarkable successes but also by As a further blow of fate must be regarded the failure
great adversities and blows of fate. The severe accident of his second board examination in 1856 which he had to
and incapacitation of his father (1838)and other misfor- call off at the very beginning because of severe illness
tunes in his family deprived him of financial support for with inability to write with subsequent irregular ap-
his education. The associated deprivations damaged his pointment as supplementary professor. Apparently his
health through inadequate nutrition, illnesses, and re- physical and emotional status had been so badly im-
duced resistance. This forced him to interrupt his philos- paired by the preceding misfortunes that he was not up
ophy studies already during the first semester exams to the additional stress of preparing for and taking the
causing him to loose his free study time. He was able to examination.
continue his education only after his sister Theresia A final blow of fate was the prolonged and wearisome,
gave him a part of her inheritance which allowed him to health-damaging conflict with the government about
complete his philosophical studies to take advantage of the religious tax which Mendel, in fidelity to his oath of
the subsequent opportunity to pursue a priestly voca- office and his conviction about the illegality of the law,
tion as a monk in the Altbriinn Chapter. Mendel believed unable to avoid.
thanked his sister by financing the education of her This externally evident professional failure of Mendel
three sons, one of whom died during his studies. must be contrasted with the fact that it was his Pisum
 Mendel 13

c e .,<&,.,,&$
1(1 ,".*.,,<,,,.,,,,,,
' -
Fig. 14. Statistics compiled by Mendel of the students and instructors of Section b of the second grade,
supervised by Mendel during the academic year 186311864,at the Oberrealschule in Brunn. The students
are listed numerically according to their nationality (Germans, Slavs), religion (Catholics, non-Catholics,
Jews), residence (Briinn, outside Briinn), tuition status (paying, free), and their performance (outstanding,
first, second, and third place). Mendel himself taught physics and natural history (botany, zoology, and
minerology). Archive of the city of Brno.

work which threw light on the fundamental basis of
hereditary and thus, on one of the most important pro-
cesses in biology, namely the transmission of traits from
generation to generation; and this is an age before scien-
tific biology had an understanding of his work and its
fundamental importance.
In this connection it must be noted that without the
repeated misfortune with his board examination he
would hardly have gotten the chance to do his almost 12
years of work with peas. He may have become a parish
priest or taken on some other pastoral duty or spiritual
function, or, as happened to several of his fellow monks,
he might soon have been entrusted with the direction of
the school. Then he would have lacked the time for his
experiments which he had available to him during the
entire period of his experiments as a substitute teacher
in the two lowest grades.
The multifaceted nature of Mendel's scientific inter-
ests were especially evident after he took the direction of
the chapter. Since the emphasis of his studies in Vienna
was on physics it is understandable that during his
entire active period of life he was engaged in meteoro-
logic work (Fig. 17). Already in 1857 he was entrusted
with the data collected since 1848by Dr. P. Olexik for the
purpose of graphic representation. And in connection
with the Natural History section of the Moravian-Sile-
sian Agricultural Society (to which he belonged since
1851 and in which he renewed his membership in 1855)
he was further appointed consultant for meteorology
[Weiling and Orel, 1967; Orel, 1970bl. In 1862163 he
submitted a comprehensive graphidtabular summary
of the meteorologic conditions of Brunn based on 15
years of data [Mendel, 18621 (Fig. 18).His bibliography
shows that until 1870 this was followed on an almost
14 Weiling
Fig. 15. Beginning of the manuscript of Mendel’s Pisum paper. The manuscript, written by Mendel
himself was found in 1900by H. Iltis. It was lost from a safe of a Brunn bank at the end of the war when the
town was plundered (1945) [Mendel, 1865b; Gedda, 19561.
annual basis with a summary of averages based on data
collected by the first meteorologic stations founded in
Moravia and Austrian Silesia [Kfiienecky, 1965; Weil-
ing, 1970~1 (by the time of Mendel’s death the number of
these stations has risen to over 170) [Weiling, 1970bl.
After the death of Olexik (1878) Mendel took over the
Brunn station himself [Weiling, 1984bl. Furthermore,
Mendel, especially in his capacity as a standing member
of the Executive Committee of the Agricultural Society,
was a t pains to spread meteorologic knowledge and to
provide efficient weather prediction to Moravian agri-
culture [Mendel, 1879; Orel, 1969; Weiling, 1982b1.
An eloquent testimony to Mendel’s meteorologic ac-
tivity are the monthly meteorologic tables preserved
today in Brno and prepared while he was the Director of
the Brunn meteorologic station for the then Central
State Institute for Meteorology and Earth Magnetism in
Vienna (Fig. 19). Their monthly preparation is docu-
mented in two sheets of calculations (these two sheets
contain other, e.g., tax notes) preserved in the Mendel
Archives in Brno [Weiling, 1970al. A good insight into
the sharpness of Mendel’s talent for observation is ob-
tained from his detailed report on a tornado which swept
over Brunn and the monastery on October 13, 1870
[Mendel, 18701.
In this connection it is important to mention the mea-
surements Mendel undertook regularly over 14 years
(1865-1878) of the ground waterlevel, and the results of
which he communicated in the meterologic monthly
summary of October 1878to the Central Institute and in
a letter of December 8, 1878 to the Chairman of the
Naturforschende Verein of Brunn, Professor Gustav
Niessl von Mayendorf (1839-1919) [Liznar, 1902; Weil-
ing, 1982131.These measurements had been instituted a t
the suggestion of the great Munich hygienist Max von
Pettenkofer (1818-1901) who suspected a relationship
between ground water levels and the occurrence of chol-
era epidemics [Weiling, 19751.
Apiculture was a special interest of Mendel. In 1870
he joined the very active and numerous Brunn Apiary
Society which he served as Vice-chairman between
1871-1873; he declined the Chairmanship after the
death of its founder Dr. med. Franz Ziwanski
(1817-1873). The apiary of the Altbrunn monastery
has a capacity of 30 hives (though sometimes Mendel
had 50 hives). It was established by Mendel in 1871 and,
after successful restoration, it is presently used by the
Brno Apiary Society [Matalova and Kabelka, 19821.
Here Mendel was engaged not only in the care of his bees
but also with the crossing of races of bees [Iltis, 1924;
Beranek and Orel, 19881 and the study of a possible
acclimatization of foreign and even tropical races of bees
[Alpatov and Orel, 19791. Mendel regularly attended
the meetings of the Society in which he reported on
 Mendel 15

