Theory Biosci.

DOI 10.1007/s12064-016-0236-8

ORIGINAL PAPER

Armin von Tschermak-Seysenegg (1870–1952): Physiologist

and Co-‘Rediscoverer’ of Mendel’s laws
Michal V. Simunek1 • Michael Mielewczik2 • Georgy S. Levit3,4 • Uwe Hossfeld3,4

Received: 5 September 2016 / Accepted: 29 September 2016

 Springer-Verlag Berlin Heidelberg 2016

Abstract The ‘rediscovery’ of Mendel’s laws in 1900 was Not only the independence but also parallelism was anal-
a turning point in modern research of heredity/genetics. ysed in the context of individual research programmes of
According to the traditional view, adopted and fostered by these three scholars. The youngest of them, Erich v.
many textbooks of genetics, Mendel’s principles were Tschermak-Seysenegg, was even excluded from the list of
presented in the first half of 1900 simultaneously and ‘rediscoverers’. The aim of this paper is to use new
independently by three biologists (H. de Vries, C. Correns, archival evidence and approximate the contribution of the
E. v. Tschermak-Seysenegg). Having thus laid the foun- physiologist and ophthalmologist Armin von Tschermak-
dations of further development, the ‘rediscovery’ continues Seysenegg (1870–1952) to the events of 1900 and 1901.
to attract considerable interest. Since the 1950s, however,
serious questions arose concerning both the chronology and Keywords Mendel’s Laws
‘Mendelism’
‘Rediscovery’

specific conceptual achievement of the scientists involved. Tschermak-Seysenegg v. Armin & Erich

This article is dedicated to the memory of Olaf Breidbach—inspiring

thinker, tireless researcher, and colleague. Introduction: in the shadow of Mendel
This article forms part of a special issue of Theory in Biosciences in The term ‘rediscovery’ is frequently used for the events of
commemoration of Olaf Breidbach.
1900, when the experimental verifications of some of
& Georgy S. Levit Mendel’s conclusions were announced publicly by three
[email protected] European botanists: Hugo de Vries (1848–1935), The
Michal V. Simunek Netherlands; Carl Correns (1864–1933), Germany; and
[email protected] Erich von Tschermak-Seysenegg (1871–1962), Austria-
Michael Mielewczik Hungary (Johannsen 1923; Iltis 1924; Roberts 1929; Grant
[email protected] 1956; Dunn 1965; Stubbe 1965; Johansson 1979; Keller
Uwe Hossfeld 2000). The ‘rediscovery’ of Mendel’s laws resulted in the
[email protected] growth of ‘Mendelism’, which was seen by contemporaries
1
as a subfield of genetics (e.g. Plate 1932; more about
Centre for the History of Sciences and Humanities, Institute
of Contemporary History of the Academy of Sciences,
Ludwig Plate in Levit and Hossfeld 2006).
Prague, Czech Republic For each protagonist, two principle questions arise:
2
Imperial College London, National Heart and Lung Institute
First—whether they discovered the principles on their own
Hammersmith Hospital, ICTEM Building, 3rd Floor Du Cane or used Mendel’s papers to interpret previously obtained
Road 2, London W12 0HS, UK results. Second—whether each of those scholars made the
3
Working Group Biology Education, Friedrich Schiller discovery independently from each other. Especially the
University of Jena, Am Steiger 3, 07743 Jena, Germany latter question touches upon the parallelism of the discovery,
4
ITMO University, Chaikovsky Str. 11, and, eo ipso, also later claims of priority and glory.
191187 Saint Petersburg, Russia Depending on the answers and the level of analysis, events of

