Wind-Thrown Trees: Storms or Management?

In order to determine if the distribution of lateral root strength is related to the shape of the system and the forces withstood, wood samples were taken from roots of various mature tree species and the strength tested. Root strength decreased along the root at different rates, depending on the type of root system present. Slightly tapered lateral roots in plate root systems were relatively stronger further away from the stem than the highly tapered laterals in heart and tap root systems. Wood strength in Norway spruce (plate system) was found to increase along the lateral roots before decreasing again. The increase in strength may coincide with the point of maximum bending of the root as the tree sways backwards and forwards in the wind. Strength was also found to increase on the underside of lateral roots in the plate systems of poplar. The undersides of these roots will experience high compressive stresses due to the weight of the tree pushing the root on to the hard bearing surface of the soil. External loading forces in plate root systems will be transmitted into the soil further away from the stem due to the lack of branches, therefore a high strength along the root will help resist mechanical stress. The high rate of branching near the stem, or large, rigid, main tap root, found in heart and tap root systems, respectively, allows a faster dissipation of forces nearer the stem, therefore a high investment in strength further along the root is not necessary.

<p>Research over the past several decades has shown that preferential flow is more the rule than the exception. However, our collective understanding of preferential flow processes has been limited by a lack of suitable methods to detect and visualize the initiation and evolution of non-uniform wetting at high spatial and temporal resolutions, particularly in real-world settings. In this study, we investigate water infiltration initiation by tree trunk and root systems. We carried out time-lapse ground penetrating radar (GPR) surveys in conjunction with a simulated stemflow event to provide evidence of root-induced preferential flow and generate a three-dimensional representation of the wetted zone.</p><p>We established a survey grid (3.5 m × 5 m, with a local slope of 10.3°), consisting of ten horizontal and thirteen vertical parallel survey lines with 0.5 m intervals between them. The horizontal lines were downslope-oriented. The grid was placed around a Quercus suber L. We collected a total of 46 (2 GPR surveys × 23 survey lines) radargrams using an IDS (Ingegneria Dei Sistemi S.p.A.) Ris Hi Mod v. 1.0 system with a 900-MHz antenna mounted on a GPR cart. Two grid GPR surveys were carried out before and after the artificial stemflow experiment. In the experiment, we applied 100 L of brilliant blue dye (E133) solution on the tree trunk. The stemflow volume of 100 L corresponded to 63.2 mm of incident precipitation, considering a crown projected area of 201 m<sup>2</sup> and a 1.3% conversion rate of rainfall to stemflow. Trench profiles were carefully excavated with hand tools to remove soil and detect both root location and size and areas of infiltration and preferential pathways on the soil profile.</p><p>The majority (84.4%) of artificially applied stemflow infiltrated into the soil, while the remaining 15.6% generated overland flow, which was collected by a small v-shaped plastic channel placed into a groove previously scraped on the downhill side of the tree. The 3D diagram clearly demarcated the dimension and shape of the wetted zone, thus providing evidence of root-induced preferential flow along coarse roots. The wetted zone extended downslope up to a horizontal distance of 3 m from the trunk and down to a depth of approximately 0.7 m. Put all together, this study shows the importance of accounting for plant and trees trunk and root systems when quantifying infiltration.</p>

Windstorms, bark beetle outbreaks and fires are important natural disturbances in coniferous forests worldwide. Wind-thrown trees promote biodiversity and restoration within production forests, but also cause large economic losses due to bark beetle infestation and accelerated fungal decomposition. Such damaged trees are often removed by salvage logging, which leads to decreased biodiversity and thus increasingly evokes discussions between economists and ecologists about appropriate strategies. To reveal the reasons behind species loss after salvage logging, we used a functional approach based on four habitat-related ecological traits and focused on saproxylic beetles. We predicted that salvage logging would decrease functional diversity (measured as effect sizes of mean pairwise distances using null models) as well as mean values of beetle body size, wood diameter niche and canopy cover niche, but would increase decay stage niche. As expected, salvage logging caused a decrease in species richness, but led to an increase in functional diversity by altering the species composition from habitat-filtered assemblages toward random assemblages. Even though salvage logging removes tree trunks, the most negative effects were found for small and heliophilous species and for species specialized on wood of small diameter. Our results suggested that salvage logging disrupts the natural assembly process on windthrown trees and that negative ecological impacts are caused more by microclimate alteration of the dead-wood objects than by loss of resource amount. These insights underline the power of functional approaches to detect ecosystem responses to anthropogenic disturbance and form a basis for management decisions in conservation. To mitigate negative effects on saproxylic beetle diversity after windthrows, we recommend preserving single windthrown trees or at least their tops with exposed branches during salvage logging. Such an extension of the green-tree retention approach to windthrown trees will preserve natural succession and associated communities of disturbed spruce forests.

