Botany in Berlin

Botany in Berlin
Author(s): Hildemar Scholz

Source: Englera, No. 7, Botany in Berlin (1987), pp. 1-3+5-15+17-103+105-217+219-288
Published by: Botanischer Garten und Botanisches Museum, Berlin-Dahlem
Stable URL: http://www.jstor.org/stable/3776724 .
Accessed: 04/04/2011 13:04
Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at .
http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless
you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you
may use content in the JSTOR archive only for your personal, non-commercial use.
Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at .
http://www.jstor.org/action/showPublisher?publisherCode=bgbm. .
Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed
page of such transmission.
JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of
content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms
of scholarship. For more information about JSTOR, please contact support@jstor.org.
Botanischer Garten und Botanisches Museum, Berlin-Dahlem is collaborating with JSTOR to digitize,
preserve and extend access to Englera.
http://www.jstor.org
ISSN 017 - 4818
ENGLERA 7
in Berlin
Botany
edited
by Hildemar Scholz
on the occasion of the
XIV International Botanical Congress
Veroffentlichungen aus dem Botanischen Garten und Botanischen Museum Berlin-Dahlem

Berlin 1987
ISBN - 3 - 921800 - 27 - 7
Herausgegeben von der Direktion des Botanischen Gartens und

Botanischen Museums Berlin-Dahlem
(Leitender Direktor: Werner Greuter)
Redaktion: Hildemar Scholz
? Botanischer Garten und Botanisches Museum Berlin-Dahlem 1987

Juli 1987
Alle Rechte, auch die des auszugsweisen Nachdrucks, der photo-

mechanischen Wiedergabe, der Herstellung von Mikrofilmen und der
Ubersetzung, vorbehalten.
Contents
Preface. 5
Elisabeth Schiemann - Life and career of a woman

scientist in Berlin, E. Lang. 17
History of phycology in Berlin, D. Mollenhauer .... 29
Contributions to the development of phytomedicine

in Berlin, W. Laux. 51
On the history of plant geography and plant ecology

in Berlin, H. Sukopp. 85
Botany at the Kaiser Wilhelm Institutes,

C. Schnarrenberger. 105
A contribution to the history of science of

heredity in Berlin, W. Plarre. 147
The collections of the Botanical Museum

Berlin-Dahlem (B) and their history, P. Hiepko .... 219
Opera magna in Berlin plant taxonomy,

H. W. Lack. 253
Name index. 283

Englera7- 1987
P r e f a c e
The present collective volume is published on the occasion of the

XlVth Botanical Congress in Berlin in 1987 almost simultaneously
with the (somewhat more voluminous) German edition. Its purpose
is to focus the attention of the public at home and abroad on what has
been achieved in Berlin in the field of botanical science. Attaining even
an approximate completeness is a demand which we had to dispence
with right from the beginning. In addition there are the various diffi-
culties of drawing borderlines thematically and geographically. This
big city - which already counted 100,000 inhabitants at the time of
the EnUghtenment - and its openness to the world make it difficult
to distinguish between the Berliner and the non-Berliner. People
come and go; science in Berlin has received ideas and impulses from
outside and has imparted them in exchange. (In the text the names of
Berlin scientists and writers are spaced out, with the exception of
references to bibliographies in brackets.) The present particular politi-
cal and administrative situation of the former "Reichshauptstadt"
has also set limits to this publication project.
The literature on the work of Berlin botanists is so extensive it
can hardly be surveyed. More recent outlines of the history of botany
at the Humboldt University (GDR) are given by several authors (Hoff?
mann et al. 1965, Bassler 1970, Natho 1977, Vent 1980, Hoffmann
1985, Hoffmann et al. 1985, Hoxtermann 1985, Jahn & Sucker 1985,
Natho & Schmidt 1985, and Walter et al. 1985), which are here expres-
sively refered to. In addition short biographies of eminent Berlin bot?
anists and biologists are pubhshed (Plesse & Rux 1977, Jahn et al.
1982, Hoxtermann et al. 1985, and Zepernick & Timler 1979);compare
also the reports on the development of "Ausseruniversitare technisch-
naturwissenschaftliche Forschungsanstalten in Berlin bis 1945" (Ruske
1979) and of the history of the Kaiser WUhelm Society, later Max
Planck Society (Gerwin 1986). A broadly conceived, extensive, supra-
regional study on the history of German universities is the work of
McClelland(1979).
Preface
The following contributions make no or only marginal mention of

the results of paleobotany (H. Potonie; W. Gothan, A. Straus),
of paleoethnobotany (G. Schwein furth, Elisabeth Schiemann)
and of research on the history of agriculture (E. Hahn, E. Werth);
also of achievements of general biological interest, e.g. protein and
sugar chemistry (E. Fischer, Nobel Prize 1902; see Fruton 1985).
Left out of account were also reports on wide areas of practice orien-
ted research done at university institutes, colleges and institutes of
different Landesanstalten before 1945, and after 1945 in the labora-
tories of the universities, of various federal agencies in Berlin (West),
and of the Academy of Sciences of the GDR. Examples are
Bacteriology: R. Koch (Nobel Prize 1905) and his school (see

Ignatius 1965, Kohler 1985); F. Schaudinn, P. Ehrlich
(Nobel Prize 1908) (see Schneider-Romheld 1955)
Botanical aspects of forestry, agriculture and horticulture: K. v.

Rumker, L. Wittmack;E. Kemmer (see Kramer 1956,
Berthold et al. 1960, Bulow 1960, Skibbe 1960, Noack 1961,
Sucker 1980, Schindler 1985, Natho & Wiedenroth 1985)
Pharmacognostics: A. Tschirch (see Langhammer & Rimpler

1965 and Hiller & Nahnhauer 1985)
Pharmacy: G. E. S tahl (see Engel 1984, Strube 1984)
Phytotherapy: R. F. Weiss
The fact that traditional special fields of botanical research in Berlin

could not be treated in this volume must be seen as a special deficiency:
Yeast and fermentation research: E. Buchner (Nobel Prize

1907), P. Lindner; S. Windisch (see Windisch 1971, 1974,
1982, Anonymous 1983, Bohme 1978)
Sugar and sugar-beet research: A. Marggraf, F. C. Achard
(see Lippmann 1929, also Kolbe 1985 and "Schriften aus dem
Zuckermuseum" 22, 1986)
Additional mention should be made of the "Jahresberichte des

Instituts fiir Systernatische Botanik und Pflanzengeographie der Freien
Universitat" and of the survey on scientific botanical activities at the
former Padagogische Hochschule (Heistermann & Heinrich 1980),
Englera7- 1987
now integrated into the Technical University. Some research results

have not been recorded yet (not even in the present contributions)
and are still awaiting thorough science-historical treatment, with
a view to the connection between "Zeitgeist" and science-organi-
zation, the activities of the Berliner Akademie der Wissenschaften
and the natural science societies or associations in East and West, the
variety of ideas and the innovative power of individual persons.
Today most people are not aware to which large extent the present
basic and textbook-knowledge of botany had its origin in the Berlin
of the 19th and the first decades of the 20th century. This is where
after some errors and confusion (see J. Hanstein 1877, Wunderlich
1982, Forster 1984) the concepts of the sexuality of plant and fungi
were estabUshed and consolidated (M. Schleiden, N. Pringsheim,
H. Kniep, M. Hartmann). The idea of the cell individual became
decisive (Th. Schwann; see Studnicka 1933) and created modern cell-
theory. In Berlin the foundation was laid for today's life cycles theory
(N. Pringsheim, A. Braun; the term "alternation of generations"
by A. v. Chamisso 1812 for metagenesis!). The comparative morpho-
logy produced top performances and gained world-wide recognition (A.
Braun, A. Eichler); see Eckardt 1957 (early opposition by C. H.
Schultz-Schultzenstein about 1850). Appreciated to this very
day are the results of the analyses of the anatomy of higher plants in
relation to their functions and environment (A. Tschirch, S. Schwen-
dener, G. Volkens, G. Haberlandt). Exemplary perfection was
reached in the further development of the Natural System (A. E n gl e r)
with 13 independent divisions of lower and higher plants (polyphyly).
(The "Grundzuge einer Theorie der phylogenetischen Systematik" were
devised by the entomologist W. Hennig 1950, then a scientific em-
ployee at the Biologische Zentralanstalt in Berlin-Dahlem.) Berlin
botanists achieved remarkable results in the field of plant serodiag-
nostics (E. Gilg in cooperation with P. Schurhoff). Pioneering
contributions to genetics, physiology of plant development and photo-
synthesis research (E. Baur, C. Correns; J. Hammerling, F. v.
Wettstein; O. Warburg, Nobel Prize 1931, and R. Willstatter,
Nobel Prize 1915) have mediated via botanical virus research (E.
Pfankuch, G. Schramm) to present molecular biology and bio-
technology. In 1828, also in Berlin, F. W ohler carried out his famous
Preface
urea synthesis, which was considered important in the vitalism contro-

versy (for critical annotations see Sapper 1941).
Berlin botanists had no little share in the botanical exploration of
the world (which certainly can never be considered finished). Above all
since the work of A. v. Hu m b ol d t (see Biermann 1984), the founder
of plant geography, and the voyages round the world by A. v. Chamisso
1815-1818 and F. Meyen 1830-1832 a long chain of successful
projects leads from the travels of Ch. G. Ehrenberg in Northeast
Africa and Arabia to J. Mildbraed's explorations 1907-1928 in
tropical Africa and ? with increasing international interlacing ? to the
immediate present. The "Vegetationskarte von Afrika" was a great
"last document of the Dahlem Africa tradition of bygone days" (co-
lonial era; W. Domke 1963, see also Domke 1966). The activities of
the Gesellschaft fiir Erdkunde in Berlin (Bader 1969) have always
been very promotional (e.g. by allocation of travel scholarships).
On reading the present collection of essays the gentle reader will

know to honour the authors' personal touch in style and mastering
of the material, mindful of the realization that "the history of science
is a history of events which are selected and interpreted normatively"
(Lakatos 1978: 17). Thus many a misbalance in the contents of the
texts may be excused. The authors, without whose unselfish efforts this
work could not have been published, may rest assured of the editor's
thanks.
References
Anonymus 1983: Professor Dr. Siegfried Windisch in den Ruhestand

getreten. - Die Branntweinwirtschaft 1983: 420-421.
Bader, F. 1969: Ruckblick nach 100 Banden der Zeitschrift der Ge-
sellschaft fur Erdkunde zu Berlin. - Die Erde 100: 93-117.
Bassler, M. 1970: Das Herbarium des Bereichs Botanik und Arboretums
des Museums fur Naturkunde an der Humboldt-Universitat. ?
Wiss. Z. Humboldt-Univ. Berlin, Math.-Nat. R. 19: 295-299.
Berthold, R. et al. 1960: Landwirtschaftlich-gartnerische, veterinar-
medizinische und forstwirtschaftliche Lehre und Forschung. ? In:
Englera7- 1987
R. Berthold (ed.), Die Humboldt-Universitat. Gestern-Heute -

Morgen. - Berlin, p. 92-103.
Biermann, K.-R. 1948: Die Akademie pflegt Humboldts Werk. -
Spektrum (Berlin) 15(4): 12-13.
Bohme, G. 1978: Autonomisierung und Finalisierung. - In: Starn-
berger Studien 1. Die gesellschaftliche Orientierung des wissen-
schaftlichen Fortschritts. - Frankfurt a.M., p. 69-130.
Bulow, F. 1960: Grundlagen, Entwicklung und Bedeutung der Land-
wirtschaftlichen Fakultat an der Friedrich-WUhelms-Universitat
Berlin bis zum Jahre 1945. - In: H. Leussink, E. Neumann & G.
Kotowski (ed.), Studium Berolinense. Aufsatze und Beitrage zu
Problemen der Wissenschaft und zur Geschichte der Friedrich-
WUhelms-Universitat. - BerUn, p. 905-919.
Domke, W. 1963: Bemerkungen zu der von L. Diels, J. MUdbraed und
G. Schultze-Menz 1939-1942 bearbeiteten Vegetationskarte von
Afrika. - WUldenowia, Beiheft 1.
- (ed.) 1966: Grundziige der Vegetation des tropischen Kontinental-
Afrika von Johannes MUdbraed f. - WUldenowia, Beiheft 2.
Eckardt, Th. 1957: Vergleichende Studie iiber die morphologischen
Beziehungen zwischen Fruchtblatt, Samenanlage und Blutenachse
bei einigen Angiospermen. Zugleich als kritische Beleuchtung der
"New Morphology". ? Neue Hefte zur Morphologie, 3. Heft. -
Weimar.
Engel, M. 1984: Chemie im achtzehnten Jahrhundert. Auf dem Wege zu
einer internationalen Wissenschaft. Georg Ernst Stahl (1659-1734)
zum 250. Todestag. - Berlin.
Forster, K.-R. 1984: Zur Entdeckungsgeschichte der Befruchtung der
Blutenpflanzen. - Zum 150. Gedenkjahr einer Pionierarbeit von
C. A. Corda. - Gleditschia 12: 193-211.
Fruton, J. S. 1985: Contrasts in scientific style. EmU Fischer and Franz
Hofmeister: Their research groups and their theory of protein struc-
ture. - Proc. Amer. PhUosoph. Soc. 129: 313-370.
Gerwin, R. 1986: 75 Jahre Max-Planck-Gesellschaft. Ein Kapitel
deutscher Forschungsgeschichte. - Naturw. Rundschau 39: 1-10,
49-62,97-109.
Hanstein, J. 1877: Die Parthenogenesis der Coelebogyne ilicifolia.
Nach gemeinschaftUch mit Alexander Braun angestellten Beobach?
tungen mitgeteUt. ? Bot. Abhandl. Morphologie und Physiologie
3, 3. Heft. - Berlin.
10 Preface
Heistermann, W. & Heinrich, G. 1980: Beitrage zur Geschichte der

Padagogischen Hochschule Berlin. - Abhandl. Padag. Hochschule
Berlin 6.
Hiller, K. & Nahnhauer, Annette 1885: Die Entwicklung der Pharma?
kognosie als Hochschulfach an der Berliner bzw. Humboldt-Univer?
sitat. - Wiss. Z. Humboldt-Univ. Berlin, Math.-Nat. R. 34: 215-222.
Hoxtermann, E. 1985: Die Botanik in der Veterinarmedizin in Berlin. -
- Dembny, H. & Lotze, Kerstin 1985: Kurzbiographien und Portrats
von Botanikern in der Geschichte der Berliner Universitat. - Wiss.
Z. Humboldt-Univ. Berlin, Math.-Nat. R. 34: 360-384.
Hoffmann, P. 1985: Zur Entwicklung der Botanik an der Humboldt-
Universitat unter besonderer Berucksichtigung der Photosynthese-
forschung. - Wiss. Z. Humboldt-Univ. Berlin, Math.-Nat. R. 34:
203-207.
- Giinther, Ulrike & Blohm, D. 1965: Die Botanik und ihre Vertreter
an der Berliner Universitat. (Von 1810-1945). - Wiss. Z. Humboldt-
Univ. Berlin, Math.-Nat. R. 14,6: 799-815.
- Kohl, J.-G. & Stahlberg, R. 1985: Konrad Ramshorn (26.1.1909-
23.11.1978) und die Entwicklung der Pflanzenphysiologie an der
Humboldt-Universitat zu Berlin. - Wiss. Z. Humboldt-Univ.,
Math.-Nat. R. 34: 255-259.
Ignatius, A. 1965: Koch, Leben und Forschung. ? Stuttgart.
Robert
Jahn, Ilse, Lother, R. & Sengbusch, K. 1982: Geschichte der Biologie.
Theorien, Methoden, Institutionen, Kurzbiographien. ? Jena.
- & Sucker, U. 1985: Zur Geschichte der Botanik an der Berliner Uni?
versitat von 1810 bis 1945. - Wiss. Z. Humboldt-Univ. Berlin,
Math.-Nat. R. 34: 189-202.
Kohler, W. 1985: Shibasaburo Kitasato und Sahachiro Hata. - Leopol-
dina(R. 3)28: 99-122.
Kolbe, W. 1985: 200 Jahre Pflanzenschutz im Zuckerrubenbau (1784-
1984). Bonn, 2nd ed. 1986.
Kramer, M. 1956: 75 Jahre Landbaustudium in Berlin 1881-1956.
Fakultat fiir Landbau an der Technischen Universitat. - Berlin-
Charlottenburg.
Lakatos, I. 1978: Pilosophical papers I. - Cambridge, Mass.
Langhammer, Liselotte & Rimpler, H. 1965: Pharmakognosie an der
Berliner Universitat. ? Deutsch. Apotheker Ztg. (105. Jg.) 42:
1468-1472.
Englera 7-1987 11
Lippmann, E. O. v. 1929: Geschichte des Zuckers seit den altesten

Zeiten bis zum Beginn der Riibenzuckerfabrikation. Ein Beitrag
zur Kulturgeschichte. ? Berlin, supplemented reprint Wiesbaden
1970.
McClelland, Ch. 1980: State, society and university in Germ any 1700-
1914. ? Cambridge, Mass.
Natho, G. 1977: Das Arboretum an der Humboldt-Universitat zu
Berlin ? Geschichte, aktueller Stand, Entwicklungsperspektiven. ?
Gleditschia 5: 37-44.
? & Schmidt, H. 1985: Zur Entwicklung der Speziellen Botanik als
Wissenschaftsdisziplin an der Berliner bzw. Humboldt-Universitat.-
- & Wiedenroth, E.-M. 1985: Zur Geschichte der Botanik an der
Landwirtschaftlich-Gartnerischen Fakultat der Humboldt-Univer?
sitat zu Berlin. - Wiss. Z. Humboldt-Univ. Berlin, Math.-Nat. R.
34: 235-245.
Noack, E. 1981: Professor Erich Maurer t- - Die Orchidee 32: 197.
Plesse, W. & Rux, D. 1977: Biographien bedeutender Biologen. Eine
Sammlung von Biographien. ? Berlin.
Ruske, W. 1979: Ausseruniversitare technisch-naturwissenschaftliche
Forschungsanstalten in Berlin bis 1945. ? In: R. Riirup (ed.),
Wissenschaft und Gesellschaft 1. - Berlin, p. 231-263.
Sapper, K. 1941: [Buchbesprechung: Bavink, Ergebnisse und Probleme
der Naturwissenschaften]. ? Blatter Deutsch. Philosophie 15: 223.
Schneider-Romheld, W. 1955: Das Steglitzer Institut fiir Serumfor-
schung. ? Steglitzer Heimat 19: 1-9.
Schindler, N. 1985: Berliner Pflanzen. Berlin und Berliner als Namens-
geber. ? Berlin. Forum 18(2).
Skibbe, B. 1960: Zur Vorgeschichte der Landwirtschaftlich-Gartne-
rischen Fakultat (1806-1881). - Wiss. Z. Humboldt-Univ. Berlin,
Jubilaumsjahrg. 9: 229-256.
Strube, Irene 1984: Georg Ernst Stahl. ? Biographien hervorragender
Naturwissenschaftler, Techniker und Mediziner 76. ? Leipzig.
Studnicka, F. K. 1933: Matthias Jacob Schleiden und die Zelltheorie
von Theodor Schwann. ? Anat. Anzeiger 76: 80-93.
Sucker, U. 1980: Zur Geschichte der Dendrologie und des Spathschen
Arboretums in Berlin (1846-1945). - Wiss. Z. Humboldt-Univ.
Berlin, Math.-Nat. R. 29, 3: 261-265.
12 Preface
Vent, W. (ed.) 1980: 100 Jahre Arboretum in Berlin. - Berlin.

Walter, Gaby, Hoffmann, P. & Seifert, Barbara 1985: Beitrage von Na-
turwissenschaftlem der Berliner Universitat zur Photosynthese-
forschung (von der Universitatsgriindung bis 1945). ? Wiss. Z.
Humboldt-Univ. Berlin, Math.-Nat. R. 34: 230-234.
Windisch, S. 1971: Ergebnisse der Forschung an Hemiascomyceten um
und seit Wilhelm Henneberg. - Kieler MUchwirtschaftl. Forschungs-
ber. 23: 139-148.
- 1974: 100 Jahre mikrobiologische Forschung. Beitrag des For-
schungsinstituts fiir Mikrobiologie der Versuchs- und Lehranstalt
fiir Spiritusfabrikation und Fermentationstechnologie. - In: 100
Jahre Institut fiir Garungsgewerbe. ? Berlin, p. 110-119.
? 1982: Forschungsinstitut fiir Mikrobiologie. ? Festschrift: 125
Jahre Versuchs- und Lehranstalt fiir Spiritusindustrie und Fer?
mentationstechnologie in Berlin 1897-1982. ? Berlin, p. 63-66.
Wunderlich, Rosalie 1982: Der wissenschaftliche Streit iiber die Ent-
stehung des Embryos der Bliitenpflanzen im zweiten Viertel des
19. Jahrhunderts (bis 1856) und Mendels "Versuche uber Pflanzen-
hybriden". - Folia Mendeliana (17) (Acta Mus. Morav. 67): 225-
242.
Zepernick, B. & Timler, F. K. 1979: Grundlagen zur 300jahrigen Ge?
schichte des Berliner Botanischen Gartens. - Englera 1.
Hildemar Scholz
EpUogue to the preface
The Nazi-era from 1933 to 1945 is also part of the history of Berlin.
In general it had practically no consequences for botany, since there
were but a few points of contact with the "Volkische Weltanschauung"
of the Nazis. It is only by a detour over the laws of inheritance formu-
lated by Mendel, the Mendelism, that connections can be made mani-
fest. Lamarckism, the theory of inheritance of acquired character-
istics, whether in man, animals or plants, was considered "Bolshevistic"
by NS-ideologists; the relations with Darwinism was ambivalent. Their
ideal was a third one: a static nature, organized strictly hierarchically,
with the Germanic race ranging top; race as fate. The order was clear
Englera 7-1987 13
and was considered a cultural obligation of the highest rank: to keep

the Germanic race from decline and fall, to practise eugenics. This was
not a new demand. A protection of the race had already been demanded
by E. B aur, then a reputable plant geneticist, in his essay "Der Unter-
gang der Kulturvolker im Lichte der Biologie" (1922, republished in
an enlarged version in 1932), written under the depressing impression
of the Peace Treaty of Versailles. In 1907 Baur (who also held a
doctor's degree in medicine) had joined the Gesellschaft fiir Rassen-
hygiene, founded by Ploetz, and was chairman of the local Berlin
branch from 1917-1919. The "Baur-Fischer-Lenz" ("Grundriss der
menschlichen Erblichkeitslehre und Rassenhygiene") was published
in 1921, Hitler studied it, and it became "the corner stone of Nazi
eugenics" (Glass 1981: 357). In this book Baur worte (p. 22): "A
people or grouping of people inferior by its hereditary disposition ?
e.g. the negroes in the United States of North America - are improved
through education and cultural influence in respect to their single
individuals, but the race as such is not changed in this way". According
to Schiemann (1935) Baur took a decisive part in the building
and staffing of the Kaiser Wilhelm Institute for "Anthropologie, mensch-
liche Erblehre und Eugenik" in Berlin-Dahlem. Baur must be counted
among the pace-makers of the application of the genetic findings
of that time on man, in race-hygiene, eugenics. "It was self-evident
that Baur had to welcome it, when the new state [1933] adopted the
endeavours of the eugenicists with a resolute hand ? in the good and
thoroughly done preparatory work, which it [the state] came upon,
Baur had an essential merit" (Schiemann 1935: 107;later Schiemann
was herself exposed to Nazi-reprisals). Still Baur was no Nazi, rather a
science-devouted idealist and a patriot. In 1933, during the last months
of his life, he is said to have strongly stood up for the members of staff
of his institute against persecution by the Nazis (Glass l.c. 363).
The idealism of Baur was not an isolated case. R. C. Muschler

(1882-1957), who had been working at the Botanical Museum Berlin-
Dahlem for a longer period of time, but had failed there as a botanist
and had later become popular as a novelist, enthusiastically called
Hitler's book "Mein Kampf' the "creed of a Messiah" (Muschler
1933). Beside there were many in different ones, a not trifling number
of opportunists (as always), but also helping hands. Today it is often
no more possible to make out, who was what. In 1938 Freiherr Fried-
14 Preface
rich v. Luetzelburg (obituary by Suessenguth 1955) became head

of the Forschungsstelle fiir Botanik of the SS-Ahnenerbe in Berlin, R.
Schutrumpf, also in Berlin, was made head of the Forschungs-
statte fiir Naturwissenschaftliche Vorgeschichte at the same insti?
tution, with a laboratory for pollen analysis (Kater 1974). L. Diels,
director of the Botanical Garden and Botanical Museum BerUn-Dahlem,
a man of strict national-conservative thinking, received the "Goethe-
MedaUle fiir Kunst und Wissenschaft" from the "Fiihrer" in September
1944 (Diels, unpublished diary notes). It was inevitable that among the
botanists many party members of the NSDAP lost their function, job
and income after 1945. Thus they experienced a minimum of what
others had been forced to suffer since 1933, only a few could emi-
grate, like R.Goldschmidtof the Kaiser WUhelm Institute.
O. Schwarz, member of the Communist Party since 1926, did not
find the position he had hoped for at the Botanical Museum Berlin-
Dahlem and lost his employment at the Biologische Reichsanstalt
Berlin-Dahlem in 1934 (Klotz 1983, Meyer 1984). F. Merken-
s c h 1 a g e r, also at the Biologische Reichsanstalt and in his "Weltan-
schauung" closely connected with Ernst Jiinger and Ernst Niekisch, as a
side line opposed the glorification of the Nordic race in various pub-
lications (e.g. 1927, 1933) and in newspaper articles, for which he was
sentenced to years in prison, among others in the KZ Dachau and the
Gefangnis Berlin-Moabit (UUrich 1970, Kirchner 1984); interventions
by Privy CouncUlors A. Bier and F. Sauerbruch could not help
(Saller 1961). Some lost their lives, above all jewish feUow-citizens; not
only acts of war led to the horrible. A. Berliner, founderof the dis?
tinguished journal "Die Naturwissenschaften" (1913, ibidem 1946
obituary by M. v. Laue) in the JuUus Springer Verlag, was driven to
suicide in 1942 (see Davidis 1985). H. L. Spath, son of the owner of
tree nurseries F. L. Spath, who had become known through his thesis
1912 on the lammas shoots (here the terms syllepsis and prolepsis in
the modern sense), was sent to the KZ Sachsenhausen in 1944, because
he had voiced his disapproval of the Nazis, and was shot there in
February 1945 (Sucker 1980). (Today the Spathsche Baumschule is the
arboretum of the Humboldt University.)
Baur, E. 1922: Der Untergang der Kulturvolker im Lichte der Biologie. -

Deutschlands Erneuerung 6: 257-268. (2nd enlarged edition 1932
in Volk und Rasse 7: 65-79)
Englera 7 - 1987 15
? Fischer, E. & Lenz, F. 1921: Grundriss der menschlichen Erblich-

keitslehre und Rassenhygiene. - Miinchen.
Davidis, M. 1985: Wissenschaft und Buchhandel. Der Verlag Julius
Springer und seine Autoren. Briefe und Dokumente aus den Jahren
1880-1946. -Miinchen.
Glass, B. 1981: A hidden chapter of German eugenics between the
two World Wars. - Proc. Americ. Pilosoph. Soc. 125: 357-367.
Kater, M. H. 1974: Das "Ahnenerbe" der SS 1935-1945. Ein Beitrag
zur Kulturpolitik des Dritten Reiches. ? Stuttgart.
Klotz, G. 1983: In memoriam Otto Schwarz. ? Wiss. Z. Friedrich-
Schiller-Univ. 32: 839-848.
Merkenschlager, F. 1927: Gotter, Helden und Giinther. Eine Abwehr
der Giintherschen Rassenlehre. ? Niirnberg.
? 1933: Rassensonderung, Rassenmischung, Rassenwandlung. -
Berlin-Steglitz.
Meyer, F. K. 1984: Otto Schwarz (1900-1983). - Haussknechtia
2: 3-6.
Muschler, R. C. 1933: Bekenntnis ? Mein Kampf um Adolf Hitler. -
In: C. A. Dreyer (ed.), Reinhold Conrad Muschler. Ein Deutscher
Weg. - Leipzig.
Saller, K. 1961: Die Rassenlehre des Nationalsozialismus in Wissen?
schaft und Propaganda. ? Darmstadt.
Schiemann, Elisabeth 1935: Erwin Baur. - Ber. Deutsch. Bot. Ges.
52:(51)-(114).
Spath, H. L. 1912: Der Joahannistrieb. Zur Kenntnis der Periodizi-
tat und der Jahresringbildung sommergriiner Holzgewachse. ?
Berlin.
Sucker, U. 1980: Zur Geschichte der Dendrologie und des Spathschen
Arboretums in Berlin. - Wiss. Z. Humboldt-Univ. Berlin, Math.-
Nat. R. 29, 3: 261-265.
Suessenguth, K. 1955: Philipp FreiherrvonLuetzelburg. -Ber. Deutsch.
Bot. Ges. 68a: 65-69.
Ullrich, J. 1970: Gedenkrede fiir Friedrich Merkenschlager. - Aus der
Spalter Heimat. Heimatkundl. Hefte 9: 25-30.
Hildemar Scholz
Englera7-1987 17
Anton Lang
Elisabeth Schiemann
Life and career of a woman scientist in Berlin
EUsabeth Schiemann was no native Berliner but she received

her entire education ? elementary school, high school, university ? in
Berlin and except for some brief stays as a visiting scientist abroad
spent her entire professional career in this city. Her life thus reflects, on
the one hand, the problems a woman encountered in entering an
academic career, and on the other hand, the very variable, sometimes
excellent, sometimes very difficult conditions under which science was
working in Berlin, in this century.
Elisabeth Schiemann was born in a Baltic-German famUy in
Livlandia, in the year 1881, and her German speech betrayed this
origin throughout her life. However, as early as 1887 the famUy moved
to Berlin where the father, Theodor Schiemann, became Professor of
Eastern-European History at the University, a post he held untU 1921.
Professor Schiemann was an archconservative man and in the last two
decades of the German Kaiserreich he played a prominent part in
extreme rightist, nationalistic circles whose political influence was not
exactly positive. However, and understandably, the famUy home had
a strong influence on EUsabeth Schiemann, an influence that lasted
throughout her Ufe. She was herself conservative but in the positive
meaning of this term. She did not oppose progress per se, in fact, she
often approached it with understanding and sympathy, but she wanted
it to maintain continuity with the positive aspects of existing concepts
and institutions. She was for evolution but against revolution. From the
parental home came also her appreciation of history, including cultural
history, which should be very helpful for her in a later period of her
life, and her interest in the arts, particularly music; her younger sister
Gertrud, with whom she was very close throughout her life, was a gifted
musician. I had the opportunity of visiting with the two sisters some
Lang: Elisabeth Schiemann
concerts and, this later, after World War II, opera performances, and
these were those occasions when Elisabeth Schiemann, in general a
reserved person, became open and communicative and her innate
warmth and the whole broad scope of her personality became most
apparent. She is also said to have been an excellent social dancer,
but this is something I know only from hearsay.
After finishing school Elisabeth Schiemann enrolled in a teacher
college for girls and having completed it taught for several years the
lower grades in a girls' school. However, she had no enthusiasm for this
activity, particularly since her interest in sciences became increasingly
pronouneed. In 1906 she enrolled as a special student in Berlin Uni?
versity, and two years later, when this University, like all universities
in Prussia, began to accept women as regular students, she registered
in this capacity - after obtaining her high-school diploma - in the
Philosophical Faculty which in those days included Natural Sciences.
Elisabeth Schiemann has described, in 1960, her time as a student

in Berlin in "Studium Berolinense", a special volume commemorating
the 150th anniversary of the Friedrich Wilhelm University of Berlin, an
anniversary which could be marked in a historically objective man?
ner only in the Western part of the now divided city. However, and this
was in line with her character, she seems to have placed more emphasis
on the positive than the negative experiences. From occasional remarks
of hers ? she did not reminisce on these matters frequently and did not
seem to like reminiscing about them ? the daily life of a "coed" was
not altogether simple in those times. It is true that the situation in
sciences was on the whole better than that in the "arts" where the few
girl students were classified by the male environment, fellow students
and instructors alike, either as workless sex horses or sexless work-
horses. However, not all professors of science were very sympathetic
towards female students, either. One such student, who later became a
world-famous scientist, was allowed to do her experimental work in the
basement of the institution in question, but was not allowed to enter
any other parts of the building. Another, when she was discovered by
the professor in the classroom, was offered his arm and, with the words,
"Respected miss (gnadiges Fraulein), may I escort you from the lecture
hall?" removed from the audience, politely but firmly. Elisabeth
Schiemann seems however not to have experienced such extreme
indignities, and it is a clear testimony for her intellect as well as her
Englera7 - 1987 19
personality that she was accepted on equal terms by some of her best
fellow students, for it is to their student years in Berlin that her life-
long ties with men like Otto Hahn and Gustav Hertz, and also with
Lise M e i t n e r ? all Nobel laureates ? are dating back.
It was under the influence of Erwin Baur, then a younginstructor

but soon to become one of the leading geneticists, and the undisputed
leader in plant breedingresearch, in Germany that EUsabeth Schiemann
became fascinated with genetics, and she did the experimental work
for her doctoral thesis in this young discipline with Baur as her major
professor. The topic of her research, induction of mutations in Asper-
gillus niger, sounds quite modern by today's standards. However,
as this mold reproduces only asexually, it was not accessible to the
method which then, and for a long time, was the touchstone of genetic
work, Mendelian analysis. The author herself had no hesitation calling
the variant forms she succeeded in isolating, but many of which were
unstable, mutants but today we would probably regard the question of
their nature, genetic or epigenetic, with more reservation. It is tempting
to wonder where the work might have lead if it had been conducted
with a mold having sexual reproduction, as it was done later, and with
such striking results, in Neurospora. However, time was probably
not yet ripe for "biochemical genetics".
Baur was at that time not on the faculty of Berlin University, but
on that of the Agricultural College of Berlin, a separate institution,
first as lecturer and then as professor of botany. "Symbioses" between
the two institutions were quite possible, however, in those days and stiU
in the days I was a student (1931-1939) and whUe Schiemann's
research was done at the Agricultural College her degree was from
Friedrich WUhelm University (today, Humboldt University) of BerUn.
However, soon after graduation she joined Baur at the Agricultural
College, as scientific assistant and later senior scientific assistant (the
first two rungs of the German academic stepladder at that time) at the
Department of Genetics, which had been created at Baur's initiative
and was the first such unit in Germany and quite likely in the world ?
certainly one of the first. In this institution she also acquired her
"habUitation", that is, became an instructor.
The environmental conditions for the new department and its staff
were, for many years, far from easy, because of World War I which
started very soon, and the difficult situation in the early post-War
20 Lang: Elisabeth Schiemann
years. The experimental work had first to be conducted on a private

plot of Baur in Friedrichshagen, a suburb to the East of Berlin. Then,
in the war, the department was moved to Potsdam where, according
to existing plans, the entire Agricultural College was to be relocated.
After the end of the war these plans had to be abandoned, however;
for some time the department was housed in the main building of the
college, in the northern part of Berlin, and it was only in 1923 that it
could move into a building of its own, spacious and well-designed,
and surrounded by ample experimental grounds, in the suburb of
Dahlem, in the southwestern part of the city. It is still in this location,
as the Institute of Applied Genetics of the Free University of Berlin
(into which some of those institutions of the Agricultural College that
happened to be in West Berlin were incorporated). Elisabeth Schie?
mann was very much involved in the preparation and organization of
these various peregrinations, and her responsibilities also included the
preparation of lectures and laboratories and the care and increase of the
department's plant stocks, particularly the valuable collection of
cereals. Above and beyond these calls of duty she was something like an
older sister to the younger coworkers of the department, tireless in
helping them to start their own research projects and assisting them
with various other problems as well.
The time in Dahlem was a golden era for the department and for
Elisabeth Schiemann. Intensive and interesting work had been done
in Friedrichshagen and Potsdam, but it was only here that the depart?
ment's research program could be pursued in full breadth and depth.
Elisabeth Schiemann developed her own, very active research,
first mainly with barley, somewhat later also with wheat, Aegilops
and Fragaria. The department, an internationally recognized center
of genetics and breeding research, was also regularly visited by some
of the most eminent plant geneticists and breeders in the world, e.g.
Nilsson-Ehle, Johannsen and Vavilov, and these contacts not only broad-
ened Schiemann's scientific outlook but also resulted in close per?
sonal relations which, as far as that was possible under the political
developments in the following years, lasted throughout the lives of
those involved. A special highlight was the Fifth International Congress
of Genetics that was convened in Berlin in 1927. Elisabeth Schiemann
was actively involved in it, too, particularly in the preparation and
supervision of scientific exhibits and as guide on at least one major
Englera7 - 1987 21
Congress trip, and again these were opportunities for establishing new
professional as well as human contacts. These extensive contacts
abroad and Elisabeth Schiemann's uncompromising attitude during
the Hitler years, which wUl be considered later in this biography, were
undoubtedly essential factors in that she was one of the first German
scientists who were invited and allowed to visit some of the former
enemy countries after the end of World War II, in her case Great
Britain in 1947.
The golden era in the Genetics Institute did not last for EUsabeth
Schiemann more than six years. Baur had been pursuing plans for
the creation of a special institute for breeding research, which he
had initiated already during World War I, with his typical energy and
persuasion, and he now succeeded, through a joint agreement between
the Kaiser WUhelm Society for the Advancement of Sciences and se?
veral leading seed companies, in bringing these plans to fruition. The
Kaiser WUhelm Institute for Breeding Research was formally founded
in 1927 and began its research work about a year later, in Miincheberg,
a small city to the East of Berlin. But in this connection there occurred
a break between Baur and Schiemann, and the latter, although
she had intensively taken part in the planning of the new institute, and
probably had expected to be appointed to a position of greater respon-
sibUity, did not move from Dahlem to Miincheberg.
The two individuals primarily concerned do not seem to have talked
about the precise reasons for this rift. In retrospect, however, one may
wonder that Baur and Schiemann had been able to cooperate
as long and as effectively as they did, for it is difficult to envisage two
individuals so different from one another, and in many respects outright
opposites. I have seen Baur personally, "in action", only at two lec-
tures and on two student field trips to the Miincheberg institute;
incidentally, on both these excursions he seemed to have, for us bloody
beginners, all the time in the world, answering all our questions, whether
smart or sUly, with outmost patience ? a trait I have noted in many
other truly eminent scientists I had the fortune of meeting. However,
from these brief encounters and from all one could hear and read about
him, Baur was not only a very imaginative and forward-looking
scientist, who thought in broad concepts and would disregard contrary
details, often to be proven right but sometimes wrong, too, but also
a fascinating speaker who had no difficulty of convincing his audience,
whether beginning students or eminent scientists in their own right,

whether business leaders or politicians, of his ideas and objectives.
Schiemann, although as already mentioned by no means against
progress, wanted to gather all facts before she was willing to commit
herself to an idea, and even then she would point out, in a very ob-
jective manner, those details that did not quite fit that idea. Herlectures
and seminars were excellent for learning all sides of a problem; but they
could not be called inspiring, and I am in fact afraid that many in her
audiences, particularly students, often were unable to see the wood
for the trees. She also had a distinct authoritarian trait which was quite
absent in Baur. In short, if one ever could anticipate a collision be?
tween an irresistible force and an immovable object it would be between
Baur and Schiemann. It is a sign of the fundamental greatness of
their personalities that they became reconciled before Baur's early
death; after his death, Schiemann wrote an outstanding obituary
which did full justice to his professional and personal qualities ("Be?
richte der Deutschen Botanischen Gesellschaft" 1935).
Also in retrospect it could perhaps be argued that the separation from
Baur helped Schiemann inasfar as it at least delayed her clash
with the Nazi regime. Baur died prematurely in 1933. Actinghead of
the Muncheberg Institute became B. Husfeld;in 1936 W. Rudorf
was appointed as the new director. Both men were, or at least pretended
to be, ardent Nazis. Within three years they had fired, under shabby
pretexts but clearly for political reasons, three of Baur's senior and
best coworkers, H. Kuckuck, R. Schick and H. Stubbe; about
half a year later another member of that "old guard", R. von Seng-
busch, was also forced to resign. Schiemann was, as already briefly
mentioned, strongly anti-Nazi; she would not have kept silent during
these events, and would most likely also have been fired by the insti-
tute's new management ? assuming she could have stayed, or wanted
to stay, that long at all. It is one of the many ironies of the "denazi-
fication", which was carried out after the end of World War II, that
whereas many "little party members" lost their jobs and some were
interned, and whereas several of the scientist dismissed from the Breed-
ing Research Institute had a difficult time finding satisfactory positions,
Rudorf remained director of the institute, now called Max Planck
Institute of Breeding Research/Erwin Baur Institute and first located
in Voldagsen near Hameln and later, on a permanent basis, in Cologne-
Englera7-1987 23
Vogelsang. While he continued in office, the research in the institute

became increasingly parochial and stagnant, a trend which only was
stopped, and replaced by a rapid and brilUant upswing, when he reached
retirement and was replaced by J. Straub (1961).
Regardless of whether or not these excursions in hindsight are justi-
fied, the break with Baur was not only a heavy psychological shock
for EUsabeth Schiemann, but also meant a drastic reduction of her
research opportunities. For reasons which are not clear to me she
decided to leave the Department of Genetics and the Agricultural
College, joining the University of Berlin where she was appointed
extraordinary (as far as the two systems can be compared, approxi?
mately between assistant and associate) professor without tenure and
was accommodated as a visiting scientist in the Botanical Museum in
Dahlem, an institution of the University. She had an office-laboratory
and field plots in the adjacent Botanical Garden (where she developed
a smaU but very instructive genetical exhibit), had additional field space
in the Kaiser WUhelm Institute for Biology, also in Dahlem, and had
support for a technician and hourly help in field work. However, all
this was a good deal less, and less effectively arranged, than what she
had before. Her economical situation was also far from easy. For
many years she lived in some of the famous or infamous "furnished
rooms with kitchen privUeges," which were then quite common in
Berlin to help paying the rent for apartments which had become too
big or too expensive, but whose landladies were usually more concerned
with the wellbeing of their bed spreads and frying pans than of their
tenants. Only later was she able to afford, together with her sister
Gertrud, a rented apartment of her own. She was able to continue some
genetical research, mainly on phylogeny and sex expression in Fragaria,
but became more and more involved in the history of cultivated plants,
with increasing attention to the prehistoric and archeological aspects.
However, whUe this development was imposed on her by circum-
stances it was fortunately one she did not resent. The new kind of
work matched the interests in history and cultural history she had
brought from her parents' home. Earlier, she had been in touch with G.
Schweinfurth, the great explorer of Africa who, on his voyages
of discovery, had paid great attention to archeology and ancient history.
Now, she estabUshed contacts with W. Andrae, the excavator of the
Ishtar Altar and now Head of the Near-Eastern Division of the Prussian
State Museums in Berlin, from whom she received, first through the
Notgemeinschaft der Deutschen Wissenschaft (an organization formed
to support German Science during the difficult post-World War I
period), fossil plant material from Mesopotamia as well as much sub-
sequent stimulation. With her usual thoroughness she rapidly acquired
considerable skills in the necessary methodology and could soon
compete with "professional" archeologists. She thus did the first steps
on the road that would lead to her most original scientific achievement,
and despite the limited conditions the period from 1930 to 1940
was among the most productive ones in her professional career and
very probably also quite satisfactory in her personal life.
This time came to an end in 1940 when, one might say at long last,
Elisabeth Schiemann did clash with the Nazi system. After this
regime had been established in 1933 virtually every body in Germany
who held some kind of job (and who was not willing and able to
emigrate) had to some extent to adjust to its demands and regulations.
Faculty members had to join the NS Docents' League (Dozentenbund),
students (except foreign ones), the NS Students' League (Studenten-
bund). Some became more or less ardent fellow travellers. Others took
part in the various activities demanded by the "Party", e.g. attending
indoctrination sessions in institutes, First of May marches and parades,
the broadcasts of Hitler's speeches to which, when during work hours,
everybody had to listen, as a group, but managed to exclude these as
much as possible from their professional and personal life, venting
their feelings at most in small groups of trustworthy friends. Elisabeth
Schiemann did not attend the Hitler broadcasts nor the obligatory
sessions of the NS Dozentenbund; she did not take part in the First
of May marches. In her classes and seminars she quoted Jewish and
Russian authors, like Aaronsohn, the discoverer of wild emmer wheat,
and Vavilov (Russian authors were also suspect, since all were "bolshe-
vists"). She was active in the Confessional Church (Bekennende Kirche),
the only major religious group in Germany that remained independent
of and opposed to the Nazis, and she not only expressed her sympathy
for persons who were persecuted for political, racial or religious reasons
but offered them advice and, as far as her, obviously limited possibili-
ties allowed, active support. As a little illustration of her attitude I
like to mention her performance in a small symposium that had been
organized by the student group of the Botanical Museum ? the section
Englera7-1987 25
of the NS Students' League which existed in each department or school -

and was to deal with the topic of hereditary and racial aspects of cul?
ture and science. The main speaker had been imported from the central
Nazi party organization of Berlin; if my memory is right he had the
truly Germanic name of Hadamitzky. He started his speech with the
declaration that there were three human races; good races, poor races,
and the Jews. The following speakers either sought to provide support
for this classificationby citing various examples, or avoided the "Jewish
question" altogether. Only EUsabeth Schiemann got up and stated,
with a clear although slightly breaking voice - as was her wont in times
of great stress or passion ? that we should acknowledge the great
contributions of different people to German culture and science ?
French, Italian, "and yes, let us say it clearly, the Jews." She went on
to name names, too, including scientists and writers who at that time
? it was 1937 or 1938 ? were harrassed by the Nazi authorities or had
already been forced to leave the country. (Interestingly, nobody
challenged her; it is true that Party Comrade Hadamitzky had already
left the meeting.)
AU this could not last too long, and in 1940 Schiemann's pro-
fessorship at Berlin University was revoked and she thus deprived of
her formal position and her ineome. Thanks to the efforts of F. von
Wettstein, Director of the Kaiser WUhelm Institute for Biology, she
was awarded a fellowship by the Notgemeinschaft, the Botanical
Museum continued to provide her with office and experimental plots,
her sister Gertrud helped her in a most unselfish manner. She could
thus pursue her studies to some extent but professionally and psycho-
logically this was very probably the most difficult period in her life,
after the break with Baur. As an ultimate blow, an AUied air raid in
1943 destroyed a large part of the Botanical Museum, including the
excellent herbarium, of which only small parts had been evacuated
to safety, and the outstanding library. Schiemann's room survived,
as if by a miracle, and unlike most of the other scientists in the Museum,
she did not lose her records, notes and materials, but further work had
become impossible. Yet in the same year a change for the better took
place. The Kaiser WUhelm Society for the Advancement of Sciences
decided to realize long-standing plans and to found a new Kaiser
WUhelm Institute for Cultivated Plants Research in Vienna. Hans
Stubbe, Elisabeth Schiemann's colleague from the years with
Baur, was appointed Director, and Schiemann herself was made

head of a Division for the History of Cultivated Plants. This creation
of a large new institute, with the war continuing and in fact its outcome
becoming quite apparent to foresighted people, may seem something
unreal; yet it made it possible to provide a measure of professional
activity and personal security to persons like Stubbe and Schie?
mann, who could not expect employment in institutions that were
under the direct control of the Nazi authorities.
Thus, once again Elisabeth Schiemann became quite busy helping
to plan an institution; once again it turned out to be an activity from
which she was not to benefit herself. For whereas Stubbe and some
other coworkersof the new institute, which had been temporarUy
housed in the vivarium in the Prater - a part of Vienna whose claim
to fame is founded on activities rather different from science, namely,
popular entertainment - and had been assigned experimental plots
in another suburb, Tuttenhofen, moved to Vienna and started some
work, Schiemann stayed in Berlin and was here when the war came
to an end. The Vienna institute was, when the Soviet armies were
approaching the city, to be transferred to Berlin but was overtaken
by the collapse of Germany on the way, in Central Germany. It was
assigned a location in Gatersleben, near Aschersleben, in the then
Soviet occupation zone of Germany and under the dynamic leader-
ship of Stubbe and with strong support by the local German and the
Soviet occupation authorities developed, despite the difficult postwar
situation, surprisingly rapidly into an excellent research center for
genetics, breeding research and related discipUnes. It was moreover the
only such institution in the Socialist part of Europe which remained
free of the "teachings" and practices of Lysenko and his followers,
and in fact did very much to prove the spurious nature of these "teach?
ings." As the Central Institute for Genetics and Cultivated Plants Re?
search it continues to be respected throughout the scientific world.
Schiemann's Division first remained as an external part of the
Institute of Cultivated Plants Research in Berlin, then became part of
the Research University (Forschungshochschule), an umbrella organi-
zation for the parts of the Kaiser WUhelm Institutes that had remained
in Berlin, and finally, in 1953, was incorporated, as a separate Research
Unit (Forschungsstelle) for the History of Cultivated Plants, into the
Kaiser WUhelm, now Max Planck Society for the Advancement of
Englera 7- 1987 27
Sciences. In 1946, Elisabeth Schiemann was appointed Professor

of the Humboldt (formerly, Friedrich Wilhelm) University of Berlin.
Thus, she was able to resume her research and teaching activities and
despite the difficult general situation in post-war Berlin carried them
out with great drive and success. She trained a number of young co-
workers and with their help conducted intensive studies on the origin
of cereals by genetical and archeological approaches, these approaches
being, and this is the major point, complementary and integrated,
and the results a good deal more than the mere sum of the individual
studies. ElisabethSchiemann had plans to make her Unit the center
of a cooperative study program on the history of cultivated plants
in all of Central Europe, but these plans remained on the drawing
board. In 1956 Elisabeth Schiemann, having reached 75 years of
age, was transferred to emeritus status and the Forschungsstelle was
disbanded. Her associates were able to continue their work in various
other institutions but the tight integration of genetics and archeo-
logy, which was characteristic for the Forschungsstelle, was obviously
lost. One cannot help regretting that this institution, unique in the
entire world and in its budgetary requirements much more moderate
than today's institutions for experimental biology, could not have been
continued in some form. However, one may ask whether among
Schiemann's young coworkers or any where else somebody with
her experience and vision for such a task could have been found.
As an emeritus Schiemann continued to be quite active. She at-

tended seminars and conferences, and maintained contacts with various
colleagues, mostly by letters composed in her large, clear and firm hand-
writing. On my last visit with her, several years before her death, she
showed me some malformations on plants on her balcony; she was much
intrigued by these and planning to report them in a short paper, but it
seems she was not able to complete this project. She received quite
a number of honors and distinctions: the Order of Merit of the Federal
Republic of Germany; honorary memberships of the French Botanical
Society and the Zoological-Botanical Society in Vienna; election to
the Leopoldina Academy of Naturalists and the Darwin Medal of this
Academy; an Honorary Doctor's degree in Agronomy from the Tech?
nical University of Berlin; Honorary Vice Presidentship of the Xllth
International Congress of Botany. She died in early 1972 and was
buried in the cemetary of the old Village Church of Dahlem, of which
she had been a faithful parishioner and whose simple but dignified ex-
terior reflects much of her own personality. Her memory will live
with us, however, because of her many and diverse accomplishments.
She showed, without ever claimihg any preferential treatment or equal
rights, that a woman can have a satisfying career in Science. She pur-
sued, despite grave set-backs and difficulties, her career with devotion
and ultimately with great success but never deviating from her prin-
ciples. She succeeded, by dedication, intuition and vision, to integrate
genetics and archeology and thus to demonstrate that "Science" and
"Arts," whose alleged incompatibility is so often deplored, can engage
in productive collaboration, provided the right problem and the right
person can be found. And she remained undaunted and uncompromi-
sing under a perverted, anethical political system, trying in whatever
way she was able to help individuals who were suffering under this
system. In all these respects she has set a shining example for scientists
in general, men and women alike.
A considerably more elaborate deseription of Elisabeth Schiemann's
life and career, written by H. Kuckuck and including a complete
bibliography, is published in "Berichte der Deutschen Botanischen Ge-
sellschaft", vol. 93, No. 2, pp. 517-537, 1980.
Address of the author:

Dr. Anton Lang, MSU-DOE Plant Research Laboratory, Michigan
State University, East Lansing, MI 48824-1312, USA.
Englera 7-1987 29
Dieter Mollenhauer
History of phycology in Berlin
Like each other part of human culture botany is influenced by the

main guidelines of each epoch of history. On the other hand, it plays
also an active role since it participates in working out our concepts of
nature which are tied to specific times. Thus, it is in the end also one
of the working agents whose cooperation results in the changing ideas
of what mankind may be. However, the botanists' part in general in-
tellectual history is only an inferior one. Very few men of plant science
have been induced by their professional work to contribute to life of
the mind. Current books on intellectual history rather seldom pay
attention to events occurring in science. The few exceptions from this
general rule are mainly men of medicine, physiology and physics whose
reflections on bearings of their professional work become popular and
attract some attention of the public. Studying plants for the sake of
plants as do people who are destined for botany is no profession to
make men whom the public considers to be engaged on the cultural
sector. Goethe is one of the very rare exceptions. Obviously the average
botanist ? apart from such genius ? lives in the honest way of normal
people with decent professions and appears not extraordinary, espe?
cially not a subject of special historical interest and worth to be docu-
mented. The discreet lives of phycologists are difficult to reconstruct.
As long as phycology has been established it has not been a crowded
profession. There have been and still are few people who spend all
their time and effort in studying algae. Homo botanicus subsp. phyko-
philos only lives in small populations and restricted to very specialized
habitats. This subspecies is an endangered one as are many with ste-
noecious life patterns. It reproduces irregularly by hybridization in an
allopatric and polytopic manner. The representatives of this taxon
grow up here and there, often unpredicted. The historian of phycology
has the hard job not only to characterize the organisms themselves and
30 Mollenhauer: Phycology in Berlin
to study their lives. He must also analyse the substrate on which they
are grown well keeping in mind that their cultivation is difficult and
often unsuccessful. In many cases he is bothered by extremely long
incubation phases after which a new specimen of the rare subspecies
may appear suddenly from nowhere.
The subspecies is a late product of evolving botany. The prerequi-
sites of phycological work have been accomplished only in the be-
ginning of the 19th century. And, of course, arising phycology was no
bread-andbutter-profession from the very beginning. Early botanists
were famUiar with all kind of plants and their different modes of life
since knowledge at that time was rather restricted. The high amount of
botanists who did much other work beside phycology is characteristic
of long periods of botanical history, up to the early 20th century. Many
of them were able to do research on rather different types of plants. So
we know L. Diels (1874-1945) as a man of high reputation for his
work on higher plants and plant geography. However, he wrote a paper
of fundamental importance on algae on bare rocks ("Die Algenvege-
tation der Siid tiroler Dolomitriffe", 1914) and, in addition, also a
chapter on Schizophyta in 1942 for "Handbuch der Biologie". C. E.
Correns (1864-1933) is one of the three rediscoverers of the Men-
delian laws and for this well-known from all textbooks. But he started
his botanical carreer with a doctoral thesis on algal cell walls in Munich
in 1889 and knew algae well so that he was enabled to describe a new
genus Naegeliella (Heterochrysophycideae). P. Graebner's opus
magnum "Die Heide Nord deutschlands" (The heathland of northern
Germany, 1901) contains much information on soil algae. Even far
beyond the frontiers of botany phycological work has been done, e.g.
by T. W. Engelmann (1843-1909), the successor on EmU du Bois-
Reymond's chair of professor of physiology. With his papers and
those by his student N. Gaidukov the long discussion on chromatic
adaptation of algae was initiated in about 1885 lasting nearly 50 years.
The debates culminated when two vehemently fighting parties had been
established. The first shared the ideas of Engelmann-Gaidukov. Accor-
ding to these an alga is best equipped to live in deepwater if its pig-
mentation is exactly complementary to quality of available light. The
opponents followed Berthold-Oltmanns who laid special stress on the
compensation point of assimilation. Its low position on the scale was
considered the decisive advantage for life in deepwater. F. Gessner
Englera 7-1987 31
(Hydrobotanik, vol. 1, 1955) has given an excellent survey of the whole

problem including its historical background and the final reconciliation
of the two theories. He showed that the seemingly incompatible con-
cepts in reality were due to overemphazisation of one among several
aspects to be considered. At present, the concept of chromatic adap-
tation is well established and plays an important role in the taxo?
nomie study of clonal strains of planktic Cyanophyta differing in
colour and growth capacities. Thus, the discussion hundred years ago
has revealed an important and prolific field of phycology.
Obviously, the whole of all phycologists includes such people with
an extremely wide scope of knowledge and interest whose achievement
is handed down to posterity as well as many average persons of algal
science.We also find the variety of coming experts represented by
people who were interested only in some few aspects of algae. They got
in touch with these plants in the course of their proper particular
studies. So the diversity of phycologists of the past includes up to the
beginning of the 20th century many types of men of plant science who
were familiar with algae and/or published phycological contributions:
morphologists, students of developmental history, taxonomists, plant
geographers, physiologists, ecologists, cytologists, here and there also
ethnobotanists, parasitologists etc. O. C. Schmidt (1900-1951) was
the first to specialize exclusively in algae by the time while he was
belonging to the staff of the Botanisches Museum Berlin-Dahlem
(1922-1939) and thus paving the way for his follower J. Gerloff
(* 1915) who was by appointment curator of algae (1956-1979). The
heterogeneity and versatility of old phycologists makes historiographer's
work in this field a very hard job. It really requires professional training
in tracing out and evaluating historical sources. This is much more than a
supplementary job for a professional phycologists as is the present
author.
Berlin has been a provincial city for long time. Only in 1700 the
famous philosopher and universal genius G. W. Leibniz (1646-1716)
suggested the foundation of an academy of sciences. Friedrich of
Prussia who later on became King Friedrich I and his wife Sophie
Charlotte took up the idea. The Friedrich-Wilhelms- Universitat (now
Humboldt-Universitat in East Berlin) has been founded in 1810. There
have been earlier institutions and further attempts to advance scien?
tific research and teaching and to exchange scientific information. These
were supplemented by societies of naturalists or circles of amateurs

meeting regularly to report on their personal research studies (e.g.
Gesellschaft Naturforschender Freunde founded in 1839). So there
were some different attempts to promote science. But together with the
whole of botany phycology got along for many years with some tran-
quillity. A first impetus to activate scientific life came with the German
Wars of Liberation (1813-15), another and more vigorous one with the
period of rapid industrial expansion in Germany (1871-73). Berlin
became by enhanced development a metropolitan city not only at-
tracting thousands of industrial workers but also many educated people
who congregated to form a unique intellectual potential of teachers,
students and scientific staff personnel.
The residenceof the Prussian kings was for some years one of the
centres of Romanticism. A somewhat detached deseription of this
cultural epoch is given by Heinrich Heine in his essay "Die romantische
Schule", written in Paris, 1836. The romantic scientists were fond of
allembracing concepts. They considered the world a hyper-organism
ruled by common laws for all subsystems in microcosms and the
macrocosm. The leading minds of the romantic "Naturphilosophie"
(Goethe, Oken, Burdach, Kieser) were living elsewhere, not in Berlin.
But their influence overcame the whole of Germany. One of their
central ideas was that of metamorphosis, so this can be traced out in
science under many different aspects. It was generally adopted that
living beings are capable of fundamental transformations causing
extreme alteration of form and shape. Polymorphism was quite usual.
The Hollandsche Maatschappij der Wetenschapen wanted to obtain
univocalness concerning metamorphosis of algae and infusoria by means
of a prize question (for the years 1835 and 1836). F. T. Kutzing,
leading phycologist of his time, was the winner of this competition with
his paper "Die Umwandlung niederer Algenformen in hohere, so wie
auch in Gattungen ganz verschiedener FamUien und Klassen hoherer
Cryptogamen mit zelligem Bau" (Haarlem 1841). One of the subjects of
discussion was "Priestley's green materia", the well-known thin green
layer of algae which covers water in flower vases etc. after being left
uncovered for some time. Mey en, otherwise a critical and far-sighted
plant physiologist, also had contributed in 1827 to the current discus?
sion among his colleagues who tried to interpret the enigmatic for-
mation of Protococcus (i.e. "primary sphere", in German "Urkugel") to
Englera 7-1987 33
filamentous algae (Confervae). A formative power was said to create in

a corresponding manner all organismic constructions on all the different
levels of integration and organization. Kiitzing's contribution by his
prize essay consisted in explaining the relativeness of the concept of
algal species and of organismic species as a whole (cp. Miiller & Zaunick
1960: 236). Thus his promising work was far ahead of his time since
the methods needed to refute or produce evidence for such theses were
not yet worked out. He pleaded for carefully registering of all the
different transformations which a given organism can undergo and to
distinguish species only on the basis of this full survey. What he had in
mind was called "Artperiode" (period of a species) by H. A. de Bary in
1858. And in 1950 the entomologist W. Henniglaid special stress on
the fact that ontogenetic changes in all organisms result in a series of
so-called "semaphoronts" which altogether constitute the respective
taxon. Today this is phycological routine work:
Careful description of complete life cycles. But the isolation and
culture techniques required for such investigations were not available in
the days of Romanticism. R. Koch was the first to use the pure culture
technique by pouring gelatine plates of bacterial culture media in Petri
dishes. And M. W. Beijerinck and G. Klebs modified the bacteriological
technique and adjusted it to phycological purposes (see Pringsheim
1954). The men of algal science in Kiitzing's days only could discuss
their microscopical observations of life material from nature and what
they could obtain in impure rough cultures. All their results were
depending on personal experience and in many respect became a matter
of opinion. Just a romanticist (however one to whom this title can be
attributed only under reserve, if Heine was correct) insisted on constant
taxonomie groups. It was A. von Chamisso (1781-1838; see Schmidt
1942), poet and biologist, participant of a scientific circumnavigation
of the world, discoverer of the alternation of generations in Tunicates,
botanist at the Royal Herbarium of Berlin. His romantic poetry is still
known to educated people of our days. However, in botany he was a
decided opponent of polymorphisms and transformation. "Of course, I
have written fairy-tales in my poetry. However, science is quite differ?
ent, and I take care to stick to pure observation and not to add phan-
tastic speculations. The kind of Nature as is pretended by the adherents
of universal transformation and metamorphosis is not my Nature. My
mind cannot live with such kind of Nature and is upset by ubiquitous
metamorphosis. There must be constant genera and species, otherwise

they have no reality." (Translated from "Reise um die Welt", 1836,
written in winter 1834/35.)
A debate with controversies of such type cannot be neutralized
and cannot result in agreement. Actually its subject is the crucial point
of all taxonomy. It is a constant challenge to find out which reasons
are decisive for distinction of species. It is the prime necessity of all
science to make these reasons known. Romanticism was a fascinating
epoch of discussion with a high standard of philosophy. Later on
science abandoned this consciousness of intellectual obligation to give
overt evidence for all conclusions. Nowadays we begin to understand
the scientificconsequences of this omission. A general feeling begins to
emerge that history and theory of science cannot longer be excluded
from actual investigations otherwise the implications remain obscure.
This is one of the reasons of the Renaissance of theoretical biology and
its historic branch which turns to Romanticism. This epoch has been
discredited by those who grew up in the time which immediately followed
this epoch and whose guidelines were deeply influenced by Neokan-
tianism and positivism. Today, Romantic science has become a model
of research bound to explicite theories while epistemologists of today
reproach actual science to operate with unrecognized philosophic
propositions and, in consequence of this, producing arbitrary know-
ledge.
About the middle of the 19th century the ideas of Romanticism had
its days. However, the thought patterns had become independent and
were perpetuated by a new generation of scientists who considered
themselves unbiassed and freed of the chains of "Naturphilosphie".
Observation and imagination which then was left in uncontrolled arbi-
trariness formed a terrible muddle. Thus we can see the troublesome
and painstaking detailed work to be done by N. Pringsheim (see his
collected papers, 1895) to unravel the true nature of cell development
and fertilization in algae. Moulded by current ideas of late Romanticism
and later on dragged into a time when his country was on the point of
emerging he could not achieve final clarity without going long wrong
tracks. Still in 1852 he considered it well possible to find spermatozoids
in Spirogyra. Under such circumstances it is evident that H. A. de Bary
in his famous "Untersuchungen iiber die Familie der Conjugaten" (In?
vestigations in the family of Conjugates, 1858) needed many pages to
Englera 7 - 1987 35
discuss all the current misinterpretations of his time, many of these

claimed by most honorable colleagues. E. g. fungi living as intracellular
parasites of algae often were mistaken for spermatozoids.
In the course of time polymorphism had become an ideology whose
adherents sometimes were rather unteachable. In the 2nd edition of his
book "Morphologie und Biologie der Algen" (vol. 3: 69) F. Oltmanns
wrote: "Whenever we find the notion polymorphism or pleomorphism
in older or even in rather recent literature dealing with fungi, bacteria
or algae this meant nothing but confusion of more or less numerous forms
and the pretention that these belonged together to form a common life
cycle though they were by no means interconnected in reality. Klebs
refers to the fact that all these false doctrines had to be uprooted suc-
cessively and each of them separately, in all three cases, i.e. for fungi,
bacteria and algae. The last to be cancelled was the polymorphism of
algae." Longevity of ideas of pleomorphism was a matter of the level
of organization. It was the tougher the lower this level was. W. Z o p f
(1846-1909) was one of the leading adherents of an extreme pleo?
morphism of bacteria (e.g. his paper in the proceedings of the Prussian
Academy of Sciences and Letters "On the genetic interrelations in
Schizophyta" in 1881). Likewise, H. Itzigsohn, physician in Neudamm
(Mark Brandenburg), cooperator and actively corresponding with Alex?
ander Braun and enthusiatic algae collector for the Rabenhorst ex-
siccata, wrote on the so-called "diamorphosis" of bluegreen algae by
which he understood an extreme plasticity resulting in long series of
connected forms which he all thought to belong to the same species.
Exaggerations of this kind provoke contradiction. Scientists of the
following generations took them as challenge for reinvestigations. Thus
these errors were rather productive for further research. It is not sure
whether less provoking conclusions by their moderate appearanee
would have entailed an equal activity. The situations was similar as with
Ehrenberg's papers on infusorian cell anatomy. Furthermore it
also should be stated that the men of metamorphosis in their way were
rather competent observers and collectors to whom we owe a lot of
discoveries. Alexander Braun was one of the most active herbarium
collectors and so enriched the collections considerably by many ex?
cursions into the surroundings of Berlin. Foreign guests often accom-
panied him as we can see from herbarium collections in the materials of
P. Magnus (1844-1914) who lived and worked in Berlin. His left
collections were bought by the Hamburg Institut fiir AUgemeine Bo?

tanik and now are stored in the Herbarium Hamburgense. There we find
collections by H. A. de Bary made in Berlin during a guided tour with
Braun. It was also Braun who discovered the first lacustrine brown
alga in the lake Tegeler See in the north of Berlin (Pleurocladia lacus-
tris, distributed in 1855 as one of Rabenhorst's exsiccata in "Algen
Sachsens").
In the same manner as elsewhere phycology in Berlin began with
geobotany: "Floras" and reports on finds of remarkable species. An
early highlight of algal research is the life's work ofC. G. Ehrenberg
(1795 1876; see Hanstein 1877) who was Hon. Secretary of the Prussian
Academy for long years. He was professor of zoology and natural
history and is famous for his pioneer work in research on infusoria, soil
biology, hydrobiology (both lacustrine and marine) and paleontology.
Part of his results was outdated rather soon since they consisted in
refutable statements concerning the internal organization of protozoa
and unicellular algae. However, it has been Ehrenberg who directed
attention to the world of protista (name given to them by Haeckel in
1866). The discussion on their structure and life and the critical rein-
vestigation of his results initiated modern cytology and protistan
research. Cell theory was for long central theme of physiology what can
be seen from contributions by R. V i r c h o w, J. M ii 11 e r, E. du B o i s-
Rey m ond and N. Pringsheim.
The early years of rapid industrial expansion in Germany and their

preliminaries gave Berlin some touch of pioneer behaviour and caused
a similar impression as the WUd West. The general aspect of the commu-
nity was governed by speculation and daring enterprises. When Heinrich
Heine came from Dusseldorf to attend the University of Berlin all this
was already about to start. It caused many preliminary and provisional
arrangements and the general impression was governed by the lack of
steady growth and solidity. It was a state of seething. Berlin was soon
in full expansion. No time was left for contemplation and introspective-
ness as in the preceding eras of classicism and Biedermeier, all was activ?
ity. Mental culture, however, declined. The spirit of the time was in-
fluenced by prosperity, progressiveness, unphUosophic or even anti-
philosophic estimation of professionalism. On the other hand the over-
crowded upstarting new metropolis was forced to be creative in solving
its growing problems of urbanization. In Berlin they came into being
Englera 7-1987 37
considerably earlier than elsewhere in Germany thus causing the inven-

tion of exemplary arrangements. In this situation the Prussian people
had to show the results of its long period of learning the hard way. It is
widely adopted to correlate with the keyword "Prussia" nothing but
distorted pictures: conceited officers, nationalism and militarism, nar-
rowminded bureaucracy and presumption. Of course, all these excesses
did exist and got into the limelight. Since they obstructed the view to
positive traits of character, viz. sense of responsibilty, restraint, dis-
cipline and altruism, these are to be demonstrated by their products.
At that time, there was a lot on in Berlin. This atmosphere of departure
was especially impressive for the students of the University which began
to flourish. Maybe it was of higher importance than the subjects them-
selves which were the matters of their education. The general feeling
moulded all who accumulated in the newly qualified capital in order
to teach and to learn. In Berlin people were open for E. HaeckePs evolu-
tionism. Here Ernst Krause (alias Carus Sterne, 1839-1903)
could successfully popularize his ideas on developing nature. In the
course of the theories of naturalism W. B61sche(1861-1939) pleaded
for a reorientation of the intellectuals in order to incorporate nature
and its rules into the cultural tradition and to consider its topics worth
to be dealt with in poetry. After Romanticism we can state a growing
indifference of poets and men of literary profession in subjects of
biology and of nature at all. Later on scepticism and even resistance
against the pair formed of nature and technology marked the scene. T.
Korner, poet of the German Wars of Liberation, is one of the examples
demonstrating the opposite in the times of Roman ticism. When he
studied in Berlin he attended the complicated philosophical lectures by
Fichte as well as those on zoology by H. Lichtenstein(1780-1857),
founder of the Zoological Garden. The estrangement between people
interested in nature and those whose culture is exclusively literary is a
matter of the 19th century. The rise of engineering science induced the
countermovement and also promoted the popularity of science and
favoured its incorporation into the subjects of instruction and edu?
cation. Realistic and naturalistic poetry again helped to make topics of
biology reputable. All such trends are typical matters of culture in
Berlin towards the end of the 19th century.
Concepts of school instruction in natural sciences were forwarded.
In 1887 a Deutscher Lehrerverein fiir Naturkunde (German Teachers'
Association for the Teaching of Natural Sciences) had been established,

even earlier, in 1885, the Berliner Lehrerverein (Berlin School Teachers'
Association) had founded a special branch Naturwissenschaftliche
Vereinigung (Natural Science Consortium). Since 1888 a special insti?
tute of adult education, the Kulturgemeinschaft Urania organized
popular lectures in science. The activities by F. v. R au m er to establish
public libraries were initiated in 1850. In 1897 the staff of the Berlin
University began to offer public lectures. Journals with the same
purpose were much older, e.g. "Die Natur. Zeitung zur Verbreitung
naturwissenschaftlicher Kenntnis und Naturanschauung" (published by
Deutscher Humboldt-Verein). AU such associations paved the way for
further specialization. This started with the foundation of the Bota?
nischer Verein der Provinz Brandenburg in 1859. Later on several
groups and associations of amateur microscopists were established,
some only local or regional, others with members all over the empire.
The best known of the latter is the Mikrobiologische Vereinigung Stutt-
gart (founded in 1907) with its well-known journal "Mikrokosmos".
Furthermore, the traditional associations as the Gesellschaft Natur-
forschender Freunde grew since people from further sections of the
population joined them, rejuvenizing the membership and broadening
the scope of interest.
With increasing prosperity and leisure time the number of amateur
phycologists increased. These always have characterized algal research
in Berlin. A big city may have a lot of them among its inhabitants.
These included many excellent specialists whose vast knowledge and
famUiarity with rich materials enabled them to write floras and mono-
graphs. Here we also can find the inspiring teachers and the enthusiastic
students. In associations of naturalists future scientists found a forum
to meet experienced specialists and contact them personally. So they
early came into contact with organismic forms in their variety. This
caused intensified knowledge about nature and gave the opportunity to
envisage many species at an age which is much more sensitive to im?
pressions of this kind than the age of a college student. Broad basis and
multiple and intensive mutual exchange of experience between ama-
teurs and professionals always has been a typical feature of phycology
in Berlin.
So phycologists in Berlin belong to rather different circles of the
population. In the beginning we see many typical scholars, e.g. C. G.
Englera 7-1987 39
Ehrenberg, H. F. Link (1767-1851) or F. J. F. Meyen (1804-

1840). They were Humboldt's contemporaries and his influence is
evident in their studies. Associated to them we find Chamisso (du
Bois-Reymond 1888, Schmid 1942). Alexander Braun was the
famous representative of idealistic (Goethean) morphology (1805-1877;
see Caspary 1877, Mettenius 1892). N. Pringsheim (1823-1894),
too, was a scholar and also a person of good standing who therefore was
free to live the life of a private savant (see Cohn 1895, Magnus 1895;
also Pringsheim 1895). He chose his subjects on his own risk and his
own responsibility. This favoured the steady and organic progress of his
investigations and enabled him to work out carefully, step by step,
a survey of structure and reproduction of algae. We can state a gradual
approximation to his final views by repeated observation and critical
reconsideration of the own interpretation. Educated with the Talmud
he was well-prepared for sophisticated intellectual exercises. Further-
more he was completely incorporated into German culture what made
him go the complicated ways unlike Thuret or other Frenchmen with
their more rational way to look upon nature.
Later on we find a considerable number of specialists, collectors and
observers, like
O.Muller (1837-1917)
L. Holtz (1824-1907)
P. Hennings (1841-1908)
M. Marsson (1845-1909)
G. Hieronymus (1846-1921)
G. Lindau (1866-1923)
R. Pilger (1876-1953)
K. Ge mein hard t (1883-1952)
WilliKrieger (1886-1954)
E.Lindemann (1888- ? )
H. Beger (1889-1955)
A. Donat (1893-1937)
H. Bethge (1885-1961)
K. H611 (* 1901)
HansKrieger (1913-1943)
Such work results in intimate knowledge of particular details. Know?

ledge of this kind can decline to sterile lexikography unless someone is
able to organize it in a didactic manner by means of handbooks, text-

books etc. One such great organizer was A. Engler (1844-1930).
K. Goebel in Munich, unlike Engler a typical individualist, liked to
tease the Dahlem crew calling them a botanical "factory" of Mr. Privy
Councillor. A botanist of the type of an assembly line worker and
manufacturer was far from Goebel's ideal. But one can also hold
completely different views concerning the work in the Botanisches
Museum Berlin-Dahlem. Many of the staff members considered them-
selves obliged to renounce on a personal career, which as we all know
is done mostly at the expence of others. They followed the Prussian
ideal to do service for the scientific community. Thus they worked out
contributions for the great handbooks which are in use up to now. Well
equipped research centres are needed to elaborate such monumental
works as well as welltrained specialists who are willing to cooperate.
Engler and his staff was not the only example. In a less spectacular
(and also a less authoritarian) manner Kolkwitz (1873-1956; see
Huber-Pestalozzi 1956) brought about similar organizational great
achievements. The work of both men resulted in the following publi-
cations which were elaborated in Berlin or directed from here:
Syllabus der Pflanzenfamilien, 12 editions (1892-1964).