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1900 might be seen not as a ‘rediscovery’ but rather as a ‘new between the two brothers, comprising as a whole 87 items
discovery’ (Jahn 1957), ‘delayed discovery’ (Zirkle 1964), of letters, correspondence cards, postcards, and telegrams.
or even a complex of ‘multiple discoveries’ (Brannigan et al. They cover the period from 1898 until 1951 (the year
1981). Issues pertaining to independence, originality, and before the death of A.T.S.). The vast majority of the extant
parallelism are thus reflected in various interpretations of the correspondence was written and sent from A.T.S. to E.T.S.
‘rediscovery’ (Roberts 1929; Stomps 1954; Jahn 1957; Dunn (83 items): only four items conversely. There are 14 pieces
1965; Stubbe 1965; Sturtevant 2001; Zirkle 1968; Olby of correspondence sent by A.T.S. to E.T.S. in the period
1985; Bowler 1989). Furthermore, given the evolving nature from March 13th, 1898, until November 19th, 1901 that
of experimental research programmes of the protagonists, it were transcribed and published recently (Simunek et al.
remains unclear what should be the extent of the period 2011a, b; Simunek and Hossfeld 2011, 2012).
covered by the term ‘rediscoverery’ (Jahn 1957). Although the mutual correspondence is clearly incom-
For a long time, Erich v. Tschermak-Seysenegg (here- plete, the information contained can help to provide more
inafter E.T.S.) had been seen as one of the ‘rediscoverers’ detailed answers to questions pertaining to their degrees of
(Johannsen 1923; Nilsson-Ehle 1924; Roemer 1941; Stubbe success in interpreting Mendel’s principles in the crucial
1941; Reinöhl 1950; Gasking 1959; Heinisch and Rudorf period of 1899–1901.
1961; Simunek et al. 2009). Some contemporaries even called An until recently unknown fact is the involvement of
him ‘‘… the very first Austrian scientist after Mendel who A.T.S. in the events of the rediscovery of the Mendelian
understood the rules of heredity’’ (Hänsel 1962: 13). It is also laws. Despite some very clear statements of E.T.S. and the
relevant that E.T.S. lived until the early 1960s. He supported assumption that the two brothers were very close (Harwood
the position of a triple parallel (simultaneous) rediscovery 2000: 1062), the extent, quality, and significance of their
already in his first papers (Tschermak-Seysenegg 1900a: 239; relationship for the early Mendelian studies remained
Tschermak-Seysenegg 1900b: 555), and consistently pro- unknown. However, it is certain that A.T.S. supported his
moted his view also in later publications (Tschermak-Sey- brother in promoting the narrative of the triple simultane-
senegg 1901b: 642; Tschermak-Seysenegg 1901d: 1029, ous ‘rediscovery’ from the very beginning and though he
1908, 1928, 1956, 1958). Implying the identity of discoveries always excluded himself from the group of Mendel’s ‘re-
in 1865 and 1900, he even mentioned ‘‘(…) a 35 years long discoverers’, his interpretations and comments may have
interval’’ when comparing in 1931 the original publication of played a crucial role in shaping Erich’s published views
Mendel’s paper and the events of 1900 (Tschermak-Sey- (Tschermak-Seysenegg 1901a, 1923).
senegg 1931: 1). Last but not least, he saw himself as a leading On a theoretical level, Armin’s contribution concerned
contributor in applying the Mendelian knowledge in the field both special issues of plant cytology (Simunek et al. 2011a,
of plant breeding (Wunderlich 1951; Harwood 1997, 2000). p. 49–50) and a general interpretation of experimentally
However, in the mid-1960s, concerns were raised, highlight- obtained results. His support of the younger brother was
ing that his original understanding of Mendel’s explanations especially important when it came to mathematical solu-
in the context of 1900 had a limited validity both in conceptual tions: ‘‘… our numerous discussions concerned primarily
and terminological terms (Dunn 1965: 76). He was dropped my work and various issues of heredity. When it came to
from the list of the ‘rediscoverers’ (Stern and Sherwood mathematical problems, I was quite dependent on his [Ar-
1956, 1978; Bowler 1989). Recently (e.g. Moore 2001), it has min’s] guidance, since that was a field where I invariably
been argued that he failed because (1) he was not able to draw failed since my earliest youth’’, stated for example by E.T.S.
any generalizations from the data he obtained, and (2) he did later in the unpublished version of his memoires.2 In
not attempt to explain the results he obtained by discovery another part of his unpublished memoires, E.T.S. even
within an appropriate theoretical framework. specified: ‘‘He read, improved, and completed almost
In the summer of 2009, a personal collection of E.T.S. everything I wrote since the age of 28 until the age of 79,
held by the Archives of the Austrian Academy of Sciences and always insisted on reading with me the proof sheets.‘‘
in Vienna was catalogued and opened for historical With few exceptions, the existing historiography of
research. At the same time, a significant part of personal genetics contains very limited information about this
possessions of Armin v. Tschermak-Seysenegg (hereinafter important representative of continental physiology and
A.T.S.), his older brother and a prominent physiologist, medicine in early and mid-20th century. A scientist, who
was identified.1 Both collections contain correspondence was especially during his stay in Prague from 1913 until
1945, deeply connected to a prominent tradition in conti-
1
The collection was part of the family archives of A.T.S.’s grandson nental European life sciences, established in the Bohemian
Dr. Armin Tschermak von Seysenegg Jr. The only two persons who
went through the correspondence in the 1980s and 1990s were Armin
2
von Tschermak-Seysenegg’s son, Wolfgang Tschermak von Sey- Collection of A.T.S. Stuttgart, transcription of E.T.S.’s manuscript
senegg, and his son, Dr. Armin Tschermak von Seysenegg Jr. ‘Mein Bruder Armin’ [My Brother Armin], p. 25.

