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Contemporary Problems of Ecology

2020 year, number 3

Ecological morphology of the protoepiphytic fern Lemmaphyllum microphyllum C. Presl in connection with adaptatiogenesis

N. M. Derzhavina
Orel State University named after I. S. Turgenev, Orel, Russia
Keywords: протоэпифиты, нанизм, суккулентизация, адаптивная стратегия, олиготрофия, protoepiphytes, nanism, succulentization, adaptive strategy, oligotrophy

Abstract >>
The study of the ecological morphology of plants is a necessary condition for solving a number of fundamental and applied problems in the field of ecology, botany, geography and physiology of plants, when conducting global and regional environmental monitoring, as well as in the selection and introduction of plants. This approach is successfully used for the study of flowering plants of different botanical and geographical zones. Ferns in this aspect tend to fall out of the field of view of botanists. The aim of this work was to study at different levels of organization from the cellular to the organizational photosynthetic apparatus of the fern sporophyte protoepiphyte Lemmaphyllum microphyllum and attempt to answer the question about the ways of its adaptation to the conditions of existence. The laboratory complex of analysis of morphology and structure of photosynthetic apparatus of plants-Siams Mesoplant-was used. It allowed us to conduct a comprehensive analysis of L. microphyllum at the level of fronds, mesophyll cells, plastid apparatus and obtain indicators: specific surface density of fronds, the number of cells per unit area of fronds, the number of chloroplasts in one cell; fronds thickness, surface area and volume of chloroplasts, area and volume of cells, etc. It was found that adapting to living conditions on the trunks and branches of trees, as well as flat rock surfaces devoid of humus, the fern involved different organs and functions. At that completely in its morphology and anatomy showed adaptations to light and humidity mode. At the same time, adaptations were developed both for the most complete capture of light, and for protection from excess light, to reduce the loss of water and its release. They have appeared at different levels of organization and include combined xero-, meso -, helio-and sciomorphic traits that allow the species to form stable populations in cenoses. It is shown that the optimal compensatory strategy was, when reducing the area of fronds, the development of water-holding capacity of tissues and cells on the basis of polyfunction of chlorenchyma (succulentization), as well as the improvement of the conductive elements of the xylem due to the appearance of vessels. These characteristics did not solve the problem of long-term dehydration, but allowed only a short-term water deficit to be tolerated. This gives reason to call such ferns hidden succulents or subsucculents. It was shown that the optimal compensatory strategy was, with a reduction in the fronds area, the development of the water-holding ability of tissues and cells (succulentization) based on the multifunctionality of the chlorenchyma, as well as the improvement of the conductive elements of xylem due to the appearance of vessels. These characteristics did not solve the problem of prolonged dehydration, but allowed only a short-term water deficit to be tolerated. This gives grounds to call such ferns hidden succulents or subsucculents.

Sporophyte biomorphology of Athyrium species in the different environmental conditions in South Siberia

I. I. Gureyeva
Tomsk State University, Tomsk, Russia
Keywords: Athyrium filix-femina, Athyrium distentifolium, папоротники, спорофит, жизненная форма, онтоморфогенез, Южная Сибирь, ferns, sporophyte, life form, ontomor