Die naturhchen Pflanzenfamilien, first edition 1887-1915, second
edition 1924-.
Hedwigia, founded by L. Rabenhorst in 1852, from vol. 32
(1893) editors from the staff of Botanisches Museum Berlin-
Dahlem, up to 1944. Tradition continued by "Nova Hedwigia",
founded in 1959 by J. Gerloff and F. Mattick, now edited by G.
Frey, U. Geissler, J. Poelt. Kryptogamenflora der Mark Bran?
denburg, organized by Botanischer Verein der Provinz Bran?
denburg (now: Berliner Botanischer Verein) in cooperation with
many staff members of Botanisches Museum Berlin Dahlem.
Publisher: Borntraeger. 2 phycological volumes: Lemmermann
(1910): Schizophyceae, Flagellates, Peridinea. Holtz (1903):
Charophyta. HUfsbuch fiir das Sammeln und Praparieren der
niederen Kryptogamen by G. Lindau. 2 editions (1904, 1922),
with cooperation from staff members of the Botanisches Mu?
seum. Publisher: Borntraeger.
Kryptogamenflora fiir Anfanger. 2 editions (1914-). G. Lindau,
R. Pilger, H. Melchior. Publisher: J. Springer Berlin.
Englera 7-1987 41
Rabenhorst's Kryptogamenflora von Deutschland, Osterreich und

der Schweiz. 2nd edition of the volumes on algae directed by R.
Kolkwitz from 1935 on. Publisher: Akademische Verlagsge-
sellschaft Leipzig (originally: Kummer publisher).
Siifiwasserflora von Mitteleuropa, new edition of the well-known
work whose 1st and 2nd edition (incomplete) has been directed by
A. Pascher (Prague). New edition started by B. Schussnig, Jena
GDR (1892-1976)
Y together with J. Gerloff (Berlin) and H. Ettl
(CSSR). Cooperatively published by VEB G. Fischer Verlag Jena
(DDR) and G. Fischer Verlag Stuttgart and New York.
Pflanzenforschung. A series of monographs, many of them with
phycological subjects (e.g. Krieger, Gemeinhardt, Donat, Hom-
feld, Canabaeus, Wichmann, H611 and others). Founded and edited
by R. Kolkwitz (1922-). Publisher: G. Fischer Jena.
Diatomeenschalen im elektronenmikroskopischen Bild, Atlas. 8
volumes, beginning in 1953, by Ursula Geissler, J. Gerloff, J.-G.
Helmcke and W. Krieger. Publisher: J. Cramer, Weinheim, Lehre,
Braunschweig.
Jahrbucher fiir wissenschaftliche Botanik. Well-known botanical
journal, founded in 1857 by N. Pringsheim. Publisher: Born-
traeger, Leipzig, Berlin.
The genius loci of Berlin is characterized by metropolitan features:

many people, many talents, teamwork and other cooperation. Other-
wise Berlin is a city lying in a beautiful landscape which also deeply
influences the inhabitants of the former capital of Germany. Principally it
is possible to do phycology wherever you want, even in dry or cold
deserts. But it is obvious that biologists are stimulated to look for
aqueous organisms in a region full of lakes, small rivers, swamps and
peat-bogs. The glacial landscape around Berlin is of this type. Even the
political island of West-Berlin has a built-up area of only 36 % of its
ground, and 6 % consist of waters (for comparison: the Federal Repu-
blic of Germany comprises 1,8 % of its ground with lakes, rivers and
other type of water, the GDR 1 %). No wonder that aqueous ecology
and limnology continue to represent main research subjects of the
scientific institutions in both parts of the divided city.
When Gleditsch began to collect material for his "Floras" the in?
habitants of Berlin already used their nice surroundings for excursions
to the countryside. In the beginning this was a privilege of well-situated
people or the unusual profession of collecting naturalists. Later on it

was everybody's pleasure to go out to the open air ("ins Griine" as we
say in Berlin). Thus all kind of water sports and also steamer excursions
became very popular and a dense system of pleasure boat services devel-
oped all around Berlin. Many lakes and lowland rivers became easily
accessible and this, in turn, was good opportunity for many collections.
Not only the resident botanists but also their visitors went out for
sampling algae. Only part of all these collections is documented in the
literature. Many additional data can be derived from reinvestigation of
the preserved herbarium specimens. This rich stock of data and herbarium
collections (some of which have to be reconstructed from double speci?
mens which have been forwarded to herbaria outside of Berlin and thus
have escaped destruction during World War II) enables scientists in
Berlin to do exemplary research in civUization and urbanization eco?
logy. Much research on the history of landscape and human settlement
has been done in Berlin. Its comparison with the stored collections and
recorded finds of former times yields rich information on bioindicators
of urbanization.Rather recently some interesting documents of algae
recorded from Berlin in the late 19th century have been detected in
Herbarium Hamburgense (Herb. P. Magnus).
Berlin is a city of lakes and forests, but also the largest industrial
agglomeration of Germany and the region with the highest population
density. Lakes and rivers in Berlin and its surroundings are therefore
deeply influenced by industrialization since more than hundred years.
Protection of the environment and urban hygiene had to do pioneer
work in Berlin.
This led to a valuable supplementation of purely scientific phyco?
logy. A wide scope of applied science includes waste water research,
hygiene of drinking water, general urban hygiene etc. Important pio?
neer work has been done by the former Preussische Landesanstalt fiir
Wasser-, Boden- und Lufthygiene (founded in 1901), now Institute
for Water, Soil and Air Hygiene at the Federal Health Office (Bundes-
gesundheitsamt, Institut fiir Wasser-, Boden- und Lufthygiene). Ob?
viously modern drinking water supply and waste water treatment
were required rather early in the industrial agglomerations. R. Virchow
who was not only professor of pathology but also deputy in the Reichstag
pleaded very early for modern urban hygiene. Resulting from the
analysis of epidemic diseases in London and Breslau F. Cohn initiated
Englera 7-1987 43
microbiological analysis of drinking water and introduced his student C.

Mez into this new field of research which was further organized by
him. Using these and several other preliminary studies R. Kolkwitz
and R. Marsson worked out a standard of biological examination of
water quality ("Grundsatze fiir die biologische Beurteilung des Wassers
nach seiner Flora und Fauna", 1902) whose further revision lead to the
well known saprobic system by which water quality is expressed by
plant and animal indicator species according their capability to stand
under polysaprobic, mesosaprobic, oligosaprobic etc. conditions. Today
this system is reliable standard of Central European water research
(Kolkwitz 1935).
Applied sciences, too, are in close relation to biology, and especially
fishery research has much in common with limnology and hydrobiology.
Berlin has a remarkable tradition in forestry and agricultural research
what can be derived from the reputation of men like J. G. Gleditsch
(1714-1786), A. S. Marggraf (1709-1782), E. Mitscherlich
(1794-1863) and, of course, A. Thaer (1752-1828). The Landwirt-
schaftliche Hochschule (Agricultural College) existed since 1881.
The Lehr- undForschungsanstalt fiir Gartenbau (Institute of Edu-
cation and Research in Horticulture) was founded much earlier, already
in 1824. In 1880 there was a first call to establish the Reichszentrale
zur Beobachtung und Vertilgung der die Kulturpflanzen schadigenden
Insekten und Pilze (Imperial Centre for Observation and Combat of
Insects and Fungi affecting Cultivated Plants). However, only in 1898
the Biologische Abteilung fiir Land- und Forstwirtschaft im Kaiser-
lichen Gesundheitsamt (Biological Division for Agriculture and For?
estry at the Imperial Health Office) was institutionalized. Its first
head was A. B. Frank (1839-1900), a man rich inideas and extremely
versatile to whom we owe the final proof of nitrogen fixation by
bluegreen algae. Since he was convinced of this fact but uncapable to
offer evidence he came back to this point again and again. Thus he
insisted on keeping this question on the agenda and his perseverance
finally became justified by K. Drewes' unquestionable experiments in
1928. The general importance of thisproper ability of Cyanophyta, not
only in paddy fields, is well known in soil science. There were many
interrelationships between general work with cryptogams as is done at
the University, the other colleges and the Botanisches Museum on one
hand and research at the Reichsanstalt on the other. Several phyto-
pathologists started their scientific carreers as phycologists. This is true

especially for H. W. Wollenweber (1879-1949) whose first publi-
cations concern Haematococcus (Thesis on this alga in 1907).
In fishery research phycology plays an important role, especially
plankton research is done by many pertinent specialists. Already in
1893 J. Frenzel (1858-1897) established the Biologische und Fische-
reiversuchsstation 'Muggelsee' (Research Station in Biology and Fishery),
later on Preufiische Landesanstalt fiir Fischerei in Friedrichshagen, now
Institut fiir Binnenfischerei der Akademie der Landwirtschaftswissen-
schaften der DDR. Frenzel was hydrobiologist and specialized in
crustacean research. Preliminary work for his research station already
began in 1886 and it was modelled after the pattern of O. Zacharias'
Biologische Station Plon (today: MaxPlanck-Institut fiir Limnologie,
Plon). A lot of important papers concerning algae in general, phyto-
plankton, lake types, classification of water bodies by algae etc. have
been published by the Friedrichshagen staff members and their guest-
workers (O. Muller, E. Lemmermann, J. Frenzel, H.-H. Wundsch,
Eva-Maria Stange-Bursche nee Bursche, Marie-Luise Albrecht,
Marlies Schluter).
Another important feature of scientific research in Berlin is the

interplay of science and industry resulting in the introduction of new
instruments and new methods. This was considerably facilitated by the
KaiserWilhelm Institutes (now research institutes of the Max-Planck-
Gesellschaft). Their special adjustment can be described as laboratories
equipped in an open-handed manner for the proper needs of eminent
scientists. They gave and stUl give them the opportunity to work in the
atmosphere of private scholars thanks to generous financial support,
mainly from industry and partly from the public. Different from
university professors they are exempt from teaching duties but free to
choose students at wUl to educate them in their labs. AU this intellec?
tual liberty favours fondness of experimenting. The Kaiser-WUhelm-
Gesellschaft zur Forderung der Wissenschaften was founded in 1911,
its Institut fiir Biologie began to work in 1914. It included among
others the departments for protist research (Head: Max Hartmann,
1876-1962) and physiology (Head: O. Warburg, 1883-1970; see
Krebs 1979). A lot of trend-setting papers in general biology, research
on sexuality, photosynthesis etc. resulted from the work in KWI fiir
Biologie Berlin-Dahlem. The contributions to "Archiv fiir Protisten-
Englera 7 - 1987 45
kunde" (founded in 1902 by F. Schau dinn, 1872-1906, coedited by

Hartmann from volume 8 onwards) came to a large extent from the
Dahlem Institute and were supplemented by further congenial studies.
The journal was for years a standard of reference of research on protists
and in general biology. Decennia of German biology got the mark of
the Hartmann era (see Nachtheim 1963) ? preponderance of a single
eminent scientist which is somewhat problematic and has caused
internal trouble (especially arising from Hartmann's complete
incompetence in the field of ecology which he categorically excluded
from scientific biology). All students of this era are directly or in di?
rectly influence by Hartman n's standard works ("Allgemeine Bio-
logie" from 1927, 4th edition in 1953; "Sexualitat" from 1943, 2nd
edition in 1956; "Die philosophischen Grundlagen der Naturwissen-
schaften" from 1948, 2nd edition in 1959). In all his publications (see
also Hartmann 1956) Hartmann refers to his studies on protists and
lower algae done by himself and by his coworkers (Hartmann:
phytoflagellates; Moewus, a man whose papers are much disputed:
Chlamydomonas and related genera with special reference to their
sexual phenomena; Hammerling and the whole "Acetabularia-
Schule"; Witta Lerche: Dunaliella; Birgithe Foyn: Ulva, Clado-
phora). A lot of current textbook knowledge has been worked out by the
KWI fiir Biologie staff, mainly by Hartmann himself, this includes
the concept of relative sexuality resulting from experiments with
Ectocarpus. There is also some relation between the KWI studies and
the work of H. Kniep (1881-1930) who during his time as professor
of plant physiology (1924-30) in Berlin wrote his book "Sexualitat der
niederen Pflanzen" (1928). O. Warburg was appointed head of
department of physiology in rather young years and remained KWI
scientist (or MPI scientist, respectively) for 56 years. His studies on cell
respiration are only part of his eminent life's work (Nobel prize 1931).
The versatile biochemist and physiologist used microalgae in several of
his studies and we owe to him the introduction of these organisms into
general laboratory practice in biochemistry.
Industrial research and Kaiser-Wilhelm-Gesellschaft also cooperated
in the development of the electron microscope. E. Ruska (* 1906,
Nobel prize 1986) is one name among others to be cited here. The new
instrument was soon tested in diatom research in Berlin by Krieger
(see Bethge 1956) and Helmcke and later on in a systematic proce-
dure by Ursula Geissler, Gerloff and G 61 z. Electron microscopic

studies in another field of cytology also have the rank of pioneer work.
During his time in Berlin with continuation in Marburg and Hamburg H.
Drawert (1910-1976) laid the foundation for further ultrastructural
investigations with bluegreen algae and desmids in cooperation with
Ingeborg Metzner-Kuster and Marianne Mix (see Mix 1978,
Schnepf 1978).
Anothertradition is less known in spite of its importance. There has

been for long time intimate cooperation between nature conservation
and phycology, and just Berlin was one of the centres of this productive
work. Based on school biology and amateur research a first attempt
to procure a scientific basis for nature conservation was institutional-
ized in the national capital (Staatliche Stelle fiir Naturdenkmalpflege
in Preussen, State Authority for Natural Monument Conservation in
Prussia, founded in 1906). Its first head was H. Conwentz (1855-
1922) succeeded by W. Schoenichen (1876-1956). The latter has
done much for practical and theoretical popularization of the ideas of
nature conservation and landscape management. He also was very active
in didactics of school biology and wrote a lot of important textbooks in
all these fields. Influenced by F. Junge, E. Haeckel, O Zacharias und R.
France he was one of the adherents of the "BUdungswert der Klein-
welt" (title of one of France's popular papers in which he pleaded for
wide popularization of microscopy for amateurs and school teachers).
Schoenichen supported these activities by reediting B. Eyferth's
(1826-1897) well known identification key for microscopic water
organisms "Einfachste Lebensformen des Tier- und Pflanzenreiches"
(first published in 1878, 5th edition in 1925). Many amateurs and
professionals used this book as a first introduction. It's by these and
parallel literary tools that phycology got a broad foundation by ama?
teur specialists.
A further activity of early nature conservation also served to advance
phycology. A generous program of regional studies of nature monu-
ments and prospective sanctuaries resulted in a broadminded pattern of
regional botanical monographs. Many of these included studies on algal
flora and vegetation. Thanks to these activities phycology in Berlin
continued to cover field work also in a time with prevaUing laboratory
research. Phycology in this city has therefore a continuous tradition of
work in nature, and many discussions to reactivate geobotanical re-
Englera 7-1987 47
search on algae and to supplement it with ecological investigations

derive from these activities (sociology of algae, plankton geography
etc, see the papers by H. Bethge, Rosemarie Buchheim-Matz, A.
Donat, K. H 611, W. Krieger and his son H. K r i e g e r, A. v. Lingels-
heim, E. Lindemann, E. Nitardy, W. Panknin, Marlies Schliiter,
Ursula Wed digen and Ursula Geissler, V. Wilkaitis).
Berlin is an inland city. However, next seashore is only 150 km

abroad from Berlin (compare Hamburg's distance from the open sea:
100 km). Nevertheless it appears somewhat farfetched to report on
marine biology in Berlin. Yet, this is only the first impression. Indeed,
important impulses for marine biology came from Berlin. Scientists
from this city initiated research on algae and marine plankton on Helgo-
land long before Britain surrendered the island to the German Empire
(Ehrenberg, N. Pringsheim, J. Miiller, Haeckel). It is not
quite correct in spite of its frequent quotation to say that it was on
Helgoland where Ehrenberg discovered the true nature of marine
phosphorescence. In reality a vast literature had been accumulated
dealing with this enigmatic phenomenon. Ehrenberg studied care-
fully all the presumptions of his time and found the correct explanation
of the matter after personal inspection of North Sea plankton. This was
in August 1835. Later on it became tradition to combine recreation and
scientific work in the course of sommer visits of the island which had
become one of the first seaside resorts already in 1826 while there was
regular steamship service from 1834. These circumstances together with
good railway service between Berlin and Hamburg favoured the installa-
tion of some kind of tradition of biological research on Helgoland by
scientists living in Berlin. N. Pringsheim successfully pleaded for a
botanical curatorship at the Biologische Anstalt Helgoland (founded
in 1892). The first man in this position was P. Kuckuck (1866-1918)
who had been student not only of Reinke in Kiel but also of M a gnu s
in Berlin. Likewise botanical research was done in Naples and Rovigno.
The Zoologische Station Neapel, founded in 1872, was the example
for all the following institutes. Among these, the Zoological Station at
Rovigno/Rovinj was founded by the Aquarium of the Zoological
Garden Berlin. All these activities were supported by the Preussische
Meereskommission (Prussian Commission of Marine Research) which
founded the Institut fiir Meereskunde an der Universitat Kiel. K. M6-
bius (1825-1908) was another scientist to maintain interest in marine
science in Berlin. He came from Hamburg and Kiel. During his profes-
sorship in Kiel he studied the wadden sea and introduced the concept
of "biocoenosis" into science. In Berlin he met several broad minded
and interested collaborators. Zoological studies in Berlin were also
concerned with protozoa and out of this borderland between zoology
and botany many useful suggestions have been forwarded. In zoology,
too, the large work of inventories of the organismic variety was on the
agenda of institutes in Berlin (Dahl, 1856-1929, founded "Tierwelt
Deutschlands und der angrenzenden MeeresteUe . . ." in 1925;Kiiken-
thal, 1861-1922, edited "Handbuch der Zoologie" since 1925). The
schedule of these handbooks included revision of the current systems
and systematic stocktaking. This, in turn, yielded new systems of
protozoa and gave evidence of the algal ancestry so that modernized
phycological systems could be elaborated (by Oltmanns, 1860-1945,
Fritsch 1879-1954, or Pascher, 1881-1945).
Berlin was the seat of the Deutsche Botanische Gesellschaft founded

in Eisenach 1883 with the cooperation of N. Pringsheim. This pro-
fessional society established a Commission fiir die Flora von Deutsch-
land (active for 20 years upt to 1905). Its members collected and an-
alyzed all pertinent literature on all the plant groups including algae.
Different publications took profit of this work. A first tangible result
was the "Kryptogamenflora der Mark Brandenburg" of which only
some volumes could be finished. One such contribution was the part on
algae (1915) by E. Lemmermann from Bremen (1867-1915) who wrote
much more than a simple "Flora" since the volume is the sum total of a
lifelong work with algae. Because of its high standard it has been
proposed to make this volume the later starting point of nomenclature
in Chroococcalean Cyanophyta.
The phycologists who were resident in or near Berlin were special-
ized in several algal groups. Some have a long series of pertinent ex-
perts, for other groups these are lacking. The survey includes diatoms
(Beger, Bethge, Ehrenberg, Geissler, Gerloff, Kolbe, W.
Krieger, Kruger, Miiller, Schluter), fUamentous and unicellular
conjugates (Braun, Gerloff, H. and W. Krieger, Meyen, Mix,
also Donat and Kolkwitz), silicoflagellates (Gemeinhardt, he
also worked on diatoms), dinoflagellates (Lindemann), Volvocales
and related groups (Gerloff, Hartmann and his students, Wollen-
weber, Reichenow), some other fUamentous green algae (Gemein-
Englera 7-1987 49
h a r d t, K n e b e 1, Lucia W i c h m a n n), marine algae (G e r 1 o f f, O. C.

Schmidt) and Charophyta (B r au n, H o 11 z).
Contrary to France or Czechoslovakia in Berlin there hasn't been a
big leader like J. Feldmann or B. Fott to work amidst an impressive
number of students directing their studies. Such type of academic
sovereign may be seen in Max Hartmann, maybe in O. Warburg,
too. But these were only interested in algae to use them as research
subjects. Furthermore, scientific work in Berlin in many cases is charac-
terized by a similar structure as that of its proper industry, by first
class workmanship and team work. Berlin's strong point lies in well
disciplined cooperation. This affects single persons who are willing to
subordinate themselves doing service in long time work or large coop?
eration. This affects the big organizers who luckily chose pertinent
specialists and were good in training skilled workers. The results, viz.
handbooks, Floras and similar surveys do not make sensation. Such
publications belong to scientific every day routine. This does not mean
that they are mass products, not at all. Phycology in Berlin has the
stamp of
many other products: solidly manufactured goods, not
extravagant, but useful.
References
Bethge, H. 1956: Willi Krieger 1886-1954. - Ber. Deutsch. Bot. Ges.

68a: 141-143.
Caspary, R. 1877: Alexander Braun's Leben. - Flora 60: 433-441,
450-457,465-471,497-507,514-519.
Cohn, F. 1895: Nathanael Pringsheim. - Ber. Deutsch. Bot. Ges. 13:
(10) - (33). - Jb. Wiss. Bot. 28: 3-32.
Cohn, P. 1901: Ferdinand Cohn. 2. ed. - Breslau.
Du Bois-Reymond, E. 1888: Adelbert von Chamisso als Naturf orscher.-
Deutsche Rundschau 61: 329-349.
Hanstein, J. 1877: Christian Gottfried Ehrenberg. Ein Tagwerk auf dem
Felde der Naturforschung des neunzehnten Jahrhunderts. ? Bonn.
Hartmann, M. 1956: Gesammelte Vortrage und Aufsatze. I. Allgemeine
Biologie. ? Stuttgart.
Huber-Pestalozzi, G. 1956: Professor Richard Kolkwitz f. ? Schweiz.
Z. Hydrol. 18: 245-253.
Kolkwitz, R. 1935: Pflanzenphysiologie. Versuche und Beobachtungen

an hoheren und niederen Pflanzen einschliefilich Bakteriologie und
Hydrobiologie mit Planktonkunde. 3. ed. - Jena.
Krebs, H. A. 1979: Otto Warburg. - Stuttgart.
Magnus, P. 1895: N. Pringsheim. - Hedwigia 34: 14-21.
Mettenius, C. 1882: Alexander Braun's Leben nach seinem handschrift-
lichen Nachlaft dargestellt. - Berlin.
Mix, Marianne 1978: Die Schriften von Horst Drawert. - Mitt. Inst.
Allg. Bot. Hamburg 16: 15-20.
Miiller, R. H. W. & Zaunick, R. (eds.) 1960: Friedrich Traugott Kutzing
1807-1893. - Leipzig.
Nachtsheim, H. 1963: Max Hartmann f. Leben und Werk (1876-1962).-
Sitzungsber. Ges. Naturf. Freunde Berlin N. F. 3: 14-20.
Pringsheim, E. G. 1954: Algenreinkulturen. - Jena.
Pringsheim, N. 1895: Gesammelte Abhandlungen. Band 1. Befruchtung,
Vermehrung und Systematik. - Jena.
Schmid, G. 1942: Chamisso als Naturf orscher. Eine Bibliographie. -
Leipzig.
Schnepf, E. 1978: In memoriam Horst Drawert. - Mitt. Inst. Allg.
Bot. Hamburg 16: 7-14.

Dr. D. Mollenhauer, Forschungsinstitut Senckenberg, AuBenstelle
Lochmuhle, D-6465 Biebergemund-Bieber.
Englera7-1987 51
Wolfrudolf Lauxi
Contributions to the development of phytomedicine in Berlin
The science of the diseases and damages of plants and their causes,
manifestations, developments as well as the measures to be taken for
the preservation of plants and harvested crops is summarized under the
term phy tomedicine. Phytomedicine stands beside human medicine and
veterinary medicine and today is generally accepted. Textbooks use this
term in their title as well as a German scientific society. It was set up in
Berlin in the twenties of this century by Otto A p p e 1 who talked about
plant medicine. Today phytomedicine includes among others phyto-
pathology, plant protection, entomology as well as mycology and viro-
logy as far as plants are concerned.
Science lives on the exchange of experiences, opinions and results.
To describe the development of a science in a restricted area would
mean that one has to do without the manyfold extrinsic influences and
transient effects which are necessary for science. On the other hand one
would have to exceed the scope of such a deseription right from the
start. Although BerUn as a result of the residence of the Biologische
Reichsanstalt fiir Land- und Forstwirtschaft (Biological Research
Centre for Agriculture and Forestry of the Reich) for many decades
had been the scientific and organizational centre of phytomedicine in
Germany the Umits of such a deseription are quite obvious.
As phytomedicine has developed since the middle of our century to
such an extensive and versatile and also complex field of science a
somewhat comprehensive deseription of its development and men-
tioning of the numerous representatives of the subdivisions of phyto?
medicine would go beyond the scope of this short deseription of the
development of phytomedicine (for further details see Bartels & Koch
\ Thanks are given to Dr. F. Geike for his assistance in translating into
English.
52 Laux: Phytomedicine in Berlin
1980, Braun 1952,1965, Mayer 1959,Ueberlein 1984, Chronik ... 1973,

50 Jahre... 1949).
Here can only be made the attempt to give a short description of the
institutions at which phytomedicine was or is done in Berlin and to
describe the scope where scientists of the different disciplines did their
job in research and teaching and organization during more than hun-
dred years. I furthermore want to describe the evolution of phyto-
medical research in this city and beyond it and I want to name some of
the leading scientists and their work that brought fundamental know?
ledge to phytomedicine. Interested readers who would like to know more
about the development of phytomedicine are referred to the very
numerous literature and to a very popular presentation of this subjectby
E. Schmidt (1948). They also can look up the original papers of the
scientists not cited in detail here but easy to find out.
Early developments
The early development in the field of phytomedicine had started

in the 18th century with efforts to classify the different symptoms of
plant diseases. In the 19th century it developed to the phytopathology
which was the precursor and part of modern phytomedicine. Only in
the second half of the 19th century possibilities for the control of plant
diseases and pests became of greater interest.
In 1727 J. L. Frisch, member of the Prussian Academy of Science
and head of the secondary school Zum Grauen Kloster, in Berlin in
"Miscellanea Berolinensia" published a paper on "Damages of crops
caused by cutworm and on the biology of Agrotis larvae". He and also
J. Gleditsch watched the occurrence of the migratory locusts and of
their parasites which he believed to be the cause of the voraciousness of
the locusts (Kolbe 1985).
In 1752 Friedrich der Grosse issued an edict concerning the
control of locusts (Facsimile in Kolbe et al. 1983) and in 1833 P.
B o u c h 6 published a paper on the "Biology of harmful and beneficial
insects in the garden and the most effective methods to control the
former".
In 1841 in Berlin a book entitled ''Pflanzen-Pathologie. Lehre von
dem kranken Leben und Bilden der Pflanzen" (Plant pathology ? The
Englera7-1987 53
science of the diseased life and forming of plants) was pubUshed from
the estate of the 1840 deceased Doctor of Philosophy, Medicine and
Surgery and Professor at the Friedrich WUhelm University of Berlin F.
J. F. Meyen. He presented a large number of symptoms especiaUy
such caused by microorganisms in order to form a system of "patho-
logy, nosology and therapy of plants". Under the heading "external
diseases", however, also damages of plants caused by insects and mam-
mals have been described. As measures for plant protection smoke of
tobacco and dispensation of sulphur besides a modification of the
culture method are discussed.
In the middle of the nineteenth century the heavy spread of blight
of potatoes (Phytophthora infestans) caused intense worries in Europe
and in Germany, too. So the docent H. Schacht (since 1860 pro?
fessor in Bonn) in Berlin was entrusted by the Prussian Royal Ministry
of Agricultural Affairs with the clarification of this nearly unexplored
disease. The results of his journeys and experiments among which he
had done unsuccessful trials on infections with spores of Phythoph-
thora infestans were summarized in 1856 in a detailed "Account to the
Royal Landes-Oeconomie-CoUegium about the potato-plant and its
diseases."
Only in the second half of the 19th century there was an institu?
tional, organizational and also scientific fusion of this special field of
plant science. EspeciaUy we owe J. Kiihn who in 1858 pubUshed thebook
"Die Krankheiten der Kulturgewachse, ihre Ursachen und ihre Ver-
hutung" (The diseases of crops ? their causes and control) that phy-
tomedical problems have been introduced in science after he received a
caU to the professorship newly estabUshed in 1862 in Halle.
From HaUe, the early centre of phytomedical research and science,
many connections lead to BerUn. M. Hollrung, the head of the ex-
perimental Station for Plant Protection of the Chamber of Agriculture
of the Province Saxony published his independent report on events
and achievements in the field of plant protection, the "Jahresbericht
uber die Neuerungen und Leistungen auf dem Gebiete des Pflanzen-
schutzes" since 1899 in BerUn and thus created a bibUographic re-
porting journal which was the precursor of the "BibUography of Plant
Protection".
Agricultural College
In 1881 the Agricultural College was founded in Berlin. Professor

of plant physiology became Albert Bernhard Frank who already
earlier was engaged in phytomedical problems in Leipzig and in 1880
had published the textbook "Diseases of plants". In Berlin he was
engaged in these problems so greatly that the institute in 1884 was
changed to Institute of Plant Physiology and Plant Protection thus
introducing phytomedical research and teaching in Berlin. Exemplary
remained his point of view that a control of plant diseases and pests
is only possible when their nature and cause is understood. By this he
postulated the inseparable connection between basic research in phyto?
medicine and practical plant protection. Besides various research
Frank was greatly involved in practical and organizational plant
protection. For all those who did practical plant protection in 1897 he
published his book "Kampfbuch gegen die Schadlinge unserer Feld-
friichte" (Fighting pests of our field crops) (see Kriiger 1901).
In the area of practical plant protection there were numerous develop-
ments. For instance the special committee for plant protection of the
German Agricultural Society (DLG) was founded in 1890 after an
attempt to install a central office for the observation and control of
insects and fungi damaging field crops had failed.
Biologische Abteilung des Kaiserlichen Gesundheitsamtes (Biological

Division of the Imperial Health Office)
One of the initiators for the foundation of the Biological Division of

the Imperial Health Office was the farmer and member of the German
Diet Dr. Dr. hc. Albert Schultz-Lupitz who on his farm Lupitz in
the Altmark had developed exemplary methods for the cultivation of
light soils and who was one of the founders of the German Agricultural
Society. On March 26th 1897 he brought forward the motion in the
198th plenary session of the German Diet in Berlin that the Diet
might pass the resolution to ask the confederated governments to set up
an agricultural technical Imperial Institute for Bacteriology and Phyto-
pathology and to provide the required money in the budget of 1898/99
(Circular No 726 of March 24th 1897). By this motion it was directly
-'
>v .,???%
Albert Bernhard Frank

1839-1900
caUed on the government to get things done. After consultations be?

tween the Imperial Health Office and the German Agricultural Society
the Biological Division for Agriculture and Forestry at the Imperial
Health Office was founded on January 28th 1898 and started to work
in autumn. A. B. Frank as Privy Senior Executive Officer in 1899
was appointed as head of the department and at the same time his Insti?
tute of Plant Physiology and Plant Protection became part of this
newly formed department. Thus studies concerning plant physiology
and plant protection became concentrated at Berlin; the Agricultural
College, however, lost the professorship of Plant Protection which was
transferred to the head of the Institute for Fermentation Industry.
By this transfer of the professorship the phytomedical research
at the Agricultural College came to an end for a long time. At the
vegetabile branch of the Agricultural Museum, however, O. Weh-
sarg was concerned with the biology of weeds and later (1954) wrote
a widely read textbook about weeds. At the Royal Zoological Museum
H.-J. Kolbe was curator. In his book "Pests and beneficial animals
of the garden" pubUshed in 1901 he explained the significance of
predatory and parasitic insects and even the mode of action of fungi
of the genus Entomophthora which are pathogenic to insects and their
potential employment for the biological control of insect pests.
At the Experimental Station of Grain Processing in Berlin E. Ber?
liner in 1911 from the Mediterranean flour moth isolated Bacillus
thuringiensis which is pathogenic to insects and suggested the employ?
ment of this bacterium for the biological control of pests in grain silos.
Bacillus thuringiensis, however, became very important only after it
had been possible to produce it industrially (in 1960) and to employ
it on a larger scale.
Phytomedical teaching was now done by members of the staff
of the Biological Division and its following institutions. They worked
as associates, associated professors, or lecturers at the Agricultural
College.
The newly founded Biological Division of the Imperial Health
Office grew fast. Farmland for experiments has been provided in the
area of the Royal Domain in Dahlem and the first laboratory has been
built. Already in May 1899 central tasks were assigned to the depart?
ment and the appropriate authorities of the individual states of the
German Reich were instructed to make reportes to the department
on all matters concerning plant protection.
Englera 7 - 1987 57
After the death of Frank in 1900 baron Karl von Tubeuf became
head of the department for only a few months before he was appointed
to a chair in Munich. He was succeeded in 1902 by Rudolf A d e r h o 1 d
who in 1905 became the First Director of the now independent Kaiser-
liche Biologische Anstalt fiir Land- und Forstwirtschaft (Imperial
Biological Research Centre for Agriculture and Forestry).
On account of an advisory board formed in 1900 the leading per-
sonalities of phytophathology and plant protection of this time, among
them Hiltner (Berlin, Munich), Hollrung (Halle), Kirchner (Hohen-
heim), Kiihn (Halle), and S o r a u e r (Berlin) came into contact with the
Division resp. Research Centre.
The Biologische Reichsanstalt fiir Land- und Forstwirtschaft (Biological

Research Centre for Agriculture and Forestry of the Reich)
The vital stage of development of the Research Centre which was

changed in 1919 into Biological Centre for Agriculture and Forestry
started in 1920 when Otto Appel who joined the Biological Division
of the Imperial Health Office in 1899 was appointed to the director
of the Research Centre. The Research Centre which had been put under
the Ministry of Food and Agriculture of the Reich was reorganized
according to the plans of A p p e 1. It founded a couple of branch offices
and extra laboratories. At the 28th special exhibition of the German
Agricultural Society in 1921 in Leipzig it came forward with the
organization of the Plant Protection Service at a total. Furthermore,
the Research Centre founded new journals like the "Nachrichtenblatt
fiir den deutschen Pflanzenschutzdienst" and the "Bibliography of
Plant Protection". In 1922 the "Ahrenschlange" with the capitals
"BRA" as a characteristic for the Biologische Reichsanstalt was taken
as a symbol for the official plant protection and was protected by
registration as a trademark in the trademarks register of the patent
office of the Reich (see Appel 1919, 1921).
In 1913 Appel had been charged to formulate a memoir on the
requirements of an act on plant protection of the Reich and in 1920
he presented his draft of an act concerning plant protection in the
German Reich to the Ministry of Economics. This draft, however, was
not realized by the Government. Only in 1937, a long time after
124 Schnarrenberger: Botany at KWI
but nevertheless he had excellent physical and chemical equipment. His

coworkers in the early years were almost exclusively technical assistants
(E. Negelein, F. Kubowitz, E. Haas, W. Christian, W. Liitt-
gens), who became world-famous through their publications. Because
of his scientific achievements and on the occasion of a lecture in
Baltimore in 1929, the Rockefeller Foundation decided in 1930 to
finance the building of the later-known Kaiser Wilhelm Institute for
Cell Physiology (and beside it, the Kaiser Wilhelm Institute for Physics)
at Garystrasse 32/Boltzmannstrasse 12/14. Warb urg's Institute was
already completed in 1931. It is built in a splendid country style and is
used today by the Max Planck Society as an Archive and a Guest
House. The old library is maintained asinWarburg's days, and can be
viewed by visitors. In front of the building, a statue of Emil Fischer
was erected in 1952 on the centenary of his birthday. In 1943, because
of the ever-increasing aerial bombardment of the city, Warburg had
to move his laboratory to Liebenberg, about 50 kilometers north of
Berlin. In 1945, after the Russian army marched in, Warb urg's entire
laboratory equipment was transported away, through an error, and was
never found again. For four years after the end of the war, the Dahlem
Institute was used as the headquarters of the American Armed Forces.
From May 1950, it was again used as a Kaiser Wilhelm Institute, and
from 1953 it became known as the Max Planck Institute for Cell
Physiology Warburg continued active scientific work almost until
the day of his death at 87 years. He died on the first of August in 1970.
Otto Warburg ranks as one of the most important German bio-
chemists. His main achievements lay in the analysis of the energy meta-
bolism of biological cells. The most important, classical work of War?
burg included a description of the enormous increase in the respi-
ration of sea-urchin eggs after fertilization (Warburg 1908), the great
difference in the respiration and fermentation of healthy and cancer
tissues/cells (Warburg 1926), the identification of the iron prophyrins
as components of the respiratory chain (Warburg's respiratory enzyme)
(Warburg 1946), the identification of the pyridine nucleotides and
flavins as hydrogen carriers in biological oxidations, the purification
and crystallization of most of the glycolytic enzymes, and the discov?
ery of glucose-6-phosphate dehydrogenase, the first enzyme of the
oxidative pentose phosphate cycle (Warburg 1948). The development
of techniques such as manometry for the measurement of the gas
Otto Appel
1867-1952
Appel had retired, the Act concerning the Protection of Agricultural

Crops was enacted exactly in accordance with A p p el's ideas.
During the buildup and development of the Research Centre the
number of scientists increased from 14 in 1907 to 42 in 1923 and 64
in 1933 when Appel retired. While being director Appel persistently
supported the German Plant Protection Service while he was in charge
of practical plant protection. Already in June 1919 during an assembly
of the representatives of the headquarters of plant protection this
institution had accepted the principles set up by A p p e 1 among which
was the introduction of the examination of plant protection products
of Germany.
It is owed to Otto Appel that the organization of plant protection
in the German Reich on which he had a substantial effect became a
very much admired example often successfully copied by other states.
The central importance of the Research Centre was not only reflected
by its publication of the "Nachrichtenblatt fiir den Deutschen Pflanzen-
schutzdienst" with the supplement "Amtliche Pflanzenschutzbestim-
mungen" but also by the fact that there was nearly no scientist in phyto?
medical research or in practice in those days who had not been for some
time at an institute or laboratory of the Biological Research Centre.
Central duties of the Biologische Reichsanstalt
The central duties of the Research Centre for instance arose when in
1917 in single countries of the Reich public measures were provided to
control the steadily spreading muskrat. The basis for these measures
was a tour of G. R 6 r i g, a scientist of the Research Centre, through
the infested areas. Until 1935 the muskrat control service was subordi-
nated to the Biological Research Centre which at least resulted in a de-
crease of the speed of spreading of this pest.
Similarily the Research Centre was involved in the control of the
Colorado beetle, Leptinotarsa decemlineata which appeared in Europe
in 1876 and was first seen in Germany in 1914 near Stade. M. Schwartz
of the Biological Research Centre was member of a committee of ex-
perts. In the following years he dealt very intensively with the biology
of this pest (Schwartz 1925) and in 1934 was appointed to the head
of the control measures by the Ministry of Food and Agriculture of
Englera7 - 1987 61
the Reich after the Colorado beetle had reappeared in Germany and
had been successfully controlled according to the "Instructions for the
control of the Colorado beetle" developed by Schwartz.
Schwartz also was appointed by the Ministry of Food and Agri-
culture of the Reich to the Authorized Representative of the Reich for
the control of the San Jose* scale (Quadraspidiotus perniciosus) after
the German Reich had become responsible for this pest after the union
with Austria. In the beginning this pest was only found in Austria.
In German it was found only after the Second World War.
In the field of entomology in 1934 a working group was founded
by the Biologische Reichsanstalt and the Deutsches Entomologisches
Institut (German Institute of Entomology) which already in 1924 had
been aimed in vain by the initiator and head Walther H o r n. This
cooperation greatly enriched the scientific work of the Research Centre.
At the German Institute of Entomology, however, there was a change
in the activity towards applied entomology and bibUographic studies.
After the Second World War the German Institute of Entomology has
been moved to Eberswalde where today it is an institution of the
Academy of Agricultural Sciences of the GDR.
The phenological service estabUshed by E. Werth and the working
in the field of quarantine were central duties, too. The working in the
area of quarantine was continued after the Second World War by K.
Ludewig, A. Harle and L. Quantz.
Although the German Plant Protection Service by an act of 1937
had been put under the Reichsnahrstand (Ministry of Agriculture
of the Reich) there remained close relations to the Biologische Reichs?
anstalt.
During the Second World War apart from some difficulties in the
number of members, by the loss of connections to international author?
ities and scientific Uterature and the destruction of some branch offices
the work in the Biological Research Centre which in 1938 had 90
scientists (Riehm 1938), continued comparatively smooth. Even
in the issue of the "Reichs-Pflanzenschutzblatt" of January 1945
there were pubUcations concerning the correlations between structure
and effectiveness of insecticides and mechanical traps for sparrows,
and then the president of the Research Centre, E. Riehm, commented
on the sparing application of pesticides owing to war and on the ration-
ing and distribution about which they had talked during a conference
of the German Plant Protection Service in December 1944.
Postwar developments
At the end of the Second World War the Research Centre in Berlin
was only slightly damaged; it was, however, separated from its branch
offices scattered all over the former territory of the German Reich and
from its departments like virology and bacteriology which had been
moved from Berlin during the war.
As Biologische Zentralanstalt fiir Land- und Forstwirtschaft (Central
Biological Research Centre for Agriculture and Forestry) it was at
first subordinated to the municipal authorities of Berlin; later it was
subordinated to the German Economic Commission that was also re-
sponsible for food and agriculture in Berlin and the then Soviet-occupied
zone. When in 1949 the political circumstances led to the division of
Berlin the remaining part of the Research Centre was subordinated to
the senate of West-Berlin while in Kleinmachnow in the then Soviet -
occupied zone in the course of the foundation of the German Academy
of Agricultural Sciences in 1951 it was started to build up the present
Institute of Plant Protection Research of the Academy of Agricultural
Sciences of the GDR (Bochow & Schumann 1985).
Simultaneously lectures on phytopathology and plant protection
started at the faculty of agricultural and horticultural science of the
Humboldt University of Berlin, and in 1961 a professorship of phyto?
pathology and plant protection was established at the Humboldt Uni?
versity. At the Technical University of West-Berlin in 1951 the Faculty
of Agriculture was founded in Berlin-Dahlem. Later it became the
Fachbereich Internationale Agrarentwicklung (Faculty of International
Agricultural Development) with which the proved cooperation with the
Biological Research Centre in Berlin -Dahlem could be continued.
Those parts of the former Biological Research Centre that had been
situated in the zone occupied by the Western Allies formed a new
Central Office in Brunswick. The now called Biologische Bundesanstalt
fiir Land- und Forstwirtschaft Braunschweig (Federal Biological Re?
search Centre for Agriculture and Forestry Brunswick) in 1950 was
subordinated to the administration of the Federal Government and in
1951 Harald Richter became President in Brunswick and simul?
taneously was Director of the Biological Research Centre in Berlin-
Dahlem. Thus the integration of the two Centres to the Federal Biolo?
gical Research Centre for Agriculture and Forestry, Berlin and Bruns?
wick (BBA) in 1954 was the only suitable consequence.
Englera7-1987 63
Of the total field of duties the Biological Research Centre of the

Reich in Berlin once had only the Mycology, Bacteriology, Zoology
(until 1976), Horticultural Virology (until 1968), Protection of Stored
Products, Pesticide Research, Diseases of Ornamental Plants (until
1985), Non-parasitic Plant Diseases, Office for Reports and Warning
(until 1970), and Economic and Legal Affaires (until 1981) after the
war remained in BerUn or came back. Added to these fields were the
Documentation Centre
for Phytomedicine being associated with the
Library and the Information Centre for Tropical Plant Protection
(1965) as well as the Division for the Examination of Chemicals (1982).
A central duty for the total area of phytomedicine in BerUn was and
stiU is the "BibUography of Plant Protection" (BibUographie der Pflan-
zenschutzliteratur) which was founded in 1921 by M. Morstatt and
was continued by J. Barner. Since 1965 it has been changed to data
processing by W. Laux and has been supplemented by new services
like the "Database PHYTOMED" and information for phytomedical
research and practical knowledge.
At the Faculty of Agriculture of the Technical University of BerUn
that later has been changed to Faculty of International Agricultural
Development the special field of phytomedicine as usual was served
by staff members of the Federal Biological Research Centre who had
a lecturership or additional professorship (W. Gerlach, K. Heinze,
A. Kloke, W. Laux, H. Muller, H. Richter, W. Sauthoff, G.
Schuhmann). It is supposed, however, that at the end of the 80ies
a full professorship wUl be established (Laux, in press).
Plant Protection Office of BerUn
Until 1919 the duties of the official plant protection service had
been assigned to the Imperial Biological Research Centre. In 1920,
however, an office for plant protection was installed in the buildings
of the Biological Research Centre which was responsible for the Pro-
vince Brandenburg and Berlin. The residence within the Research Centre
supported the close contact between the office and the Research
Centre. These relationships remained when the office because of lack
of rooms in BerUn-Dahlem had to move to the building of the Chamber
of Agriculture of the Province Brandenburg and later on was trans-
ferred to Potsdam-Luisenhof. Apart from this move of the office a

couple of duties concerning Gross-Berlin, for instance the monitoring
of plants, remained at the Research Centre. In 1946 the Institute
for Biological Investigations and Plant Protection was formed to do
the tasks of the Plant Protection Service in Berlin. After several changes
of the name of the office it became the Pflanzenschutzamt Berlin
(Plant Protection Office Berlin) in 1953. Under its directors F.
Zacher, G. Fichtner and H.-P. Plate there has been also a good
collaboration with the Biological Research Centre and the Federal
Biological Research Centre, respectively.
The Otto Appel Commemorative Medal
One of the most important tasks of the president of the Federal

Biological Research Centre at that time is the patronage of the Otto
Appel Commemorative Medal. This award for valuable scientific or
organizational work concerning plant protection was offered on the
occasion of the 85th birthday of the Privy Councillor Prof. Dr. Dr. h.c.
Otto Appel and he was the first to receive this award. Since that time
this medal has been awarded to 21 outstanding representatives of
phytomedicine and plant protection at home and abroad. With the
Otto Appel Commemorative Medal German Phytomedicine does not
only honour one of its most outstanding figures but after Otto
A p p e Ps death also puts honourable works and persons in the field
of phytomedicine in relation to him.
Otto Appel
Otto Appel who was the head of the Biologische Reichsanstalt

from 1920 until 1933 has given decisive impulses to the Research
Centre, to the German Plant Protection Service, and to phytomedical
research and teaching (Schlumberger 1956). In research he has not
only suggested but also has done research himself. The focus of his
interest was the potato. The detection of the pathogens of black leg of
potatoes that after a preliminary note was described by him in detail
in 1902 was not only a very important finding for potato growing in
Otto Appel Medal
Germany but simultaneously was a basic contribution to the problem

whether bacteria could be pathogens of many plant diseases at all
which at that time was very controversial. In 1903 Appel preliminarily
named this pathogen Bacillus phytophthorus. By this finding Appel
introduced bacteriological research in phytomedicine.
Already in 1902 Appel proved his relation to practical work when
he presented a detailed paper "About studies on pitting of potatoes"
before he laid the foundations of his scientific reputation by his fun-
damental work on black leg in 1903.
The scope of scientific tasks of a scientist in those days can be seen
from a view into the journal "Arbeiten aus der Biologischen Reichs?
anstalt fiir Land- und Forstwirtschaft" of 1902 where Appel besides
the paper on the pitting of potatoes, already mentioned above, pub?
lished a paper "On the significance of the Friihlings-Kreuzkraut Senecio
vernalis as a weed" and together with A. J a cob i a paper "On obser?
vations and experiences on the plague from rabbits and its control".
A further vital branch of research was estabUshed by Appel in
1907 with the edition of the leaflet "Potato leaf roll". Basing on the
experiences made in Denmark during a serious occurrence of this dis-
ease he differentiated in this leaflet between potato leaf roll and the
already known crinkle of the potato. At that time, however, he did not
recognize that the potato leaf roll was caused by a virus.
The dispute on the question whether potato leaf roll and other
diseases had to be attributed to an ecological weakening or an in-
fectious disease caused by virus occupied the Biological Research
Centre and many scientists all over the world for many decades.
Friedrich Merkenschlager and the theory of ecological weakening
In Berlin these studies were especially associated with Friedrich

Merkenschlager who came to Berlin in 1925. In the experimental
field of the Research Centre with Dahlem sandy soil and "Dahlemer
Abbauboden" (soil exhaustion), respectively, known from the litera-
ture and other experimental stations Merkenschlager and his
coUeagues were engaged to solve the problems of potato diseases.
According to the theory of weakening the value of the seed potatoes
depended on the environmental conditions which under European
Englera 7 - 1987 67
culture conditions were not adequate. A continuous cultivation of

potatoes, therefore, inevitably led to a decrease in yield and to symp-
toms of diseases. Extensive studies on the metabolism of proteins and
carbohydrates in potatoes and on enzymes which were very difficult
in those days did not bring a satisfactory solution of the problem.
Although not refusing a certain ecological factor at the occurrence
of the disease the supporters of the theory that viruses caused these
symptoms of disease, on the other hand, were not able to substan-
tiate their theory by experiments sufficiently. Only in 1935 Stanley
was able to establish the proteinaceous character of the tobacco mosaic
virus.
The field ofMerkenschlager's scientific interests was far
beyond these problems just mentioned
as can be seen from his "Tafeln
zur vergleichenden Physiologie und Pathologie der Kulturpf lanzen"
published in 1927 and from his contribution on "Plant nutrition and
plant diseases" in the 6th edition of Sorauer's "Handbuch der
Pflanzenkrankheiten" published in 1933 (see also Merkenschlager &
Klinkowski 1933). Extensive studies in the field of plant nutrition
have been done by Merkenschlager as well as papers on anthro-
pology (Ulsamer 1970).
His remarkable career at the Biologische Reichsanstalt at Berlin
only lasted eight years. On the basis of his papers concerning anthro-
pological problems he was dismissed in 1933 and even was imprisoned
in 1937. After the war Merkenschlager did not return to Berlin
(Ullrich 1970).
Erich Kohler and plant virology
The studies on potato diseases which had been attributed to ecolo?

gical weakening by Merkenschlager were continued by Erich
Kohler who joined the Biological Research Centre in 1921 and
first was engaged in the problems of wart diseases of potatoes. In 1932
he turned to plant virology and was able to disprove the theory of eco?
logical weakening in potato culture by his accurate and versatile studies.
In a series of papers on the virus disease of potatoes Kohler ini-
tiated plant virology at the Biological Research Centre. In the follo?
wing years and decades plant virology spread from the potato to
numerous plants. K 6hier

cultivated especially devoted himself to the
transmission of viruses to the spreading and systematics of viruses
of the Solanaceae. He also devoted himself to research on problems
of resistance of the plant and the methods of the detection of viruses
with test plants.
From plant virology the favourable situation of Berlin for scientific
research is obvious: By the cooperation of G. A. Kausche and E.
Pfankuch of the Biological Research Centre and H. Ruska (the
brother of the winner of the Nobel prize in 1986) of the Laboratory
for Electron Microscopy of the Siemens & Halske Corporation it
was possible for the first time to image geometrically exact macromole-
cules of the tobacco mosaic virus and of the X-virus of the potato
in the Biological Research Centre by using the newly developed Siemens
electron microscope (Klinkowski 1960). Calculations of the size of the
macromolecules using other methods could be confirmed by appli?
cation of electron microscopy. By these studies electron microscopy
has been introduced in plant virology where it has become an important
tool in research in the following years and decades. These new findings
initiated a biochemical approach to virus diseases and to the problem
of inactivating viruses which was discussed by Pfankuch (1943),
while Kausche took them as a basis for his studies on transmissi-
bility of viruses by seeds.
After the Second World War K 6hier worked in Brunswick where
he was head of the Instituteof Virology of the Federal Biological
Research Centre untU 1955. In 1966 he was awarded by the Otto
Appel Commemorative Medal. Even at an advanced age of 80 years he
surprised colleagues with remarkable papers.
In Berlin the tradition of virology was continued by H. A. Usch-
draweit in the field of horticultural virology until 1968.
Mycological research
While studying virus diseases of plants was a relatively new branch

of research investigations on fungi as pathogens of plant diseases dated
back to the nineteenth century.
Already F r ank was engaged in research of fungus diseases when he
studied cherry leaf scorch and identified Gnomonia erythrostoma as
pathogen of this disease.
Englera 7 - 1987 69
As he was able to gain deepened knowledge of the biology of the

fungus he was able to suggest measures for its control that were prompt-
ly adopted in practice. A further discovery with far-reaching conse?
quences was the description of the fungus Phoma betae that at that
time was believed to be the pathogen of the heart and dry rot of sugar -
beet. It is notable, however that Frank in this connection made the
reservation of a certain disposition of the plants for an infection which
was confirmed forty years later when it was shown that a deficiency
of boton was the primary cause of heart and dry rot and that this on
the other hand was an important factor for an infection with Phoma.
Frank was also very active in the field of entomology. When
Germany was faced with the threat of an introduction of the San
Jos6 scale he was intensively concerned with scales.
The mycological problems that were taken up by Frank at the
turn of the century for the moment were continued by R. Ader-
hold who came from the Pomological Institute of Proskau to the
Biological Research Centre and devoted his special interest to the
diseases of fruit. He very thoroughly studied the biology of the pa?
thogen of the scab and the until then controversial questions con?
cerning the dependance of the Fusicladium species on a host. Together
with W. Ruhland he studied the brown rot of apples and they were
able to diagnose the individual Sclerotinia species and to define them.
The beet was a very important matter of investigation, too, at the
Research Centre. From 1906 to 1911 with it Phytium, Aphanomyces,
and Phoma were identified as pathogens of the black leg and its me-
chanism of transmission was partly elucidated.
Hans Wilhelm Wollenweber
In 1920 a basic development started with the newly established myco?

logical laboratory under Hans Wilhelm Wollenweber. Wollen?
weber in 1908. came to the Research Centre for the first time. After
studies in the U.S.A. and other countries he became a member of the
Research Centre in 1918. Here Wollenweber determined the
development of mycological research beyond the Research Centre for
decades. He introduced pioneering methods into mycology like the
pure culture of fungi without host plants. Besides the tasks related to
Hans Wilhelm WoUenweber
1879-1949
Englera7-1987 71
practice special attention was paid to the morphology and systematics

of fungi pathogenic to plants. Especially for WoUenweber and his
successors, H. Richter who was honoured in 1977 by the award of
the Otto Appel Commemorative Medal and W. Gerlach, Fusarium
was the centre of interest during their work. These studies on Fusarium
started in 1908 with research of A p p e 1 and WoUenweber on the
Fusarium of potatoes and led to the first monograph on the genus
Fusarium in 1913. Soon the studies on the disease caused by Fusarium
were extended by WoUenweber and his collaborators to other
cultivated plants Uke Gramineae, Cyclamen, Cajanus indicum, Lupi-
nus, and other plants. During these studies the economic significance
and possibilities of the control of the disease caused by Fusarium
became more and more important. Therefore, WoUenweber and
Reinking reviewed these problems in 1935. From 1916 to 1935
the comprehensive treaties "Fusaria autographice delineata" of
WoUenweber containing 1200 plates had been pubUshed and in 1931
and 1943 a monograph on Fusarium consisting of two parts in which
67 species, 58 varieties, and 24 formae of the genus Fusarium have
been described was published. These studies were carried on with great
intensiveness by Richter and finally after the Second World War
by Gerlach. They culminated in the monograph on Fusarium by
Gerlach and Nirenberg (1982).
The methodical and biological findings obtained during these studies
on Fusarium have been reflected in the research on many other fungal
diseases of cultivated plants. The most important aim was to find
relations between the teleomorphs and anamorphs of Ascomycetes.
For these experiments on more than 50 species were carried out in?
cluding studies to find out the influence of nutrition, light, and tem-
perature on the formation of teleomorphs. WoUenweber and
Richter not only were engaged in the then unsolved problem of
Dutch elm diseases
the pathogen of which, Graphium ulmi (Ophio-
stoma ulmi), already in 1927 had been isolated by WoUenweber
but were also engaged in the susceptibiUty of the different species
and varieties of elms to fungal infection and in the pathogen Rhab-
docline pseudotsugae of the needle east of Pseudotsuga.
The basis for the control of fungal diseases always was the eluci-
dation of the biology of the pathogen but it also was necessary to do
morphologie-systematic studies for its unequivocal identification and
definition. In this connection the studies of Richter on Fusarium

diseases and rust of fodder legumes in the thirties should be mentioned
(Richter 1938). The studies on Rhizoctonia solani of K. O. Miiller
in the preceeding decennium finally led to the development of a control
of Rhizoctonia by a treatment of the seed potatoes.
In 1985 the monograph on Phytophthora and Pythium by H.
Krober as a result of decades of research was put besides the mono?
graph of Gerlach and Nirenberg on Fusarium. Already in the
early sixties Krober by his studies on Peronospora tabacina had
decisively contributed to the control of the epidemic blue mould
disease of tobacco in the Federal Republic of Germany (6 Jahre Blau-
schimmelkrankheit . . . 1966). G. Schuhmann (1966) studied the
physiological specialization of Tilletia spp. and the resistance of wheat
varieties against this pathogen and in the late seventies Roswitha
Schneider published a monograph on the genus Pyrenochaeta.
Extensive mycological research was also done by Walter Saut-
h o f f in the field of ornamental plant culturing, the economic signi?
ficance of which enormously increased after the Second World War.
Of the numerous studies that led to an identification of many diseases
of ornamental plants and recommendations to horticultural practice
we only want to stress the elucidation of the Erica wilt. This disease
which for a long time was regarded as a consequence of mistakes
during culture or as an effect of different fungi could be definitively
attributed to Phytophthora cinamoni and simultaneously methods
could be elaborated to avoid this disease. In the same way H. G. Pag
was able to identify Phytophthora cryptogea as pathogen of the foot
rot of Gerbera. In this department particular attention was also paid
do diseases of orchids.
Karl Otto Miiller and the phytoalexins
With Karl Otto M ii 11 e r experts in phytomedicine associate phyto?

alexins. This complex of problems inevitably was a result of his exten?
sive mycological studies. While working on his 1921 thesis he had
already studied developmental physiology of the mycelium, and some
time later he became research assistant at the Biologische Reichsan?
stalt. At the suggestion of Appel to whom he was united in friend-
Englera 7-1987 73
ship for many years he maintained good relations to the Agricultural

CoUege. In 1924 he qualified himself as a lecturer at the CoUege and
in 1925 got a lectureship in phytopathology and applied mycology.
In 1928 Muller was appointed associate professor at the Agricul?
tural College (Linskens 1978).
At the Biological Research Centre he at first worked on Phytoph-
thora and studied the appearance of different strains and types being
morphologicaUy and physiologically distinguishable. During these
studies he came across a phenomenon which determined aU his further
research. He describes this phenomenon in the following way: "Al?
though the pathogen invaded the host plant it died after a short time
without being able to fructify."
In the course of his further research he approached the problem of
degeneration disease of potatoes from the point of view of breeding
for resistance. Many potato hybrids which are resistant or tolerant
towards virus diseases have been due to his breeding efforts. His in-
creasing activities in plant breeding, however, which also included
pine and cereals were considerably restricted after 1933. These restric-
tions made him concerning himself with the originaUy theoretical
question: "What does prevent the pathogen from growing further when
it has invaded the resistant host plant?" The solution of this question,
however, could only be tackled after the Second World War.
Although M ii 11 e r did not return to the Federal Biological Research
Centre he was closely attached to the German phytomedical research
even during his numerous researches abroad. During his studies in
Cambridge and AustraUa he was able to prove the antitoxins in plants
which were formed after the pathogen had invaded the plants and
which he had postulated. He called these antitoxins phytoalexins.
The term "phytoalexin" for the first time is mentioned in a paper
pubUshed in 1958 in the "AustraUan Journal of Biological Science"
("Studies on phytoalexins I") (Muller 1958). In the sixties coworkers
of Muller were able to elucidate the structure of two phytoalexins
and in 1964 it was possible to show by tracer technique that the
formation of phytoalexins is induced by metaboUtes of the pathogen
which are released into the host cell.
Bacteriological research
After Appel by his discovery .of the pathogen of the black leg of
potatoes in 1903 had introduced bacteriological research into the
Biological Research Centre bacteriological studies were continued for
many years by Carl Stapp. Besides many other studies including
mycological research Stapp in 1939 combined different bacterial
strains pathogenic to potatoes to the Bacillus phytophthorus group.
This group, now known as Erwinia carotovora, also today is of utmost
importance in agriculture. During his studies he applied obviously for
the first time in phytomedicine serological methods which later on
especially in virology became very important.
In the field of bacteriological research not only damaging bacteria
have been studied. Stapp for instance has made valuable contri-
butions to the biology of nodule bacteria. He and his collaborators
have also carried out extensive research on the metabolism of bacteria
in soil. In 1958 his services to phytomedicine were honoured by the
award of the Otto Appel Commemorative Medal.
His successor H. Bortels carried on this research. He was able to
show that molybdenum and vanadium were specific cofactors in nitro-
gen fixation. Until the fifties Bortels carried on bacteriological
research. He especially tried to find potential influences of the weather
on microbial activities, for it might be possible that bacteria could be
inhibited by the rise of cyclonic weather situations while anticyclonic
weather situations might activate bacterial life.
Bortel's successor, H. Stolp especially became well-known by
the detection of the predatory bacterium Bdelovibrio bacteriovorus
in 1965. For this he has been honoured as the first scientist not being
a physician by the award of the Robert Koch Medal.
In the seventies and eighties H. P e t z o 1 d and R. M a r w i t z carried
out extensive studies on mycoplasms which were only recognized in
the sixties as being pathogenic to plants. To identify these pathogenic
organisms very extensive methodical and cytological research was
necessary, especially an application of the scanningelectronmicroscope.
Pesticides
The study of pesticides simultaneously began with the adoption