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capital already by Jan (Körting 1968, 120–1; Maaß 1971, 1911, he was at the age of forty-four appointed a court
60, 81). counsellor (Hofrat), thus becoming the youngest person in
the whole Habsburg monarchy who at that time held this
title. In the meantime, he married in Berlin on June 6th,
Life and professional career 1911, Ilse Charlotte Penck (1886–1951), daughter of
(Friedrich) Albrecht K. Penck (1858–1945), well-known
Armin (Eduard Gustav) was born in Vienna on September professor of geography and geology in Berlin. Armin and
21, 1870, as the second child of Gustav Tschermak Ilse had four children (one daughter died prematurely)
(1836–1927), professor of mineralogy, and his wife Her- (Fig. 1).
mine, born Fenzl. She was a daughter of Eduard Fenzl On May 28, 1913, A.T.S. was appointed here a full
(1808–1879), an Austrian botanist and examinator of G. professor of physiology effective as of June 1, 1913, where
J. Mendel in Vienna. he followed in the footsteps of Ewald Hering. In the end, he
Armin had an older sister, Silvia (1868–1923), and the stayed in Prague until 1945. Tschermak’s strongly pro-
already mentioned younger brother Erich. Silvia (married German feelings surfaced during the WWI, when in 1915
Hillebrand) was a bacteriologist and an active scientist in he volunteered for military service in the Austro-Hungarian
her own right. For example, in 1918–19, she shortly army in the rank of assistant physician. First, he served in
worked as a laboratory assistant under Karl Landsteiner the 2nd army, later in the Emperor Franz Joseph I Guards.
(1868–1943), the discoverer of the blood groups A, B, AB, He was also deployed with the group of the Knights of
and 0 from 1909. Malta, 1st surgical unit in Vielgereut and Malga Belem,
The relationship between the brothers remained very Tyrolia, and later in Dolina, Galicia. During his military
close throughout their lives. When their father retired service, he distinguished himself mainly as a capable sur-
(1906), he was knighted and chose the aristocratic pred- geon but also played an important role in advocating a
icate ‘Edler von Seysenegg’. In 1881–98, A.T.S. attended reform of Austrian military healthcare. After the end of
the Catholic gymnasium in Kremsmünster, Upper Austria.
In October 1889, he started his medical studies at the
Viennese university. In the summer term of 1893 and
winter term of 1893–94, he stayed in Heidelberg, Ger-
many. A.T.S. eventually graduated from the university in
Vienna on July 1, 1895, whereby he became a doctor of
general medicine. Later, he also qualified as a general
practitioner.
Already early on, A.T.S. decided to aim at an academic
career. He started by working for a few months as a
demonstrator at the Institute for General and Experimental
Pathology under Salomon Stricker (1834–1898). Then, he
transferred as a volunteer to the Institute of Physiology of
the Royal Saxonian United Friedrich University in Halle-
Wittenberg under the supervision of the famous physiolo-
gist Ewald K. K. Hering (1834–1918). On January 23,
1899, based on his work ‘Colour Sense in Indirect Vision’
he was named Privatdozent. In 1900–06, he worked as
assistant of Julius Bernstein (1839–1917). In 1902, he
undertook a short study trip to Sankt Petersburg, Russia,
where he collaborated with Ivan P. Pavlov (1849–1936).
Later on, he also stayed at marine station for zoology of
Anton Dohrn (1840–1909) in Naples, Italy. On August 1st,
1903, A.T.S. was named extraordinary professor. In early
1906, he accepted a chair at the Veterinary University
(Hochschule) in Vienna. Though his primary interest was
in human physiology, he there worked as a full professor of
physiology and veterinary physics. In 1908, he served as
the managing pro-rector of the school, and in the following Fig. 1 The siblings Tschermaks, app. 1880s (Photo Archive Michal
year, he became its first elected rector. On November 5th, Simunek, Prague)

