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Alpine and Arctic treelines are assumed to be shifting toward higher latitudes and altitudes as a consequence of climate warming. Here, we compared the survival and growth of 1264 silver birch (Betula pendula Roth.) trees representing nine half-sib families. The trees were planted in two arboreta situated in distinct altitudinal environments in northern Finland in 1976 and 1977. The arboreta were located 9 km from each other and approximately 60 km north from the species' most northern natural growth site at that time. They were fenced to prevent vertebrate grazing, which is known to be among the most important factors limiting the expansion and regeneration of forests in European treeline ecotones. Overall, 90% and 81% of the trees were alive five and 40 years after planting in the two arboreta, respectively. Survival of trees varied among the half-sib families, especially in Arboretum 1, situated in a lower altitudinal environment characterized by soils with lower levels of nutrients, a longer growing season, and harsher winter temperatures. Trees were distinctively bigger in Arboretum 2: 50% taller (6.2 m vs. 4.4 m) and 68% thicker (9.5 cm vs. 5.6 cm) compared to trees in Arboretum 1. Furthermore, the performance of half-sib families varied depending on the garden they were grown in. These results demonstrate that the acclimation capacity of B. pendula allows its distribution to expand north from the present range; however, local abiotic environmental conditions (soil fertility and winter temperatures) and other selection pressures (herbivory) are likely to affect the genetic structure and growth of B. pendula populations., Competing Interests: Declarations. Competing interests: The authors declare no competing interests., (© 2025. The Author(s).)

Scenarios for future climate predict an increase in precipitation amounts and frequency of rain events, resulting in higher air humidity and soil moisture at high latitudes, including in northern Europe. We analysed the effects of artificially elevated environmental humidity (air relative humidity and soil moisture) on leaf gas exchange, water relations, growth and phenology of silver birch (Betula pendula) trees growing at the free air humidity manipulation experimental site situated in the hemiboreal vegetation zone, in eastern Estonia, with no occurring water deficit to the trees. The environmental humidity manipulation did not significantly affect the water relations traits but did affect some leaf gas exchange parameters, growth and phenology of the trees. Elevated air humidity (H) did not influence photosynthetic capacity and stomatal conductance, while the trees exhibited higher stomatal sensitivity to leaf-to-air vapour pressure difference compared with the trees at ambient conditions (C) or at elevated soil moisture (I). H trees demonstrated reduced height growth and foliage biomass, increased allocation to stem radial growth and prolonged leaf retention in autumn compared with the C trees. Increased air humidity supports longer leaf retention and growth period, but this does not translate into increased growth parameters at the tree level. The changes in tree growth in response to increasing atmospheric humidity could plausibly be explained by (i) retardation of foliage development and (ii) changes in resource allocation, causing a shift in the ratio of photosynthetic to non-photosynthetic tissues in favour of the latter. Under high atmospheric evaporative demand, higher stomatal sensitivity in H trees induces faster stomatal closure, which may result in carbon starvation. A future rise in atmospheric humidity at high latitudes may lead to reduced tree growth and forest productivity, in contrast to the predicted future of forests., (© The Author(s) 2024. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

Keymessage The study revealed the major biological processes occurred at three developmental stages and identified candidate genes involved in primary vein development of birch plants. Vascular tissues usually mirror the surrounding leaf shape and its development plays a fundamental role in plant performance. However, the information of vascular development in birch trees, especially primary vein development, remains unclear. Therefore, we conducted the anatomical observation on primary veins from leaves at different development stages in Betula pendula 'Dalecarlica'. With the development of primary vein, dynamic changes in mechanical tissue thickness and primary vein diameter were consistent with each other, and the sum of phloem, xylem and cambium thickness was significantly varied. Transcriptome analysis indicated that primary vein development could be divided into three stages, namely Stage I, II and III, which were in aggreement with anatomical observation. Expression of marker genes associated with vascular tissues revealed that pro-vasculature development occurred at Stage I and II, and phloem development occurred at Stage III. GO enrichment analysis of differentially expressed genes (DEGs) showed that shared DEGs at Stage II were mainly engaged in cell division and cell cycle, and shared DEGs at Stage III were mainly engaged in phosphorylation. Decreased cell division and cell cycle as well as activation of lignin biosynthesis might contribute to primary vein development. Combining phenotypic traits, we performed weighted gene co-expression network analysis and identified a cytochrome P450 84A (CYP84A) family gene (BpF5H1). Based on analyses of gene families, expression patterns and yeast-two hybrid assay results, we proposed a potential electron transfer pathway involving BpF5H1 and three cytochrome b 5 proteins during primary vein development in B. pendula 'Dalecarlica'. These results could shed some light on which biological processes occurred during primary vein formation and provide some valuable clues for vascular morphogenesis in woody plants., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024. Published by Elsevier B.V.)