Englera7-1987 75
of the mandatory testing of pesticides in Germany. This testing of

pesticides was done at the Chemical Laboratory of the Research Centre
where J. Houben was employed since 1921. Houben simulta-
neously had a professorship of chemistry at the BerUn University. W.
Fischer continued Houben's work. The extension of the testing
of pesticides, however, is especially owed to H. Muller. This manda?
tory testing and approval today is a legal task of the Federal Biological
Research Centre (in Brunswick). Fischer in the fifties extended the
testing of the effect of pesticides on organisms. By this extension of
the research of this department he laid the foundation for the in-
vestigation of the crucial problems of the effects and side effects of
pesticides which has been continued by W. Ebing under ecochemical
and ecotoxicological aspects. Special attention is paid by I. Schup-
han to the analysis of residues in the ecosystem and to the develop?
ment of models of ecosystems. During these studies many methodical
and technical problems had to be solved. The extensive studies in the
field of gas chromatography of pesticides, however, are reflected in
bibUographic pubUcations (Ebing 1986).
Non-parasitic plant diseases
The field of non-parasitic plant diseases became particularly impor?

tant when A. Kloke joined the Federal Biological Research Centre in
1959. He and his colleagues worked on many diseases of vegetables
caused by a deficiency of nutrients (among others celery, white cab-
bage, lettuce). These diseases have been identified as a deficiency of
calcium and boron, respectively. Already in the sixties great attention
was given to damages of plants caused by radiation and mineral oil.
Also the effects of lead originating from exhaust gases of automobiles
on the vegetation along the roads have been studied. These studies
at last led to the passing of the Petrol-Lead-Law. Kloke (1982)
established guidelines for upper limits of the contamination of soUs
by heavy metals originating from industrial emissions, sewage sludge,
and garbage. These guidelines later have been partly revised (Kloke
1985) and are used in the Federal RepubUc of Germany and other
countries for legislative measures to protect soil and environment.
In the seventies H.-O. Leh studied the effect of deicing salts on
trees at the streets in towns and on the trees and shrubs along the hig-
ways and roads. By his studies he was able to preserve these plants in
towns and landscape, for in many towns the application of deicing
salts has been prohibited (Leh 1977). According to these studies the
median strip of highways now is planted with shrubs resistant to
deicing salt. These shrubs not only improve the scenery of the highways
but are also effective glare shields.
The main emphasis of the work of Kloke and his colleagues,
however, laid in the field of the effect of heavy metals in the environ-
ment, especially on vegetation. These studies have been carried on by
G. Schonhard who analyzed the pollution of many allotments and
agricultural economic plants with heavy metals.
Pests
Although pests undoubtedly are part of phytomedicine they only

shall be touched within the scope of this report on the development of
phytomedicine in Berlin. When looking at early papers from the Biolo?
gische Reichsanstalt it is obvious that most of them dealt with rodents,
birds, and parasitic nematodes to elucidate their biology in order to
find out methods for their control. In contrast to this the papers
dealing with entomological problems were rare, especially if one com-
pares the number of species dealt with. The reason for this discrep-
ancy may be the fact that at the beginning of this century entomolog?
ical research as a whole was not at its hight. The education at the
universities within the department of zoology was absolutely inade-
quate and entomologists when had not dealt with taxonomical prob?
lems did not have access to agriculture (with the exception of forest
protection).
Due to the lack of phytomedical education which later was deman-
ded with emphasis by A p p e 1 in the Biologische Reichsanstalt the first
scientists in most cases were botanists and pharmacists. Even H. Hase,
one of the most prominent zoologists of the Biological Research
Centre in 1920 got his international reputation by developing and
oranizing lice control during the First World War. This, of course,
was a field of public health rather than of agricultural zoology. Despite
this there have been many papers in Berlin dealing with entomological
Englera7-1987 77
problems. One of the first was the study of Frank who by his paper
on the pea beetle and its economic significance paved the way for a
modern phytomedical entomology. Important entomological studies
at the Biological Research Centre, however, have not been performed
in Berlin but in the branch offices of the Research Centre, so in the
field of research on grape Phylloxera (J. Moritz) and on aphids (C.
Borner). Research on problems arising from tropical pests which had
to be performed by the Biological Research Centre mainly was in the
field of morphology and systematics. Scientists like Morstatt who
had studied the pests of coconut trees and sorghum, however, pubUshed
the results of their research after having returned from the colonies
to Germany after the First World War.
In the forties and after the Second World War K. H e i n z e worked
on the green peach aphid Myzus (Myzodes) persicae and other aphids
which were related to the stUl relevant potato virus. These studies lead
to systematics of the virus transferring aphids. K. Mayer became
well known from his extensive research on dipters damaging plants.
He also was very interested in history and in 1959 pubUshed the book
"4500 Years of Plant Protection". Besides the pests of oUferous plants
Dora Godan especially dealt with slugs and snaUs detrimental to
plants. Her extensive knowledge has been recorded by her after her
retirement in the remarkable monograph "Pest slugs and snails. Bio?
logy and control" (Godan 1983).
Of utmost importance especially to practice of plant protection
was the nomenclatural work of G. Schmidt over many years. Al?
ready before the Second World War he pubUshed a comprehensive
compilation of the common names of insect pests in different countries.
After the Second World War he dealt with the German names of insect
pests which he pubUshed under the title: "Die deutschen Namen
wichtiger Arthropoden" (The German names of important arthropods")
(1970). In 1980 he pubUshed a supplement to this booklet.
In the field of stock protection F. Z a c h e r and later W. F r e y were
studying pests of stored agricultural products and food. They were
studying pests of textiles, too. After the Second World War the studies
have been directed towards pests in grain storage, their biology, and
control for instance by fumigation. These studies are of utmost econo?
mic importance and have been continued by R. Wohlgemuth and
his colleagues.
Sorauer and the "Handbook of Plant Diseases"
Finally we want to pay tribute to a leading personality of phyto?

medicine who has been still effective in our time (Wittmack 1917).
The greater part of his life P. Sorauer has not been in Berlin but was
occupied at the Agricultural Experiment Station Dahme, Brandenburg,
and the Royal Pomological Institute at Proskau near Oppeln. In 1893,
however, Sorauer moved to Berlin, gave lectures at the Humboldt
Academy and qualified as a university lecturer at an age of 63. Here he
worked almost until his death in 1916. His manifold activities in the
field of science and organization, however, were closely associated with
Berlin. He was for instance member of the Advisory Board of the
Imperial Biological Research Centre for Agriculture and Forestry as
well as member of the Special Committee for Plant Protection of the
German Agricultural Society. Here he cooperated very closely with
Frank who later became the first head of the Imperial Research
Centre. Together with Frank (and later with Rorig) Sorauer
edited on behalf of the German Agricultural Society the work: "Pflan-
zenschutz. Anleitung fiir praktische Landwirte zur Erkennung und
Bekampfung der Schadigungen der Kulturpflanzen". He also founded
the journal "Zeitschrift fiir Pflanzenkrankheiten" (Journal of Plant
Protection) and edited it until his death.
His most important work, however, already was initiated in 1874
when Sorauer in one volume edited the "Handbuch der Pflanzen?
krankheiten" (Handbook of plant diseases) which was published by
the publishing company Wiegand, Hempel & Parey (since 1881 Paul
Parey Publishers) in Berlin. In this first edition the damages caused
by sulphur dioxide which today it is much argued about already have
been described. The second edition, published in 1886, already consisted
of two volumes and the third edition, published in 1908-1913, had
three volumes. Today this "Handbook" consists of six volumes with 26
parts on the whole which are partly in the seventh edition. The authors
of the "Handbook" are well-known scientists in their field and they
have met over decenniums to create this book. So it can be said with
every justification that this "Handbook" is the standard work of phyto?
medicine of the German linguistic area and far beyond it.
Efforts to tackle those fields of operation which later have been
called registration and warning service originated from Sorauer who
Paul Sorauer
1839-1916
paid special attention to statistics of diseases and pests. With emphasis

he asked for more information about the circumstances of the arising
of diseases, about the weather etc. EspeciaUy he stressed the indivi?
dual predisposition of the plants for an attack of the parasites. And
the suggestion to breed strains of cultivated plants being resistant
to diseases and pests was supported by Sorauer, too. This field
even today is a problem of immediate interest for plant breeding.
Outlook
"Dahlem that meant ampleness of thinking and experimentation

and at the same time it was a code word for concentration and inte-
gration. Dahlem was a form of academic life which up to that time
certainly nowhere in history of research became evident." These words
of Friedrich Merkenschlager (on the quarter Dahlem in Berlin
where research institutes were concentrated) are undoubtedly not only
related to phytomedicine but to the whole of the research units in this
part of Berlin like the then "Kaiser-Wilhelm-Institute", the different
research centres, the Botanical Garden and Museum, and other places
of research.
Today the situation of research and teaching has changed in many
respects. The concentration, however, of institutes and personaUties
of science in Berlin, especiaUy in Dahlem, is stUl existing. Here we
find institutes of the Max Planck Society, Federal Research Centres,
institutes of agricultural sciences of the Technical University, and
research estabUshments of the Land Berlin. The Free University with its
numerous institutes impressively dominates the scenery. FinaUy the
significance of the infrastructure which is of great importance for
scientists and research should not be forgotten. Here we find libraries,
information offices, and computing centres which are concentrated
nowhere else like in BerUn.
There remains the hope, however, and we have to appeal to all
those who have the responsibUity that those who work in the field
of phytomedicine should have the chance to work in research and
teaching as Merkenschlager has stated for the twenties.
Englera 7-1987 81
References
Appel, O. 1919: Die Zukunft des Pflanzenschutzes in Deutschland. -

Angew. Botan. 1: 1-15.
? 1921: Die Organisation des Pflanzenschutzes im Deutschen Reich.?
Arb. DLG314: 119-136.
Bartels, R. & Koch, W. 1980: Wegbereitende deutsche Phytopatho-
logen. - Mitt. BBA H. 196.
Berliner, E. 1911: iiber die Schlaffsucht der Mehlmottenraupe. ?
Zeitschr. Gesamt. Getreidewesen 3: 63-70.
Bibliographie der Pflanzenschutzliteratur. - Berlin, Hamburg. 1914/
19(1921)- 1958(1962/63). - N. F. 1. 1970 ff. revised by W. Laux
et al. to 1939 by H. Morstatt, to 1958 by J. Barner.
Bochow, H. & Schumann, K. 1985: Die Entwicklung der Phytopatho-
logie und Pflanzenschutzwissenschaft an der Humboldt Universitat
zu Berlin. - Wiss. Z. Humboldt-Univ., Math.-Nat. R. 34: 223-
229.
Braun, H. 1952: Pflanzenschutz. - In: O. Keune (ed.), Manner die
Nahrung schufen. ? Hannover, p. 264-286.
? 1965: Geschichte der Phytomedizin. ? Berlin, Hamburg.
Chronik zum 75jahrigen Jubilaum der Biologischen Bundesanstalt
fiir Land- und Forstwirtschaft. 1973. - Mitt. BBA H. 148.
Ebing W. 1986: Gaschromatographie der Pflanzenschutzmittel XIV. -
Mitt. BBA H. 231.
Frank, A. B. 1897: Kampfbuch gegen die Schadlinge unserer Feld-
fruchte. ? Berlin.
50 [fiinfzig] Jahre Deutsche Pflanzenschutzforschung. Festschrift zum
fiinfzigjahrigen Bestehen der Biologischen Zentralanstalt fur Land-
und Forstwirtschaft in Berlin-Dahlem. 1949. - Berlin.
Gerlach, W. & Nirenberg, H. 1982: The genus Fusarium, a pictorial
atlas. - Mitt. BBA H. 209.
Godan, D. 1983: Pest slugs and snails. Biology and control. ? Stuttgart.
Handbuch der Pflanzenkrankheiten 1874-1985: Begr. von P. Sorauer.
In 6 volumes and several editions. ? Berlin, Hamburg.
Bd. 1: Die nichtparasitaren Krankheiten.
Bd. 2: Die virus- und bakteriellen Krankheiten.
Bd. 3: Pilzliche Krankheiten und Unkrauter.
Bd. 4: Tierische Schadlinge und Nutzpflanzen, 1. Teil.
Bd. 5:2.TeU.
Bd. 6: Pflanzenschutz.
Jahresbericht iiber die Neuerungen und Leistungen auf dem Gebiete
des Pflanzenschutzes. - Erstattet von M. Hollrung. ? Berlin. ?
1. 1898 (1899)-3. 1900(1902).
Contin. - Jahresbericht iiber die Neuerungen und Leistungen auf dem
Gebiete der Pflanzenkrankheiten. - 4. 1901 (1903) - 7. 1904
(1905).
Contin. - Jahresbericht iiber das Gebiet der Pflanzenkrankheiten. -
8. 1905(1907)- 16. 1913(1917).
Klinkowski, M. 1960: Gustav-Adolf Kausche. - Nachrichtenbl. Deutsch.
Pflanzenschutzd. N. F. 14: 196-199.
Kloke, A. 1982: Erlauterungen zur Klarschlammverordnung. - Land-

wirtschaftUche Forschung, Sonderheft 39: 302-308.
- 1985: Richt- und Grenzwerte zum Schutz des Bodens vor Uber-
lastungen mit Schwermetallen. ? Forschungen zur Raumentwick-
lung: 14: 13-24.
Kolbe, H. 1901: Gartenfeinde und Gartenfreunde. Die fiir den Garten?
bau schadlichen und niitzlichen Lebewesen. - Berlin.
Kolbe, W. 1985: 200 Jahre Pflanzenschutz im Zuckerriibenbau (1784-
1984). -Bonn.
? Bussmann, R. & Winter, F. 1983: Landbau und Ernahrung. ? Bonn.
Krober, H. 1985: Erfahrungen mit Phytophthora de Bary und Py-

thium Pringsheim. - Mitt. BBA H. 225.
Kruger, F. 1901: Albert Bernhard Frank. - Ber. Deutsch. Bot. Ges.:
19:(10)-(36).
Laux, W.: Die Lehre im Fachgebiet Phytomedizin. In: 200 Jahre

Landbauwissenschaften in Berlin, Schriftenreihe des Fachbereichs
Internationale Agrarentwicklung. [in press]
Leh, H.-O. 1977: Die Gefahrdung des Straftenbaumbestandes in Berlin

durch EinwUkung von Auftausalz. ? Berliner Naturschutzblatter
60:256-264.
Linskens, H. F. 1978: K. O. Muller 1897-1978. - Ber. Deutsch. Bot.

Ges. 91: 399-405.
Mayer, K. 1959: 4500 Jahre Pflanzenschutz. - Stuttgart.

Englera 7-1987 83
Merkenschlager, F. & Klinkowski, M. 1933: Pflanzliche Konstitutions-

lehre. Dargestellt an Kulturpflanzen. - Berlin.
Meyen, F. J. F. 1841: Pflanzen-Pathologie. - Berlin.
Miiller, K. O. 1958: Studies on phytoalexins. I. The formation and
immunological significance of phytoalexin produced by Phaseolus
vulgaris in response
to infections with Sclerotinia fructicola and
Phytophthora infestans. - Austr. J. Biol. Sci. 11: 275-300.
Pfankuch, E. 1943: Die Bedeutung unserer Kenntnisse von Aufbau
und Eigenschaften der Virusproteine fiir eine Chemotherapie der
Viruskrankheiten. - Mitt. BBA H. 67: 46-57.
Richter, H. 1938: Lupinenkrankheiten. - Mitt. BBA H. 58: 87-101.
Riehm, E. 1938: 40 Jahre Biologische Reichsanstalt. ? Nachrichtenbl.
Deutsch. Pflanzenschutzd. 18: 49-51.
Schacht, H. 1856: Bericht an das Konigliche Landes-Oeconomie-
Collegium iiber die Kartoffelpflanze und deren Krankheiten. ?
Berlin.
Schlumberger, O. 1956: Otto Appel 1867-1952. - Ber. Deutsch.
Bot. Ges. 68:211-215.
Schmidt, E. W. 1948: Die kranke Pflanze. - Kleinmachnow.
Schmidt, G. 1970: Die deutschen Namen wichtiger Arthropoden. ?
Mitt. BBA H. 137.
Schuhmann, G. 1966: Untersuchungen iiber die physiologische Spe-
zialisierung von Tilletia caries (DC.) Tul. und Tilletia controversa
Kiihn in Deutschland und das Resistenzverhalten von Weizensorten. ?
Mitt. BBA H. 117.
Schwartz, M. 1925: Anleitung zur Bekampfung des Kartoffelkafers. ?
Nachricht enbl. Deutsch. Pflanzenschutzd. 4: 15-16.
6 [sechs] Jahre Blauschimmelkrankheit des Tabaks in der Bundes-
republik Deutschland (1959-1964). 1966. - Mitt. BBA H. 120.
Stapp, C. 1939: Bakterielle Pflanzenerkrankungen. - Zentralbl. Bak-
teriol. I. Abt., Supp. 144: 94-108.
Stolp, H. 1965: Isolierung von Bdellovibrio baeteriovorus. ?Zentralbl.
Bakteriol. I.Abt., Suppl. 1: 52-56.
Ueberlein, J.-H. 1984: Notizen zur Geschichte der Phytomedizin. ?
Jahrb. Verein Geschichte Berlins 33: 93-103.
Ullrich, J. 1970: Gedenkrede fiir Friedrich Merkenschlager. - Aus der
Ulsamer, W. 1970: Prof. Dr. Friedrich Merkenschlager. ? Aus der
Wehsarg, O. 1954: Ackerunkrauter. Biologie, allgemeine Bekampfung

und Einzelbekampfung. ? Berlin.
Wittmack, L. 1917: Paul Sorauer. - Ber. Deutsch. Bot. Ges. 34: (50)-
(57).
WoUenweber, H. W. 1916-1935: Fusaria autographice delineata. -
Berlin: Selbstverlag, 1200 Tafeln.
? 1931: Fusarium -Monographie. Fungi parasitici et saprophytici.
Z. Parasitenkd. 3: 269-516.
? 1943: Fusarium -Monographie. II. Fungi parasitici et saprophytici.
Zentralbl. Bakteriol. Parasitenkd. Infektionskr., II, 106: 104-105,
171-202.
? & Reinking, C. A. 1935: Die Fusarien, ihre Beschreibung, Schad-
wirkung und Bekampfung. ? Berlin.

Prof. Dr. Wolfrudolf Laux, Biologische Bundesanstalt fiir Land- und
Forstwirtschaft, Kdnigin-Luise-Strafce 19, D-1000 Berlin 33.
Englera 7 - 1987 85
Herbert Sukopp
On the history of plant geography and plant ecology in Berlin
The estabUshment of ecology as a scientific discipline around the

middle of the last century was closely Unked to the breakthrough of the
descendence theory and the selection theory formulated by Darwin
(1809-1882) to explain the evolution of the organic world. In addition
to study of the internal and external structure of organisms, which
had hitherto been at the centre of attention, demands were now made
to investigate environmental requirements of plants and animals, in
order to be able to interpret the adaptations of organisms. These
considerations led the zoologist Ernst Haeckel (1834-1919) to the con-
cept of a branch of science "ecology", a "science of adaptation", which
he drew up in 1866 in his "General morphology of organisms":
"When we spoke of the external existence conditions above, our

eye was mainly on the inorganic, the influence of Ught, warmth,
humidity, the inorganic nutrition etc. Much more important than
these, however, and much more influential on the alteration and
adaptation of species are the organic, i.e., the interactions among
aU organisms. Every individual type of organism is dependant on
many others that exist with it in the same place, and which can
be either harmful, neutral or beneficial. Every organism has
enemies and friends among the others, which may either threaten
or faciUtate its existence. The former may deprive it of nutrition,
e.g. parasites, the latter may nourish it, e.g. host plants. Obviously
the number and characteristics of all the organic individuals
which Uve together in the same place influence one another, and
every alteration of the number and characteristics of a single
species must have an effect on the others in interaction with it.
The fact that these mutual interactions of all neighbouring
organisms are highly important, and have a much great er influence
on the alteration and adaptation of species than the inorganic
86 Sukopp: Plant geography and plant ecology
existence conditions, was first clearly determined by Darwin.

Unfortunately these highly complicated interrelationships of
organism are for the most part completely unknown, since they
have hitherto received almost no attention, so that here an
enormous, highly interesting and important field for future
investigations discloses itself. The
ecology, or the science of
nature's economy, a part of physiology, which has not yet been
included as such in the textbooks, promises in this respect to bear
the most wonderful and surprising of fruits."
18 Century
The 18th Century saw the development of the fundamentals of

Natural History, the precursor of plant geography and ecology. Natural
History was concerned with the description and classification of living
organisms, and with their distribution. Zoological and botanical in?
vestigations on all continents led to the "discovery of diversity" (Mayr
1982). The number of known plant species, for example, grew from some
3,000 at the beginning of the 17th Century (Tabernaemontanus) to
10,000 at the beginning of the 18th Century (Tournefourt), to 30,000
by the beginning of the 19th Century (A. P. de Candolle) and reached
90,000 by the middle of the 19th Century. In Brandenburg, the Berlin
botanist Johann Gottlieb Gleditsch (1714-1786) had already listed
1,200 plants; he was aware of the existence of various floristic elements.
Alongside native German plants there were also those "(which) are
common in Russia beyond Moscow, also in Livonia around Riga, in
some areas of the Ukraine, in North American districts, in Meifien,
Thuringia, Bohemia, northern and middle regions of France, in Switzer-
land, on the lower and middle Alps and on the Harz Mountains. The
poor sand and heath soil has, furthermore, numerous plants unique to
that area."
He was also aware that each landscape in Brandenburg had its own
specific species in addition to those common to all districts, but that
there were no endemic taxa in Brandenburg.
His studies were prompted by the various environmental problems
at a time of overexploitation of natural resources. Gleditsch worked
in a number of areas of management: agriculture, forestry, bee-keeping
Englera7-1987 87
and crop protection (Krausch 1977, cf. also Natho 1973, 1975). A
special problem faced at that time was the unstable sandy areas which
in those times covered considerable land areas and threatened to result
in the deteriorationof surrounding land. For 1782 Kloden (1832) Usted
no fewer than 23 such sandy areas in the Middle Mark which were
larger than 100 acres (25 ha). Gleditsch gave a detaUed account of
their origin and character as well as options for stabUizing these sandy
areas.
At the centre of education in natural sciences in BerUn in the 18th
Century was the CoUegium medico-chirurgicum, originaUy founded in
1685 for the training of miUtary doctors, surgeons and pharmacists.
Finances were provided by the Academy of Sciences, whose forerunner
the Society of Sciences had administered the Court and Vegetable
Gardens since 1718. Thus it was that the Director of the Botanical
Garden had the responsibUity for pharmacology and botany at the
CoUegium medico-chirurgicum. Gleditsch held this position from
1744 to 1786. It was during this period that LinnS's effort to system-
atically classify all organisms focussed attention on the nature of sexual-
ity, especially of plants. In 1749 Gleditsch conducted his "ex-
perimentum beroUnense" in the Botanical Garden to Schoneberg, which
involved the artificial pollination of a palm (Chamaerops humilis). The
development of the fruit verified the sexuaUty of plants. Gleditsch
held lectures on forestry at the Bergakademie (founded in 1770); he
created the basis for regulated forestry practice in Prussia.
It was not only in government institutions that medical expertise
and botanical interest were coupled. A special role was played by Ernst
Ludwig Heim, Town Doctor for Spandau, who became famous asthe
tutor of the brothers Alexander and WUhelm v. Humboldt. Heim
also stimulated the love of botany in the Rector of the "Large School"
in Spandau, Christian Conrad Sprengel (1750-1816) (see Meyer
1953, 1967; Stocker 1979), of whom Carl Ludwig Willdenow
(1765-1812) wrote in the historical part of his "Grundrifi der Krauter-
kunde" (WUldenow 1810, 5th ed., p. 58): "Christian Conrad Sprengel,
formerly Rector now private savant in BerUn, discovered with pains-
taking observation the true method by which nature has provided for
the fertUization of plants."
Sprengel stated clearly that the size, shape and colour of petals,
the arrangement and relative position of the flower organs, and the
production of fragrance and nectar have a specific importance for the life
of the plant, and are closely related to the transfer of pollen by insects
or the wind. In the introduction to his book on "The revealed secrets of
nature in the construction and fertilization of flowers" (1793) he
wrote (p. 2):
"Examining the flower of the Geranium sylvaticum I found that

the lower part of the petals were covered with fine, soft hairs on
the inner side and at the edges. Convinced that the wise Creator
of nature had not produced even a single hair to be placed with?
out precise design, I considered the possible function such hairs
might have. It soon occurred to me that if one assumed that the
five drops of sap secreted by just as many glands are determined
to provide nourishment for some insects, then one could not find
it surprising if precaution had been taken to prevent this sap from
being spoilt by rain, and that the hairs had been placed there with
this intent."
Thus Sprengel became the founder of floral ecology. His obser?

vations formed a basis of knowledge which was first rediscovered by
Darwin, and on which Darwin repeatedly drew (Wichler 1936). S p r e n-
g e l's floral ecology was without rival in Berlin for a long period.
In 1773 the Berlinsche Naturforschende Gesellschaft (Berlin Nature
Research Society) was founded (Becker 1973, Herter & Bickerich
1973). Around the end of the century the meetings of the Society
clearly reflected activity in the field of natural sciences (Leopold v.
B uch, Alexander v. Humboldt, Willdenow, Heim). It is the
oldest German society for natural sciences still in existence, and gave
birth to the Botanical Society of the Province Brandenburg (since 1980
the Berlin Botanical Society) and to the German Botanical Society.
The academic activity in this period was not restricted to the basic
sciences, but also concerned itself with practical problems of agriculture
and forestry. The beginnings of scientifically based agricultural practices
are directly attributed to Albrecht Dieter Thaer (1752-1828), a
physician from Celle. In 1804, T h a er was given an estate in Moglin by
the Prussian King for teaching and research purposes. In 1810, this
institution was coupled with the Berlin University. T h a e r was granted
a professorship for Agriculture and Fiscal Sciences, which he occupied
until 1819. Thaer must be credited with the application of natural
Englera7-1987 89
sciences to agriculture, e.g., introducing crop rotation, and promotion

of potato and sheep farming.
Forestry science prior to the founding of the BerUn University was
influenced by Gleditsch and Willdenow. Lectures were held in
the Bergakademie. After the death of Gleditsch, forest botany and
forest science in BerUn were continued by August Ludwig von Burgs-
dorf f (1747-1802). He took over the Forest Inspection Tegel in 1777
and also administered part of the forest. In 1783 he pubUshed the first
part of a treatise on tree species, on the beech, with a foreword by
Gleditsch. In 1787, the second part appeared on "Native and exotic
oaks". The same year saw the pubUcation of his two-part "Guide to
the growth and appropriate planting of native and exotic timbers which
survive outdoors in Germany and in similar climates". In 1788 the first
part of a forestry handbook was pubUshed, which eventually resulted in
three editions. The second part followed in 1796. Burgsdorff's
importance lay in his encouragement of the cultivation of exotic
woody cultivars with seeds obtained from North America. From Tegel
he marketed seeds and plants (Scamoni 1955).
In 1821 a Forestry College was founded by Friedrich Leopold P f e il
(1783-1859), in cooperation with the University. In 1830 this was
discontinued and the forest science tradition in Eberswalde as the
Hohere Forstlehranstalt (after 1868 Forest Academy) was contin?
ued.
Developments foUowing the founding of the BerUn University
The development of botany in the 19th Century was greatly in?

fluenced by the founding of the BerUn Friedrich WUhelm University in
1809. The former Court and Academy Garden in Schoneberg, which
Willdenow (1765-1812) directed from 1801 to 1812, was incor?
porated into the University as a botanical garden in 1810 (Timler &
Zepernick 1978). Willdenow received the Chair of Botany in the
Faculty of Philosophy. During W i 1 ld e n o w' s pharmaceutical appren-
ticeship his uncle Gleditsch had taken over the botanical instruction.
"At Sprengel's side he got to know the location of several rare plants
around Berlin. Uncle Gleditsch knew how to nurture this scientific
curiosity and to direct it along the right paths, especially through the
lessons that he gave the apprentice pharmacist. It is from this time that
the beginnings of the Herbarium date. He continued this endeavor ener-
getically until his death" (Konig 1898). He published his "Grundrifc der
Krauterkunde" in 1792, which went through many editions, was widely
read, and had great significance for plant geography, among other
things. The chapter "History of plants" can be regarded as a foundation
for this branch of science. The first paragraph of the book (in the 5th
edition 1810) runs:
"The history of plants involves the influence of climate on

vegetation, the changes of climate endured by plants and how
they have been preserved by Nature, migration of plants, and
finally their distribution over the globe."
Probably for the first time - at least in a prominent place ? we find

"history" used in a modern sense in a work of Natural History. The
term history (Geschichte) had until then not had a temporal perspective,
but referred to the "reporting of facts" ? in contrast to "philosophy"
which sought "reasons for facts" (Trepl 1987). In this tradition, Cha?
misso (Endtmann 1983) describes in 1827 the effect of human
culture after his world trip on the Rurik 1815-1818:
"Wherever humans settle, the face of nature is changed. His

domesticated animals and plants follow him; the woods become
sparse; and animals shy away; his plants and seeds spread them-
selves around his habitation; rats, mice and insects move in under
his roof; many kinds of swallow, finch, lark and partridge seek his
care and enjoy, as guests, the fruits of his labor. In his gardens
and fields a number of plants grow as weeds among the crops he
has planted. They mix freely with the crops and share their fate.
And where he no longer claims the entire area his tenants estrange
themselves from him and even the wild, where he has not set
foot, changes their form."
This quotation is taken from an instructional work "Botany for the
non-botanists" which Chamisso wrote for the Culture Ministry "A
survey of the most useful and harmful plants, whether wild or culti?
vated, which occur in North Germany. Including views of botany and
the plant kingdom", a title in the tradition of Natural History.
Willdenow had developed the ideas of plant geography together
Englera7 - 1987 91
with Alexander v. Humboldt (1769-1859), who had been a close

friend since 1788(Jahn 1966, Meyer-Abich 1967). Humboldt's
fundamental work on plant geography appeared in 1805 and 1807 in
Paris as "Essai sur la Geographie des Plantes", and 1807 in Tiibingen as
"Ideen zu einer Geographie der Pflanzen". In his "Ideen zu einer
Physiognomik der Gewachse" in 1806 Humboldt introduced the
concept of the "association" for plant communities and recognized the
significance of Ufe forms for the deseription of vegetation for the earth.
Humboldt writes in 1806 about the "Heathland, this 'Gruppie-
rung' (or in the French association) of Erica vulgaris, Erica tetralix,
Lichen icmadophila and Lichen haematoma".
More significant than the concepts he introduced was his under-
standing of science. Humboldt "strived ... for an overview of the
perceptible world in its entirety and at gaining the broadest possible
comprehension of the interaction of the forces" (Schmithiisen 1957).
In "Naturgemalde der Tropenlander" (1807) we find: "In the great
Unkage of causes and effects, no substance, no activity may be con?
sidered in isolation. The equiUbrium which rules amongst the pertur-
bations of apparently antagonistic elements results from the free
play of dynamic forces, and a complete overview of nature, the final
purpose of all physical studies, can only be achieved if no force and no
formative process is neglected, so that for the philosophy of nature a
broad, promising field of study wUl be formulated."
Here Humboldt adopts an idea of Friedrich WUhelm S c h e 11 i n g
(1775-1854), the head of the German "Naturphilosophie", that aU
natural phenomena are based "on the never ending conflict of the
opposing basic forces of material" (Leps 1980: 31). Humboldt's
"hoUstic" and "physiognomic" approach had a more aesthetic than
scientific meaning. The latter was contained in reformulations by some
plant geographers in the 19th Century, above aU Griesebach. Thus
Humboldt's conception became the starting point for the (further)
development of ecology (Trepl 1987, cf. Hard 1970). Humboldt not
only set out a definitve basis for the development of ecology, he
founded plant geography in an institutional sense, which in the BerUn
tradition was continued by E n g 1 e r and Diels.
The continuation of the plant geography work of Willdenow and
A. v. Humboldt BerUn provided a great impetus for the study of the
native flora. Although the natural distribution of plant species provides
no particular justification for treating Berlin as an independent area, it

does play a special role in the research.
At the time of Aschersona number of Berlin flora works already
existed, beginning with the purely illustrative work of H e c k e r (1742,
1757/8) through the Latin works of Willdenow (1787), Reben-
tisch (1805), Kunth (1813), Chamisso (1815) and Brandt
(1825) to the first "Flora" in German by D iet rich (1824). Ascher?
son published his "Studiorum phytographicorum de Marchia Branden-
burgiensi specimen" in 1853, and layed down the fundamentals of
floristics in the Mark in numerous studies. He followed his "Flora of
the Provinz Brandenburg with a Special Flora of Berlin" (1864), which
with its comprehensiveness and accuracy represent the pinnacle of
floristics in the Mark. One of the earliest vegetation scientists was
Schweinf urt h, who published in 1861/62 an "Attempt to outline the
vegetation around Strausberg and Blumenthal near Berlin". Research on
the flora was later pursued by Graebner, until with the publication
of Waldenburg's treatise on the phytogeographic position of the Mark
Brandenburg a stage of completion was more or less reached.
From the industrial revolution to 1945
Humboldt, whose extensive contributions to natural science had

greatly influenced the first half of the century, died in 1859. A com-
ment he made in a talk at the opening session at the Assembly of
German Natural Scientists and Physicians in Berlin in 1828 on the
founding of this society is characteristic of him, namely to have "united
all branches of physical knowledge in a true and deep feeling for unity
of nature."
The decades following the Humboldt era up to the present are
characterized by an ever-increasing specialization of knowledge. In
1866 Ernst Haeckel (1834-1919) presented a categorization of the
various component areas of zoology in his "General morphology", and
this was followed by many other similar attempts. This led directly to
further specialization. Today not only are the individuals engaged in
systematics specialized into small taxonomie groups, but also general
botany and zoology have split into separate fields which though parallel
in many respects, scarcely recognize one other and then often with
Englera7 - 1987 93
some disdain (Tembrock 1959). Some of the most notable aspects of

this complicated development are described for the situation in Berlin.
The year 1878 was of major importance for the development and
extension of botanical teaching and research at the Berlin University.
In addition to the professorship for Systematic Botany, which was
compled with to the directorship of the Botanical Garden and Museum,
a new chair and a Botanical Institute were estabUshed (from 1914 the
Plant Physiological Institute).
Adolf Engler (1844-1930; Stafleu 1981) was offered the Chair
for Systematic Botany in 1889. Under his editorship (in part with Karl
Prantl) 19 volumes of "NatiirUche PflanzenfamiUen" appeared from
1887 to 1909. In addition to systematics, plant geography was a second
field of interest for Engler, demonstrated by his account of plant
geographie formations on the land to be occupied by the new botanical
garden in Berlin-Dahlem. Engler was responsible for plant geography
gaining a permanent place at the BerUn University, and he did pioneering
work towards a synthetic approach in this field. His "Versuch einer
Entwicklungsgeschichte der Pflanzenwelt" (1879, 1882), which por-
trayed plant distribution and differentiation as evolutionary processes
in geological periods was in many ways a very influential work. The
series of volumes edited together with Drude under the title "Vege?
tation of the Earth" is a "worthy predecessor of present-day ecological
research" (Stafleu 1981: 33).
With Eugen Warming's (1841-1924) textbook (1895; in German
1896: "Lehrbuch der Okologischen Pflanzengeographie") ecology
achieved the status of a mature science. In BerUn, Paul Graebner
(1871-1933) curator of the Botanical Garden, pubUshed his detaUed
descriptions of vegetation in this vein, e.g. in 1895 in his dissertation on
the Hundekehlefenn in Grunewald.
In 1922 the BerUn Botanical Garden initiated the mapping of plant
distributions in Germany. FoUowing an appeal byJ.Mattfeld(1895-
1951) hundreds cooperated in the mapping of plant species on the basis
of the grid system of the German ordinance (cartographie sheets) from
1932 to 1942 under Fr. Mattick. The results of this work were
almost entrrely destroyed in the Second World War. The remaining
original material serves today as a source for the floristic mapping of
Middle Europe. The curators of the Botanical Museum also included
Friedrich Markgraf (1897-1987) and Eberhard Ulbrich (1879-
1952; Mildbraed 1954), who dedicated themselves expressly to the

vegetational science which developed from plant geography.
The discussion of natural philosophy problems ? especially the

problem of the holistic treatment of living systems, was strongly
influenced in Berlin by the Director of the Kaiser Wilhelm Institute for
Biology, Max Hartm ann (1876-1962; Nachtsheim 1963). In an
essay in 1938 entitled "Zum Begriff der 'Ganzheit' lebender Sys-
teme", Heinrich Jakob Feuerborn (1883-1979), curator at the
Zoological Institute at the University, followed Hartmann (1935) in
rejecting the view of "Ganzheit" as a superior operative principle, and
emphasized that unity and entirety did not solve any problems - it was
a way of indicating a problem. He included both organisms and living
communities in his considerations.
The concept of the living community as an entity played a special
role in forestation. It led Alfred Moller (1860-1922), who had been the
Director of the Forstakademie in Eberswalde, to develop the concept
of "permanent forest", which he expressed at length in his book "Der
Dauerwaldgedanke, sein Sinn und Bedeutung" (1922), written shortly
before his death. The development of the forest should be disturbed as
little as possible by the management practices and the "constancy of
the forest system" must be retained. Moller's successor at the Fore?
station Institute of the Forstliche Hochschule Eberswalde, to which
the status of the Academy had been raised, was Alfred Dengler (1874-
1944). After the death of Moller he engaged himself energetically in dis-
putes concerning the issue of "permanent forests", warned against
exaggeration and inadmissable generalizations and repeatedly emphasized
the importance of local site factors in all forestry practices. His lectures
were compiled in 1930 to form the standard textbook "Waldbau auf
okologischer Grundlage" (cf. Kollektiv der Mitglieder der Fakultat 1961).
The increasing tendency toward specialization in science was con-

fronted with the necessity to solve practical problems arising from the
stormy industrial development, population growth and the develop?
ment of urban centres. These problems required the application of a
broad spectrum of scientific and technical knowledge. Thus the Govern-
ment of Prussia or of the Empire felt it necessary to create a number of
institutes in which scientists from various disciplines could work
together addressing practical questions. "The Land and Reichs Institutes
Englera7-1987 95
thus realized at the turn of the century the concept of interdisciphnary

cooperation on distinct practical problems" (Ruske 1979).
Three examples demonstrate this cooperation of scientists to solve
practical problems:
? Water supply ? sewage purification
? Pest control
? Nature conservation
The most pressing problems arising from the rapid industrial devel?
opment of the German Empire were those of water supply and sewage
disposal in the urban areas. A major contribution to the solution of
questions of city hygiene in Berlin in the second half of the 19th
Century was made by the Rudolf Virc how (1821-1902). He pursued
questions of epidemiology and pubUc health intensively (Ackerknecht
1957, Burmeister et al. 1985). He felt that at the heart of all measures
to combat epidemics such as abdominal typhus or to reduce the cata-
strophic infant mortality lay in the installation of sewage systems. He
designed such a system for BerUn, the first section of which was com-
pleted in 1877. This system did not deposit waste water into the Spree
River but instead used special sewage-farms (cf. Kolkwitz 1909) whose
hygienic control was a special concern of V i r c h o w.
In 1901, the Prussian State Government founded a Royal Research
and Testing Institute for Water Supplies and Sewage Disposal located
in BerUn (today, Institute for Water, Soil and Air Hygiene at the
Federal Health Office) (Naumann 1961).
Richard Kolkwitz (1873-1956) and Maximillian Marsson
(1845-1909) were appointed as biologists in the Institute, and the prin-
ciples they developed for the biological assessment of water based on its
flora and fauna were of particular importance. Kolkwitz and Marsson
recognized the importance of indicator organisms to evaluate the purity
of water. The new results were elaborated in their "saprobiotic" system
(Kolkwitz & Marsson 1902, 1908, 1909b). This system is the basis for
the current ecological evaluation of water quaUty (cf. Elster 1982,
Leschber 1986). Under the direction of Kolkwit z, the first succesful
rehabiUtation of a lake was carried out. K o lk w it z (1909a, 1914) had
investigated the mass growth of algae in the Lietzensee and came to the
conclusion that the decisive factor for algae production lay in the
almost continual introduction of nutrients from the mud at the bottom
of the lake. He developed a method to remove these by swilling out the

nutrient-rich mud-layers at the bottom of the lake with nutrient-poor
water ? a method which proved to be a success in the Lietzensee. For his
fundamental contributions to biological-ecological water analysis he
was awarded an honorary medical-doctorship at Berlin University in
1926 (Leps & Burmeister 1987).
The disposal of industrial and domestic waste water, and its rele-
vance for fisheries led to the establishment of a research station in
Friedrichshagen at Miiggelsee by Johannes Frenzel and his successor
Paulus Schiemenz (1858-1936) which was later taken over by the
Prussian State and was known, from 1920 on, as the Prussian Institute
for Fishery. Primarily under the direction of Hans-Helmut W u n d s c h
(1887-1972) a series of important works were written ("Beitrage zur
Fischereibiologie markischer Seen"), which dealt with metabolic
processes in the lakes and the typology of the lakes (Schaperclaus 1957,
Mayer-Waarden 1970).
Pest control in agriculture and forestry was the goal of the Imperial
Biological Research Centre for Agriculture and Forestry, which was
established in 1905 from a former department of the Imperial Health
Office. This Centre was a particularly good example of interdisciplinary
ecological research, the need for which had already been recognized.
The tasks of the biological section included "research on environmen-
tal requirements of agricultural animal and plant pests in order to
establish a basis for their systematic control, and at the same time to
study beneficial organisms, such as pollinating insects and the natural
enemies of pests; furthermore to study environmental factors detri-
mental to agriculture and forests, such as the pollution of the air by
smoke or waste gases; and finally, the collection and publication of
research from the Institute Centre and other sources".
This programme signified from the very beginning a need to estab?
lish interdisciplinary research in the Centre, since successful control of
pests and diseases could only be achieved when all the biological and
environmental parameters were taken into consideration (Ruske 1979).
The Biologische Reichsanstalt gained an international reputation
(Schlumberger 1947), particularly under the direction of Otto Appel
(1867-1952).
The Prussian State directed attention toward nature conservation
Englera7 - 1987 97
at the turn of the century (Schoenichen 1954, Klose 1957, Auhagen &
Sukopp 1984). Following a classic memorandum by Hugo Conwentz
(1855-1922) in 1904 on "Die Gefahrdung der Naturdenkmaler und
Vorschlage zu ihrer Erhaltung" a state advisory board was founded in
1906 in Danzig for the preservation of natural areas and was transfered
six years later to BerUn. In numerous lectures and essays, coUected by
Wahnschaffe et al. in the "Grunewald-Buch" (1907, 1912) the
"importance of conserving the Grunewald moors" (Beitrage zur Natur-
denkmalpflege 1910, p. 135-143) was emphasized. Kurt Hueck
(1897-1965), one of the most important individuals in the develop?
ment of plant geography and vegetation science, was employed at the
State (later Imperial) Office from 1924-1944 (Kostler 1955, Berger-
Landefeldt 1965, Grosser 1966). He produced the first example of
vegetation cartography in Germany based on phytosociological records.
The condensed results of this work appeared in the three-volume "Die
Pflanzenwelt der deutschen Heimat" (1928-1933). As a continuation
of this work on Germany's vegetation the "Pflanzengeographie Deutsch?
lands" was pubUshed in 1937. The conclusion of this work were the
vegetation maps for Middle-Europe (1938), the German Reich, Berlin
Sheet (1943) and Lower Saxony (1948).
Developments since 1945 in West BerUn
Research after 1945 was concerned with the rapid changes of

flora and vegetation under human influence aU over the world, but
especially in an area such as Berlin including applications in nature
conservation (Auhagen & Sukopp 1984, Sukopp & Schneider 1981).
This was emphasized more in the BerUn research than was the case
for most other ecological research centres. This developed from general
considerations of human influence on vegetation (Sukopp 1969, Blume
& Sukopp 1976) to the production of "Red Books" of endangered
species (Sukopp & Elvers 1982, Klawitter & Schaepe 1985) and to an
assessment of plant and animal communities for biotope mapping in the
Landscape Program of Berlin (Arbeitsgruppe Artenschutzprogramm
BerUn 1984).
The special situation of BerUn led to a particular emphasis on urban
ecology. Study of ruderal flora and vegetation began with the work of
S c h o 1 z (1960) and was continued by several projects: Sukopp et al.

(1971),Bornkamm& H ennig (1982). Recent development in plant
ecology includes experimental ecology, emission ecology, succession
research, population biology of plants in the field, experimental sites
and the laboratory of the Institute of Ecology (Bornkamm & Kohler
1987).
In 1972 the institutes for Applied Botany (Director Ulrich Berger-
Landefeldt, 1912-1967), Soil Science and Horticulture from the
Faculty of Landscape Planning at the Technical University made
application to be united together as an Institute for Ecology (Sukopp &
Schneider 1979). Later on the Institute was complemented by limno-
logical and climatological groups (Sukopp 1983).
The conflict between the increased specialization in the biological
and ecological sciences, which began before the turn of the century,
and the need for interdisciplinary approach to practical problems has
continued to the present day. The current problems of environmental
protection, and landscape preservation require new forms of cooper-
ation among disciplines. Therefore an interdisciplinary research group
"Ecology and Environmental Research" was founded in 1972 (Sukopp
1977, Bornkamm 1983).
In the Botanical Museum Dahlem, the tradition of the Engler era was
continued after 1945 with the publication of further volumes of "Na-
turliche Pflanzenfamilien" and the 12th reworked edition of the "Sylla-
bus der Pflanzenfamilien". In the sixties and seventies a "Flora of
Togo" (in French) was published. Since 1978 an critical inventory of
mediterranean flora (Med-Checklist, 1984-) and a South Aegean Project
were conducted (by W. Greuter) as well as a study of rain forest
mosses ("Bryotrope").
Plant geographical research in the Instituteof Plant Taxonomy and
Plant Geography concentrated on pollination ecology and an atlas work
on Near East (Frey & Kurschner 1985). Berlin tradition in vegetation
mapping of Africa (Mildbraed) was continued by contribution to
the mapping project of the Deutsche Forschungsgemeinschaft (German
Research Community) (Bader 1967).
This account of developments after 1945 is restricted to West Berlin
since a condensed history of research in East Berlin has recently been
published (Leps 1985).
Englera7- 1987 99
References
Ackerknecht, E. H. 1957: Rudolf Virchow. Arzt, PoUtiker, Anthro-

pologe. ? Stuttgart.
Arbeitsgruppe Artenschutzprogramm BerUn, Leitung: Sukopp, H.,
Red.: Auhagen, A., Frank, Heidemarie & Trepl, L., 1984: Grund-
lagen fiir das Artenschutzprogramm BerUn. ? Landschaftsentw. u.
Umweltforsch. 23: 1-993.
Ascherson, P. 1864: Flora der ProvinzBrandenburg, der Altmark
und des Herzogthums Magdeburg, 2. Abt. Specialflora von BerUn. ?
Berlin.
Auhagen, A. & Sukopp, H. 1984: GeschichtUches zum Berliner Natur?
schutz. - Berlin. Naturschutzbl. 1984: 4-14.
Bader, F. 1967: Die Vegetationsgeographie auf dem Blatt Lake Victoria
im Afrika-Kartenwerk der Deutschen Forschungsgemeinschaft. ?
Ber. Dtsch. Bot. Ges. 80: 291-297.
Becker, K. 1973: Abriss einer Geschichte der Gesellschaft Naturfor-
schender Freunde zu BerUn. - Sitzungsber. Ges. Naturf. Freunde
BerUn N. F. 13: 1-58.
Berger-Landefeldt, U. 1965: Kurt Hueck. - Nachrichtenbl. Natursch.
Landschaftspfl. 36: 45-46.
Blume, H.-P. & Sukopp, H. (1976): Okologische Bedeutung anthro-
pogener Bodenveranderungen. ? In: H. Sukopp, & W. Trautmann
(ed.), Veranderungen der Flora und Fauna in der Bundesrepublik
Deutschland. - Schr. R. Vegetationskunde 10: 75-89.
Bornkamm, R. 1983: 10 Jahre Projektgruppe Okologie und Umwelt-
forschung. ? In: H. Sukopp (ed.), 10 Jahre Institut fiir Okologie. -
Berlin, p. 61-68.
- & Hennig, U. (1982): Experimentell-okologische Untersuchungen
zur Sukzession von Ruderalpflanzen-Gesellschaften auf unterschied-
lichen Boden. I. Zusammensetzung der Vegetation. ? Flora 172:
267-316.
- & Kohler, M; 1987: Ein Naturgarten fiir Lehre und Forschung. -
Landschaftsentw. u. Umweltforsch. 45: 1-133.
Burmeister, K.-J., Grosser, J. & Leps, G. 1985: Urbane Okologie im
gesundheitspolitischen Konzept Rudolf Virchows (1821-1902). -
3. Leipziger Symposium urbane Okologie, 13. November 1985.
Conwentz, H. 1904: Die Gefahrdung der Naturdenkmaler und Vor?
schlage zu ihrer Erhaltung. ? BerUn.
Elster, H.-J. 1982: Zur Definition der "Gewasser"- bzw. "Wasser-

giite" und iiber die limnologischen Grundlagen ihrer Beurteilung
in Vergangenheit und Zukunft. ? Schriftenreihe Ver. WaBoLu
54:21 ff.
Endtmann, K. J. & M. 1983: Adelbert von Chamisso (1781-1838) -
ein markischer Botaniker. ? Gleditschia 10: 13-27.
Feuerborn, H. J. 1938: Zum Begriffe der "Ganzheit" lebender Sy-
steme. - Naturwiss. 26: 761-771.
Frey, W. & Kiirschner, H. 1985: SFB 19, Tiibinger Atlas des Vorderen
Orients, Abschlufibericht des Faches Botanik 1974-1985. - Berlin.
Graebner, P. 1895: Studien iiber die norddeutsche Heide. Versuch
einer Vegetationsgliederung. - Bot. Jb. 20, 500-654.
Grosser, K. H. 1966: In memoriam Kurt Hueck und Friedrich Solger. -
Naturschutzarbeit in Berlin und Brandenburg 2: 22-25.
Haeckel, E. 1866: Generelle Morphologie der Organismen. ? Berlin.
Hard, G. 1970: Der "Totalcharakter derLandschaft". Re-Interpretation
einiger Textstellen bei Alexander von Humboldt. ? Geograph. Z.,
Beih. A. v. Humboldt.
Hartmann, M. 1935: Analyse, Synthese und Ganzheit in der Biologie.?
Sitzber. Preuss. Akad. Wiss., Phys.-Math. Kl., 135: 366-393.
Herter, K. & Bickerich, R. 1973: Die Mitglieder der Gesellschaft Natur-
forschender Freunde zu Berlin in den ersten 200 Jahren des Be-
stehens der Gesellschaft 1773-1972. - Sitzungsber. Ges. Naturf.
Freunde Berlin N. F. 13: 59-157.
Jahn, Ilse 1966: Carl Ludwig Willdenow und die Biologie seiner Zeit. ?
Klawitter, J. & Schaepe, Annemarie 1985: Gefahrdung und Riickgangs-
ursachen der Moose in Berlin (West) ? Eine Rote Liste. ? Verh.
Berl. Bot. Ver. 4: 101-120.
Kloden, K. F. 1832: Beitrage zur mineralogischen-geognostischen
Kenntnis der Mark Brandenburg. Fiinftes Stiick. ? Programm . . .
der Gewerbeschule, p. 1 -72.
Klose, H. 1957: Fiinfzig Jahre staatlicher Naturschutz. ? Giessen.
Konig, C. 1898: Willdenow. - Allgemeine Deutsche Biographie 43. -

Leipzig, p. 252-254.
Kostler, J. N. 1955: Professor Kurt Hueck. - Forstwiss. Cbl. 84: 387-
388.
Englera7- 1987 101
Kolkwitz, R. 1909a: Uber die Planktonproduktion der Gewasser,

erlautert an Oscillatoria Agardhii Gom. ? Landw. Jb. 30, Suppl.
5:449-472.
- 1909b: Die Rieselfelder und ihre Pflanzenkulturen. - Verh. Botan.
Ver. Prov. Brandenburg 51: 141-144.
- 1914: Uber die Ursachen der Planktonentwicklung im Lietzensee. ?
Ber. Deutsch. Bot. Ges. 32: 369-666.
- & Marsson, M. 1902: Grundsatze fiir die Beurteilung des Wassers
nach seiner Flora und Fauna. ? Mitt. Kgl. Priifungsanstalt Wasser-
versorgung und Abwasserbeseitigung 1: 33-72.
- 1908: Okologie der pflanzlichen Saprobien. ? Ber. Deutsch. Bot.
Ges. 26a: 505-519.
- 1909: Okologie der tierischen Saprobien. ? Intern. Rev. Gesamt.
Hydrobiol. 2: 126-152.
KoUektiv der Mitglieder der Fakultat 1961: Die ForstwirtschaftUche
Fakultat Eberswalde. - Wiss. Z. Humboldt-Univ. BerUn, JubUaums-
jahrg. 9(1959/60): 257-319.
Krausch, H.-D. 1977: Das Wirken von Johann GottUeb Gleditsch
auf dem Gebiet der Landeskultur. - Gleditschia 5: 5-35.
Leps, G. 1980: ProblemgeschichtUch-philosophische Analyse der
aquatisch-okologischen Wissenschaftszweige unter besonderer Be-
rucksichtigung des Lebenswerks von Karl August Mobius (1825-
1908) und August Thienemann (1882-1960). - Diss. BerUn.
- 1985: Zur Geschichte der Okologie in BerUn. - Wiss. Z. Humboldt -
Univ. BerUn, Math.-Nat. R. 34: 342-359.
- & Burmeister, K.-J. 1987: Richard Kolkwitz (1873-1956) zum
Gedenken. - Biol. Rundschau 25 (3).
Leschber, R. 1986: Das Saprobiensystem - Frucht der Zusammenarbeit
zweier Naturforscher. Eine Wiirdigung der gemeinsamen Arbeiten
von Richard Kolkwitz und MaximiUan Marsson. ? WaBoLu-Hefte
1: 51-64.
Mayr, E. 1982: The growth of biological thought. - Cambridge, Mass.
Meyer, D. E. 1953: Biographisches und BibUographisches iiber Christian
Conrad Sprengel. ? Mitt. Bot. Garten Mus. BerUn-Dahlem 1:
118-125.
- 1967: Goethes botanische Arbeit in Beziehung zu Christian Konrad
Sprengel (1750-1816) und Kurt Sprengel (1766-1833) auf Grund
neuer Nachforschungen in Brief en und Tagebuchern. - Ber. Deutsch.
Bot. Ges. 80:209-217.
Meyer-Abich, A. 1967: Alexander von Humboldt. In Selbstzeugnissen
und Bilddokumenten ? Reinbek.
Meyer-Waarden, P. F. 1970: Aus der deutschen Fischerei, Geschichte

einer Fischereiorganisation. ? Berlin.
Mildbraed, J. 1954: Eberhard Ulbrich. - Willdenowia 1: 154-174.
Nachtsheim, K. 1963: Max Hartmann f. Leben und Werk (1876-1962).-
Sitzungsber. Ges. Naturf. Freunde Berlin N. F. 3: 14-20.
Natho, G. 1973: Johann Gottlieb Gleditsch (1714-1786). - Gleditschia
1: 7-15.
? 1975: Das Schriftenverzeichnis von Johann Gottlieb Gleditsch. ?
Gleditschia 3: 5-27.
Naumann, E. 1961: 60 Jahre Institut fiir Wasser-, Boden- und Luft-
hygiene. ? Schriftenr. Ver. Wasser-, Boden-, Lufthygiene 18.
Ruske, W. 1979: Aufieruniversitare technisch-naturwissenschaftliche
Forschungsanstalten in Berlin bis 1945. - In: R. Riirup (ed.),
Wissenschaft 1. - Berlin, p. 231-263.
und Gesellschaft
Scamoni, A. Uber
1955: den gegenwartigen Stand der forstlichen
Vegetationskunde. ? Sitzungsber. Deutsch. Akad. Landwirtschaft -
wiss. Berlin 4 (6): 1 -22.
Schaperclaus, W. 1957: Ein Leben fiir die Fischerei. Prof. Dr. H. H.
Wundsch zum 70. Geburtstag. - Z. Fischer. Hilfswiss. N. F. 1-7:
3-37.
Schlumberger, O. 1947: Otto Appels Verdienste um den deutschen
Pflanzenschutz. - Festschrift Otto Appel. ? Berlin, p. 1-2.
Schmithiisen, J. 1957: Anfange und Ziele der Vegetationsgeographie. ?
Petermanns Geogr. Mitt. 101.
Schoenichen, W. 1954: Naturschutz, Heimatschutz. Ihre Begriindung
durch Ernst Rudorff, Hugo Conwentz und ihre Vorlaufer. ? Stutt?
gart.
Scholz, H. 1960: Die Veranderungen in der Ruderalflora Berlins. Ein
Beitrag zur jiingsten Florengeschichte. ? Willdenowia 2: 379-397.
Schweinfurth, G. 1861/62: Versuch einer Vegetationsskizze der Um-
gegend von Strausberg und des Blumenthals bei Berlin. Nebst einer
phytotopographischen Karte. ? Verhandl. Bot. Ver. Prov. Branden?
burg 3/4: 91-126.
Stafleu, F. A. 1981: Engler und seine Zeit. - Bot. Jb. Syst. 102:
21-38.
Stocker, O. 1979: Okologie als existentiales Problem im Viererschema
der biologischen Wissenschaften. ?Flora 168: 13-52.
Sukopp, H. 1969: Der Einflufi des Menschen auf die Vegetation. ?
Vegetatio 17: 360-371.
? (ed.) 1977: Interdisziplinare Arbeitsgruppe (Projektgruppe) Okologie
Englera7 - 1987 103
und Umweltforschung 1972-1976. - Z. Tech. Univ. BerUn, TUB,

9:278-322.
? 1983: 10 Jahre Institut fur Okologie. - Berlin.
? Dapper, H., Zimmermann-Jaeger, S., De Santo-Virzo, A. & Born-
kamm, R., 1971: Beitrage zur Okologie von Chenopodium botrys L. ?
Verhandl. Bot. Ver. Prov. Brandenburg 108: 3-74.
? & Elvers, H. (ed.) 1982: Rote Listen der gefahrdeten Pflanzen und
Tiere in BerUn (West). Ergebnisse des CoUoquiums iiber Riickgang,
Gefahrdung und Schutz der Flora und Fauna in Berlin (West) vom 4.
bis 6. 6. 1980. - Landschaftsentw. u. Umweltforsch. 11: 1-384.
? & Schneider, C. 1979: Zur Geschichte des Instituts fiir Okologie und
der okologischen Wissenschaften in BerUn. ? In: R. Rurup (ed.),
-
Wissenschaft und Gesellschaft 2 Berlin, p.: 239 -246.
? -1981: Zur Geschichte der okologischen Wissenschaften in Berlin.
-Verh. Ges. Okol. 9: 11-19.
Tembrock, G. 1959: Zur Geschichte der Zoologie in Berlin. - Wiss.
Z. Humboldt-Univ. Berlin, Math.-Nat. R. 8: 185-196.
Timler, F. K. & Zepernick, B. 1978: Der BerUner Botanische Garten.
Seine 300jahrige Geschichte vom Hof- und Kiichengarten des
Groften Kurfursten zur wissenschaftUchen Forschungsstatte. ?
Berlin. Forum 11 (7).
Trepl, L. 1987: Geschichte der Okologie. - Frankfurt a. M.
Wahnschaffe, F., Graebner, P. & Dahl, F. 1907: Der Grunewald bei
BerUn. Seine Geologie, Flora und Fauna. ? Jena.
? & Hanstein, R. v. 1912: Der Grunewald bei BerUn. Seine Geologie,
Flora und Fauna. 2. ed. ? Jena.
Waldenburg, I. 1935: Die botanischen Erforschung Brandenburgs. ?
Verhandl. Bot. Ver. Prov. Brandenburg 75: 177-238.
Wichler, G. 1936: Kolreuter, Sprengel, Darwin und die moderne Bliiten-
biologie. - Sitzungsber. Ges. Naturf. Freunde BerUn 1935: 305-341.
WiUdenow, C. L. 1792: Grundrifi der Krauterkunde zu Vorlesungen
entworfen. ? BerUn., 5. ed. 1810.

