Theoretical Biology in Estonia
Urmas Sutrop, Kalevi Kull
The following is a slightly edited and completed
version of a published small book:
Sutrop U., Kull K. 1985. Theoretical Biology in Estonia. Tallinn: Valgus, 28 p.
What is theoretical biology?
Theoretical science in an ecologically oriented culture
On the history of theoretical biology in Estonia
Philosophical Problems of Biology
Theory of Evolution
Typology, Taxonomy, Theory of Classification
General physiology, Ontogenetics, Genetics
Theory of geobotany and plant productivity
Mathematical methods and modelling
Conferences, associations and research groups
10 Spring Schools of theoretical biology in Estonia 1975-84
If we accept that theoretical thinking is a promotor and a source of ideas for all science, we should allot
theoretical biology a particularly important place in the science off today. The ecological challenge in
the contemporary world has vastly added to its significance.
In Estonia theoretical biology
has been an area of contact for specialists in related fields, and has enabled them to regard their own
specific problems in relation with other spheres of biology. In this country work in theoretical biology
has been conducted mainly by individual investigators. The scientific institutions and research groups
set up for this purpose are a comparatively new development.
The present synopsis is
designed as a brief survey of the studies in the field of theoretical biology carried out by Estonian,
Baltic German, Russian and other local investigators in Estonia. The survey covers the period between
the inception of science and the present time. The work is supplemented by a bibliography of the more
important publications of those investigators (but not more than one or two items per author
What is Theoretical Biology ?
All branches of sciences may be looked
upon as having two facets : the theoretical and the empirical one. Theoretical biology as a whole
comprises almost all theoretical parts of biology. While empirical biology deals with measurements,
observations, descriptions, experiments, facts and collecting of data, theoretical biology is concerned
with models, concepts, rules, laws, principles, calculations, generalizations and conclusions arrived at
both by induction and deduction.
The theoretical facet of biology is as old as the science of
biology, but the term 'theoretical biology' has been in use only since the beginning of this century
(Johannes Reinke,"Einleitung in die theoretische Biologie", Berlin, 1901).
C.H. Waddington, the theory of general biology is the most interesting and important component part
of theoretical biology. The various branches of biology, such as bacteriology,
botany, zoology are
rather loosely related to each other and thus the theoretical studies are also of necessity confined to a
single branch. This is why writing an all-embracing survey of theoretical biology is quite a
Theoretical Science in an Ecologically Oriented Culture
underlying principles of ecological life standards are those of ecological stability. It presupposes
minimization of long-distant transport of materials and maximization of recycling in economy, a good
exchange of information and cultural integrity of small territories.
The main and
principal task of science, from the ecological and cultural points of view, is to collect information
about the mechanisms which quarantee stability of culture and life. Accordingly, local cultures need
specific scientific organizations for the collection of this particular information. The need for such
information is very urgent.
In any branch of science its theoretical part is the main
organizer of its facts. The preservation and use of the information is impossible without theoretical
methods (formulae, models, theories, rules, etc.). Therefore, highly developed theoretical thinking both
in biological and social sciences is a precondition for the existence of a stable culture in a civilized
Proportionally developed science in local cultures implies that there exist sufficient
numbers of specialists necessary for those cultures. In biology there is always a need for experts who
can determine the taxa of organisms inhabiting local ecosystems. Presumably the experts of theoretical
methods in biology are also perpetually needed.
From the ecological point of view, the
principal task of biology is to get to know the mechanisms that keep the biosystems in balance. The
power of self-reproduction which is a fundamental quality of life, gives biosystems a possibility of
unlimited growth. These are the mechanisms of biological equilibrium.
responsible for the investigation of many levels of organization, which renders it a very sophisticated
science. Indeed, the subject being very complex, the theory about that subject cannot be simple either.
Part of the difficulties, but also its attractiveness, arise from borderline position of biology between the
exact sciences and the humanities. Accordingly, the evolvement of the theoretical facets of the science
of life requires more attention and very careful work.