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WWI and creation of Czechoslovakia, he returned to Pra-

gue, where he opposed the new development and deman-
ded the establishment of a German University in Prague
and was a spiritus movens of several incidents of the rad-
ical German circles (Hoensch et al. 1999: 120–122;
Osterloh 2006: 110–111). Nevertheless, he remained in
Prague and went on to establish his own school (among his
colleagues or students we find, e.g. Richard Kahn, Eugen
Steinach, Steffan Jellinek, and Viktor F. Hess, the winner
of Nobel Prize in physics). He was an important member of
the local German scientific community. In 1920–21 and
then again in 1925–26, he served as Dean of the German
medical faculty in Prague. In 1929, he became the chair-
man of the newly founded society, which later became
known as ‘‘Deutscher Verein für Familienkunde und
Eugenik in der Tschechoslowakischen Republik’’, in which
he published together with several of the main represen-
tatives of the early Sudeten German eugenics movement
such as Bernhard Brandt, Friedrich Breinl and Otto Grosser
in a yearbook on genealogy, genetics and eugenics
(Simunek 2015) (Fig. 2).
A.T.S. retired at the end of February 1939 with various
honours. Nonetheless, after the German occupation of
Bohemia and Moravia and transfer of the Prague German
University under the competence of the Reich Ministry of
Science, Education etc. (Reichsministerium für Wis-
senschaft, Erziehung und Volksbildung), A.T.S. returned
from retirement and headed the institute de facto until the
end of the WWII in May 1945. His continuing academic
career in the Third Reich was doubtless aided by his
political conformity. In summer 1942, at the age of
72 years he volunteered and joined the German Navy
(Kriegsmarine). He served first as chief staff surgeon, later
as squadron physician and advisor for optics of Kriegs-
marine in Kiel, Saßnitz, and Swinemünde, where he took
Fig. 2 Armin von Tschermak-Seysenegg as an extraordinary profes-
part in the research on night vision. In 1946, however, he
sor in Halle, 1901 (Photo Archive Michal Simunek, Prague)
decided to officially move to Straubing, Bavaria. Despite
his advanced age, he tried to get involved in Bavarian
academic life, and advocated the establishment of another issues of electric phenomena in living organisms, questions
university in Regensburg. In consequence of very stressful of pre-existence of electric charge in body organs,
negotiations, he suffered a brain haemorrhage on February including intact ones, issues of relation between the size of
9, 1950. He died two years later, during a regular walk on a muscle and the magnitude of the electric process, but also
October 9th, 1952, after suffering another stroke. for example the thermodynamics of electric discharge in
He was interested in the physiology of muscles and the particular case of electric ray (Maaß 1971, 62–5).
nerves (e.g. the so-called Tschermaksche Druckversuch, Another large area of A.T.S.’s interest was connected with
that is, Tschermak’s pressure test), bio-electrics and psycho-physiology, a scientific discipline whose foremost
physiology of the brain, but also in general physiology, in representative was another important teacher of him, Ewald
particular in physiological optics (physiology of the eye Hering. Continuing in his footsteps, A.T.S. focused mainly
and sight), and even in issues of physics and chemistry as on the notion of a living substance surviving under con-
well as cellular physiology and biochemistry (digestion ditions of two balanced contrasting phenomena (so-called
enzymes and fermentation, generation of heat during assimilation and dissimilation), which led to his work on
enzymatic splitting, etc.) (Maaß 1971, 62–5). Tschermak’s tonic innervation of autonomous organs. In somewhat
first teacher, Julius Bernstein, attracted him especially to related line of research, A.T.S. also studied the general

