Nomogenetic biology and its Western counterparts

Sabine Brauckmann, Kalevi Kull

Published in:
Naumov R.V., Marasov A.N., Gurkin V.A. (eds.) 1997. Lyubischevskie Chteniya 1997. Ul’yanovsk: Ul’yanovskij gosudarstvennyj pedagogicheskij universitet, pp. 72-77.

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The history of theoretical biology has not yet been extensively reviewed by the historians of science, and particularly the comparison between the work done in different cultures is still a white spot on the map of biological sciences.

In contrary to T.Kuhn's view on the development of science as a periodical replacement of alternative paradigms, in the case of a more close and detailed view on biology there could be seen a permanent and parallel co-existence of alternative long-lasting approaches, which differ remarkably by their dominance and spread in scientific community.

In such a way, e.g., von Baer's (1876) critics of Darwin and the antinomies of these two approaches (Darwinian and Baerian) have been discussed in Russian biology for a long time, in some cases with rather dramatic counter-effects, but a comprehensive analysis of it is still absent (cf. Beloussov 1994; Graham 1993; Mikulinskij, Polyanskij 1983). The non-Darwinian or anti-Darwinian views of the the turn of the century have been quite well analysed by historians of science (Bowler 1992), what is not the case concerning all the following decades. This is true especially for the nomogenetic view formulated by the Russian geographer Lev Berg (1926) and later developed by Aleksandr Lyubischev (1968; 1982; Svetlov, Meyen 1982), Sergej Meyen (1979; 1988; Meyen et al. 1977), and others (Chajkovskij 1990), even though without a wider recognition yet, serving as a remarkable participant in the diversity of types of biological explanation.

If one wants to frame this Russian approach and to show on which path it is leading, one could state as follows. From the point of view of making emphasis on typology, it is close to structuralism (Schreider 1977). Emphasising the logical correctness in biological discourse, and the exact definition of terms, it is close to Joseph Woodger's (1929) early approach to deduce formally biological theory from a few principles. Stressing the independence, or at least importance, of the laws of form, it reminds D'Arcy Thompson, or later B.C.Goodwin, W.F.Gutmann, A.Lima-de-Faria. Looking for rules and predictability in evolution, it may remind B.Rensch (1961; 1970), or T.Eimer. Discussing with physicalists and liking systemic holists, it is close to H.Driesch, and may be to L.v.Bertalanffy. Criticizing the importance of natural selection in the evolution, it is close to M.Kimura, or N.Eldredge and S.J.Gould (1977), or H.Paterson (1993). In later or recent years, somewhat similar to the nomogenetic school views have been represented by R.Thom (1972; 1975; 1983), S.Kauffman (1993), and also S.Newman (1994), G.Müller, J.Mittenthal, et al. There has, however, probably not been any corresponding equally strong anti-Darwinian school in Western evolutionism after 1930s.

Due to the limited space we will confine our remarks about the Western parallels to R.C.Lewontin and B.C.Goodwin, choosing them as the most terse biotheoreticians of an anti-Darwinian approach, and also considering their educational background - Lewontin is a trained evolutionary geneticist and Goodwin a mathematical biologist - thus representing two divergent realms of biology. Both of them are influenced by the Cambridge Club of Theoretical Biology, especially by Needham, resuming this biotheoretical concept of the 1920s and 30s, and further developed it. Therefore, it is appropriate to give a short sketch of the most fundamental statements of Needham.

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J.Needham connected his approach of law- determined evolution with thermodynamics. His basic statements on evolution governed by laws set that during evolution the quantity of organic components being necessary for development (ontogeny) enlarges and that simultaneously the degree of complexity of the morphological forms and geometric relations between the components increases qualitatively. By this process of complexity the components of lower levels of organisation are enfolded like envelopes by more complex ones. During the long periods of evolution the control functions of the organisms became efficient, so that organisms become autonomous entities towards their environment. Due to this autonomy they are qualified to expand their activities in space to the extent that enables them to construct their own environment. “A living organism is both a ‘patterned mixed-up- ness’ and a ‘patterned separatedness’. ... The point is that the works of organisation have a certain similarity at all levels of their operation” (Needham 1976, 178).