Hieracia Nsegeliana ed. A. Peter.

13. Hieracium auriculiforme Fr.

snbsp. Mendelii 1. striatnm.
Plaota hybrida inter H. Pilosella subsp. bruennenae et H. Auricula
anbsp. Auricula a. geuuiuem 1. normale 0, a el. Mendel foecundatioue artificiali
procreata, in horto botanic0 Monacensi culta. - 1.
VI. VIL. VIII. r X . 2 .

Hieraoh Naegaliana od. A. Peter.

21. Hieracium pyrrhanthes n. hybr. ba

rabsp. .pyrrhmthes a. geoninnm 1. obtasum.
Planta hybrida inter H.aurantium 1. norniale e t H. Auricula z.genu-
innm 1. nnrmale 9 a cl. Mendel (Bruenn) foecundatione artificiali prncreata.
in horto botanic0 Monseensi culta. - VI. VII. VIII. 1X.
c
#rIt e o r o l o 8isi
22. Hieracium pyrrhanthes n. hybr.
rrabsp. ohryaoobronm.
Plant0 hybrida inter H. aurantisoum 1. normale st H. Auriciila

-
s. genninam 1. normale 0, a cl. Mendel (Brnenn) foeciindatione artificinll
procreata, in horto botanico Monacensi c u b . - VI. VII. VIII. 1X

Hieroolo N8qaiiana ed. A. Peter

23. Hieracium pyrrhanthes n. hykr.

snbsp. rsripilnm.
Planta hybrida inter H.a u r a n t i n m 1. normale et H. Auricula a.genu-

-
inum 1. nonnale 7, a cl. Meudel (Ilruenn) foecnndatione artiticiali prncretita.
iii liorto Mania, blonacensi nllta. - VI.. VII. ’ V ,Smtitr UafIogc.