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physiology of senses and ophthalmic physiology. In these Saale or the Section for Mathematics and Natural sciences
areas, he focused on sight and colour vision—where he was of the German Society for Science and Art for the Cze-
inspired by the Purkinje phenomenon —but also on the choslovak Republic in Prague. As a devout Catholic, he
stereoscopy of colours, colour blindness, mutual interfer- was especially honoured when, as one of just a handful of
ence of colours, photosensitivity, night vision, etc. (Maaß German scientists (e.g. M. Planck, C. Delouge, E. Abder-
1971: 81). A central place among several hundreds of halden, E. Schrödinger, and E. F. Petritsch), he was in 1936
original papers belongs doubtless to his synthetic work offered membership in the Vatican academy Academia
‘General Physiology’ (Tschermak-Seysenegg 1916–24) Pontificia de las Ciencias.
(Fig. 3).
During his long and uncommonly prolific scientific
career, A.T.S. won numerous awards and distinctions. In Hereditary/genetic issues
1911, he received the Japanese Meiji Imperial Cross of
Merit together with the badge of 2nd class of the Order of In the area of hereditary studies, A.T.S. was among the
the Rising Sun (Commander), he was named Grand Officer pioneers of a line of research that was inspired by Gregor J.
of the Order of the Crown of Italy with a star, Knight Mendel’s studies from 1850s and 1860s, which after 1900
Commander of the Order pro Merito Melitensi with a cross, became known as ‘Mendelism’ (Simunek et al. 2012;
Grand Officer of the Yugoslav Order of St. Sava with a Hossfeld et al. 2015a, b, 2016).
star, and received various Austrian distinctions. He was a The mutual collaboration with his brother started
member of many scientific societies, such as the German already before 1900. In March 1898, A.T.S. encouraged his
Academy of Natural Scientists (Leopoldina) in Halle an der younger brother to write his second thesis (Habilitation) in
botany with the clear goal of pursuing an academic career
(Simunek et al. 2011a: 39–40). As a research topic, he
suggested the phenomenon of doubleflowered varieties and
the study of natural and artificial processes such as the use
of irritant liquids (organic acids) that may be used to
stimulate this phenomenon. At that time, there already
existed a large number of both original papers and reviews
on this topic (Masters 1886; Penzig 1890–1994). Interest-
ingly, A.T.S. also brought to his brother’s attention the
possibility of appearance of double-flowered varieties due
to the influence of parasites such as nematodes. This
research direction was quite unique at that time because
reasons behind the phenomenon were still unknown
(Korschinsky 1901). A.T.S. advised his brother on the
methodology of research, offered help with preparation
techniques, and suggested the assistance of Adolf von
Liebenberg. During the following year, 1899, he encour-
aged his brother to focus primarily on his habilitation (2nd
degree in Germany, Dr. sc.), and suggested they would
meet in the city of Halle, Germany, where he stayed, to
discuss further details (Simunek et al. 2011a: 41).
The first extant letter on hybridisation was written on
April 4th, 1900, after the brothers spent the Christmas sea-
son of 1899/1900 together in Vienna (Simunek et al. 2011a:
42–5). At this time, A.T.S. translated and reviewed for his
brother de Vries’s notes Sur la fécondation hybride de
l’albumen [On the Hybrid Fertilisation of the Albumen]
published in the end of 1899 in the Parisian journal Comptes
Rendus de l’Académie des Sciences (De Vries 1899). He
also recommended to Erich’s two further papers authored by
Hugo de Vries and Carl Correns (Correns 1899; De Vries
Fig. 3 Armin von Tschermak-Seysenegg giving his lecture at the
Institute of Physiology of the Faculty of Medicine of the German 1900). Although he focused primarily on Sergej Nawa-
University in Prague, 1920s (Photo Archive Michal Simunek, Prague) schin’s and Jean-Luis Guignard’s theories of fertilisation,