ON THE HISTORY
OF THEORETICAL BIOLOGY IN ESTONIA
In 1691, J.Salbom defended at
Academia Gustavo-Carolina his dissertation which may be considered as predecessor of theoretical
biology in Estonia. The vitalism of J. Salbom had some original traits, regarding life not as a function
of spirit, but as an expression of some fundamentals of life (principii vitalis).
disputation of A.Arvidi on plants (1647) and the dissertation of S.Matthiae on the structure of animal
body (1698) were of a more concrete nature, but worth mentioning as starting points of life science in
Philosophical Problems of Biology
The years 1811-14
witnessed the work of Karl F. Burdach (1776-1847) at Tartu University. He was K.E.Baer's teacher
and later, in Köningsberg (1817-34), his colleague. In his works K.E.Burdach evolved ideas about
progressive development in nature in the spirit of the philosophy of F.Shelling. His course of lectures
"Naturwissenschaft des Menchen" was in fact an introduction to general biology. K.F.Burdach, by the
way, also made use of term 'biology' which he employed to designate the science which included the
anatomy, physiology and psychology of man in 1800. Later, in 1802, G.R.Trevianus and J.B.Lamarck
extended the sphere of use of the term to include the whole of living nature.
important figure among the theoretical biologists in Estonia is undoubtedly Karl Ernst von Baer (1792-
1876). He studied at Tartu University (1810-14) and spent the last decade of his life (1876-67) in
Tartu, having kept in close contact with Estonia all through his life. K.E.von Baer made a great impact
on the development in his life time and long after his death. He has touched upon very many problems
of general biology. One of his greatest contributions to the theory of biology was the evolvement of the
typological approach, relating it to the doctrine of individual development; he was one of the founders
of the doctrine. Baer is also known to have been the first to put forth the antimony of Darwinism and
Gustav Teichmuller (1832-88), professor of philosophy at Tartu University
since 1871, was interested in Baer's type theory, concentrating on the philosophical interpretation and
critical analysis of Darwinism.
Jakob Johann Uexküll (1864-1944), born in Estonia, a
graduate from Tartu University (1884-88), is the author of one of the first monographs entitled
"Theoretical biology " (1920). His philosophical views were influenced by I. Kant and biological ones
by J.Müller, K.E.Baer and H.Driesch. He is considered to be the founder of zoosemiotics and one of
those who paved the way to ethology and comparative physiology of non-vertebrates and the study of
physiological archetypes. Some concepts (the reflex circle, the perceptron) first formulated by
J.Uexküll were rediscovered by biocybernetics several years later.
doctrines both in this country and elsewhere rose mostly from the works of Gustav P.A.Bunge (1844-
1920). He was a graduate from Tartu University and later its professor of physiology. Neovitalism was
a progressive trend in biology of that time. Postneovitalist trends are reflected to some extent in the
works of Nikolai Kuznetsov (1864-1932); he worked as a botanist at Tartu University from 1895 to
1914, J.Uexküll, philosophers Hermann Keyserling (1880-1946) and Nicolai Hartmann (1882-1950; he
studied medicine at Tartu University in 1901-02 and wrote among his numerous philosophical papers a
book on the foundations of biology).
Evald Oldekop (1885-1952), a meteorologist by
speciality, published a very interesting biological and philosophical paper concerning the principle of
hierarchy in nature and its relation to the antinomy of mechanism and vitalism (1930).
Alfred Koort, philosopher (1901-1956), wrote a review about the concept of type (1936,
Over the period following Word War 2 problems of the philosophy of biology have
been discussed at some length by H.Haberman, K.Paaver, M.Makarov, H.Kallak, M.Valt-Remmel,
T.Sutt, T.Loit, a.o. in their publications. Biophilosophical problems have been touched upon by L.Valt,
J.Rebane,V.Tohver, J.Kaplinski, H.Trass, L.Näpinen et al.
Some kind of evolutionary views are noticeable in the works of J.K.Eduard
Eichwald (1795-1875), who supported the idea of alternation of species under the influence of external
factors and the previous taxa. The theme of his venia legendi was "On the Limits of the Animal
Kingdom and Stages in its Development" (1821).
Alexander Keyserling (1815-1891) who
was first and foremost a geologist and a paleontologist had a very wide circle of interests, and he
advocated evolutionist ideas. C.Darwin in his "Origin of Species" mentioned A.Keyserling and
K.E.Baer among his predecessors. A.Keyserling in his item of 1853 put forward a hypothesis about the
evolution of organisms under the influence of special molecules spreading epidemy-like. This may-be
considered as an anticipation of the gene-transfer evolutionary mechanism. According to B.Raikov, the
historian, Christian H.Pander (1794-1865, stud. med. at Tartu University 1812-14), K.F.Burdach and
M.H.Rathke were among the pre-Darwinian evolutionists.