a3fcr, 1860.
questions of bee care and stimulated the other apiarists
to document observations and undertake investigations. !Bilbtfm braurntiller,
iWUaP1rr prl .I f brht U P kr IJdrrl Sfrbnk kr 5liiinibrllrv
At one of these meetings he reported on a 4-year period
of observations concerning the flight of a selected stock Fig. 17. The textbook used by Mendel on Meteorology by A.
of bees with the aim of estimating the amount of honey Kunzek. It contains numerous annotations and marginal notes in Men-
produced from the accumulated data [Mendel, 1877; del’s del’s handwriting, as well as the motto on the title page (also in Men-
handwriting): “Whoever cannot be alone is not reconciled with
Weiling, 1982bl (Fig. 20). himself‘ [Weiling and Orel, 19671.
In 1859Mendel participated with his own choice vege-
tables in a horticultural exhibition in Brunn, and in
1863 he was elected into the horticultural section of the
Moravian-Silesian Agricultural Society whose affairs
he represented till his death [Iltis, 19241. Here also he “epochal” importance [Anonymous, 1884; Matalova,
combined practical interests such as advances in hor- 19841.A story is told about JosefMaresch, Mendel’s long
ticulture (a.0. through participation in fairs as sponsor term gardener and servant, a simple man who knew
of a prize, as judge, or exhibition of own produce) with Mendel exceptionally well and who used to say: “Ah my
efforts to improve cultivated plants such as flowers dear chap. Now the Lord Prelate, there was a gardener!
(Fuchsias) and, above all, fruit. With respect to the latter All the other gardeners should really take an appren-
he was particularly interested in the combination of ticeship with him” [Doupovecl.
quality and disease resistance in apples and pears or As Abbot of one of the largest of the monasteries of
their adaptation to ecologic conditions (i.e., late flower- Moravia, Mendel had to function as spiritual director of
ing) [Vavra, 1965; Orel and Vavra, 1968, Vavra and the monastic community, administrator of the property
Orel, 1971a, bl. After his death the horticultural section of the monastery together with several domains, and
honored him and his horticultural work with a detailed perform public functions. Thus, he was a member of the
obituary which also referred to his work on peas and its Moravian legislature and not surprisingly, a person-
 Meteorologische
Fig. 19. Portions of a monthly meteorologictable made out by J.G.
Mendel for the Meteorological State Station in Vienna (here August
1882).The illustration contains the head of the table with the dates of
the first 5 days of the month (first pentad). For purposes of simplifica-
tion he left off the first digit of the values for barometric pressure (the
barometer reading reduced to zero degrees-in millimeters), thus: 48.5
should be read as 748.5). In addition, the bottom portion of the table
with the sums of the data of each individual monthly pentad, the
monthly means, as well as the extreme values of barometric pressure
and air temperature, the frequency distribution of the observed wind
directions and finally summaries ofthe kind, amount, and frequency of
precipitation.
18 Weiling
1 Hr. 5. I Bai 1873. i ,%$rgung VII.
YJIitt~eiIungai ber 8ereine:Qeituiig.
Bti her oin 16. Xprif 1873 abqt.lpltcntti 81.rma[tungera14g.Bigung be8
miibrifdpn Bicnt.np$tl *%erfiite@, bci mtl@er ber $c&la6rbigftc inf. abt @regor
Fig. 20. The head of the monthly publication of the Moravian Api-
ary Society: “The Honeybee ofBrunn.”(In addition t o this German one,
a Czech edition was also published with in part individual contribu-
tions (Vcela brngnska, Brno).
ality in great demand in scientific, cultural, and general
social associations and always desired and welcomed as
a member or in a leading function. Sajner has summa-
rized no less than 8 scientific and 26 non-scientific soci-
eties and associations to which Mendel belonged and to
which he made contributions over some or many years.
[Sajner, 1971a, bl. The latter included a.0. religious,
literary, agricultural, horticultural, charitable, human-
itarian, and school associations. Here we shall cite only
the scientific organizations: the Austrian Zoological-Bo-
tanical Society in Vienna (member since 1853);the Soci-
ety for the Dissemination of Natural Historical Knowl-
edge in Vienna (as of 1873);the Naturforschende Verein
in Briinn (as of 1861; Vice-chairman, 1868-1870); the
Austrian Meteorologic Society in Vienna (since 1868,
“supportingmember” on the basis of payment of a larger