The core regulators which are required for primary root growth and development also function in lateral root development or lateral root stem cell niche maintenance. The primary root systems and the lateral root systems are the two important root systems which are vital to the survival of plants. Though the molecular mechanism of the growth and development of both the primary root systems and the lateral root systems have been extensively studied individually in Arabidopsis, there are not so much evidence to show that if both root systems share common regulatory mechanisms. AP2 family transcription factors such as PLT1 (PLETHORA1) and PLT2, GRAS family transcription factors such as SCR (SCARECROW) and SHR (SHORT ROOT) and WUSCHEL-RELATED HOMEOBOX transcription factor WOX5 have been extensively studied and found to be essential for primary root growth and development. In this study, through the expression pattern analysis and mutant examinations, we found that these core regulators also function in lateral root development or lateral root stem cell niche maintenance.

Soil compaction is known to limit plant growth by reducing soil macroporosity and restricting gas, water, and root movement. Recent evidence from study sites across the United States and Canada, however, suggests that tree growth is not universally affected by soil compaction from forest harvesting practices. Our observational study examined rooting patterns in mixed conifer plantations in the central Sierra Nevada of California to determine whether tree roots use continuous pathways or channels to overcome the physical restrictions of compacted soil. Replicate plots were established 20–25 years earlier to compare compacted and uncompacted treatments after clearcut harvesting. Fine and lateral root counts were taken at multiple depths in soil trenches. Rooting was extensive in compacted plots despite consistently high soil strength values (~3 MPa). No differences in rooting patterns or in fine or lateral root numbers were detected between compaction treatments. The results indicated long-term tolerance by conifer roots to soil compaction without clear use of preferential channels or uncompacted zones.

ABSTRACTThe rice root system consists of three types of roots; main root, L-type and S-type lateral root (LR). These component roots are morphologically and histologically different, which is termed as heterorhizy. Root system hydraulic architecture is related to the unique features of component roots. We hypothesized that each component root contributes in different degrees to water uptake of the whole root system. Rice varieties IRAT 109 and Taichung 65 were grown in pots filled with soil under continuous waterlogged (CWL) and drought (CD) conditions until two weeks after heading. Morphology and histological structures of roots, which may regulate radial water movement, were compared among the three component roots. Moreover, hydraulic conductivity (Lpr) of the root system, which represents the water uptake ability, were measured with a pressure chamber. Based on a model that Lpr of the whole root system is a product of Lpr of each of the component roots and their surface areas, we found that the differences in Lpr between the two varieties and the plants grown under different soil water conditions for any of the component roots did not support the corresponding differences in the measured Lpr of the whole root system. In contrast, a significant and positive correlation was found between Lpr of the whole root system and the percentage of surface area of S-type LR but not for the other component roots. These results indicate S-type LR might have a higher contribution to Lpr of the whole root system than the other component roots.