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the connection to hybridization experiments was apparent. wrote on February 18th, 1901, about the collaboration with
Last but not least, A.T.S. explicitly discussed de Vries’s his brother, and mentioned explicitly that some corrections
knowledge (or ignorance) of ‘Mendel’s teaching’. Com- Erich adopted had the effect of ‘‘… saving us from disgrace
menting on de Vries’s experiments with xenia in Zea Mais, in front of that vicious Correns’’ (Simunek et al. 2011a:
A.T.S. wrote at the beginning of April 1900: ‘‘But he [de 53–57). On March 10th, 1901, A.T.S. welcomed the reprint
Vries] doesn’t know the teaching of Mendel! Ha! Ha!! But of Mendel’s original papers (Mendel G. J./Tschermak E.
please don’t offend him: He will need to learn it from you!’’ 1901), which he perceived unambiguously as offensive to
(Simunek et al. 2011a: 42–5). More than ten days later, on Correns (Simunek et al. 2011a: 56–8). On May 18th, 1901,
May 16th, 1900, A.T.S. first commented on the corrections he helped Erich with the question of whether endosperm
of Erich’s thesis (Simunek et al. 2011a: 46–8). At that time, fertilisation has the same validity as a real fertilisation
Armin suggested that Erich should publish a short report (Simunek et al. 2011a 58–9). On May 19th, 1901, he
(Autoreferat) on his results in the Biologisches Centralblatt. commented on his experiments with Primula, and informed
This idea, in the end, did materialise in a further early ‘re- his brother of his intention to work on cotyledon formation
discovery’ paper (Tschermak-Seysenegg 1900c), which has (Cotyledonenkeimungsplan), a subject Erich which also
been often overlooked (Mielewczik et al. 2016). Using some touched previously (Tschermak-Seysenegg 1901b) (Simu-
of Mendel’s terminology, A.T.S. returned to his study of de nek et al. 2011a: 60–1). On October 19, 1901, A.T.S. rec-
Vries’s work (in particular, to the hybridization of Chier- ommended that Erich should present his 1901 Zeitschrift für
athus). He expressed his views on current research of vari- das landwirthschaftliche Versuchswesen in Österreich arti-
ation and especially heredity as follows: ‘‘The entire theory cle as an ‘important correction of the views of Mendel and
of heredity needs critical examination. So far [underlined in Rimpau’, that is, not as their pure repetition (Simunek et al.
the original], it offered no proof that a hereditary transfer of 2011a: 62–3). He also instructed Erich on the presentation of
traits of, for example, a species or an individual from a Mendel’s laws in Vienna. When congratulating him on his
parent organism to the offspring takes place in such a way birthday on November 13th, 1901, A.T.S. expressed his
that the differentiation of the former causally determines view that Erich’s chances for a successful academic career
that of the latter. What is a fact is just the conformity, the were at that stage promising (Simunek et al. 2011a: 66–7).
sharing of certain traits: one part involves the living sub- He also mentioned some further unspecified metaphorical