Polyploid plants typically exhibit phenotypes distinct from diploids. Understanding the mechanism underlying plant polyploid phenotype variation is a critical prerequisite for effectively utilizing polyploid resources. In this study, we induced and obtained autotetraploid birch along with its diploid parent. Comparative analysis revealed significant differences in the morphology of leaves and stems between them. Transcriptome analysis indicated that 3.86% of genes showed significant differential expression between diploids and tetraploids. The genes significantly downregulated in tetraploids were primarily associated with functional terms related to hormone regulation, plant development, and morphogenesis. Notably, a group of genes involved in brassinosteroid (BR) biosynthesis was downregulated in tetraploids, and the level of active BRs in tetraploids was significantly lower than that in diploids. A Cas9/gRNA gene editing method was used to perform functional deletion mutations on BpCYP90C1. The knockout of BpCYP90C1 resulted in limited biosynthesis of 6-deoxoCS and CS in plants. Compared with wild-type (WT) plants, the bpcyp90c1 mutants exhibited a significant increase in leaf epidermal cell and a decrease in the number of leaf epidermal cells. The bpcyp90c1 mutants had curled leaves with obvious serrated edges and cordate base. Furthermore, the height and internode spacing of the bpcyp90c1 mutants were shorter. These phenotypic variations were similar to those of tetraploid birch. This indicates that the decrease in active BR levels was an important factor affecting the variation of tetraploid leaves and stems. Our research provides important insights into the molecular mechanisms of phenotypic variation in autopolyploid plants., (© 2025 Scandinavian Plant Physiology Society.)

Expansins (EXPs) are unique plant cell wall proteins with the ability to induce cell wall expansion and play potential roles in xylem development. In the present study, a total of 25 BpEXP genes were identified in Betula platyphylla. Results of bioinformatics analysis described that BpEXP gene family was highly conserved in the process of evolution. All these genes were clustered into four groups, EXPA (Expansin A), EXPB (Expansin B), EXLA (Expansin-like A) and EXLB (Expansin-like B), according to phylogenetic analysis and BpEXPA1 was highly homologous to PttEXP1 and PttEXP2. The results of RT-qPCR showed that BpEXPA1 was expressed higher in stems and preferentially expressed in the first internodes, followed by apical buds and the third internodes, promoter expression analysis with GUS assay demonstrated that it was expressed in developing xylem, suggesting that BpEXPA1 might be involved in the development of the primary stems of birch. Overexpression of BpEXPA1 can promote cortex cell expansion and then enlarge the cortex cell area and layer, however inhibit the secondary cell wall deposition and result in the thinner cell wall and larger lumens of xylem fiber in transgenic plants. This study will provide information for investigating the regulation mechanism of BpEXP family genes and gene resources for birch genetics improvement., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier GmbH. All rights reserved.)