In the years 1863-64, lectures
in Darwinism and plant physiology at Tartu University were delivered by Matthias J. Schleiden (1804-
81), the founder of cell theory.
Carl J. Seidlitz (1798-1885, graduated from Tartu
University in 1820) is an author of several articles popularizing Darwin's works.
Georg C.M.Seidlitz (1840-1917, student of Tartu University 1858-62, son of C.Seidlitz) was
an advocate of Darwinism and a theoretician of international renown. G.Seidlitz delivered a series of
lectures on Darwinism at Tartu University in 1870-77. The lectures, later published a book, were one
of the first of its kind in the world.
Friedrich Berg, a plant-breeder (1845-1938) wrote an
article on the predetermination of the variability of species.
who specialized in paleontology and geology was a Darwinist; his work was on the role of bacteria on
In the years 1898-1902 lectures in zoology at Tartu University were
delivered by Aleksei N.Severtsov (1866-1936), the most celebrated representative of generations of
biologists in the family, founder of the doctrine of phylembryogenesis.
physiologist and histologist (1827-1906), C.E.Mercklin, paleobotanist (1821-1901), F.Schmidt,
paleontologist (1832-1908), A.Rauber , embryologist (1841-1917, in Tartu 1886-1911),
G.P.Mihhailovski, paleontologist (1870-1912, in Tartu 1905-17), P.A.Poljakov, comparative anatomist
(1862-?, in Tartu 1902-1911) and V.A.Afanasjev, patoanatomist (1859-1942, in Tartu 1894-1918)
have touched upon problems of evolution of organisms in their works.
(1892-1932) graduated from Tartu University in 1918 and worked here as a lecturer, his greatest
contribution being the study and popularization of Darwin's theory. The first Estonian professor who
dealt with problems of evolution before WW2 was Johannes Piiper (1882-1937).
Elmar Leppik (1898-1978), graduate from Tartu University (1926) is the author of several
publications concerning problems of general biology, including Vavilov homological series.
As for the authors who have dealt with problems of evolution in their publications in the last
decades, mention should be made of K.Paaver (microevolution of mammals and general problems of
theory of evolution), T.Sutt (directedness of evolution ), M.Valt-Remmel (views of K.E.Baer;
ontogeny and phylogeny), H.Kallak (history, teaching, textbooks), V.Masing (evolution of
ecosystems), M.Viikmaa (evolution of man), K.Põldvere (evolution of tissues), H.Mikelsaar (origin of
life), H.Haberman, A.Heinaru, Ü.kosk, A.Palumaa et al. Raik Mikelsaar has put forth an archigenetic
hypothesis of cellular evolution.
Typology, Taxonomy, Theory of
Johann F. Eschscholtz (1793-1853, at Tartu University 1819-31) published
a work on the general problems of vertebrate classification.
Between 1828 and 1835 the
Chair of Physiology and Pathology was headed by Martin H. Rathke (1793-1860). He made several
important generalizations in comparative anatomy, such as the discovery of gill slits in the foetus of
birds and mammals, disproval of the thesis of comparability of crustaceans and vertebrates put forth by
The rules formulated by K.E.Baer, known as the Baer Laws and
the Cuvier-Baer type theory were a remarkable contribution to biological thinking of their time. The
Baer Laws reflect the regularities of morphogenetic processes, they develop further the
morfotaxonomic system of the type theory.
J.Uexküll was one of the first to distinguish
physiological (functional, behavioural) archetypes besides the morphological ones. The general
problems of typology in connection with biology have also been treated by G.Teichmüller, A. Koort,
M. Remmel, K.Paaver, V.Masing.
At the present time, A.Raitviir and E.Parmasto are
working on the numerical taxonomy of fungi. V.Masing, H.Trass, J.Paal and M.Zobel have obtained
some results in the theory of classification of plant communities.
physiology, Ontogenetics, Genetics
Carl E.H.Schmidt (1822-94) who studied under
J.Liebig and dealt with the comparative physiology of invertebrates under F. Wöhler and R. Wagner in
Göttingen, worked in Tartu from 1846 on. In collaboration with Georg F. Bidder (1810-1894, Rector
of Tartu University 1858-65), he published the results of an extensive research into the nutrition
theory. Among the famous pupils of C.Schmidt in Tartu were also G.Bunge and W.Ostwald, the
winner of Nobel prize.