Prof. Dr. Herbert Sukopp, Technische Universitat Berlin, Fachbereich 14,
Institut fur Okologie, Schmidt-Ott-StraBe 1, D-1000 Berlin 41.
Englera 7-1987 105
Claus Schnarrenberger
Botany at the Kaiser Wilhelm Institutes
History of Dahlem as a scientific center
Beside the old University of Berlin, which developed in the center of

the city, another part of Berlin, Dahlem, achieved great significance in
several respects for science at the beginning of this century. Not the
last to profit from this development was also botany.
Today, Dahlem presents itself as one of the best residential districts
of Berlin, with many splendid villas from the first half of the century.
Here, on the edge of Steglitz toward Dahlem, lie the Botanical Garden
and the Botanical Museum. Here the world-renowned Kaiser Wilhelm
Institutes were established, which today have their inheritance as Max
Planck Institutes, in newer buildings and other places. Here lie the
Institutes for Agriculture and for Horticulture. Here the Free University
of Berlin was established, after the Second World War. And not last,
an Institute for Gene-Biological Research was recently established
here by the Schering Company and the Senate of Berlin.
The history of Dahlem has been wonderfully collected and presented
also with respect to scientific history, in Michael E ngel's "Geschichte
Dahlems" (1984). The story of the early settlement reaches back into
the thirteenth century. For several centuries, Dahlem was a small village
with an estate of relatively large area, and a church, the St. Anne's.
This church was founded in the fourteenth century. From 1832 to
1852, it was the location of the second station of an optical telegraph
line, with 57 stations from Berlin to Ehrenbreitstein (Koblenz), until
the transmission of news was taken over by an electromagnetic proce?
dure. Today, the cemetery around St. Anne's Church has been consider-
ably extended. Numerous important scientists lie buried there, so that
a visit already reminds one of the earlier significance of Dahlem.
The second to the last owner of the estate of Dahlem was, by the way,
the lawyer Friedrich von Beyme (1765-1838). He was an intimate

friend of Friedrich WUhelm III., and a member of his cabinet. From
1808, he was even Minister of Justice with the title Lord High Chancellor
(Reichskanzler). Among other things, Beyme was concerned with much
preliminary work for the founding of the Friedrich WUhelm University,
although the actual foundation in 1810 was not his job, but became
the responsibUity of WUhelm von Humboldt. After Beyme's death,
the estate was transferred to his daughter Charlotte, who, however,
sold it in 1841 to the Prussian crown land treasury for 200,000 thalers.
At that time, Dahlem was still a small village with a completely rural
character. It was stUl surrounded by fields and by the then much larger
Grunewald. From 1846, it was leased out, together with the Domain
StegUtz. The seat of both estates became the Domain Dahlem, which,
with some surrounding buildings, is still in existence today.
The contemporary development of Dahlem began towards the end
of the last century. In 1873, Dahlem obtained a better connection to
Berlin, with the construction of the Berlin -Potsdam-Magdeburg rail-
road. In 1889, a paved road was built from the hunting lodge Grune?
wald to Steglitz. This is the present day Konigin-Luise-Strasse and
Grunewaldstrasse. The hunting lodge Grunewald was often the starting
point for hunting parties of the royal family. The constructional
development of Dahlem was nevertheless considerably delayed in order
to make greater pro fits from the rising prices of house lots. FinaUy, in
1898, the construction of the Botanical Garden was begun, and the first
streets were projected: Altensteinstrasse, EngleraUee, Podbielskiallee,
RheinbabenaUee, Habelschwerdter Allee, Thielallee, CecilienaUee
(today, PacelliaUee). FinaUy in 1901 building began.
At the beginning of this century, from 1873 to 1911, in the Ministry
for Agriculture, Domains, and Forests, the Ministry Director, Professor
Hugo Thiel (1839-1918) was responsible for the partition of the
Royal Domain in Dahlem. Thiel himself was a leading agricultural
scientist. He was akeady authoritatively involved in the planning of
the Agricultural College in 1881, and was planning at the turn of the
century to move this CoUege to Dahlem. This first took place, however,
between 1921 and 1924. Today, the fields on both sides of Lentzeallee
still bear witness of this event. Still, Thiel managed to move the Royal
Horticulturists School from Potsdam-WUdpark to Dahlem, at the pre?
sent address, Konigin-Luise-Strasse 22. The partition of the Domain
Englera 7 - 1987 107
was carried out by a 5-membered committee under the chairmanship

of Thiel. He first commissioned the architect W. Kyllmann, and in
1917 the architect W. Schweitzer, to prepare a plan for building. To
a not inconsiderable extend, the ideas of the city-planner H. Jansen,
entered the picture. In addition to private houses, these plans included
the construction of several public buildings: the Materialpriifungs-
anstalt, the Astronomisches Recheninstitut (today, the office of the
President of the Free University of Berlin), the Museum of Ethno-
logy, the Secret State Archive, schools, churches and scientific insti?
tutes, the School for Horticulturists, the Botanical Garden, the old
Institute of Plant Physiology, and the Kaiser Wilhelm Institutes. For
employees, officials, and scientists, comfortable rental apartments
were built in the southern part of Dahlem, in the neighborhood of Habel-
schwerdter Ailee. In fact, it was possible to win the interest for villas
in Dahlem of the wealthy: architects, artists, scientists, directors of
industry, merchants, bankers, physicians. This situation eventually
meant that practically every old villa in Dahlem can be associated with
an important personality. Not seldom, important architects have made
these villas a witness to their ability.
The picture of Dahlem as a scientific center was finally strongly
influenced by a man, Friedrich A11 h o f f (1839-1908), who apparently
had nothing to do with the committee for the partition of the Domain
Dahlem. Althoff, a lawyer, was already importantly involved in the
founding of the University of Strassburg in 1872. In 1882, he was
appointed to the Prussian Ministry of Culture, and obtained in a short
time the responsibilities for the Universities (construction excepted),
technical colleges, other scientific establishments, libraries, and the
higher schools. Because of his abilities, he soon became the dominating
personality in the Ministry of Culture. Among many other great achieve-
ments of this man, belong his plans to create in Dahlem a science center
outside of the city center of Berlin. These plans matured only slowly,
and undoubtedly required very much persuasion of other personalities,
offices, ministries, the Kaiser, etc, in order to be realized. Originally,
Althoff seems even to have had the idea of moving to Dahlem the
entire University, or at least all of the scientific university institutes
including anatomy and other medical institutes. This truly grand idea,
however, encountered strong reservations from all sides, above all,
also from the Ministry of Finance.
These ideas, therefore, had to be dropped. Some of Althoff's

notions could nevertheless be reaUzed. In 1897, it was established by
law that the Botanical Garden -and the Botanical Museum (Konigin-
Luise-Strasse/corner of Altensteinstrasse) should be transferred to
Dahlem, and that a new Institute of Pharmacy should be built for the
University (Konigin-Luise-Strasse 2-4). These projects were completed
successively in the following years. That the Institute of Plant Physio?
logy arose on the other side of the street from the Botanical Garden
(Konigin-Luise-Strasse 1-3), was not the result of long planning
and deUberation, but the wish of G. Haberlandt, appointed in
1910. He insisted that the original Botanical Institute (later Plant
Physiological Institute) of his predecessor, S. Schwendener, should
be moved from the center of Berlin to the neighborhood of the Bo-
tanial Garden.
In conjunction the expansion of the Imperial Health Office
with
(founded 1876), a Biological Division for Agriculture and Forestry was
estabUshed in 1898, to investigate the then current problems of plant
protection. In 1910, this Division obtained an experimental field of
10 hectares area at Konigin-Luise-Strasse 17/19 (Government Experi?
mental Garden). The spatical separation of the Biological Division and
the experimental field soon made necessary a separate service and
laboratory building, which was erected in 1902-1905. From this arose,
in 1905, the Imperial Biological Research Centre for Agriculture and
Forestry.
The Kaiser WUhelm Institutes
The founding of the Kaiser Wilhelm Society in the year 1911 is

certainly one of the most briUiant scientific-poktical achievements
in the German-speaking countries, if one measures this by the signif-
icance of the scientific activities of this society and of its successor,
the Max Planck Society. The roots of this idea go back to Althoff,
who, in his Ufetime, always elicited discussions and expert opinions
regarding the development of scientific institutions in Prussia. The
professors at the University of Berlin were already then complaining
about the lack of space for the increasing numbers of students. In various
expert opinions, it was suggested that pure research institutes should
Englera 7-1987 109
be established. It was obviously, however, a great problem to reconcile

the often conflicting opinions, to obtain the agreement of different
ministries, to ensure the financing, either from the state or from private
or from mixed sources, and finally, also, to win the good will of Kaiser
Wilhelm II. On the occasion of the approaching centennial celebration
of the founding of the University, an additional reason appeared
to commence planning a new conception of the research facilities.
Unfortunately, Althoff retired in 1907 and died one year later,
without leaving behind detailed plans for the establishment of pure
research institutes.
After Althoff's death, it was F. Schmidt-Ott who, together
with the theologian A. H a r n a c k, the chemist E. Fischer, the
physiologist M. Rub ner, the bacteriologist A. von Wassermann,
and many others, within a short period of time articulated ideas for the
founding of the Kaiser Wilhelm Society. The basis for action became
eventually a "confidential memorandum" of A. von Harnack, which
was laid before the Kaiser on Nov. 21, 1909. In this memorandum, it
was emphasized, among other things, that research results were of great
value for national prestige, and that a leading role in the natural sciences
had great political, national and economic value. This was done toassure
the agreement of the Kaiser. In sum, a "Royal Prussian Society for the
Advancement of the Sciences" was proposed, with a seat in Dahlem. It
should consist of founders and scholars.
The meeting for the foundation of the Kaiser Wilhelm Society
took place on Jan. 11, 1911. A. Harnack became the first president;
F. Krupp and the banker L. Delbriick became the vice-presidents.
The first institutes were the Kaiser Wilhelm Institute for
founded
Chemistry, and the Institute for Physical Chemistry and Electroche-
mistry. Among others, the Royal Chemical Society, with E. Fischer
as president, was decidedly involved, both conceptually and finan-
cially, in the founding of the Institute for Chemistry. The first members
of the Institute were E. Beckmann, R. Willstatter, and O.
H a h n. The Institute for Physical Chemistry and Electrochemistry was
aided by the initiative and a grant from L. Koppel (owner of the
Berlin banking house, Koppel and Co.). F. Haber became the first
director.
In addition to the earlier suggestions for research institutes for
chemistry and physics, the Greifswald er paleontologist, O. Jaeckel,
particularly, suggested a research institute for pure biology. He indi?

cated that revolutionary results had been achieved precisely in biology,
and that further research of basic principles promised to become
particularly successful, although practical results could not be promised
immediately. At first, however, no uniform and applicable conception
for a Kaiser WUhelm Institute for Biology could be found. In a modern
conception, genetics as well as the physiology and development pro-
cesses should be taken into account. Finally it was agreed, first to find
outstanding scientists, and then to build these men institutes, for which
a number of grants from banks were on hand. It was cause of great
regret that the Wiirzburg physiologist, Th. Boveri, withdrew his original
acceptance, on the grounds of poor health. Instead, the botanist C.
C o r r e n s was appointed, who carried out the building plans of Boveri
for the Kaiser WUhelm Institute for Biology. Further appointments
included the zoologist Hans Spemann, the geneticist R. Gold-
schmid t, the protozoologist M. Hartmann, and the physiologist O.
Warburg. In 1915, Correns, Spemann and Hartmann were the
first to begin their work.
Already before the founding of the Kaiser Wilhelm Society, BerUn
maybe regarded as fortunate that some of these personalities seized the
initiative in the founding of new scientific institutes. Many actions
certainly went outside of official channels; and personal interests as
well as opportunity for economic gain could certainly not be ruled out.
The final touchstone, however, was mostly the good faith which
justified the various plans. The successes of later years provided the
final justification for the founding of the Kaiser Wilhem Society.
Willstatter
Richard Martin Willstatter (1872-1942), born in Karlsruhe,

came from an old and tradition-rich Jewish family, which included
rabbis, scholars and respected merchants. In 1883, when his father went
to the U.S.A. for several years, his mother moved with him to Niirn-
berg, to be in the neighborhood of her father. After preparatory school,
Willstatter attended the Technical College of Munich, where he
studied chemistry, which had already captivated his interest. He ob-
tained his Ph. D. with Prof. A. Einhorn at the institute of the great
Richard Willstatter
chemist A. von Baeyer, with a dissertation on the structural elucidation

of cocaine. Baeyer, whom he honored highly, and who recognized him
as an excellent scientist, then urged him to qualify only two years
later as a university lecturer, with his research on "Investigations of
the Tropin Group". At the age of 24, Willstatter thus became
the youngest university lecturer. In 1902, he received a position as
extraordinary professor in Munich until he finally was appointed
professor at the laboratory for general and analytical chemistry of the
Federal Technical College of Ziirich. In 1912, he followed an invitation
to join the Kaiser Wilhelm Institute for Chemistry in Berlin, left it
again, however, in 1916 to become the successor of his teacher A.
von Baeyer in Munich. In 1924, Willstatter resigned his professor-
ship, though he continued to direct, from Munich-Bogenhausen, the
studies of his coworker, Margarete Rohdewald. She performed further
studies on enzymes in the laboratory of Wieland, Willst att er's
successor at the Chemical Institute. Willstatter had to emigrate in
1939 and settled in Moralto-Locarno in Switzerland. There he composed
his autobiography, among other things, and died in 1942.
Willst att er's autobiography was first edited for publication in
1949, by his student A. Stoll. It describes very impressively and
completely the heights and depths of his life as a research scientist.
This biography is in fact a piece of scientific history par excellence,
and already for this reason an exceptionally worthwhile bit of reading
matter.
Willstatter was essentially a chemist. His teacher, A. von Baeyer,
stimulated his interest in natural products. During his time in Ziirich,
Willstatter investigated, among other things, the green pigments
of plants. He published this work in book form in 1913, together
with one of his best and most trustworthy students, A. Stoll. With
the methods of alkaline and acid hydrolysis available at the time, and
with the use of various solvents, he succeeded in identifying Mg2+ as a
component of chlorophyll. He separated chlorophylls a and b, carotene,
and xanthophylls qualitatively and quantitatively, and identified the
phytol residue of chlorophyll. He showed that these plant pigments
were the same in all plants, including green algae and that they occurred
in the same combinations. Only in brown algae did he find a substan-
tially different composition. By further chemical degradation in acid
and alkaline media, he finally arrived at well defined chlorophyll
Englera7-1987 113
derivatives such as the phaeophytins, phaeophorbides, phytochlorines,

phytorhodins, phyllins, porphyrins, and various derivatives of pyrrol.
Enormously impressive are also his studies at that same time of the
enzyme chlorophyllase, present in the entire plant kingdom (including
algae). This enzyme catalyzes the hydrolytic splitting (or the transesteri-
fication in an alcohol solution) of the phytol moiety of chlorophyll,
leaving free chlorophylUde. In contrast to the chlorophylls, the free
chlorophyllide can easily be crystalUzed. The action of chlorophyllase
was thus responsible for the occurrence of crystals in microscopic leaf
sections, as described by the chemist and musician J. Borodin already in
1882. Such endogenous activities as that of chlorphyllase on chlorophyll
could accordingly also be used intentionally, in order to obtain well
defined degradation products by the use of enzymes.
Already very early, Willstatter recognized that it was necessary
to use a tissue in good condition in order to isolate substances from it
in unaltered form. In this connection, he was the first to develop the
process of lyophylization, that is, the vacuum drying of frozen tissues.
Because of the use of fresh plant material, W illst at t er was credited
with bringing plants into the chemical laboratory. This principle was
regarded as quite essential by Willstatter and Stoll in the che-
mistry of natural products, and was later often used in other investi-
gations (StoU 1933). The structural formula for chlorophyll developed
at that time by Willstatter and Stoll naturaUy still contained errors.
This was related among other things to the fact that at that time ring
systems with more than 8 components were regarded as unstable, or
not capable of existence, according to the strain theory of Baeyer (StoU
1933).
W illst atter's studies of chlorophyll from his time in Zurich had
already at that time made a great impression, so that he received the
Nobel Prize in 1915 for his "Investigations of pigments in plant king?
dom, above all, of chlorophyll". This was the first award of a Nobel
Prize to a member of the stUl very young Kaiser WUhelm Society.
At the time, Willstatter had already been in BerUn for three
years, he applied himself among other things to the principles of
photosynthesis in green plants, and to the analysis of the anthocya-
nins. With their work on photosynthesis, Willstatter and Stoll
once again made valuable contributions, which they have collected in
the book "Investigations of the assimilation of carbonic acid" (1918).
The first two sections of this book are a great pleasure to read, also
for present-day plant physiologists, because of the elegance of the
experiments therein described, and their logical development.
First, Willstatter and Stoll established the constancy of the
chlorophyll content of illuminated leaves. Thereby, they repudiated
the concept frequently expressed at the time, that the chlorophyll
was degraded or transformed during CO2 assimilation. Then they
pointed to the five limiting factors for photosynthesis, determined by
Blackmanin 1905:
1. the available carbon dioxide concentration

2. the available amount of water
3. the intensity of the available light energy
4. the amount of chlorophyll present
5. the temperature of the chloroplasts
The yield of photosynthesis is determined by that factor which is

present in limiting amount. Since, up til that time, no quantitative pro?
cedure for chlorophyll determination was available except their own,
Willstatter and Stoll made, with their new analytic procedures,
quite fundamental discoveries regarding the quantitative contribution
of chlorophyll to the yield of photosynthesis. First, they determined
that under optimal condition, the yield of photosynthesis for normal
green leaves lay within 5 and 12 g CO2 per hour per gram of chlorophyll
(defined as the assimilation number). They observed deviations from
these values in younger leaves, which showed a somewhat higher
assimilation number than fully differentiated leaves. Tissues with little
chlorophyll showed quite substantial deviations from the above-men?
tioned values. During the yellowing in the fall, they observed in most
leaves a parallel decrease of chlorophylls a and b, whereas the yellow
pigments changed little. Depending on whether the chlorophyll de-
clined rapidly or slowly in a particular plant, the assimilation number
could then either remain the same (e.g. poplar, plane tree, strawberry)
or decline substantially (e.g. Robinia, lime tree). The investigations
of yellow -leaved varieties with little chlorophyll (aurea mutants with
about 10 to 50 % of the chlorophyll content of normally green plants)
showed at high light intensity, assimilation number 10 to 20 times
higher than those of the green (wild-type) plants, so that the abso-
lute rate of assimilation per leaf of these mutants was almost compa-
Englera7 - 1987 115
rable with that of the green plants. Only in yellow-leaved varieties with
quite low chlorophyll content, did the absolute assimilation rate be?
come substantiaUy lower. In weak Ught, on the other hand, the assimi?
lation rate for yellow varieties was much lower than for green plants.
To explain these results, it was clear that not only the Ught-absorbing
chlorophyll but also enzymatic factors contribute to the photosyn-
thetic yield. In green plants in strong Ught, an excess of chlorophyll
is present, and the enzymatic factors are limiting. In yellow-leaved
varieties, chlorophyll itself becomes limiting only at very low illu-
mination levels. This phenomenon is particularly apparent in the
yellow-leaved varieties in weak Ught. Whereas the chlorophyll content
of green plants is stiU present in sufficient amount (in weak Ught), this
becomes the Umiting factor to a much greater extent in yellow-leaved
varieties. Also on iUumination of etiolated seedUngs, Willstatter and
Stoll observed quite high assimilation numbers, as in the leaves of
yellow varieties. They interpreted this result thus, that "absence of light
(= growth in darkness) inhibits the formation of chlorophyll, but not
the formation of the enzymatic factors necessary for the photosyn-
thetic yield".
They made a similarly brilUant analyses of the temperature depen-
dence (Qio) of photosynthesis in green and yellow plants, and pointed
to the Umiting duaUsm of chlorophyll content as well as enzymatic
factors for optimal photosynthetic yields. Their reflections about light
scattering and absorption in green and yeUow leaves are downright
astounding. This is a problem which has only been solved in a manner
approaching satisfaction by plant physiologists today. For the photosyn?
thetic quotient CO2/O2, they determined a value of one, with quite
smaU deviations. For succulent plants (Phyllocactus and Opuntia),
however, they determined values between 0.44 and 0.89. Here, as in
further analyses, one observes the critical understanding of Willstatter
and Stoll, not only for chemistry but also for the biology of plants.
In aU this work, the accuracy of their analyses is striking, in view of
the methodology then available.
The concept of the mechanism of CO2 assimilation adopted by
Willstatter and Stoll was, like that of their contemporaries, cha-
racterized by the assumption that chlorophyll reacts directly with CO2,
which is possibly converted to formaldehyde by way of several interme-
diates, and eventuaUy forms glucose. Willstatter and Stoll were

Englera 7 - 1987 117
thus intimately concerned with measurements of C02-binding by

chlorophyll, under the most varied conditions. This work led to the
conclusion, however, that no intermediates of CO2 assimilation could
be detected. As we know today, the notion that the oxygen formed
during CO2 assimilation arises from water and not from CO2, was first
developed by C. B. van Niel in 1924. Not before 1949 did Calvin and
his coworkers formulate the Calvin cycle.
The second large complex problem that Willstatter solved

during his sojourn in Dahlem, was the analysis of another class of
plant pigments, the anthocyanins. He was the first who succeeded
in isolating and crystallizing these pigments from plants, in converting
them into their components (aglycone, sugar, and methanol), and in
determining their chemical structure on the basis of well-defined
decomposition steps. Particularly interesting is how he was able, by
comparison of anthocyanins from various plants (bachelor buttons,
roses, lingonberries, pelargonia, larkspur, grapes, blueberries, hollyhocks,
peonies, saleria, winter- and summer-asters, cherries, and wild plums),
to relate the great variety of colors to only three building stones (antho-
cyanidines). These compounds differ only by the OH-groups in the
3'-, 4'-, and 5'-positions (pelargonidin, cyanidin, and delphinidin),
and additionally by a sugar on the OH-group in the 3- and 7-positions,
or by methoxy groups on one or several OH-groups of the molecule.
The breadth of color variability of each anthocyanin or anthocyanidin
was competently demonstrated by variation of solvent, pH, or tem-
perature. Not only for him, but also for many visitors of his rather
remote institute, it was a joy to admire the splendidly colored flowers
grown in the neighborhood of his laboratory.
In his work on anthocyanins, as in his work on chlorophyll, Will?
statter was particularly interested in the constitution of the pigment as
it was present in the plant; for the investigations made at that time in
his and other laboratories, on dried plant material often resulted in
degradation products from which the compounds present in the living
plant could not be reconstituted.
Willstatter pursued many other aspects of chemistry than the
chlorophylls and the anthocyanins. Thus, he succeeded in the synthesis
of several tropene-alkaloids, of cyclotetraene, quinones and quinonei-
mines. He was the first to succeed in the saccharification of wood by
concentrated hydrochloric acid, a procedure which was later further
developed by F. Bergius for the technical preparation of glucose. For

his equally famous scientific friend, Fritz H ab e r, at the Kaiser Wilhelm
Institute for Physical Chemistry and Electrochemistry, Willstatter
developed filters containing urotropine for gas masks. Interesting for
biochemists is also his work on the purification of enzymes (peroxi-
dase, lipase, trypsin, amylase) by adsorption procedures (among others,
with aluminum hydroxide) and by chromatographic procedures. This
work was done later, however, in Munich.
In his autobiography (1949), Willstatter said that work and

antisemitism had determined his Ufe. That scientific work was a ful-
fUlment for him, is not in question. That he was able, however, to
achieve so much productivity and originaUty, is certainly due to his
very special talents. It is impressive, besides, how he took care of his
many coworkers, who often later became famous scientists themselves.
Willstatter's great scientific achievements were the reason for
calling him from Zurich to the Kaiser Wilhelm Institute for Chemistry
in BerUn. E. Fischer, Germany's other great chemist (beside A. von
Baeyer) traveled speciaUy to Zurich in order to induce the at first
reluctant Willstatter to come to Berlin. And Emil Fischer had
success. The fact that Willstatter received at the same time the
position which had been occupied by the famous chemist, van't
Hoff at the University of Berlin, filled him with additional pride.
His colleagues at the Kaiser Wilhelm Institute for Chemistry, the Otto
Hahn Building of today (ThielaUee 63) were at that time Otto H a h n
and Lise M e i t n e r on the ground floor, and E. Beckmann on the
floor above him. At other Kaiser Wilhelm Institutes in Dahlem were
Correns, Spemann, Goldschmidt, Neuberg, Wassermann,
and above aU, Fritz Haber, with whom Willstatter developed a
very intimate friendship. Willstatter's and Haber's villas lay on
opposite sides of Hittorfstrasse (Faradayweg 8 and 10). Willstatter's
decision to move to Munich after four years in Berlin was certainly due
to his veneration for his teacher, A. von Baeyer, whose successor he
became. Baeyer himself had proposed Willstatter's name to the
Munich faculty, and obtained unanimous approval.
Willstatter's private life was overshadowed several times by
disappointments and blows of fate, which he was mostly able to over-
come with admirable strength. In 1903, he decided to found a family
with Sophie Leser; but his marriage was of tragically short duration.
Englera 7-1987 119
In 1908, his wife suffered an attack of appendicitis and died. Sponta-

neously, he dedicated himself to the education of his two children,
Ludwig and Margarete. Again, tragedy struck, when his ten-year-
old son died quite unexpectedly, in 1915.
The second factor determining his life, antisemitism, was experienced
by Willstatter already in his childhood in Karlsruhe, and especially
as a schoolboy, during his days in Niirnberg. Munich presented him
finally with the great test of his Jewish belief. It had already hit him
hard when A. von Baeyer preferred to recommend his son-in-law
Piloty 1899 for a university appointment, rather than Willstatter.
Baeyer, himself a Jew who had converted to Christianity, attempted
at another time to get W i 11 s t a 11 e r to do likewise, since there might
otherwise be considerable difficulties with his career. Willstatter
was probably simply a believing Jew, deeply anchored in the tradition
of his religion. For him, there could therefore be no discussion of this
point. His time in Zurich was not burdened in this respect. Berlin was
also less problematical, since he was only one of many Jewish scientists
there. In any case, the problem of antisemitism was less evident in Berlin
than in Munich, and at the Kaiser Wilhelm Institutes, it did not exist for
him. But with his return to Munich, it again became very acute, when
on signing his appointment, King Ludwig III warned the responsible
minister: "But that is the last time I will agree to signing for a Jew".
In Munich, antisemitism was quite general in the population, and it
was also quite clear, though hidden, among his colleagues on the
faculty, and Willstatter felt it repeatedly. It reached a climax
first in 1924, when the faculty three times in succession rejected
highly qualified Jewish scientists for faculty positions. After the third
time, Willstatter spontaneously handed in his resignation, to the
horror and regret of many colleagues and students. His emigration to
Switzerland can probably be classified as the more sensible behavior,
in view of the antisemitic attitude which became dominant in 1939
under the Hitler regime.
Stoll
An intimate coworker of Willstatter was Arthur Sto 11 (1886-

1971). He was born in Schinznach (Switzerland) and studied chemistry
at the Federal Technical College in Zurich. Already in 1909, before

completing his studies, he had accepted the invitation of R. Will?
statter to work with him on.chlorophyll. He obtained his Ph. D.
degree in 1911, with a thesis "On chlorophyllase and chlorophyllide".
This work led to the first structural clarification of the chlorophylls.
In 1912, when Willstatter went to the Kaiser Wilhelm Institute
for Chemistry in
Berlin-Dahlem, Stoll followed Willstatter
and became his head assistant. In Berlin, Stoll continued his investi?
gations of the chlorophylls, and also devoted himself to the more
fundamental phenomena of photosynthesis and CO2 fixation. All
of these studies were pubUshed together with Willstatter in book
form (Willstatter & Stoll 1913, 1918), and have aUeady been described
in the chapter on Willstatter. When Willstatter was called to
Munich in 1916, Stoll followed, but accepted a position with the
Sandoz Company in Basel already in the following year. His respon-
sibility there was to build up a pharmaceutical section.
Like Willstatter, Stoll was primarily a chemist. What he accom-
plished, however, for botany, during the years with Willstatter,
places him side by side with the great botanists of his time, and justifies
honoring him here with his own chapter. Of course there are some
arguments about why he never appeared to be a botanist. It plays
a part, that it was difficult for such a young scientist as Stoll to
step out of the shade of such a brilliant personaUty as Willstatter.
Secondly, the field "Biochemistry of Plants", under which one would
classify Stoll's work, was simply not occupied sufficiently by the
botanists of the day, to provide discussion partners for Stoll. Thirdly,
one must take into account that Stoll never held a regular professor?
ship at an university, where he could build his own school. The fact
that Stoll and Willstatter achieved such great importance for
botany, although they were no botanists, can not lastly be ascribed
to their long engagement with the biochemistry of plants. Of course
there were, in Willstatter's laboratory, many other brilliant investi-
gators besides Stoll, scientists who worked on the chlorophylls or,
above all, on anthocyanines. Nevertheless, their engagement with
botany was rather brief, although they also became important in-
vestigators in other fields of biology or in the field of chemistry.
The great scientific potential of Stoll came to full flower with
the Sandoz Company. There he succeeded in isolating a number of al-
Englera 7- 1987 121
caloids from ergot (Claviceps purpurea) and of cardioactive glycosides

from sea onion (Scilla maritima), foxglove (Digitalis purpurea, lanata
and ferruginea), Strophanthus kombe and garlic (Allium sativum), all
these compounds being of great therapeutical values in medicine up
til today. Also worthy of mention is Stoll's work on the synthesis
of compounds of the tropane series. Some of these compounds are
important antispasmotics. In the case of the majority of materials
prepared, one could usually recognize a clearly established procedure:
isolation of the natural compound from the plant, degradation with
endogenous (seldom exogenous) enzymes, chemical degradation to
welldefined products, and finally the determination of structure (Stoll
1933). In this way, Stoll achieved for botany a decisive insight into
the chemistry and biology of the natural products of plants.
Stoll furthermore continued with his coworker E. Wiedemann
the work on chlorophyll begun with Willstatter. In 1935, he
succeeded in formulating the total structure of chlorophyll in almost
final form. In 1938, he showed that the chlorophylls, with the carot-
enoids, are bound to a cell fraction with protein and lipid (thylakoid).
He termed this chromoprotein "chloroplastin". But just at this time,
the botanists had begun to develop more fundamental ideas about the
anchoring of the chlorophylls in the chloroplast.
Because of his outstanding scientific achievements, Stoll received
numerous offers of university positions. He was honored by academies
and societies as well as universities. In 1955, he became President of
the International Union for Pure and Applied Chemistry. Also within
the Sandoz Company, he climbed high in the chain of command, as
the Director of the Company (1923), as a Vice President of the Ad-
ministrative Board (1935), as President of the Board of Directors of
Sandoz (1949-1956), and as President of the Administrative Board
(1964-1967). Stoll died in Dornach in 1971. The life and work of
Stoll have been summarized in exemplary fashion in a publication
honoring him on his 70th birthday.
Warburg
Otto Warburg (1883-1970) was one of the scientists working

longest in Berlin. He was born in Freiburg/Breisgau, and stemmed on
his father'sside from a large, north-German Jewish family, which

included numerous significant personaUties in the fields of science,
banking and business, and art. In addition to the biochemist Otto
Warburg, there was, incidentally, another Otto Warburg (1859-
1938), a botanist who, among other things, pubUshed a 3-volume
textbook of botany (Warburg 1913-1922). Otto Warburg's father,
Emil Warburg (1846-1931) was a well-known professor of physics,
first at the University of Freiburg, and from 1895 at the Friedrich
WUhelm University of Berlin, until he became Director, from 1906
tUl 1922, of the Federal Bureau of Standards in Berlin-Charlottenburg.
There he determined for the first time the quantum yield of photo-
chemical reactions. The Ufe and work of Emil Warburg were eval-
uated and honored in 1931 by his student, James Frank. War-
burg's mother, Elisabeth, born Gaertner, was from a south-German
family.
Otto Warburg began his study of chemistry in 1901 at the Uni?
versity of Freiburg, and transferred in 1903 to BerUn. In 1906 he ob-
tained his Ph. D. under Emil Fischer, a chemist whom he honored
highly, with a thesis on the synthesis of opticaUy active compounds
and the hydrolysis of animo acid esters by pancreatic enzymes. After
this, Warburg went to the University of Heidelberg to study medicine.
There, he worked in the laboratory of Ludolf von Krehl, an outstanding
physician and the author of the reference work "Pathological physio?
logy". Warburg obtained his Ph. D. degree in medicine in the year
1911, and remained in Heidelberg until 1914.
On the first of April, 1914, Warburg was named a member of the
Kaiser WUhelm Society, above all at the instigation of the geneticist
Theodor Boveri and of his earlier teacher, Emil Fischer. Warburg
thereby became a leader of a research section, with Ufe tenure, and
could devote himself completely to his research interests, without
teaching or administrative duties. His early career was only interrupted
once, when he volunteered for miUtary service in an elite cavalry
regiment in World War I, in which he also was wounded. Only a personal
appeal from the physicist Albert E inst ein, moved to this by War-
burg's mother, caused him to return to research seven months before
the end of the war. Warburg's laboratory was located at first on the
fourth floor of the Kaiser Wilhelm Institute for Biology at Boltzmann-
strasse 3 in Berlin-Dahlem. He could only hire 6-10 assistants, there,
Otto Warburg


exchange in respiration, fermentation, and photosynthesis, the optical

tests for measuring enzyme activity by following changes in the oxi-
dation-reduction state of the pyridine nucleotides, and the t issue -
slice method for determination of the respiration rate and other reac-
tions in normal and cancer tissue, were methods which were developed
by Warburg and have given valuable service to generations of scientists.
In addition to the work mentioned above, Warburg also investi-
gated several very basic problems of the energy metabolism of plants.
The introduction of the (Chlorella) as an object for metaboUc phys-
iological investigations (Warburg 1919) formed the basis of his botanical
research. Suspensions of similar algal cells were mostly much more
suitable than whole leaves for the quantitative photo chemistry of
photosynthesis.
Quite early, Warburg (1920) observed that 02 inhibits the photo?
synthesis of algae. This phenomenon has been caUed the Warburg
effect of photosynthesis. The cause of it was first recognized fourty
years later as the process of light respiration and of the glycolate
pathway (Wang & Waygood 1962, Tolbert 1963). In 1960, Warburg
and Krippahl (1960 a) first determined the conditions important for
glycolate formation, i.e. low CO2 and high O2 partial pressures. It
was more than ten years later that the key reaction of glycolate for?
mation, the ribulose-l,5-bisphosphate oxygenase reaction was identi-
fied (Bowes et al. 1971, Andrews et al. 1973).
In an early botanical study on the "Reduction of nitric acid in green
cells" (Warburg & Negelein, 1920), Warburg first made a quantitative
analysis of nitrate assimilation in algae. After optimizing the conditions
for nitrate assimilation, he first determined that the amount of ex-
creted ammonia was dependent on the amount of nitrate added.
Accompanying manometric measurements gave a dark respiratory
quotient CO2/O2 = 1,6. He explained this by pointing out that in
addition to normal respiration (CO2/O2 = 1), a part of the carbon
(carbohydrate) was oxidized to CO2 (extra CO2) by the oxygen of the
nitrate. On the basis of the equation
HNO3 + H20 + 2C = NH3 + 2C02
Warburg expected the formation of 2 moles of CO2 per mole of ex-
creted ammonia. Since he found only half of this amount, he postulated
that the missing ammonia was built into the cells. As a consequence, he
was able to show that nitrogen-poor cells incorporated more ammonia
Englera 7-1987 127
than nitrogen-rich cells. On the basis of their different sensitivity to

cyanide, he differentiated the chemical reactions of nitrate reduction and
ammonia formation from the reactions of CO2 uptake. That the
reduction of nitrate to ammonia occurs in several steps, he concluded
from the fact that, at low O2 concentrations, additional nitrous acid
(and only this compound) was excreted, and the nitrite formation, in
contrast to the ammonia formation, was not inhibited by HCN. He
attributed the toxic action of the lower O2 concentrations to a poi-
soning by nitrite. In evaluating this work, one must keep in mind that
the enzymes responsible for nitrate reduction to ammonia, nitrate and
nitrite reductase, were first clearly identified by Evans and Nason
(1953) and by Hageman, Cresswell and Hewitt (1962), respectively.
Finally, Warburg found only a small development of O2 in light if no
CO2 was added, but a twenty fold increase on addition of nitrate,
whereas, in the dark, comparatively little extra O2 was liberated for the
ammonia formation. Thus, nitrate may be regarded as the first so called
Hill reagent. On setting up the energy balance for the formation of
ammonia from nitrate according to the above equation, Warburg
argued that sufficient free energy would be supplied by the photo-
synthetic refixation of 2 CO2 (-230,000 cal/mol) to overcome the
positive free energy of the reduction of nitrate to ammonia (+ 162,000
cal/mol) in a dark reaction. ? The latter considerations may be taken to
show how little was known about such problems at that time. The time
was indeed too early for a more detailed modern interpretation.
Warburg was one of the first scientists to attempt an analysis of
photosynthesis as a photochemical process (Warburg & Negelein 1922,
1923). He came to the conclusion that the evolution of one molecule
of O2 required 4 quanta of light. In the fifties, these values became the
object of violent discussions and quarrels because of the difficulties
experienced in other laboratories in reproducing them.
For a summary of Warb urg's views on photosynthesis, we refer
to the didactically excellent presentation of W. Bladergroen (1959).
Since this work is mainly of historical interest, we will only point
out here, that Warburg regarded the primary problem in photosyn?
thesis as the identification of the one-quantum reaction that lay at
the heart of the system. Warb urg had learned the procedures for the
measurement of light intensity and absorption from his father, in whose
laboratory the necessary measuring devices were developed. And the
identification of an underlying one-quantum process was a cornerstone

of early photochemistry, which Warburg had learned from Nernst.
That the primary photochemical "step in photosynthesis led immediately
to O2 evolution from a CO2 derivative was a notion borrowed, perhaps
unconsciously, from Willstatter. Warburg failed to reaUze that
this might be an assumption which was incorrect.
In connection with his investigations of photosynthesis, Warburg

made a number of interesting observations. Thus, he was the first to
show that quinone, besides Fe^+, could be used as Hill reagent (War?
burg & Luttgens 1944). In addition, he showed that CO2 has a role in
the photosynthetic electron transport chain of the Hill reaction (War?
burg & Krippahl 1960b). It is known today that the binding site for
CO2 is on the herbicide-binding protein.
Warburg received many honors for his achievements. The greatest,
the Nobel prize for medicine and physiology, he received in 1931,
for the discovery of the catalytic role of the iron porphyrins in biolo?
gical oxidations. Already in 1927, he had been suggested for a Nobel
prize together with Fibiger, for his work on the metaboUsm of cancer
cells, but this honor was then given to Fibiger alone for his work on the
discovery of Spiroptera carcinoma (later discredited). In the year
1944, Warburg was once more suggested for the Nobel prize for his
work on the identification of flavins and nicotinamide as hydrogen
transferring agents in biological oxidations, but this prize was never
conferred, since German citizens were not, at that time, aUowed by
Hitler to accept the Nobel prize.
Warburg's scientific achievements must certainly be ascribed to

his enormous talent. He himself often emphasized that the best basis
for the career of a young scientist was personal contact with the great
investigators of the time, and he cited for himself: Emil Fischer,
his father, Emil Warburg, and Ludolf von Krehl. Of additional
importance in Warburg's case was the fact that Berlin before the
Second World War was a center without equal in the area of the natural
sciences, and the personal contacts between outstanding scientists
must have been especiaUy stimulating. Thus, Warburg had contact
already in his early years with personaUties such as Walter Nernst,
Jacobus Henricus van't Hoff, Theodor WUhelm Engelmann,
Max Planck, Albert Einst ein, Fritz Haber, and many others.
Englera 7 - 1987 129
Beyond this, he always had contact with the best scientists on his
own travels, or from visitors to Berlin. Occasional criticism that he was
neglecting the education of young scientists was answered with the
comment that three of his students, Meyerhof, Theorell, and Krebs,
had received the Nobel prize.
In his laboratory, Warburg mostly had technical assistants whom
he trained himself and who became famous because of their outstanding
work. These were, at first, E. Negelein, F. Kubowitz, E. Haas,
W. Christian, and W. Liittgens. After the Second World War, one
can particularly mention G. Krippah 1 and H.-S. Gewitz.ToWar-
b urg's academic coworkers belonged: H. Hartmann, A. Reid,
H. Theorell, E. G. Ball, E. St. French, J. N. Davidson, Th.
Bucher, and H. Tiedemann. From a botanical point of view,
the visits of Hans Gaffron (1925-1927), Dean Burk (1949-1950),
D. I. Arnon, E. St. French and B. Vennesland should be particularly
mentioned. Of these, Dean Burk made a special effort to spread know?
ledge of Warb urg's result s in the U.S.A.
For his entire life, Warburg can be described as a typical hermit.
He was very critical of his environment. In science, he made the greatest
demands on himself and on others. He was certainly in every respect
a fanatic for truth in pure science. Nevertheless, he was not always
free from prejudice, above all, when he met criticism of his own con?
clusions. From the middle of the fifties, this criticism became increas-
ingly strong, above all with regard to his work on photosynthesis,
and this led several times to quarrels with friends. This was apparently
caused by the fact that Warburg did not take into sufficient conside-
ration the results of other scientists, so that he was left mostly alone
with his outdated theories and his polemics.
There is much else of interest and importance to tell about War-
b u r g's life and work. Hans Krebs, who was a good friend, has written
a fine biography; and Nachmansohn (1979) has described Warburg
as a man and as a Jew. Warb urg was one of the very few Jews who
remained unmolested in the Third Reich.
After Warb urg's death, his laboratory received, from a botanical
point of view, a continuation in the Forschungsstelle Vennesland.
Birgit Vennesland, born in Kristiansand in Norway in 1913, studied
biochemistry at the University of Chicago and obtained her Ph. D. there
in 1938 under the sponsorship of M. Hanke. During the following
two years, she worked in the laboratory of B. Hastings at the Harvard

Medical School in Boston on the utilization of ^C for the study
of mammalian carbohydrate metaboUsm. During this work, it was
demonstrated that the carbon of HCO2 entered newly formed liver
glycogen when fasted rats were fed lactate or glucose. In the following
years at the University of Chicago, Vennesland studied a series
of carboxylating and decarboxylating reactions in non-photosynthetic
plant tissues. Part of the point here was a comparative study of such
reactions in animal and plant tissues. Her main achievement was the
demonstration, together with F. Westheimer, of the direct and stereo-
specific transer of hydrogen between cofactor and pyridine nucleotide
in a broad range of pyridine nucleotide dehydrogenase reactions (West?
heimer et al. 1951; Vennesland 1958).
After Warburg had demonstrated the stimulatory effect of CO2
on the HU1 reaction, Vennesland substantiated and extended his
findings. In the year 1961, she accepted Warburg's invitation to
come to Berlin for a year of research in his laboratory. This was followed
by a number of shorter visits in subsequent years, until she moved per-
manently to Berlin in 1968. Because of difficulties in Warburg's
laboratory, the Max Planck Society set up the Forschungsstelle Vennes?
land at Harnackstrasse 23, about the time of Warburg's death.
Vennesland's main interests were the isolation and characteri-
zation of the nitrate reductase of Chlorella vulgaris. Of special interest
in this connection was the inactivation of the enzyme by NADH and
HCN (Lorimer et al. 1974). As a possible source of HCN in the algae,
D-histidine was identified, which decomposed to give HCN ina reaction
catalyzed by a D-amino acid oxidase in the presence of peroxidase.
High light intensity and high partial pressures of oxygen favored the
production of HCN (Pistorius et al. 1977). HCN production was also
demonstrated in cyanobacteria (Anacystis nidulans), which was shown
to excrete HCN under appropriate conditions (Pistorius et al. 1979).
In this case, the favored precursor was L-histidine, in the presence
of a basic amino acid oxidase with unusuaUy high activity for an enzyme
of this class. Pistorius has continued her work on this enzyme at
the University of Bielefeld where she went when Vennesland retired
in 1981, and has shown that the enzyme or a closely related derivative
is present in photosystem II particles, where it possibly plays an essen-
tial role in the process of photosynthetic O2 evolution.
Englera 7 - 1987 131
Vennesland was one of the first women in the history of

biochemistry and has received world wide recognition. She has gathered
a personal view of her scientific pasture in her "Recollections and small
confessions"(1981).
Hartmann
Max Hartmann (1876-1962), born in Lauterecken (Pfalz), began

his studies with forestry, at the Bavarian College of Forestry in Aschaf-
fenburg, but he soon transferred to the University of Munich to study
biology. After obtaining a Ph.D. under the zoologist R. Hertwig with a
thesis on "Egg formation and egg maturation in the sea star Asterias
glacialis", he stayed for short periods in Strassburg, in Heidelberg, and
in Giessen where he finally qualified as a university lecturer in 1903.
His studies at that time were concerned with "Investigations of gene?
ration shifts of the Dicyemides", a work which was published in 1906
and for which he received the prize of the Belgian Academy of Sciences.
Hartmann's further life and work was often influenced by scien?
tists with whom he was befriended. He began to work on the Protista
with R. Hertwig, one of the most important German zoologists of that
time, with whom he remained on friendly terms. After he left Munich,
it was the zoologist O. Buetschli and the botanist G. Klebs, who gave
him many impulses for his work. Friendships with Theodor Boveri and
with Fritz Schaudinn also go back to that time.
In 1905, at the instigation of F. S chaud inn, Hartmann went
to the Royal Institute for Infectious Diseases (the Robert Koch Insti?
tute of today) in Berlin. There he established a new section for proto-
zoology. His work on the structure and the development of pathogenic
protozoa such as Entamoeba histolytica, trypanosomes, etc. soon ob-
tained for him the reputation of a leading protozoologist. In the year
1909, Hartmann accepted an invitation to Brazil, to the Institute
Oswaldo Cruz, where he built up a protozoology section and was able,
together with Chagas, to extend his research to new tropical protozoa.
At the suggestion of Th. Boveri, Hartmann was invited in 1914
to the newly founded Kaiser Wilhelm Institute for Biology in Berlin-
Dahlem, as leader of an independent section of protozoology. Until
his death in 1962, he remained associated with this Institute, first as
o
?
5-
I
Englera7- 1987 133
one of its directors, and after his retirement in 1955, as a scientific

associate. During World War II, the Kaiser WUhelm Institute for Biology
was transferred to Hechingen, and after the end of the war continued
there as the Max Planck Institute for Biology. During his time in BerUn,
Hartmann belonged to the old BerUn University, from 1909 as
Extraordinary Professor, and from 1921 a Honorary Professor. After
the World War II, he became Honorary Professor at the University of
Tubingen. During the last years of his Ufe, Hartmann withdrew to his
country home in Buchenbiihl (Allgau), where he died in 1962.
In accordance with his scientific training, Hartmann was a zoo-

logist, not a botanist, but his work at the Kaiser WUhelm Institute for
Biology was also of great importance for botany. It was often said,
very telUngly about Hartmann, that he was the last great scientist
who could represent "General Biology". The scientists of later times
can apply themselves only to branches of biology, as a result of in-
creasing speciaUzation. This characteristic of Hartmann was parti?
cularly Ulustrated in his textbook "General biology. An introduction
to the science of Ufe" (Hartmann 1927, 1953). Under general biology,
he understands "directly and mainly the knowledge of the life of the
cell", including "aU processes which occur in Uving bodies". Under
this heading, he then subsumes also aU currently active fields of biology,
such as cell biology, metaboUc and stimulus physiology, ontogeny,
sexuaUty, evolution etc. Such comprehensive works are today only
written by author-collectives, and then edited in a manner far less
satisfying intellectuaUy.
In his main scientific work, Hartmann dedicated himself to the
nature and rules of sex and the determination of sex in the animal
and plant kingdoms. He has assembled his knowledge on this subject
in his book "Die Sexualitat" (1943; 1959), which he dedicated to
Carl Correns. His own theory of sex and fertiUzation consisted of
three laws:
1. The law of general bipolar bisexuality. The law states that in aU
cases of fertiUzation, a female and a male sex ceU copulate, also when
these ceUs can not be differentiated externally.
2. The law of general bisexual potency. This law states that every or?
ganism and every cell (irrespective of whether it is diploid or haploid)
is in principle bisexual. This is among other reasons founded on the fact
that the female haploid genome of Sphaerocarpus is able, after X-ray

irradiation, to develop normal male sex organs (antheridia).
3. The law of the relative strength of male and female determination.
The first definitive proof of the validity of relative sexuality (third
law) was provided in his work (Hartmann 1925 a, b) on Ectocarpus
siliculosus, in which not only female and male gametes appeared to
copulate with each other, but also strong males with weak males, as
well as strong females with weak females. Unfortunately, the prin?
ciple of relative sexuality turned out to be incorrect and was already
critizied by contemporarians (Mainx 1933). In his analyses Hart?
mann encountered that male gametes formed groups of cells surroun-
ding female gametes before subsequent cell fusion and zygote for?
mation. Such group formation was, then, used as a measure for a
fertilization reaction without further proof. Miiller, in 1976, showed
that group formation, as observed by Hartmann with Ectocarpus
siliculosus, can easily be caused by several other factors, but does not
necessarily lead to zygote formation so that the concept of relative
sexuality had to be abandoned again.
Nevertheless, the discussion on sexuality, as propagated by Hart?
mann, turned out to be very fruitful and led finally to the discovery
of gamones. Already in 1933 F. M o e wus, acoworkerof H artm ann,
showed that mt+ and mt-- gametes of Chlamydomonas eugametos gave
off sex-specific substances into the medium. Through the mutual
interaction of these substances, group formation (copulation) occurred.
This then opened the possibility of analyzing the factors for sexual
attraction between gametes in the Kaiser Wilhelm Institute for Medical
Research in Heidelberg, in the laboratory of E. Kuhn (Moewus 1938,
Kuhn et al. 1938, 1939). Finally the carotenoids cis- and trans-cro-
cetin were identified as fertilization factors. Shortly thereafter Hart?
mann (Hartmann et al. 1939, Kuhn & Wallenfels 1939), together with
E. Kuhn and K. Wallenfels, also succeeded in demonstrating the gyno-
and andro-gamones in sea-urchin and later also in rainbow trouts
(1944, 1947).
The originally grandiose advance in the demonstration of gamones
in Chlamydomonas eugametos was clouded by doubts expressed mainly
by American scientists. It developed that Moewus, who had in the
past many valuable scientific contributions to his credit, never obtained
pure compounds for further delivery to Kuhn. This case of deception
Max Hartmann
finaUy entered the newer history of science as one of the first great
scandals. What may have moved Moewus to scientific fraud in this
and other cases (Renner 1958) remains open. Hartmann finaUy
succeeded in demonstrating that a glycoprotein was a gamone in Chla-
mydomonas eugametos in 1956, in one of his last pubUcations, with
Foerster and Wiese.
One can evaluate Hartmann's achievements in the field of sexuaUty
only when one takes into account the fact that at the turn of the century
there did not exist any coherent picture of sexuaUty in biology. At that
time, three were stUl under discussion:
hypotheses a rejuvenation
hypothesis of Buetschli
and Maupas, the amphimixis or germ plasma
mixing theory of Weismann, and the sexuaUty hypothesis of Buetschli
and Schaudinn. The rejuvenation theory stated that, after the
appearance of ageing, fertUization led to rejuvenation. The ground
was withdrawn from this hypothesis when Hartmann (1921) suc?
ceeded in growing the alga Eudorina purely vegetatively for a period of
ten years over 3,000 generations, without the appearance of any signs of
physiological depression. The amphimixis (see above) can on the other
hand, as A. Weismann himself emphasized, only be the result of ferti?
Uzation and therefore never the cause of the first steps in the process.
Nevertheless, the amphimixis theory counted as the ruUng concept,
well into this century. The sexuaUty hypothesis was first formulated by
Buetschli in 1887/89, and then again forgotten in favor of the amphi?
mixis and rejuvenation theory, untU F. Schaudinn in 1904 again
developed ideas simUar to BuetschU's. It was mainly the credit of
Hartmann that the discussion about sexualdifferentiation was much
stimulated in subsequent decades. Hartmann has, incidentaUy,
authored a summary of his work in "Bericht iiber 37 Jahre Forschung
am Kaiser-Wilhelm -(Max-Planck)- Institut fiir Biologie" (Hartmann
1951).
FinaUy it should be stated that Hartmann was one of a few
biologists with an exceUent access to phUosophy. Hartmann devel?
oped clear concepts in articulating the methodological and theoretical
basis of biology and the natural sciences, and in formulating meaningful
boundaries between the natural sciences and the humanities. In this
theory of cognition he distinguishes between a generaUzing and an
exact induction and deduction method and points out that the two
methods of induction and deduction are intimately interconnected by
Englera 7 - 1987 137
analytic and synthetic procedures. The strongest scientific achievements

may originate only if all components of this "fourfold method texture
of induction", i.e. induction, deduction, analysis and synthesis, are
applied altogether although only one method may appear to be the
predominant in a particular example. A short summary on Hartmann
as a philosopher has been presented by Reinboth (1964). His major
publication in this area is "The philosophical bases of the natural
sciences" (Hartmann 1948, cf. 1959). Fifteen of his publications in this
field have been collected in his "Collected lectures and essays II.
Natural philosophy" (Hartmann 1956b). Two of these contributions
deal with the Philosophy of Nature by Nicolai Hartmann. The biologist
and the philosopher Hartmann were namesakes and had an extensive
exchange of thoughts.
Hammerling
One of the most important students of Max Hartmann's at the

Kaiser Wilhelm Institute for Biology was Joachim Hammerling
(1901-1980). From 1920 to 1924, Hammerling studied in Berlin.
He took botany with Haberlandt, and in 1921, a semester of
zoology in Marburg. He obtained his Ph. D. under Max Hartmann
with a thesis on the asexual reproduction of the lower sweet water
worm Aeolosma.
At Hartmann's Institute, and as his assistant, Hammerling
carried out his first basic work with the single-celled alga, Acetabularia,
which later brought him unforgotten fame. Shortly, before the be?
ginning of the Second World War, he was appointed Director of the
German-ItaUan Institute for Marine Biology in Rovigne d'lstria. At
the end of the war, he moved back to Germany, first to an orphaned
Kaiser Wilhelm Institute in Langenargen/Bodensee. Later, he was at-
tached to a division of the Dahlem Kaiser Wilhelm Institute for Bio?
logy, which had in the mean time been moved to Tiibingen. In 1947,
he then received a building from the city of Wilhelmshaven, which was
assigned to the Max Planck Society as the successor of the Kaiser
Wilhelm Society. Here Hammerling established the well-known
Max Planck Institute for Marine Biology which was renamed 1968 as
Max Planck Institute for Cell Biology. At his own request, he was
retired in 1970. An extensive biography onJ. Hammerling has been

presented by Harris (1982).
Acetabularia is a one-celled giant alga from the family Dasyclada-
ceae. It consists of a rhizoid and a stalk. After several weeks in culture,
a hair whorl and a cap develop on the end of the stalk. Until cap
formation, the nucleus continuaUy increases in size. It finally breaks
up. The daughter nuclei wander into the single chambers of the cap
and form cysts. Two experiments with Acetabularia enabled Hammer?
ling to answer a series of basic questions of biology. By removal of
the rhizoid, including the nucleus present in it, he was able, with the
remaining stem, also without a nucleus, to observe the formation of a
whorl and a cap under suitable conditions. Secondly, he succeeded in
transplanting into a type of Acetabularia (A. mediterranea), the nucleus
of another type (A. wettsteinii) and observed the formation of a cap
of A. wettsteinii in the cytoplasm of A. mediterranea, and vice versa.
From his results, he concluded that there must exist morphogenetic
substances for cap formation in Acetabularia, which increased in a
gradient from the rhizoid to the tip of the stem. A reciprocal gradient
of morphogenetic substances must exist for the regeneration of the
rhizoid. These substances must be formed in the nucleus and then
transferred to the cytoplasm, as demonstrated by the nucleus-specific
cap formation in the transplantation experiments (Hammerling 1932,
1934 a-d, 1936).
Hammerling once termed it a stroke of luck that he happened
on this experimental system. Nevertheless, the exciting thing in his time
was the demonstration of a substance (material) which transferred
information for cap formation from the nucleus to the cytoplasm.
Today one naturally speaks of a long-Uved mRNA as the chemical
basis of this substance. To the cytologists and geneticists of that time,
such matters were substantiaUy unknown. Hammerling, not well
versed in biochemistry, always expressed himself very carefully re?
garding this point. Nevertheless, he was fully conscious of the principal
meaning of his conclusions.
From our current point of view, the scientific stimulus to work with
Acetabularia lies now as then in the simplicity of the one-ceUed system
with its long-lived mRNA. In spite of the fact, however, that several
scientists have formed the IRGA Society (International Research
Group on Acetabularia) in 1973, most of the biochemical aspects of
Englera 7- 1987 139
the morphogenic substances are today still unclear, or they are just
now becoming soluble by the methods of molecular biology.
Hammerling has earned special mention in the history of botany
in Berlin, also for the reason that three of his students in Wilhelms-
haven were closely associated with the Free University. Professor H.-G.
Schweiger (1927-1986) finally became Director at the Max Planck
Institute for Cell Biology, and this Institute was moved in 1977 to
Ladenburg near Heidelberg. In 1970, Schweiger received the title of
Honorary Professor at the Free University of Berlin. Two further
coworkers, Heinz Clauss and Giinter Werz, later became professors
at the Institute for Plant Physiology of the Free University of Berlin.
Clauss ceased in 1980. All three have more or less intensively followed
up problems ofAcetabularia.
Schramm
At the Kaiser Wilhelm Institute for Biochemistry, Gerhard Schramm

applied himself for several years to a botanical problem: the first
analysis of a plant virus.
Schramm, the son of a Hamburger family, was born in Yokohama
in 1910. He studied chemistry in Munich and Gottingen. InGottingen,
he became acquainted, among others, with Butenandt. With the
latter, Schramm went to Danzig in 1933, obtained hisPh. D. degree
there, and then followed Butenandt in 1936 to the Kaiser Wilhelm
Institute for Biochemistry in Dahlem. Shortly before the outbreak
of the war, he was chosen to take responsibility for chemistry at a
workplace for virus research, established jointly by the Kaiser Wilhelm
Institute for Chemistry and for Biology.
In Berlin, Schramm was substantially involved with the develop?
ment of the first German air-driven ultracentrifuge (1941). In the
course of his scientific work he carried out the first basic analyses of
tobacco mosaic virus. Initially, this was done in collaboration with his
biological colleagues, G. Melchers, G. Bergold, H. Friedrich-
Freska, R. Danneel, A. Lang and others. Nothing was known at
that time about the viral reproduction mechanism. First, Schramm
showed that the protein component of this RNA-virus was not respon-
sible for its reproduction in the infected cell. For the virus protein
could be modified or partiaUy removed, but the thereby modified

virus particles were still able to cause the reproduction of the original
virus particles in infected cells. In 1943, Schramm succeeded in
separating small identical protein subunits from tobacco rnosaic virus
by treatment with weak base, and then, by treatment with weak acid,
he could reconstitute the original particles, except that the RNA was
now absent. These reconstituted particles were no longer infectious.
Thereby it was proved that the protein component of the virus does
not contain the information for viral replication, although it possesses
a significant function in biological morphogenesis. This important
finding was at first not recognized abroad, because of the political
events in Germany at that time, and because of the war. Butenandt
(1971) has cited it as a good example of the fact that at some institutes
in Germany, in spite of difficult conditions, good research could stUl be
done, indepent of the regime.
In 1943/44, because of increasing aerial bombardment, the Kaiser
WUhelm Institute for Biochemistry was moved to Tiibingen. Schramm
with his work group was first given space at the Hygienic Institute
of the University of Tiibingen. Later he received his own section in the
new Max Planck Institute for Viral Research. In 1956, Schramm,
together with Gierer (1956 a, b), succeeded in demonstrating that the
isolated RNA contained aU the information necessary for the repli?
cation of the virus. Mutations were caused by treatment of the viral
RNA with nitrite (Schuster & Schramm 1958). After determination
of the amino acid sequence of the viral proteins, it could be shown
that the mutation was associated with a change in the amino acid
sequence.
With his work in Berlin, and especiaUy with his later investigations,
Schramm laid a strong foundation for our knowledge of the tobacco
mosaic virus. In this field, he was a pioneer of molecular biology. He
died at the early age of 59, during the preparation of a Friedrich
Miescher Symposium in Tiibingen.
The Ufe and scientific merits of other botanists at Kaiser WUhelm

Institutes or Max Planck Institutes, primarily geneticists, are pre?
sented in the articles by W. Plarre and by A. Lang.
Englera 7-1987 141
Acknowledgements
I would like to thank Dr. B. Vennesland for the translation of the

German manuscript of this article and Prof. Dr. K. M. Hartmann,
Erlangen, for some helpful discussions. Thanks are also due to Dr. E.
Henning for giving access to photographs of the former Kaiser Wilhelm
Institutes for Biology and for Cell Physiology, of Hartmann and of
Warburg in the "Archiv zur Geschichte der Max-Planck-Gesellschaft".
Mrs. S. Dobre-Seidenzahl has kindly provided the photograph of the
former Kaiser Wilhelm Institute for Chemistry. The photograph of
Willstatter was taken from "Willstatter, R. 1949: Aus meinem Leben.
Von Arbeit, Mufie und Freunden."
References
Andrews, T. J., Lorimer, G. H. & Tolbert, N. E. 1973: Ribulose di-

phosphate oxygenase. I. Synthesis of phosphoglycolate by fraction-I
protein of leaves. ? Biochemistry 12: 11-18.
Bladergroen, W. 1960: Problems in photosynthesis. - Springfield, 111.
Bowes, G., Ogren, W. L. & Hagemann, R. H. 1971: Phosphoglycolate
production catalyzed by ribulose diphosphate carboxylase. -
Biochem. Biophys. Res. Commun. 45: 716-722.
Butenandt, A. 1971: Vorwort. ? In: G. Schramm, Bauplane des Lebens.
Probleme und Ergebnisse der Biochemie. ? Miinchen (with a com?
plete bibliography of Schramm's publications), p. 7-14.
Engel, M. 1984: Geschichte Dahlems. - Berlin.
Evans, H. J. & Nason, A. 1953: Pyridine nuceotide-nitrate reductase
from higher plants. - Plant Physiol. 28: 233-254.
Festschrift Arthur Stoll zum siebzigsten Geburtstag: 8. Januar 1957.
Arbeiten aus dem Gebiet der Naturstoffe. ? Basel 1957 (with a
complete bibliography of Stoll's publications).
Frank, J. 1931: Emil Warburg zum Gedachtnis. ? Naturwiss. 19.
993-997.
Gerwin, R. 1986: 75 Jahre Max-Planck-Geseilschaft. Ein Kapitel
deutscher Forschungsgeschichte. Teil 1: Griindungsgeschichte und
erste Aufbauphase der Kaiser-Wilhelm-Gesellschaft. ? Naturwiss.
Rundschau 39: 1-10.
Gierer, A. & Schramm, G. 1956a: Die Infektiositat der Nucleinsaure

aus Tabakmosaikvirus. - Z. Naturf. 11b: 138-142.
- -1956b: Infectivity of ribonucleic acid from tobacco mosaic virus. -
Nature 177: 702-703.
Hammerling, J. 1932: Entwicklung und Formbildungsvermogen von
Acetabularia mediterranea. ? Biol Zentralbl. 52: 42-61.
- 1933: Uber formbildende Substanzen bei Acetabularia mediterranea,
ihre raumliche und zeitliche Verteilung und ihre Herkunft. - W.
Roux' Archiv Entwicklungsmech. Organismen 131: 1-81.
- 1934a: Regenerationsversuche an kernhaltigen und kernlosen Zell-
teilen von Acetabularia wettsteinii. ? Biol. Zentralbl. 54: 650-665.
- 1934b: Uber Genomwirkungen und FormbUdungsfahigkeiten bei
Acetabularia. ? W. Roux' Archiv Entwicklungsmech. Organismen
132:424-462.
- 1934 c: Entwicklungsphysiologische und genetische Grundlagen der
Formbildung bei der Schirmalge Acetabularia. - Naturwiss. 22:
829-836.
- 1936: Studien zum Polaritatsproblem. I-III. ? Zool. Jahrb. 56:
439-486.
Hagemann, R. H., Cresswell, C. F. & Hewitt, E. J. 1962: Reduction of
nitrate, nitrite and hydroxylamine to ammonia by enzymes extracted
from higher plants. - Nature 193: 247-250.
Harris, H. 1982: Joachim Hammerling. - Biogr. Mem. FeUows Roy.
Soc. 28: 114-124 (with a complete Ust of Hammerling's pubU-
cation).
Hartmann, M. 1921: Die dauernd agame Zucht von Eudorina elegans. -
Arch. Protistenkunde 43: 223.
- 1925a: Uber die relative SexuaUtat bei
Ectocarpus siliculosus. -
Naturwiss. 13: 975-980.
- 1925b: Untersuchungen iiber die relative SexuaUtat. I. Versuche
an Ectocarpus siliculosus. ? Biol. Zentralbl. 45: 449-467.
- 1927: Allgemeine Biologie. Eine Einfiihrung in die Lehre vom
Leben. - Jena, 4th edition 1953.
- 1943: Die SexuaUtat. Das Wesen und die
Grundlagen desGeschlechts
und der Geschlechtsbestimmung im Tier- und Pflanzenreich. ? Jena,
2 nd edition 1956.
- 1944: Befruchtungsstoffe (Gamone) bei Fischen (Regenbogen-
forelle). - Naturwiss. 32: 231.
Englera 7-1987 143
- 1948: Die philosophischen Grundlagen der Naturwissenschaften.

Erkenntnistheorie und Methodologie. - Jena, 2nd edition Stutt?
gart 1959.
? 1951: Bericht iiber 37 Jahre Forschung im Kaiser-Wilhelm- (Max-
Planck)-Institut fiir Biologie. ? Jahrbuch 1951 der Max-Planck-
Gesellschaft zur Forderung der Wissenschaften e.V.: 199-228.
? 1956a Einfiihrung in die allgemeine Biologie und ihre philosophi?
schen Grund- und Grenzfragen. ? Sammlung Goschen 96. ? Berlin.
? 1956b: Gesammelte Vortrage und Aufsatze II. Naturphilosophie. ?
Stuttgart.
- Forster, H. & Wiese, L. 1956: Neue Befunde iiber die Befruchtungs-
stoffe (Gamone) und ihre theoretische Bedeutung. ? Naturwiss.
43:313-317.
- Medem, F. Graf von, Kuhn, R. & Bielig, H.-J. 1947: Uber die
Gynogamone der Regenbogenforelle. ? Naturwiss. 34: 25-26.
- Schwartau, O., Kuhn, R. & Wallenfels, K. 1939: Uber die Sexual-
stoffe der Seeigel. ? Naturwiss. 72: 433.
Krebs, H. A. 1979: Otto Warburg. Zellphysiologe, Biochemiker, Medi-
ziner. ? Stuttgart (with a complete list of Warburg's publication).
Kuhn, E., Moewus, F. & Jerchel, D. 1938: Uber die chemische Natur
der Stoffe, welche die Kopulation der mannlichen und weiblichen
Gameten von Chlamydomonas eugametos im Lichte bewirken. -
Ber. Deutsche. Chem. Ges. 71: 1541-1547.
- - & Wendt, G. 1939: Uber die geschlechtsbestimmenden Stoffe
einer Grunalge. - Ber. Deutsche. Chem. Ges. 72: 1702-1707.
? & Wallenfels, K. 1939: Uber die chemische Natur des Stoffes, den
die Eier der Seeigel (Abracia pustulosa) absondern, um die Sperma-
tozoen anzulocken. ? Ber. Deutsch, Chem. Ges. 72: 1704-1713.
Lorimer, J. H., Gewitz, H.-S., Volker, W., Solomonson, L. P. & Vennes?
land, Birgit 1974: The presence of bound cyanide in the naturally
inactivated form of nitrate reductase of Chlorella vulgaris. ? J.
Biol. Chem. 249: 6074-6079.
Mainx, F. 1933: Die Sexualitat als Problem der Genetik. Versuch eines
kritischen Vergleiches der wichtigsten Theorien. ? Jena.
Moewus, F. 1938: Carotinoide als Sexualstoffe von Algen. ? Jahrb.
Wiss. Bot. 86:753-783.
? 1933: Untersuchungen iiber die Sexualitat und Entwicklung der
Chlorophyceen. - Arch. Protistenkunde 80: 469-526.
Muller, D. G. 1976: Relative sexuaUty in Ectocarpus siliculosus. A

scientific error. - Arch. Microbiol. 109: 89-94.
Nachmansohn, D. 1979: German-Jewish pioneers in science, 1930-
1933.-Heidelberg.
Pistorius, K., Gewitz, H.-S., Voss, H. & Vennesland, Birgit
Elfriede,
1977: Cyanide formation from histidine in Chlorella. A general
reaction of aromatic amino acids catalyzed by amino acid oxidase
systems. - Biochim. Biophys. Acta 481: 384-394.
- Jetschmann, K., Voss, H. & Vennesland, Birgit 1979: The dark
respiration of Ancystis nidulans. Production of HCN from histidine
and oxidation of basic amino acids. ? Biochim. Biophys. Acta
585:630-642.
Reinboth, R.: Der Biologe Max Hartmann als Naturphilosoph. ?
Philosophia Naturalis 8:3-8.
Renner, O. 1958: Auch etwas iiber Moewus, Forsythia and Chlamy-
domonas. - Z. Naturforsch. 13b: 399-403.
Schramm, G. 1941: Die luftgetriebene Ultrazentrifuge. ? Kolloid-Z.
97: 106.
- 1943: Uber die Spaltung des Tabakmosaikvirus in niedermolekulare
Proteine und die Riickbildung hochmolekularen Proteins aus Spalt-
stiicken. ? Naturwiss. 31: 94-96.
Schuster, H. & Schramm, G. 1958: Bestimmung der biologisch wirk-
samen Einheit in der Ribonucleinsaure des Tabakmosaikvirus auf
chemischem Wege. - Z. Naturf. 13b: 697-704.
StoU, A. 1933: Ein Gang durch biochemische Forschungsarbeiten. -
Berlin.
Tolbert, N. E. 1963: Glycolate pathway. ? In: Photosynthetic me-
chanisms in green plants. ? Publication 1145, Nat. Acad. Sci. USA,
Nat. Res. Council, p. 648-662.
Vennesland, Birgit 1958: Stereospecificity of hydrogen transfer in pyri?
dine nucleotide dehydrogenase reactions. ? Federation Proceedings
17:1150-1157.
- 1981: Recollections and small confessions. ? Ann. Rev. Plant
Physiol. 32: 1-20.

Wang, D. & Waygood. E. R. 1962: Carbon metaboUsm of C14-labeled
amino acids in wheat leaves. I. A pathway of glyoxylate-serine
metaboUsm. - Plant Physiol. 37: 826-832.
Englera 7- 1987 145
Warburg, O. 1913-1922: Die Pflanzenwelt.I. Protophyten, Thallo-

phyten, Archegoniophyten, Gymnospermen und Dikotyledonen;
II. Dikotyledonen; III. Dikotyledonen und Monokotyledonen. ?
Leipzig.
Warburg, O. H. 1908: Beobachtungen iiber die Oxidationsprozesse
im Seeigelei. ? Hoppe-Seyler' Z. Physiol. Chemie 57:
? 1919: Uber die Geschwindigkeit der photochemischen Kohlen-
saurezersetzung in lebenden Zellen. - Biochem. Z. 100: 230.
? 1920: Uber die Geschwindigkeit der photochemischen Kohlen-
saurezersetzung in lebenden Zellen II. ? Biochem. Z. 103: 188-217.
? 1926: Uber den Stoffwechsel der Tumoren. ? Berlin.
? 1946: Schwermetalle als Wirkungsgruppen von Fermenten. -
Berlin, 2nd edition 1948.
? 1948: Wasserstoffiibertragende Fermente. ? Berlin.
? & Krippahl, G. 1960a: Notwendigkeit der Kohlensaure fiir die
Chinon- und Ferricyanid-Reduktion in griinenGrana. - Z. Naturf.
15b: 367-369.
-1960b: Glykolsaurebildung in Chlorella. - Z. Naturf. 15b:
197-199.
? & Liittgens, W. 1944: Chinon als Hill-Reagenz. ? Naturwiss. 32:161.
? & Negelein, E. 1920: Uber die Reduktion der Salpetersaure in
grunen Zellen. ? Biochem. Z. 110: 66-115.

? ? 1922: Uber den Energieumsatz bei der Kohlensaureassimilation. ?
Z. Physikal. Chem. 102: 235-266.
? ? 1923: Uber den Einflufi der Wellenlange auf den Energieumsatz
bei Kohlensaureassimilation. ? Z. Physikal. Chem. 106: 191-218.
Westheimer, F. H., Fisher, H. F., Conn, E. E. & Vennesland, Birgit

1951: The enzymatic transfer of hydrogen from alcohol to DPN. ?
J. Am. Chem. Soc. 73: 2403.
Willstatter, R. 1949: Aus meinem Leben. Von Arbeit, Mufie und
Freunden (A. Stoll, ed.). ? Weinheim/Bergstrafie (with a complete
bibliography of Willst att er's publications).
? & Stoll, A. 1913: Untersuchungen iiber Chlorophyll. Methoden und
Ergebnisse. ? Berlin.
? ? 1918: Untersuchungen iiber die Assimilation der Kohlensaure. ?
Berlin.