Background: Salt stress is one of the major environmental factors affecting plant growth and productivity. BRI1-EMS suppressor 1/brassinazole-resistant 1 ((BES1/BZR1) plays an important role in responding to abiotic stress in plants. Although the impacts of BES1/BZR1 on plant growth and resistance have been documented, the potential mechanisms are not fully elucidated in Betula platyphylla. This work contributes to the understanding of how BES1/BZR1 promotes stress tolerance in woody plants., Results: Six BES1/BZR1 family members were identified from Betula platyphylla. Cis-element analysis showed that the promoters of six genes were rich in ABA-responsive element (ABRE), MYB and MBS cis-acting elements, which are reported to be involved in abiotic stress responses. Quantitative real-time polymerase chain reaction (qRT-PCR) analysis indicated that BpBZR1-6 (BPChr10G06000) could be induced by salt stress, ABA and BRs. BpBZR1-6 was localized in the nucleus and had transactivation activity. Ectopic expression of BpBZR1-6 enhanced Arabidopsis tolerance and decreased abscisic acid (ABA) sensitivity under salt treatment. Specifically, the seed germination rate, root length, fresh weight and chlorophyll content were significantly higher in BpBZR1-6-overexpressing (OE) transgenic plants than in wild-type (WT) plants after salt stress (P < 0.05). Additionally, BpBZR1-6 overexpression showed enhanced the reactive oxygen species (ROS) scavenging capability under salt stress, including increasing the activities of antioxidant enzyme, resulting in a decrease in O 2 - and H 2 O 2 accumulation, and reducing malondialdehyde (MDA) content. Meanwhile, the expression levels of six antioxidant enzyme genes were higher in OE plants than in WT plants after stress., Conclusion: BpBZR1-6 overexpression enhanced the salt tolerance of transgenic plants by modulating antioxidant enzyme gene expression and ROS scavenging, which may provide underlying strategy for breeding of salt-tolerant plants., Competing Interests: Declarations. Ethics approval and consent to participate: Birch (Betula Platyphylla) and Poplar (Populus trichocarpa) specimens were obtained from Northeast Forestry University, Harbin, China. The voucher specimens were stored in Northeast Forestry University. The plants were identified by Yao Chi. All plant materials used in this study were owned by the authors and/or no permissions are required. All the plants complied with national guidelines and legislation, and did not deposit in a public available herbarium and involve in any endangered or protected species. Consent for publication: Not applicable. Competing interests: The authors declare no competing interests., (© 2024. The Author(s).)

Background: Amalgamation of metal-tolerant plant growth promoting rhizobacteria (PGPR) with biochar is a promising direction for the development of chemical-free biofertilizers that can mitigate environmental risks, enhance crop productivity and their biological value. The main objective of the work includes the evaluation of the influence of prepared bacterial biofertilizer (BF) on biometric growth parameters as well as physiological and biochemical characteristics of rapeseed ( Brassica napus L.) at copper action., Methods: The prepared BF was based on novel metal tolerant strain of PGPR Buttiauxella sp. EA20 isolated from the rhizosphere of orchid Epipactis atrorubens and birch wood biochar (BC). The pot-scale experiments included six treatments: peat-containing control substrate (CS); CS + 200Cu (200 mg Cu/kg of soil); CS + 5% BC ( v/v ); CS + 5% BC + 200Cu; CS + 5% BF ( v/v ); CS + 5% BF + 200Cu., Results: Single Cu treatment caused the decrease in rapeseed leaf area, shoot and root length, fresh and dry biomass, as well as an increase in water saturation deficit, possibly due to damage of cell membranes by lipid peroxidation. Addition of BF or BC alone mitigated these harmful effects of copper. Application of BF, regardless of Cu addition, increased the rapeseed leaf area (1.6 times on average), plant fresh and dry biomass (2.5 times on average), and photosynthetic pigment content (1.8 times on average). In addition, BF treatment along with Cu enhanced the antioxidant activity of B. napus due to the accumulation of non-enzymatic antioxidants such as carotenoids, free proline and soluble phenolic compounds, including flavonoids. Moreover, plant enrichment with copper and essential macronutrients such as nitrogen, phosphorus and potassium was observed., Conclusions: The study concludes that application of complex biofertilizer based on metal tolerant PGPR strain Buttiauxella sp. EA20 and birch wood biochar mitigated the harmful effects of copper, enhanced the rapeseed growth and increased its biological value. Future perspective includes evaluation of the potential for using the resulting biofertilizer to improve the growth and biofortification of other crop species., (© 2024 The Author(s). Published by IMR Press.)

The process-based xylem formation model is an important tool for understanding the radial growth process of trees and its influencing factors. While numerous xylogenesis models for conifers have been developed, there is a lack of models available for non-coniferous trees. In this study, we present a process-based model designed for xylem formation and ring growth in broad-leaved trees, which we call the Broad-leaved Tree-Ring (BTR) model. Climate factors, including daylength, air temperature, soil moisture and vapor pressure deficit, drive daily xylem cell production (fibers and vessels) and growth (enlargement, wall deposition). The model calculates the total cell area in the simulated zone to determine the annual ring width. The results demonstrate that the BTR model can basically simulate inter-annual variation in ring width and intra-annual changes in vessel and fiber cell formation in Fraxinus mandshurica (ring-porous) and Betula platyphylla (diffuse-porous). The BTR model is a potential tool for understanding how different trees form wood and how climate change influences this process., (© The Author(s) 2024. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