Carl B.Reichert (1811-83), K.E.Baer's disciple in the field of
embryology wrote a book on histogenetics (1845), in which he refuted the theory of free cell formation
in liquid cytoblasteme and championed the theory of all cells developing from previously existing
In the years 1896-1902 the professorship of histology, embryology and comparative
anatomy at Tartu University was held by Nikolai K.Tschermak (1856-1903). In his book "On the
structure of Living matter" (1895) he evolved his hypothesis of the whirl-like motion of the particles of
living matter. He regarded the open vortexes as self-regulating elements whose reciprocal influences
determine the properties of protoplasm. K.Saint-Hilaire has mentioned him as a forerunner of the
biophysical trend in cytology.
Essential problems of general biology were also treated of
by Edmund F. Russow (1844-97) and Vladeslav Rothert (1863-1916), botanists of Tartu
One of the founders of biorhythmology, Nikolai Pärna (Paerna, 1878-1923),
who was of Estonian descent, but spent most of his life in St.Petersburg, also dealt with some
theoretical problems of physiology.
Alexander Lipschütz (1883-1974) was professor
of physiology from 1919 to 1929 at Tartu University. In his numerous works he treated of many
problems related to general physiology.
Hans Richter (at Tartu University since 1923)
has dealt with the tendency of spiral development and relations between form and function in living
Ants-Peep Silvere (1933-83) developed theoretical principles of micromorphology,
especially the principle of discretess (compartmentation) of life.
of symbiosis and parasitism are investigated by U.Riispere (plants and nematods), A.Turovski (insects,
marine organisms), T.Tiivel (insects and microorganisms), U.Sutrop (plants and bacteria) a.o.
R.Vilu has get some theoretical results in the modelling of procaryotic cell growth.
Theoretical aspects of genetic processes are investigated by T.Soidla, T.Orav, I.Toots,
J.Remme, O.Toompuu, E.Raukas et al, some general problems of embryogenesis by J.Kärner,
Theory of Geobotany and Plant Productivity
geobotanists make up a theoretical school of international renown. Its representatives, Teodor Lippmaa
(1892-1943), August Vaga (1893-1960), L.Laasimer, V.Masing, H.Trass, T.Frey, J.Paal a.o. have to a
smaller or larger extent treated general and theoretical problems of biology in their works. Their
particular results are connected especially with geographical and spatial aspects of plant
In the last decades, much work has been done in the field of theory and
modelling of plant productivity. This trend may be named theoretical school of J.Ross. Particularly, the
theory of radiation distributions in plant communities is progressed by J.Ross, T.Nilson, V.Ross,
A.Koppel. Models of photosynthesis have been developed by A.Laisk, J.Viil, O.Keerberg. Partition of
assimilates and effects of water deficit are examined by H.Moldau. Models of crop production (e.g. for
potato) are worked out by H.Tooming and J.Sepp. Much computational work with dynamical models
of plant productivity are done by Z. and I. Bichele. Spatial distribution of plants in community is
investigated by T.Frey, K.Lõhmus, T.Oja. The model of evergreen tree growth and theoretical
problems of size effects are examined by O. and K. Kull.
One of our first scientists to apply mathematical methods in biology was
Arthur Oettingen (1836-1920, at Tartu University 1863-93), Professor of physics, who progressed the
analytical methods in plant phenology. G.V.Kolossov (1867-1936) was interested in the mathematical
theory of evolution. Statistical methods were applied early if forestry (A.Mathiesen) and anthropology.
K.Regel (in 1920s) dealt with statistical investigation of meadow vegetation. Mathematical problems
of biology were analysed to some extent also by K.R.Kupffer (1872-1935) and A.Kärsna (1907-
In recent decades statistical and other methods for biological data processing have
been applied and evolved by L.Võhandu, S.Veldre, T.Möls, A.Nilson, T.Frey, V.Ross, E.Tiit, J.Paal,
K.Lohmus, A.Raitviir, V.Kask, R.Aps, A.Kiviste, M.Zobel, R.Leht a.o.