sum); and the Moravian Apiary Society (since 1870;
Vice-chairman, 1871-1873); the German Pomological
Society in Braunschweig and (as of 1873) the Pomologi-
cal Society of Reutlingen; and last but not least, the
Imperial-Royal Moravian-Silesian Society for the Pro-
motion of Agriculture and Knowledge of Land and Na-
ture (the Agricultural Society).
This society was founded in 1770 and reorganized
after the revolution of 1848;its Vice-Director (as of 1849)
and General Director (as of 1865) was Mendel’s prede-
cessor Prelate Napp at whose death more than 8,000
members belonged to the mother and the 18 daughter
associations distributed over the length and breadth of
Moravia [Weiling, 19681. After 1850 the Society com-
prised six independent sections: Forestry (founded
18421, Natural Sciences (founded 18491, Agriculture
(1849), Historical-Statistical (1850), Fruit, Vini- and
Horticultural (18501, and Apiary (1854, independent as
of 1869 as the Moravian Apiary Society).Mendel was an
active member of all five of the last-named sections.
Soon after his election as Abbot he became a member of
the parent organization (Gesamtgesellschaft) and in
1870 he was elected to its 12 member Executive Com-
mittee. In this position he was reconfirmed every 3 years
during the regular elections [Orel, 1970al. In his posi-
tion Mendel, from time to time, acted for the Vice-Direc-
tor or the General Director of the Society; however, his
most important effect was through ideas (i.e., establish-
ment of the weather forecasting system), expert consul-
tations, and through notices or reports in the organ of
the society (the “Communications”) [ Mittheilungen,
1868-1883; Orel, 1971bI.l
The late Czech law professor Vybral reported that in
his position Mendel finally got into serious political dif-
ficulties [Vybral, 19711. On the basis of land ownership
the Altbriinn chapter was represented by its prelate in
the electoral chamber of the landowners. The group in-
cluded between 160 and 170 voting members and could
send 25 of the total of the 100 delegates to the Moravian
legislature. For such positions there were candidates of
two parties. The first was the Conservative-National-
Autonomous party of the nobility interested in equality
of the nationalities to which belonged practically all
members of the hierarchy of Moravia. The second was
the German-Liberal party which was, put simply, pri-
marily concerned about the interests of the German
speaking part of the population. During the elections for
the Moravian legislature of 1870/1871Mendel voted for
the German-Liberal (also called the constitutional)
party. He continued to express his preference for that
party even after the results of the election (which de-
cided in favor of the conservative party) were being
contested. This put the then still junior prelate Mendel
into an exceedingly uncomfortable position since he had
made enemies of several senior clerics and other person-
alities. This steadfast action on behalf of the constitu-
tional party on the part of the one “cleric among those
entitled to represent the class of the large land owners in
exercising their electoral right for the Moravian Legis-
lature” was the occasion for the Governor of Moravia,
Count von Thun, to propose Mendel 21/2 months after
the last election (February 28, 1872) as grand com-
mander of the Order of Franz-Josef. These circum-
stances are well documented by Richter [1943] on the
Mendel wrote numerous pieces for the “Communications.” His
authorship was identified as “M”, “m”, or “ G M . They are predomi-
nantly on agricultural innovations or harvest reports. Occa-
sionally he wrote a longer piece. One of these is Mendel, 1879. In
view of the recent peaceful cultural revolution in Czechoslovakia, a
three-page essay “Against Communism and Socialism” is of inter-
est [Mittheilungen, 1877, p 6-81; for obvious reasons it was for-
merly little noted in that country. After a brief introduction “M’
asks the rhetorical question: “How do we keep the teachings of
communism and socialism away from our rural workers?” in order
to introduce a n essay that had appeared earlier in the Luneburger
Agricultural News (Luneburger Landwirtschaftliche Zeitung).
After a description of the historical development of such political
views the writer of the essay recommends the building of homes for
the workers with a piece of land for personal agricultural use as
well as the establishment of vocational schools or training centers
with obligatory attendance between age 14 and 17 years.
 Mendel 19