Prof. Dr. Claus Schnarrenberger, Institut fiir Pflanzenphysiologie,
ZeUbiologie und Mikrobiologie, Freie Universitat Berlin, Konigin-
Luise-Strafie 12-16a, D-1000 Berlin 33.
Englera 7 - 1987 147
Werner Plarre
A contribution to the history of science of heredity in Berlin
Introduction
Although the term "science of heredity" is not very popular, it was

chosen for the title, because of the historical importance in Berlin and
in Germany. In trying to recapitulate the historical events in this
science, only local research projects and results will be mentioned.
Therefore it simply would be anachronistic to speak of genetics in this
way. The change in the term will be pointed out in the chapter on Carl
C o r r e n s.
At present, the knowledge of modern genetics is rapidly increasing
and will continue to do so furthermore. To realize the importance
of historical studies in this field, one should stop for a moment and look
into the past. Therefore, it is more important to focus on the findings
from earlier history rather than on the scientific results of the past
decades. These results are continuously and extensively published,
and those publications are easily accessible. All of this has been taken
into consideration in the following.
More than 25 years ago Stubbe (1961) published a "History of
genetics up to the point of the rediscovery of Mendelian rules" (Kurze
Geschichte der Genetik bis zur Wiederentdeckung der Vererbungs-
regeln Gregor Mendels). A second edition appeared in 1964. Mentioned
were ideas, results, experiments, hypotheses, theories and philosophical
thoughts on the various characters inheritable in plants, animals and
man. Outstanding personalities were deemed worthy of recognition.
Their works have led to advancement in heredity and applied field
work. Extraordinary discoveries which led to developments in new
areas were outlined. In such a framework the following may, could and
should be regarded from the same point of view.
Although not strictly followed, the outline of this essay is divided
into time periods. Chronological overlapping is due to the necessity of
148 Plarre: Science of heredity in Berlin
recording the ideas and directions of research work of specific institutes

over time. In honouring particular personaUties, it was not regarded
as important to account all experiments and results but rather to relate
other particulars such as the environmental and political conditions
under which their work had been done. Also, in order to create a more
colourful picture of the scientists, their personal and private Ufe wUl
be mentioned. Biographical studies were as important as personal
impressions. I stUl tried to be as objective as possible.
Supporting contributions of the Botanical Garden and Museum
If one had to write the history of the science of heredity in Hei?

delberg and not in Berlin, one would have to step further back into
the past by several centuries. One would have to begin with the discovery
of a splitted leaf mutant of Chelidonium majus laciniatum in a pharma-
ceutical garden in 1590. At that time Heidelberg already was a well-
known university city, and Berlin was only a smaU residence city
(Mark Brandenburg). The special deviated form of Ch. majus laciniatum
appears in natural habitats also in Berlin. But it has only recently been
discovered and has not yet been described. This mutant is similar
(analogous) to that of the bipinnatifid leaf type (mutant) of Sambucus
nigra, found in the garden of the Institute of Applied Genetics in Dahlem
in 1972 and bred true since that time as a clone. The Ch. majus la?
ciniatum of Heidelberg was tested and confirmed as a pure Une: the
dissected leaves have remained inheritably constant up to the present.
If one had to look in Berlin for an example par excellance that only
in the broadest sense had something to do with genetics, one would
find nothing until 1749. But this would then be a spectacular event,
which under the name "experimentum berolinense" created quite a
sensation among the botanists of those days (Zepernick & Karlsson
1979).
Johann Gottlieb Gleditsch 1714-1786
In the 17th and 18th century a series of fundamental experiments

were accomplished which provided proof of sexuaUty in plants, i.e.
Englera 7 - 1987 149
the existence of female and male sex organs (Stubbe 1964: 78). When
Gleditsch pollinated a female plant of the dioecious palm Chamae-
rops humilis in Berlin with pollen which he had procured from the
Leipzig Botanical Garden from a male one, Gleditsch had success
in fruit setting. He had realized, that this "experimentum berolinense"
belonged to the ranks of the greatest experiments of botanists of that
time. Those, who today work with artificial pollination know how
shortlived is pollen under natural storage conditions. The palm fruits,
which had been produced in Berlin, furnished definiteproof of sexuality
in plants. It is doubtful whether it was known throughout the botanical
world, how the Assyrians, around 850 B. C, pollinated their dioecious
date palms. Later, Chamisso reported it in full details.
The fruit set, which Gleditsch obtained, assured him of satisfac-
tion surely. It was much later, that some one in Berlin became con?
cerned about the heredity of sex in plants and animals. But it is remark-
able that Gleditsch was a forerunner in this field of research.
Adelbert von Chamisso 1781-1838
"There will be something new!"
With this sentence Chamisso announced to his friend, the bo-

tanist de la Foya, his scientific work "De animalibus quibusdam e
classe vermium Linnaeana: De Salpa" (On the genus Salpa), which
appeared in 1819 in Berlin. The investigations of this work were carried
out on board of the "Rurik" from 1815-1818 during the Romanzoff
Research Expedition, in which Chamisso participated as a natural
scientist. In his publication with coloured drawings Chamisso des?
cribed and documented exactly for the first time the "alternation of
generations" in animals. This discovery earned Chamisso the Honorary
Ph. D. of the newly founded Friedrich Wilhelm University. However,
his opponents with cynical criticism gave him the title of "scientific
fairytale teller". The discovery achieved international recognition,
first in 1842 through the Danish zoologist Steenstrup as well as through
the British scientistsand Hensly in 1848-1852
Forbes (Schneebeli-
Graf 1983). He was an honest and precise scientist, who was way
ahead of his time.
Chamisso became in 1819 an assistent in the Botanical Garden,

and in 1833 he took over as the first administrator of the Herbarium.
His bontanical works primarily dealt with the sexuaUty of plants.
He reported that the palm Chamaerops humilis which had been polli-
nated by Gleditsch could stUl be seen with its offsprings in the
Botanical Garden. In detaU he expressed his opinion on the asexual
reproduction by animals (polyps etc.) and plants. Above all, he advises
that all the characteristics of completely developed plants (colour,
chlorosis, double corolla, etc.) continue in cuttings; a new independent
life can develop only from the seed itself. Thereby Chamisso comes
close to the idea that a clone corresponds to one and the same idiotype
(genotype) and only with sexual reproduction can new "develop-
ments" (changes) be expected.
Worth mentioning are also his examples of the immigration of
plants and animals as followers of human culture, especiaUy of cereal
farmers. He provides precise data on the distribution of cultivated
crops and their accompanying flora (weeds) like the colonization of
islands or the expansion up to the northern cultivation border. He writes
that barley was cultivated at 70?N in Lapland, and that the corn-
flower (Centaurea cyanus) has its border at the 64 N near Umea
(Sweden). He was familiar with how the date palm culture was carried
out in the Levant, "preferably the female trees are cultivated, and one
tended to bring cut male flower panicles to the plantations and to hang
these panicles on the fruit trees. The insects execute the pollination"
(Schneebeli-Graf 1983). Therefore Chamisso touches upon a field
of research, that of the domestication of cultivated plants, which in-
deed has a causal connection to applied genetics.
Chamisso was 15 years old when he emigrated from France to
Berlin with his mother and his brothers and sisters. Later, he repeatedly
returned. Yet, he always sensed the feeling of homelessness: neither
to be French or German. FinaUy, after his departure from Prussian
military service he found an inner peace through his broad scientific
and Unguistic studies, as well as in poetry. He became famous as a poet
but his scientific pubUcations are just as outstanding. These scientific
pubUcations (about 2,500 printed pages) amount to double in compari-
sion to his poetic writings. In the Anglo-American speaking world
they are more respected than in Germany (SchneebeU-Graf 1983).
Nevertheless, in the entrance hall of the Botanical Museum hangs a
bronze memorial plague with his head in reUef.
Englera 1- 1987 151
Georg Schweinfurth 1836-1925
With a historicalreflection on the research projects that deal with

the evolution(domestication) of cultivated plants, one finds very early
in Berlin an area of agrarian-geographical and paleoethnobotanical
work. Recognition that were won for later genetic experiments on
cultivated crops were very valuable. The initial steps are to be found
in the first half of the 19th century with Chamisso. In the second
half of the 19th century one encounters Schweinfurth, the Africa
explorer, who obtained great merit in this field.
On one of his travels in 1870 Schweinfurth discovered the wild
forms of Ricinus communis and Citrullus lanatus (water-melon) in the
jungle of the White Nile (Schiemann 1938). In his ability to integrate
all his results in an overall and general consideration, he made reference
in 1891 to the development of culture, which has a close connection
with the domestication of plants. In the "cosmopolitan nature of agri?
culture" he sees a common link, that unites mankind, and that all
people are dependent upon one another. He recognizes that a national
economical development should follow the "law of differentiation
and integration", and therefore should be handled also on an inter?
national level. If one looks upon economic negotiations in our days,
no one can recognize that politicans have learned anything from former
cognitions.
Schweinfurth not only collected living plant material but
also worked as a paleoethnobotanist. The plant material which he
brought from his winter trips was then cultivated in Berlin, in the
summer, and stored in the Herbarium of the Botanical Museum. In
1943, during the war, the Herbarium was destroyed by fire. But fortu-
nately precious objects, namely neeklaces, headbands and other pieces
of jewelry made out of plant material (flower, fibres) which were found
by Schweinfurth in the royal ancient Egyptian tombs could be
saved. The "mummy wheat" (Triticum turgidum var. dicoccum) which
was also brought out of the tombs failed to germinate after several
attemps. There exists an amusing anecdote about the "mummy wheat".
Schweinfurth repeatedly tried to get that wheat to germinate by
planting in the garden. One day some seedlings actually appeared.
How they got to be there, however, was due to Schweinfurth's
factotum who had placed a few fresh seeds between the others. "I
only wanted to do the professor a favour, because Herr Professor so

much wished to discover germinating wheat kernels." Another often
told story about Schweinfurth is not humorous at all. It shows
more his ignorance the need for preserving original nature in Africa,
for he prided himself on shooting down masses of hippopotamuses
during his trips to the NUe areas.
Elisabeth Schiemann 1881-1972
The works on the history of cultivated plants have been carried on

into the 20th century in Berlin from a broader base in a special and
wonderful way here in the Botanical Garden. From 1930-1943 in
order to better clarify the phylogeny of cereals, especiaUy the Uneage
and origin of wheat, E. Schiemann followed through many experi?
ments and genetic analyses. Demonstrating the lines of descent of "the
wUd to the cultivated forms" of different species, E. Schiemann was
able to point out numerous mutations which foUowed Mendelian segre-
gation, e.g. in barley, tomatoes, snapdragon, and other ones. Her goal
was to depict through the incorporation of taxonomic examinations
a comprehensive "History of cultivated plants", involving their origin
and evolution from wUd types to high bred cultivars.
In her long succesful working life, E. Schiemann became widely
noted. She did research with very exact precision in her working pro-
grammes. But her function in the Botanical Garden was unfortunately
only an intermezzo. The heavy bomb attack, which destroyed the ex?
tensive herbarium material of the Museum in 1943, brought her work
here to an end. Since this time very Uttle genetic work has been done
on cultivated plants other than that which was reported by H. Scholz
(1986) in the last years on the origin of weed types in Panicum and
barley.
The life work and personality ofE. Schiemann are also mentioned
and praised in another contribution (see Lang's chapter). A biography
by her student, H. Kuckuck, was pubUshed in 1980. To this I only
want to add, from personal meetings with this highly honoured lady of
science, what appears worthy of mention to me of her research works.
Her "thoughts on VavUov's Gene-Centres-Theory" (1939) were for me
Englera 7 - 1987 153
very meaningful. With her description of the processes of the evolution

of crops, like those in "secondary centres" she corrected Vavilov's
hypothesis of regarding Ethiopia as a centre of origin for wheat, as it
is much more a "withdrawal centre" of the tetraploid wheat. Originally,
diploid forms were missing here. E. Schiemann held the view that
also in recent times gene-centres originate in small areas. Asan example,
she cited the northern Alpine border where a large variability of hexa-
ploid wheats is found. Those are, in spite of their morphological dif-
ferences, well adapted to the specific ecological conditions. Today
these landraces are still valuable gene resources. In consideration of
these questions the discussion of migration plays a greater role (Schie?
mann With rye, for example, the problem arises, how did
1939).
Secale africanum reach into the highland of South Africa? A migration
from Orient out (Arabian Peninsula), over East Africa along the chains
of mountains and gaps up to the Cape is imaginable (Hoffman et al.
1985: 140). With her critical position E. Schiemann led the way
for other scientists who worked in this field of gene-centres (Zhukowsky
1956, Kuckuck 1962, Plarre 1972, Schachl 1984).
An encounter with E. Schiemann on the occasion of her award
of the Honory Doctor of Berlin's Technical University in 1962 is
helpful to me in characterizing her personality. What impressed me
the most was-the sovereignty with which the 80 year-old lady accepted
this award. With a sure voice she responded, freely speaking to the
greeting and laudatory speech given by the Vice-Chancellor of the
University and the Dean of the Faculty of Agriculture. She mentioned
that she felt, she had been chosen for the field of the science of heredity,
and to some extent, for the direction of her work. She explained
it in the following manner: "Because my studies occurred at a time
when genetics was young, I have been able to experience at first hand
its stormy development almost from the beginning. Decisive in my
life were, above all, the encounters with great personalities, of which
I name three in particular: Erwin Baur, Georg Schweinfurth and Walter
Andrae, who provided us with the beautiful Ischthartor in the Berlin
Museum". Even in her old age, E. Schiemann was always a per?
sonality who radiated greatness upon her colleagues. This can be seen
in the photograph of the 85th birthday.
Englera7 - 1987 155
The development up to 1945
Friedrich WUhelm University
After Oscar Hertwig discovered the fertiUzation of the egg of the sea-
urchin, Toxopneustes lividus, in vivo (1875) the science of heredity
received a strong impetus in Germany. At the same time research was
done on the clarification of the fertiUzation process on botanical
objects as well. In Berlin it appeared that this new knowledge, which
had finaUy contributed to the new science of heredity, would only
cause the zoologists to profit. Oscar Hertwig (1849-1922) in 1888
was the new professor in "General Anatomy, Histology, Developmental
and Comparative Anatomy" at the Friedrich WUhelm University
untU 1921. During his time in BerUn he did the fundamental investi?
gations of ovo- and spermatogenesis (1890). He was able to prove
explicitly the "reductional division" (meiotic reduction) during the
gametogenesis in Ascaris megalocephala. In 1918 Oscar Hertwig
concluded and summarized his research work: "Dokumente zur Ge?
schichte der Zeugungslehre" (Stubbe 1964: 248/249).
The brothers Oscar and Richard Hertwig, the former active in
Berlin, the latter in Munich, towards the end of the 19th century,
opened the crucial door to new aspects for the science of heredity with
their developmental, physiological and karyological investigations of
the fertiUzation process. At the same time there was not much done in
this field in botany at the Friedrich WUhelm University. The professor-
ships for both General Botany (BerUn NW., Dorotheenstrasse) and
Systematic and Plant Geography (BerUn N., InvaUdenstrasse) were
occupied by notable scholars, Schwendener and Engler. Gene?
tic investigations, however, even in the broadest sense, were not carried
out (Leussink et al. 1960: 847). That changed in 1903 when Erwin
Baur became the assistent of Schwendener.
Through the rediscovery of Mendel's laws in 1900 the ground was
prepared for the rapid development of the science of heredity. Baur
started with experiments in a smaU greenhouse in the institute in
the Dorotheenstrasse. He executed the first attempt at clarifying the
infectious spread of variegation in leaves of Malvaceae. Chlorotic
forms of snapdragon (Antirrhinum) representatives of the so-called
aurea-group, became his next subjects. Baur proves that the 2: 1
segregation aurea: typica is attributable to a lethal factor ? the one
group of homozygotes dies in the embryonic stage or in the early

seedling stage. Therefore the work led to a plant which Baur had
followed for quite some time, and from which he had obtained many
hereditary secrets. This species Antirrhinum majus has become the
example par excellance for the science of heredity in botany. It is from
this species that Baur has obtained his nickname: "Baur mit dem
Maul eines Lowen" (Baur with the snap of a dragon). In 1907 the first
important publication on Antirrhinum was concerned with a form only
viable as hybrid. Soon in the university garden the experimental field
no longer sufficed. Therefore, Baur established his breeding garden
on his private grounds in Berlin-Friedrichshagen. This garden was not
abandoned until 1916 when a larger area was made available in Pots?
dam. It was namely here that a new institute was planned for him.
Baur later handed over to co-workers, whom he entrusted to
compile the genetic themes from horticultural and agricultural practice,
a broad er assigment from basic research. The snapdragon could never be
excluded. By doing so he wanted to emphasize a botanical object
parallel to Drosophila. Nearly 30 works on Antirrhinum have been
published by him. They were summarized in 1924 with detailed pedi-
grees, illustrations and nice coloured plates. When one reads this publi?
cation, one experiences aesthetic enjoyment. One also recognizes the
amount of devotion with which Baur accomplished his work as well
as the amount of enthusiasm he had for the variation of colours and
formations of the various blossoms (Baur 1924: 167). What would
it have meant to him if he had experienced what has emerged from a
combination of the mutants divaricata x hemiradialis in practical
breeding? The result of this combination is a wide opened and butter-
fly-like bloom. In 1966 the new cultivars "Bright Butterfly" and
"Juliwa" were prizewinners of the All-America-Selection! One gram
of the seeds cost then 50 DM. At the end of the 40s the first strains
were produced at the Max Planck Institute in Ladenburg/Rosenhof
(Knapp 1966).
While Baur was an assistant, and took the qualifying examination
to become a lectureship (habilitation 1904), he offered already several
lectures on heredity until 1911, when he became a professor. His first
textbook was then published: "Einfiihrung in die experimentelle
Vererbungslehre" (Introduction into experimental heredity). The book
became a standard work which quickly sold out and regularly had to
Englera7 - 1987
157
be re-published. UntU B aur's death it appeared in 11 editions. Because

he was convinced of the significance of the science of heredity to the
poUtical economy Baur made clear it in his lectures and practical
courses. Under the title: "WissenschaftUche Grundlagen der Pflanzen-
ziichtung, ein Lehrbuch fiir Landwirte, Gartner und Forstwirte" (The
scientific fundamentals of plant breeding, a textbook for farmers,
gardners, and foresters) he provided a valuable teaching book for
students and experts. At that time, stUl engaged at the Faculty of
Philosophy, Baur gave out dissertations on genetics. In 1908 with
Carl Correns and Valentin Haecker he founded the first scientific
periodical: "Zeitschrift Abstammungs- und Vererbungs-
fiir induktive
lehre" (Journal for the inductive science of descent and heredity).
The periodical soon received international attention.
The rediscovery of Mendel's laws evoked a strong response in Germany
up untU World War I, but genetics had no foothold at the universities.
Only then could a young science develop for a length of time ? as
Nachtsheim (Leussink et al. 1960: 858) ascertained in retrospect -
if it could be taught at universities. At the Friedrich WUhelm Univer?
sity this was not possible for one reason or another. After World War I
the University prided itself on having engaged an Honorar Professor
Carl Correns, who since 1914 had been the Director of the Kaiser
Wilhelm Institute for Biology. However, at Friedrich WUhelm Uni?
versity no such institute was founded. Other German universities did
not see the necessity either. Only at the LandwirtschaftUche Hoch-
schule of BerUn (Agricultural CoUege) one was estabUshed. Here it
should especiaUy be mentioned that during the "Third Reich" from
1933-1945 no institute of genetics was founded at any German uni?
versity, if one disregards a professorship for genetics which was estab?
Ushed in 1941 at the Reichs University of Strassburg (Alsace) and of
which Edgar Knapp became the director. He moved from the Kaiser
WUhelm Institute of Breeding Research in Miincheberg (Telschow
1942). Otherwise at this time heredity teachings at the universities
would have been misused and biased. Professorship for Racial Hygiene,
Racial Biology and even for Racial Psychology were estabUshed. It
is unfortunate that there were not only profiteers, but also a few
well-known human geneticists and anthropologists who offered their
services and risked their good reputations (Leussink et al. 1960: 860).
Agricultural College
In Berlin the untiring efforts of an Erwin Baur, who in 1911

became the University Professor in Botany at the Agricultural College,
succeeded in founding an institute which, when the construction plans
were completed, was to be especially equipped for him in 1914. The
war interrupted this and for the time being, only a temporary building
was erected in Potsdam. Just after the war the Institute of Heredity
and Breeding Research was then built in Berlin-Dahlem and in 1922-23
was ready for student admissions. The Institute is furnished in the
most modern way, with laboratories and glass-paneled bay (here
terrariums, aquariums and cages were located). Stables were built for
larger animals (chickens, rabbits). It was most important, however, that
greenhouses and an experimental field of 3,5 ha became available.
Below, Erwin Baur, his colleagues and successors at this Institute, as
well as the most important research work will be described in more
detail.
In connection with the foundation of the Institute the historical
development of the Agricultural College should be mentioned briefly
and especially the changing membership which this Research Institute
of Heredity and Breeding, the first one in Germany, pointed out.
The College had been founded in 1881 as a "Royal Prussian" one, the
Friedrich Wilhelm University already 1810. However, Albrecht Thaer
as Professor of agricultural science also belonged with his institute
to the University at that time. The beginning of agricultural studies
on academic level can be dated in Berlin back to 1810. The designation
"Royal Prussian" was naturally abolished after World War I. The College
lost its independence in 1934 when it was incorporated into the Fried?
rich Wilhelm University, at first as the Agricultural Veterinary Faculty
and from 1937/38 onward as the Agricultural Horticultural Faculty
(the science of horticultural had been added).
The development from 1945-1961
Humboldt University
After World War II the Friedrich Wilhelm University was reopened

and in the Faculty of Science a Chair of Genetics was created and an
Englera7-1987 159
institute founded. Hans Nachtsheim was appointed Director of the

Institute for Genetics. In addition, the Institute had been set up by
Baur at the Agricultural Horticultural Faculty survived. The Uni?
versity was renamed and became known as the Humboldt University.
The "poUtical administration unit Berlin" only existed for a short
while after 1945. In the late forties the city was divided and at uni?
versity level an intellectual struggle became increasingly apparent.
As far as biology was concerned, both in basic and applied research
the doctrine of Lysenko, which had emerged during the StaUn era
and later was found to beeing pseudo-scientific, had a disastrous
role in this dispute. It claims that acquired characteristics can be in-
herited and attaches importance to the appUcation to certain methods
of treatment and rearing (the influence of a mentor) during the onto-
genesis of a Uving creature, in order to induce specific, permanent
changes in its characteristics. A huge propaganda campaign was launched
by Soviet and some German scientists and by the organs of their
Communist parties and their administrations. The teaching and research
programmes of the Lysenko "school" were to adopted without question,
and the more than questionable results were to be emphasized accord-
ingly. Results which, as it later turned out, had in some cases even
been forged. Other attempts were also made to influence teaching and
research at the Humboldt University. The autonomy of the University
was at risk. This resulted in a number of professors ceasing to co-operate
and giving up their posts. One of these was Hans Nachtsheim in
1949. His Chair for Genetics at the Humboldt University was not
fiUed again and the Institute was closed (Leussink et al. 1980: 857).
EUsabeth Schiemann, who had been Professor of Genetics and the
History of Cultivated Plants since 1946, also left the University in
1949. She had, it is true, reached retiring age but she had always
fought uncompromisingly against the adoption of Lysenko's doctrine.
In 1940, during the Third Reich, she had already been expeUed from
this University because of her opposition to Nazi poUcies.
Free University
In December 1948 in the western part of the city the Free University
was estabUshed. In 1949/50 the emigration of professors and students
from East-Berlin increased evenmore. The biologists found a new home

in the newly created Faculty of Mathematics and Natural Sciences at
the Free University (FU). Here a Chair for General Biology and Genetics
was established with an institute (Leussink et al. 1960: 862).
The agriculturists and horticulturists did not join the FU. They
founded in 1951 a Faculty for Agriculture (Landbau) at the Technical
University (Technische Universitat), which for some time had its
institutes and administration in Charlottenburg (West-Berlin). It must
be mentioned that 11 of the 27 institutes belonging to the Agricultural-
Horticultural Faculty of the Humboldt University were located in the
western part of the city, mainly in Berlin-Dahlem. There the construc?
tion of a new faculty could be undertaken under favourable conditions,
in other words with good institutional arrangements, as far as was
possible in the post-war time. For example substantial war damage
existed in Dahlem at the Research Institute of Heredity and Breeding.
How the development of this institute ensued since its foundation, will
be discussed in detail below.
Humboldt University (GDR)
The lack of staff at the Agricultural Horticultural Faculty could be

soon overcome. However, it was harder to replace the lost institutes.
In 1956 the faculty again disposed of 27 institutes (Skibbe & Gocht
1956). Among them, like one in West-Berlin, an Institute of Heredity
and Breeding Research exists. It was opened in 1953 outside the city
in Blumenberg. The first Director was Alois M u d r a. He was summoned
from the well-known Institute of Breeding Research in Miincheberg
(GDR). This Institute formerly belonged to the Kaiser Wilhelm Society
and in 1951 had been incorporated into the German Academy of
Agricultural Science of Berlin (GDR). Mudra's research concerned
practical breeding's purposes. In the foreground stood the work on
corn, sunflower, and oil-pumpkin. Next to that was the accomplish-
ment of more precise experimental field trials on mathematical bases.
In 1955, after Mudra's emigrations, Karl Zimmermann became
his successor. In practical breeding he obtained great merits, and worked
for many years, also in Miincheberg. Here he made a valuable contri?
bution to the breeding of the alkaloid-free lupines (Siifi-Lupine) with
Englera7-1987 161
indehiscent pods, whereby fundamental genetic problems must be

solved. In his broad research and breeding works forage grasses played
an important role (Skibbe & Gocht 1956).
In Berlin in the late 50s with the existence of two institutes for
heredity research, which had one and the same name exactly, the sep-
aration formally took place, but the loyal attachment remained and
a direct exchange of thoughts was still possible at this time via mutual
visits. The separation became final in 1961 by the hermetic demarcation
of East-BerUn from West-BerUn through the erection of the Wall.
The year 1961 consequently is regarded as a caesura for historical
consideration on the development of the genetics in BerUn. From this
year on the deseription must be separate. Furthermore, I do not see
myself as competent to report on the history of genetics after 1961
in the eastern part of BerUn.
The Institute of Heredity and Breeding Research
The history of the Institute began 75 years ago, when in 1912

Erwin Baur became Professor of Heredity Research for the first
time in Germany. The founding and the estabUshment of the Institute
have previously been explained above as weU as primary Baur's
works and teachings. In the following the development of the Insti?
tute through its directors and other scientists (co-workers) wUl be
explained, whereby the most important research work wUl be dealt
with in detaU.
Erwin Baur 1875-1933
"That our cultivated plants and domesticated animals are thor-

oughly and systematicaUy studied and precisely analysed, is the
most important task of the science of applied genetics."
With this sentence from the first edition of his textbook, which
appeared in 1911, Baur defined one of the tasks of his research work.
In addition, he further realized that such analytical studies could only
carried out in an appropriate state institute. The actual breeding of new
cultivars and races should always be the task of the professional plant
breeder. Baur clarified this jobsharing with a comparision of the

institutes of chemistry and of the production of chemical goods in
factories: "Likewise in the days ahead the institutes of heredity research
and the practical breeder will begin to cooperate". He was supported
in this efforts by his colleagues, the agronomist v. R ii m k e r and the
botanist Wittmack. Wittmack was the first who had described
the wheat-rye-bastard (1899), which was discovered in 1888 by
Rimpau, and which today is named by taxonomists as Triticosecale
Wittmack (Hoffmann et al. 1985: 67). Baur's acquaintanceofNilsson-
Ehle and his visit to the Swedish Seed Association in Svalof were
extremely influential and their methods of research left a deep im?
pression.
In order to have more land available for his experiments on breeding,
Baur obtained a farm in Dahmsdorf (about 40 km distant from Berlin).
In cereal breeding he purposed his goal of breeding wheat for sandy
soils. He attached great importance to cross-breeding. Therefore
he collected an extensive assortment of different genetic types. In
1911 he wrote to Schweinfurth asking him to bring him samples
of African cereals. On his South American trip in 1930 he collected
with his co-worker Schick frost-resistant wild potatoes. He brought
some to Germany, one of it still is much used in cross-breeding today:
Solanum acaule (Hoffmann 1985: 212). The grapevine he wanted to
improve and to attain resistance against Phylloxera vastatrix through
cross-breeding with the American grapes. In Dahlem the breeding
garden was surrounded with wild fruit trees and primitive cultivars.
A "pear orchard" was installed. There are 14 genotypes of apples and
9 of pears saved by grafting that are now serving as gene reserves. As
well a special walnut tree still exists, which is frost-resistant, because
its first buds do not open until the middle of May, when frost no longer
threatens.
The extreme success in breeding alkaloid-free lupines must be
attributed to Baur's foresight. This task, completely solved by his
co-worker, von Sengbusch, in 1931, is a classic example of how
important theoretical reflections are in order for experiments to be
successful. Baur had started from "the law of homologous series"
and intentionally adopted the plan of breeding alkaloid-free lupines,
i.e. "sweet lupines". Only a few mutants were found, but within
a few years a large amount of seeds was available. With the inclusion
Englera7 - 1987 163
of an additional area for seed multiplication in the former Palastine

two harvests were obtained in one year. Baur's last candidate for
a doctor's degree, H.-J. Troll, had taken over this difficult assign-
ment and successfuUy solved.
The example of the "sweet lupines" shows how thoroughly Baur
concerned himself with the genetic fundamentals of the objects of
his research. Theoretical knowledge he obtained from the investigation
of mutants, in particular of the snapdragon. He researched the segre-
gation of the Mendel-factors, recorded linkage groups and estabUshed
chromosome maps. He was not interested here in wether the mutations
were recessive or dominant. In his opinion the latter category was
responsible for most of the basic differences which have led, in even
changing combinations, to the increase in the variabiUty of cultivated
plants and domestic animals. On the theoretical side he attached
importance to the fact that in test plants and test animals very strange
"homologous series" of mutations were always observed. These mu?
tations, which are famiUar to us in one species, frequently occur in a
homologous way in related species. In his textbook (Baur 1919: 293)
he pro vides a Ust of examples of "homologous mutating" in plants
and animals. These include the case of the domestication of the field
mouse (Microtus arvalis), during which essentially the same colour
variations are obtained by mutations as these known in other species
of mice or in the rabbits. He concluded ? taking another comparison
from chemistry of the homologous series of different chemical con-
figurations ? that in organisms homologous mutations have evolved
from homologous changes in the chromosornes. Thus, as early as 1919,
Baur had already postulated unequivocaUy the "law of the homo?
logous series", which was precisely formulated and proved in the case
of cultivated plants by a large number of examples by Vavilov in 1922.
In his first studies after 1900 Baur had also already worked on non-
Mendelian characters. Like Correns he beUeved that there are genetic
carriers outside the cell nucleus. In his investigations of green and white
variegated Pelargonium zonale he reaUzed that he has concerned with
periclinal chimeras which could be demixed. His work on the graft -hy?
brids of Laburnum and Crataegus led to the recognition that somatic
ceU conjugation (cell fusion) had not taken place there, as had been
assumed elsewhere.
Sexual species hybridization in the snapdragon finaUy led him to
engage on experimental studies in the field of evolutionary genetics.

In his textbook he dealt in detail with theories of speciation. He was
also very interested in zoological objects, especially domestic animals.
Baur was aptly described as an ardent supporter of the selection
theory by Ernst Mayr(1976: 351), one of the leading investigators of
the science of evolution, who studied in Berlin in the 20s (Harwood
1985, see also Baur 1919: 345). Baur did not want the empirical
findings from his experiments on the snapdragon to be understood to
be generally valid. He states that sometimes differences between species,
and in particular between higher systematic units, are not the same as
the sum of small basic differences caused by mutations. Nevertheless he
saw, for the time being, in the theory of selection the only serious
attempt to explain evolution.
Erwin Baur was not only an able and successful teacher and research
scientist.
He also had great organizing talent. At the end of thetwenties,
from his Institute in Dahlem, he felt the need to contribute on a larger
scale of practical breeding. Schiemann (1935) describes at length
what motivated him and how he was led to found the Kaiser Wilhelm
Institute of Breeding Research in Miincheberg in the Province of Bran?
denburg. The Institute was built in 1927 and inaugurated in September,
1928. Baur remained Director of the Dahlem Institute until 1931.
In 1927, however, he accomplished another, quite different task in
the field of organization: under his presidentship the Vth International
Congress on Genetics was held Berlin, the scientific and also the political
significance of which can not be overestimated. The names of highly
distinguished figures in the visitor's book bear witness to this. The
findings of four scientists dominate this conference. Vavilov gave a
lecture on his theory of gene centres. Muller reported on experiments
with x-rays in which he had attempted to produce mutations arti-
fically in Drosophila. Correns gave a basic paper on non-Medelian
inheritance. Baur presented his assortment of mutants from the
snapdragon.
To include reference to the time spent by Baur in Miincheberg
would be to go beyond the limits of this report. Only 5 years were
left to him to work at his new research post. He died of angina pec-
toris on December 2, 1933, at the early age of 59.
Englera7 - 1987 165
Hans Nachtsheim 1890-1979
In 1921, before the Dahlem Institute was completed, Hans Nachts?

heim came from Munich, when he had taken hisPh. D. in 1913 under
Richard Hertwig and was appointed to an university lecturer in 1919,
to Berlin to take up a post at the Agricultural College. Baur attached
great importance from a theoretical as well as from a practical point
of view, to the genetics of domestic animals. Nachtsheim, who had
untU worked with bees, Drosophila and annelids, chose larger animals
for his experiments. He also changed his methods by investigation
and analysis. From cytological investigations, preferentiaUy on sex-
determination, he went on to analyse the results of hybridization. He
began by carrying out studies on the heredity of pigs. Nachtsheim
found support for his work from the animal breeders at the neigh-
bouring institute. From 1921-1936 Carl Kronacher (1871-1936)
was Director of the Institute of Livestock Breeding and Domestic
Animal Genetics who has carried out intensive studies on the science
of domestic animal heredity since the rediscovery of Mendel's laws
(Skibbe & Gocht 1956: 41). In Dahlem he instaUed a "Race- and
Research Stable" for cattle, which is stUl of importance today. At
that time L. Heck (1934) "bred back into existence" the extinct
aurochs in the BerUn Zoo.
At present in the Institute of Animal Production, under the di-
rection of J. We niger, P. Horst and W. Schloter, breeding prin-
ciples are currently being developed to produce breeds which are
adapted to tropical conditions (heterosis, genotype environment
interaction) in cattle, sheep and goats, and the tropical relevance
of the major genes are being tesfed in the chicken. Work is being
done in the tropics and also climatic chambers in Berlin. The mouse
serves as a model to test stress reactions.
At the Agricultural College Nachtsheim found moreover two
other zoologists who were concerned with the science of heredity.
One was Ludwig Plate, an expert on the selection theory and a
Neolamarckist, the other, at Baur's Institute, was C. Kosswig, who
was working on the genetics of fur pigmentation in the rabbit and
on pigment formation in cyprinodonts (Zahnkarpfen), Xiphophorus
and Platypoecilus.
From the mid-twenties onward Nachtsheim also worked on the
Erwin Baur: looking far into the future with his penetrating gaze
V . r I
:ft
flMAr*^-:/%?<,*.
fal
***<?%*?
jo?*#VM'?v&.
'uS.
From the visitor's book of the Institute of Heredity and Breeding

Research. The heading was the work of Elisabeth Schiemann. Well-
known and famous names from all over the world can be found there.
Beatrice Bateson could be William Bateson's daughter. It is certainly
not his wife, whose Christian name began with A
fr*
'W^
Erwin Baur (centre) in his field, pleased with his grain crop, which he
is proudly showing his co-worker, H. Nachtsheim (right) and his
colleague from the Kaiser WUhelm Institute for Biology, R. Gold-
schmidt. Goldschmidt pleasantly lost in thought is playfully fingering
an ear of "sand wheat"
rabbit. He had found an ideal test animal for both investigations of the
genetics of evolution and also for the special analysis of heredity. The
history of the process of domestication could easily be followed and
even reconstructed. The inheritance of many economically important
features such as the quaUty of the fur, its colour or even the genetics
of malformations and of the well-known Parkinson's disease could
be explained. Gene combinations could be achieved specifically on
these analyses. The race formation was in harmony with the genetic
findings. Corresponding conclusions could also be drawn concerning
the gene effects. Homologous mutations could observed in various
domestic animals during the process of domestication. One result of
this research work on the rabbit was the book "Vom Wildtier zum
Haustier" (From wUd to the domestic animal), pubUshed in 1935. A
third revised edition was procured by Nachtsheim and his student
Stengelin 1977.
In his extensive breeding experiments Nachtsheim came often
across with sick animals, to which he devoted much attention, beginning
in 1934, with methodical hybridization experiments (see also Parkinson's
disease). He succeeded in defining epilepsy, and Pelger's anomaly
(leucocytes polymorphic nuclei) as hereditary phenomena. In other
cases (cataracts in rabbits) he diagnoses a genetic disposition and the
expression, the phenotypic abnormaUty, is determined by the in?
fluence of specific environmental factors. Nachtsheim sees in the
interaction of a genetically conditioned constitution with unfavourable
environmental parameters a basis for explaining the existence, even in
humans, of many congenital deformities. When a Department of Ex?
perimental Pathology of Heredity was founded at the Kaiser Wilhelm
Institute of Anthropology in Dahlem 1941 Nachtsheim became
its director.
The first leading woman research workers in the field of genetics
In a historical appraisal of the development of the Dahlem Institute

special mention must be made of the work of three women engaged in
research. They are amongst those geneticists who are not only famous
for their services to science but who were also important personaUties
who experienced the birth of the new science of heredity. Notwith-
Englera 7 - 1987 169
standing their commitment and hard work it was not easy for them
as women at the beginning of the 20th century in Berlin to complete
their academic training. Elisabeth Schiemann (1891-1972) aptly
summed up the situation when she wrote (Leussink 1960: 845);"ghis
in Germany who strove to achieve more than an average education and
to follow an independent career, and who were not artistically gifted,
were actually left no other choice than to enter the teaching profession,
which was normaly the culminating pront of three years at a teacher's
training college". It was possible in Berlin for a woman to enrole at
the Friedrich Wilhelm University for the first time (later than in other
German states) in the winter term of 1908/09. Whether or not they
were allowed to attend lectures, however, was still left to the personal
discretion of the professors. The scientists, however, were not so hard
on woman students as members of other faculties. Once she had been
able to enrole in 1908, Elisabeth Schiemann became one of the
first woman students in Berlin. She had, however, already attended
scientific courses before as an occasional student. From Baur, who
was at that time still teaching at the Botanical Institute of the Fried?
rich Wilhelm University, she soon received a subject for a dissertation
and was "at long last able to start working independently". She then
began to grapple with the problem of mutations in Aspergillus niger.
She took her doctorate in 1912 and became B aur's scientific assistant
in 1914. Thus the ranks were opened to woman scientists and she
soon received reinforcement from Luise vonGraevenitz and Emmy
S t e i n. In the early years E. Schiemann's responsibilities were mani-
fold. Erwin Baur was obliged to serve as physician in the German Navy
Department, and organisational duties at the Potsdam provisorium
(see above) made scientific work extremely difficult.
When research in Dahlem was allowed to continue E. Schiemann
helped to make up the linkage groups on the snapdragon. She contri-
buted to the publication (Baur et al. 1929) which appeared on this
subject. Information on the investigations of the evolution of culti?
vated plants, which she had already begun at that time, is cited above.
Mention need otherwise only be made here of her work as senior
assistant and university lecturer (qualifying examination: 1924) and
her appointment as an associate professor in 1931. Lang (see his
contribution in this volume) has reported at length on E. S chie?
rn an n's career. He also elaborates on the situation which led her to
leave the Institute in 1930. Details of her research work after the World
War II wUl be given separately.
Emmy Stein (1879-1954), who came from Dusseldorf, trained
to become a gardner at the Horticultural School in Marienfelde in
Berlin. She studied at the University of Jena and took her Ph. D. in
botany in 1913 under the supervision of Ernst Stahl. During the World
War I she joined Erwin Baur, was also, therefore, in Potsdam, where
she worked as scientific assistant together with E. Schiemann and
Luise von Graevenitz. After 1923 she worked in Dahlem on search
into mutations. She tried, on the snapdragon of course, to induce
mutants by irradiating germinating seeds with radioactive rays. For this
purpose she used a radium compound with very "high energy" 7-rays.
These 7-ray emitters, the handling of which was not exactly without
risk to the person concerned, continued to be used at the Institute
until after World War II. They were not finally handed over to the
authorities until 1961.
One of the important results of many years of intensive research,
relating to analyses in fields of genetics, cytology and the physiology
of embryonic development (ontogenetic development) was the creation
of complex mutations in somatic cells, which manifested themselves
phenotypically as a cancerous proUferation of tissue, as so-called
phytocarcinoma.
This proUferation did not necessarily prove to be hereditary, could
however be passed on via vegetative propagation as a "Dauermodifi-
cation" according to Jollos' definition (1935). FinaUy, with later
generations of clones it was possible, however, to select mutants with
monofactorial inheritance for tissue degeneration. The analyses proved
that recessive genes (canc and cai) had to be involved, which were
passed on via the gametes. In another case it was possible to isolate
the dominant gene K for extremely cancerous radiomorphosis from a
complex mutant.
In 1940 Emmy Stein moved to the Kaiser WUhelm Institute for
Biology in Dahlem. There she worked under Fritz v. Wettstein with
her radium mutants on the hormonal influence of scion on its stock.
In 1948 she went with the department when it moved to West
Germany, where she worked until her death in 1954. In her last pubU-
cation on the growth of the cell nucleus and chromosome proUferation
she dealt with the occurrence of endomitosis in conjunction with the
Englera7 - 1987 171
pathological development of organs and thus touched on general

questions concerning the development of cancer.
Of the three women who, once they had left school, set out bravely
and with determination to take up a career via a long course of uni?
versity study, Paula Hertwig (1889-1983) was to a certain
extent
at an advantage in that she was able to start immediately on experi?
mental scientific work at the institute of her father, Oscar Hertwig.
She had also the advantage to be a BerUner born as late as 1889, and
the formal restrictions regarding university entrance for women had
akeady been Ufted when she took her school leaving examinations. It is
nonetheless surprising that her first scientific work on the cell division
of Ascaris, by means of irradiation with radium, was published as early
as 1911. In subsequent years other works were to follow, which reported
on findings relating to the germ cells and embryos of amphibians and
fish, which had been exposed to irradiation with radium. In 1916 she
presented her dissertation. She had been influenced not only by the
work of her father and brother Gunther but also by the courses given
by B a u r, which she had attended. It became clear that from this would
follow a career in research, dedicated to the science of heredity. Paula
Hertwig followed this course throughout her long and successful life.
She began to work with Erwin Baur before the Institute in Dahlem
was completed in 1922/23. She thus contributed to the reputation and
authority of the young woman scientists already working with him.
Since Hans Nachtsheim had already begun to work with Baur
in the early 20s the importance of animal genetics increased. For
Paula Hertwig mice and rats acquired the same importance that
rabbits had for Nachtsheim. She was, however, also responsible for
genetic analyses with chickens. She carried out extensive radiobiological
investigations, performed heredity analyses on the house mouse,
and also worked out Unkage groups. It was possible, for example,
to trace the pathological heredity phenes induced by means of radiation,
e.g. semi-steriUty, over many generations (Hertwig 1940). Radiobio-
logy developed during this period in BerUn into a branch of genetics
and became increasingly important. Basic investigations of the causes
of mutations were carried out by Timofeeff-Ressovsky (1947).
It was more likely to be possible to draw conclusions about the
effects of radiation on man by using the mouse as a test-animal to
determine the radiation dose and subsequent genetics damage than
by basing results on the genetics of Drosophila. Paula Hertwig's

research work on mice was taken up, confirmed and continued by
many other scientists. The new discovery that in the spermatogonia of
the mouse mutations occur 10 to 15 times more often than in com-
parable Drosophila experiments with the same radiation dose was of
far-reaching importance. In a final study of the history of radiabio-
logical research Paula Hertwig left us not only a general survey of
radiogenetics but also a legacy, a warning of the threat to man posed by
radioactivity.
Paula Hertwig and Elisabeth Schiemann were, as professors
during the "Third Reich", confronted with the hostility of the ruling
powers of the day as a result of the integrity and liberal turn of mind
they revealed in their political lives. E. Schiemann was even banned
from the University in 1940. After the war women were once again
aware of where their duties lay when false doctrines emerged from the
biological theories of Lysenko. With clear arguments in discussions and
at university courses they warned students and younger colleagues of
false prophets. After the war P. Hertwig accepted a professorship
in Halle/Saale. She remained and worked there until she retired from
academic life. From this time I have a very pleasant memory of her as
an admirable research worker and teacher. The years of her retirement
she spent in Villingen in the Federal Republic.
HansKappert 1890-1976
Thanks to the efforts of B aur and his colleagues the Dahlem Insti?
tute had reached a position of world reknown by the time the former
left for Miincheberg. The short interim period until Hans Kappert took
over as professor and director of the Institute was bridged by the
authoritative powers and excellent leadership by the likes of Elisabeth
Schiemann, Paula Hertwig and Hans Nachtsheim. To assume
the responsibility of running such a prestigious institute was no easy
task. The way in which Kappert mastered it and fulfilled the ex-
pectations placed in him earnt him national and international recog-
nition. The visitor's book alone, in which the names of a number of
famous foreign scientists appear, particularly in conjunction with the
Vth Congress on Genetics, bear witness to this. The proportion of
Englera7 - 1987 173
foreign visitors, especially from Asian countries but also from America
and Europe, was high in the thirties too. However, scientists from USSR,
who had previously come in such large number, no longer appeared.
The disastrous poUcies of the Nazis finaUy east their shadow over
the exchange of scientific ideas and development of international
projects.
Hans Kappert's career can be described as foUows: in 1914 he
took his Ph. D. under Carl Correns with a dissertation from the
field of appUed genetics (the investigation of sugar-peas and other
varieties of peas: wrinkled and round types and their hybrids). Cor?
rens took him with himself to BerUn when he became Director of the
Kaiser WUhelm Institute for Biology! Kappert discovered there not
only his preference for formal genetics based on an exact, mathematical
approach to the subject but also for appUed genetics in the field of
breeding research. After he had grappled for 10 years with the practical
problems of breeding - first in the Institute of Bast Fibre Research and
then as Director of seed cultivation with the firm "Dippe" in QuedUn-
burg ? he had aU the necessary qualifications which enabled him as
Baur's successor to maintain continuity in the work and Une of
research already established. Kappert carried out basic research on
other species than those by Baur. Analyses of Unkage groups were
carried out on Matthiola incana. The analysis of tetrads in Salpiglossis
confirmed that all the germ cells coming from one pollen tetrad can
function efficiently. Thus the conclusion reached by zoologists, derived
from the manner of sexual inheritance in hermaphrodite animals,
namely that only two efficient gametes are produced from one germ
cell tetrad, had to be rejected as not appUcable to hermaphrodite
plants. Investigations on dioecious plants to determine the practical
use of only one sexual partner, as with Asparagus, the male plants
of which yield better, were aimed at developing suitable methods
of breeding and selection for the practical breeder. Studies on the
inbreeding-heterosis problem (rye, radish) or on the segregation of
heterozygous polyploids (Cyclamen) were as much a part of Kappert's
work as investigations of lethal factors (Matthiola) or of the inheri?
tance of quantitative characters (the size in tomatoes).
Of the research findings which were of importance for the practi?
cal breeder two examples should be given by way of clarification.
A breeder was unhappy with the considerable variations in colour
of a released pink Cyclamen-cultivar, which ranged by propagation

through seeds from very pale pink to dark red. The analysis of the
heredity showed a segregation of this tetraploid cultivar, which had
originated as a duplex type into 5 genotypes, as is normally to be
expected in the intermediary inheritance of a 4x-heterozygous geno-
type. It was possible to recommend the breeder to handle the problem
of maintaining this cultivar by continually crossing simplex and quad-
ruplex types. - A very neat solution was found to the problem of how
to breed a throughout double Matthiola economically, which has
become completely sterile as a result of an abundance of corolla leaves
producing double flowers. This even amazed the experienced breeder
Schreiber, who came from a large firm of flower growers, when in 1938
in Dahlem he was able to convince himself of the perfect functioning of
a method of maintaining such a cultivar which had been developed
there. Kappert had succeeded in combining heterozygous SC/sc
genotypes, which had simple corollas (S) and dark green leaves (C),
with a pollen sterility factor which is absolutely linked both of the
dominant alleles (SCX-link age). Of the male gametes only the sc-pollen
are efficient and these fertilize the SCX- and sc-egg cells, so that the
progeny of SCX/sc heterozygotes are always either dark green with
simple corollas (SCX/sc genotypes) or light green with double corollas
(sc/sc genotypes), and they occur in a ratio of 1: 1. When they are
transplanted the dark green seedlings are put into the seed-bed for seed
production, and the light green ones in the flower-bed to be sold. -
Kappert himself referred to his field of research as "applied and
theoretical genetics in particular the studies of the problems of plant
breeding". The precise formulation of research aims, approved methods
and the application of mathematical principles were characteristic of his
research work. The fundamentals of Mendelian genetics were consol-
idated by his findings, not least by his explanations of exceptions to the
rule.
As a teacher he was strict but fair. He even examined students
in air-raid shelters. He founded his own "school" and those who
took their doctorate under his supervision still proudly call themselves
"Kappert"-students. Two editions of his textbook "Die vererbungs-
wissenschaftlichen Grundlagen der Ziichtung" (1953) (The funda?
mental principles of the science of heredity in breeding) have appeared.
It is not easy to read but it is an excellent reference book for lectures
Englera7 - 1987 175
and exercises. Kappert's special heritage deserved our deep gratitude.

He has left us with an historical appraisal of genetics which presents
the experience of four decades. His students, W. H o r n and G. W r i c k e,
took his manuskript, added to it a biography and a comprehensive
bibliography and pubUshed it after his death (Kappert 1978).
To complete the picture of Hans Kappert's character it should
be said that his dedication to his Institute and his coUeagues was parti?
cularly apparent in the last years of the war and thereafter. He made
a considerable effort, for example, to relieve human suffering by using
the institute's breeding garden for the production of food.
He was opposed to obscure and totaUtarian teachings and did not
merely ignore the results of the so-caUed "re-education experiments",
carried out by the Lysenko school, he interpreted them correctly and
exposed the mistakes which had been made in the execution of the
experiments (Kappert 1953: 309). This was a great contribution to the
fact that, especiaUy among students of agriculture, the Lysenko doc?
trine found no support. Disaster was also thus averted as far as breeding
and practical plant cultivation were concerned. In the USSR, on the
other hand, it was not only genetic research which suffered a setback as
the result of Lysenko's biological theory but also practical breeding and
the effects on the development of production in agriculture were also
very negative.
Walter Hoffmann 1910-1974
In 1958 Walter Hoffmann became Kapp ert's successor. He was

a very versatile scientist and had akeady acquired a reputation in
serveral institutes (Roemer & Rudorf, Handbook of plant breeding
1961, vol. 5: 204, 264). He had taken his Ph. D. in 1934 at the Bo?
tanical Institute in Heidelberg with a dissertation on the subject of
sprouting behaviour in mature cereal kernels, especiaUy in barley
(Auswuchsverhalten beim Getreide, besonders bei Gerste). Soon after
he moved to H. Kuckuck's department at the Kaiser WUhelm Institute
for Breeding Research in Miincheberg (Hartmann 1936: 281) and from
there in 1942 to the department of fibre plant breeding in Mahrisch-
Schonberg in the Sudetenland, which was then part of the German
"Reich". After the war he was Director of the Institute of Plant Bree-
ding at the Agricultural Faculty of the Martin Luther University in

Halle/Saale until his appointment in Dahlem.
In the third and final part of his professional career in Dahlem
Hoffmann devoted his time to work which can be classified as
belonging to the fields of breeding methodology and cultivated plant
evolution. His aim was to develop improved methods of selection
for the practical breeder, taking into account the laws of genetics.
In order to obtain a higher rate of gene exchange in linkage groups
more emphasis was to be placed on backcrossing methods. Not only
one partner was to be used, however, but in Bi (Fi) it was to be a
quite different one and in B2 (Fi) a third was to be involved, so that
the combination possibilities might be better exploited. It was an
advantage, in the artificial creation of mutants, to subject the process of
selection to "one-kernel-populations" (Einkornpopulationen), i.e. of
each mature M\ plant one seed should be taken for M2 and this pop-
ulation should be treated a second time (Hoffmann et al. 1971: 149).
Allogamic populations are particularly suited for this purpose. Hoff?
mann provided a model example with the radish (Raphanus sativus
var. sativus).
According to a more recent definition breeding is referred to as
evolution directed by man. In this sense Hoffmann was concerned
in breeding research with questions of evolutionary genetics. In the
development of new fodder crops used as green fodder and/or green
manure importance was attached to creating greater variability with
regard to improved adaption for the cultivation in intercropping between
two main crops. Hoffmann achieved great success with a poly-
ploid hybrid from cross between Brassica campestris subsp. pekinensis x
B. campestris subsp. oleifera (Chinese cabbage x turnips) with 2n = 4x=
40 chromosomes. Since 1969 a cultivar known as "Perko" has been
released, which is still cultivated today at home and abroad. Hybrid
forms which are the result of a cross between species and genera of
Triticum and Aegilops also add to the increased variability in the
range of cultivated plants.
To ensure high fibre production in combination with good seed
production Hoffmann also devoted his attention to the special
genetic question of sex-inheritance in the dioecious hemp (Cannabis
sativa). He bred a so-called "simultaneously maturing hemp" (gleich-
zeitig reifender Hanf) which only contains female types concerning
Englera7-1987 177
growth habit, but has a different proportion of male flowers (in odd
cases hermaphrodite flowers are found). In the inheritance of dioecy it
was assumed that it is based on a heterogametic/homogametic system.
This was supported by the proof of the existence of an XY pair of
chromosornes (male determining). In the creation of monoecists in
hemp which have very different male and female types as far as their
growth habit is concerned, polymeric genes, which are evidently located
in the autosomes, play an important part. With this hypothesis Hoff?
mann made a valuable contribution to the general theory of sex-in?
heritance (Roemer &Rudorf l.c: 251). It is in accordance with Correns'
basic findings on subdioecists and complements the findings of Hart?
mann and Goldschmidt on sex-determination.
Like his predecessors, Kappert and Baur, Hoffmann can also
claim to have a "school". Many of his students were or still are success-
ful abroad and are often involved in development aid projects. He
provided many Ph. D. students from Third World Countries with neces?
sary equipment for their professional careers and thus contributed
to a transfer of knowledge to these countries. I am grateful to have
been one of his first Ph. D. students in HaUe/Saale.
Interim period ? managing directorate 1974-1984 at the Institute of

Applied Genetics
As its last full professor Hoffmann upheld the tradition of the

Institute with his research and guaranteed success and continuity in
its development and achievement. In applied genetics it is an advantage
if a certain Une of research is continued over a longer period. For 60
years, until Hoffmann's death, the Institute only had 3 different
directors, each of whom had determined the Une of research.
Since the ref orm of the university system at the beginning of the
seventies the Berlin universities have no longer had faculties, but
instead smaUer units, known as subject departments (Fachbereiche).
The directors of institutes are elected to manage their affairs for a
limited period. Decisions are made within the institutes by managing
directorates.
At the time of this ref orm regulations were introduced Umiting
the agricultural and horticultural degree courses. These measures were
responsible for the renaming of the Institute which, in 1972, became

known as the Institute of Applied Genetics and was affiliated to the
Subject Department of Biology of the Free University.
After H o f f m a n n's death in 1974 four working groups were set up
by the professors then in office to complete Hoffm ann's unfinished
work, to co-ordinate other independently carried out studies and to
continue the line of research hitherto pursued. The names of the groups
give an idea by the work which they took on; greater specialization within
the Institute became apparent; the work groups were devoted to: the
cytogenetics of plants, endogenous conditioned causes of mutations by
Gertrud Linnert (1978); biochemical selection criteria in rye, barley
and lupines, gene reservoirs in situ by W. Plarre (Hoffmann et al.
1985: 137, 185); synthesis and genetic analysis of new cultivated
plants, in particular Triticale-types by K. D. Krolow (Hoffmann
et al. 1985: 67), and breeding methodology (composite crosses) and the
analysis of specific features in wheat, peas and rape by W. Oden-
bach (Hoffmann et al. 1985: 51).
The increase in the number of members of staff at the time of the
move to the Free University had a positive effect on the biology degree
course. As the research programmes, which deal exclusively with prob?
lems of applied genetics, have undergone no fundamental changes they
help considerably to extend the framework of possible subjects for
first degree dissertations and Ph. D. theses in biology. The study of
economic and cultivated plants has acquired greater importance than it
used to have for all students of biology, including those training to
become teachers. This means that qualified biologists are now able to
choose from a wider range of posts.
Unconventional methods of plant breeding since 1985
A further development in genetic research in the service of plant

breeding was introduced with the appointment as Professor of Otto
S c h i e d e r who moved to Dahlem from the Max Planck Institute
of Breeding Research in Vogelsang, Cologne. The committee respon?
sible for making this appointment had advertised a post in the field
of unconventional plant breeding. With my retirement from academic
lifte it became possible to re-introduce the former positions of full pro-
Englera7 - 1987 179
fessor, which are now grouped together and referred to as C4 ? position.

Otto Schieder thus became the successor of Walter Hoffmann.
In plant breeding it wUl be necessary in the future to create new
basic material for selection by different new kinds of methods. Solutions
to problems are being sought with the help of traditional (conventional)
methods (sexual hybridization, mutation induction). The expanding
of genetic variabUity and the introduction of entirely new kinds of
gene-combinations require more and more the use of speciaUzed la?
boratory techniques. Protoplasts which have been won from cell or
tissue culture are now, for example, being isolated from various plants
with the help of biochemical and/or biophysical agents and fused with
one another. Work is at present also being done to improve these
methods, so that the number of fusion products can be increased. The
peculiar features of the physiology of development of the regenerates
are being investigated. Cytogenetic structures still have to be ascertained.
Using tissue cultures it is now possible to clone new kinds of geno-
types, which are easier to handle. Their suitabiUty in this respect for
use in practical breeding is being tested on a large scale. Single chromo?
sornes or parts of genomes can be removed from fused or other cells
using micromanipulators. Moreover it is possible to transfer genetic
structures from donor ceUs to other species by means of vectors (plas-
mids, viruses).
It can be concluded from the research findings to date that investi?
gations should be carried out on the types of plants which are most
suited to this kind of genetic engineering. These include genera, for
example, in the family of Solanaceae, such as Nicotiana, Petunia,
Solanum (potato) or Datura (Schieder 1987). It could be of greater
practical significance to increase the genetic variance of staple food
plants such as cereals or to produce genotypes with a specific aim, for
example, ones with multiple resistance to diseases, pests and herbi-
cides. It is obviously not only appropriate techniques (methods of
gene-technology), which are needed to manipulate cereals, there is also
a general lack of knowledge of processes involved in the physiology
of development in the regeneration of caUi to make them complete
plants. Gramineae in connection with evolution clearly belong to the
"younger" plant families.
With another group of considerable economic importance, the
Brassica species, tissue cultures in Petri-dishes can be used for the
above-mentioned large-scale tests. It is possible to select a number

of genotypes which are resistant or sensitive to the toxins of the Phoma
fungus. This was very well demonstrated by the work of Maria Dolores
Sacristan. With the Brassica species it is also possible to breed new
variants of economic importance via sexual hybridization, and even
if they used directly they may serve as further partners for
cannot
cross-breeding. Other new hybrids which have recently been produced
at the Dahlem Institute were obtained by crossing Brassica carinata
(2n = 34) and B. napus (2n = 38) with 2n = 36 chromosomes. They
include lines which are resistant to Phoma and which were backcrossed
with rape so that the resistance could be transfered (Sacristan & Gerde-
mann 1986).
Institute of General Genetics
Shortly after the foundation of the Free University (Freie Universitat)

in December, 1948, an institute of genetics was set up in Berlin. Hans
Nachtsheim (1890-1979), who had given up his professorship
at the Humboldt University in East Berlin, became its first Director
in 1949 (see above and below). After his retirement from academic
life in 19 5 5 Herbert Liiers (1911-1979) was appointed as his successor
and was Director until his retirement in 1976. Liiers achieved great
success not only as teacher and research worker but also as Dean and
Rector (1963-1965) of the Free University.
After obtaining his doctorate in 1934 Herbert Liiers became
scientific assistant in the genetic department of the Kaiser Wilhelm
Institute of Brain Research in the suburb Buch (see below), where
questions concerning population genetics in insects and mice were
being investigated and mutation research carried out (Telschow 1942).
He remained there with short breaks until 1951, finally in the position
of Head of Department, then worked at the Humboldt University
until 1953, when he became Department Head at the Max Planck
Institute of Comparative Biological and Pathological Heredity in West
Berlin. There he took over the direction of the Institute of General
Genetics at the Free University in 1955.
Whilst working in Buch Liiers made valuable contributions to the
study of Drosophila genetics, involving amongst other things organ
Englera7-1987 181
transplants. The necessary techniques he had learnt during a long stay

in Paris 1937. At the beginning of the fifties he went on to research
questions of hereditary damage caused by cytostatic agents, used in the
treatment of tumours. He finaUy carried out a series of investigations
on the effects of chemical and physical mutagenic agents not only in
Drosophila but also in ceU tissue or in vitro in human leucocytes. In
interdisciplinary research Liiers proved to be highly suited to team-
work and showed, as for example in the study of complex questions
such as that of mutagenicity, how problems can be best tackled jointly
by introducing a division of labour. He worked together on questions
of human genetics with F. Vogel, a speciaUst in this field then working
in BerUn. Liiers' student (Ph. D.) and colleague, K. Sperling, later
concerned himself solely with research into human genetics.
Liiers had dealt extensively with his colleagues (1974) with general
problems such as "Genetics and Evolution Research". In the final
years B. E. Wolf, H. J. Belitz, H. Nothel, G. Obe, H. and G.
Rohrborn and K. Sperling worked with him. Studies on chemical
and physical mutagenic agents and on stress factors in Drosophila have
produced important findings. General conclusions can be drawn from
the result of the expression of exactly locaUzed lethal and resistance
factors about the physiological function of such genes in other or?
ganisms. Conclusions from "induction r6sum6" ? production of facts
to prove a general statement ? represent theoretical knowledge of far-
reaching epistemological importance. The interspecific differences in
radiosensitivity, which can be observed in species of higher organisms
and which reveal a genotypical dependence in cases of damage caused
by radiation, are, for example, also worthy of mention (Nothel 1970).
From my personal experience of Herbert Liiers I must add that
on the occasion of the farewell party given on his retirement from
University life he approached me with a suggestion which honoured
me: that we should together undertake the task of writing the history
of genetics in Berlin. I was to be responsible for the "botanical section"
and he was to taken on the part devoted to zoological genetics (in?
cluding human genetics). This idea had its roots in the fact that in the
first decades of the twentieth century highly significant contributions
to the worldwide development of the young science of genetics had
been made in BerUn. At that time, in 1976, this was not clearly re?
cognized internationally. It can, however, now be proved by the latest
publications of Harwood (1984, 1985) and Saha (1981). The con?

centrations of institutes and famous scientists made Berlin, at least
at the time of the Vth International Congress, into the Mecca of the
geneticists (see above). For Herbert Liiers, however, it was also a
determining factor that we had both direct contact and a close re?
lationship with many important figures, and had worked or were still
working in their institutes. They had been our teachers and we had
experienced how the science of heredity had been influenced ideo-
logically by a variety of schoolsof thought and under these circum-
stances we had been able to follow the examples set by scientists of
integrity. Last but not least, in 1976 it was still possible to have a
clear view of the development of the history of genetics in Berlin.
Herbert Liiers' death was untimely, his memory and his ideas
are still with us. I can unfortunately only realize one part of this
project, however. To establish links between botany and zoology,
between basic and applied research, I must "intrude" into the field
of research of my colleague Liiers, but I cannot produce a com-
prehensive account. I trust the reader will show understanding for
these shortcomings. I am therefore only able to report briefly on
recent developments in the Institute which Liiers administered until
1976 and on the present organization of research projects.
The Institute of General Genetics, which is run by a managing
directorate, is divided into three departments: Anthropology and
Human Biology (Carsten Niemitz); Biophysicsand Radiobiology
(Ingolf Lamprecht, Wolfgang Laskowski); Genetics (Giinter
K o r g e, Horst K r e s s, Horst N 61 h e 1, Giinther O b e). As in the past,
work on the study of plants is still not carried out at this institute.
Institute of Human Genetics
In 1976 this Institute was founded as part of the Free University

and became affiliated to the University Hospital in Charlottenburg in
Berlin. With this third institute at the Free University the third im?
portant branch of the science of heredity, the study of human heredity,
was now represented at the University and assumed a position of equal
importance with those of botanical and zoological genetics. This devel?
opment which saw a division of teaching and research into these fields
Englera7-1987 183
was the inevitable consequence of the studies and knowledge which had
been accumulated during the preceding decades. Our thanks are due to
those responsible for the organization and decision-making involved
in the creation of this tripartite system in the field of teaching as weU
as that of research and also for the fact that it at the same time took
care completely of all concerns and interests of the subjects of gene?
tics. During the "Third Reich" the study of human heredity was
severely exploited. The term "eugenics" was used, and an official
pubUcation bore that name. The subject was allowed to degenerate
into the pursuit of an one-sided racial doctrine. It must be remembered,
however, that scientists such as Erwin Baur, had nevertheless rightly
recognized the importance of the science of heredity as a means to
further the weU-being of the whole of mankind. As Renner (1935)
wrote, however, in an obituary for Baur, the laws of heredity are
shrouded in the threefold mystery of the long duration of generations,
the smaU number of offspring and the large number of chromosornes.
For the pure and the applied science there was not sufficient know?
ledge of human heredity at that time to make it possible to carry out
famUy hygienics successfully. Today so much information is available
that this field is given priority at the Institute of Human Genetics in
BerUn and genetics is applied in the diagnosis of hereditary diseases.
About 2,500 investigations of chromosornes are carried out each year
(Sperling: verbalcommunication).
This Institute is administered by a managing directorate. Karl
Sperling, who studied under Liiers, was appointed full Professor
in 1977. Three other professors are also attached to the Institute:
Joachim K1 ose, Jiirgen Kunze und Reinhard Luhrmann.
In research priority is given to cytogenetic questions, such as the
question of chromosome condensation. Great importance is also at?
tached to the analysis of the heterogenity of somatic cell hybrids. The
important research work done by Sperling & Marcus (1984) on
the mapping of genetic activity on mammaUan chromosornes, by
K1 o s e on biochemical questions and by K u n z e on clinical genetics
has brought the Institute international acclaim. This was also the
reason why, although it had only existed for ten years, the Vllth
International Congress on Human Genetics was held in 1986 in West
BerUn and the Institute was entrusted with its organization. 2,000
scientists attended this conference.
Kaiser Wilhelm Institute in Berlin, later known as the Max Planck

Institute
With the proclamation of the foundation of a Kaiser Wilhelm Society

for Advancement of Sciences, which Kaiser Wilhelm II read in 1910 on
the occasion of the 100th anniversary of the inauguration of the
Friedrich Wilhelm University in the assembly hall of Berlin University,
the way was paved for the future of scientists and scientific develop?
ment and prospects were also opened up for the generous support of
free research. The constituent assembly of the Society took place
in January 1911, in the conference room of the "Akademie der Kiinste"
(the Berlin Academy of Arts). The first paragraph of the constitution
states: "The purpose of the Society is to promote science, in parti?
cular by setting up and maintaining scientific research institutes"
(Staab 1986: 40). There were various reasons for setting up research
institutes outside the University. These included the fact that the burden
of teaching left little time for research, also the fact that equipment
was becoming increasingly expensive. Moreover there were signs that
it was becoming difficult to keep up with research being done abroad.
These arguments apply equally today and were also quoted in a similar
fashion after the Second World War by the Max Planck Society (MPG),
which was set up to succeed the Kaiser Wilhlem Society (KWG) (Staab
1986: 38).
It was and still is the duty of the institutes (KWI and MPI) to conduct
research programmes in the interests of the common good, to con-
centrate mainly on basic research and, in particular, to introduce new
lines of research.
Kaiser Wilhelm Institute for Biology
As early as 1912 it was decided to found an Institute for Biology

as part of the Kaiser Wilhelm Society (see also Schnarrenberger in this
volume). According to M elchers (1961: 115) the history of the
Institute can with the help of "the most important dates be taken from
the files more or less completely intact". Considering more closely the
period around 1912 Melchers refers to the difficulty of viewing
history objectively. As far as the state of research in genetics, the
Englera 7 - 1987 185
physiology of development and other disciplines is concerned it is

possible, but when we are dealing with the people involved, the appoint?
ment of individuals, why, for example, a certain scientist was chosen
and not another, then it becomes more difficult, especially, as in the
case of the Kaiser Wilhelm Institute in Dahlem, when the structure
of the Institute is not intended to be hierarchical. The Institute was
opened in 1914/15 with 5 independent departments. C. Correns
took on the department of the biology of plants, together with the
direction of the whole Institute. He was succeeded in 1934 by F.v.
Wettstein. H. Spemann was responsible for the matter of animal
development, later O. Mangold, R. Goldschmidt for the science
of animal heredity, later A. Kiihn. Research into plants and animal
protists was in the hands of M. Hartmann, who was later succeeded
by H. Bauer. O. Warburg was in charge of general physiology.
At this state I must limit this account: in order to remain within
the prescibed framework. My attention should be directly firstly
towards genetics research, in particular in the field of botany, and
secondly to the period from the opening of the Institute in 1914/15
to its transfer to West Germany between 1943 and 1945. As I am not
able to go into great detail ? information can be obtained from the
Kaiser Wilhelm Society annual reports (in particular the writings of
Hartmann 1936, 1951 and Melchers 1961) - a choice has tobe made.
Melcher's (1961)
report contains some very critical passages
on the projects selected in the initial years and refers to
research
failures in the field of genetics research, for example, to the fact that
our "German pioneers of genetics" considered the question of the
analysis of the localization of genes much too early as completely
settled. No "Morgan School" emerged in Germany. Did this apply
in general or just to the Kaiser Wilhelm Institute in Dahlem? Were
there reasons for it? In the next section the question will be dealt
with at the end in more detail. My main aim, however, must be to
concentrate on the genetic research, which our Dahlem scientists
from the Kaiser Wilhelm Institute were engaged in until 1945 and which
did, after all, earn them international recognition for their services to
science, and to the understanding of the processes of inheritance.
Carl Correns 1864-1933
"Now it is clear from the outset and generaUy agreed even among
the most enthusiastic supporters of the nucleus monopoly that
plasma is necessary for the production of every MendeUan char-
acter" (Correns 1928).
When he was appointed Director of the Kaiser WUhelm Institute in

