Constructive organization of forest-forming tree species growing in the middle zone of European Russia

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Abstract

The purpose of this report is to reveal features of structural organization of 12 species of trees of the temperate zone, which constitute the first layer of phytocenoses, from the standpoint of the concept of C. Edelin (1991). The structural organization of the studied species is based on the principle of hierarchic construction of the crown, i.e. the subordination of lateral skeletal axes to the main axis, the trunk. In flowering plants, sympodial growth is widespread, in one of its variants there are prerequisites for the appearance of elements of polyarchic organization in the crown – two equivalent axes forming a fork, which sometimes leads to a bifurcation of the trunk. Subsequent differentiation of the fork axes into the leading and subordinate can lead to the formation of branches that exceed the branches of the hierarchic organization in terms of durability and power. The appearance of elements of a polyarchic organization is less determinate than of the elements of a hierarchic one. Among the studied species, the forks appear less often in Picea abies, and more often in Salix alba and S. pentandra. In shade-tolerant species under low light conditions, the inclusion of elements of a polyarchic organization in a hierarchic structure underlies the formation of an umbellate crown, and in open space in all species, except Picea abies, it allows forming more spreading crown. No correlation was revealed between the structural organization of the studied species and their role in the successional system, as well as between their longevity and shade resistance.

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About the authors

M. V. Kostina

Sevastopol State University

Author for correspondence.
Email: mv.kostina@mpgu.su
Russian Federation, Universitetskaya Str., 33, Sevastopol, 299053

N. S. Barabanshchikova

Moscow Pedagogical State University

Email: ns.barabanshchikova@mpgu.su
Russian Federation, Kibalchich Str., 6/3, Moscow, 129164

O. I. Nedoseko

Lobachevsky State University of Nizhny Novgorod – National Research University

Email: nedoseko@bk.ru
Russian Federation, Arzamas Branch K. Marks Str., 36, Arzamas, 607220

M. N. Stamenov

Samara Federal Research Scientific Center RAS, Institute of Ecology of the Volga River Basin RAS

Email: mv.kostina@mpgu.su
Russian Federation, Komzin Str., 10, Togliatti, 445003

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2. Fig. 1. Shoot systems formed within two years in the species with sympodial growth: а – hierarchic shoot system; б – polyarchic system with opposite leaf arrangement and two replacement shoots forming a “fork”; в – polyarchic system with alternate leaf arrangement and two replacement shoots forming a “fork”; г – inexplicit polyarchic shoot system. 1 – bud ring; 2 – lateral shoots; 3 – dead shoot top; 4 – replacement shoot.

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3. Fig. 2. Shoot systems formed within two years in the species with monopodial growth: а – hierarchic shoot system formed as a result of monopodial growth with a pronounced length predominance of the shoot from the apical bud over lateral shoots; б – inexplicit hierarchic shoot system formed as a result of monopodial growth with slight length predominance of the shoot from the apical bud over lateral shoots; в – hierarchic shoot system formed as a result of slowing growth of the shoot from the apical bud and the formation of a replacement shoot; г – polyarchic system formed as a result of slowing growth of shoot from the apical bud and the formation of two replacement shoots; д – polyarchic system formed as a result of equal growth rates of the shoot from the apical bud and a lateral shoot. 1 – bud ring; 2 – lateral shoots; 3 – a shoot from the apical bud, exceeding lateral shoots in length and thickness; 4 – a shoot from the apical bud, lagging in development from lateral shoots; 5 – lateral shoots outgrowing the shoot from the apical bud and forming a fork; 6 – a shoot from the apical bud forming a fork with a shoot from a lateral bud; 7 – a lateral shoot not lagging in growth from the shoot from the apical bud.

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4. Fig. 3. Hierarchic and polyarchic systems formed as a result of sylleptic branching: а – hierarchic system with sylleptic shoots not outgrowing the maternal one; б – hierarchic system with one of the sylleptic shoots outgrowing the maternal one; в – polyarchic system with two sylleptic shoots outgrowing the maternal one and forming a fork. 1 – zone with inactive buds; 2 – zone of sylleptic shoots; 3 – completion of growth; 4 – sylleptic shoots; 5 – shoots of regular innovation; 6 – crown bud; 7 – replacement shoots, the function of which is performed by sylleptic shoots.

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5. Fig. 4. Main variants of fork development within five years: а – formation of two equal powerful axes developing according to the trunk program for four years; б – formation of a powerful axis of II order, formed as a result of a fork partition in the third-fourth year after its appearance; в – formation of an axis of II order hardly differing from other axes appeared from the buds of regular innovation. 1 – fork formed within five years; 2 – powerful side axis of II order; 3 – axis performing the function of trunk; 4 – side axis of II order formed as a result of differentiation of fork axes in the next year after appearance.

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6. Fig. 5. Hierarchic structural organization of trees with elements of polyarchic (а, б) and hierarchic organization of the crown (в). 1 – trunk bifurcation; 2 – dying axes; 3 – powerful axes formed as a result of differentiation of fork axes; 4 – growing axes.

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7. Fig. 6. Umbellate crown of Acer platanoides under the forest canopy (a), appearance of a leading axis after formation of a light gap (b). 1 – fork consisting of plagiotropically oriented axes; 2 – axis changed the direction of growth from plagiotropic to orthotropic; 3 – axis formed from an axillary bud.

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8. Fig. 7. The percentage of Betula pendula trees growing in an open space (а) and in plantations (б) (sampling of 30 trees for each variant of growth): without trunk bifurcation, with trunk bifurcation, with formation of a powerful branch. Blue – Trunk not bifurcated, orange – Trunk bifurcated, gray – Heavy branch from the trunk.

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9. Fig. 8. Constructive organization of the studied species from the standpoint of the C. Edelin’s concept of crown organization: а – hierarchic constructive organization of Populus tremula, Betula pendula, and B. alba with rapid leveling of frequently appearing forks; б – hierarchic constructive organization of Salix alba and S. pentandra with a high share of polyarchic elements; в – hierarchic constructive organization of Acer platanoides with regularly appearing polyarchic elements; г – hierarchic constructive organization of Picea abies with rapid leveling of rarely appearing forks.

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