So far the
best results in mathematical modelling have been obtained by the team working on the modelling of
plant productivity (see the previous chapter). The scientists who have dealt with mathematical
modelling in other branches of biology include R.Vilu, R.Teinberg, O.Toompuu, J.Männik, E.Raukas,
A.Aruja, J.Engelbrecht, M.Kahru, K.Ross, V.Aladyev
CONFERENCES, ASSOCIATIONS AND RESEARCH
K.E.Baer has made a lasting impact on the development of
biological thought in Estonia. The birthdays of the scientist are marked by scientific sessions (1877-
1926, renewed after 1972). Conferences were held to celebrate his 175th and 180th birthday (1967 and
1972) and the centenary of his death (1976). In 1975 the Estonian Naturalist's Society started
publication of "Folia Baeriana", an aperiodical collection of research papers. In Tartu there has been
opened Baer Museum. Many works devoted to the theoretical views of the scientist have
Several all-Union conferences held here in the past ten years testify to a
growing interest in theoretical biology in Estonia. These include the symposium "Weather, Harvest,
Mathematics" (1975); the 4th Winter School of Theoretical biology and the conference "Biology and
the Linguistics" (1978); it was organized in cooperation with the group of theoretical biology of
Moscow University, directed by A.P.Levich); the series of symposia on the evolutionary biology-
"Theory of Evolution and the 'Man-Nature' problem" (1978), "Micro- and Macroevolution" (1980),
"Methodological problems of Evolutionary Theory" (1984).
Mention should be made of
some other conferences: "Problems of Contemporary Darwinism" (1982); the section "Synergetics and
cooperative phenomena in solids and macromolecules" (1982); the section of theoretical ecology in the
conference "Problems of Contemporary Ecology" (1978); "Session on Theoretical Biology:
Theoretical Systematics" (1976) a.o. Problems of mathematical methods in biology have been
discussed in some sessions (e.g.1966) and publications of the Section of Exact Sciences of the Estonian
The associations of theoretical biologists came into being in the
beginning of the 1960s when the Section of General Biology, Genetics and Selection was formed in the
Estonian Naturalists' Society. In 1967 working team of ecological theoreticians "Ökotoorium" attached
to the Republican Committee for International Biological Program was set up. Both displayed marked
activity in their work until 1972.
In 1970s, a research group of medical cybernetics of the
Computing Centre of the Estonian Ministry of Health dealt with the qualitative modelling of the
development process of biosystems (H.Lippus, J.-T.Tevet).
On February 10, 1974 some of
the then students of biology convened and decided to organize a seminar of theoretical biology. That
was succeeded by the first spring school of theoretical biology in Estonia (1975). It has become an
annual event. The Scientific Society of Students registered a new circle, the group of theoretical
biology. It was the members of that group who in 1977 set up the section of Theoretical Biology
attached to the Estonian Naturalists' Society. The students group of theoretical biology continues its
work at Tartu University.
The group of theoretical biology of the Students' Scientific Society
and the section of theoretical biology attached to the Estonian naturalists' Society are at the moment the
main bearers of theoretical thought in life science in Estonia and the organizers of the spring schools of
The professional work in the field of theoretical biology is done
now in the Department of Evolutionary Problems (K.Paaver, T.Sutt) of the Institute of Zoology and
Botany. The work of many other research groups is also connected with the development of
mathematical models and the investigation of various theoretical problems in biology, e.g. plant
physiology group in the Institute of Astrophysics and Atmospheric Physics (J.Ross, A.Laisk,
H.Moldau, I.Bichele, T.Nilson); group of mathematical modelling (T.Oja), Laboratory of Plant
Ecophysiology (K.Kull) and Department of Bird Ecology (V.Lilleleht, R.Leht, J.Keskpaik) in the
Institute of Zoology and Botany; Department of Biochemistry (R.Vilu) and Department of Molecular
Genetics (M.Saarma, J.Remme, A.Nigul) in the Institute of Chemical Physics and Biological Physics;
Department of Marine Optics (M.Kahru) in the Institute of Thermo- and Electrophysics; Department of
Plant Physiology (O.Keerberg, J.Viil) and Department of Molecular Biology (A.Aruja, E.Raukas) in
the Institute of Experimental Biology; the Estonian Laboratory of Agrometeorology (H.Tooming,
J.Sepp, P.Karing) a.o.
course of lectures in 'theoretical biology' has been delivered by H.Kallak at Tartu University to biology
students specializing in the Department of Genetics and Cytology (previously Genetics and
Darwinism) since 1973. A course in 'biomathematics' has been delivered by T.Möls to biologists since
1969 (in 1977 it was renamed to 'mathematical methods in biology'). In some other courses (e.g.
biophysics by J.Simisker, ecology by T.Frey and M.Zobel) more specific mathematical models of
biosystems are dealt with. Theoretical problems are given much space also in evolutionary biology
(H.Kallak), in some ecological disciplines (H.Trass, V.Masing), in ontogenetics (M.Viikmaa,
10 SPRING SCHOOLS OF THEORETICAL BIOLOGY
IN ESTONIA 1975-84
The first Spring School of Theoretical Biology (STB) was
held at Rutja in May 8-10, 1975. The circle of topics of the school was not restricted. The problems
directedness of evolution (lecturer T.Sutt), theory of polymorfism (T.Paaver), theory of classification
(S.V.Chebanov, Leningrad), formalization of general qualities of biosystems a.o. were discussed.