1913 Carl Correns already had a successful career behind him. Of
the 3 scientists who rediscovered MendePs laws his part was the most
important because of the very extensive and sound interpretation of
their results. His colleague, Emmy Stein (1950) was not the only one
to draw attention to this. (Gaestel & Niirnberg 1986 deemed worthy
Correns' work and life at great length.) It is also evident from a
more recent American pubUcation which points out in particular
that we should speak not of "rules" but of laws of inheritance. It can
be clearly demonstrated how this knowledge was obtained, how the
sequence of hypothesis - theory ? law emerged. This was Carl Cor?
rens greatest achievement (Monaghan & Corcos 1984).
Before Correns moved to Dahlem he had analysed a great number
of results of Mendelian segregations, i.e. chromosome-setssegregations -
he worked in all on about 340 plant genera ? but had also tackled
the problem of the role of cytoplasma in his studies of inheritance.
Although he was concerned to a large extent with research into genetics
of chromosornes, as in the investigation of the sex-inheritance of
dioecious plants (Melandrium), he began to devote more time to non-
MendeUan inheritance. It was partly his work on Mirabilis, which he
nicknamed "MiserabiUs" because of the trouble he had crossing them ?
one ovule in the ovary ?, and partly work on Pelargonium, which led
him to investigate the questions of extranuclear inheritance. Here
he is also concerned with the physiology of development in the inter-
action between cell nucleus and plasma, in which the green and white
colour of the leaves (variegation) plays an important part (among the
writings of Wett stein (1938) pubUcations on this subject by Cor?
rens (1919/20) are quoted).
To return to the sentence quoted above from the pubUcation of
1928 it should be mentioned that this was taken from the general
lecture which Correns held at the Vth Congress on Genetics in
1927 in Berlin. He went on here to explain that a character is formed
Englera 7 - 1987 187
without the involvement of a gene, but not without the involvement

of plasma. In this way he extends the idea of gene expression to include
certain genetic mechanisms which are localized in the cytoplasma. In
this study the role of the plastids as genetic carriers is also discussed.
The development of the ideas on cytoplasmic inheritance at this time
and in the period until 1945 was described in detail by Harwood in
1984.
Experiments in inheritance carried out on plant families with varie-
gated leaves were for Correns of prime importance in the under?
standing of the non-Mendelian inheritance. Besides the Mendelian
inheritance he finds cases in which the white colour is only passed
on by the mother (Mirabilis). Not all his results could be published.
A very important publication on non-Mendelian inheritance appeared
after his death in 1937 (Wettstein 1938), found among the mass of
the work he left, which Emmy S t ein (1950) put in order and Margaret
S. Saha (1981) more closely evaluated. In 1961 Renner referred to this
highly informative summary as a "canon for a great length of time".
The knowledge which Correns himself gained from this work
Wettstein (1938) described very subtly as follows: "The process
of inheritance became for him, at the point where others were often
no longer able to fathom the formal chromosome mechanisms, an
ingenious game with the forces of distribution and also the success of
the effect of this distribution . ..".
As mentioned above, during his career at the Kaiser Wilhelm Institute
Correns continued to carry out intensive studies on sex-inheritance
and sex-determination. He investigated above all the question of how
the regular sex ratio 1:1? refers to it as mechanical ? could be shifted.
He worked with varying amount of pollen, cut off the pistils, allowed
then the pollen to become old or exposed it to the effects of alcoholic
fumes. With the dioecious plant Trinia glauca (Umbelliferae) he in?
vestigated the death rate of the two sexes and found that the male ones
were more likely to succumb to an infection than the females. He saw
the possibility of purposefully influencing the sexes and warned of the
dangers which arise if these matters were put into the hands of man to
decide. He also asked whether it might be possible, even in human
beings, to disturb the hand of fate (Stein 1950) in this way.
Subdioecists, plants with no stricly marked separation of the sexes,
Correns was able to divide into the subandroecists and subgynoe-
?iîE^j^^2^3322a^^^-ia
A portrait of Correns, an impressive picture of the great research

scientist, the visionary and introvert
Englera 7 - 1987 189
cists. The former category only produces male offspring, the latter only
female. Among the subdioecists there are intermediary stages which
are referred to as intersexes and can be compared to the animal organisms
(butterflies) which Goldschmidt investigated simultaneously at the
Institute.
In order to show Correns as a man of his day in his style of
research, the effect he had and the way he coped with his work a
very colourful presentation used by Renner (1961) can be drawn
upon. In a very original account he compared the lives and careers
of the two great geneticists Bateson and Correns. During the same
period they both made a decivise contribution to the progress of
the young science. We have Bateson to thank for the fact that every-
where today we now nearly always speak of "genetics" and no longer
of "heredity". He coined and publicized this name for the new science.
Renner writes: "Among the leading geneticists none is more different
from the Bateson than the austere Correns." Symbolically he compared
the recluse Correns with St. Hieronymus accomplished by the lion
and Bateson with St. George slaying the dragon.
In addition to the comparison between Correns and Bateson,
one should perhaps make another one, namely between Correns
and Baur. Both worked at the same time, not far from one another,
and the achievement of both was great. One concentrated mainly on
applied genetics, the other did basic research without concerning
himself in the least with practical matters. Correns had nothing
to do with horticultural and even less with agricultural plant breeding.
In my research I found hardly any reference to meetings between the
two Dahlem scientists. However, they surely must have complemented
each other well! Baur with his snapdragon mutants and gene analyses
almost appears to be a "Morganist" and in the following years at the
Kaiser Wilhelm Institute in Miincheberg work was intensified in order
to produce a comparision, an outstanding example, for Drosophila
genetics, from the plant world. Correns' task was certainly the more
difficult of the two. At the Congress on Genetics he provokingly
expresses his doubts with regard to the nucleus monopoly in inherit?
ance, and emphasized the significance of cytoplasma as a carrier of
genetic material. This may, in my opinion, have been the result of a
tacit agreement. It did in any case earn for both institutes and their
directors prestige and recognition; the number of foreign guests greatly
increased in the subsequent period. Six years of intensive research were

left to Correns in Dahlem, for Baur in Muncheberg, the time was
no longer. Many followed eagerly in Correns' footsteps and worked
on cytoplasmatic inheritance. Amongst the botanists there was
Wettstein, Michaelis, Renner, Oehlkers etc, among the zoolo-
gists Goldschmidt, Nachtsheim and Kuhn. Harwood (1984)
gives a detaUed account of this. The continuation on a large scale of
Baur's work on the snapdragon ? and there are indications that this
was intented ? was probably not possible after his sudden death since
his coUeagues were not sufficiently well informed of his plans (Renner
1935).
Fritz von Wettstein 1895-1945
Wettstein was appointed first Director and Head of Department

as Correns' successor. He came from Munich, had been a scientific
member of the Kaiser WUhelm Society since 1931 and had already
made a name for himself in Berlin as Correns' young scientific
assistant, from 1919 to 1925. In 1923 he passed the qualifying exami?
nation to become Professor. During this interim period in Dahlem he
had set up for himself a very varied field of genetic research with ex?
periments in which he crossed polyploid mosses (Bryatae) included
crosses with different species and genera (Wettstein's pubUcations,
see Stubbe 1950/51). His work was later classified in the Kaiser WU?
helm Institute and divided into 3 main categories. They are concerned
with questions of the effects of genes (which involves plasmatic in?
heritance), sex-determination and sex-inheritance and finaUy speciation,
i.e. evolution of species in which great importance is attached to
polyploidization.
During the continuation of the work on mosses, which had begun
in the 20s, extensive morphological and cytological investigations are
carried out. With the help of a simple knack ? diploid stems of moss
capsules are cut up and stimulated to regenerate mossy cushions which
form gametogonia and are fertUized ? polyploid series can be produced
from an initially haploid form. The physiology of development can be
observed in the interaction between the number of genomes and the
cell size in the different polyploids: if the number of genomes increases
Englera 7 - 1987 191
at a ratio of 1: 2: 3: 4, the size of the cell increases in geometric progres-

sion, dependent on a constant measuring unit within a certain taxono?
mie group for each increase in growth. This is determined by individual
genes. With hybrid species the rate of growth is more favourable than
with crossed lines of one species with different ploidies. Wettstein
was the first to gain insight into the quantative interaction of the same
and different genes and its effects on cell growth and organ develop?
ment in such hybrid populations. Here a fact of far-reaching signifi?
cance became the more distantly related the crossed species
evident:
are, the more the plasma from the egg cells, as well as the different
genomes, is involved in character formation. Wettstein also worked
in this sphere on flowering plants. Encouraged by these findings and
other information on non-Mendelian inheritance he summerized
the entire mechanisms of heredity with the following formula: idio-
type = genotype + plastidotype + plasmotype. This definition can be
traced back to 1927. It was published by Correns in 1928 but did
not gain significance, in particular in the field of applied genetics
and breeding research, until the 30s and later. It can be varied without
difficulty to take account of new information on the effects of mito-
chondria and plastids; it has been included in German textbooks
(Hoffmann et al. 1971:33).
Via his experiments on mosses Wettstein's attention was of
necessity directed towards the question of sex-determination. In a
normally genotypically dioecious species (Bryum caespiticium), in
which haploid male and female mossy cushions occur, it was possible
to produce hermaphrodites by means of polyploidization using the
above-mentioned method. Different ploidy levels contained a dif?
ferent number of sex-determining factors for male and female; it
was also possible to establish the influence of the amount and the
time of the occurrence of male or female gametogonia.
As a result of the above-mentioned studies which, although they
had been carried out on a specific object, were nevertheless part of
two different fields of genetic research, Wettstein devoted more and
more of his time to questions of evolutionary genetics (publication
list, see Stubbe 1951). He was able to indroduce excellent new experi?
mental species: from the haploid/diploid, B. caespiticium, he produced
the diploid/tetraploid species, B. corrensii. Initially it was a victim of
the unfavourable giant growth habit, after ten years,however, perhaps
due to favourable gene mutations, a competive small cell form with a

constant number of chromosornes developed, Uke that which is also
found growing wUd (Renner 1946). In his work on the formation of
species Wettstein naturaUy also concerned himself with other
objects. Together with Stubbe he investigated the role of micromuta-
tions and macromutations. Here they turned again to the snapdragon
(references see Stubbe 1951). Their aim was to carry out experimental
evolution research. It is clear how far W et t st ein's ideas concerning
the questions of polyploids as a basic phenomenon of evolutionary
genetics reached, if we consider his final, very substantial pubUcation,
in which he discussed the significance of Hofmeister's alternation of
generations within different classes in the plant kingdom.
For Wettstein it was no coincidence that with the evolutionary
steps in the plant kingdom, the final Unks of which are the angio-
sperms, the diploid generation (sporophyte) has repressed the haploid
phase (gametophyte) more and more. He saw the explanation for this
in the fact that in the diploid state, when there are two genomes, reces-
sive alleles of different mutated genes can be enriched until by chance
recombinants with a selective advantage are brought together. These
can then spread quickly (Wettstein 1943).
The experiments on polyploids were to be continued on a large
scale with flowering plants. The war, however, the transfer of the
Institute to various remote corners and the concern about the fate of
colleagues made it impossible for him to work in peace ? and then
there was his health. He had to take on a long journey under the most
difficult conditions. He never recovered properly from a fit of pneu-
monia contracted in 1944. Already sick again in February, 1945, he
started his last journey to Tyrol to wait for and see the end of the war
at his home in Trins. This was not to be!
Considering the photo of Wettstein with his family (see illustra?
tion) and knowing his children as adults I have come to know well
and admire, I feel it is appropriate to refer briefly to F. v. Wettstein
as an individual and to his private Ufe. If I quote here from the many
obituaries the words of Otto Renner (1946), it is because I know
how carefully he chose his words and, although he was a down-to-
earth scientist, how well he was able to express deep feelings. This
is evident in his obituary for Fritz Wettstein and was here so fitting:
"the nobleman with the gentle voice and the strong heart lives on in
Englera 7- 1987 193
Von Wettstein's happy family in Berlin-Dahlem; a smile is quivering

even on the lips of the pensive father
us as oX|3ioc; happy in his father's house which was inspired by the

gifts of all the muses and also in the company of a prospering family . . .,
happy as a loved and respected teacher and equally as an esteemed
scientists,... ."
Richard Goldschmidt 1878-1958
"When I look back on the abundant years have I not every reason
to be grateful for this life and to impact these feeUngs to my
family, all my friends and my colleagues and students scattered
all over the world? "
Goldschmidt wrote these words in March, 1958, shortly before

his death but still in good health, in the foreword to his memoires,
which originally appeared under the title, "In and out of the Ivory
Tower" (1958). The German edition entitled "Im Wandel das Bleibende"
was published in 1963. Reading this book with its deseription of all
his pleasant but also very sad memories one is able to follow and under-
stand how Goldschmidt experienced Ufe as a Jewish emigrant.
It includes not only a full account of his personal fate but also reports
on his professional career and gives detaUs of his research work and
all its problems.
When Goldschmidt signed his contract as Head of Department
(the science of animal heredity) in 1914 in Berlin ? he left us with a
very detaUed deseription of the whole story and of the negotiations
between the five designated department heads and the President of the
Kaiser WUhelm Society, Adolf Harnack (Goldschmidt 1963: 96) -
he was overjoyed to discover that he would be able to undertake a
study tour not only to Japan but around the world, and then war
broke out ... .Goldschmidt was arrested in the U.S.A. and did
not return until 1919. In 1921 he took on the post of second Director,
which he retained until 1936, in spite of the many injustices he suf-
fered. He was then given an appointment at Berkeley.
Before Goldschmidt began his investigations of sex-inheritance
on the gypsy moth, Lymantria, in Dahlem on a large scale ? he had
travelled to Japan to solve the problems of the different races of this
moth ? he had already coUected a wealth of analytical material since
1906 in Munich. By 1914 he had earnt himself with this not only a
Englera 7 - 1987 195
Richard Goldschmidt sits before his microscope at the Kaiser Wilhelm

Institute in Berlin-Dahlem looking serious and thoughtful. Is he looking
into a distant, uncertain future, and is the thought of this weighing
heavily on his mind?
reputation as a successful research scientist but had also made a name

for himself as a teacher. Hans Nachtsheim was one of his students at
that time. In the same year as Baur, 1911 he published his first text-
book "Einfiihrung in die Vererbungswissenschaft" (An introduction to
the science of heredity). By 1936 Goldschmidt was able to solve
the problems of sex-inheritance in Lymantria. One of this perhaps most
important discoveries was the "balance theory of sex-determination".
It states that sex-differentiation is determined by a quantitative pro-
portion (balance or lack of balance) between the realizators(determining
genes) for male and female present in both sexes. The sex-chromosome
mechanism is responsible for the shift in balance in favour of one or
other of the realizators.
A conclusion was drawn by induction with which all the facts
concerning sex-determination in the plant and animal kingdom were
to agree. All findings concerning sex-intergrades (intersexes) can be
explained by means of this theory (Goldschmidt 1936).
In the course of his experiments on evolution genetics Goldschmidt
bred millions of gypsy moths of different geographic races in order to
obtain important information: geographic variability only represents
a process of microevolution, it does not contribute to the evolutionary
formation of the species, it leads to a dead end. With these studies
Goldschmidt won great recognition at the Vlth Congress on Ge?
netics in 1932 in Ithaca (Goldschmidt 1963: 340). He had given his
lecture in German and at Morgan's request had to repeat it in English!
From then on the problems of evolution occupied him constantly.
What had to be added to the findings of Darwin in order to explain
macroevolution? Goldschmidt sees the answer in the occurrence of
macromutants which in rare cases drastically influence the early em-
bryonic processes (hopeful monsters).
As a genetic zoologist Goldschmidt also worked with Drosophila
during his time in Dahlem. In experiments in which he tried to create
mutations by means of temperature shocks he came to the conclusion
that very many known mutants can be induced as non-hereditary
copies as a function of time, the genotypicalbackground and the kind of
strain. For this phenomenon he coins the term "phenocopy", which has
since been properly defined and became part of the vocabulary of
genetics. Many examples of phenocopies can be found amongst ani?
mals and plants. They could be referred to as "clever" modifications.
Englera7 - 1987 197
During the Lysenko era they were often considered to be hereditary

deviations. This led to false hopes caused in breeding research.
In 1931 the studies on mutagenicity in Drosophila came to be
published in the "Suddeutsche Zeitung", thanks to the dangerous,
irresponsible reporting of a BerUn journaUst. He referred to the 50,000
fruit fUes used in the experiments and maintained that in 500 years it
might be possible to create a race consisting only of geniuses. This
journaUst had gained access to the Institute via the Kaiser WUhelm
Society, had visited it with one of the scientific assistants and only
had a short interview with Goldschmidt. Angered by theshameless
conclusions drawn in the article which appeared under the heading,
"Neues aus der Wissenschaft" ("New scientific findings") he com-
plained to the Kaiser Wilhelm Society. Apparently no attempt was
made to retract. Was this the work of a Nazi ideologist?
During his career in Dahlem Goldschmidt was already well-
known as the great theoretician. Beside his "balance theory of sex-
determination", his "physiological theory of inheritance" should be
mentioned. It forms the basis of later studies on patterning in butter-
flies and genetic fixation during moulting and similar processes (Gold?
schmidt 1963:341).
Goldschmidt was able to work for more than another two
decades at Berkeley, teaching and engaged in research. He had to
emigrate from Berlin after losing his German citizenship, which had
deeply hurt him. In 1936 he was dismissed from the Institute. It was
not easy for him to leave Germany but California became a real home
to him (Goldschmidt 1963: 328). He recalls his time in Dahlem ? the
part before 1933, that is ? with gratitude. He praises the excellent op-
portunitiesfor research at the Kaiser WUhelm Institute and the pleasant
atmosphere in which he had worked with his coUeagues there. Those
who know him from this time and also younger coUeagues who were
impressed by his research and foUowed in his footsteps showed their
great recognition in 1953 by conferring on him two honourable dis-
tinctions: he became an Honorary Member of the German Zoological
Society, and the Free University (Faculty of Mathematics and Sciences)
gave him the degree of Honorary Doctor. It was also a great inter?
national honour for him to be made Honorary President of the IXth
Congress on Genetics in 1953 in Bellagio; he even held a lecture there.
On that occasion his German friends visited him in Switzerland.
Alfred Kuhn 1885-1968
In 1937 Alfred K iihh was appointed as second Director and G o ld-

sc hm id t's successor. He had already gained a reputation as a genetic
Zoologist in Gottingen, in particular as a result of his studies on questions
of gene physiology in butterfly mutants. Kiihn and his school con-
tributed to a great extent to the discovery of the effects of genes
in polyphenes in the formation of chemical phenes. He continued to
carry out research in the field which Goldschmidt had paved the way
for in theory. Thanks to his ability to experiment (transplantations) he
succeeded in establishing a chain, even a network of gene actions in his
objects: Drosophila, Ephestia, Ptychopoda and Plodia (small moths). It
became possible to develop general models of gene expression. These
fundamental studies can only be referred to briefly here. Some of them
were carried out in conjunction with the Kaiser Wilhelm Institute for
Biochemistry. Butenandt and his colleagues isolated, analysed and
synthesized the gene dependent substances, e.g. "kynurenine" which is
formed during pigment synthesis in Ephestia and Drosophila (Butenandt
1951, further literature on this subject).
In another direction K ii h n pursues the question of the formation of
morphological patterns with the evaluation of mutations and with
experimental intervention at important stages of development (sensitive
periods during the pupa stage). Like Goldschmidt he succeeded in
inducing phenocopies using the effects of heat.
AlfredKiihn was only able to remain with his department in
Dahlem until 1943. It was then moved to Hechingen (Wiirttemberg).
He was later able to continue his successful research at the Institute
which changed its name and became Max Planck Institute for Biology
in Tiibingen.
Max Hartmann 1876-1962
Max Hartmann was appointed as Director of the Department of

Plant and Animal Protist Research in Dahlem shortly after the Institute
was founded in 1914. In Giessen, and after in Berlin at the Institute
of Infection Diseases (later the Robert Koch Institute), he had made
a name for himself as an expert in the field of Protozoa research.
Englera7 - 1987 199
Hartmann was concerned with 3 areas of research: the phys?

iology of reproduction (fertiUzation and sexuaUty in protists); the
fundamentals of cytology; and questions of inheritance with regard to
problems of developmental physiology. A lengthy report on the phys-
iological investigations and the information
obtained concerning various
objects can be found elsewhere
(see Schnarrenberger in this volume).
Mention need only be made here of some cytological and genetic
research studies.
Hartmann's coUeagues, in particular K. Belaf, were able to show
convincingly in the 20s that there was a basic conformity between the
construction and behaviour of the chromosornes in protists and those
in more highly developed plants and animals. This applies to the normal
nucleus and cell division (Hartmann 1951: 224). Other important re?
search findings include the proof of a heterophasic alternation of gen-
erations in green algae, which was already generaUy known in the case
of mosses and ferns. After fertiUzation a diploid generation forms which
is then subject to meiotic when the zoospores develop,
cell division
and sex-plants subsequently arise and form gametes (Hartmann 1951:
226). This discovery is reminiscent ? and this should perhaps be men?
tioned in a historical review of this subject in BerUn ? of the important
work done 100 years earlier by Chamisso on the alternation of gen-
erations in tunicates (Salpa) (see above). Viktor Jollos carried out
extensive investigations on variabUity and the processes of inheritance
in uniceUular organisms. He found changes in his Unes, which were
neither mutations nor simple modifications. He referred to them as
"dauermodifications" (Hartmann 1951: 265, Harwood 1985).
Hartmann's findings in connection with the sex-determination
of protists (including algae and fungi) marked the climax of his research
career. Initially he had assumed in theory that they were bisexuaUy
potent; he then went on to verify this hypothesis. He was able to prove
experimentally in the case of the brown algae (Ectocarpus) that strong
male gametes mate with weak female ones and strong females with
weak males. Relative sexuaUty was also confirmed in other algae and
fungi, in a flagellate even in conjunction with morphological differ-
entiation (Hammerling 1963).
Hartmann was not there to experience the transfer of the Institute
at the end of the war. In 1938 he offered his services in the develop?
ment of a new institute in Greece. After the war, however, he was back
in Tiibingen at the Max Planck Institute where he worked until his

retirement from academic life in 1955.
Colleagues and guests
In an institute like the Kaiser Wilhelm Institute, in which the in?

dividual departments work independently, scientists are only given
very broad guidelines for work on basic biological problems and they
are very free to choose their subjects and methods of research. This
freedom even existed at the Kaiser Wilhelm Institute in Dahlem within
individual departments, which meant that scientific assistants had the
opportunity to undertake large-scale projects and to make a name for
themselves. They were not bothered in any way by the administration
(unlike those working in university departments). In the Dahlem
Institute it was also possible to some extent to cut oneself off from
the political influence during the "Third Reich". The directors offered
certain protection. Jewish colleagues, however, were not allowed
to continue working there.
Well-known colleagues such as Charlotte Auerbach, Curt Stern
and Viktor Jollos also had to leave the country. Shortly after
Correns' death Mrs. F. A. Lilienfeld joined Kihara (Tokyo),
and a very talented scientist of the younger generation, F. G. Brieger
went to London (John Innes Horticultural Institution). In 1927,
at the age of 27, Brieger had joined Correns as a scholarship
holder. He left the Kaiser Wilhelm Institute in 1933 as senior lecturer
in botany and the science of heredity. In 1936 he moved to Sto Paulo
where he acquired a great international reputation, working as a teacher
and research geneticist. In 1958 he contributed a chapter on "Popu-
lation genetics" to the German "Handbuch der Pflanzenzuchtung"
(Handbook of plant breeding). He died in 1985 in West Germany,
three years after his return.
Hans Stubbe, a scientist who had been expelled from another
institute, was able to find work in Dahlem. Like Kuckuck and
S c h i c k he had to leave the Kaiser Wilhelm Institute in Miincheberg
in 1936 because of his political views. It is on record that at the Kaiser
Wilhelm Institute in Dahlem Wettstein and Kiihn had tried to get
their industrious and efficient colleagues exempted from military
service (Melchers 1961: 126, Kiihn 1961: 135).
Englera7 - 1987 201
In the reports of the Kaiser WUhelm respectively Max Planck So-

cieties the heads of department praised the work of their coUeagues
(Goldschmidt 1936: 251, Hartmann 1951, Kuhn 1961: 135). A more
comprehensive account of the work of all the departments is provided
by Melchers (1961: 122). Repeated reference is made in the above
passages to important scientists and their works. Tribute should be paid
today to most of the scientists, although some of them were very young
when they started work at the Kaiser WUhelm Institute and some of
them never returned from the war. As their representative here, as it
were, let Karl Pirschle be mentioned, who was scientific assistant
under Wettstein and who lost his Ufe fighting in the "territorial
army" (Volkssturm) in the last days of the war. He was to be made
independent director of the Department of Physiology at the Kaiser
WUhelm Institute of Cultivated Plant Research (Director: Hans Stubbe).
His experiments using ultra-violet rays as cUmatic and selective factors
opened the way for new research.
There are few scientists from the Kaiser Wilhelm Institute days be?
fore 1945 who can stiU look back today on their careers as research
geneticists in Dahlem. Those who are stiU aUve include Hans Stubbe,
born in 1902, whom Otto Renner referred to as "the athlete of mu-
tation research" and to whom we owe a monograph on the genetics and
cytology of the snapdragon (1966), and Georg Melchers, born in
1906, who became well-known for his transplantation experiments
performed in Wettstein's department. He caused Hyoscyamus to
flower by using a graft partner which was capable of flowering, although
it should not actuaUy have flowered before a certain need for cold
(vernalization), which is conditioned by heredity, had been fulfiUed.
With Anton Lang (born 1913) he tried to find the hormones affecting
flowering (Lang 1980) and with Gerhard Schramm (1910-1969)
he tried to discover the structure of virus particles. FinaUy there is
also Joseph Straub, born 1911, who made a considerable contri?
bution to research into polyploids. The coUeagues who made names
for themselves in the departments of Goldschmidt and Kiihn
and Hartmann and who are stiU aUve today include Hans Bauer
(born 1904) who acted as Department Head for Max Hartmann
from 1940 on, Viktor Schwartz (born 1907) who was famous for his
investigations of the physiology of development in Protozoa and in?
sects, and Jakob S e i 1 e r who may stiU be aUve today in Zurich. A very
honourable woman zoologist, whose 90th birthday we celebrated on

the 4th February, 1987 in Berlin, with lectures held in her honour,
must not be forgotten: Dr. Katharina Heinro th. At the beginning of
the 30s she worked in Correns' Institute under Knapp on regen-
eration problems in mosses (Marchantiatae) and springt&ils (Collembola).
From 1945-1956 she was the Director of the Berlin Zoo.
The list of famous guests is very long. They came from all parts of
the world. Many scientists from overseas, especially from Japan, worked
in Goldschmidt's department. They concentrated mainly on
cytological and cytogenetic investigations. The woman cytologist,
Barbara McClintock (Ithaca), who was already very well-known at
that time worked in Dahlem on "her" maize chromosomes. She is still
alive today (born 1902). As rather belated recognition of her work on
the understanding of jumping genes (transposons) in the 40s she re?
ceived a Nobel prize in 1983. Transposable genetic elements are DNA-
pieces which can often be interchanged, i.e. which jump about, within
one chromosome and also between different chromosome during
ontogenesis; many characteristics, which, where colour is concerned,
cause the variegation, can be explained by these findings. They are
commented on in deatil in a study by Nevers et al. (1986) which
appeared recently. The example of easily mutable alleles (labile genes)
investigated by B a u r and C o r r e n s are also quoted here.
Kaiser Wilhelm Institute for Biochemistry 1936-1943/44
This Institute developed from the Kaiser Wilhelm Institute of Ex?

perimental Therapy which had been founded in Dahlem 1912. It did
not receive its new name until 1936 when Butenandt (born 1903)
was appointed Director. With his investigations of the enzymatic
biogenesis of sexual hormones and carcinogenic substances Bute?
nandt soon entered on a fruitful joint venture with Alfred Kiihn
(Kaiser Wilhelm Institute for Biology), who was working on gene con-
trolled substances in insects. In 1951 Butenandt reported at length
on the success of this inter-departmental project. This basic research
into gene physiology - the isolation, analysis and synthesis of products,
the explanation of enzymatic processes governed by genes (the one
gene-one enzyme-hypothesis) ? found international acclaim. Together
Englera7 - 1987 203
with other important results on hormone research they led to Bute-

na nd t being awarded the Nobel prize for chemistry in 1939.
Important contributions to work on the viruses were made in the
Virus Research Study, which was officially founded in 1941 but
which had aUeady existed in the form of a workshop since 1937.
It consistedof four departments. The Botany Department was under
G. Melchers from the Kaiser WUhelm Institute for Biology, G.
Schramm was responsible for chemistry, R. D a n n e e 1 from the
Kaiser WUhelm Institute for Biochemistry for zoology and G. B e r g o 1 d
for entomology, who was working in an outside branch.
In the few years which remained before the Institute moved in?
vestigations were carried out on plant viruses in tomatoes and tobacco
and results obtained which have greatly added to our genetic know?
ledge of viruses, e.g. of their mutability and of the resistance or sen-
sitivity of host plants. The achievements of Melchers, Bergold,
Danneel, Friedrich-Freska and in particular of Schramm
deserve our praise. A complete list of the pubUcations of this period
can be found in the book of Schramm (1971).
In 1942 and 1943 Schramm was able to prove with his experi?
ments on the infectiousness of the tobacco mosaic virus (dissection
into protein units, in vitro synthesis but without nucleic acid) that
such protein particles are not infectious i.e. they are not capable
of reproduction. It thus follows that the protein components do not
primarUy contain the genetic information for reproduction. Final
proof that it is the nucleic acid of the virus alone which is the genetic
carrier of the information for reproduction was suppUed by 1955. This
finding which was so crucial for the further development of Molecular
Biology and genetics was not accepted or recognized abroad at the
time (pubUcation 1943, see Schramm 1971). It was difficult to imag-
ine that exact, correct results had been obtained. In his memoires
Watson writes that for most people it was inconceivable that in an
institute like the Kaiser WUhelm Institute such complicated experi?
ments were being carried out during the war years. It was easier to
imagine that for some reason or other the Nazis had given the work
their direct support and that the analyses were not correct (foreword
from Butenandt to Schramm 1971: 10). According to Butenandt
for a long time after the war the Kaiser WUhelm Institute suffered
as a result of such foreign prejudice. ? The work was later continued
in Tiibingen and then in Munich.
Researches on the history of cultivated plants
The above sections make it clear how extensively and in how many
different fields genetic research was carried out at the Kaiser Wilhelm
Institute for Biology. At the end of the war research was started on
cultivated plants, and this meant that not only general questions
concerning the genetics of economic plants were investigated but also
those relating to the genetics of evolution. At Tuttenhof near Vienna
building began in 1942 on an institute that was to be directed by Hans
Stubbe. A Department of the History of Cultivated Plants was
planned with Elisabeth Schiemann as Department Head. This
department was able to start working, although only in a very limited
way, after the war in 1945 in West Berlin. It was not able, however, to
develop properly until 1948 when it received appropriate funding. It
became incorporated into the Max Planck Society in 1953 as a Re?
search Establishment (Forschungsstelle), but it was liquidated soon
after in 1956 when Elisabeth Schiemann retired from academic
life for good.
Above there is a full account of E. S c hiem a n n's research into the
genetics of cultivated plants, which she carried out until 1943, and
tribute was paid to her as one of the important scientific figures of her
day (see also report by Lang in this volume). It only remains to refer to
some of the results of her work. In the studies on Fragaria (straw-
berries) speciation and sex-differentiation were investigated and it was
possible to solve the problem of the sex-shift. Beside the distribution of
sex determining genes over several chromosomes there was also a case
with a shift which was plasmatically conditioned. With cereals the same
questions concerning descent and speciation were investigated. Of her
colleagues Ursula Nurnberg-Kruger and Giinter Staudt should be
mentioned. When the research establishment was closed the in complete
work and material were sent to Vogelsang in Cologne (MPI).
In the last years of her working life E. Schiemann developed
interesting methods for the determination of the prehistoric remains
of cultivated plants. Amongst other things 3,000 grain seed impressions
were produced with the help of plasticine. The well-known paleo-
botanist, Maria Hopf (1954), from the Roman-German Central
Museum in Mainz, who was working with E. Schiemann at the
time, made a considerable contribution to the evaluation of these
studies. Prehistoric finds were also analysed for foreign museums.
Englera7 - 1987 205
Kaiser Wilhelm Institute for Brain Research (Hirnforschung) 1931-1945
NikolaiW. Timofeeff-Ressovsky 1900-1981
The Institute was akeady founded in 1914 but building was held
up because of the war and, with Oskar Vogt as its Director, research
finally began at this Institute in Buch in the north-east of Berlin in
1931. As director Vogt was able to enlist the services of the very
talented young scholar, Timofeev-Resovskij (Russian spelUng).
Vogt had always had close contact with the USSR. In the "Third
Reich" he was reprimanded for being pro-Communist. He lost his
professorship at the Friedrich WUhelm University in 1935 but was able
to stay on as director in Buch untU 1937. Together with his wife
Ce"cile he then took over the Institute of Brain Research in Neustadt in
the Black Forest, which the two of them made famous.
That Timofeeff-Ressovsky came to Germany was partly due
to the fact that Vogt ran the State Institute of Brain Research in
Moscow and had dissected Lenin's brain in complete paraffin sections
in order to be able to analyse it better (Spatz 1961). In return he was
able to engage Timofeeff-Ressovsky, who was very interested in
genetics, to work at the Kaiser WUhelm Institute. The foundation of the
Department of Experimental Genetics was opposed by coUeagues at the
Kaiser WUhelm Institute for Biology (Eichler 1982). In the end the
research programmes were clearly defined. In Buch (later in his own
institute) four main areas of research were to be dealt with: 1) the
investigation of the variabiUty of special, changing features under
natural Uving conditions, so that variation graphs could be charted. Here
work was done at first on insects. Mutation experiments were combined
with experiments involving population genetics and were carried out on
Drosophila, ladybirds (Coccinellidae) and mice. Fish were later also
used (Betta splendens)', 2) the investigation of the penetrance and
expressivity of character formation, i.e. studies on less manifest genes
("weak genes"); 3) studies on the ex6genous influence of germ plasma;
4) a thorough genealogical exploration of the diseases in the adjacent
clinic
From the mutation experiments with x-rays and neutron rays it
was possible to derive information about radiation doses in man. Com-
parative studies on the specific effects of different amounts and differ?
ent kinds of radiation were carried out on a large scale on chromo-
somes. Timof6eff-Ressovsky became famous for his exact

charting of target curves for many different methods of treatment.
With his colleague, Karl G. Z immer, he formulated the "target
theory of the biological effects of radiation", which has been inter-
nationally recognized (Timof eeff-Ressovsky & Zimmer 1947). As
early as 1935 he published a comprehensive paper on the nature of
gene mutations and gene structure, together with Zimmer and Max
Delbriick (1906-1981) from the Kaiser Wilhelm Institute for Che?
mistry in Dahlem, the molecular geneticist who was later to become
famous (Nobel prize 1969).
Timofeeff-Ressovsky chose to stay in Buch in 1945. He was
arrested and sent back to the USSR (Spatz 1961). He certainly did not
collaborate with the Nazis. One of his two sons died in the concentration
camp in Mauthausen in 1942. He was actively engaged in scientific
research until very late in life; his last post was at the Institute of
Medico-Biological Research in Moscow. As in Berlin, there too, he
had his own circle of scholars. Herbert L ii e r s, the last full Professor
of genetics at the Free University was one of them (Eichler 1982).
Bernhard Patzig 1890-1958
The specification "human heredity" was also coined in the course

of the thirties and with it came the assignment to carry out genetic
research. Patzig tried to make use of the findings from Timofeeff-
R essovsky's department in the field of genetics, for example, the
fact that so-called weak genes with incomplete penetrance and differ-
ing expressivity occur in certain congenital diseases, such as the extra-
pyramidal-motor diseases. These also include Parkinson's disease and
St. Vitus* dance (Chorea Huntington). St. Vitus' dance was defined
as a dominant heritable disease with high penetrance but widely varying,
partly very weak expressivity. Patzig also worked on the genetic
investigation of the right-side/left-side problem in patients with speech
defects taking into account the lateral localization in the brain and on
the elementary genetics of epilepsy (Vogt & Vogt 1936). These studies
clearly show how in this Iinstitute, i.e. in the Department of Research
for Human Heredity and Constitutional Diseases (menschliche Erb- und
Konstitutionsforschung), 1931-1945, basic research found practical
application.
Englera7-1987 207
After the war genetic research was no longer continued along these
lines neither in Buch (Berlin) which is now an institute of the German
Academy of Sciences of the GDR, nor at Frankfurt am Main at the
Max Planck Institute of Brain Research in the Federal RepubUc of
Germany.
Kaiser Wilhelm Institute for Anthropology, the Science of Human

Heredity and Eugenics 1927-1945, and its successor organizations
During the Vth Congress on Genetics in Berlin in 1927 the future

of the Institute was decided. As there was no institute of human
genetics in Germany this is what it was primarily intended to be. In
addition to a Department of Anthropology (Eugen Fischer, who was
also Director) there was also a Department of Eugenics (H. Mucker-
mann) and another concerned with human genetics (Otmar v. Ver-
schuer). Two other Departments were later opened, for the Psycho-
logy of Inheritance and for Experimental Genetic Pathology.
Until the Nazis came into power the Institute had a good reputation
as a result of the human genetic research carried out there (the in?
fluence of heredity on the nature and course of tuberculosis in twins,
the inheritabiUty of psychical characteristics, the development of a
theoretical basis for the genetic, anthropological proof of descent
needed by the judiciary). At the end of the 30s Verschuer con?
centrated on phenogenetic investigations, i.e. on the question of the
effect of genetic factors on the development on the organism, on its
form and function (Vogel 1961). He was particularly interested in the
relationship between pathogenetic phenes and processes of the physio?
logy of development. As man cannot be used as the object of such
experiments, a model was needed. In 1941 Hans Nachtsheim
took over the new Department of Experimental Genetic Pathology;
he took his rabbits with him.
In the middle of the 30s, however, the ideologists of the "Third
Reich" succeeded in extending their influence and making the Institute
and its employees serve their poUtical purpose. It remains to be seen
whether pressure was exerted on them whether opportunists offered
their services. In retrospect is must be said that the reputation of
human genetics in Germany has suffered greatly. It is possible to dis-
cover what happened by consulting institute reports. The present

President of the Max Planck Society, Heinz A. Staab, quoted from
them in 1985. He read that scientists of that Institute had been
"responsible to a great degree for the racial laws passed in the Third
Reich", and had been present at "discussions concerning the establish?
ment of legal rules of administration". Heinz Staab made the remark on
this subject that there had been scientists from the Kaiser Wilhelm
Society, too, who had obviously broken the fundamental ethical rules
of science, and that we all have this burden to bear. We must accept it
without glossing the matter over or only seeing it, from one angle, just
as it is important to generally encourage acceptance of the truth of our
past.
When Nachtsheim (1890-1979) began work in the Department
of Experimental Genetic Pathology in 1941 he already had a "genetic
fund" on which to draw: a wide range of breeds of rabbit. They in?
cluded some which could be used in experiments to compare genetic
manifestations, which can also be observed in man. This applies to
Parkinson's disease, the rabbit cataract (opacity of the crystalline lens)
and to Pelger's disease (leucocyte anomaly).
In 1945 Nachtsheim took over the direction in Berlin of the
remains of the Institute under the name Institute for Comparative
Biology and Pathology of Heredity, later handed over to the German
Academy of Sciences of the GDR and to the Max Planck Society. He
was no longer able to work much with rabbits. Of 19 inbred strains he
lost 17 in the last year of the war in an air raid. He therefore began to
concentrate more on human genetics, in particular on the application of
genetic findings within the framework of family hygienics. He was also
concerned with questions relating to population policy. In a discussion
with members of the German Parlament he was once asked whether he
wanted to meddle in the affairs of the Creator. During the Nazi era he
was a clearthinking, responsible research scientist, who could in no way
be reproached for his conduct. From the security of this moral basis he
was able to demand the resumption of human genetic research in Berlin.
In 1953 Friedrich Vogel became head of a Department of Human
Genetics. He was concerned with the mutation rate of individual
genes and in addition with special genetic problems. His colleague,
W. Helmbold, investigated the relations between blood groups and
epidemics. Problems of hereditary, physiological variability were also
Englera7-1987 209
investigated with regard to reactions to external stimuli. Georg Geipel

worked on the devemopment of tactile carinas in human limbs, in
particular in limb deformities.
In addition, research of a quite different nature was also carried
out, namely the genetic investigation of microorganisms (W. Stein, W.
Harm, W. Laskowski, W. Klingmiiller, P. Knolle). In 1954
when Herbert Liiers joined them a Department was opened in which
work was carried out on the chemogenetics of Drosophila.
Microorganisms were also the objects of research of Fritz Kaude-
witz, who took over the direction of the Institute as successor in
1960. He worked on gene dependent biosynthesis in bacteria and
Neurospora and also on the genetics of an RNS-phage. These program-
mes were continued until 1965. Then the Institute was taken over by
scientists who have been working since 1964 on problems of mole-
cular genetics and who continued to administer the institute under
another name.
Max Planck Institute for Molecular Genetics since 1965
With the foundation in 1964 of this Institute, which developed

from the Max Planck Institute for Comparative Biology and Pathology
of Heredity and in which in 1965 three new heads of departments
took up their posts, a new era of basic genetic research began in Berlin.
The past is well and truly over, the future has begun. With the foun?
dation of the new Institute the courses were also reorganized. The ob-
ject of the Institute is to investigate genetic processes and to apply
the genetic information obtained at a molecular level. Nucleic acid and
protein molecules were taken as materials and biochemical, biophysical,
genetic and immunological methods used for the investigations. The
material was obtained from the organella of bacteria, viruses and
phages, with which the experiments were naturally carried out directly
(Max Planck Society 1985: 245).
A new buUding in Dahlem, Ihnestrasse 63-67, was made avaUable to
this Institute between 1969 and 1970. At present it houses three de?
partments. As information on the progress of research and the results
achieved can easUy be found at any time in the pubUcations of the
Max Planck Society by describing its organization and Usting the fields
of research it covers.
The directors of the departments take it in turns to be on the

managing directorate. In addition to the departments there are also
a few independent work groups. The Max Planck Society yearbook
for 1985 lists the following fields of research:
Department Heinz S c h u s t e r: DNA replication in bacteria, plasmids,
bacteriophages. The genetics and biochemistry of replication enzymes,
RN A - tumour- viruses.
Department Thomas A. Trautner: biochemistry of the regulation of
DNA-synthesis, biological activity of isolated nucleic acids, restriction/
modification of penicillin induced changes in pneumococci. Genetic
analysis of the photodifferentiation in Neurospora.
Department Heinz-Giinter Wittmann: biochemical, physical, genetic
and immunological investigations of the structure and function of
ribosomes and of the molecular mechanism of protein biosynthesis.
In 1985, of 87 scientists not less than 50 were guests or scholarship
holders.
Instituteof Gene Biological Research (Genbiologische Forschung)

Berlin GmbH since 1986
Convinced of the rapid progress which is to be made in the field

of genetics with regard to the practical application of new findings
in the next decades scientists, economists and politicians are all equally
interested in contributing their share. The possible uses of genetic
engineering in industry, medicine and agriculture have become in-
creasingly apparent. Since there is, however, also an awareness of the
fact that research must continue to be generously supported, thought
is being given to the matter of how best to do this.
The genetic research institute was founded as a limited company
(German: GmbH). It is financed 50 % by the Berlin Government
and 50 % by Schering AG, a well-known industrial firm producing
pharmaceutics, which also sells biocides for agriculture and forestry.
The Institute is closely bound to the Free University and the neigh-
bouring Max Planck Institute of Molecular Genetics by a co-operation
contract. Lothar Willmitzer, who was appointed Director on 1st
October, 1986, is at the same time full Professor (C4-position) at the
Institute for Biochemistry and Molecular Biology at the Free Uni-
Englera7 - 1987 211
versity. In 1985, in the Institute of Plant Physiology, Cell Biology

and Microbiology Barbel Friedrich began research on "genetic
determinants and their expression for unusual metabolisms in bacteria".
The aim is to present the individual processes of bacteria when they are
exposed to hydrogen oxidation, denitrification and heavy metal resist-
ance. This also, and above aU, has led to the creation of ajoint project
with the Free University Institute of Applied Genetics, where Otto
Schieder is engaged in unconventional plant breeding. From the
Technical University in BerUn the biotechnological research field
shaU also to be included in this joint project. This means the Institute
of Biotechnology with its four full C4-professors (at present these are
Hanswerner D e 11 w e g, Roland D r e s s e 1, Ulf S t a h 1 and one is yet to
be appointed) and one associate professor (Matthias Reuss). Other
scientists from the Free and TechnicalUniversity, biologists, chemists,
botanists and medicalresearch workers enganged in basic research are
already working together on a special research project entitled "Re-
gulation Structurs in Nucleic Acids and Proteins".
The scientific work done at the genetic institute solely serves basic
research. No research is therefore being "alone there for Schering".
The only advantage of the institute for the firm is that when patents
are being registered and Ucences granted for new methods of pro?
duction, it has a "right of option on normal market terms" (priority
over other interest ed parties). At present the research work covers two
main fields:
1) Molecular biological investigations of higher developed plants ? the
main object of this research is the potato, and the aim, the regulating
of gene activity, i.e. gaining a better understanding of the regulating of
genetic information (gene isolation and characterization, the transfer
of genetic material to help change the amino acid composition or
improve resistance to diseases and pests).
2) Molecular genetic investigations of microorganisms ? which interact
with higher developed plants. The aims are the same as those described
in 1) with relation to the gene activity of fungi and bacteria which have
either a harmful or a beneficial (symbiotic) effect on plants. Research
is to be carried out on the interaction at a molecular biological and
in particular at a genetic level.
These studies are to make use of new methods of isolation, identifi?
cation and the transfer of genetic material, which themselves are stUl
in need of improvement. Technological questions play an important

part in all these research projects (they deal after all with gene techno-
logy, i.e. genetic engineering).
With this Institute, which will employ about 60 scientists in time,
Berlin has been endowed with a highly modern research institute, the
likes of which only exist so far in a few other university towns, namely
Heidelberg, Cologne and Munich.
Future prospects
The present Senator for Science in Berlin, Georg Turner sees, in the
current possibilities for development, the conditions fulfilled which
will win recognition for Berlin and the position as the fourth German
centre of genetic research, which will enable her to be able to compete
successfully with other university towns. Here the question arises: must
the idea always prevail that it is important to pursue development and
engage in research in order to remain competitive? Will this not arouse
in even more people the suspicion that modern genetic research could
be abused and subjected to commercial interests, that it might only
serve certain commercial organizations, expecially when patents are
being awarded, or that it could favour the establishment of monopolies
and lead to new positions of strength (Rousselet 1985) ? ? Criticism of
the practical application of methods of gene technology in the drug
industry, in medical science and in agriculture is often voiced by
those who have doubts on moral, ethical or ecological grounds. Since
the possibility of manipulating genetic material can now be forseen
we must ask if there is a danger that this development might get out
of control! These are the questions scientists must discuss. Society
must exercise control on the basis of international agreements, to
prevent research findings being misused. Frank and open co-operation
is more important here than confidentially resulting from competi-
tiveness.
In Berlin the positon is very favourable so that basic and applied
research scientists should be able to complement each other and work
together. In the field of plant genetics or plant breeding this is quite
obviously the case. In animal breeding research, too, there are oppor-
tunities for the application of basic genetic findings. In the Institute
Englera7 - 1987 213
of Animal Production of the Technical University methods of gene

technology in the development of adapted stocks.
are justified
FinaUy, whilst still studying, biologists and scientists in BerUn
already have the opportunity of doing genetic research in all disci-
plines. They can choose between human, animal and plant genetics.
All three courses are equaUy well attended. As the microbe and mole?
cular geneticists also have highly modern equipment at their disposal
it can be said that all the branches of genetics are well catered for in
research and teaching. Thus it should be possible to continue the old
tradition which developed when the science of heredity was flourishing
in Berlin: of carrying out research which is as versatile and comprehen-
sive as possible. May this development be fruitful to all those involved
and, in addition, contribute to the well-being of us all.
References
Baur, E. 1924: Untersuchungen iiber das Wesen, die Entstehung und

Vererbung von Rassenunterschieden bei Antirrhinum -
majus.
Leipzig.
? Herzberg-Frankel, O., Husfeld, B., Saulescu, N. & Schiemann, Elisa?
beth 1929: Kopplungserscheinungen bei Antirrhinum majus. ? Z.
ind. Abst.- Vererbungslehre 50: 314-343.
Butenandt, A. 1951: Uber die Wirkungsweise der Erbfaktoren. -
Jahrbuch 1951 der Max-Planck-Gesellschaft: 160-174.
Correns, C. E. 1928: Uber nichtmendelnde Vererbung. ? Suppl. Z.
ind. Abst.-Vererbungslehre: 131-168.
Eichler, W. 1982: Zum Gedenken an N. W. Timofeeff-Ressovsky
(1900-1981). - Deutsch. Entom. Z., N. F. 29: 287-291.
Gaestel, M. & Niirnberg, P. 1986: "Labore non lauro laetor!" (Arbeite

und erliege nicht der Verfiihrung des Lorbeers!) Carl Erich Correns
(1864-1933). - Gleditschia 14: 333-342.
Goldschmidt, R. 1936: 25 Jahre Kaiser-Wilhelm-Gesellschaft 2. KWI

fur Biologie in Berlin-Dahlem: 251 -260.
- 1963: Im Wandel das Bleibende. - Berlin.
Hammerling, J. 1963: Max Hartmann. ? Naturwissenschaften 51:

365-366.
Hartmann, M. 1936: 25 Jahre Kaiser-Wilhelm-GeseUschaft 2. KWI fur

Biologie in Berlin-Dahlem: 261-266.
- 1951: Bericht uber 37 Jahre Forschung im KWI(MPI) fur Biologie -
Jahrbuch 1951 der Max-Planck-GeseUschaft: 199-228.
Harwood, J. 1984: The reception of Morgan's chromosome theory in
Germany: Inter-war debate over cytoplasmic inheritance. - Medizin-
hist.J.19:3-32.
- 1985: Geneticists and the evolutionary synthesis in interwar
Germany. - Ann. Sc 42: 279-301.
Heck, L. 1934: Uber die Neuzuchtung des Ur oder Auerochs. - Ber.
der Intern. Ges. zur Erhaltung des Wisents 3: 225-294.
Hertwig, Paula 1940: Vererbbare SemisteriUtat bei Mausen nach Ront-
genbestrahlung. - Z. ind. Abst.- Vererbungslehre 79: 1-27.
- 1957: Zur Geschichte der strahlenbiologischen Forschung und ihre
Bedeutung fiir die Gegenwart. - Wiss. Z. Univ. HaUe, Math.-Nat.
6:405-412.
Hoffmann, W., Mudra, A. & Plarre, W. 1971: Lehrbuch der Zuchtung
landwirtschaftUcher Kulturpflanzen 1. ? BerUn.
- 1985: Lehrbuch der Zuchtung landwirtschaftUcher Kulturpflanzen
2, 2nd edition. ? BerUn.
Hopf, M. 1954: Anatomische Untersuchungen an Weizenspelzen und
-kornern verschiedener Polyploidiestufen als Vorarbeit fiir die
Bestimmung prahistorischer Funde. ? Zuchter 24: 174-180.
Jollos, V. 1935: Studien zum Evoiutionsproblem 2. ? Biol. Zentralbl.
55: 390-436.
Kappert, H. 1978: Vier Jahrzehnte miterlebte Genetik. ? Berlin.
Knapp, E. 1966: Eine Klarstellung: Antirrhinum Juliwa und Bright
Butterflies. - Gartenwelt 66: 506-507.
Kuckuck, H. 1962: Vavilovs Genzentrentheorie in heutiger Sicht. ?
3. Kongrefi Eucarpia Paris 1962: 177-196.
Kuhn, A. 1961: Jahrbuch der Max-Planck-GeseUschaft 1961, 2. Teil.
MPI fiir Biologie in Tiibingen: 135-141.
Lang, A. 1980: Some recoUections and reflections. - Ann. Rev. Pl.
Physiol. 31: 1-28.
Leussink, H., Neumann, E. & Kotowski, G. 1960: Studium Bero-
linense. ? Gedenkschrift der Westdeutschen Rektorenkonferenz
und der Freien Universitat Berlin zur 150. Wiederkehr des Griin-
dungsjahres der Friedrich-WUhelms-Umversitat zu BerUn. - Berlin.
Englera 7-1987 215
Linnert, Gertrud 1978: Genetische Analyse von instabilen Mutationen

bei Antirrhinum majus L.: Genetische und somatische Reversion
am incolorata-2-Locus. - Biol. Zentralbl. 97: 513-526.
Liiers, H., Sperling, K. & Wolf, B. E. 1974: Genetik und Evolutions-
forschung bei Tieren. - In: G. Heberer (ed.). Die Evolution der
Organismen II/l. - Stuttgart, p. 190-363.
Max-Planck-Gesellschaft 1985: Jahrbuch 1985. - Miinchen.
Mayr, E. 1976: Evolution and the diversity of life. - Cambridge, Mass.
Melchers, G. 1961: Jahrbuch der Max-Planck-Gesellschaft 1961, 2.
Teil. MPI fiir Biologie in Tubingen: 111 -134.
Monaghan, F. & Corcos, A. 1984: On the origins of the Mendelian
Laws. - J. Hered. 75: 67-69.
Nachtsheim, H. & Stengel, H. 1977: Vom Wildtier zum Haustier. -
Berlin.
Nevers, P., Shepherd, N. S. & Saedler, H. 1986: Plant transposable
elements. - Advances Bot. Res. 12: 103-198.
Nothel, H. 1970: Investigations on radiosensitive and radioresistant
populations of Drosophila melanogaster. ? Mutation Res. 10:
463-474.
Plarre, W. 1972: Die Entstehung eines Genzentrums in Ostafrika. ? Z.
Pflanzenziicht. 68: 124-148.
Renner, O. 1935: Gedenkworte auf Erwin Baur. ? Z. ind. Abst.- Ver.
erbungslehre 70: 351 -357.
- 1946: Friedrich Wettstein Ritter von Westersheim 24.6.1895 bis
12.2.1945. - Naturwissenschaften 33: 97-100.
- 1961: William Bateson und Carl Correns. - Sitzungsber. Heidel-
bergerAkad.Wiss.,Math.-Nat. Kl., Jahrg. 1960/61: 159-181.
Rousselet, W.-P. 1985: Institut fiir Gentechnologie, Pharma-Riese
kooperiert mit Universitaten. - Berlin FU-info 13: 12-13.
Sacristan, Maria Dolores & Gerdemann, Maria 1986: Different behavior
of Brassica juncea and B. carinata as sources of Phoma lingam
resistance in experiments of interspecific transfer to B. napus. -
Pl. Breeding 97: 304-314.
Saha, M. S. 1981: The Carl Correns papers. ? The Mendel newsletter.?
Archival Resources for the History of Genetics & Allied Sciences
21: 1-6.
Schachl, R. 1984: Ein eiszeitliches Genzentrum fiir Getreideausgangs-
formen. - Aktuelle Probleme der Landwirtsch. Forschung, 9.
Seminar 1982, Landw.-chem. Bundesversuchsanstalt Linz/Donau:

50-58.
Schieder, O. 1987: Zellfusion und Hybrid her stellung bei Pflanzen. ?
Sitzungsber. Ges. Naturf. Freunde Berlin (N. F.) 27: 119-126.
- Ber. Deutsch. Bot. Ges.
Schiemann, Elisabeth 1935: Erwin Baur.
52:51-114.
? 1938: Georg Schweinfurths Bedeutung fiir die Kulturpflanzen-
forschung. - Ziichter 10: 18-21.
? 1939: Gedanken zur Genzentrentheorie Vavilovs. ? Naturwissen?
schaften 27: 377-383,394-401.
Schneebeli-Graf, R. 1983: Adelbert von Chamisso . . . Und lassen
gelten, was ich beobachtet habe. ? Berlin.
Scholz, H. 1986: Die Entstehung der Unkmut-Gerste Hordeum vulgare
subsp. agriocrithon emend. - Bot. Jahrb. Syst. 106: 419-426.
Schramm, G. 1971: Bauplane des Lebens. ? Munchen.
Sengbusch, R. von 1931: Bitterstoffarme Lupinen II. - Ziichter 3:
93-109.
Skibbe, B. & Gocht, H. 1956: Berliner Forschung und Lehre in den
Land wirtschaf tswissenschaften. ? Festschrift zur 75-Jahr-Feier. ?
Berlin.
Spatz, H. 1961: Jahrbuch der Max-Planck-GeseUschaft 1961, 2. Teil
MPI fiir Hirnforschung in Frankfurt/Main, Gottingen und Koln:
405-421.
SperUng, K. & Marcus, M. 1984: Mapping of genetic activity on mam-
maUan chromosornes. - Chromosornes Today 8: 169-178.
Staab, H. A. 1986: Kontinuitat und Wandel einer Wissenschaftsorgani-
sation: 75 Jahre Kaiser-WUhelm-/Max-Planck-GeseUschaft. -
MPGSpiegel4:37-53.
Stein, Emmy 1950: Dem Gedachtnis von Carl Erich Correns nach
einem halben Jahrhundert der Vererbungswissenschaft. ? Natur?
wissenschaften 37: 457-463.
Stubbe, H. 1951: Nachruf auf Fritz von Wettstein. ? Jahrb. Deutsch.
Akad. Wiss. BerUn 1950/51: 1-12.
? 1966: Genetik und Zytologie vonAntirrhinum L. sect. Antirrhinum. -
Jena.
Telschow, E. 1942: Jahrbuch 1942 der Kaiser-WUhelm-GeseUschaft. -
Aus der Arbeit der KWI vom 1. April 1941 bis 31. Marz 1942:
9-48. - Aus dem Mitgliederkreis: 63-77.
Englera 7-1987 217
Timofeeff-Ressovsky, N. W. & Zimmer, K. G. 1947: Das Treffer-

prinzip in der Biologie. ? Leipzig.
? ? & Delbriick, M. 1935: Uber die Natur der Genmutation und der
Genstruktur. ? Nachr. Ges. Wiss. Gottingen, Math.-Phys. Kl.,
Fachgr. Biol. N. F. 1: 190-245.
Vavilov, N. 1922: The law of homologous series in variation. ? J.
Genet. 12:47-69.
Vogel, F. 1961: Jahrbuch der Max-Planck-Gesellschaft 1961, 2. Teil,
MPI fiir vergleichende Erbbiologie und Erbpathologie in Berlin-
Dahlem: 291-300.
Vogt, C. & Vogt, O. 1936: 25 Jahre Kaiser-Wilhelm-Gesellschaft, 2.
KWI fur Hirnforschung in Berlin-Buch: 387-399.
Wettstein, F. von 1936: Biologie. ? 25 Jahre Kaiser-Wilhelm -Gesell?
schaft, 2: 105-126.
? 1938: Carl Erich Correns. - Ber. Deutsch. Bot. Ges. 56: 140-160.
? 1943: Warum hat der diploide Zustand bei den Organismen den
grofieren Selektionswert? ? Naturwissenschaften 31: 574-577.
Zepernick, B. &Karlsson, Else-Marie 1979: Berhns Botanischer Garten.?
Berlin.
Zhukowsky, P. M. 1956: DieEntstehungder Kulturpflanzen.? Deutsch.
Akad. Landwirtschaftswiss. Berlin, Sitzungsber. Bd. 5, H. 23.

