Is Natural Selection still have to be Regarded A Foundation Stone of Evolutionary Process ?

Natural selection is a buzzword used to describe the main driving force of evolution. Its creative role is believed to be based on: a) an unlimited variety of organisms caused by hereditary variation and b) a direct connection between hereditary changes and their phenotypic expression. These are the two requirements that can lead to the genetically based changing modalities of characters through “iterations” of natural selection in the series of successive generations. Are these two requirements fulfilled in the nature, however? The present study focuses on the analysis of these two “foundation stones” of natural selection. Firstly, hereditary variation is shown to be essentially non-homogenous. New hereditary characteristics of individuals fall onto a narrow “strip of land” in the sea of potential possibilities. Secondly, the consequences of changes in the genotype of an organism are involved into a system of hierarchical multiple compensation, from the molecular to the biocenotic level. In a way, the signal of hereditary change passes through a series of “system filters” at epigenetic, ontogenetic, physiological, behavioural, populational and biocenotic level. Each filter is represented by multiple feedbacks maintaining the integrity of systems at each level and at all the hierarchical levels taken together. It is in these “system filters” the adaptive nature of characters is formed representing the every individual as a subject to the Law of Multilevel Self-Organization. The emerging understanding of this provides a strong reason to change the evolutionary paradigm from the mainly selectogenetic to the mainly orthogenetic one. DOI : COMING SOON Corresponding Author: Andrey I. Granovitch, Department of Invertebrate Zoology, Faculty of Biology, St. Petersburg State University, Universite tskaya nab. 7/9, St. Petersburg, 199034, Russia, E-mail: a.granovich@spbu.ru Running title: Is Natural Selection a foundation stone of evolution?


Introduction
Natural selection, laconically expressed by Spenser and Darwin as "…the survival of the fittest" 23 14,20,29,74 . We may, therefore, be fairly sure that the contribution of individuals into the gene pool of the next generation is unequal. Whether this unequal contribution is connected with evolution, a vector process involving series consisting of hundreds and thousands generations is, however, far from being evident. Such a connection, if any, has to be separately proved. It may well turn out that we observe mere stochastic or cyclic populational-genetic processes 1 rather than changes at the evolutionary scale allegedly caused by "natural selection". So, the differential contribution of individuals into the gene pool of the next generation as such does not prove that the mechanism of natural selection is at work. This begs the question: can natural selection be considered as the main driving force of evolution? This is Note: In our opinion, the notion of selection should not be expanded to mean the universal synthesis of the oppositions of "prohibition and permission" (see 77 , p. 252). Such an expansion makes it impossible to disentangle the processes of structural self-organization at various levels and the processes ruled by the mechanism of If found, a distinct correlation of this kind would provide compelling proof of the effectiveness of natural selection as an evolutionary factor.

Potential Material for Natural Selection
Stating the Question Heterogeneity of organisms in respect of reproductive characters -random, unlimited and independent of the environment -is the cause of differential input of individuals into the next generations. The major mechanism of transformation is selection of the "fittest", or natural selection. Cumulatively, in the series of generations, it results in adaptive changes of the modal species characteristics. The environment determined the direction of selection, and therefore the model is ectogenetic: it admits transformism under the influence of external factors. The model is also idiographic, as it focuses on the uniqueness of transformation acts and their probabilistic character. The ideas of indirect adaptogenesis underlie the "modern evolutionary synthesis" of the 20 th century.

Direct adaptogenesis.
The impact of the environment results in adequate changes in the morpho-functional inheritable characteristics of the organism. In this way, adaptive changes of organisms are not mediated by selection but form directly under the influence of the environment in a series of generations. This means that the model admits the existence of ectogenesis, transformation under the influence of external factors. The model is idiographic: each transformation act, being determined by a unique combination of the environmental conditions, is considered as unique. The mechanism of direct adaptogenesis underlies the ideas of various Lamarckian evolutionary hypotheses.
The model considers internal patterns of morpho -functional organisation of living systems as the driving force of transformation. Its logic can be characterized as "self-assembly" (rather than selection!) of increasingly complex systems. Therefore, it is the only model that consistently admits the existence of autogenesis: the change based on internal patterns of the organisms' structure. According to this model, variation of organisms is strictly directional rather than random. The model is nomothetic: it searches for strict laws of evolutionary transformations and envisages the possibility of predictive interpretations. On the whole, it admits the existence of internal directionality of the evolutionary process, in other words, orthogenesis. The main problem of orthogenetic interpretations of evolution is associated with the explanation of the formation of adaptations in the course of evolution, that is, their correspondence to the environment 70 . categories of a higher rank. In all these cases it has been shown that only a minor proportion of the potential diversity is actually implemented in the nature 18,75,78,100,101,102,113,118,119,121 . This discrepancy between the potential and the actual diversity may be an argument in favour of essential structural constraints of the origin of forms (that is, the primary "nonhomogeneity" of variation). At the same time, this assessment is not necessarily incompatible with a different viewpoint: the initial variation is reasonably "homogenous" and the subsequent non-homogeneity is the result of the mechanism of differential mortality, that is, natural selection. ("anti-cryptic selection"?) 16  Hereafter the term "system filter" will be used to It is only from this viewpoint that its significance can be assessed.

Conclusion
Two major components should be considered when discussing whether the mechanism of natural selection does act in reality: the structure of variation and the complex of hierarchical "system filters".
As concerns the structure of variation: we can conclude that new hereditary characteristics of individuals fall onto a narrow "strip of land" in the sea of potential possibilities (see Fig. 1). Hereditary variation is non-homogenous, and very much so. This means that it is the characteristics of variation that determine the direction of potential evolutionary changes. It is utopian to think that natural selection has a virtually unlimited range of hereditary variations at its disposal. It does not, and this is due, first of all, to the character in which variation is manifested, its non-homogeneity. We have to admit that in the continuum of possible evolutionary mechanisms ranging from orthogenetic (a narrow range of the potentially possible variation is implemented) to selectogenetic (much of the potential variation is implemented) orthogenesis rules the day (see Fig. 1).
Just how effective can selection be within the range of we have represented this process as passage through a series of "system filters" (Fig. 2). Each filter is represented by multiple feedbacks, compensatory reactions, which maintain the integrity of systems at the epigenetic, ontogenetic, physiological, behavioural, populational and biocenotic level. The first system filters plane. Therefore, it would be better, for the sake of clarity, to call it "differential mortality" rather than continue using the term "natural selection", strongly linked with the idea of "creative evolutionary potential".

Acknowledgments
I am eternally grateful Natalia Lentsman for her invaluable contribution to the English translation of the