basis of the award documents. Indeed, Mendel received grown some 2,500 F, Pisum plants which, according to
the Komturkreuz of that Order as early as March 21, Rasmusson’s estimate [personal communication],would
1872 [Kfiienecky, 19651. have occupied about half of his garden plot. We may
Especially grave were the consequences of Mendel’s further assume from his letter to Nageli of April 18,
position when in May 1874 the very party he favored 1867 that during 1865 and 1866 he gave the remaining
initiated the legislation which, a.0. levied a church tax seeds from his published Pisum experiments to other
to cover the needs of the Catholic religion in Austria. investigators in an effort to stimulate “confirmatory
The conflicts during the legislative debate, the burden experiments” [Correns, 19051. Correns did not include
to be imposed on his chapter and the position ofthe other the highly informative eight-page list for Nageli of the
members of his chapter exerted an extraordinary emo- 147 types of seeds from 1864 in the publication of the 10
tional pressure on Mendel which did not leave him be- letters of this correspondence,but rather gave it as a gift
fore the end of his life and which made him a decisive in 1920 t o the Swedish Mendel Society a t the occasion of
opponent of that law. To the end he sought with all his its 10th anniversary. The Society started volume 100 of
strength to invalidate the law and stubbornly refused to its journal Hereditas with this list in facsimile form and
obey the State’s demand for payment of the imposed tax thus made it accessible again to the world of science (Fig.
with the consequencethat the government put a lien on 21). In his own words, as of 1872 Mendel had to “neglect
the income of the monastic domains. completely” his experimental work with plants and to
During this impasse the Moravian German-Liberal cease the genetic analyses requiring several years [Cor-
party was evidently at pains to mediate a financial rens, 19051, in part probably because of the oppressive
compromise by offering Mendel a public position. Thus, political situation. Nevertheless, the intellectual con-
in April 1876 Mendel was elected by the Moravian Leg- frontation with the segregational behavior of his plants
islature Vice-Director and member of the Board of continued repeatedly after this point in time.
Trustees of the newly founded Mortgage Bank of Mo-
ravia; in October 1881 he was elected Director of the
Bank. According to Vybral, this solution may have been
influenced by Josef Auspitz (1812-1889) the former
Principal of the Oberrealschule and Mendel’s long-time
superior in the school system [Vybral, 19681. In 1869
Auspitz had been appointed Inspector General of the
school system; he resigned 2 years later. As of 1873 he
devoted himself exclusively to newspaper work a t the
Mahrische Tagesbote whose editorship he assumed in
1879. Politically, he was a leader of the German-Liberal
party of Moravia. He discovered Mendel’s pedagogic
gifts early on and in spite of lacking board certification
was able to maintain Mendel as supplementary Pro-
fessor in the Oberrealschule. In addition, he was proba-
bly one of the few persons in Brunn who recognized the
fundamental importance of Mendel’s experiments as is
evident in the detailed obituary in the Tagesbote whose
author was probably Auspitz himself [Weiling, 1984dl.
As noted above, except for one additional cross of peas Iu‘