Abstract >>
The aim of the work to present the biomorphological characteristics of ferns of the genus Athyrium Roth, occurring in South Siberia, and to identify changes of the life form during ontogenesis and in individuals of the same biological age in different ecological and coenotic conditions. Biomorphological study of plants is an important task since both the organization of populations of species and their stable existence in cenoses depend on the life form of the species. The main thing in the identification of life forms of sporophytes of ferns is the study of the formation of its perennial part, which is the rhizome. Based on the structural features of the sporophyte, we define the life form of the sporophytes of the Athyrium species living in South Siberia as summer-green polycyclic densely-rosette ferns with monopodial-dichotomously growing underground-aboveground rhizomes bearing rosette of monomorphous fronds at the apical end. The life form of the different Athyrium species is represented by two variants: (1) large-rhizome orthotropic weakly branching vegetatively non-growing monocentric ferns ( Athyrium filix-femina (L.) Roth., A. monomachii Komarov, A. sinense Rupr.), and (2) large-rhizome ascending weakly branching and weakly vegetatively growing monocentric or implicit-polycentric ferns ( A. distentifolium Tausch ex Opiz). Our study showed that the rhizomes and fronds size is increased, and the ability of rhizomes to branching is realized during sporophyte ontogenesis. The branching is dichotomous: the daughter branches are formed inside the terminal bud of the maternal branch and become distinguishable only after several years. It was determined, that rhizome remains uniaxial throughout life in sporophytes with low vitality, and rhizomes tend to branch in sporophytes with high vitality. Branching of the rhizome starts in young spore-bearing sporophytes. The largest number of branches of the rhizome, and consequently rosettes of fronds is characteristic for middle-aged spore-bearing sporophytes. At the end of the life, non-specialized disintegration (senile partitioning) occurs in the sporophytes having the branched rhizomes. This is expressed in the disintegration of the rhizome into the separate branches. According to the results of the study in various phytocenoses of the Kuznetsk Alatau, the intensity of rhizome branching in sporophytes of the same ontogenetic stage depends on environmental conditions. The rhizomes of A. filix-femina produce a large number of branches in the lighter mixed (fir-aspen and fir-birch) forests; in A. distentifolium , the intensity of the rhizome branching increases from the forest belt towards the subalpine one.

Shoot formation model as basis for adaptations of flowering plants

N. P. Savinykh, S. V. Shabalkina
Vyatka State University, Kirov, Russia
Keywords: модель побегообразования, побег, модуль, условия переменного обводнения, поливариантность развития, адаптация, shoot formation model, shoot, module, conditions of variable watering, development multivariance, adaptation

Abstract >>
The work is aimed at the description of biomorphology of plants living in variable flooding conditions. The results expand the concept of modular organization as a constructive and representative way of comparative morphological analysis and explanation of the mechanisms of biomorph evolution. Structural organization of biomorphs in the gradient of change in the stability of humidification from floodplain meadows to coasts to water cuts and beaches: Solanum ulcamara , Thalictrum simplex , Lysimachia vulgaris , Lycopus europaeus , Stellaria palustris Persicaria amphibia , Erysimum cheiranthoides is studied from the perspective of a systemic and complementary approach. It has been determined that acceleration in the vegetative and generative spheres leads to the appearance of monocarpics in the hygrophilic line of evolution. It is shown that ontogenesis of individuals of seed and vegetative origin is reduced to a single fruiting and the specificity of the shoot formation model is preserved in plants in all considered biotopes. Specific signs identified: the presence of long internodes in the substrate, in the supersubstrate parts during flooding, in grass rags and dense grass under low light; sympodial growth by formation of shoots of substitution. On the model species, it was determined that plant adaptability is provided by changes in individual structural elements of monocarpic shoots, and the structural and functional zonation of monocarpic shoots determines the autonomization of parts of an individual. Our results allowed us to confirm that the monocarpic shoot and the system formed on its basis is one of the categories of axes of a herbaceous plant as part of an architectural unit (as understood by D. Barthelemy and Y. Caraglio). The data obtained showed that the settlement of reservoirs with flowering plants was possible from long-shoot long-rhizome grasses and subshrub lian in the following directions: 1) the emergence of monocarpic annuals; 2) the emergence of young and annual vegetative origin with the formation of acentric and clearly polycentric stolon-forming and root-shoot biomorphs.