stance (the personal part) immediately, the other (germinal) ‘preparation’ (Präparation) of Hugo de Vries’ person, and
only later, usually only after some infusion of foreign plas- commented in this context that ‘‘Correns will keep his peace
mas, after a ‘fertilisation’. This conformity may be for the now’’ (Simunek et al. 2011a: 66–7). A week later, he
main part related to some particular difference between the thanked Erich for a paper on cereals (Tschermak-Seysenegg
cytoplasm and the nucleus…’’ (Simunek et al. 2011a: 46–8). 1901c); he ranked this paper very high (Simunek et al.
In this context, he called Mendel ‘blessed’ [Segen bringend]. 2011a: 68). Later, both A.T.S. and E.T.S. supported the
Congratulating Erich on June 23rd, 1900, A.T.S. answered interpretation line of simultaneous and triple ‘re-discovery’
his question about the origin of the fabric of the Pisum corns (Simunek et al. 2016).
(Simunek et al. 2011a: 49–51). He recommended to his This rough overview rises the following question:
brother to notify Hugo de Vries of certain corrections in his, Which version of early ‘Mendelism’ E.T.S. presented in
that is, Erich’s larger paper (Tschermak-Seysenegg 1900b) around 1900? He focused primarily on traits in hybrids, in
by a postcard, and added explicitly that Carl Correns should particular on data he obtained in F1 and F2 generation by
not be included. Referring to Correns, he wrote: ‘‘… that heteromorphic xenogamy (Xenogamy means the transfer of
squabbler will be eventually proven wrong by a (later) pollen grains from the anther to the stigma of a different
correction’’ (Simunek et al. 2011a: 49–51). He also strongly plant) (Tschermak-Seysenegg 1901b). In Mendel’s original
discouraged the adoption of de Vries’s segregation [Spal- work, he appreciated the analysis of traits (Merkmals-
tung] hypothesis. He advised his brother: ‘‘The segregation analyse), or ‘decomposition of the whole habitus into
of traits is a schematic speculation of de Vries: I emphati- individual traits’ (Tschermak-Seysenegg 1901d: 1030). He
cally warn against believing it!’’ (Simunek et al. 2011a: also analysed the expression of various traits including the
49–51). On October 15th, 1900, A.T.S. told E.T.S. that he so-called vegetative traits (vegetative Merkmale), and
should finish the second paper on heredity of peas and beans concluded that some traits are realised in hybrids only in an
(Tschermak-Seysenegg 1901b), and discussed de Vries’s alternating (alternierende) form. He saw this as a proof of
theory of mutation (Simunek et al. 2011a: 52). In the first ‘different valuation’ of the traits. Based on this observa-
half of 1901, A.T.S. was apparently very actively working tion, he introduced his own terms for recessive (‘under-
on interpreting the results of Erich’s hybridization experi- valued’/unterwertig) and dominant (‘over valued’/
ments. Correcting their crucial misunderstanding of Men- überwertig) traits. The ‘valence of a trait’ was thus ‘be-
del’s original notion of a ‘first generation of hybrids’, he haviour/comportment of a trait’ (Tschermak-Seysenegg