This study is part of an international mission in the Amazon rainforest, involving researchers from the Federal University of Rio de Janeiro, Universities of Leeds, Linköping, British Columbia, and coordinated by Prof. Vincent Gauci and Dr. Sunitha Pangala. The primary objective is to reconcile top-down methane (CH₄) emission estimates, derived from remote sensing data over Amazon floodplains, with bottom-up measurements obtained from field studies. Previous satellite observations indicated a discrepancy of 20 million tons of CH₄ emitted annually, a significant gap that could not be fully explained by ground-based sources. This project aimed to resolve this difference by integrating both remote sensing satellites and field data using ABB’s Off-Axis Integrated Cavity Output Spectroscopy (OA-ICOS) to understand and bring to light the different dynamics involved in methane emissions in the Amazon rainforest.Reconciling top-down and bottom-up carbon budgets can be particularly challenging in specific ecosystems where topography complicates site access and sampling. Under such conditions, the availability of compact and rugged cavity-enhanced laser-based analyzers offering sub-ppb precision is invaluable for environmental scientists. ABB’s portable greenhouse gas ultraportable analyser, GLA132-GGA (47 cm × 35.56 cm × 17.78 cm) is capable of monitoring CH4 with a 1 s precision of 1.4 ppb, which can be improved to 0.2 ppb with 100 s averaging time. The analyser is based on OA-ICOS technology that combines high precision capabilities through enhanced cavity path length and robustness to mechanical vibrations, which is crucial to field applications. Scientists used semi-rigid custom chambers wrapped around the trunk of floodplain trees and connected them to the GLA132-GGA to measure individual CH4 emissions from 2357 trees in 13 floodplain sites.The findings provide a crucial link to reconcile the 20-million-ton discrepancy in Amazon's methane budget. Scaled estimates of methane flux emitted from floodplain trees align closely with the missing methane observed in previous satellite data. During the rainy season, when Amazon tree roots become submerged, trees have evolved specialized adaptations to enhance oxygen supply to their roots by enlarging pores in their stems. This adaptation inadvertently facilitates the release of methane, produced by microorganisms in the waterlogged soil, through the same pore openings. Floodplain trees thus function as natural chimneys, venting substantial quantities of methane into the atmosphere. These large emissions from floodplain trees play a pivotal role in closing the Amazon methane budget.Furthermore, a second campaign revealed that methane produced deep within the soil column can also escape to the atmosphere via tree roots, even when the water table is below the surface. Regression analysis demonstrated that, while methane emissions show negligible response to increased flood levels above the soil surface, there is a clear dependence of whole-tree methane emissions on the presence of submerged roots. This highlights the importance of floodplain trees in regulating methane fluxes across varying hydrological conditions, underscoring their significant role in the global methane cycle. 

An experimental study was performed to test the model of buttress development proposed by Claus Mattheck: that the high rates of growth occurring along the top of the junction between the trunk and the lateral roots are stimulated by the mechanical strains set up by wind forces which are concentrated in these regions. Strain gauges were attached around the base of the trunk and the lateral roots of young rainforest trees which were developing buttresses; strains were measured during simulated wind events when the trunk was bent over using a winch. Results supported Mattheck's theory: strains were concentrated along the top of developing buttresses and on the trunk above them – the places of maximum growth – and were negligible on the sides of the buttresses and at the base of the trunk where no growth was occurring. Buttress development seems to be controlled by the mechanical environment of the tree. Once the root system, in which lateral roots are attached to the subsoil well away from the trunk by sinker roots, has formed the growth of buttresses occurs automatically. This produces a mechanically efficient anchorage system in terms of the carbon investment. Trees which do not form buttresses instead transfer forces into the ground using a tap root or sinkers positioned close to the trunk.

Lateral roots (LRs) occupy a large part of the root system and play a central role in plant water and nutrient uptake. Monocot plants, such as rice, produce two types of LRs: the S-type (short and thin) and the L-type (long, thick, and capable of further branching). Because of the ability to produce higher-order branches, the L-type LR formation contributes to efficient root system expansion. Auxin plays a major role in regulating the root system development, but its involvement in developing different types of LRs is largely unknown. Here, we show that auxin distribution is involved in regulating LR diameter. Dynamin-related protein (DRP) genes were isolated as causative genes of the mutants with increased L-type LR number and diameter than wild-type (WT). In the drp mutants, reduced endocytic activity was detected in rice protoplast and LRs with a decreased OsPIN1b-GFP endocytosis in the protoplast. Analysis of auxin distribution using auxin-responsive promoter DR5 revealed the upregulated auxin signaling in L-type LR primordia (LRP) of the WT and the mutants. The application of polar auxin transport inhibitors enhanced the effect of exogenous auxin to increase LR diameter with upregulated auxin signaling in the basal part of LRP. Inducible repression of auxin signaling in the mOsIAA3-GR system suppressed the increase in LR diameter after root tip excision, suggesting a positive role of auxin signaling in LR diameter increase. A positive regulator of LR diameter, OsWOX10, was auxin-inducible and upregulated in the drp mutants more than the WT, and revealed as a potential target of ARF transcriptional activator. Therefore, auxin signaling upregulation in LRP, especially at the basal part, induces OsWOX10 expression, increasing LR diameter.