The sympodial model of bulb growth in Amaryllidaceae: a comparative morphological approach

V. V. Choob
Botanical Garden of Lomonosov Moscow State University, Moscow, Russia
Keywords: предлист, низовые чешуи, морфология луковиц, симподиальное возобновление, prophyll, lower scales, bulb morphology, sympodial innovation

Abstract >>
For a long time Amaryllidaceae served as a textbook example of monopodially growing bulbs. The main argumentation in favor of this viewpoint was the position of the first leaf of the innovation bud in Galanthus : it was not placed with its abaxial side to the inflorescence stalk. The discussion on monopodial or sympodial innovation was continued for more than 150 years, and was completed by the victory of the sympodial model. Nevertheless, the variability of phyllotaxis of the innovation bud in Amaryllidaceae did not receive satisfactory explanation by nowadays, that brings actuality to our research. We have reviewed and re-discovered the prophylls of the innovation bud throughout all the Amaryllidaceae family, leading to strict homologization of these leaves in different clades of the family. By the means of comparative structural morphology we rendered the universality of the sympodial model of bulb innovation in all the members of Amatyllidaceae. Based on the results of the provided study, we have postulated, that the prophyll of the innovation bud undergoes the reduction from well-developed green leaf (photoprophyll) through lower scales (cataprophylls) to complete ablation (phantom prophyll). Moreover, we revealed, that the prophyll morphological characters are synapomorphies for the principal geographical clades of Amaryllidaceae (established according to molecular phylogeny data). The major variability of the prophyll structure we documented for the African tribes of Amaryllidaceae. It is important to mention, that the postulated row of morphological reduction of the prophyll could not be referred to as the row of evolutionary events. As for the common ancestor of Amaryllidaceae, we propose the polymorphic state with different degree of the prophyll development. Further evolution lead to stabilization of the prophyll morphology in the majority of geographical clades, but African Amaryllidaceae. Correspondingly, our data undoubtedly proved the model of sympodial innovation for the bulbs in all the Amaryllidaceae family.

On architecture models of annual astragali (Astragalus L., Fabaceae)

A. K. Sytin, L. V. Ryazanova, D. D. Slastunov
Komarov Botanical Institute of RAS, St. Petersburg, Russia
Keywords: annual astragali, architectural models, evolutionary morphology, adaptation, taxonomy

Abstract >>
The typological problem of revealing architectural models of annual astragali ( Astragalus L.) has emerged a close relationship between the shoot system and the structure of inflorescences. The features of the selected four patterns of architectural models are discussed. The synthesis of approaches has given us insight into the evolutionary trends. The application of the method of architectural models provide the means of classifying of life forms of annual astragali. The synthesis of molecular data, taxonomy and biomorphology confirms the concept of mosaic evolution and heterochronism (heterobatmy) of characters.

Parallelism in the development life forms species of the genus Kudrjaschevia (Lamiaceae): ontogenesis, architectural analysis

V. A. Cheryomushkina1, A. Yu. Astashenkov1, D. S. Saidov2
1Central Siberian Botanical Garden of SB RAS, Novosibirsk, Russia
2Institute of Botany, Physiology and Genetics of Plants of AS RT, Dushanbe, Tajikistan
Keywords: архитектура растений, Kudrjaschevia, онтогенез, жизненные формы, architecture of plant, ontogenesis, life form

Abstract >>
The study is aimed at studying the herb life forms of closely related narrowly localized endemic species of the genus Kudrjaschevia in Central Asia. A detailed description of life forms will make it possible to identify morphological rearrangements of plants associated with the adaptation of species to certain ecological conditions. Our results will expand our understanding of the main directions in the evolution of morphs, which is practically not studied in plants of the Central Asian flora. In the process of somatic evolution, individuals of this group adapted to various specific habitats: rocks - K. nadinae , K. grubovii , K. pojarkoviae ; mountain scree - K. allotricha ; rocky outcrops of the mountain slopes - K. korshinskyi . Using an architectural approach to the description of plants, it was found that various ecotopic and ecological-coenotic conditions did not affect the nature of the development of life forms of related taxa. Parallelism in the development of life forms in the studied species is manifested in the passage of the same phases of morphogenesis: primary shoot, main axis, primary bush (clone) and in the preservation of a monocentic biomorph. In ontogenesis in individuals of closely related species, similar reiteration of axes is formed. Each axis characterized as an annually dying branching / non-branching with radial symmetry, having opposite phyllotaxis, the apical meristem of which is realized in inflorescence. The long-term basis of plants in all representatives of Kudrjaschevia was built as a result of sympodial sequential articulation of the short basal parts of the axes (residues) according to the monochasial / dichasial types. Despite the difference in the aerial parts of the studied rock ( K. nadinae , K. grubovii , K. pojarkoviae ) and mountain slope species ( K. allotricha , K. korshinskyi ), the shoot formation and the common long-term basis of the individuals are the same. The diversity of the aerial parts of the axes reflects the adaptation of plants to specific ecological conditions, while maintaining a genetically fixed, general development program.