with differential flowering time, Mendel finished his

Pisum experiments in 1863 “in order to test through
experiments with other plants the experiments gained
with Pisum.” However, he had learned from the crosses
performed for this purpose in 1863-1864 “that it may
not be easy t o find plants suitable for a comprehensive
series of experiments, and that in an unfavorable case
years may pass without the desired results” [Correns,
19051. On the basis of his correspondence with Nageli
which Mendel discontinued after his 10th letter of No-
vember 18,1873 (without answering two further letters
of Nageli from 1874-18751, we may conclude with Get1
[1971] that these preliminary experiments involved Fig. 21. Page 5 of the eight-page enumeration sent by Mendel to
crosses between (at least) 34 species of 14 different iden- Professor Nageli with the 147 bags of seed for purposes ofverification of
tified plant genera. In his second letter Mendel provided the results of his Pisum experiments. This page contains the “ge-
notype” designations of Mendel of 24 Fz plants (one plant to a bag of
Nageli with seeds (peas) from six different parental seeds) from the tetrahybrid cross bcDg x BCdg together with the indi-
plants and from an additional 141 F2plants from eight cation that in addition to the yellow seeds in bag no. 976 the green ones
different crosses of his 1864 material for control pur- ought to yield pure (constant) offspring with all four traits. The charac-
ter traits involve: B, b: cotyledon colors (yellowor green); C, c: seed coat
poses. The numbers of these F, plants ranged from 90 to color (brown or white); D, d: pod form (even or constricted); and G, g:
2,171. We may therefore assume that in 1864 he had stem length (long or short).
20 Weiling
Only a few pages of notes escaped the destruction of
the unbound material after Mendel’s death; these in-
clude a page with genetic notes (Fig. 22). It is frequently
reproduced and has been discussed and analyzed by
several authors. Richter [1924, 19431 considers the
figures segregation data of a di- or trihybrid cross in-
volving a blue or violet-floweredplant probably the cross
Linaria vulgaris x Linaria striata mentioned in his cor-
respondence with Nageli. However, Frimmel [19241
thinks these data refer to flowers or seeds of Lathyrus
odoratus. R.A. Fisher interprets the data (343:92:166)
as a 9:3:4 relationship between two interactive genes
with recessive epistasis of an F, generation involving
seed color presumably of beans [Darlington and Mather,
19501. Lamprecht is also of the opinion that these data,
which are in good agreement with his own segregational
data, represent segregations of seed color of Phaseolus
vulgaris [Heimans, 19681. However, Olby, largely fol-
lowing the views of Fisher, thinks the data more likely
relate to segregation in L . vulgaris x L. striata. In the
second edition of his book he cites the opinion of Hei-
mans which, however, does not seem entirely convincing
to him [Olby, 19661. Heimans seems to demonstrate
with, to me rather convincing arguments, that these
data refer to the trihybrid experiment by Mendel with
peas [Heimans, 19681. This experiment is based on the
seed forms roundlwrinkled, the seed colors yellowlgreen,
and the graybrownlwhite colors of the seed coat. In his
detailed descriptions of traits accepted for experimenta-
tion Mendel described the color of the seed coat, in case it
was not white, as “gray, graybrown, leather brown, with
or without violet spotting.” It is evidently this differen-
tial pigmentation of the seed coat that Mendel refers to
when he writes of the 687 seeds of his 24 F1hybrids that
their seed coats “were all spotted graybrown or gray-
green” [Mendel, 1865a,bl.Indeed, the number 166 ofthe
F2plants with white seed coat of his publication agrees
with that of the note sheet. However, the number of the
other forms is too low by 38 (343 + 92 = 435 [note sheet],
contrasted with 639 - 166 = 473 fertile plants with non-
white seed coat in the trihybrid experiment). Heimans
deals with this difference by pointing out that in his
supplementary attempt to explain genetically the evi-
dently non-uniform appearance of the non-white seed
coat, Mendel had to base himself on the labels on his
seed bags, some of which may have been lost by then
[Heimans, 19681. At this point we shall refrain from
further consideration of the question on how to deal with
the missing 38 entities indicated in the figures on the
note sheet.
Only Richter [1924,19431 and Heimans [19681 have
considered the not unimportant question of when it was
that Mendel made these genetic notes. Both refer to
notes Mendel made on the tax problem on the same
sheet of paper. Since they contain reference to the tax
law of May 7, 1874 with citation of this date, they con-
clude that the genetic notes have to be dated after this
date. Heimans adds that the tax-related notes may refer
to a memorandum Mendel submitted to the Government
on August 15,1876, so that the notes have originated in
the latter year. However, the question remains, as also
for Richter and Frimmel, whether the genetic notes were
made before or after the remarks pertaining to the tax
problem.
Contradictory data by Richter and Heimans pertain-
ing to the size of this note sheet and on the placement on
the sheet of the genetic and tax notes caused me to
request a copy of the front and back sides of the sheet
from the Division of Genetics of the Moravian Museum
in Brno. I am most grateful for the prompt response to
my request.
This sheet measures 3 3 . 3 ~ 2 1 . 0cm and contains
notes on the tax problem on both sides. On one side
(front) these notes occupy half, on the other about 3/4 of
the sheet. At a certain point of time the sheet was folded
in half creating a blank portion from the bottom half of
the front and was covered at right angle to the tax notes
with genetic notes (Fig. 22). The sheet evidently repre-
sents a draft with a false beginning of an official letter
by Mendel. This is evident from the 314 covered (back)
side of the sheet which is written hastily with lead or
copy-pencil. Apparently no one has ever noticed before
that between lines 12 and 13 ofthis text there is inserted
a line carefully written in ink: (‘Zu der am 26 Juni
abgehaltenen Baum.” These words evidently refer to a
meeting held on June 26 of an unknown year that had to
do with trees or fruit trees. Thus, it was not difficult to
determine on hand of the short report in the Proceedings
of the Moravian-Silesian Agricultural Society [Mit-
theilungen, 1868-18831 concerning the monthly meet-
ings of its Executive Committee, of which Mendel was a
member, that the note referred to an examination for
fruit tree inspectors held that day under Mendel’s
commissary supervision and reported a t the session of
the Society of July 7,1880 and in its Proceedings. After
further inquiries to the Genetics Division of the Mora-
vian Museum in Brno with the request for an additional
look into the State Archives, I received shortly after-
wards a copy of a six-line letter of Mendel (Fig. 23) which
begins with the same words together with the protocol
concerning the evaluation of this examination at the
session of the executive committee of July 7,1880. From
this it can be concluded that Mendel made the genetic
notes only some time after June 26, 1880 and after he
drafted the tax notes on both sides (one side probably not
being a continuation of the other). In substantial agree-
ment with Heiman’s interpretation, it seems obvious
that 15 years after completion of his Pisum work and
some 3 years before his death, Mendel was still inten-
sively involved with an unsettled aspect of his work,
namely the possible determination of the color of the
seed coat of his peas by two interactive “factors.” From
the copy of this sheet it is further evident that after it
was folded to serve as free space for the genetic notes, the
upper half stuck out a bit and was trimmed resulting in
loss of the upper portion of a line of text. It also shows
that the crease that resulted from folding the sheet is
torn over a length of about 13 mm at the right lower
corner and that the portion of the sheet with the genetic
notes is slightly rolled at that point (Fig. 22) suggesting
that Mendel held the sheet frequently in his right hand
while pondering the segregation ratios [Weiling, 19901.
These conclusions are supported by the report of the
Abbot Franz Sal. BaEina (1863-1943) who was received
 !
LA ----I ..__
.-..-..-.._. -. ....
A_ .. ... - - . . .-.. . .. . .. .