PDF HTML阅读 XML下载 导出引用 引用提醒 毛竹种群向针阔林扩张的根系形态可塑性 DOI: 10.5846/stxb201401190143 作者: 作者单位: 浙江农林大学浙江省森林生态系统碳循环与固碳减排重点实验室,浙江农林大学浙江省森林生态系统碳循环与固碳减排重点实验室,浙江农林大学浙江省森林生态系统碳循环与固碳减排重点实验室,浙江农林大学浙江省森林生态系统碳循环与固碳减排重点实验室,浙江农林大学浙江省森林生态系统碳循环与固碳减排重点实验室,浙江农林大学浙江省森林生态系统碳循环与固碳减排重点实验室,浙江农林大学浙江省森林生态系统碳循环与固碳减排重点实验室 作者简介: 通讯作者: 中图分类号: 基金项目: 国家自然科学基金项目(31170594;31270497);浙江省自然科学基金项目(LY14C160010);浙江农林大学科研发展基金资助项目(2009FR060);临安市科技发展计划项目(05067) The response of root morphological plasticity to the expansion of a population of Phyllostachys edulis into a mixed needle-and broad-leaved forest Author: Affiliation: Zhejiang Agriculture and Forestry University,Zhejiang Agriculture and Forestry University,,,,, Fund Project: 摘要 | 图/表 | 访问统计 | 参考文献 | 相似文献 | 引证文献 | 资源附件 | 文章评论 摘要:为了弄清毛竹(Phyllostachys edulis)向针阔林扩张过程中根系的形态可塑性反应,在浙江天目山自然保护区毛竹向针阔林扩张的典型过渡地带,连续区域上设置毛竹纯林、针阔-毛竹混交林(以下简称过渡林)、针阔林3种样地。用根钻法采集样地毛竹根系、针阔树根系并比对其生物量密度、细根比根长、相邻同级侧根节点距等形态特征参数变化。结果表明:随着毛竹的扩张程度增加,林内根系生物量密度增加;且与针阔树竞争过程中毛竹将更多的根系放置于表层;同时在水平方向上随离样株距离的增加未出现明显变化,而针阔树根系则随离样木距离的增加而逐渐减少;毛竹根系比根长明显增加,平均增幅15%;一、二级侧根节点距则均有所下降,毛竹侧根数量增多。这些结果表明毛竹种群可通过根系生物量密度、细根比根长、相邻同级侧根节点距等形态可塑性方式实现向周边森林扩张。 Abstract:Moso bamboo(Phyllostachys edulis), an economically important plant, is often cultivated not only for its delicious shoots and versatile culms, but also as an important biomass resource in southern China. However, with its robust growth and strong rhizomes, Moso bamboo populations have been expanding rapidly into adjacent forests. Root plasticity may enable the bamboo to efficiently forage for nutrients in heterogeneous soils. The ability of a Moso bamboo plant to respond to competition through root plasticity may be an important trait for the species, but little information is available on how root systems respond to belowground competition during colony expansion. To provide baseline information on the morphological plasticity of the response of Moso bamboo roots during population expansion, we continuously sampled three forest types. These were 1) Moso bamboo forest; 2) mixed bamboo, needle-and broad-leaved forest(or transition forest); and 3) mixed needle-and broad-leaved forest. This study was conducted in a typical transition zone in the TianMu Mountain Nature Reserve, Zhejiang Province, China, where there is ongoing bamboo expansion. Rhizomes and roots from Moso bamboo, and roots from mixed needle-and broad-leaved trees were collected by the soil core method to analyze root morphological parameters, including root biomass density, special root length, and internodes length of lateral roots of the same order. The results showed that(1) the root/rhizome system biomass of the three forest types could be ranked as Moso bamboo forest(Moso bamboo root/rhizome system biomass=1630.46 g/m2) > Moso bamboo in the transition forest(902.76 g/m2) > the mixed needle-and broad-leaved forest(the needle-and broad-leaved trees root system biomass=412.76 g/m2) > the needle-and broad-leaved trees root system biomass in the transition forest(272.36 g/m2). As Moso bamboo expanded into a mixed needle-and broad-leaved forest, Moso bamboo culm density increased gradually, with a corresponding increase in root biomass.(2) In a vertical direction, the root system biomass of Moso bamboo in the 0-10 cm soil layer increased from 44% of all root biomass in the Moso bamboo forest to 56% in the transition forest. Simultaneously, the root biomass of needle-and broad-leaved trees shifted from the upper to lower soil layers. The root system of Moso bamboo managed to place more biomass in upper soil layers when it encountered other trees, allowing it to gain a competitive advantage.(3) In the horizontal direction, the root biomass of Moso bamboo did not significantly decline with an increase in distance from the sample tree, while the root biomasses of needle-and broad-leaved trees decreased with increasing distance from the sample tree.(4) The special root length of Moso bamboo in the transition forest increased by an average of 15% compared to the Moso bamboo forest, while the internodes length of first and second lateral roots decreased. The number of bamboo lateral roots also increased. Moso bamboo, with its ability to increase the special root length and decrease the length of the lateral root internodes, would outcompete trees in the belowground environment. These findings indicate that the morphological plasticity of Moso bamboo, shown by its root biomass, special root length, and internode length of the same order lateral roots, easily results in the expansion of Moso bamboo populations into surrounding forest. 参考文献 相似文献 引证文献