Modification of the architecture of the dwarf subshrub Thymus seravschanicus (Lamiaceae) in Republic of Tajikistan

E. B. Talovskaya
Central Siberian Botanical Garden of SB RAS, Novosibirsk, Russia
Keywords: адаптация, архитектурная единица, полукустарничек, Thymus seravshanicus, Республика Таджикистан, adaptation, architectural unit, dwarf subshrub, Thymus seravshanicus, Republic of Tajikistan

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Using an architectural approach - one of the modern methods of detailed study of the structure of plants, the life form of the dwarf subshrub Thymus seravshanicus is characterized. The Species is widely distributed in different ecotopes in the Republic of Tajikistan (Hissar, Zeravshan, Hazretishokh ridges). For the first time in the structure of individuals, an architectural unit (AU) was identified and characterized, which consists of the main compound skeletal axis (n order), compound skeletal axes of the 1st (n+1) order, branching shoots and ephemeral shoots. Depending on the position of the main (n order) compound skeletal axis in space, a variety of AU was revealed: orthotropic, ascending, and mixed type. The features of their development are characterized. It is shown that in different ecological and cenotic habitat conditions, the structure of individuals is formed by repeated repetition of three or one architectural unit at the same time. In the mixed-grass steppe (black forest belt) and mixed-grass meadow (subalpine belt) communities, the structure of individuals is formed by a combination and repeated repetition of all AU. In areas with a lack of free substrate, the development of individuals begins with the formation of orthotropic AE, a dense shrub is formed; in areas of free substrate, the development of individuals begins with the formation of an ascending AE, a clump is formed. In the steppe community (juniper belt), the structure of T. seravshanicus individuals consists only of ascending AE and a clone is formed. The polyvariance of shoots development was revealed: 1) in the conditions of the black forest belt, intensive branching is characteristic for individuals, the duration of monopodial growth of shoots is up to 3 years; 2) in the conditions of the subalpine belt - rosette shoots, the duration of their monopodial growth up to 6 years; 3) in the conditions of the juniper belt in the structure of shoots there are no short metamers, the duration of monopodial growth up to 2 years. The identified features are mechanisms of adaptation of T. seravshanicus to growing conditions and do not lead to changes in the life form of the dwarf subshrub. The obtained data expand the understanding of the structure of dwarf subshrub, their morphological plasticity and serve as a basis for modeling the ways of morphological transformation of dwarf subshrub in different habitats.

Role of latent buds in crown architecture in coniferous and deciduous trees of the temperate zone

M. V. Kostina1, N. S. Barabanshchikova1, O. I. Yasinskaya2
1Moscow Pedagogical State University, Moscow, Russia
2Tsytsin Main Moscow Botanical Garden of RAS, Moscow, Russia
Keywords: деревья, крона, архитектурная модель, скелетные оси, ветви, реитерационные комплексы, спящие почки, trees, crown, architectural model, skeletal axis, branches, reiterated complexes, latent buds