Fig. 22. Mendel’s“genetic note sheet.” (Reducedt o 0.8 of the original).This shows that the genetic notes were written on the free space of a
larger, legal sheet (for details see text).

into the order by Mendel 3 months before his death, that
up until his death Mendel had been full of confidence
that: “the time will come when the validity of the laws
discovered by me will be recognized” [Richter, 19431.
This confidence as well as the ongoing preoccupation
with his discovery during the last years of his life evi-
dently were so strong that conscious of his impending
death and contemplating his past life, Mendel spoke the
following sentences at the occasion of the vesting of the
latest novices:
“I have experienced many a bitter hour in my life.
Nevertheless, I admit gratefully that the beautiful, good
22 Weiling

Fig. 23. Upper part of the ’%back”of the genetic note sheet with the words: ‘Zuder am 26.Juni abgehaltenen Baum” between lines 12 and 13 of a
legal draft pertaining to the tax problem. Below that the corrected complete statement beginning: “Zu der am 26. Juni abgehaltenen Priifung fur
Obstbaum-Warter . . . ” (with respect to the exam given on 26 June for fruit-tree inspectors . . . ”) found in the State Archives of Brno in the files of
the Moravian-Silesian Agricultural Society (minutes of the session of the Executive Committee of July 7, 1880). In his first try Mendel had
probably intended to use the collective concept “Baumzucht” (tree cultivation, q.v. Fig. 6 ) when be became aware that the examination referred
solely to the part involving fruit trees. Mendel reports that two candidates Clemens Schuppler and Emil Zapletal presented themselves for the
exam. “Both absolved the exam completely satisfactorily with the grade ‘very useful.’ ”