According to Lewontin's (1985) approach, the organisms are defined as subjects of evolution and objects for evolution. They are subjects, because they can autonomously select what is relevant for them, e.g. their food. Furthermore, they are able to exploit physical signals from the environment. As an example may serve the phenomenon of external fluctuations of temperature transformed by inner organs into (bio)chemical signals. That is the reason, why neither environment nor organisms could be comprehended as distinct and separated entities. Hence, both represent an interlocked network. Applied to the feature of evolution, it means that the relation between organisms and environment determines also the feature of selection. According to Lewontin, ontogeny is a branching set of developmental pathways taking organisms as unifying particles of external and internal forces. It characterises organisms as the place of the interactions, by which their survival is simultaneously conditioned. Every organism here constructs its own environment, and participates to its self-construction and to the creation of its environment being conditional for its survival. This does not contradict to the statement that organisms are created by interaction of gene and environment. Therefore, adaptation alone is not able to base and explain morphological structure. Consequently, he concludes, one should speak of construction instead of adaptation (Lewontin 1985, 104).

To formulate it in a more detailed way, one could state, concerning the relationship between gene and environment, that morphology, physiology, metabolism, behaviour - shortly the phenotype - is the product of genes (genetics) and environment. And exactly to this product the organism has developed itself according to its spatial and temporary constraints. “...Information from the environment modulates the biosynthetic pathways in a way that matches the rate of activity to the demand for the product. ... The arrows of causation point from gene and environment to organism. In fact, however, the organism participates in its own development ... because it is the determinant of its own milieu” (Lewontin 1985, 94, 96).

Lewontin criticizes classical Darwinian population genetics for two reasons: firstly, it ignored the organism, and secondly, it ascribed fitness to a given gene. These failures or desiderata are caused by the initial question for which the theory was constructed, namely, could a gene conferred a slight selective advantage spread through a population and displace the former wild-type gene? For him the entire body of theory is epistemologically incomplete (Lewontin 1974). To elude such shortcomings he postulates a theory mapping the genotype to the phenotype, by which the phenotype is related to fitness and by which changes in phenotype refer to resulting changes in genotype.

In a programmatic article written by him and S.J.Gould (Gould, Lewontin 1979, 581) they introduce against the adaptationist programme the statement, that organisms

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must be analysed as integrated wholes controlled by Baupläne, which on the other hand are constrained by phyletic heritage. In this evolutionary model pathways of development and morphological structure called architecture constraint themselves by delimiting mutational changes. That means, these specific pathways are more important for evolution than selective force that may mediate change when it occurs. For Gould and Lewontin the adaptationist programme fails to distinguish current utility from reasons for origin as well as to consider alternatives to adaptive stories. Likewise they criticise it, because it forgets to consider adequately such completing themes as random fixation of alleles, production of non-adaptive structures by developmental correlation with selected features (e.g. allometry), the separability of adaptation and selection, multiple adaptive peaks, and current utility as an epiphenomenon of non-adaptive structures. Therefore, the adaptationist programme is named Panglossian paradigm.

In this context they refer to German paleontology, namely to O.H.Schindewolf, A.Remane, and P.P.Grasse. For all of them natural selection can only explain superficial modifications of the Bauplan that fit structure to environment. But the construction of the Bauplan itself and transitions between Baupläne must involve internal mechanisms. According to Gould and Lewontin such a strong argument is too close to an appeal to mysticism. Therefore, they propose a weaker argument by emphasising the importance of constraints for developmental programmes (Gould, Lewontin 1979, 594). The constraints themselves are divided in two categories, i.e. phyletic and developmental being also a subcategory of phyletic restriction. Here Gould and Lewontin refer explicitely to A.Seilacher (Seilacher 1970), for whom structural restrictions cause architectural constraints for development, that never were adaptations, but rather the necessary consequences of materials and designs selected to build basic Baupläne.