At the 2nd STB - "Theory of Evolution of Biosystems" (May 7-9, 1976, Rutja) -
were discussed the general characteristics of the mechanism of evolution (M.Viikma et al.), problems
of epigenetic factors in evolution (M.Remmel et al.), statistical models in biology (T.Möls), problems
of evolutionary progress a.o.
The 3rd STB - "Theory of Organism" (May 6-9, 1977, at
the ornithological station of Puhtu) - was dedicated to Jakob Uexküll. His works as well as the
organismic theories in biology in the first half of the 20th century were reviewed; contemporary
problems of modelling of organisms and self-reproduction were also discussed (A.Laisk, R.Leht et al.).
A special discussion of the paper "On the concept of organism" ( by R.Rosen, Canada) took
At the 4th STB - "Theory and Speculations in Molecular Biology " (May 6-9, 1978,
at the biological station of Tipu) - were discussed models of molecular recognition, protein synthesis,
structure and organization of chromosomes, active centres of enzymes, sets of biochemical reactions et
al. A special discussion was held on the ethics of the scientist. At the same time it was the first meeting
of Estonian molecular biologists.
At the 5th STB - "Theory of growth" (May 11-14, 1979,
Simisalu) the main problems were connected with modelling of plant growth (J.Ross, H.Moldau,
A.Nilson, A.Kiviste, T.Frey, T.Oja et al) and growth limiting and regulating mechanisms (R.Vilu,
A.Koppel, T.Timmusk, K.Kull et al.); the problems of stability in growth models (M.Kahru) and
relations between growth and morphogenesis (T.Neumann) a.o. were also discussed.
the 6th STB - "Rhythms of Life" (May 8-11, 1980, Muuksi) various theoretical problems of periodical
processes and wave phenomena were discussed, especially classification, genetics, ontogenesis and
modelling of biological rhythms (T.Orav, M.Epler, R.Vilu et al.).
At the 7th STB (May
8-11, 1981, Järvselja) the problems of self-organization (synergetics), homeostasis and self-regulation
in organic systems were discussed.
The 8th STB - " Theory of Behaviour" (May 6-9, 1982,
Puhtu) was also the first meeting of Estonian ethologists. Various semiotic (J.Lotman),
ecophysiological (J.Keskpaik), genetic (M.Viikmaa), a.o. aspects of animal behaviour and the
problems of formal language for the description of behaviour (A.Lotman) were discussed.
At the 9th STB - "Theory of Cell" (May 6-9, 1983, Haeska) were discussed and analyzed the
characteristics of elementary living systems, role of discreteness and compartmentability ( A.-P.
Silvere), structure of minimal cell, endosymbiotic cells (T.Tiivel), archigenetic hypothesis of cell
evolution (R.Mikelsaar), modelling of cell physiology and growth (R.Vilu et al.).
10th STB - "Theory of Organic Form" (May 7-9, 1984, Rõuge) - was devoted to theoretical aspects of
morphology and morphogenesis. The problems of quantitative morphology in paleontology (I.Puura),
the history and development of morphological concepts (A.Palumaa, P.Veromann), morphological
analysis of population (R.Mänd), theoretical problems of morphogenesis (H.Kallak, K.Kull) and
morphogenesis of nervous system ( T.Neumann), criteria of symmetry and the relations between
symmetry and entropy of structures (J.-T.Tevet), the paradigm of K.E.Baer in morphology
(M.Remmel), problems of morphology of symbiotic systems (T.Tiivel, U.Sutrop), types and levels of
biological organization (M.Viikmaa), psychophysics of form perception (T.Bachmann) were
The materials of the first five STBs were published in vol. 69 of the
Yearbook of Estonian Naturalists' Society (Problems..., 1984 - see Bibliography), the papers of the 9th
(Raku..., 1983) and of the 10th STB (Orgaanilise..., 1984) were published separately. Selected papers
of the participants of these STBs have been issued in a separate book (Teooria...,
Aladyev V.Z. 1980.