hours far outnumbered the others. My scientific work
brought me much satisfaction and I am convinced that
the entire world will recognize the results of these
studies” [Nachtweh, 1922; Richter, 1924, 19431.
From these two sentences one can conclude that his
education, the planning and execution of his “epochal”
experiments [Anonymous, 1884; Weiling, 1984d1, and
the discovery and late analysis of the laws stamped
Mendel’s life in an essential manner.
Subsequent developments showed Mendel was right.
However, it was impossible for him to foresee a further
“blow of fate.’’ Because when R.A. Fisher determined
that Mendel’s results showed a greater-than-expected
agreement with expectation on the basis of the Chi
square test developed since then [Fisher, 19361,Mendel
was accused by uncritical journalists and scientists of
knowingly falsifying data [Weiling, 19851. However, in
the meantime the basis for such accusations has been
removed since it was determined that the statistical
model underlying the Chi square test is inappropriate
for the conditionsunderlying Mendel’s segregation data,
so that Fisher’s conclusion does not apply [Weiling,
19891. The following brief remarks must suffice:
Mendel’s ratios are not random samples of adequately
large and thus binomially distributed totalities as pre-
supposed by the Chi square test but rather the sum of
several or numerous ratios based on single plants (in the
case of seed characteristics) or ratios of offspring (in the
case of plant characteristics). The data of each single
plant-segregation and of each descent-segregation are
in their turn the sums of several individual segregations
based on the segregation within the seedpods of single
plants. And this segregation occurring in the seed pods
or ovaries of individual plants is in its origin not bino-
mial but decidedly hypergeometrical. In this connection
it must also be noted that the figures reported by Mendel
for individual experiments are not gross but net totals
since they also reflect losses through abortion of zygotes,
failures to germinate or for a plant to reach reproductive
maturity, and even selection on the part of the investiga-
tor (as in case of 2: 1 ratios).
All of this results in the fact that on one hand the
variance of the reported summarized segregation ratios
of Mendel’sactual experiments is reduced in comparison
with the variance of the segregation compounds (the
single basic ratios) and also in comparison with a ran-
dom binomial sample of that reported totality i.e., if the
ratio of the reported totality really would be distributed
binomially). On the other hand Mendel’s data are re-
sults of completely different experiments: segregation
data from a first, a second, or a third generation etc.,
which all derived from an initial cross in a P-generation.
And these ratios data are diminished by the above-
named and sometimes recurrent different losses (in part
significantly) different in their variance. However, this
fact makes it impossibleto add the Chi square values for
the different experiments on the results on which Fisher
based his judgement.
Fisher’s objection to the supposed greater-than-
chance fit between observed ratios and expected num-
bers is therefore based on the assumption that Mendel’s
data are ratios of random binomial samples while they
Mendel 23
represent in fact net sums of initially larger numbers of
individual segregations of a hypergeometrical type in
the fertilized seeds diminished by zygote, seedling,
growth, orland other losses.
Fisher 119361 makes the assumption that 5.63% or
0.751° of the samples of 10 plants of each a heterozygous
offspring will be classified as homozygotes. He calcu-
lates this probability using a binomial model (i.e., based
on samples from infinite totalities) while the number of
seeds of Mendel pea plants was on the average 30. There-
fore it cannot be excluded that he took his 10 seeds only
from a part of the pods of a plant (i.e., from seed numbers
less than 30) [Weiling, 1989 and papers there cited).
Nevertheless, it is possible to show in Mendel’s data
that the variance of the segregation of individual plants
and individual offspring corresponds approximately to a
binomial distribution, so that its Chi square values can
be regarded as approximately free of bias. In other
words, a too-good agreement between observation and
expectation is seen only when the data of several single-
plant segregations or segregations of individual off-
spring (or their Chi squared values) are added.
ACKNOWLEDGMENTS
I am grateful to the staff of the University Libraries of
Brno, Prague, and Olomouc in Czechoslovakia; of the
University Library and National Library in Vienna,
Austria; of the State Archives in Brno, Vienna, and
Olomouc; of the City Archives of Brno, and especially of
the Genetics Division of the Moravian Museum (Men-
delianum) in Brno for their active support over the last
25 years of my investigations of the background and life
of Johann Gregor Mendel and his scientific work. This
paper was translated from German to English by J.M.
Opitz. It was typed by Vicki L. Smith.
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~ Which points~are incorrect in R. A.
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