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
Selected bibliography


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).

According to 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 challenge.

Theoretical Science in an Ecologically Oriented Culture

The 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 word.

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.

Biology is 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.


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).

The 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 Estonia.

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.

The most 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 nomogenesis.

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.

The neovitalist 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, 1938).

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.

Theory of Evolution

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.

N.I.Andrussov (1861-1924) who specialized in paleontology and geology was a Darwinist; his work was on the role of bacteria on evolution.

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.

F.Ovsjannikov, 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.

Aleksander Audova (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 Classification

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 E.G.Saint-Hilaire.

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.

General 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 cells.

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 University.

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 nature.

Ants-Peep Silvere (1933-83) developed theoretical principles of micromorphology, especially the principle of discretess (compartmentation) of life.

Theoretical problems 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, T.Neuman a.o.

Theory of Geobotany and Plant Productivity

Estonian 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 communities.

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.

Mathematical Methods and Modelling

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- 42).

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 a.o.


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 appeared.

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 Naturalists' Society.

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 theoretical biology.

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.


A short 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, T.Neuman) etc.


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 place.

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.

At 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.).

The 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 discussed.

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..., 1984).

Selected bibliography

Aladyev V.Z. 1980. Mathematical theory of homogeneous structures and their applications. Tallinn: Valgus, 268.

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- 470.

Arvidi A. 1647. Disputatio physica de plants. Dorpati: J.Vogelius.

Audova A. 1931. Der wirkliche Kamf ums Dasein. - Acta et Commentationes Universitatis Tartuensis (Dorpatensis), A22 (3), 1-180.

Baer K.E.v. 1876. Reden. St.Petersburg: H.Schmitzdorff. Bd. 1, 1864, vii+296; Bd. 2, xxv+480.

Berg F. 1885. Prädisposition beim variiren der Arten. - Gaea, 4- 5.

Bidder F., Schmidt C. 1852. Die Verdauungssaefte und der Stoffwechsel. Mitau: G.A.Reyher, x+413.

Bunge G. 1887. Vitalismus und Mechanismus. - In: Lehrbuch der physiologischen und pathologischen Chemie. Leipzig: F.C.W.Wogel, 3- 15.

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.

Eschscholtz 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.

Haberman H. 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.

Kahru M. 1983. Phytoplankton patchiness generated by long internal waves: a model. - Mar. Ecol. Progr. Ser. 10, 111-117.

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- 358.

Keyserling A. 1893. (Über die Darwinsche Theorie.) - Sitzungsberichte der Naturforscher-Gesellschaft bei der Universität Dorpat, 10(1), 1892. Dorpat, 46- 66.

Keyserling H. 1910. Prolegomena zur Naturphilosophie. München: J.F.Lehmann, xii+160.

Koort A. 1936. Beiträge zur Logik des Typusbegriffs. Teil I und II. - Acta et Commentationes Universitatis Tartuensis (Dorpatensis), B 38 (4), 1-138, 139- 263.

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