Mathematical theory of homogeneous structures and their applications. Tallinn: Valgus,
Aruja A., Vilu R., Raukas E. 1982. Detection of periodic patterns in RNA
sequences: the first encapsidated region of TMV RNA. - J. theor. Biol., 94(2), 457-
Arvidi A. 1647. Disputatio physica de plants. Dorpati:
Audova A. 1931. Der wirkliche Kamf ums Dasein. - Acta et
Commentationes Universitatis Tartuensis (Dorpatensis), A22 (3), 1-180.
K.E.v. 1876. Reden. St.Petersburg: H.Schmitzdorff. Bd. 1, 1864, vii+296; Bd. 2,
Berg F. 1885. Prädisposition beim variiren der Arten. - Gaea, 4-
Bidder F., Schmidt C. 1852. Die Verdauungssaefte und der Stoffwechsel. Mitau:
Bunge G. 1887. Vitalismus und Mechanismus. - In: Lehrbuch
der physiologischen und pathologischen Chemie. Leipzig: F.C.W.Wogel, 3-
Eichwald E. 1821. De regni animalis limitabus atque evolutionis gradibus.
Dorpat: J.C.Schünmann, x+121.
Engelbrecht J. 1981. On theory of pulse
transmission in a nerve fibre. - Proc. R. Soc. London, A375, 195-209.
J.F. 1819. Ideen zur Aneinanderreichung der rückgrathigen Thiere, aus vergleichende Anatomie
gegründet. Dorpat: J.C.Schünmann, 51.
Folia Baeriana. Tallinn: Valgus. Vol. 1,
1975, 184; vol. 2, 1976, 164; vol. 3, 1978, 252; vol. 4, 1983, 112.
1960. Populatsioonide vordlevast uurimisest evolutsiooniteooria alusel. [On comparative investigation
of populations on the basis of the theory of evolution.] - Proceedings of the Academy of Sciences of
the Estonian SSR. Biology, 9(2), 82-96.
Haberman H. 1968. Elu olemusest. [On the nature of life.] Tallinn: Valgus, 127.
Hartmann N. 1912. Philosophische
Grundfragen der Biologie. Göttingen: Vanderhoek & Ruprecht, 172.
1983. Phytoplankton patchiness generated by long internal waves: a model. - Mar. Ecol. Progr. Ser. 10,
Kallak H.-H. 1984. Darvinism, selle eelkäijad ja edendajad. [Darwinism,
its forerunners and promotors.] Tallinn: Valgus, 120.
Keyserling A. 1853. Note sur
la succession des etres organises. - Bulletin de la Societe geologique de France, 2. Ser., 10, 355-
Keyserling A. 1893. (Über die Darwinsche Theorie.) - Sitzungsberichte der
Naturforscher-Gesellschaft bei der Universität Dorpat, 10(1), 1892. Dorpat, 46-
Keyserling H. 1910. Prolegomena zur Naturphilosophie. München:
Koort A. 1936. Beiträge zur Logik des Typusbegriffs. Teil I
und II. - Acta et Commentationes Universitatis Tartuensis (Dorpatensis), B 38 (4), 1-138, 139-
Leppik E. 1970. Phylogeny, hologeny and coenogeny, basic concepts of
environmental biology. - Acta Biotheoretica 23(3-4), 170-193.
Lippmaa T. 1939.
The unistrated concept of plant communities (the unions). - Am. Midl. Nat., 21, 111-
Lipschütz A. 1915. Allgemeine Physiologie des Todes.
Matthiae S. 1698. Circuli anatomico-physiologici minoris,
segmentum primum; seu dissertatio prima de oeconomia corporis animalis, hoc, est, cogita,
functionum animalium potissimarum formalitatem et causas concernentia. Dorpati: J.Brendeken,
Mikelsaar H. 1975. A concept of amino acid archaeorelation: Origin of life and
the genetic code. - J. theor. Biol., 50(1), 203-212.
Mikelsaar R. 1981. C-A base
pairs in transfer ribonucleic acids and codon-anticodon complexes. - J. theor. Biol., 92(2), 163-
Mikelsaar R. 1983. Human mitochondrial genome and the evolution of
methionine transfer ribonucleic acids. - J. theor. Biol., 105(2), 221-232.