Abstract >>
The study was aimed to investigate the latent buds localization and the characteristics of the structures arising from the buds in conifers and deciduous trees in response to age-related changes and various growing conditions. The crown analysis was performed using architectural models and reiteration concept. It was found that in deciduous trees, the latent buds were usually at the bases of annual growth, and in conifers the latent buds were at the bases of annual growth, in the bud scales axils, or along the entire length of the shoot in the needle axils and in the shoot’s distal part. It was revealed that the short shoots’ terminal buds could function as latent buds. In deciduous trees, the skeletal axes’ natural aging is the endogenous factor of shoots’ development from latent buds. In evergreen conifers it is the plant’s response to the needles’ aging. This leads to the growth induction of latent buds and to the acropetal appearance of the buds on the mother axis in conifers. In deciduous trees, the reiterated complexes form in response to the axis development cycle completion, which causes the initiation of latent buds in the centripetal direction. In conifers, the reiterated complexes look like bundles: such structures form due to the latent buds’ easy initiation, which leads to the formation of the other orders’ shoots. In deciduous trees, the reiterated complexes are characterized by prolonged growth, acrotonic branching mode and orthotropic growth direction. In deciduous trees, the crown of the old generative and senile plants almost entirely consists of reiterated complexes. It has been found that conifers have a different degree of latent buds participation in the crown formation, which can vary significantly within the same genus. In Picea abies (L.) Karst., the reiterated complexes replace small twigs. In some other species (for example, Picea schrenkiana Fisch. et E. Mey., Abies sibirica Ledeb., Pinus sibirica Du Tour.), the reiterated complexes can also replace the extending from the trunk skeletal branches. In Pinus sylvestris L., no secondary crown is formed.

Effects of lighting and gender differences on the architecture of the boreal species of Salix L

O. I. Nedoseko
Arzamas Branch of Lobachevsky University, Arzamas, Russia
Keywords: Salix, жизненные формы, архитектурные модули, гендерные отличия, life forms, arc

Abstract >>
The work proposes an algorithm of analysis of structural and functional organization of crowns of Salix species. It can be used to study the architecture of various species of trees and shrubs. The boreal species of Salix are insufficiently investigated from the perspective of structural and biological diversity. The study of plant organs based on the concept of “architectural models” is particularly relevant. On the example of three-year escape systems (TYTS) we first studied the architecture of crowns of Salix species. Depending on gender differences and lighting conditions a new technique to study the architectural modules of Salix boreal species has been developed. From the 16 species studied 11 life forms have been revealed. They refer to two major categories: trees and shrubs. For some species, stannic life forms have also been described. 7 architectural modules have been identified: 3 modules in trees and tall shrubs, 2 modules in medium-sized shrubs, and 2 modules in low shrubs. Life forms of individuals of Salix sub-species were formed on the basis of three architectural modules, and Vetrix sub-family are based on six architectural modules. We have proved that female individuals of Salix species (other than S. acutifolia ) have a greater variety of TYTS types compared to male ones (assimilating “year-long” shoots as a part of TYTS in female individuals are 1.3 to 1.7 times larger than in male ones. It provides them with a larger assimilating surface. The crowns of female individuals have been found to be more branched. Year-long shoots developing from sleeping buds are found in low shrubs 3-6 times more often than in trees. Different parts of the crown of trees and high shrubs species differ in the ratio of year-long assimilating shoots: the largest number of them are in the middle and upper parts of the crown, and the smallest are - in the lower part.

Architectural analysis of representatives of the order Celastrales: the structure and rhythm of development of the shoots in connection with adaptations of species to different environmental conditions

I. A. Savinov
Moscow State University of Food Production, Moscow, Russia
Keywords: архитектурные модели, модели побегообразования, побеговые системы, побеги, аксиллярные комплексы, порядок Celastrales, адаптации, экология, architectural models, models of shoot formation, shoot systems, shoots, axillary complexes, Celastrales, adaptations, ecology