Root systems play a pivotal role in water and nutrient uptake from soil. Lateral root (LR) growth is promoted to compensate for inhibited main root growth. Compensatory LR growth contributes to maintaining total root length (TRL) and hence water and nutrient uptake in compacted soils. However, it remains unclear how shoot and root phenotypic traits change during the compensatory growth and whether there are genotypic variations in compensatory root growth. This study analyzed shoot and root morphological traits of 20 rice genotypes, which includes mutants with altered root morphology, during the vegetative stage using a semihydroponic phenotyping system. The phenotyping experiment detected large variation in root and shoot traits among the 20 genotypes. Morphological changes induced by root cutting were analyzed in six selected genotypes with contrasting root system architecture. Root cutting significantly affected root distribution along vertical sections and among diameter classes. After root cutting, more roots distributed at shallower depth and thicker LRs developed. Furthermore, genotypes with deeper root growth without root cutting allocated more compensatory roots to deeper sections even after root cutting than the genotypes with shallower rooting. Due to the compensatory LR growth, root cutting did not significantly affect TRL, root dry weight (RDW), or shoot dry weight (SDW). To analyze the interaction between crown root (CR) number and compensatory root growth, we removed half of the newly emerged CRs in two genotypes. TRL of YRL38 increased at depth with CR number manipulation (CRM) regardless of root tip excision, which was attributed to an increase in specific root length (SRL), despite no change in RDW. Taken together, the tested rice genotypes exhibited compensatory root growth by changing root distribution at depth and in diameter classes. Reducing CR number promoted root development and compensatory growth by improving the efficiency of root development [root length (RL) per resource investment].

The root system is responsible for water and nutrients uptake from the soil, and therefore, its extension is basic for an efficient acquisition. The maize root system is formed by different types of roots, and the lateral root branching substantially increases the surface for nutrient uptake. Therefore, the regulation of lateral root formation is fundamental in the development of root functions. Root architecture is basically controlled by auxin and cytokinins, which antagonize in the formation of lateral roots (LR) along the primary root axis, with auxin, a stimulator, and cytokinins inhibitors of LR development. This interaction has been analyzed in several zones along the primary root where LRs in different developmental stages were located. The root has been divided into several zones, such as meristem, elongation zone, and mature zone, according to the developmental processes occurring in each one. As Arabidopsis root elongated more slowly than maize root, these zones are shorter, and its delimitation is more difficult. However, these zones have previously been delimitated clearly in maize, and therefore, they analyze the effect of exogenous hormones in several LR developmental stages. The inhibitory effect of cytokinin on lateral root formation was observed in already elongated primary root zones in which initial events to form new lateral roots are taking place. Contrarily, auxin increased LR formation in the primary root segments elongated in the presence of the hormone. The inhibitory effect of cytokinin was reversed by auxin in a concentration-dependent manner when both hormones were combined. However, auxin is unable to recover LR development in primary root zones that have been previously elongated only in the presence of cytokinin. This antagonistic auxin-cytokinin effect on LR development depended on the balance between both hormones, which controls the root system architecture and determines the formation of LR during the process of initiation.