Prof. Dr. W. Plarre, FurstenstraBe 28, D-1000 Berlin 37.
Englera 7-1987 219
Paul Hiepko
The collections of the Botanical Museum Berlin-Dahlem (B)

and their history
Herbaria and other scientific collections are ? in addition to a large

library ? essential parts of a research institute specializing in taxonomie
botany. The fate of such an institute is, therefore, closely linked with
the fate of its collections. The destruction of the greater part of the
herbarium of the Botanical Museum Berlin-Dahlem in 1943 was there?
fore a far-reaching event in the history of this institution. One may
think that an historical account of the herbarium of the Botanical
Museum could omit its development up to March 1, 1943, because
the old holdings were totally destroyed. But, fortunately, this opin-
ion is completely wrong, even if it was ? prompted by publications
like that of Merrill (1943) ? widely circulated during the first decades
after World War II (see also Shetler 1969: 707). It was estimated that
about one-half million specimens were saved (Sleumer 1949: 173)
and among this remaining stock there are at least single specimens
from nearly all old collections, especially among the surviving types.
I will, therefore, give a short account of the entire history of the
Botanical Museum which can be divided into three periods:
1. From the beginning until 1913. - Since Urban(1881, 1909, 1916)
has discussed the history of the Botanical Museum up to 1913 in several
publications (including comprehensive lists of the collections), this
period can be dealt with concisely.
2. 1913 to March 1, 1943. ? For the following period up to the de?
struction of the Herbarium building there is no account of the develop?
ment of the holdings of the Botanical Museum. A summary will be
given based on the annual reports published in the "Notizblatt des
Botanischen Gartens zu Berlin-Dahlem".
3. March 2, 1943 to present. ? In this chapter, the surviving parts of
the old collections that formed the basis for the rebuilding of the her?
barium will be covered.
220 Hiepko: Collections of the Botanical Museum
From the beginning until 1913
The earliest herbarium in Berlin was set up by J. S. Elssholz

around 1657, even before the founding of the Botanical Garden in
1679. This herbarium was kept in the Royal Library, but it is no
longer extant. Other small collections of plants were kept in the Na-
turaUenkabinett of the Sozietat der Wissenschaften zu BerUn founded
in 1700 (since 1744 the Royal Academy). These collections which were
for some time looked after by J. G. G 1 e d i t s c h, in 1770 contained ca.
5,600 species.
Another collection of dried plants was maintained in the Cabinet of
the GeseUschaft Naturforschender Freunde zu Berlin since the foun?
dation of the Society in 1773. This herbarium ultimately contained
1,042 species with 1,902 specimens (Urban 1916: 12).
These old est plant collections in Berlin were not well maintained
and were hardly used for scientific purposes. The first intensively
used herbarium was built by C. L. Willdenow who was Director
of the Berlin Botanical Garden from 1801 until his death (1812);
from 1810 he was also Professor at the newly founded Friedrich Wil?
helm University. of Willd enow's
Because important and volu-
minous pubUcations his outstanding herbarium contains very many
types and was one of the largest collections in its time (Eckardt 1965).
In connection with the increasing importance of herbaria at the

beginning of the 19th century an institutionalization of the formerly
private collections can be observed (Stafleu 1987). The origins of the
Royal Herbarium ? later Botanical Museum ? go back to this time,
thus, it is not possible to give a precise date of foundation. FoUowing
Urban (1916) and Eckardt (1966) I consider the year of Link's
assumption of the directorship, i.e. 1815, as the year of foundation
because Link soon decided to build a regular herbarium (Urban
1916: 12). This herbarium at first consisted of several smaU collections
of exotic plants. In 1818 the extremely important collection of Will?
denow was bought for the herbarium by order of Friedrich Wilhelm III,
King of Prussia. The Herbarium WUldenow, fortunately, was not de-
stroyed in 1943 and is stUl kept separate (B-W). It contains ca. 38,000
specimens comprising 20,260 species of vascular plants (the 6,000
Cryptogams mentioned by Urb an are not extant, except 27 species of
Englera 7 - 1987 221
Chara). Details on the composition of the collection are given by U r b a n

(1916: 412 ff.) and H i e p k o (1972).
The extraordinarily valuable specimens of the historic Herbarium
Willdenow are now ? like those of many other historic collections -
not sent out on loan. In 1971, therefore, a microfiche edition was
produced by IDC, Zug, including a systematic index. Somewhat later
a printed alphabetical index was also published together with intro-
ductory chapters (Hiepko 1972; see also Stafleu 1972). The following
papers on parts of the Herbarium Willdenow were published: Plants
of J. R. & G. Forster (Hiepko 1969), Mexican plants ofHumboldt&
Bonpland (Ern 1976, Appendix), collections of Ruiz et al. (Lack
1979), plants from Tournefort's journey to the Orient (Wagenitz
1962), and the genus Aster (Jones & Hiepko 1981).
Through the acquisition of this herbarium the collections became

so large that it was necessary to comply with the second important
requirement for a working institutional herbarium, i.e. the appoint?
ment of a professional staff. In 1819 thebotanistD.F.L. von Schlech?
tendal was employed as "Aufseher der offentlichen Krautersammlung"
(supervisor of the public collection of herbs), and was even assisted by
a servant in poisoning and mounting the specimens. At the same time
A. von Chamisso, known as a poet and scientist, was appointed as
"Mitaufseher des Botanischen Gartens" (co-supervisor of the Bota?
nical Garden) to start a herbarium of plants cultivated in the garden.
But Chamisso soon withdrew from the Botanical Garden and
studied with Schlechtendal the plants collected during the latter's
voyage around the world in 1815-1818 that was initiated by Romanzoff.
After the appointment of Schlechtendal to the University at
Halle in 1833 Chamisso succeeded him as curator of the herbarium
and remained for 5 years. When he retired shortly before his death in
1838 J. F. Klotzsch, who was assistant in the herbarium from 1834,
was appointed curator and continued to hold the position until he died
in 1860. During this period he was supported by several assistants. An
important innovation introduced by Klotzsch was the sending of
specimens on loan to foreign botanists. Through this policy the number
of types increased considerably since many specialists described new
taxa based on specimens from the Berlin Herbarium.
Also during this time some extremely valuable private collections
were purchased. The largest and most important collection was that
of C. S. Kunth, Vice-Director of the Botanical Garden, who died in

1850. Before he took over the position in Berlin, he had Uved in Paris
from 1815 to 1828 whUe working on the plants coUected by A. von
Humboldt and A. Bonpland in America. Kunth's herbarium was
a coUection of ca. 70,000 specimens, comprising about 54,500 species,
and contained ca. 3,000 types of taxa described in the "Nova genera
et species . . ." as well as many duplicates from the herbarium in Paris,
and plants from the botanical gardens in Paris and BerUn and other
important coUections.
The herbarium of Link, purchased after his death in 1851, was not
as large as that of Kunth (only 3,113 species of Cryptogams and
16,382 species of Phanerogams), but it also contained many types of
taxa described by Link (fungi as well as plants from the Botanical
Garden).
Another large addition was a part of the herbarium of Nees von
Esenbeck acquired in 1855 (nearly 10,000 species of Cyperaceae,
Gramineae, Juncaceae, and Restionaceae).
At this time there was stiU no adequate building to house the rapidly
growing collections. First the herbarium was housed in some
rooms of a building belonging to the Academy in Berlin, but in 1822
it was moved to a smaU residential building in Neu-Schoneberg, i.e.
near the Botanical Garden. Because of lack of space 35 years later the
herbarium had to be transfered to a building of the University in Berlin.
From this place, however, it was in 1871 again forced out and moved
to a private house with unheated rooms in the centre of Berlin. During
the last two moves Link's successor, the famous morphologist A.
Braun was already Director (1851-1877). Braun tried with great
effort to get a new building especiaUy for the collections in the area of
the Botanical Garden at Schoneberg. FinaUy he succeeded, but he died
before the construction was started in 1878. The new building was
inaugurated by the new Director, A. W. Eichler, with the official
name "Konigliches Botanisches Museum" (Royal Botanical Museum). In
this Museum for the first time a pubUc department was estabUshed in
which, besides other objects, larger specimens such as fruits, wood
samples, and spirit coUections were exhibited.
After Klotzsch's death in 1860 J. Hanstein was appointed
"Erster Kustos" (first curator), but only 5 years later he accepted a
chair at Bonn. He was succeeded by the former second curator, A.
Englera 7 - 1987 223
Garcke (1819-1904), author of the well-known "Illustrierte Flora von

Deutschland" which was published in several editions. His collaborators
wereP. Ascherson (1834-1913) and F. C. Dietrich(1805-1891).
In 1884 Ascherson was replaced by K. Schumann(1851-1904).
The successor of Braun, A. W. Eichler (1839-1887, Director
since 1878) brought the colossal research project "Flora brasiliensis" to
Berlin which he had taken over from Martius at Munich in 1868.
Therefore, Eichler's herbarium, purchased shortly before he died,
was especially rich in Brazilian plants (over 12,000 specimens from
Glaziou).
In the 1880s several collections made by German expeditions to
Africa came into the possession of the Botanical Museum; these collec?
tions, however, were studied only in part at that time.
OnOctober 1,1889 A. Engler(1844-1930)wasappointedDirector.
Under his direction the Botanical Museum reached the zenith of its
development at the beginning of the 20th century (Eckardt 1966:
168 ff., Stafleu 1981). Here the role of I. Urban(1848-1931) hasto
be emphasized as he has already been mentioned as author of the
history of the Botanical Museum. Urb a n's scientific achievement
was equal to Engl er's and in his position as "Unterdirektor" (Sub-
Director) he was decisively involved in the moving of the Botanical
Garden and the Museum from Schoneberg to Dahlem. Through this
moving of the collections (which had grown explosively during the
1890s) into a spacious new building the best environment for fruitful
research was created.
Along with studies for the main works edited by Engler, "Die
Naturlichen Pflanzenfamilien" (with K. Prantl), "Das Pflanzenreich"
and "Die Vegetation der Erde" (with O. Drude), the study was pushed
forward of the immense collections which came to Berlin at the end
of the last century from the young German colonies in East Africa,
Cameroon, Togo, S. W. Africa, New Guinea etc. (cf. Timler & Zeper?
nick 1987). In this connection E ngler edited in his "Botanische Jahr-
biicher" between 1892 and 1913 for instance 42 "Beitrage zur Flora
Afrikas" (Contributions to the Flora of Africa) comprising more than
6,900 total pages.
Through the fast growth of the collections, the Museum building
was again too small only a few years after the opening. In October
1906 the construction of the new building in Dahlem with a fourfold
increase in space was finished, and the transfer of the collections
ended six months later.
224 Hiepko: CoUections of the Botanical Museum
1913 to March 1,1943
The year 1913, the last year covered by Urban's history of 1916,
coincidentaUy was also the last year in the rapid development of the
Botanical Museum during Engler's directorship. After the outbreak
of World War I in 1914 the number of staff members was soon reduced,
and the stream of collections from the colonies dried up very quickly.
Nevertheless, the study of plant specimens already in the herbarium
resulted in 12 more "Beitrage zur Flora von Afrika" (Contributions
to the Flora of Africa) pubUshed in the "Botanische Jahrbiicher"
(vols. 51-62, 1914-1929) with more than 1,700 pages edited by
E ngler.
Shortly before the war another series was started, "Beitrage zur
Flora von Papuasien" (Contributions to the Flora of Papuasia) edited
by C. Lauterbach and pubUshed in the same journal. This series of
papers was at first based on collections ofC. Ledermann, R. Schlechter,
H. HoUrung, C. Lauterbach, O.Warburg and many others (see Timler &
Zepernick 1987), mainly from the northeastern part of New Guinea
(Kaiser-WUhelmsland). For the later pubUcations in thisseries, edited
by L. Diels, new coUections of Brass, Clemens and others were also
studied. The 25 parts of the series were pubUshed between 1912
and 1940 and comprise over 3,300 pages.
After retiring in 1913, Urban worked exclusively on the plants

from the West Indies assembled in the Herbarium Krug & Urban. These
collections were continuously expanded during the years 1915-1935
by the coUections of E. L. Ekman who added more than 24,000 num?
bers.
Engler retired in 1921 at 77 and his pupU L. Diels(1874-1945)
was appointed Director. Diels had been Vice-Director of the Botanical
Garden and Museum at Dahlem since 1913. During the last 7 years
of Engler's directorship several other large collections came into
the possession of the Museum. The most important was the herbarium
of the Naturhistorisches Museum zu Liibeck which contained several
thousand specimens including many old collections not present in
Berlin before (see Anon. 1916: 405 f. for list of collectors). The most
important part of the "Herbar Lubeck" was the original S. African
collection of J. Drege with about 8,000 specimens that had originaUy
belonged to E. H. F. Meyer (Konigsberg, f 1858).
Englera 7 - 1987 225
Other notable acquisitions during that time were the herbaria of

H. Dingler (including the original Greek collection of W. v. Spruner),
H. Kinscher, E. Koehne, M. Schulze (Jena), and J. Utsch (for details
see Appendix A).
The transition of the directorship to L. Diels- R. Pilger (1876-

1953) was D i e l's successor as Vice-Director ? at first resulted in hardly
any change in the research program. In addition to studies for the
second edition of "Die Natiirlichen Pflanzenfamilien" and further
monographs for the "Pflanzenreich" new collections were studied
and the results, especially descriptions of new and rare taxa, were
published. Between 1914 and 1939 Diels edited for the journal
"Botanische Jahrbiicher" five contributions to the Flora of Micronesia
and Polynesia. In the "Notizblatter des Botanischen Gartens und
Museums zu Berlin-Dahlem" papers on several notable collections
were published as series: Luetzelburg (Brazil), Steinbach (Bolivia),
Tessmann (Peru), and Schlieben (E. Africa, with ca. 500 new taxa!).
When the "Flora brasiliensis" was finished in 1906, no large new

floristic project existed in the Museum. U r b a n all his life had attended
to the flora of the West Indies as editor and main author of "Symbolae
antillanae", and P. Graebner worked until his death in 1933 onthe
"Synopsis der mitteleuropaischen Flora" originally founded by Ascher-
so n. Then in 1939 a new institutional project was started ? a Flora of
S. W. Africa (Anon. 1940: 284). Two years later the treatments of sev?
eral families were already finished, but most of the manuscripts of the
flora burned in 1943 (Werdermann 1954) and the project was resumed
after the war at Munich.
Collecting trips by staff members of the Botanical Museum to
European countries or to the tropics were not possible during World
War I and the following years. The first expedition from 1923-1927
was led by E. Werdermann to Chile and Bolivia. Several collections
were also made during short trips to the Balkans, the Near East, the
Alps, and to the tropics: Cameroon, Brazil, Ecuador etc. (see Zepernick &
Timler 1979: 14 f.).
In the 1920s many collections from China came to Berlin (those of
Hu, Klautke, Schneider, Sin, and others) which were studied by Diels
and later by visiting Chinese botanists. For example, Y. C. Wu worked
for more than 4 years (1936-1940) at the Botanical Museum.
The Royal Botanical Museum at Berlin-Dahlem, ca. 1910; right:
entrance to the Botanical Garden
?fî?S5!^î#?W?-
?#?*;}??
?V?ift?&Mii&s?
The same building after reconstruction, 1978

Englera7 - 1987 227
Among the many important collections that were added to the

herbarium during the 1920s and 1930s those of A. W. Roth (1757-
1834), W. Becker (vouchers and types of his work on Viola), E. H. L.
Krause, C. Mez, F. A. Kornicke, H. Sandstede (Cladonia), A. Peter
(E. Africa), and K. H. Zahn (Hieracium) are some of the most notable
(for details see Appendix A). Another noteworthy acquisition was the
herbarium of the Naturhistorischer Verein fiir die Preussischen Rhein-
lande und Westfalen at Bonn received in 1936 (Anon. 1937). This
huge coUection of about 200,000 specimens containing the important
herbarium of L. C. Treviranus was evacuated to Miincheberg east to
BerUn and did not return to Dahlem after the war. It is now in East
Berlin at the herbarium of the Humboldt University: BHU (cf. Bassler
1970 and Hiepko 1979).
At the beginning of the 1940 s the herbarium contained about 4

million specimens (Eckardt 1966: 159); only ten years previously the
Phanerogam collection alone had been estimated at 2,2 million sheets
(Anon. 1934: 11). Just after the beginning of World War II the collec?
tions of Cryptogams stored in the attics of the Herbarium building were
moved to safer locations in the cellar, in corridors, and other rooms
of the building. Only the Herbarium WUldenow was evacuated to a
bank vault. Around the beginning of 1943 a sorting out of types and
other authentic
material was begun. Unfortunately, this effort was
not finished when the Herbarium building was destroyed by fire in a
bombing raid on the night of March 1-2, 1943.
"The loss of the BerUn herbarium is a catastrophe of major pro-
portions to world botany" wrote MerrUl (1943), who continued:
"This herbarium, one of the largest and most important in the world, . . .
contained the basic historical collections of Germany outside of those
at Munich. Scores of thousands of type specimens from aU parts of the
world were thus destroyed". Fortunately about 20,000 types of the
Phanerogam collcetions have been saved (see part III).
The dimensions of this loss can only be visuaUzed in comparison
with the significance of the old BerUn herbarium. It was founded
relatively late. In the territory of the later German Reich seven in?
stitutional herbaria existed before 1815, including the Royal Herbarium
at Munich, founded in 1813. Many herbaria in other European coun?
tries - e.g. the Netherlands, France, Great Britain, and Sweden - were
estabUshed much earUer.
While most herbaria soon tended to be specialized geographically,

the Berlin herbarium did not show any limitations regarding either
groups of plants or geographical regions. The fact that plants from the
area surrounding Berlin were relatively rare in the collections for more
than 40 years after its foundation is characteristic for this herbarium.
The neotropics, however, were a special field of interest right from
the beginning and in the 1930s the Botanical Museum Berlin-Dahlem
consequently "housed the world's largest collection of neotropical
types" (Grime & Plowman 1986: 932).
The growing political importance of Prussia in the 19th century,
the fast development of Berlin as capital of the German Reich from
1871, and the key position of the Botanical Museum for the exploration
of the new German colonies in Africa and Oceania caused the Berlin
herbarium to become the largest in Germany and in German-speaking
Europe. With ca. 4 million specimens, it ranked with Kew, Leningrad,
and Paris as one of the four largest herbaria of the world.
March 2, 1943 to present
The extent of devastation revealed on the morning of March 2,

1943 was described by Pilger (1953a: 4). The eastern part of the
Botanical Museum housing the library and the bulk of the herbarium
had been burnt out, as well as the workrooms in the central part of
the building. Only those collections that had been kept in the basement
and in the west wing of the Museum were saved.
Since it was not possible to store the collections that survived in
the less damaged parts of the building this material was evacuated
eventually to seven different locations (Pilger 1953a: 18 f.). The most
important collections - the Herbarium Willdenow, the ca. 20,000
types of Phanerogams that were saved, duplicates of valuable collections
etc. ? were stored in a mine at Bleicherode in Thuringia. Also kept at
the same place were the very large collections that had been handed
over to the Berlin Museum in 1944 from the Zoologisch-Botanische
Gesellschaft in Vienna and the Botanical Institute of the University
of Vienna, namely large parts of the broken up herbarium of the
Zoologisch-Botanische Gesellschaft (cf. Lack 1980), Herbarium Ulle-
pitsch (Lack & Wagner 1985), Herbarium Reuss and other collections.
Englera7- 1987 229
The fate of each of the evacuated collections was different: The

Hieracium collections moved to Eberswalde and the Lichen herbarium
of J. Hillmann stored in the church of Buckow were destroyed by
fire. The large herbarium of the Naturhistorischer Verein fiir die Preussi-
schen Rheinlande und Westfalen evacuated to the Erwin Baur Institute
at Miincheberg was removed by the Soviet authorities to an unknown
destination and, in 1964, presented to the Humboldt University in
East Berlin.
The extraordinarily valuable coUections at Bleicherode, which were
also missing since March 1946, turned up again in the former Staats-
bibliothek in the Soviet sector of BerUn at the end of 1947. Because
the Botanical Museum was located in the American sector it was se-
parated in March 1946 from the University, the center of which was in
the Soviet sector. Because of this situation the authorities of the
University did not want to return the coUections to the Botanical
Museum. After some difficult negotiations the material came back in
March 1948 in exchange for the Ubrary of the Institute of Physics
of the BerUn University which had been evacuated to Dahlem during
the war. The entire coUection of Pteridophytes came back even later.
The Director of the Museum at that time was R. Pilger, successor
to Diels who had died in November 1945. Pilger - who in 1946
was already 70 years old - bore the burden of post-war rebuilding un?
tU the end of 1950, aided only by some of the old staff members.
The restoration of the two surviving wings of the Museum was very
slow and only more or less finished in the middle of the 1950s. There?
fore it was not possible to appoint a new director before 1955. The
scientific staff had been reduced to 8 persons. From 1950 to the
present there were nine changes in the directorship. In addition to E.
Werdermann (1892-1959), H. Melchior (1894-1984), and W.
Domke (1899-), Th. Eckardt (1910-1977) who led the Botanical
Museum for 13 years should be mentioned especiaUy. His was the
longest period of directorship after the war.
Since 1960 it was possible to enlarge the scientific staff to 17
botanists. However, the reconstruction of the building for the her?
barium and Ubrary could not ? despite of several attempts - be turned
into reaUty. During the tercentenary celebration of the BerUn Bo?
tanical Garden in 1979 a new building was again promised and, finaUy,
in 1987, under the directorship of W. Greuter, the coUections were
moved into the new wing.
The rebuilding of the herbarium was started soon after the 1943
destruction, from some European collections that remained at Berlin
(e.g., the herbaria of Engelhardt, B eg e r, and B o t h e). The fast growth
of the holdings of the Botanical Museum in the following years was
only possible through generous gifts from friendly institutions and
botanists. For example, specimens came from the collections of the
Botanical Institute of the Faculty of Agronomy of the Berlin University
and from the University at Marburg as well as large collections from
Vienna. Other sources of new material were specimens generously sent
as gifts or in exchange soon after the war from American and British
herbaria. The large herbarium of Bornmiiller, including many types
especially from S. W. Asia, is one of the most important collections
in the new Berlin herbarium (cf. Wagenitz 1960).
Collection trips organized by the Botanical Museum were for a long

time taken only to European countries. Since 1972 several expeditions
have been carried out in the Republic of Togo, to contribute to the
exploration of this botanically relatively poorly known West African
country. The result of this project, a "Flore analytique du Togo",
was published in cooperation with the University at Lome (Brunel
etal. 1985).
The herbarium of the Botanical Museum Berlin-Dahlem (B) now
contains about 2,5 million specimens. In the following pages, a survey
of holdings of the different plant groups in the general herbarium will
be given, emphasizing the parts of the old collections that were saved
in 1943 (cf. Hiepko 1978) which according to Sleum er (1949: 173)
was estimated to be "more than half a million sheets."
Nevertheless it must be stressed that because of the confusion during
and after the war this compilation cannot be complete and that it is
necessary to check each individual case in the herbarium to be sure
whether or not a type was saved.
In the following text only a few important collections are mentioned.
More information on these and additional collections is given in Appen?
dix A.
Algae
The Algae collections of the general herbarium were completely
destroyed in 1943. Only 57 sheets of the old herbarium are extant,
having been on loan during the war. However, some fascicles of old
Englera7 - 1987 231
material were found in the basement including 600 sheets of the

Kutzing herbarium acquired in 1894. In the extant spirit coUection
there are ca. 480 bottles of Algae from Cameroon (Ledermann), An-
nobon (Mildbraed), Madagascar (Voeltzkow), and the Kerguelen
Is.(Werth).
Today the coUection of Algae comprises about 34,500 sheets which
are numbered throughout.
Fungi
All of the material of Uredinales and Fungi imperfecti was saved,
while the remainder of the herbarium of Fungi was destroyed. Several
additional types from different groups of Fungi were saved because
they were kept in the pubUc department of the Museum, e.g. types of J.
F. Klotzsch and P. Hennings (cf. Kohlmeyer 1962a, 1962b).
Other important collections were saved because they were kept in safer
parts of the building. Among these are the herbaria of C. G. T. Preuss
(cf. Julich 1974) and Th. Nitschke (cf. Gerhardt & Hein 1979).
Lists of extant types of Fungi described by W. Kirschstein and
G. Otth were pubUshed by Hein & Gerhardt (1981) and Hein
(1985) respectively. In 1979 the Fungus collection of the Biologische
Bundesanstalt at Berlin-Dahlem was presented to the Botanical Museum.
This was a coUection of more than 50,000 specimens of mostly parasitic
fungi from aU over the world (but mainly C. Europe).
The Fungus herbarium of the Botanical Museum comprised about
250,000 specimens according to an estimate of J. Kohlmeyer in
1962. Today there are more than 300,000 specimens.
Lichens
The Lichens of the general herbarium were nearly completely
destroyed, only two loans of several specimens came back to Berlin
after the war (see Appendix B). In addition to this material the smaU
herbaria of J. Lahm, F. W. Zopf (including a coUection of Uchen sub?
stances, cf. Huneck et al. 1973), and H. Zschacke were saved (Mattick
1954).
The largest coUections acquired after the war are those of G. Lettau
(38,000 nos.), O. Behr (11,000 nos.), and V. Grummann (10,000
nos.). Today the Lichen herbarium comprises about 100,000 specimens.
Bryophytes
Only some fascicles of the genera Dicranella and Campylopus were
saved in 1943. The remainder of the general herbarium was destroyed.
Fortunately some important old collections are extant because they
were kept separately. The most valuable herbarium is that of S. E. de
Bridel which is still not included in the general herbarium (Schultze-
Motel 1977). Other old or newly acquired collections came from Andres,
M. Fleischer, Hintze, Loeske, K. Osterwald, H. Reimers
and many others (see Appendix A).
A list of extant types of C. Warnstorf's herbarium was published by
Schultze-Motel (1962), who also reported on a collection of
bryophytes made in 1829 by Ch. G. Ehrenberg and A. von Hum?
boldt in Siberia (Schultze-Motel 1963). The list of an extant old col?
lectionfrom Peru (leg. Raimondi) has just been published (Menzel &
Schultze-Motel1987).
The Bryophyte collection now contains more than 250,000 specimens.
Pteridophyta
The Pteridophyte herbarium is - apart from the small families
Marattiaceae and Ophioglossaceae - completely extant. This collection
comprised over 1,000 fascicles in 1943 and today gives us a good im?
pression of the importance of the old Berlin herbarium. In this group
the list of collections
published by Urb an (1916) is still valid without
any reservations.
In 1961 the number of sheets was estimated to be 275,000 (Meyer
in Potztal 1962: 39; Meyer 1968: 307). Today the fern collection
of the Botanical Museum may be the world's largest herbarium col?
lection of pteridophytes, comprising ca. 300,000 specimens (see also
Morton 1969: 20).
Phanerogams (seed plants)

The surviving parts of the old Phanerogam herbarium are composed
of the following:
a) For various reasons (separate storage, on loan, etc.) the material
of some plant families, genera, and a few entire special collections
escaped the destruction completely or to a great extent; some examples
are Begoniaceae, Pandanaceae, Antirrhinum, Herbarium Willdenow,
collections of cones and fruits, Dingler's Rosa collection, etc. (see Ap-
Englera7- 1987 233
pendices A and B). The extant remnants of larger coUections, such

as C. Koch's herbarium (cf. Lack 1978b), may also be mentioned
here.
b) As has already been mentioned, shortly before the bombing raid
type specimens had begun to be extracted from the general herbarium
and put into the basement of the Museum. Through this action about
20,000 types and other old sheets were saved in families of Gymno-
sperms, Monocotyledons, and some of the Dicotyledons. Unfortu-
nately this activity was not completed. Because not aU types were marked
several were overlooked. On the other hand some of the saved sheets
are not type specimens. Lists of the surviving famiUes were pubUshed
by Sleumer (1949) and Pilger (1953b: 27 f.). The dates of these
pubUcations were checked in the herbarium and are Usted in combi?
nation with other data in Appendix B.
c) A third category of extant material is the group of duplicates of
different collections saved in 1943. Among this material many isotypes
were found, and thus almost aU famiUes contain some old specimens.
A few of these collections include A. Peter, H. J. SchUeben, and A.
Stolz (E. Africa), G. Zenker (Cameroon), C. Ledermann (New Guinea,
Caroline and Palau Is.), M. S. Clemens (New Guinea), and C. Troll
(BoUvia and Africa).
Since the records of the distribution of duplicates from the BerUn

herbarium were also destroyed the pubUcation by Leeuwenberg (1965)
on the distribution of coUections from Africa is very useful in tracing
isotypes from the old Berlin collections. The neotropical types of the
Berlin herbarium ? 15,800 according to MerriU (1943) - were photo-
graphed in the 1930 s by J. F. Macbride for the Field Museum of Na?
tural History in Chicago and these photographs are avaUable there
(Grime & Plowman 1986).
Another source of isotypes present today in the Botanical Museum
was herbaria received after 1943 that contained dupUcates of old BerUn
collections, e.g., the herbaria of R. Gross (see Pilger 1953b: 28), K.
Dinter, H. J. Schlieben, and from the botanical institute of Marburg
University (Pilger l.c: 35).
The pubUshed lists of types from certain families of flowering
plants of the Berlin herbarium show that even in groups known to be
totally destroyed in 1943 some old material can be found: Cucurbita?
ceae (Lack 1978 a), Flacourtiaceae and related famiUes (Zepernick
1978 &1979), and Orchidaceae (Butzin 1978, 1980 & 1981).

Because of the complex situation after the destruction of the her?
barium building the present holdings of the herbarium are extremely
heterogeneous. Besides families containing hundreds of types there are
other groups without hardly any type material.
The holdings of the Phanerogam herbarium at present total ap-
proximately 1,5 million sheets of world-wide collections, but especially
from Europe, S. W. Asia, and Africa. The material is now stored in
compactors in a windowless new building.
Special collections
Since so many types of the general herbarium were destroyed the
extant special collections are very important.
The extensive spirit collection was almost completely saved. The
greater part of this collection comprises specimens of flowering plants
(fruits, large flowers, entire plants, etc). There are also several bottles
of algae, fungi, ferns, and gymnosperms. This special collection contains
many type specimens or fragments of types that were destroyed in the
herbarium (cf. list of type specimens of Cactaceae by Leuenberger
1978 & 1979). The spirit collection comprises ca. 13,000 bottles
(Potztal 1962: 42).
The collection of gymnosperm cones (also twigs and male strobili)
is completely extant (ca. 1,500 nos.).
The old collection of fruits and seeds (ca. 7,000 nos. of dried spe?
cimens) is also largely extant and includes many type fragments. After
World War II a few other collections (Vilmorin-Andrieux, Wittmack,
Schiemann, etc.) came into the possession of the Museum. The
collection of fruits and seeds comprises ca. 15,000 nos.
The greater part of the collection of wood samples has also been
saved. It totals ca. 8,200 samples and comprises several larger extra-
European collections (cf. Potztal 1962: 41 f. and the citation of wood
samples in Brunel et al. 1985).
The collection of galls (cf. Urban 1916: 416) is completely extant
(66 fascicles). Some additional large collections not yet inserted are
those of F. T h o m a s, L. Geisenheyner, H. Harms and others.
There is also a small collection of fossils with about 300 specimens
from Central Europe comprising all periods of geological history. Most
fossils have been in the Museum since before 1943.
Fortunately G. Schweinfurth's amazing collection of vegetal
Englera7-1987 235
remains from Egyptian tombs was saved; it comprises 392 samples

(Germer 1986).
Zepernick (1985) reports on another paleo-ethnobotanical
collection of the Botanical Museum containing plant remains from
NeoUthic and Bronze age lake dwelUngs in C. Europe.
Last but not least two additional extant old coUections should be
mentioned: a rather large collection of resins, balsamic resins, lacs,
copals, latexes, etc. (with many samples from Schweinfurth and
other famous collectors), and a large collection of plant fibres.
Acknowledgements
I express my thanks to those coUeagues at the Botanical Museum BerUn -

Dahlem who gave valuable informations on different parts of the
collections. My sincere thanks are due to Dr. L. E. Skog and Dr. R. A.
DeFUUps, U.S. National Herbarium, who undertook to edit the EngUsh.
References
Anon. 1916: Bericht iiber den Botanischen Garten und das Botanische
Museum zu Berlin-Dahlem vom 1. April 1915 bis zum 31. Marz
1916. - Notizbl. Konigl. Bot. Gart. BerUn-Dahlem 7(61): 398-415.
? 1934: Bericht iiber den BotanischenGarten und dasBotanische Museum
zu Berlin-Dahlem vom 1. April 1933 bis zum 31. Marz 1934. ?
Notizbl. Bot. Gart. BerUn-Dahlem 12(111): 1-28.
? 1937: Bericht iiber den Botanischen Garten und das Botanische
Museum zu BerUn-Dahlem vom 1. AprU 1936 bis zum 31. Marz
1937. - Notizbl. Bot. Gart. BerUn-Dahlem 13(119): 531-561.
? 1940: Bericht iiber den Botanischen Garten und das Botanische
Museum zu BerUn-Dahlem vom 1. AprU 1939 bis zum 31. Marz
1940. - Notizbl. Bot. Gart. BerUn-Dahlem 15(2): 279-301.
Bassler, M. 1970: Das Herbarium des Bereiches Botanik und Arboretum
des Museums fiir Naturkunde an der Humboldt-Universitat zu Ber?
Un. - Wiss. Z. Humboldt-Univ. BerUn, Math.-Nat. R. 19(2/3):
295-299.
Balick, M. J. 1980: The current importance of the Palm Herbarium at

Berlin-Dahlem. - Principes 24: 33-36.
Brunel, J. F., Hiepko, P. & Scholz, H. (eds.) 1985 ('1984'): Flore analy-
tique du Togo. Phanerogames. - Englera 4: 1-751.
Butzin, F. 1978: In Berlin vorhandene Typen von Schlechters Orchi-
deenarten. - Willdenowia 8: 401-407.
- 1980: Typenstudien im Berliner Orchideen-Herbar: Arten von
Kranzlin und Mansfeld. - Willdenowia 10: 147-149.
- 1981: Typenstudien im Berliner Orchideen-Herbar: Diverse mar-
kierte Typen. - Willdenowia 11: 119-121.
Eckardt, Th. 1965: Zum Gedenken an den 200. Geburtstag von Carl
Ludwig Willdenow *22.8.1765 f 10.7.1812. - Willdenowia 4:
1-21.
- 1966: 150 Jahre Botanisches Museum Berlin (1815-1965). - Will?
denowia 4: 151-182.
Ern, H. 1976: Descripcion de la vegetacion montaiiosa en los estados
Mexicanos de Puebla y Tlaxcala con un apSndice "Las plantas Mexi-
canas en el herbario Willdenow de Berlin-Dahlem". - Willdenowia,
Beih. 10.
Gerhardt, E. & Hein, B. 1979: Die nomenklatorischen Typen der von
Th. Nitschke beschriebenen Arten im Pilzherbar des Botanischen
Museums Berlin-Dahlem. - Willdenowia 9: 313-330.
Germer, Renate 1986: Die Pflanzen des alten Agypten. - Botanisches
Museum, Berlin-Dahlem.
Grim6, W. E. & Plowman, T. 1986: Type photographs at Field Museum
of Natural History. - Taxon 35: 932-934.
Hein, B. 1985 ('1984'): Originalmaterial und Hinweise zu den von G.
Otth beschriebenen Fungi-Arten und subspezifischen Taxa im
Herbar des Botanischen Museums Berlin-Dahlem. ? Willdenowia 14:
413-416.
- & Gerhardt, E. 1981: Die nomenklatorischen Typen zu Kirschsteins
Namen von Gattungen, Arten und subspezifischen Taxa im Pilz?
herbar des Botanischen Museums Berlin-Dahlem. ? Willdenowia 11:
123-163.
Hiepko, P. 1969: Von J. R. und G. Forster gesammelte Pflanzen im
Herbar Willdenow in Berlin. - Willdenowia 5: 279-294.
- 1972: Herbarium Willdenow - alphabetical index. - IDC, Zug.
- 1978: Die erhaltenen Teile der Sammlungen des Botanischen Museums
Englera7 - 1987 237
Berlin-Dahlem (B) aus der Zeit vor 1943. - WUldenowia 8:

389-400.
? 1979: Das Schicksal des "Ausserrheinischen Herbariums" des Natur-
historischen Vereins fur die Preussischen Rheinlande und Westfalen.?
WUldenowia 9: 207-208.
? 1980: Das Herbar von Rudolf Baschant. - WUldenowia 10: 83-86.
Huneck, S., Follmann, G. & Redon, J. 1973: Mitteilungen iiber Flech-
teninhaltsstoffe XCVI. Identifizierung einiger Flechtenstoffe aus der
Belegsammlung Friedrich WUhelm Zopfs. ? WUldenowia 7: 31-45.
Jones, Almut G. & Hiepko, P. 1981: The genus Aster s.l. (Asteraceae)
in the Willdenow Herbarium at Berlin. - WUldenowia 11: 343-360.
Julich, W. 1974: Liste der im mykologischen Herbar von C. G. T. Preuss
(t 1855) vorhandenen Arten. - WUldenowia 7: 261-331.
Kohlmeyer, J. 1962a: Die PUzsammlung des Botanischen Museums
zu Berlin-Dahlem (B). - WUldenowia 3: 63-70.
? 1962 b: Index alphabeticus KlotzschU et RabenhorstU herbarU myco-
logici. ? Beih. Nova Hedwigia 4: 1 -231.
Lack, H. W. 1978 a: Typen der Cucurbitaceae im BerUner Generalher?
bar. - WUldenowia 8: 425-429.
? 1978b: Das Herbar C. Koch. - Willdenowia 8: 431-438.
? 1978 c: Die AbbUdungsvorlagen (Skizzen, Herbarbelege) zu Zahns
Hieracium -Monographie. ? WUldenowia 8: 439-442.
? 1979: Die siidamerikanischen Sammlungen von H. Ruiz und Mitar-
beitern im Botanischen Museum BerUn-Dahlem. ? Willdenowia 9:
177-198.
? 1980: Das Herbar der zoologisch-botanischen GeseUschaft in
Wien. - WUldenowia 10: 67-81.
? & Wagner, Daniela 1985 01984'): Das Herbar UUepitsch. - WUlde?
nowia 14: 417-433.
Leeuwenberg, A. J. M. 1965: Isotypes of which holotypes were destroyed
in Berlin. - Webbia 19: 861-863.
Leuenberger, B. E. 1978: Type specimens of Cactaceae in the Berlin-
Dahlem Herbarium. - Cact. Succ. J. Gr. Brit. 40: 101-104.
? 1979: Typen der Cactaceae in der Nasspraparate-Sammlung des Ber?
Uner Herbars. - Willdenowia 8: 625-635.
Mattick, F. 1954: Verluste der deutschen Lichenologie 1943-1951:
J. HUlmann, C. F. E. Erichsen, G. Lettau und H. Sandstede. - WUl?
denowia 1: 175-197.
Mendes, E. J. & Balsas, J. 1981: Type numbers of the H. J. SchUeben
collection known to exist at LISC, 1980. - Bol. Soc. Brot., s6r. 2,
53: 1223-1231.
Menzel, M. & Schultze-Motel, W. 1987: Studies on Peruvian Bryophytes

II. The mosses collected by Antonio Raimondi between the years
1868 and 1879. - WUldenowia 16: 511-525.
Merrill, E. D. 1943: Destruction of the Berlin Herbarium. - Science
98 (2553): 490-491.
Meyer, D. E. 1968: Systematik der Farnpflanzen. ? In: Fortschritte der
Botanik 30: 302-313. - Berlin.
Morton, C. V. 1969: The Fern collections in some European herbaria
II: Botanical Museum and Garden, Berlin. ? Amer. Fern. J. 59:
11-22.
Pilger, R. 1953a: Bericht iiber den Botanischen Garten und das Bo?
tanische Museum Berlin-Dahlem vom 1. Marz 1943 bis 31. Marz
1947. - [Willdenowia] Mitt. Bot. Gart. Berlin-Dahlem 1: 1-21.
? 1953b: Bericht iiber den Botanischen Garten und das Botanische
Museum Berlin-Dahlem, April 1947 - Dezember 1948. - [Willde?
nowia] Mitt. Bot. Gart. Berlin-Dahlem 1: 22-31.
Potztal, Eva 1962: Bericht iiber das Botanische Museum und den Bota?
nischen Garten Berlin-Dahlem fiir das Jahr 1961. ? Willdenowia
3:33-62.
Rosenthal, Margarete 1956: Nachruf auf den Mykologen Wilhelm
Kirschstein (1863-1946). - Willdenowia 1: 606-611.
Scholz, H. 1977: Notizen zu einigen Axonopus-Arten (Gramineae-
Paniceae) nach Berliner Typenmaterial. ? Willdenowia 8: 93-99.
? 1979: Die Typen der Namen der von Alexander Braun beschriebe?
nen Penicillaria-Taxa. (Pennisetum americanum). ? Willdenowia 9:
203-206.
Schultze-Motel, W. 1960a: Georg Kiikenthal (1864-1955). - Willde?
nowia 2: 316-373.
? 1960b: Afrikanische Cyperaceae aus den Sammlungen von A.
Peter. - Willdenowia 2: 495-518.
? 1962: Das Moosherbar von Carl Warnstorf. -Willdenowia 3: 289-313.
? 1963: Bryologische Ergebnisse der Reise von Alexander von Hum?
boldt, Ehrenberg und Rose in den Ural und nach Sibirien (1829). ?
Nova Hedwigia 5: 79-90.
? 1977: Bryology at the Botanical Museum, Berlin-Dahlem. ? Taxon
26: 150-151.
Shetler, S. G. 1969: The herbarium: past, present, and future. ? Proc.
Biol. Soc. Wash. 82: 687-758.
Sleumer, H. 1949: The Botanical Gardens and Museum at Berlin-Dah?
lem. - Kew Bull. 1949: 172-175.
Englera7-1987 239
Stafleu, F. A. 1972: The WUldenow Herbarium. - Taxon 21: 685-688.

- 1981: Engler und seine Zeit. - Bot. Jahrb. Syst. 102: 21-38.
- 1987: Die Geschichte der Herbarien. - Bot. Jahrb. Syst. 108:
155-166.
Timler, F. K. & Zepernick, B. 1987: The colonial botany of Ger?
many. ? Ber. Deutsch. Bot. Ges. 100 (in press).
Urban, I. 1881: Geschichte des Konigl. botanischen Gartens und des
Konigl. Herbariums zu Berlin nebst einer Darstellung des augen-
bUckUchen Zustandes dieser Institute. ? Jahrb. Konigl. Bot. Gart.
Berlin 1: 1-164.
- 1909: Kurze Geschichte der Entwickelung der Sammlungen. - In:
Der KonigUche Botanische Garten und das Konigliche Botanische
Museum zu Dahlem: 11 -30. ? BerUn.
? 1916: Geschichte des KonigUchen Botanischen Museums zu BerUn-
Dahlem (1815-1913) nebst Aufzahlung seiner Sammlungen. ? Dres?
den. (Also in Beih. Bot. Centralbl. sect. 1, 34: 1-457. 1917.)
Wagenitz, G. 1960: Joseph Bornmuller 1862-1948. - WUldenowia
2: 343-360 (Das Herbar Bornmuller: 357-360).
? 1962:Pflanzen von der Orientreise Tourneforts im Herbar WUlde?
now in BerUn. ? WUldenowia 3: 109-136.
Werdermann, E. (ed.) 1954: Beitrage zur Flora von Siidwestafrika ? 1.
Gramineae von R. PUger f. - WUldenowia 1: 198-274.
Zepernick, B. 1978: Typen und Typoide der Flacourtiaceae im General?
herbar des Botanischen Museums BerUn-Dahlem. - WUldenowia
8:409-424.
? 1979: Typen der FlacourtUnae (ausgenommen Flacourtiaceae) im
Generalherbar des Botanischen Museums BerUn-Dahlem. - WUlde?
nowia 9: 203-206.
- 1985: ('1984'): Material aus den Pfahlbauten Robenhausen, M6-
ringen, Schussenried und Mondsee im Botanischen Museum BerUn-
Dahlem. - WUldenowia 14: 405-408.
? & Timler, F. K. 1979: Grundlagen zur 300jahrigen Geschichte des
BerUner Botanischen Gartens. ? Englera 1: 5-303.

Prof. Dr. Paul Hiepko, Botanischer Garten und Botanisches Museum
Berlin-Dahlem, Konigin-Luise-Strafce 6-8, D-1000 Berlin 33.
Appendix A
Selective list of collections acquired by the Botanical Museum Berlin-

Dahlem (B) between 1914 and 1986
The following list contains the more important and larger collections
acquired by the Botanical Museum Berlin-Dahlem (B) between 1914
and 1986 and assigned to certain collectors (or owners). It was not
possible to include all collectors. It should be borne in mind that col?
lections mentioned
as "original herbarium" very often contain spec?
imens collected by other persons than the owner. Additional names
of collectors represented at B are given in Hiepko (1980), Lack (1980)
and Lack & Wagner (1985).
Some collections from the time before 1914 (see lists in Urban
1916) are mentionedif they were at least partly saved in 1943. The
collections made by the present staff members are not included.
In accordance with the Collectors Index (part II of Index Herbanorum)
the indication "r." in front of a year marks the date at which the
collection was received at the Museum. Thus, it is possible to see whether
the material was destroyed in 1943. If material received before March
1943 was saved, this fact is indicated. It should be stressed again
that in the groups listed in Appendix B certain specimens from almost
all old collections are extant.
The dates of receipt are taken from the published annual reports
which comprise the period from April 1 of each year to March 31 of
the following, i.e. the financial year formerly used. The dates given
refer to the first year of the period mentioned. Thus, all collections
received since 1943 (r. 1943 and later) came to the Botanical Museum
after the destruction of the herbarium building.
For the selected Exsiccatae of different groups of Cryptogams
included in the list, the dates of receipt are not mentioned; these
collections are all extant.
Allorge, P.: Bryotheca iberica, no. 1-150. Paris 1928-1938.

Allorge, V. & P.: Bryophyta azorica. Paris 1939.
Andres, H.: Bryophyte herbarium, r. 1968 (t 18,000, mostly Europe,
especially Rhineland).
Ascherson, P.: Small parts of the original herbarium (r. 1895) extant.
Barros, J. J. de: Portugal, r. 1939 (? 2,000, partly extant).
Englera7 - 1987 241
Baschant, R.: Original herbarium, r. 1959 (? 25,000, mostly C. & S.

Europe, some Senegal, Brazil and specimens of other continents [ex-
change]; many duplicate of old collections, cf. Hiepko 1980).
Becker, W.: Viola herbarium, r. 1929 (? 4,300).
Beger, H.: Original herbarium, r. 1941 (? 12,000, col. 1906-1939, most?
ly C. Europe), almost completely extant.
Behr, O.: Lichen herbarium, r. 1962 (? 11,000).
Berger-Landefeldt, U.: Original herbarium, r. 1968 (? 23,000 + 686
Cryptogams; mostly C. and S. Europe).
Beyer, R.: Original herbarium, r. 1925 (over 30,000, C. Europe, espe?
ciaUy WestphaUa), partly extant.
Borgmann, E.: N. E. New Guinea (Mt. Wilhelm) col. 1960/61, r. 1971
(? 950).
Bornmuller, J.: C. & S. Europe, r. 1914-1937 (3,002, incl. 2,150 nos.
Hieracium)-, Macedonia, r. 1919-1924 (3,331 ex herb. HBG); Iter
graecum a. 1926, r. 1927 (504); Iter anatoUcum quintum a. 1929,
r. 1931 (621); Balkan a. 1931, r. 1932 (79); TripoU & S. Italy
a. 1933, r. 1934 (381). - The private herbarium of J. Bornmuller
("ca. 600 sehr starke Faszikel") was purchased for the Botanical
Museum in 1938 and moved to BerUn in several parts since that
year. The material received before March 1943, therefore, has been
largely destroyed (Ust of destroyed families in Wagenitz 1960:
358). However, the greater part of the herbarium came to BerUn
after World War II and is extant. Most plants were coUected by
Bornmuller himself in S. W. Asia, S. & C. Europe, and Macaronesia;
many specimens are from other coUectors: e.g. Th. Strauss, Gabriel,
Gauba, Sintenis and several Russian coUectors (see also Wagenitz
1960).
Bothe, H.: Original herbarium, r. 1944 (? 250 fasc, mainly C. Europe,
including many different coUectors).
Brade, A. C: Costa Rica and Brazil, r. 1921-1939 (3,079), incl. 500
ferns).
Brass, L. J.: New Guinea, r. 1928-1939 (914).
Bridel, S. E. de: Bryophyte herbarium, r. 1829 (? 1,000 species,
extant and stiU kept separately).
Carr, C. E.: New Guinea, r. 1935/36 (5,032, many specimens extant).
Clemens, J. & M. S.: N. Borneo, r. 1931-1936 (6,904), r. 1944 (1,328);
New Guinea, r. 1935-1939 (8,772 of which? 2,600 nos. with
duplicates were saved).
Crouan, H. M. & P. L.: Aigues marines du Finistere. 3 fasc. (404 nos.).

Degener, O. (p.p. with Degener, L): Hawaii, r. 1928-1938 (1,930);
N. America, r. 1933 (902); Hawaii, USA, Fiji, Australia, New Zea?
land, USSR, r. 1950-1986 (3,468).
Desmazieres, J. B. H. J.: Plantes cryptogames de France. Fasc. 1-16
(no. 1-800). Lille 1853-1860.
Diels, L.: Europe, r. 1919-1944 (217); N. America, r. 1933 (345);
Ecuador,r. 1933(925).
Dingler, H.: Original herbarium, r. 1914 (? 30,000, Anatolia, Syria,
Palestine, and S. Germany; incl. herb. Spruner, Greece), "Flora
orientalis" col. 1873-1874 partly extant. - Rosa herbarium, r.
1936 (?14,000, chiefly collected by Dingler himself, C. & S. Europe,
completely extant).
Dinter, K.: S. W. Africa, r. 1929-1941 (1,565). - Original herbarium
S. W. Africa), r. 1946 (? 8,000).
Domke, W.:Cameroon, r. 1938 (? 1,500); N. W. Libya,r. 1939 (? 500).
Drege, J.: Original herbarium, r. 1915 (? 8,000, ex herb. Liibeck;origi-
nally belonging to E. H. F. Meyer, Konigsberg, 11858).
Eckardt, Th.: Switzerland (chiefly Puschlav), S. Europe, Madeira
Is.,USAr. 1977 (? 3,150).
Ehrenberg, Ch. G.: Ural & Sibiria, col. 1829: 216 Bryophytes extant
(Schultze-Motel 1963).
Ekman, E. L.: Weast Indies, r. 1915-1935, via I. Urban (24,127).
Elmer, A. D. E.: Philippines, r. 1924& 1938(2,900);Borneo,r. 1925/26
(? 1,800).
Engelhardt, V.: Original herbarium, r. 1943 (? 20,000), mainly collected
himself: Triest, Istria, Dalmatia, Italy, S. E. Alps, Rhodos.
Fahrendorf, E.: Original herbarium: Fungi, r. 1960 (? 4,800).
Fiedler, O.: Original herbarium, r. 1950-1965 (7,100, Europe and from
overseas).
Fleischer, M.: Macedonia, r. 1917 (1,800, incl. 600 Bryophytes); r.
1919-1927 (600, different origin). - After World War II a large
amount (over 2,000 nos. with many duplicates) of mostly unde-
termined mosses from tropical Asia, Oceania, Australia, and New
Zealand have been found; this material obviously was intended for
a continuation of Fleischer's exsiccatae "Musci (frond.) archipelagi
indici (et polynesiaci)" which are nearly complete at B.
Fries, E. M.: Scleromyceti Sueciae. Lundae 1819-1834 (? 200).
Englera7- 1987 243
Gabriel, A. & A.: Iter irano-gedrosiacum a. 1937 (many specimens

ex herb. Bornmuller).
Gauba, D. E.: Plantae Gaubeanae iranicae 1935 (many specimens ex
herb. Bornmuller).
Geisenheyner, L.: CoUection of galls, r. 1921; completely extant.
Gluck, H.: Original herbarium, r. 1940 (71 fasc: Europe, 53 fasc:
N. America; 429 boxes fruits of Trapa). The Trapa fruit collection
is extant.
Gossweiler,J.: Angola, r. 1924-1939(? 2,100);r. 1950(571).
Gross, L.: Original herbarium, r. 1929 (350 fasc, mostly Europe).
Gross, R.: Original herbarium, r. 1946 (? 20,000, including 3,000
Cyperaceae with many isotypes from B and other important col?
lections, e.g. 600 leg. Jurgens (S. Brazil), 800 leg. Stolz (E. Africa).
Grummann, V.: Original herbarium: Lichens, r. 1968 (? 10,000).
Hamel, G. (p.p. & Hamel-Joukow, A.): Aigues de France. 5 fasc.
1927-1931 (250). - Aigues des AntiUes francaises. 1931 (150).
Heinrichs, E.: Ecuador, r. 1932-1936 (1,000).
Hennings, P.: Original herbarium: Fungi, r. 1887-1908, over 1,000
types and other specimens extant (from holdings of the old pubUc
department of the Botanical Museum and saved groups of Fungi).
Herter, W. G.: Fungi, r. 1916 (? 2,650, Europe and S. America).
Phanerogams, r. 1917-1937 (3,900, Europe, N. Africa, S. America)
+ 1,300 Cryptogams and 1,080 ferns.
HUimann, J.: Original herbarium: Lichens, r. 1944 (14,000); the
collection was evacuated to Buckow (east to BerUn) and was de?
stroyed there in 1945.
Hintze, F.: Original herbarium: Bryophytes, r. 1961 (7,000, Pome-
rania).
- Mosses,
Hoppe, D. H.: Fungi epiphylU. No. 1-10. Ratisbonae 1809.
r. 1944 (? 300, including types of Hoppe & Hornschuch).
Hu,H. H.: China, 1921-1929 (6,800).
Huber, H.: Fungi, mostly Basidiomycetes, r. 1913-1943 (3,678, Austria).
Hubschmann, A. v.: Bryophytes, r. 1982 ff. (3,496, Europa, Macaro-
nesia, Ceylon; including specimens of other coUectors).
Husnot, P. T.: Musci GaUiae, 12 fasc. 1870 ff., r. 1944 (600).
Huth, E.: Original herbarium, r. 1941 (190 fasc, Europe and Orient).
Jahn, E.: CoUection of Myxophyta, r. 1944 (? 3,000).
Jurgens, C: S. Brazil, r. 1923-1934 (? 750); ca. 600 ex herb. R. Gross
(r. 1946).
Kalbreyer, W.: Herbarium, r. 1915 (276, W. Africa col. 1877; 1,650,

Colombia col. 1877-1884, many palms).
Kinscher, H.: Rubus herbarium, r. 1918 (33 large fascicles, mostly
C. Germany), completely extant.
Kirschstein, W.: Original herbarium: Fungi, r. 1945 (10,000, chiefly
Ascomycetes), cf. Rosenthal (1956), Hein & Gerhardt (1981).
Klautke,P.: China and Korea, r. 1925-1929(1,572).
Klotzsch, J. F.: Herbarium vivum mycologicum. 14 Cent. (incomplete),
cf. Kohlmeyer (1962b).
Koch, C: Original herbarium, r. 1913. For saved parts see Lack (1978b).
Koehne, E.: Dendrological herbarium, r. 1919 (22,000, mostly cult.).
Kornicke, F. A.: Original herbarium, r. 1934 (? 20,000, mostly C.
Europa, including Fungi). ? Europe, r. 1983 (? 4,200 Phanero-
gams,? 1,100 Fungi).
Kopsch, A.: Bryotheca saxonica. 1919 ff. No. 1-300.
Krause, A. & A.: Alaska, r. before 1914 (360 extant, on loan during
the war).
Krause, E. H. L.: Original herbarium, r. 1929 (? 60 fasc);Bryophyte
herbarium, r. 1935 (? 2,000). - Fungi, r. 1941 (22 fasc. extant,
cf. Kohlmeyer 1962 a).
Krause, K.: AnatoUa and other regions, r. 1926-1940 (5,478).
Krieger, K. W.: Fungi saxonici. Fasc. 1-50 (no. 1-2,500). Konigstein/
Elbe 1885-1919.
Kukenthal, G.: Caricoideae herbarium, r. 1931; remaining Cyperaceae,
r. 1944 and 1953 (? 6,000, cf. Schultze-Motel 1960a); Rubus,
r. 1948(6,540).
Kiitzing, F. T.: Algae, 600 sheets of the orig. herb. purchased in 1894
have been saved (but no types).
Lahm, J.: Lichen herbarium, r. 1936 (ex herb. Munster), extant.
Ledermann, C: 950 duplicates of the collection received before 1914
were saved (New Guinea, Carolines, Palau Is.).
Lettau, G.: Original herbarium: Lichens, r. 1952 (38,000, incl. 1,000
Fungi; chiefly East Prussia and S. W. Germany) + 38 fasc. Phanero?
gam s.
Loeske, L.: Bryophytes, r. 1934 (250); today many Bryophyte collec?
tions at B which came to Berlin with other herbaria.
Ludwig, A.: Original herbarium, r. 1958 (305 fasc. Phanerogams;
92,000 Cryptogams, including ca. 50,000 Fungi), mostly C. Europe.
Englera 7-1987 245
Markgraf, F.: Balkan (Albania and N. Greece), r. 1924-1941 (3,000);

Brazil, r. 1939 (? 900).
Mattfeld, J.: Balkan (Bulgaria, Greece, Turkey-in-Europe), r. 1924-
1930 (? 5,800); C. Europe, mostly Alps, r. 1921-1939 (? 2,000).
Mattick, F.: Lichens: Brazil, r. 1953 (1,070); Chile, r. 1961 (? 1,500).
Mauczka, V.: Original herbarium, r. 1944, from Vienna (120 fasc,
many cult.).
Melchior, H.: S. Alps, r. 1929-1940 (7,778); N. Iran (Elburs), r. 1936
(650).
Menzel, P.: Leaf herbarium, r. 1928 (?) (36 fasc, leaves mostly ex herb.
Berlin), completely extant.
Meyer, D. E.: C. Europe, r. 1955-1973 (3,900, predominantly Pteri-
dophytes).
Mez, C: Original herbarium, r. 1933 (125 fasc, including many types:
Gramineae, Bromeliaceae, Lauraceae, Myrsinaceae).
Morton, F.: Original herbarium, r. 1962 (? 9,000, Alps, Guatemala,
Ethiopia).
Miiller, O.: Diatoms, r. 1914 (? 3,000).
Nitschke, Th.: Original herbarium: Fungi, r. 1936 (ex herb. Munster),
extant, cf. Gerhardt & Hein (1979).
Nordstedt, O. & Wahlstedt, L. J.: Characeae Scandinaviae exsiccatae,
3 fasc. (120 nos.).
Osterwald, K.: Original herbarium: Bryophytes, r. 1923 (large collection,
C. Europe, chiefly Brandenburg and Alps), completely extant.
Otth, G.: Fungi, ca. 100 boxes ex herb. Kornicke and herb. Nitschke;
cf. Hein (1985).
Peekel, G.: Neumecklenburg (New Ireland), r. 1908-1938 (? 1,200).
Peter, A.: S. & E. Africa (Tanzania), r. 1935 (47,000),? 28,000 nos.
with many duplicates extant (cf. Schultze-Motel 1960b).? Hieracium
collection, r. 1937.
Pilger, R.: Mattogrosso, r. 1944 (250 ex herb. Jena).
Polunin, O.: Kashmir (and Libanon), r. 1959 (849); Pakistan, r. 1961
(439).
Prelinger, M.: Original herbarium, r. 1944 (65 fasc, C. Europe, Austria).
Preuss, C. G. T.: Original herbarium: Fungi, r. 1936 (1,848, ex herb.
Munster), extant, cf. Kohlmeyer 1962; Jiilich 1974.
Pritzel, E. G.: W. Australia, col. 1910: many duplicates extant.
Rabenhorst, L. & Pazschke, O.: Fungi europaei et extraeuropaei. Cent.
1-50. Dresdae/Lipsiae 1859-1905 (1-4,500), nearly complete.
Raimondi, A.: Peru herbarium p.p., r. 1930 (some types and 230 Bryo?
phytes extant, cf. Menzel & Schultze-Motel 1987).
Rambo, B.: S. BrazU, "Herbarium Anchieta", r. 1954-1964 (? 8,000).
Range, P.: Sinai, r. 1916 (304); Nigeria, r. 1929 (31); S.W. Africa,
r. 1940 (? 600). - Original herbarium, r. 1951-1953 (? 8,000,
C. Europe).
Rechinger, K. H.: Greece, r. 1939 (1,437); Greece, r. 1955-1984
(? 3,600); Iran,r. 1970-1984 (? 9,000).
Reimers, H.: Cameroon, r. 1928 (681). - Original herbarium (C. Eu?
rope), r. 1961 (Algae: 2 fasc, Fungi: 53 fasc, Bryophytes: 100 fasc,
Phanerogams: 61 fasc).
Reitz, R. & Klein, R. M.: Brazil, r. 1959-1969 (2,553).
Reuss, A. L. v.: Original herbarium, r. 1944 from Vienna (30,000,
Austro-Hungarian Empire).
Roemer, F.: Original herbarium, r. 1939 (100 fasc, incl. 28 fasc.
Rosa; chiefly from Pomerania).
Roth, A. W.: (1757-1834): Original herbarium, r. 1925, ex Natur-
historisches Museum zu Oldenburg (10,000, with many duplicates
from botanists of his time).
Rottenbach, H.: Original herbarium, r. 1949,exNaturwissenschaftUches
Museum Coburg (Europe, Phanerogams: 5,600, Fungi & Lichens:
400; many different coUectors).
Ruiz, H. et al.: Peru and ChUe (specimens in Herb. WUldenow [B-W:
109] and in the general herbarium [B:? 350]; cf. Lack 1979).
Sandstede, H.: Cladonia herbarium, r. 1934 (ca. 60,000, mostly Eu?
rope). - After 1943: Cladonia exsiccatae p.p. ex herb. Th. Rein-
stein and herb. H. Zschacke.
Schack, H.: Hieracium herbarium, r. 1934 (? 6,000), completely
extant.
Schiffner, V.: Original herbarium, r. 1944 (Algae, Lichens and Bryo?
phytes).
Schlechter, R.: S. Africa, r. 1961, ex herb. Zurich, Z (711).
Schlichtkrull, P.: Original herbarium, r. 1969 (? 3,000, C. & S. Europe).
Schlickum, A.: Hieracium herbarium, r. 1949 (4,168).
SchUeben, H. J.: E. Africa (Tanzania), r. 1931-1935 (7,962 nos. with
10 duplicates each). - Original coUection: E. Africa, r. 1944(3,532);
S. Africa, r. 1956 (796); Madagascar, r. 1960 (240); Madagascar
and S.W. Africa, r. 1968 (280). - Mendes & Balsas 1981: List of
isotypes present in LISC.
Englera 1- 1987 247
Schmid, B.: India, Nilgiri Hills, col. 1831-1837, r. 1932, ex herb

Jena(? 2,000).
Schneider, C. K.: China, r. 1919-1921 (3,616). - Balkan, col. 1907,
extant (ex herb. Bornmuller).
Schultze-Rhonhof, A. & H.: Ecuador, r. 1937-1939 (1,497).
Schulz, O. E.: C. Europe, r. 1944 (20 fasc).
Schulz, Roman (brother of O. E. Schulz): Original herbarium, r. 1945,
ex Markisches Museum Berlin (large collection, chiefly Branden?
burg and Alps).
Schulze, M. (Jena): Original herbarium, r. 1915 (? 30,000, Europe,
many Orchids and Rosae). Partly extant, e.g. Potentilla spp.
Schwarz, O.: S. Carpathian Mts., r. 1928 (690); W. Anatolia 1932/
33, r. 1939 (820, many specimens extant).
Schweinfurth, G.: Original herbarium, r. 1925 (over 18,000). Many
specimens from Egypt extant (ex herb. Bornmuller, Muschler etc.).-
The collection of vegetal remains from Egyptian tombs extant.
Sehnem, A.: S. Brazil, r. 1963/64 (? 450, Pteridophytes + many
Phanerogams).
Seydel, R.: S.W. Africa, r. 1954-1967 (? 2,000 nos. with duplicates).
Siehe, W.: Anatolia, r. 1895-1928 (?2,000), 647 extant.
Sin, S. S.: China, r. 1927-1929 (3,317).
Sintenis, P.: Puerto Rico 1886: partly extant; Balkan and Orient
1889-1896: many specimens extant (ex herb. Bornmuller and
herb. Ullepitsch).
Sponheimer, J.: Original herbarium: Fungi, r. 1939 (? 9,000, Europe,
mainly W. and S. Germany).
Spruner, W. v.: Greece, r. 1914 (with Dingler's herbarium).
Steinbach, J.: Bolivia, r. 1925-1929 (? 7,100).
Stolz, A.: E. Africa (Nyassa Highland), col. 1909-1913 (many dupli?
cates extant and? 800 ex herb. R. Gross, r. 1946).
Strauss, Th.: Iran (large collection extant, ex herb. Bornmuller).
Sydow, H.: Fungi exotici exsiccati. (Nos. 1-1,300).
Tessmann, G.: Peru, r. 1924-1926 (2,545).
Thomas, F.: Collection of galls, r. 1920 (?6,000, completely extant);
Fungi, r. 1921 (20 fasc).
Touton, K.: Hieracium herbarium, r. 1929 and 1936 (? 19,000, com?
pletely extant).
Treviranus, L. C: Original herbarium, r. 1936 (ex herb. Naturhisto-
rischer Verein fiir die Preussischen Rheinlande und Westfalen, Bonn

(? 30,000). This herbarium is now at the Humboldt University
Berlin (BHU), cf. Bassler 1970 and Hiepko 1979.
Troll, C: BoUvia 1926-1929,. r. 1930 (3,400); NE., E. & S. Africa, r.
1933-1937 (1,840); Nanga Parbat, r. 1937 (1,194). - Extant
duplicates: Bolivia (? 2,300), Africa (? 630).
Turckheim, H. v.: Original herbarium, r. 1920 (90 fasc, chiefly Guate-
mala and West Indies).
Ule, E.: Original herbarium, r. 1915 (Europe: 1,300, Brazil and Peru:
2,500; Cryptogams: 800).
UUepitsch, J.: Original herbarium p.p., r. 1944, from the botanical
institute of the Vienna University (15,000, Austria, Balkan etc),
cf. Lack & Wagner 1985 (with Ust of coUectors).
Utsch, J.: Original herbarium, r. 1916 (? 20,000, Europe and N.
America; including 1,300 Rubi from WestphaUa).
Vaupel,F.K. J.:Samoa, r. 1905-1919 (? 1,300), partly extant (? 600).
Volk, H.: Afghanistan, r. 1952 (? 1,400).
Wagenitz, G.: Anatolia (with H.-J. Beug) 1957 (527); C. Europe,
some from Lapland, r. 1959-1969 (1,036); Tunisia 1968 (279).
Walter, H. & E.: Original herbarium, r. 1970 (12,200, Australia, Africa,
Europe, AnatoUa, S. America).
Wangerin, W.: Original herbarium, r. 1939 (15,000, mostly own collec?
tions, Europe).
Warnstorf, C: Bryophytes, r. 1944 (?), probably with the herbarium
of his son J. Warnstorf (? 3,000, including many types, cf. Schultze-
Motel 1962).
Warnstorf, J.: Original herbarium, r. 1944 (Bryophytes: 112 fasc,
other Cryptogams: 20 fasc).
Werdermann, E.: Chile, r. 1924-1927 (1,199, duplicates of 989 nos.
extant); BoUvia, r. 1928 (689, duplicates of 310 nos. extant);
Brazil, r. 1932 (1,291). - S. & S.W. Africa (with H. D. Oberdieck),
col. 1958/59 (2,457). - For types of Cactaceae in the spirit collec?
tion see Leuenberger (1978, 1979).
WUldenow, C. L.: The historical Herbarium WiUdenow (r. 1818) con-
taining 20,000 species and ca. 38,000 sheets was saved; see part
1 of text and Hiepko 1972.
Winter, G.: Fungi, r. 1875-1888, many specimens extant in the saved
groups of Fungi (Uredinales and Fungi imperfecti).
Englera 7-1987 249
Wittrock, V. & Nordstedt, O.: Algae aquae dulcis exsiccatae. 35 fasc,

1877-1903(1,612).
Wolff, H.: Herbarium p.p., r. 1915 (? 7,000, mostly Europe), material
partly extant.
Zahn, K. H.: Hieracium herbarium, r. 1941 (25,000, incl. ? 2,800
S. Germany and W. Alps). - Vouchers of illustrations for the
types;
monograph of Hieracium extant (cf. Lack 1978 c).
Zopf, F. W.: Lichens and lichen substances, r. 1936 (ex herb. Munster);
the material is extant (cf. Huneck et al. 1973).
Zschacke, H.: Lichens (Corsica, Switzerland), r. 1920-1926 (721);
r. 1948(400).
Appendix B
Extant collections of the Botanical Museum Berlin-Dahlem (B) from the

time before 1943
In the following list the saved material of all groups of the general
herbarium is compiled. It should be borne in mind that in almost all
families of Phanerogams additional type material from other sources
can be found. The material of the saved special collections was not
taken into account.
types = types (and/or other old specimens) extant

loan = specimens on loan during the war, returned to Berlin between
1951 and 1983
Algae Pteridophytes
Ca. 600 sheets of herb. Kutzing The entire Pteridophyte her?
(received 1894) extant; loan: barium - except Marattiaceae
57 sheets and Ophioglossaceae ? extant
Fungi Phanerogams
Uredinales and Fungi imperfecti Acanthaceae, loan: 311 sheets
completely extant; furthermore Achatocarpaceae, types
several types kept in the public Aizoaceae, types
department of the Botanical Mu? Amaranthaceae, types; loan: 119
seum and in the herbaria of sheets
Nitschke, Preuss, etc. extant (see Amaryllidaceae, types; loan: 309
App. A) sheets
Annonaceae, types; loan: 545
Lichens
sheets
Herbaria of Lahm, Zopf, and
Apiaceae ? see Umbelliferae
Zschacke extant (see App. A);
Aponogetonaceae, types
loan: 2246 (mainly Lecanora,
Araceae, types
Lecidea, Verrucaria); types of
Arecaceae - see Palmae
Stictaceae
Asclepiadaceae, loan: 125 sheets
Bryophyta (Dischidia, Hoya, mostly types
Herbarium Bridel completely ex? of Schlechter)
tant Asteraceae ? see Compositae
Dicranaceae, some fasc (Campy- Balsaminaceae, material ? excl.
lopus and Dicranella p.p.) extant Eurasia - extant
Englera 7- 1987 251
Begoniaceae, completely extant of A. Peter see Schultze-