Goodwin formulates a structuralistic approach to biology which formally presupposes to static structure a processual event borne by function. This is explained by the statement, that such a forming process guarantees systemic wholeness, holoblastic cleavage patterns, self-regulation and heredity. “Functionalism is thus included in the dynamic structuralism that I have described because a stable life cycle, an attractor in morphogenetic space, includes the environment as part of the field dynamic that generates the form of the life history. ... It [structuralism] involves developing a theory of organisms as processes - life cycles ...” (Goodwin 1990, 238, 240). One should note here that heredity is interpreted by memory, which causes a process of reproduction. “In other words, genomic DNA is functionally and structurally as flexible and changeable as the rest of the organisms” (Ho 1989, 32). Premise is, that a limited number of realizable biological structures, determined by physico- chemical constraints and being inherent in living organisms, are manifested in evolution. The biogenic structure results from necessity determined by the environment. In this context Goodwin refers to Goethe's dynamical archetype, which is interpreted as a proto-element of a simply structured set of transformations. “Thus, process structuralism can be viewed as a marriage of rational morphology and neo-Darwinism” (Resnik 1994, 9).

The structuralistic approach generates a dynamically floating morphologies, because they are focused on the physiological variables - in contrary to Turing's mathematical model for chemical morphogenesis. Formulated in mathematical terms, dynamics causes a geometric form (gestalt). This morphological pattern reacts to the former dynamical state, and by this process, a topological geometry is constructed. For Goodwin, it is objectively wrong to assume, that a set of chromosomes alone constitutes a finite class of instructions or informations for determining a temporary order of events and the details of morphology. Furthermore, he corrects the hypothesis, that DNA of an organism is self-replicating, by referring to S.Spiegelman's (1967) experimental data.

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Concerning the problems of selection, it results from this approach that the features of living systems are feasible by selection and could be stabilised by it, too. “We could, if we wished, simply replace the term natural selection with dynamical stabilization, the emergence of the stable states in a dynamical system” (Goodwin 1994, 53). Consequently, the Darwinian selection defined as a blind game by chance is not qualified to press ahead the evolution. “This accounts for the equifinal nature of development (Bertalanffy 1952), and also for the non-random nature of phylogeny” (Lambert, Hughes 1989, 69).

To sum it up, the objective of biological structuralism is to uncover the unity of dynamical process, by which the structured manifoldness of forms (gestalten) is constructed and on which the laws and principles of organization conceptually are based.

On the place of conclusion, we will make an attempt to list the most fundamental ideas of the nomogenetic school.

(1) Taxonomy of organisms should be something like Mendeleyev's table of chemical elements, i.e. non- phylogenetic, combinative classification. The natural classification should reflect the internal laws of structure and variation without being phylogenetic, while phylogenetic relations reflect history.

(2) Congruent to taxonomy dealing with classification of organisms is meronomy, which structures components of holistic systems, e.g., organs in organisms.

(3) Natural selection plays the secondary and mainly negative role in evolution. (Using a literary expression - in contrary to Darwinism, environment cannot select, because the ability to select means ability to choose, and it assumes subject; a subject, i.e. organism and not environment, is able to make choices, to select. There is more than only a difference in metaphor).

(4) A natural system could be built, although evolution is often polyphyletic, and in many cases almost non- reconstructable.

(5) Organisms of different taxa have different archetypes, which determine their possible ways of evolution.

(6) Morphogenetic rules rather than ecology is responsible for the limits and paths of variation.

(7) The problems of form rather than function have the primary importance.

An important note is that the adaptive evolution may be important primarily at species and intraspecies levels, whereas the statements of nomogenetic view work for higher taxa. For the nomogenetic school the neo-Darwinian theory of evolution formally considers the phenotype and genotype to be existing simultaneously. It means that there are no any important theoretical conclusions made which could result from the time difference of changes in phenotype and genotype. In this sense, what is proposed is actually not a refutation of the neo-Darwinian theory, but a more general view in which the possible time difference is added. It also means that the theoretical object of biology is grasped in a basically more diverse way. When the time difference is becoming equal to zero, i.e. the system is simplified, we may get exactly the existing theory as a special case. Such a generalisation is needed for finding a real place for nomogenetic factors in evolution.


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