1978. Matemaatika bioloogidele 1. [Mathematics for biologists.] Tartu,
Oettingen A.J.v. 1879. Phänologie der Dorparter Lignosen: Ein Beitrag zur
Kritik phänologischer Beobachtungs- und Berechnungsmethoden. Dorpat,
Oldekop E. 1930. Über das hierarchische Prinzip in der Natur und seine
Beziehungen zum Mechanismus-Vitalismus-Problem. Reval: F.Wassermann,
Oldekop E. 1933. Le Principle de Hierarchie dans la Nature et ses rapports avec
le probleme du Vitalisme et du Mecanisme. (Trad.: M.Nicolas.) Paris: J.Vrin,
Orgaanilise vormi teooria. [Theory of organic form.] (T.Oja, ed.) Tartu, 1984,
Paerna N. 1923. Das Wellenphänomen des Lebens: Periodische Schwankungen
im Seelenleben des Menschen. Leipzig.
Piiper J. 1937. Arenemisõpetuse ajaloo
põhijooni. [On history of theories of evolution.] Tartu: Noor-Eesti, 163.
of contemporary ecology. (T.Frey, ed.) Tartu, 1978, 136.
Problems of the
theoretical biology. Year-book of the Estonian Naturalists' Society, 69. (T.Frey, ed.) Tallinn: Valgus,
Quantitative methods of vegetation study. (T.Frey, ed.) Tartu, 1969,
Raku teooria. [Theory of cell.] (T.Tiivel, ed.) Tartu, 1983,
Regel K. 1921. Statistische und physiognomische Studien an Wiesen. - Acta et
Commentationes Universitatis Dorpatensis A1(4), 1-88.
Reichert C.B. 1845.
Bemerkungen zur verleichenden Naturforschung im Allgemeinen und vergleichende Beobachtungen
über das Blindegwebe und die verwandten Gebilde. Dorpat: E. J. Karow,
Richter H. 1924. Zwei Grundgesetze (Funktion- und Strukturprinzip) der
Lebendingen Masse. - Acta et Commentationes Universitatis Dorpatensis A6(12), 1-
Richter H. 1929. Die Relation zwischen Form und Function und das
teleologische Prinzip in den Naturphänomen. - Acta et Commentationes Universitatis Tartuensis
(Dorpatensis) A18(4), 1-34.
Ross J. 1981. The radiation regime and architecture of
plant stands. The Hague: W.Junk, 391.
Salbom J. 1691. Dissertatio medica,
exhibens spiritus corporis humani pathologiam. Dorpati: J. Brendeken,
Schneider G. Dogma und Problem. Riga : Jonk & Poliewski, 1909,
Seidlitz C.v. 1865. Über die Vererbung der Lebensformen, Eigenschaften und
Fähigkeiten organischer Wesen auf ihre Nachkommen in Bezug auf Physiologie und praktische
Heilkunst. St.Petersb., 72.
Seidlitz G. 1871. Die Darwin'sche Theorie. Dorpat: C.
Seidlitz G. 1876. Beiträge zur Descendez-Theorie. Leipzig:
W. Engelmann, iv+176.
Teichmüller G. 1877. Darwinismus und Philosophie.
Dorpat: C. Mattiesen, vii+89.
Teooria ja mudelid eluteaduses. [Theory and models
in life science.] (T.Tiivel, K.Kull, T.Neumann, U.Sutrop, eds.) Tartu, 1984,
Toompuu O.G., Shcherbakov V.P. 1980. Genetic recombination: formal
implications of a crossed strand-exchange between two homologous DNA molecules. - J. theor. Biol.
Uexküll J.v. 1920. Theoretische Biologie. Berlin: Paetel,
Uexküll J.v. 1928. Theoretische Biologie. 2te gäntzl. bearb. Aufl. Berlin:
Uexküll J.v. 1980. Kompositionslehre der Natur. Frankfurt a.M.:
Vaga A. 1940. Fütotsönoloogia põhiküsimusi. [On some
fundamental problems in phytocenology.] - Acta et Commentationes Universitatis Tartuensis
(Dorpatensis) A35(6), 1-153.
Valt M. 1977. K.E.v.Baer ja Darvinism. [K.E.v.Baer
and Darwinism.] Tallinn: Valgus, 176.
Vilu R. 1983. Calculation of probabilities of
random occurrence of different type of palindromes. - J. theor. Biol., 102(2), 261-
Back to K.Kull's homepage