[Morphogenetic bases and parametric system of trichome evolution in plants of the genus Draba L]

Using quantitative morphological analysis of light microscopy data, the normal variation of trichome morphogenesis is studied in six whitlow grass species (Draba L.) and the morphological variation of adult trichomes in 11 species. The evolution consists in the transition from a radial morphogenesis pattern to bilateral and replacement of complex (branched) trichome rays with simple (unbranched) rays. A parametric system is constructed for classification of the ray morphology; this system includes two parameters-the ratio of the numbers of complex to simple rays, characterizing the probability of secondary branching of primary buds, and the number of primary buds, characterizing the probability of primary branching on the surface of the trichome cell. In this parametric space, all of the studied species fit well a third-order curve consisting of two ascending branches displaying a positive correlation between the primary and secondary branchings and a descending branch, located between them, where the primary and secondary branches are negatively correlated. The deduced evolutionary direction is almost independent of the size of the trichome cells and is explained exclusively by the mechanics of morphogenesis: acceleration in the development of the primary bud of the ray decreases the probability of its own branching and creates additional elastic extension of the cell surface, preventing other buds from branching. The morphogenesis itself appears to be a mechanically nonholonomic system, filtering in a selective manner the fluctuations of the same sign, which explains the directed pattern of its evolution. In the evolutionarily initial state, trichome ontogenesis is absent because its modules (primary buds) are formed by a mirror duplication. The ontogenesis commences when mirror symmetry in the formation of modules is lost and replaced with an axial pattern; thus, the change in the morphological type of buds is a direct consequence of the emergence of ontogenesis and its further evolution. Its main material is intraindividual variation, the only source of which is the mechanics of morphogenesis itself. It is found that morphological evolution can take place at an initially zero heritability and zero adaptive value of morphological differences.