Abstract >>
In despite of the considerable interest of many researchers in the Celastrales order, which is largely due to the great diversity of structural characters and almost cosmopolitan distribution (with the exception of some arid regions), a comprehensive biomorphological analysis of its representatives has not yet been carried out. The aim of this work is to analyze the structural and rhythmological characters of the shoots and shoot systems (vegetative and reproductive) of representatives of the order Celastrales, the volume of which is taken according to the latest data of molecular phylogenetics [APG IV]. The main working tool for this analysis was the concept of architectural models. Among the studied representatives of the order, the following architectural models were identified [according to the terminology of Hallé et al., 1978]: Rauh, Attims, Champagnat, Mangenot, Scarrone, Stone. The shoot systems are of three types: 1) polycarpic polycyclic with monopodial or acrosympodial growth and intercalary or pseudo-terminal inflorescences, with orthotropic, plagiotropic and / or mixed orientation of shoots in space, according to the rhythm of development - silleptic and proleptic shoots; 2) monocarpic mono-and dicyclic, terminal inflorescences; 3) monocarpic monocyclic - once blooming, with sympodial growth and terminal inflorescences, with elongated or shortened shoots. In all these cases, the nature of the blooming of flowers is basipetal. For grassy and semi-woody representatives ( Parnassia , Stackhousioideae), they additionally used shoot formation models identified by Russian biomorphologists [Serebryakova, 1981]: monopodic rosette (generative shoots are monocarpic monocyclic, flowers single, terminal) and sympodial semi-rosette (generative shoots are monocarpic monocyclic, but with acropetal blooming of flowers in inflorescence). For the taxon as a whole, the formation of axillary complexes is very usually, which are very different in structure and rhythm of development, that allows to successfully use these characters in the characterization of individual species and in establishing evolutionary trends. Thus, the structural diversity of shoot systems, which is the basis for the formation of different biomorphs, has been revealed for representatives of the Celastrales order. A high degree of ecological and morphological plasticity is manifested in individual and intraspecific shoot polymorphism, the dependence of the ratio of different types of shoots from environmental conditions, the possibility of changing the model during ontogenesis, the formation of transitional models (in Celastrus , Tripterygium ). The possibility of rapid transformation of the all shoot system under extreme environmental conditions was shown using the example of the genus Parnassia and the subfamily Stackhousioideae.

The characters of a biennial shoot system as a unit for modeling the crown development of ulmus glabra huds

I. S. Antonova1, V. A. Bart1,2
1Saint Petersburg State University, St. Petersburg, Russia
2Almazov National Medical Research Centre, St. Petersburg, Russia
Keywords: побег, зональность побега, экспоненциальная модель, многомерный дисперсионный анализ, пространственно-временные единицы строения кроны, двулетняя побеговая система, shoot zonality, exponential model, multivariate analysis of variance, spatio-temporal units of the crown structure, biennial shoot system

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The article presents the results of a study of the structural organization of the crown of Ulmus glabra . Knowledge of the types of shoot systems that regularly change in the tree crown in ontogenesis of an individual is of theoretical and practical value. In our studies, we used the architectural (modular) approach, which allowed us to describe the spatio-temporal program of tree crown development. The results obtained allowed us to determine that the main structural unit of the crown, resistant to changes in climatic factors, is a biennial shoot system. The choice of this unit is due to the fact that the zonality of the maternal shoot is only detected in the second year of its life. To identify the spatio-temporal structure of a biennial shoot system, the characters of shoots of the first and second years of life are compared. The biennial shoot systems of undergrowth plants of U. glabra were studied in the forest-steppe oak forest of the Belogorye reserve. 100 individuals of the same ontogenetic state were studied. In the studied trees, shoot systems were differentiated into large (“growth”) and small (“main”) by having the same location in the crown. Based on the cross-correlation function, which depends on the number of internodes on the maternal shoot, a regression model of the distribution of side shoot lengths was constructed. It was shown that, when numbering the internodes on the maternal shoot from the top, the dependence of the lengths of the side shoots of the “growth” and “main” biennial shoot system of U. glabra on the number of internodes is consistent with the exponential model. It is established that the development of a biennial shoot system is associated with the conditions of the irradiation and position in crown. Comparison of the two samples allowed introducing the shift parameter into the model, which defines a special zone on the maternal shoot.