The development of a plastic root system is essential for stable crop production under variable environments. Rice plants have two types of lateral roots (LRs): S-type (short and thin) and L-type (long, thick, and capable of further branching). LR types are determined at the primordium stage, with a larger primordium size in L-types than S-types. Despite the importance of LR types for rice adaptability to variable water conditions, molecular mechanisms underlying the primordium size control of LRs are unknown. Here, we show that two WUSCHEL-related homeobox (WOX) genes have opposing roles in controlling LR primordium (LRP) size in rice. Root tip excision on seminal roots induced L-type LR formation with wider primordia formed from an early developmental stage. QHB/OsWOX5 was isolated as a causative gene of a mutant that is defective in S-type LR formation but produces more L-type LRs than wild-type (WT) plants following root tip excision. A transcriptome analysis revealed that OsWOX10 is highly up-regulated in L-type LRPs. OsWOX10 overexpression in LRPs increased the LR diameter in an expression-dependent manner. Conversely, the mutation in OsWOX10 decreased the L-type LR diameter under mild drought conditions. The qhb mutants had higher OsWOX10 expression than WT after root tip excision. A yeast one-hybrid assay revealed that the transcriptional repressive activity of QHB was lost in qhb mutants. An electrophoresis mobility shift assay revealed that OsWOX10 is a potential target of QHB. These data suggest that QHB represses LR diameter increase, repressing OsWOX10 Our findings could help improve root system plasticity under variable environments.

A soil and root system sampling technique that accurately measures root distribution within the soil profile without causing excessive damage to experimental plots would improve the efficiency of root system research. A monolith mapping root sampling technique is described that combines the positive attributes of the soil monolith and profile wall methods with a less destructive hand tool sampling protocol. The objective of this field study was to compare monolith mapping with a standard monolith washing technique for the purpose of measuring root distribution in the upper 30 cm of the root zone. Root systems of maize (Zea mays L.) (V6 stage of development) from plots (Vienna loams oils; fine‐loamy, mixed Udic Haploborolls) treated with broadcast or banded P were sampled using the modified monolith method. The roots present in the soil monolith were mapped (monolith mapping method), and the maps used to represent two‐dimensional root distribution. The soil monolith was then grid‐sectioned into 5.1 by 5.1 by 7.6 cm rectangular blocks and washed to separate roots from the soil. Root length was then measured using a line‐intersect method (monolith washing method). Monolith mapping and monolith washing methods both detected differences in root system distribution in the soil profile. A significant linear relationship (r = 0.87) between root length density and root number as measured by the two methods allows conversion of the numerical data collected by the monolith mapping method into root length density values. We conclude that monolith mapping works well for determining the spatial differences in root distribution in the upper 30 cm of the root zone. The relative time requirements, accuracy, and less destructive nature of the technique (compared with the trench profile and framed monolith methods) result in more efficient collection of data on root system characteristics.

Genetic transformation of most dicotyledonous plants by Rhizobium rhizogenes (also known as Agrobacterium rhizogenes) results in production of composite plants plants consisting of wild-type shoot and transgenic root system. Composite plants are the suitable model for investigation of hormonal mechanisms related to development of the root system as regulatory links between the root system and the shoot maintains in such plants.
 In most plants initiation of lateral root primordia occurs above the elongation zone [1]. However, in cucurbits and some other species, including important cereal crop buckwheat (Fagopyrum esculentum Moench), lateral root primordia initiation and development occurs in the apical meristem of the parental root [2, 3].
 The phytohormone auxin is a key regulator of lateral root development. Fusions of auxin-responsive promoters and reporter genes can be used to study the role of auxin in the development of root system of non-model plants such as cucumber (Cucumis sativus L.) and buckwheat [4].
 The agrobacterium mediated transformation technique of cucurbits [5] has been adapted for buckwheat. R. rhizogenes strain R1000 was used in all transformations. Set of binary vectors based on pKGW-RR-MGW or pKGW-MGW was developed to study auxin response maxima (DR5::mNeonGreen) or auxin transport (fusions of genes encoding auxin efflux proteins PIN and mNeonGreen).
 Pattern of auxin response maxima was similar in both species and included quiescent center and initial cells, columella, xylem cell files and lateral root primordia on all stages of development. Members of CsPIN1 (CsPINb and CsSoPIN1) group contributed unequally in generation of auxin maximum required for lateral root primordium initiation.
 The research was supported by the RFBR grant 20-016-00233-a.