(on loan during the war at Motel 1960 b
Irmscher) Dioscoreaceae, types
Berberidaceae, types Droseraceae, types
Bignoniaceae, 38 sheets
loan: Elaeocarpaceae, types of Knuth
Brassicaceae ? see Cruciferae (leg. Clemens, New Guinea)
Bromeliaceae, types Eriocaulaceae, types
Bruniaceae, types Euphorbiaceae, loan: 208 sheets
Burmanniaceae, types Gentianaceae, loan: 179 sheets
Cactaceae, types in the spirit (Centaurium)
collection (Leuenberger 1978 Gnetaceae, types
& 1979) Gramineae, types (p.p., excl.
Campanulaceae, loan: the greater Bambuseae); loan: 476 sheets.
part of Lobelioideae (30 Types of Axonopus: Scholz
fasc) 1977; types of Penicillaria:
Capparaceae, types Scholz 1979
Casuarinaceae, types Grubbiaceae, types
Centrolepidaceae, types Gyrostemonaceae, types
Cephalotaxaceae, types Hamamehdaceae, types
Ceratophyllaceae, types Hernandiaceae, types
Commelinaceae, types p.p. Hydrocharitaceae, loan: 20 sheets
Compositae, loan: 82 sheets (Ottelia)
(Senecio). Several old Hiera- IlUciaceae, types
cium collections extant, e.g. Iridaceae, types; loan: 482 sheets
Schack and Touton (see App. Labiatae, loan: 271 sheets (Thy-
A) mus & Mentha)
Convolvulaceae, loan: 25 sheets Lactoridaceae, types
(S. America) Lamiaceae ? see Labiatae
Corsiaceae, types Lauraceae, types p.p.maj. (types
Crassulaceae, loan: 859 sheets of some neotropical genera
Cruciferae, types missing)
Cunoniaceae, types Lihaceae, types
Cyperaceae, types p.p. (Cyperus Loranthaceae, types
s.l. and Eleocharis p.p.),many Magnoliaceae, types
isotypes ex herb. R. Gross. Malvaceae, loan: 10 sheets(Noto-
On the Cyperaceae herb. of triche)
G. Kiikenthal see Schultze- Marantaceae, types p.p.
Motel 1960 a; on Cyperaceae Mayacaceae, types
Meliaceae, old material incl. (Ranunculus)

many isotypes from Reliquiae Rapateaceae, types
Harmsianae Resedaceae, types
Menispermaceae, types Restionaceae, types
Misodendraceae, types Rhamnaceae, loan:78 sheets
Moraceae, types; loan: 198 sheets Rosaceae, types p.p.min.; loan:
(Sorocea) 660 sheets (Rosa); Dingler's
Musaceae, types p.p.min. Rosa herbarium completely
Myrothamnaceae, types extant
Nepenthaceae, types Santalaceae, types
Nyctaginaceae, types Sapindaceae, more than 100 du?
Nymphaeaceae, types; loan: 36 plicates of S. American col?
sheets lections
Octocnemataceae, types Saxifragaceae, types p.p. (e.g.
Olacaceae, types Escallonia, Polyosma, Quin-
Onagraceae, loan: 964 sheets tinia)
(Jussiaea) Schisandraceae, types
Opiliaceae, types Scrophulariaceae, loan: 1598
Orchidaceae, some types (cf. sheets (incl. aU material of
Butzin 1978, 1980 & 1981) Antirrhinum and Kickxia)
Palmae, collection of palm fruits Simaroubaceae, loan: 31 sheets
completely extant; the greater Solanaceae, loan: 37 sheets
part of the herbarium spe? Sparganiaceae, types
cimens is also extant (cf. Stemonaceae, types
Balick 1980) Taccaceae, types
Pandanaceae, all material extant Taxaceae, types
Papaveraceae, types Tovariaceae, types
Phytolaccaceae, types Typhaceae, types
Piperaceae, types Ulmaceae, types
Poaceae ? see Gramineae UmbelUferae, loan: 28 sheets
Podocarpaceae, types (Ferula)
Podostemonaceae, types Urticaceae, loan: 258 sheets
Polygonaceae, types Velloziaceae, many old speci?
Portulacaceae, types mens extant (incl. types)
Proteaceae, types Winteraceae, types
Ranunculaceae, loan: 16 types Xyridaceae, types
(Clematis, Delphinium from Zingiberaceae, types (p.p.) and
tropical Africa); 643 sheets other old material
Englera 7-1987 253
H. Walter Lack
Opera magna in Berlin plant taxonomy
Chicago on the Spree
"Chicago on the Spree" was one of Berlin's nicknames at the turn of the
century; brief and concise, it expresses several basic characteristics of this
metropolis: it is a young city of immigrants with a strong economy and a
dynamic, almost American, way of life standing in marked contrast to the old
centres of Europe.
Among the European capitals Berlin is a late-comer; whereas in the course
of history places like London or Paris became political centres as early as the
Middle Ages, there was never a single focal point in Central Europe, but sever?
al. Although situated within the Holy Roman Empire, Berlin lay at its fringe,
in the northeastern corner at a distance of only some dozen kilometers from
continuous Slavonic settlement area. Here many developments started late -
the oldest written record of Berlin goes back to the year 1237, it took more than
two and a half centuries until the small merchant town on the Spree River
became the place of residence of the Electors ofBrandenburg. Over the centu?
ries territory after territory was added to the possessions of the Electors, later
to become also Kings in Prussia - most of these territories lying east of Berlin.
The political expansion of the Kingdom of Prussia in the nineteenth century
finally made the late-comer one of the main political, economic and cultural
centres of Central Europe, but still there were several others of major impor?
tance like Vienna, Hamburg and Munich, and Berlin only became capital of
the newly founded Deutsches Reich as late as 1871.
From then onwards the city was intimately connected with the fate of this
political structure and experienced all stages of rise and fall leading to almost
complete destruction, division and agony followed by a period of slow recon-
struction and recovery under most unusual, in some respects bizarre, condi-
tions. After the Nazis had brought endless harm, pain, misery and wrong from
their headquarters in Berlin over all of Europe and after this city had in turn
been bombed, set on fire and ruined, it astonishingly lives and - although
254 Lack: Opera magna
divided - continues to play a role in Europe, no longer as a political and

economic, but still as a cultural metropolis.
This historical background is reflected by the inhabitants of Berlin, "an
audacious breed of men" as Goethe called them. They came as immigrants
from all parts of Central Europe, in particular from the territories of the Kings
of Prussia, often ofticially welcome for economic reasons - the underpopu-
lated, under-developed wilderness should become prosperous. This immigra-
tion mentality is apparent in many fields - there are few family dynasties with
an unbroken tradition over centuries, few top positions were and are filled with
people born in Berlin. Even more important is a second aspect: the collapse of
the Deutsches Reich and its capital broke human ties to an extent unknown
anywhere else in Europe - literally hundreds of thousands fled Berlin,
hundreds of thousands arrived and started a new development. There is little
continuity, tradition plays a comparatively subordinate role: in short an almost
American way of life predominates.
Again starting late, Berlin became one of Europe's major economic cen?
tres; although relying almost completely on imported goods and raw materials
it became - as centripetal political forces grew stronger in the late nineteenth
century - the main industrial city between Paris and Moscow. The spectacular
concentration of funds and manpower resulted in very considerable spending
of money on science and technology unparalleled in other parts of Central
Europe. But the time of glory was relatively short - ending in catastrophe,
bombs and the deportation of whole factories. Chicago on the Spree stood in
ruins. Although rebuilt to a considerable degree after the Second World War,
the economy of Berlin has lost momentum, the pace-makers and trend-setters
of economic development have moved elsewhere.
Sciences in Berlin
These more general aspects have and had a profound influence on the
sciences in Berlin. In this field, too, major developments started late: more
than 450 years had passed since the universities of Prague, Cracow and Vienna
had been founded, when in 1810 the Friedrich Wilhelm University opened in
Berlin. The Society of Sciences, later to be called the Royal Academy of
Sciences came into existence more than a century after the foundation of the
Accademia dei Lincei in Rome, half a century after the foundation of the Royal
Society in Oxford and the Imperial Academy of Natural Sciences in Schwein-
furt. In this respect, too, further development was rapid indeed, making Berlin
Englera 7-1987 255
quickly one of the foremost places for sciences in the world: as early as 1910
more than 9,000 students were enrolled at the Friedrich Wilhelm University,
about 30 Nobel prizes were attributed to scientists working in Berlin. In the
thirties Max Planck, Albert Einstein, Otto Hahn and Lise Meitner -just
to give an example from the field of physics - worked simultanously in Berlin.
It is, however, characteristic, that all four were immigrants, having been born
in Kiel, Ulm, Frankfurt am Main and Vienna respectively, all four spending
their retirement and old age outside Berlin. Very diverse reasons had brought
them to Berlin, but one aspect is evident: the extraordinary facilities for scien?
tific work available in an economically strong metropolis were attractive, simi?
lar conditions could only be found with difficulty elsewhere in Europe.
A city like Berlin, however, did not only attract people, it also attracted
objects - mummies, paintings, coins, books, plant specimens; over the centu-
ries this city became one of the European centres of museums and libraries
with famous institutions like the Royal Library Unter den Linden and the
agglomeration of art museums on the Spree Island in the very heart of the city.
Again several peculiarities should be noted: compared to museums and librar?
ies in Paris or London, the institutions in Berlin were late-comers, which
reached their climax after the nse of the Deutsches Reich. They have never
become the focal points in Central Europe - there have always been several -
the German Library in Leipzig, the Imperial collections in Vienna, the Royal
galleries in Dresden and Munich have not only been strong competitors, but
have sometimes surpassed the museums and libraries in Berlin. The aston-
lshing, breath-taking growth of these institutions in the first half century of the
Deutsches Reich was backed by substantial funds coming directly or indirectly
from a strong economy, but were followed by an equally astonishing, breath-
taking collapse and a fragmentation of the collections in the following years.
Fortunately much less was actually destroyed than is usually assumed: but the
colossus is now split into two halves resulting in the somewhat strange and
anomalous situation of most scientific institutions now existing in duplicate -
starting with the old Friedrich Wilhelms University split now into the Hum?
boldt University in Berlin (East) and the Free University in Berlin (West) and
ranging to the Egyptian Museum split into two collections, one in the Bode
Museum in Berlin (East) and one in Berlin (West). The natural history collec?
tions form a rare exception to this general rule: the very extensive zoological,
paleontological, mineralogical and geological material is conserved undivided
at the Museum for Natural Sciences in Berlin (East) whereas the botanical
material is kept almost undivided in Berlin (West).
Another striking fact is the apparent lack of a main field. Whereas the old
political centres had universal collections, but usually with a strong emphasis
on traditional spheres of influence in and outside Europe, areas later to
become their colonies, Berlin with its peculiar circumstances and the late
acquisition of the few, comparatively small Protectorates of the Deutsches
Reich did not possess such specialities. Consequently, many collections here
are characterized by a remarkable encyclopedic breadth both in time and
space.
A Mecca for botanists
In contrast to many other fields of science, botany had a relatively early

start in Berlin. When the Royal B otanic Garden in Schoneberg near B erlin was
founded in 1679 few similar institutions existed north of the Alps, but develop?
ment was rather slow. At a time when the Jardin du Roi in Paris and the Hortus
Medicus in Amsterdam flourished, the Royal Botanic Garden in Schoneberg
was in a poor state having been destroyed in the Seven Years' War by Russian
and allied troops. Some years later the situation of the garden was still so bad
that the proposal was made by the Royal Academy of Sciences to sell the land
and give up the institution altogether. This suggestion was not followed, but
when Carl Ludwig Willdenow was appointed Director of the Royal Botanic
Garden in 1801 he took over "a desert scarcely meriting the name of garden".
And when Alexander v. Humboldt returned from his great voyage to the
Americas no herbarium of any size existed in Berlin besides Willdenow's
small, private collection. The facilities for botanic research were then so much
more attractive in Paris that Humboldt decided to have his rich material from
South and Central America studied there, the results being also published in
the French capital.
With the foundation of the Friedrich Wilhelm University in 1810, the inte-
gration of the Royal Botanic Garden into this new centre of science and the
foundation of the Royal Herbarium later to become the Royal Botanical
Museum, an era of rapid development started. "It seems that every botanist
will have to go on a pilgrimage to Berlin - like the Mussulman to the tomb of
the prophet - if he wishes to die botanically happy", was the comment of a
distinguished visitor, Josef August Schultes from Bavaria, as early as 1822.
The next 120 years were a time of steady growth for the Royal Botanic Gar?
den in Schoneberg, later in Dahlem (both suburbs ultimately becoming parts
of Gross-Berlin) and made the institution the centre for plant taxonomy in
Englera 7-1987 257
Central Europe. A constant flow of living and dried specimens as well as

botanical literature from all over the world arnved in Berlin, the number of
staff members steadily increased, the scientific output reached several
hundred pnnted pages per year.
Again several peculiarities merit attention: in contrast to the situation in
London, St. Petersburg or Vienna, where two or even three major botanical
institutions existed or exist in nvalry side by side, there had always been only
one place for plant taxonomy in Berlin: the Royal Botanic Garden, later to
become the Botanical Garden and Museum, all collections, all working capaci-
ties were concentrated there, even pnvate scholars used it as a base and depos-
lted their collections there. In contrast to the examples mentioned above, the
institution in Berlin always formed part of the University and over the years
was led by a succession of scientists, who were simultanously professors of
botany and directors of the Royal Botanic Garden, thus having to fulfill com-
mitments in research, academic teaching and the administration of an ever-
growing scientific collection. Although this meant an extremely heavy burden,
the triple construction gave extraordinary results: directors like Adolf Engler
and Ludwig Diels not only contributed very substantially to science by
publishing a long series of original papers, but also successfully managed a
major research institute and taught systematic botany to the students of the
Friedrich Wilhelm University. It is interesting to note that all professors of
botany in Berlin until 1945 - with the only exception of Willdenow - were
immigrants, who had not been born in Berlin; but the facilities for botanical
research in this city were so attractive that none of them gave up his position to
work at another university.
The growing political, economic and cultural strength of Berlin is well
reflected by the transfer of the Royal Botanic Garden to Dahlem: truly royal
in size it now covered 42 hectares of ground and thus had become by far the
biggest botanical garden in Central Europe. Glasshouses of a colossal size
were constructed - then only surpassed by the Royal Botanic Gardens at Kew
and the Imperial Botanical Garden in St. Petersburg - as well as the biggest
botanical museum then in existence: the huge red-brick building on the Koni-
gin-Luise-Strasse housed about four million herbarium specimens, numerous
collections of seeds, fruits, wood specimens, fibres from all over the world and
a rich botanical library covering four centuries. When the Royal Botanic Gar?
den and Museum in Dahlem was officially opened in 1910 the young capital of
the Deutsches Reich was enriched by a botanical centre surpassing most
equivalent institutions in the old capitals of Europe.
Further growth slowed down somewhat in the twenties and thirties, but it
was the inferno of the Second World War which almost brought an end to the
Botanical Garden and Museum in Berlin-Dahlem. In 1943 the peaceful Bo?
tanical Museum was set on fire by phosphorus bombs resulting in an almost
complete destruction of one of the finest botanical collections and libraries of
the world. Later the glasshouses were ruined and a few days before the uncon-
ditional surrender of the Deutsche Wehrmacht, the Botanical Garden itself
turned into a battlefield: tanks made their way across the flower beds, trenches
were dug, dozens of German and Soviet soldiers died in action.
The prospects for the future were dim; potatoes and other vegetables were
grown for years in the Botanical Garden, it took a long time until the few
surviving collections were returned from their evacuation sites. Although the
Botanical Garden and Museum were put by the American Military Govern-
ment under the administration of the Magistrat of Berlin as early as 1946 thus
ending 136 years of association with the Friedrich Wilhelm University, the
reconstruction and consolidation process was very slow: it took about a quarter
of a century until the big tropical conservatory was rebuilt and nearly half a
century will have passed until reconstruction work on the Botanical Museum
will be finished. The splitting of Berlin, the blockade of the access routes to the
western sectors of Berlin, the workers' revolt in 1953 in the eastern sector of
Berlin brought further uncertainty, the Berlin Wall finally fixed in concrete a
most abnormal political situation. In these difficult years botany could not
flourish; within ten years of the end of the Second World War a whole gener-
ation of botanists from the Engler era had died, emigration continued, and it
was quite difficult to find suitable replacements. The sixties and seventies saw
political stabilization and the foundation of two new, though small centres for
plant taxonomy in Berlin: at the Free University an Institute for Systematic
Botany and Plant Geography was founded, while at the Humboldt University a
similar institute and a modest botanical garden were created. Again fully inte-
grated into the network of international cooperation with a significant share in
several major research projects, Berlin - and in particular the Botanical
Garden and Botanical Museum Berlin-Dahlem - has become a Mecca for
botanists again, although now within the context of a somewhat unusual
general situation.
Englera 7-1987 259
Opera magna
Over the centuries a considerable number of most prestigious journals in

the field of plant taxonomy has been founded and edited in Berlin - ranging
from "Linnaea" (founded by Diederich Franz v. Schlechtendal in 1826, now
discontinued) and "Botanische Jahrbiicher fiir Systematik, Pflanzenge-
schichte und Pflanzengeographie" (founded by Engler in 1881, then still in
Kiel) to "Repertorium specierum novarum regni vegetabilis" (founded by
Friedrich Fedde in 1906) and "Nova Hedwigia" (founded by Johannes
Gerloff and Fritz Mattick in 1959). For many years these periodicals were
connected with the Royal Botanic Garden, later Botanical Garden and
Museum, since the editors were working there. Currently the last three jour?
nals are still edited or co-edited by members of staff of the Botanical Garden
and Botanical Museum Berlin-Dahlem and of the botanical institutes of the
Humboldt University and the Free University. Although these journals make
Berlin the centre for editorial work in plant taxonomy in Central Europe, they
will not be dealt with here.
From the point of view of scientific development the original research
papers produced in Berlin have to be regarded as more important; going into
thousands and occupying a considerable number of meters in library shelves,
the topics range from Willdenow's studies on the genus Amaranthus to
D iels' s contributions to the plant geography of Western Australia, from Hein?
rich Friedrich Link's pioneer work on fungi to current studies on the flora of
the South Aegean Islands. There is no leitmotiv, but a constant search for a
better understanding of plant life - without any main field or special emphasis
on a particular group or geographical region. No attempt is made here to
summarize this extensive literature totalling tens of thousands of printed
pages and scattered over many periodicals, serials and independent publi?
cations.
Whereas it is possible - although often only with great difficulty - to write
original research papers in plant taxonomy at more modest institutes with
their reduced facilities, major works, like Floras covering a wide area, multi-
volume conspectuses covering greater parts of the plant kingdom, can only be
produced at major institutes. In this way, e. g. the "Flora of Tropical Africa"
and its successors the "Flora of West Tropical Africa" and the "Flora of Tropi?
cal East Africa" were and are produced at the Royal Botanic Gardens at Kew,
just in the same way as the "Flora of the Soviet Union" is intimately linked with
the Komarov Botanical Institute in Leningrad and the new " FloraNeotropica"
with the New York Botanical Garden. Outside such institutions with their
combination of scientific collections, library holdings and manpower these
opera magna are impossible to achieve. Such mammoth works provide most
helpful syntheses, containing a wealth of new information and a critical evalua-
tion of the literature, thus becoming indispensable reference tools, often for
generations of plant taxonomists.
Berlin and the botanical collections eonserved there became the basis for
several opera magna - Engler's "Pflanzenfamilien" probably being the most
widely known and, even today, 72 years after its completion, forming a corner-
stone in plant taxonomy. Out of a series of possible candidates six works have
been selected here.
Willdenow's "Species plantarum"
Like a trumpet call the series of opera magna of Berlin plant taxonomy
starts with Willdenow's "Species plantarum", an astonishing one-man work
totalling more than seven thousand printed pages with an ambitious goal: to
provide a synopsis of all the genera and species of plants known to the author
and to give brief descriptions, notes and references to the literature.
In 1753 Linnaeus had published a synopsis of all the genera and species of
plants known to him, the famous "Species plantarum", totalling 1,200 printed
pages; after a slightly extended second edition prepared by Linnaeus and a
reissue of the latter, usually regarded as third edition, Willdenow produced
the fourth edition of "Species plantarum": but this is a completely mdepend-
ent work to be attributed to Willdenow only, with more than six times as
many pages and starting to appear in 1797. Its approach is stnetly Linnaean, the
complete adtierence to the sexual system, the arrangement of genera, even the
typography follows the famous model. There are many more notable similan-
ties: both authors were relatively young when their "Species plantarum" start?
ed to appear in press: Linnaeus had reached the age of 46, had been elected
member of the Royal Academy of Sciences in Stockholm and appointed Pro?
fessor at the University of Uppsala, Willdenow had reached the age of 32, had
been elected member of the Royal Academy of Sciences in Berlin and was
appointed Professor at the CoUegium medico-chirurgicum a few months later.
Linnaeus had already become Director of the Botanical Garden of the Univer?
sity of Uppsala, Willdenow was appointed Director of the Royal Botanic
Garden of the Royal Academy of Sciences four years after the first part of the
Englera7- 1987 261
"Species plantarum,, was published. Both, Linnaeus and Willdenow, had to

start from scratch - at the beginning of their careers in Uppsala and Berlin
research facilities for plant taxonomy were poor: no herbarium, no botanical
Ubrary, no collections of living plants meriting attention were available. So
both had to build up their own private herbarium and Ubrary; with the help of a
network of correspondents, both brought together collections of living and
Carl Ludwig WiUdenow, Director of the Royal Botanic Garden in Schoneberg

and first Professor of Botany at the Friedrich Wilhelm University. Copper
engraving. - Deutsche Staatsbibliothek, Berlin.
dried plants from the remotest parts of the world then known without having
ever travelled outside Europe themselves. In this respect Willdenow's "Spe?
cies plantarum" differs from all other opera magna: when started, no founda-
tions for phytotaxonomic work existed in Berlin, Willdenow laid them and
the acquisition of his herbarium by the King of Prussia is usually regarded as
the starting point for the Royal Herbarium in Berlin. In providing specimens
from overseas Willdenow's friend Alexander v. Humboldt was most prolif-
ic, sending material of about 3,600 species from the South and Central Ameri?
can flora, part of them described by Willdenow in the "Species plantarum"
as new to science. Collections by Hipolito Ruiz and his collaborators from the
Real Expedicion Botanica to the vicekingdom of Peru, by Friedrich August
Freiherr von Marschall von Bieberstein from the Russian Empire and by
Johann Reinhold and Johann Georg Forster from Captain Cook's second cir-
cumnavigation and by many others were equally important. Consequently
Willdenow's "Species plantarum" is based on herbarium specimens, some?
times on living material coming mainly from the Royal Botanic Garden, but
often on references to the literature only. It should be noted that coverage was
not complete and critics published long lists of omissions, but this is hardly
surprising: Willdenow, like Linnaeus, worked with remarkable speed under
conditions far from ideal and could only dream of the working facilities
available at the time in the library of Sir Joseph Banks in London or in the
Jardin des Plantes in Paris. Although even outdated in systematic approach at
the time of publication Willdenow's "Species plantarum" has to be regarded
as the last almost completed major conspectus of the plant kingdom in
Linnaean tradition.
Of the twenty-four Linnaean orders Willdenow treated twenty-three and
published the first part - the ferns - of the Cryptogamia; less than two years
after his appointment as first Professor of Botany at the newly founded Fried?
rich Wilhelm University Willdenow died of cancer at the early age of forty-
seven. His successor as Director of the Royal Botanic Garden, Link, born in
Hildesheim, then immediate bishopric, and the only head of this institute later
to be decorated with the order Pour le merite, provided the account for various
fungal groups, but all the other cryptogams were left untreated.
"Flora brasiliensis"
When August Wilhelm Eichler born in Neukirchen m Kurhessen was
appointed in 1878 Professor of Botany at the Friedrich Wilhelms University
and Director of the Royal Botanic Garden in Schoneberg and moved from Kiel
Englera7- 1987 263
'?'^k'^î^m^Mî.M'f
^fjkk^*'
3HM!
C. L. Willdenow, "Species plantaûm,^ Vol. 1, Berlin, 1797. Title page. - Bota?

nischer Garten und Botanisches Museum Berlin-Dahlem, Library.
to Berlin he took with him a major project: the "Flora brasiliensis", a multi-
volume work of outstanding scientific importance. Judging from the number
of species treated, it is the biggest flora ever written, comprising when finished
no less than 20,733 pages (actually columns, two on each large page) and 3,811
full page illustrations. The project has an unusual background which is briefly
summarized here: this seems necessary since the "Flora brasiliensis" is usually
associated only with Munich and its first editor Carl Friedrich Philipp v.
Martius; however, careful analysis shows that more pages were edited in Berlin
than in Munich and that many more contributions were written in the former
city.
In 1817 a solemn wedding took place in Vienna: Leopoldine, archduchess
of Austria, was married in procuration to Dom Pedro, Crown Prince of Portu-
gal, Algarve and Brazil. Her father, Emperor Franz I of Austria, gave orders
that a group of naturalists should accompany the bridal voyage to Rio de
Janeiro and collect plants, animals and minerals for the Botanical and Zoologi?
cal Gardens of the Emperor as well as for his Natural History Cabinets. Follow?
ing suggestions from Ferdinand III, Grand Duke of Tuscany, and Maximilian 1
Joseph, King of Bavana, three additional naturalists joined the party, among
them the botanists Martius and Guiseppe Raddi. Travelling on different routes
and penetrating deep into the virtually unknown interior of Brazil, the
naturalists were extremely successful in assembling a vast amount of material,
which today forms part of the treasures kept in the Natural History Museums
in Vienna, Munich, Brussels and Florence, with duplicates widely distnbuted.
The scientific study and evaluation of the botanical specimens were not
accomplished, as one might have expected, in Vienna or Florence. Johann
Emanuel Pohl had returned home from Brazil as a sick man and after his early
death the "Plantarum Brasiliae icones et descriptiones" published in Vienna
remained a torso. Raddi's "Plantarum Brasiliensium nova genera" published
in Florence was left equally unfinished when its author died on the homeward
journey from a tour to Egypt. It was Martius who became the botanical hero of
the whole expedition and made Munich for same decades a centre for the
study of the Brazilian flora. Enobled on the day of his arrival in Munich, later
elected member of the Royal Academy of Sciences in Munich and appointed
Professor of Botany at the Ludwig Maximilian University and Director of the
Royal Botanic Garden in that city, Martius spent his whole life analyzing and
describing the plant world of the Empire of Brazil. After having published
several extensive precursor studies - notably the "Nova genera et species plan-
Englera 7- 1987 265
tarum", "Historia naturalis palmarum" and "Herbarium florae brasiliensis" -

Martius started the Flora brasiliensis project in 1840. From the beginning it
was an international endeavour standing under the protection of three
sovereigns mentioned on the title-pages of all 130 instalments: Ferdinand I,
Emperor of Austria, a son of Franz I, Ludwig I, King of Bavaria, a son of Maxi?
milian I Joseph, and Pedro II, Emperor of Brazil, a son of Pedro I. Martius
acted as editor, contributions coming from the kingdoms of Prussia, Bavaria,
Saxony, Belgium and Great Britain, from Switzerland, to name just the most
important places. Although placed under the auspices of the "Imperial and
Court Museum in Vienna" and supported financially from several sides, the
bulk of the editorial work was done in Munich. Martius himself contributed
only about 2 percent of the accounts to the whole work, but edited the first 46
instalments, the collections in Munich and Vienna as well as in Berlin, Kew,
London, Paris and St. Petersburg forming the basis for this truly opus
magnum.
After the death of Martius in 1868, it was only natural that his private
assistant and later curator at the Royal Botanic Garden in Munich Eichler,
who since 1861 had helped with the preparation of the manuscripts, became his
successor as editor of the "Flora brasiliensis". Describing his involvement with
the project as "a big and heavy, but also beautiful task" Eichler wrote about 4
percent of the accounts, but edited the second third of the instalments, starting
from part 47 and ending with part 99. Of these 53 fascicles only six were pub?
lished during Eichler's stay in Munich, nine during his appointment as
Professor of Botany at the k. k. Technical University and Director of the
Botanical Garden and Herbarium of the Johanneum in Graz, followed by
fourteen fascicles edited during his years as Professor of Botany at the
Christian Albrecht University and Director of the Botanical Garden in Kiel.
Under very favorable circumstances, subsidized by the government of the
Brazilian Empire with no less than 10,000 Mark per year, the Flora brasiliensis
project finally moved to Berlin; starting with fascicle 77 all subsequent parts
were edited here. Eichler's heavy commitments in Berlm, m particular the
reorganization of the Royal Botanic Museum, which was opened during his
time in office, as well as several rearrangements in the Royal Botanic Garden,
led to a sharp decrease in his own research activities on the flora of Brazil. It
was therefore understandable that Eichler looked out for an editorial assist?
ant, whom he found in Ignaz Urban, first assistant at this institute, who had
already published two treatments when still a high-school teacher in Lichter-
felde near Berlin (today Berlin-Lichterfelde).
.**.i.?/3.'3.
?
-?'
,.?. ?i'-i-'
C. F. Ph. v. Martius, A. G. Eichler & I. Urban (ed.), "Flora brasiliensis", Part

130, Leipzig, 1906. Title page. - Botanischer Garten und Botanisches Museum
Berlin-Dahlem, Library.
Englera 7- 1987 267
The Royal Botanic Garden and Museum in Schoneberg was a good place
for the project at that time due to the presence of extensive collections from
Brazil gathered by Friedrich Sellow, Franz Julius Ferdinand Meyen and
Ernst Heinrich Ule as well as extremely rich duplicate series received from all
major museums; excellent facilities for literature work were available at the
Royal Library, then still in the so-called "Kommode" next to Unter den
Linden. When at the early age of 47 Eichler died in 1887 of cancer after along
illness, little more than half of the "Flora brasiliensis" was finished. It was
again only natural that Urban became Eichler's successor as editor. Within
the next nineteen years he managed to bring the project to a happy end, this
being a remarkable acceleration compared to progress made under Martius
and Eichler.
In some respects the situation in Berlin was notably different from the
circumstances in Munich, Graz and Kiel. Whereas Urban contributed like
Martius only little more than 2 percent of the total number of accounts, he was
able to find among his colleagues at the Royal Botanic Garden and Museum
some most active and competent collaborators - notably Karl Schumann,
born in Gorlitz in Prussian Silesia, who contributed no less than 1,407 pages,
thus ranking second in importance after Andre Cogniaux, high-school teacher
in Verviers in Belgium, who besides Melastomataceae had treated the 142
genera and 1,765 species of Orchidaceae in a total of 3,105 pages. Taking into
consideration the accounts written for "Flora brasiliensis" in Berlin - by
Schumann, Urban, Otto Berg, born in Stettm in Prussian Pomerama(now
Szczecm in Poland), Max Giirke, born in Beuthen in Prussian Silesia (now
Bytom in Poland), and others - the Reichshauptstadt clearly ranked as number
one in Central Europe for "Flora brasiliensis". The leading position of Berlin in
neotropical plant taxonomy is further stressed if the accounts written for this
project by people who later moved to Berlin like Eichler and Engler are
added as well as by contnbutors who had graduated from the Friedrich Wil?
helm University, but were living elsewhere when writing their treatments for
this opus magnum.
Thus "Flora brasiliensis", published like Willdenow's "Species planta?
rum" in Latin and written by authors who for the overwhelming part had never
set foot on Brazilian soil, was neither a one-man work, nor a mostly Bavarian,
Austrian or Prussian undertaking; but the share of the Royal Botanic Garden
in Schoneberg was considerable. Although taxonomie opinions have changed
over the last eighty years, so far no modern Flora of the vast area of Brazil has
been produced. The approximately 250 copies of the work, impressive in its
folio size and twice reprinted, therefore remain a lasting monument to

Martius, Eichler and Urban.
Engler's "Pflanzenfamilien"
When Adolf Engler, born in Sagan in Prussian Silesia (now Zagan in

Poland), and appointed Professor of Botany at the Friedrich Wilhelm Univer?
sity and Director of the Royal Botanic Garden in Schoneberg in 1889, moved
from Breslau (now Wrocfaw in Poland) to Berlin he also took with him a major
project: "Die Naturlichen Pflanzenfamilien", like "Flora brasiliensis" a multi-
volume work of outstanding scientific importance. But the situation was differ?
ent in many respects: "Pflanzenfamilien" is not a Flora, but a handbook writ?
ten with the aim of providing a synopsis of the knowledge then existing in the
field of plant taxonomy, thus not being restricted geographically and dealing
with both phanerogams and cryptogams. Therefore the "Pflanzenfamilien" is
synthetic in approach, treats systematic entities to the rank of genus, but does
not consider infrageneric categories in detail. Whereas "Flora brasiliensis" is
written in Latin, the lingua franca of plant taxonomy, the "Pflanzenfamilien"
is published in German, and whereas only less than half of the fascicles of
"Flora brasiliensis" were edited in Berlin, 149 out ofa total of 186 instalments
of the "Pflanzenfamilien" were edited in this city. The latter work with its four
divisions and 32 independently paginated volumes may therefore be regarded
as a genuine opus magnum of Berlin plant taxonomy.
The "Pflanzenfamilien" was edited by two men: Engler, who took over the
responsibility for the phanerogamic part, i. e. divisions 2-4, and Karl Prantl,
who was in charge of the cryptogamic part, i. e. division 1. The first idea of the
"Pflanzenfamilien" may have been discussed in Munich in 1871 by Engler and
Prantl, both of whom had positions at the Royal Botanic Garden of the Ludwig
Maximilian University in that year. Then their roads parted: after having been
Eichler's successor in Munich, Engler also became Eichler's successor in
Kiel and after five years as Professor of Botany and Director of the Botanic Gar?
den of the Friedrich Wilhelm University in Breslau, he finally became Eich?
ler's successor in Berlin. Prantl moved to the Julius Maximilian University in
Wurzburg, then as Professor to the Forestry Academy in Aschaffenburg and
finally became Engler's successor in Breslau. But Prantl's share in the "Pflan?
zenfamilien" was relatively small: he died of tuberculosis at the early age of 44
after having written a few treatments for the phanerogamic part and edited
Englera 7-1987 269
only the first two fascicles of the cryptogamic part. He had no successor in his
position as co-editor of this handbook. Although the "Pflanzenfamilien" is
often simply called "Engler-Prantl" this abbreviation clearly places too little
weight on the leading figure of the project, Engler, who edited by far the
greater part of this opus magnum and wrote the treatments for no fewer than 60
families.
Characteristically it was also Engler who gave an outline in the preface to
the general index of the phanerogamic part, stating the "underlying tendencies
of the whole work", namely "to describe all more important morphological and
anatomical conditions of the single families with reference to their biological
peculiarities, giving special attention to the relationship between living and
extinct forms". Furthermore it was Engler's "very special wish, that as far as
possible a natural division of the families based on relationships should be
aimed at, and that for this purpose peculiarities should be taken into consider-
ation not only in flower morphology but also in anatomy. A division of the
genera into subgenera, sections and groups on a basis as natural as possible
and taking geographical distribution into account was appreciated." This
sounds very much like Engler's credo, the theoretical basis of his new syn-
thetic approach to plant taxonomy.
Of course the "Pflanzenfamilien" had precursors, notably the "Genera
Plantarum" by George Bentham and Joseph Dalton Hooker, published in
London in the years 1862 to 1883, and the "Histoire des plantes" by Henri Bail-
lon, published in Paris in the years 1866 to 1895. But Engler's concept was
much broader, taking into consideration information from many more fields.
Most importantly, the "Pflanzenfamilien" was not a one-man work or the work
of two authors, but the result of teamwork. The first two fascicles may act as
an example: the accounts of various monocotyledons were written by Franz
Buchenau, high-school teacher in Bremen, Oscar Drude, Professor of Botany
at the Polytechnikum and Director of the Royal Botanic Garden in Dresden,
Engler, then still in Breslau, and Ferdinand Pax, one of Engler's assistantsin
Breslau and later Prantl's successor in that town.
No fewer than a total of 57 contributors was involved in the phanerogamic
part. The result is remarkably homogenous, due to Engler's strict editorial
policy and the fact that about one sixth of the accounts was written by himself,
whereas the greater part of the remaining proportions was contributed by his
colleagues and students in Breslau and Berlin. Foreign contributors included,
among others, Richard Ritter von Wettstein, who wrote several treatments
when Privatdozent at the Rudolph University in Vienna and later as Professor
S25BS:
,;Lr:i? "lV?*J,*ft
FLAIltZiuiNFAialliiililN
31, ??
imiir MlfetfktaiittmtffrfifrfrnyliirrariiiiiiMr fMiiAlifftffiif
.&*W0&r: ^;***^ te ^Mtaflft
-?3fc
*-V?
;&:---^;^Vi^
??- ? , **>-^"*?- >vf ^ *'?
^ Vi f fe k> Jf* * *V
m>^
?Wm
A. Engler (ed.), "Die Naturlichen Pflanzenfamilien,,, Gesamtregister zum II.

bis IV. Teil, Leipzig, 1899, Title page. - Botanischer Garten und Botanisches
Museum Berlin-Dahlem, Library.
Englera 7- 1987 271
of Botany and curator of the botanical garden of the Karl University m Prague,
as well as Robert Chodat, Professor for Systematic and Pharmaceutical Botany
at the University of Geneva.
Thus the "Pflanzenfamilien" was an international project like "Flora brasi?
liensis", but the Berlin part was much bigger in size and importance. In the
same way as the "Genera plantarum" were based, as the title reads, "ad exem-
plaria imprimis in herbariis kewensibus servata" (primarily on the herbaria
kept at Kew) and the "Histoire des plantes" on material conserved in the
Museum National d'Histoire Naturelle in Paris, the "Pflanzenfamilien" had
the living and dried collections as well as the library of the Royal Botanic Gar?
den and Museum in Schoneberg as its basis.
The production speed of the "Pflanzenfamilien" was breath-taking; it took
less than 12 years to complete the phanerogamic part, i. e. 24 volumes with a
total of 6,608 pages; the completion of the cryptogamic part, i. e. 8 volumes
with a total of 4,978 pages, however, was not reached until 1909.
When the phanerogamic part was finished in 1899, Engler was a man of
only 55 years, holder of the most influential position in plant taxonomy in Cen?
tral Europe, a member of the Royal Academy of Sciences in Berlin, still very
active and characterized best by his two categories of priority: urgent - tomor-
row morning, very urgent - should have been done yesterday. Not surprisingly
four independently paginated supplements were produced between 1897 and
1915, and in 1924, at the age of 80, Engler started to edit a second extended
edition of the "Pflanzenfamilien", incorporating new data but otherwise
following the scheme of the first edition.
Even more astonishingly, Engler was able to write the accounts for no
fewer than 22 families of flowering plants before he died in 1930 at the age of
86, when 11 volumes of the second edition had been published. Six of these
dealt with cryptogams comprising among others treatments of parts of the
lichens by Alexander Zahlbruckner, Director of the Department of Botany at
the Natural History Museum in Vienna, and of parts of the mosses by Viktor
Ferdinand Brotherus, high-school teacher in Helsinki; the remaining five
volumes contained accounts on various phanerogamic groups, the greater part
of which was written by Engler's team in Dahlem. Volume 14a of the second
edition of the "Pflanzenfamilien" may be regarded as his swan song: in 167
pages Engler gives a "Short commentary on the situation of flowers and repro?
duction in angiosperms" and "Principles of the systematic arrangement of
plant families with special reference to the angiosperms".
Hermann Harms, scientific officer in the service of the Prussian Academy

of Sciences in Berlin, became Engler's successor as editor of the second edi?
tion of the "Pflanzenfamilien"; until 1943 he was able to produce nine more
volumes, this clearly indicated a decreasing scientific out-put in plant taxon?
omy starting as early as the thirties. In the first decade of the second edition 15
volumes were published, followed by 8 in the second decade, a single volume
in the third decade, two in the fourth decade, none in the fifth and a single one
in the sixth decade. This last volume appeared in press in 1980 and is the first
part published in English, the new lingua franca in plant taxonomy. Thus a
torso remains, no doubt of outstanding importance as evident by the fact that
the work was reprinted in 1959 to 1960, but it seems very doubtful if this second
edition of the "Pflanzenfamilien" will ever reach completion.
Ascherson & Graebner's "Synopsis der mitteleuropaischen Flora"
To a degree difficult to imagine today, Berlin occupied a central position in

Central Europe in the years around 1900. It was therefore only natural that it
also offered ideal conditions for writing a Flora of Central Europe. Starting in
1896, Paul Ascherson, born in Berlin, then Professor of Botany at the Fried?
rich Wilhelm University, and Paul Graebner, born in Aplerbeck in Prussian
Rhineland, then assistant at the Royal Botanic Garden in Schoneberg, made
use of these facilities and produced the ?Synopsis der mitteleuropaischen
Flora", scheduled for at least 19 independently paginated volumes, but com-
prising vascular plants only.
Like in its precursor, Ludwig Reichenbach's "Flora germanica excursoria",
the area treated is vast including all territories of the Deutsches Reich, the
Austro-Hungarian Monarchy (including Bosnia and Hercegovina), the king-
doms of Poland (then part of the Russian Empire), of the Netherlands and of
Belgium, Switzerland, the Grand Duchy of Luxemburg, the Principality of
Montenegro as well as the French and Italian parts of the Alps. The collections
then kept in the Royal Botanic Museum and the herbaria of the two authors
were so rich, that their treatment could be based primarily on these holdings.
External contributors were comparatively few, among them private scholars
like Wilhelm O. Focke, physician in Bremen, who treated Rubus, and Karl Her?
mann Zahn, Professor at the Staatstechnikum in Karlsruhe, who wrote the
account of Hieracium.
Englera7-1987 273
Compared to other Floras of Europe, notably the new "Flora Europaea"

published in five volumes in the years 1964 to 1980 in Cambridge, Ascherson
& Graebner's "Synopsis" is extremely detailed, rich in annotations and con?
tains descriptions and keys for numerous infraspecific categories; full synony-
mies, ecological notes and commentaries are also provided. Not surprisingly,
the work remained incomplete: when Ascherson died in 1913 only six vol?
umes were finished, when Graebner died two decades later five more vol?
umes had been published; Paul Graebner filius continued work until the out-
break of the Second World War, but several volumes hat not even started to
appear by that time. A second, revised edition, published parallel to the first,
also remained a fragment. Still important as a source for floristic data, the
"Synopsis" may well be regarded today as outdated in several respects, having
been superseded by Hegi's "IUustrierte Flora von Mitteleuropa" and, notably,
by "Flora Europaea".
Urban's "Symbolae antillanae"
Ignaz Urban, born in Warburg in Prussian Westphalia, was of a much

more sedentary nature than Eichler or Engler. He had studied at the Fried?
rich Wilhelm University in Bonn as well as at the Friedrich Wilhelm Univer?
sity in Berlin and been appointed head assistant at the Royal Botanic Garden
and Museum in Schoneberg. Having successively refused offers for the posi?
tion of a Professor of Botany and Director of the Botanic Garden at the Georg
August University in Gottingen, the Friedrich Wilhelm University in Breslau
and the Akademisches Gymnasium in Hamburg, Urban became, together
with Engler, the leading personality of plant taxonomy in Berlin around 1900.
As curator and finally as Assistant Director of the institute he played a vital
role in the transfer of the Royal Botanic Garden and Museum from Schone?
berg to its present site in Dahlem and edited the last part of "Flora brasilien?
sis". His own scientific interests, however, centred on the flora of the West
Indies.
Although Urban never visited this region himself, he published a few
thousand pages on its flora and validated the names of several hundred new
taxa he knew only from herbarium material. Being based at one of the centres
for plant taxonomy in Europe, Urban was able to study the constant stream of
specimens coming into the Royal Botanic Museum from the Caribbean -
collected by Leopold Krug and his associate in Puerto Rico, Domingo Bello y
Espinosa, by Paul Sintenis, who had been hired to gather plant material, by
Heinrich Baron v. Eggers, by Erik Ekman and many others.
Besides his "Sertum antillanum" published in "Feddes Repertorium spe-
cierum novarum" and his "Plantae haitienses et dominguenses" published in
"Arkiv for Botanik", Urban made his findings primarily known in "Symbolae
antillanae", comprising no less than 5,644 pages and thus clearly outnum-
bering all his other contnbutions on the flora of the West Indies. Like Engler,
an extremely hard working man, Urban wrote most of the treatments himself,
thus making "Symbolae antillanae" a true opus magnum of Berlin plant taxon?
omy.
It should be noted, however, that this work is clearly not a Flora, but rather
a miscellany: it comprises various treatments on vascular plants from this part
of the neotropics with a few scattered notes on bryophytes, a most valuable
botanical bibliography of the West Indies, a detailed, annotated list of botani?
cal collectors active in the region, as well as chapters on the plant geography of
individual islands. Volume 4 bears the subtitle "Flora portoricensis" (Flora of
Puerto Rico), volume 8 "Flora dominguensis" (Flora of Haiti) whereas volume
9 is devoted exclusively to Ekman's collections from Cuba.
Twenty collaborators contributed to "Symbolae antillanae", but their share
in the work is rather small compared to Urban's input. As in "Flora brasilien?
sis" Cogniaux wrote the account of Orchidaceae; Otto Eugen Schulz, high-
school teacher in Berlin, studied Solanaceae, Cruciferae and Compositae,
ranking second in importance, followed by several others.
When in 1931, less than three months after Engler, Urban died at the age
of 83, the "Symbolae antillanae" was successfully completed and Berlin had
lost its second grand old man in plant taxonomy.
"Das Pflanzenreich"
The Royal Academy of Sciences in Berlin celebrated in 1900 the bicente-

nary of its foundation by Friedrich III, Elector of Brandenburg. Engler made
successful use of this occasion to launch a new project, to be financed jointly by
this learned society and the Royal Ministry of Culture: "Das Pflanzenreich", a
series of monographs in plant taxonomy aiming at a much more detailed analy?
sis than the "Pflanzenfamilien" had offered. Engler was to be responsible for
the whole undertaking, but found in Harms, born in Berlin and at that
time assistant at the Royal Botanic Garden and Museum in Schoneberg, a
Englera 7- 1987 275
The glory of Berlin plant taxonomy; from left to right: Ignaz Urban, Assistant
Director of the Royal Botanic Garden and Museum in Schoneberg, later in
Dahlem; Ernst Friedrich Gilg; Adolf Engler, Director of the Royal Botanic
Garden and Museum in Schoneberg, later in Dahlem, and Professor of Botany
at the Friedrich Wilhelm University (1889-1921); Paul Graebner; Ludwig
Diels, Director General of the Botanic Garden and Museum in Berlin-Dahlem
and Professor of Botany at the Friedrich Wilhelm University (1921-1945);
Friedrich Fedde; Robert Pilger, Director of the Botanic Garden and Museum
in Berlin-Dahlem (1945-1950); Otto Christian Schmidt; Hermann Harms.
Photograph taken in summer 1924. - Botanischer Garten und Botanisches
Museum Berlin-Dahlem, Portrait collection.
"Redakteur", who shouldered the full editorial responsibilities. From 1901 to

his death Harms was in a way the soul of the "Pflanzenreich", first with
Engler, and after the latter's death with Diels as editor.
It is with some hesitation that the "Pflanzenreich" is included among the
opera magna since its character as a series of revisions is clearly evident. How?
ever, all its 106 volumes were edited in Berlin, the plant material kept in the
Royal Botanic Garden and Museum in Schoneberg, later in Dahlem, acted as
the backbone for the whole work, and a considerable number of accounts was
written by members of staff of this institute. More important still: the format
outlined by Engler in the preface of the first volume was strictly adhered to for
over forty years - keys and descriptions in Latin, full synonymies, notes and
commentaries in German, selected specimens only to be cited. Right from the
beginning, it was clear that such a mammoth undertaking aiming at a complete
deseription of the plant world to specific level not limited geographically and
thus standing in the direct tradition of Willdenow's "Species Plantarum" and
Augustin Pyramus de Candolle's "Prodromus systematis naturalis regni vege-
tabilis" would take very long time to complete successfully. Therefore the
publication followed no sequence fixed in advance, each volume was indepen-
dently paginated and provided with a separate index, thereby in a way
obscuring the general structure underlying the whole work.
Like in the "Pflanzenfamilien" production speed was extraordinary: during
the first decade of the "Pflanzenreich" no fewer than 44 volumes were
published - comprising among the more extensive treatments parts of Araceae
by Engler, Primulaceae by Pax and Reinhard Knuth and Zingiberaceae by
Schumann.
The second decade of the "Pflanzenreich" saw 30 more impressive vol?
umes, but then - like in the second edition of the "Pflanzenfamilien" - a no-
ticeable decrease in scientific productivity occurred: only 22 volumes were
published in the third decade, followed by nine in the fourth and a single in the
fifth decade.
Taking the "Pflanzenreich" as a whole 52 contributors worked together,
most of them being based in Central Europe. It is interesting to note that some
of the most voluminous accounts were produced by private scholars - starting
with Knuth, high-school teacher in Berlin, who wrote treatments of seven
families (including Geraniaceae and Oxalidaceae), followed by Zahn, who
dealt with the giant genus Hieracium, the only part of Compositae published in
the "Pflanzenreich", and Georg Kukenthal, a protestant clergyman in Coburg,
whose studies on Cypereae are still of fundamental importance for the
Englera 7-1987 277
understanding of this group. Members of staffof the Royal Botanic Garden

and Museum in Schoneberg, later in Dahlem contributing to the "Pflanzen?
reich" include besides the three directors Engler, Diels, born in Hamburg,
Free and Hanseatic Town, and Robert Pilger, born in Helgoland, then still
British, and the latter's successor, several curators and assistants. To these a
few names of specialists living in Berlin should be added: Schulz, who had
collaborated with Urban for the "Symbolae antillanae", wrote parts of Cruci?
ferae, Fedde, then high-school teacher in Charlottenburg (today Berlin-Char-
lottenburg), contributed parts of Papaveraceae, Hermann Wolff, veterinarian
in Berlin, parts of Umbelliferae.
Indirectly, however, the Berlin share in the "Pflanzenreich" was even big-
ger: Carl Mez wrote the contributions for this work when at the Vereinigte
Friedrich University Halle-Wittenberg and later as Professor for Plant Physiol?
ogy at the Albertus University in Konigsberg (now Kaliningrad in the Soviet
Union), but had received his doctorate in Berlin after having written a thesis
under the supervision of Urban. Gustav Schellenberg wrote his account
of Connaraceae when publisher of a newspaper in Wiesbaden, but he had
worked as a research assistant at the Royal Botanic Garden and Museum in
Dahlem, to name but a few examples. Even in the time of slowdown the stand?
ards in academic life in Berlin remained very high - volume 104 of the "Pflan?
zenreich", a monograph of Tnuridaceae on a world-wide basis, was a thesis
submitted to obtam the doctorate at the Friedrich Wilhelm University.
Like in the "Pflanzenfamilien" a few specialists from outside of the Deut?
sches Reich contributed to this last opus magnum of Berlin plant taxonomy:
Cogniaux, who had collaborated for "Flora brasiliensis" and "Symbolae antil?
lanae", wrote the account of part of the Cucurbitaceae, Alfred Barton Rendle,
then assistant at the British Museum in London, dealt with Najadaceae, Giin-
ther Beck Ritter von Mannagetta und Lerchenau, Professor of Botany and
curator of the Botanical Garden of the Karl University in Prague, contributed
a monograph of Orobanchaceae, and notably Franz Wimmer. His monograph
of Lobelioideae and Cyphioideae which forms volume 106 of the "Pflanzen?
reich" illustrates in a most striking manner the tragedy of plant taxonomy in
Berlin caused by the Second World War.
Wimmer, a catholic priest and private scholar in Wampersdorf in Lower
Austria, had sent the first part of his manuscript to Harms, who after having
made it ready for the press passed it on to the publisher Wilhelm Engelmann in
Leipzig. When this first part appeared in April 1943, Harms had died and an
event had happened, which immediately became known in botanical circles as
Lack: Opera magna
**V**
&M1
ft '**
3*SSi
"*
-^::**r?; ^vfttKjT'
*? ' V :%--,
is*^-4***-*?<
:*rrr *-\' :'*:'*~-'-*-*ii**z%ii
^H
A. Engler (ed.), "Das Pflanzenreich", Vol. 1, Leipzig, 1900. Title page. - Botani?
scher Garten und Botanisches Museum Berlin-Dahlem, Library.
Englera 7-1987 279
the Dahlem catastrophe: during the night of March 1st to March 2nd 1943 -
exactly one month after the surrender of the last troops of the Wehrmacht in
Stalingrad - the Botanical Museum Berlin-Dahlem was for the most part
destroyed by one of the early air raids on Berlin. The basis of "Pflanzenfami?
lien", "Synopsis", "Symbolae antillanae" and "Pflanzenreich", about four
million herbarium specimens of outstanding scientific value annotated by
generations of plant taxonomists as well as one of the finest botanical libraries
in Europe, was reduced to ashes. "Immediately after the first high explosive
bomb a second fell on the roof of the herbarium wing, broke it open and set the
herbarium on fire. At the same time a number of phosphorus cans fell. Due to
the strong wind, which blew that night, the fire very quickly spread. Soon the
whole herbarium was in flames and the fire passed on to the library... After a
few hours the tragedy had ended: the fire, which shining over a long distance
had produced smoke and heat, died down in a wet and cold night of early
spring; the walls of the burnt-out building stood black against the night sky." -
was an eye-witness's report.
Shortly after the Dahlem catastrophe, the greatest loss which ever hap-
pened to plant taxonomy anywhere on the world, the stock of the first m-
stalment of Wimmer's monograph was also largely destroyed in an air raid on
Leipzig. When the second part of his manuscnpt arnved in Berlin in 1950, the
Deutsches Reich had de facto ceased to exist, Berlin was occupied by the Four
Alhes, the Friedrich Wilhelm University and also the city was split; it took
three years until this second instalment was published by the German Acad?
emy of Sciences, based in the Soviet sector of the city and here acting as the
successor of the Prussian Academy of Sciences. When shortly before his death
in 1961 Wimmer finished a supplement as well as the treatment of Cyphioideae
and the manuscript was finally sent to Berlin, the Wall had already been built
and the future of this city was most uncertain indeed. More than seven years
had passed after Wimmer's death when this last instalment of volume 106 was
finally published; not a single volume of the "Pflanzenreich" has appeared
since. Thus another torso remains.
The material sent on loan to the Natural History Museum in Vienna for
study by Wimmer, however, had survived and was duly returned to the Botani?
cal Museum Berlin-Dahlem. Together with Willdenow's herbarium - found
according to oral tradition in a railway wagon in Saxony years after the
armistice - the bulk of the fern collection and material of several families of
flowering plants this loan represents the only major portion of the general her?
barium left completely intact by the destruction caused by the Second World
War (for further details see Hiepko's paper published in this volume).
Epilogue
After the collapse of the Deutsches Reich, the inferno of the battle of Ber?
lin, silence followed. It was a time when priorities were totally reversed, when
potatoes for food and coal for heating ranked first, not the sciences and botany
in particular. More than nine years passed until "Notizblatt aus dem Botani?
schen Garten und Museum Berlin-Dahlem", founded by Engler in 1899 and
afterwards renamed "Willdenowia", reappeared after the last war-issue had
been published in 1944. Most appropnately it started with volume one: in
many respects plant taxonomy in Berlin started again from scratch.
The silence had lasted very long: in the second city badly hit by the Second
World War and then also jointly occupied by the Four Allies, in Vienna, the
recovery was distinctly quicker: it took only three years until "Annalen des
Naturhistorischen Museums in Wien" reappeared, the extremely rich and
more or less intact holdings of the Natural History Museum and the Austrian
National Library had already been brought back in the first post-war winter.
The return of the scattered remaining herbarium material from the evacuation
sites to Berlin was on the contrary not completed until 1948, and - hard to
believe - more than thirty years passed until the greater part of the botanical
literature formerly kept by the Prussian State Library was brought back to this
city.
The prerequisites for opera magna in plant taxonomy simply no longer ex-
lsted in Berlin. Consequently not a single major scientific project comparable
to e. g. the "Pflanzenfamilien" was started after the end of the Second World
War, nor were the existing works finished. All this is sad, but hardly surprising.
On the other hand, it was possible within the four post-war decades, the first
two of which were at least fairly critical for the future of Berlin, to build up
excellent working facilities for plant taxonomy with a major centre, the Botani-
cal Garden and Botanical Museum Berlin-Dahlem, and two minor centres at
the Free University and the Humboldt University, all three being full mem?
bers of the international scientific community.
After 1945 plant taxonomists in both parts of the city contributed to several
major scientific projects, monographs, revisions and a long list of various
research papers were published. A "Flora of Togo" was finished in 1985, parts
of the revised edition of Hegi's "Flora von Mitteleuropa" were edited here.
Thus in a way an atmosphere of business as usual prevails. The only possible
post-war candidate for opera magna in plant taxonomy is the Med-Checklist
project currently in progress with three research teams in Berlin, Geneva and
Englera7- 1987 281
Montpellier, headed by Werner Greuter, the present First Director of the

Botanical Garden and Botanical Museum Berlin-Dahlem, Herve Burdet and
Gilbert Long, respectively. Two out of a total of eight volumes scheduled have
been published so far, the last in 1986, being as the subtitle reads "A critical
inventory of vascular plants of the circummediterrranean countries".
Willdenow's "Species Plantarum", "Florabrasiliensis", Engler's "Pflanzen?
familien" and Urban's "Symbolae antillanae" represent the glory of plant
taxonomy in Berlin, the torso character of Ascherson & Graebner's
"Synopsis" and "Das Pflanzenreich" stands for its decline and fall. May plant
taxonomy and plant taxonomists in Berlin not experience another catas-
trophe, continue to contribute to the growth of knowledge and be able to work
in peace.
Note
This paper is based on an extensive survey of the relevant historical and bio-
graphical literature summarized in the following two standard bibliographies:
Stafleu, F. A. & Cowan, R. S. 1976-1986: Taxonomic literature, 2nd edition,
1-6. - Regnum Veg. 94, 98,105,110,112,115.
Zepernick, B. & Timler, F. K. 1979: Grundlagen zur 300jahrigen Geschichte
des Berliner Botanischen Gartens. - Englera 1.
Full references will be given in the German edition of this paper.

Priv. Doz. Dr. H. Walter Lack, Botanischer Garten und Botanisches Museum
Berlin-Dahlem, Konigin-Luise-StraBe 6-8, D-1000 Berlin 33.
282 Name index
Name index
Achard, F. C. 6 Bothe 230

Aderhold, R. 57, 69 Bouche, P. 52
Albrecht, M.-L. 44 Brandt 92
Althoff, F. 107-109 Braun, A. 7, 35, 36, 39, 48, 49, 222,
Appel, 0. 51, 57, 59, 60, 64-66, 71, 223
72, 74, 76, 96 Bneger, F. G. 200
Ascherson, P. 92,223,225,272,273, Buch, L. v. 88
281 Bucher, Th. 129
Auerbach, Ch. 200 Buchheim-Matz, R. 47
Barner, J. 63 Buchner, E. 6
Ball, E. G. 129 Burgsdorff, A. L. v. 89
Bauer, H. 185, 201 B utenandt, A. 139,140,198,202,203
Baur, E. 7, 13, 19-22, 25, 26, 155- Chamisso, A. v. 7, 8, 33, 39, 90, 92
159, 161-167, 169-173, 177, 183, 149-151, 199, 221
189, 190, 194, 196, 202 Christian, W. 124, 129
Beckmann, E. 109, 118 Clauss, H. 139
Beger, H. 39, 48, 230 Conwentz, H. 46, 97
Belaf, B. 199 Correns, C. E. 7, 30, 110, 118, 133,
Behtz, H. J. 181 147,157,163,164, 173, 177, 185-
Berg, O. 267 191, 200, 202
Berger-Landefeldt, U. 98 Dahl 48
Bergold, G. 139, 203 Danneel, R. 139, 203
Berliner, A. 14 Davidson, J. N. 129
Berliner, E. 56 Delbruck, M. 206
Bethge, H. 39, 47, 48 Dellweg, H. 211
Bier, A. 14 Diels, L. 14, 30, 91, 224, 225, 229,
Bolsche, W. 37 257, 259, 275-277
Borner, C. 77 Dietnch, F. C. 92, 223
du Bois-Reymond, E. 30, 36 Domke, W. 229
Bornkamm, R. 98 Donat, A. 39, 47, 48
Bortels, H. 74 Drawert, H. 46
Englera 7-1987 283
Dressel, R. 211 Gemeinhardt, K. 39, 48

Ebing, W. 75 Gerhardt 231
Eckardt, Th. 7, 220, 229 Gerlach, W. 63, 71, 72
Ehrenberg, Ch. G. 8, 35, 36, 39, 47, Gerloff, J. 31, 46, 48, 49, 259
48, 232 Gewitz, H.-S. 129
Ehrlich, P. 6 Gilg, E. 7, 275
Eichler, A. W. 7, 222, 223, 262, Gleditsch, J. G. 41, 43, 52, 86, 87,
265-268, 273 89,148-150, 220
Einstein, A. 122, 128, 255 Godan, D. 77
Elssholz, J. S. 220 Golz 46
Engel, M. 105 Goldschmidt, R. 14, 110, 118, 167,
Engelmann, Th. W. 30,128 177, 185, 189, 190, 194-198, 201,
Engler, A. 7, 40, 91, 93, 155, 223, 202
224, 257, 259, 267-278, 280,281 Gothan, W. 6
Eyferth, B. 46 Graebner, P. 30, 92, 93, 225, 272,
Fedde, F. 259, 275, 277 273, 275, 281
Feuerborn, H. J. 94 Graebner, P. filius 273
Fichtner, G. 64 Graevenitz, L. v. 169, 170
Fischer, Emil 6, 109, 118, 122, 124, Greuter, W. 98, 229, 281
128 Gross, R. 233
Fischer, Eugen 207 Grummann, V. 231
Fischer, W. 75 Gurke, M. 267
Fleischer, M. 232 Haas, E. 124, 129
Foyn, B. 45 Haber, F. 109,118,128
Frank, A. B. 43, 54-57, 68, 69, 77, Haberlandt, G. 7,108,137
78 Haeckel, E. 47
Frank, J. 122 Hammerling, J. 7, 45,137-139
French, E. St. 129 Harle, A. 61
Frenzel, J. 44, 96 Hahn, E. 6
Friedrich, B. 211 Hahn, O. 19,109,118, 255
Friedrich der GroBe 52 Hanstein, J. 222
Friednch-Freska, H. 139, 203 Harm, W. 209
Fnsch, J. L. 52 Harms, H. 234, 272, 274-277
Frey, W. 77 Harnack, A. 109,194
Gaffron, H. 129 Hartmann, H. 129
Garcke, A. 223 Hartmann, M. 7, 44, 45, 48, 49, 94,
Geipel, G. 209 110, 131, 133, 134-137, 177, 185,
Geissler, U. 46-48 198,199, 201
284 Name index
Hase, H. 76 Kemmer, E. 6
Heck, L. 165 Kirschstein, W. 231
Hecker 92 Klingmuller, W. 209
Heim, E. L. 87, 88 Kloke, A. 63, 75, 76
Hein, B. 231 Klose, J. 183
Hemroth, K. 202 Klotzsch, J. F. 221, 222, 231
Hemze, K. 63, 77 Knapp 202
Helmbold, W. 208 Knebel 49
Helmcke 45 Kniep, H. 7, 45
Hennig, U. 98 Knolle, P. 209
Hennig, W. 7, 33 Knuth, R. 276
Hennings, P. 39, 231 Koch, C. 233
Hertwig, 0. 155, 171 Koch, R. 6, 33
Hertwig, P. 171, 172 Kohler, E. 67, 68
Hertz, G. 19 Kornicke, F. A. 227
Hiepko, P. 221 Kohlmeyer, J. 231
Hieronymus, G. 39 Kolbe, H.-J. 48, 56
Hillmann, J. 229 Kolkwitz, R. 40, 43, 48, 95
Hiltner 57 Korge, G. 182
H611, K. 39, 47 Kosswig, C. 165
Hoff, J. H. van't 118, 128 Krause, E. 37
Hoffmann, W. 175-179 Krebs, H. 129
Hollrung, M. 53 Kress, H. 182
Holtz, L. 39, 49 Krieger, H. 39, 45, 47, 48
Hopf, M. 204 Krieger, W. 39, 47, 48
Horn, W. 61, 175 Knppahl 126, 129
Horst, P. 165 Krober, H. 72
Houben, J. 75 Krolow, K. D. 178
Hueck, K. 97 Kronacher, C. 165
Humboldt, A. v. 8, 39, 87, 88, 91, Kruger 48
92, 221, 222, 252, 256, 262 Kubowitz, F. 124, 129
Humboldt, W. v. 106 Kuckuck, H. 22, 28, 152, 154, 175,
Husfeld, B. 22 200
Jacobi, A. 66 Kuhn, A. 185, 190, 198, 200-202
Jollos, V. 170, 199, 200 Kukenthal 48
Kappert, H. 172-175, 177 Kunth, C. S. 92, 222
Kaudewitz, F. 209 Kunze, J. 183
Kausche, G. A. 68 Lamprecht, l. 182
Englera 7-1987 285
Lang, A. 139,169, 201 Michaehs 190

Laskowski, W. 182, 209 Milbraed, J. 8, 98, 231
Laue, M. v. 14 Mitscherhch, E. 43
Laux, W. 63 Mix, M. 46, 48
Leh, H.-O. 75 Mobius, K. 47
Lerche, W. 45 Moewus, F. 45,134, 136
Lichtenstein, H. 37 Moritz, J. 77
Lihenfeld, F. A. 200 Morstatt, H. 63, 77
Lindau, G. 39 Muckermann, H. 207
Lindemann, E. 39, 47, 48 Mudra, A. 160
Lindner, P. 6 Muller, H. 63, 75
Lingelsheim, A. v. 47 Muller, J. 36, 47
Link, H. F. 39, 220, 222, 259 Muller, K. 0. 72, 73
Linnert, G. 178 Muller, O. 39, 44, 48
Ludewig, K. 61 Muschler, R. C. 13
Liiers, H. 180-183, 206, 209 Nachtsheim, H. 157, 159, 165, 167,
Liihrmann, R. 183 168, 171, 172, 180, 190, 196, 207,
Luttgens, W. 124,129 208
Luetzelburg, F. v. 14 Negelein, E. 124,129
Magnus, P. 35, 47 Nernst, W. 128
Mangold, O. 185 Neuberg 118
Marcus 183 Niemitz, C. 182
Marggraf, A. S. 6, 43 Nirenberg 71, 72
Markgraf, F. 93 Nitardy, E. 47
Marsson, M. 39, 43, 95 Nothel, H. 181, 182
Marwitz, R. 74 Nurnberg-Kruger, U. 204
Mattfeld, J. 93 Obe, G. 181, 182
Mattick, F. 93, 259 Odenbach, W. 178
Mayer, K. 77 Osterwald, K. 232
Mayr, E. 164 Pag, H. G. 72
McChntock, B. 202 Pankmn, W. 47
Meitner, L. 19,118, 255 Patzig, B. 206
Melchers, G. 139,184,185, 201, 203 Petzold, H. 74
Melchior, H. 229 Pfankuch, E. 7, 68
Merkenschlager, F. 14, 66, 67, 80 Pfeil, F. L. 89
Metzner-Kuster, i. 46 Pilger, R. 39, 225, 228, 229, 233,
Meyen, F. J. F. 8, 32, 39, 48, 53,267 275, 277
Mez, C. 43, 277 Pirschle, K. 201
286 Name index
Pistonus, E. 130 Schlechtendal, D. F. L. v. 221, 259

Planck, M. 128, 255 Schlechter, R. 224
Plarre, W. 178 Schleiden, M. 7
Plate, H.-P. 64 Schloter, W. 165
Plate, L. 165 Schluter, M. 44, 47, 48
Potonie, H. 6 Schmidt, E. 52
Pringsheim, N. 7, 34, 36, 39, 47, 48 Schmidt, G. 77
Quantz, L. 61 Schmidt, 0. C. 31, 49, 275
Raumer, F. v. 38 Schmidt-Ott, F. 109
Rebentisch 92 Schneider, R. 72
Reichenow 48 Schonhard, G. 76
Reid, A. 129 Schoenichen, W. 46
Reimers, H. 232 Scholz, H. 98, 152
Reinkmg 71 Schramm, G. 7, 139, 140, 201, 203
Reuss, M. 211 Schurhoff, P. 7
Richter, H. 62, 63, 71, 72 Schutrumpf, R. 14
Riehm, E. 61 Schuhmann, G. 63, 72
Rohrborn, G. 181 Schultz-Lupitz, A. 54
Rohrborn, H. 181 Schultz-Schultzenstein, C. H. 7
Rong, G. 60, 78 Schultze-Motel, W. 232
Rubner, M. 109 Schulz, O. C. 274, 277
Rudorf, W. 22 Schumann, K. 223, 267, 276, 278
Riimker, K. v. 6, 162 Schuphan, 1. 75
Ruhland, W. 69 Schuster, H. 210
Ruska, E. 45 Schwann, Th. 7
Ruska, H. 68 Schwartz, M. 60, 61
Sacnstan, M. D. 180 Schwartz, V. 201
Sauerbruch, F. 14 Schwarz, O. 14
Sauthoff, W. 63, 72 Schwemfurth, G. 6, 23, 92,151,152,
Schacht, H. 53 162, 234, 235
Schaudinn, F. 6, 45, 131, 136 Schwendener, S. 7, 108, 155
Schellenberg, G. 277 Seiler, J. 201
Schelling, F. W. 91 Sellow, F. 267
Schick, R. 22, 162, 200 Sengbusch, R. v. 22, 162
Schieder, O. 178, 179, 211 Sleumer, H. 230, 233
Schiemann, E. 6,13,17-28,152-154, Sorauer, P. 57, 67, 78-80
159, 164, 169, 170, 172, 204, 234 Spath, F. L. 14
Schiemenz, P. 96 Spath, H. L. 14
Englera 7-1987 287
Spemann, H. 110, 118, 185 Virchow, R. 36, 42, 95

Sperhng, K. 181, 183 Vogel, F. 181, 208
Sprengel, Ch. C. 87, 88 Vogt, 0. 205
Stahl, G. E. 6 Volkens, G. 7
Stahl, U. 211 Wahnschaffe, F. 97
Stange-Bursche, E.-M. 44 Waldenburg92
Stapp, C. 74 Warburg, E. 122
Staudt, G. 204 Warburg, O. 122, 224
Stein, E. 169, 170, 186, 187 Warburg, 0. H. 7, 44, 45, 49, 110,
Stein, W. 209 121-124, 126-130,185
Stengel 168 Wassermann, A. v. 109, 118
Stern, C. 200 Weddigen, U. 47
Sterne, Carus 37 Wehsarg, O. 56
Stoll, A. 112-115, 119-121 Weiss, R. F. 6
Stolp, H. 74 Weniger, J. 165
Straub, J. 201 Werdermann, E. 225, 229
Straus, A. 6 Werth, E. 6, 61, 231
Stubbe, H. 22, 25, 26,147,192, 200, Werz, G. 139
201, 204 Wettstein, F. v. 7, 25, 170, 185-187,
Sukopp,H. 98 190-193, 200, 201
Thaer, A. D. 43, 88, 158 Wichmann, L. 49
Theorell, H. 129 Wilkaitis, V. 47
Thiel, H. 106, 107 Willdenow, C. L. 87-92, 220, 256,
Thomas, F. 234 257, 260-263, 267, 276, 279,281
Tiedemann, H. 129 Willmitzer, L. 210
TimofeefT-Ressovsky, N. W. 171, Willstatter, R. M. 7, 109-115, 117-
205, 206 121, 128
Trautner, Th. A. 210 Windisch, S. 6
Troll, H. J. 163 Wittmack, L. 6, 162, 234
Tschirch, A. 6, 7 Wittmann, H.-G. 210
Tubeuf, K. v. 57 Wohler, F. 7
Ulbrich, E. 93 Wohlgemuth, R. 77
Ule, E. H. 267 Wolf, B.E. 181
Urban, i. 219-221, 223-225, 232, Wolff, H. 277
265-268, 273-275, 277, 281 WoUenweber, H. W. 44, 48, 69-71
Uschdraweit, H. A. 68 Wricke, G. 175
Vennesland, B. 129-131 Wu, Y. C. 225
Verschuer, O. v. 207 Wundsch, H.-H. 44, 96
288 Name index
Zacher, F. 64, 77
Zepernick, B. 235
Zimmer, K. G. 206
Zimmermann, K. 160
Zopf, W. 35

Botany in Berlin

Uploaded by

Document Information

Original Description:

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Botany in Berlin

Uploaded by

Copyright:

Available Formats

Botany in Berlin

Author(s): Hildemar Scholz

on the occasion of the

XIV International Botanical Congress

Veroffentlichungen aus dem Botanischen Garten und Botanischen Museum Berlin-Dahlem

Herausgegeben von der Direktion des Botanischen Gartens und

Redaktion: Hildemar Scholz

? Botanischer Garten und Botanisches Museum Berlin-Dahlem 1987

Alle Rechte, auch die des auszugsweisen Nachdrucks, der photo-

Elisabeth Schiemann - Life and career of a woman

History of phycology in Berlin, D. Mollenhauer .... 29

Contributions to the development of phytomedicine

On the history of plant geography and plant ecology

Botany at the Kaiser Wilhelm Institutes,

A contribution to the history of science of

The collections of the Botanical Museum

Opera magna in Berlin plant taxonomy,

Name index. 283

The present collective volume is published on the occasion of the

The following contributions make no or only marginal mention of

Bacteriology: R. Koch (Nobel Prize 1905) and his school (see

Botanical aspects of forestry, agriculture and horticulture: K. v.

Pharmacognostics: A. Tschirch (see Langhammer & Rimpler

Pharmacy: G. E. S tahl (see Engel 1984, Strube 1984)

The fact that traditional special fields of botanical research in Berlin

Yeast and fermentation research: E. Buchner (Nobel Prize

Additional mention should be made of the "Jahresberichte des

now integrated into the Technical University. Some research results

urea synthesis, which was considered important in the vitalism contro-

On reading the present collection of essays the gentle reader will

Anonymus 1983: Professor Dr. Siegfried Windisch in den Ruhestand

R. Berthold (ed.), Die Humboldt-Universitat. Gestern-Heute -

Heistermann, W. & Heinrich, G. 1980: Beitrage zur Geschichte der

Lippmann, E. O. v. 1929: Geschichte des Zuckers seit den altesten

Vent, W. (ed.) 1980: 100 Jahre Arboretum in Berlin. - Berlin.

EpUogue to the preface

and was considered a cultural obligation of the highest rank: to keep

The idealism of Baur was not an isolated case. R. C. Muschler

rich v. Luetzelburg (obituary by Suessenguth 1955) became head

Baur, E. 1922: Der Untergang der Kulturvolker im Lichte der Biologie. -

? Fischer, E. & Lenz, F. 1921: Grundriss der menschlichen Erblich-

EUsabeth Schiemann was no native Berliner but she received

Elisabeth Schiemann has described, in 1960, her time as a student

It was under the influence of Erwin Baur, then a younginstructor

years. The experimental work had first to be conducted on a private

whether beginning students or eminent scientists in their own right,

Vogelsang. While he continued in office, the research in the institute

of the NS Students' League which existed in each department or school -

Baur, was appointed Director, and Schiemann herself was made

Sciences. In 1946, Elisabeth Schiemann was appointed Professor

As an emeritus Schiemann continued to be quite active. She at-

Address of the author:

History of phycology in Berlin

Like each other part of human culture botany is influenced by the

(Hydrobotanik, vol. 1, 1955) has given an excellent survey of the whole

were supplemented by societies of naturalists or circles of amateurs

filamentous algae (Confervae). A formative power was said to create in

metamorphosis. There must be constant genera and species, otherwise

discuss all the current misinterpretations of his time, many of these

collections were bought by the Hamburg Institut fiir AUgemeine Bo?

The early years of rapid industrial expansion in Germany and their

considerably earlier than elsewhere in Germany thus causing the inven-

Association for the Teaching of Natural Sciences) had been established,