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1906: 882). He did not, however, see these characteristics the principle of a regular quantitative valence (Satz von der
as generally valid but rather limited to vegetative traits. In gesetzmässigen Mengenwertigkeit), which concerned the
relation to both breeding practice and a ‘racial history’, he number of ‘carriers’ of one particular trait in comparison
was especially interested, most probably for very practical with other corresponding traits; and (3) the principle of
reasons, in the historically understood ‘age of a trait’ regular hereditary valence (Satz von der gesetzmässigen
(Tschermak-Seysenegg 1901d, 1906: 882). He implied that Vererbungswertigkeit) or ‘segregation of traits’ (Spaltung
dominant traits might be older than recessive ones, how- der Merkmale), which could be derived from regular
ever clearly stating himself, that there are certainly many quantitative valuation in a chain of generations.
exceptions (Tschermak-Seysenegg 1906: 882). This would Hereditary ‘factors’ or ‘elements’ as well as the theory of
not support his inclination to hereditary factors as ‘atoms of a ‘purity of gametes’ did not play an explicitly significant
heredity’ in the mechanistic materialistic sense at this time role in his/their early analysis, i.e. in association with a pair
(Allen 2002). Even later, he distinguished between ‘racial of determiners which segregate to individual reproductive
or mendeling traits’ and ‘special traits’ in accordance with cells as they are formed. In papers published by E.T.S. in
H. de Vries (Tschermak-Seysenegg 1906). Later on, E.T.S. 1900 and 1901, there is only one detailed remark on Men-
described his original approach as purely ‘descriptive’ and del’s view, namely, that he ‘… even deduced some elements
‘phenomenological’ ‘‘… in order not at once to anchor the to be in the special organs of the cell or parts of the plasm’
newly-beginning experimental phase of the doctrine of (Tschermak-Seysenegg 1901d: 1030). Especially the theory
heredity—as had happened inexpediently with Darwin- of factors was later attributed by both Tschermak-Seysenegg
ism—to definite theoretical terms‘‘(Olby 1985: 123). In the brothers to C. Correns, W. Bateson, R. Punnett and L.
words of Tschermak’s contemporaries, he dealt primarily Cuénot. In several places, the so-called ‘trait carriers’
with the ‘external mode of heredity’ [äußere Ver- (Träger der Merkmale) were used in connection with the
erbungsweise] (Roemer-Bromberg 1914) (Fig. 4). ratio of 3:1. As a part of ‘further development of Mendel’s
In 1900, and even more in 1901, E.T.S. resumed his legacy’ A.T.S. further added the theory of cryptomery,
views into ‘theory of regular differential valence of traits in which was, for example, understood by both as a parallel
heredity’ (Lehre von der gesetzmässigen Verschiedenwer- line of genetical thinking to the theory of cumulative and
tigkeit der Merkmale für die Vererbung) (Tschermak- simultaneously acting factors (Herman Nilsson-Ehle)
Seysenegg 1901b: 37–8, 1901c: 643–47). It was seen as (Tschermak-Seysenegg 1923: 704, 1942: 228).
consisting of three basic postulates: (1) The principle of a The mutual collaboration of Tschermak-brothers on
regular dimensional valence (Satz von der gesetzmässigen hereditary/genetic issues clearly did not end in 1901. In the
Masswertigkeit), related to an ‘absolute dimension’ (das already quoted manuscript of his memoirs, E.T.S.
absolute Ausmass) or a proportional ‘relative dimension’ acknowledges that ‘‘(…) he [A.T.S.] was from the begin-
(das relative Ausmass) with a clear ‘prevalence’ (Präva- ning to the end of my scientific career my collaborator and
lenz) or ‘undervaluation’ of one or almost of both traits; (2) advisor’’.3 And added that for example, they commented
together on W. Johannsen pure-line theory (Tschermak-
Seysenegg and Tschermak-Seysenegg 1927). A.T.S.’s
probably most important single contribution was his ‘the-
ory of weakening of traits through hybridisation’, also
known as a ‘theory of hybridogenous genasthenia’
(Tschermak-Seysenegg 1918). It was based on his own
experimental research with the crossbreeding of poultry
(Tschermak-Seysenegg 1918). At his physiological insti-
tute in Prague, A.T.S. carried out hybridisation experi-
ments, including reciprocal ones, using 5 species of poultry
and 161 hybrids, until winter 1916–17. In three/four gen-
erations, he followed and correlated 32 traits. This theory
was closely related both to the theory of cryptomery—best
known in E.T.S.’s formulation—and described a precursor
of a true disappearance of a hereditary trait, that is, of
genophthisis (Tschermak-Seysenegg 1935: 192). Last but
not least, A.T.S. referred the results of hybridisation

3
Fig. 4 Last known common photo of the brothers A. T. S. And E. Collection of A.T.S. Stuttgart, transcription of E.T.S.’s manuscript
T. S., 1950 (Photo Archive Michal Simunek, Prague) ‘Mein Bruder Armin’ [My Brother Armin], p. 1.

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 Theory Biosci.

experiments also to the area of eugenics, or ‘Fami- ‘rediscoverers’ and original Mendel’s conclusions was
lienkunde’, as part of which it was then called in German. substantially smaller than it has been generally assumed
In the 1920s, it was in his view both already possible and previously.
desirable to develop a systematic study of heredity in
humans (Tschermak-Seysenegg 1931). From a medical Acknowledgments This paper was published as part of a research
project HO–2143/8–1, 8-2 supported by the German Research
point of view, he emphasised especially the hereditary Foundation (DFG). GL‘s work is supported by the Russian Founda-
character of blood diseases, but was interested also in the tion for the Humanities, project 16-03-00555.
heritability of diseases of the eye (such as colour blindness
for greed and red colours or albinism of the eye), which he
found particularly relevant. He spoke explicitly of ‘patho- References
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