Plant metabolism faces a challenge of investing enough enzymatic capacity to a pathway without overinvestment. As it takes energy and resources to build, operate, and maintain enzymes, there are benefits and drawbacks to accurately matching capacity to the pathway influx. The relationship between functional capacity and physiological load could be explained through symmorphosis, which would quantitatively match enzymatic capacity to pathway influx. Alternatively, plants could maintain excess enzymatic capacity to manage unpredictable pathway influx. In this study, we use photorespiration as a case study to investigate these two hypotheses in Betula papyrifera. This involves altering photorespiratory influx by manipulating the growth environment, via changes in CO 2 concentration and temperature, to determine how photorespiratory capacity acclimates to environmental treatments. Surprisingly, the results from these measurements indicate that there is no plasticity in photorespiratory capacity in B. papyrifera, and that a fixed capacity is maintained under each growth condition. The fixed capacity is likely due to the existence of reserve capacity in the pathway that manages unpredictable photorespiratory influx in dynamic environments. Additionally, we found that B. papyrifera had a constant net carbon assimilation under each growth condition due to an adjustment of functional rubisco activity driven by changes in activation state. These results provide insight into the acclimation ability and limitations of B. papyrifera to future climate scenarios currently predicted in the next century., (© 2024. The Author(s).)

Pollution is an integral part of global environmental change, yet the combined and interactive effects of pollution and climate on terrestrial ecosystems remain inadequately understood. This study aims to explore whether pollution alters the impacts of ambient air temperature on the population dynamics of herbivorous insects. Between 1995 and 2005, we studied populations of two closely related moths, Eriocrania semipurpurella and E. sangii, at eight sites located 1 to 64 km from a large copper‑nickel smelter in Monchegorsk, Russia. We found that pollution and temperature influence the performance of Eriocrania larvae mining in the leaves of mountain birch, Betula pubescens var. pumila, through multiple pathways. This is evident from the unconsistent changes observed in larval and frass weight, mine area, and leaf size. We found increases in both leaf quality and larval weight with decreasing pollution levels at both spatial and temporal scales and attributed these to the impact of sulphur dioxide, rather than trace elements (nickel and copper). The quality of birch leaves increased with spring (May) temperatures, enabling Eriocrania larvae to achieve greater weight while consuming less biomass. During the larval growth period (early June to early July), Eriocrania larvae increased their consumption with rising temperatures, presumably to compensate for increased metabolic expenses. Contrary to our expectations, the per capita rate of population change did not correlate with larval weight and did not vary along the pollution gradient. Nevertheless, we detected interactive effects of pollution and climate on the rate of population change. This rate decreased with rising winter temperatures in slightly polluted and unpolluted sites but remained unchanged in heavily polluted sites. We conclude that pollution disrupts mechanisms regulating the natural population dynamics of Eriocrania moths., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier B.V. All rights reserved.)

Functional traits regulate plant response to environmental changes, with consequences on population dynamics. However, how plant functional traits impact population dynamics, including growth, mortality, and recruitment, remains elusive in temperate forests across different successional stages. In this study, we compiled data on population dynamics and eight functional traits, encompassing hydraulic, wood, and leaf traits, from 35 species commonly found in a secondary poplar-birch forest and a broad-leaved Korean pine forest in Northeast China. We quantified the intrinsic relationships between plant population dynamics and assessed how plant functional traits influenced these dynamics. The results demonstrated a gradual increase in the correlation among population dynamics as forest succession progressed. In the secondary forest, tree growth rate and mortality rate were negatively correlated, while growth-death rate and growth-recruitment rate were not related. Conversely, in the broad-leaved Korean pine forest, there was a significant negative correlation between tree growth rate and mortality rate, as well as between growth rate and recruitment rate, while tree mortality rate positively correlated with recruitment rate. Additionally, functional traits effectively predicted population dynamics, but the predictive ability varied across successional stages. Functional traits, particularly xylem hydraulic traits ( e.g ., Huber value) and anatomical traits ( e.g ., mean xylem conduit diameter), were stronger predictors of tree growth, mortality, and recruitment rates at the late successional stage compared with the early stage. These findings indicated that population dynamics and functional traits exhibited strong regularity in the late successional stage of broad-leaved Korean pine forests.

Betula utilis exhibits intriguing characteristics and interactions with its environment and has specific adaptations that enable it to thrive in various water conditions. Drought has a prominent role in influencing the growth and development of vegetation, while temperature serves as a crucial determinant of species distribution in high-altitude environments. The investigation was centered on the eco-physiological dimension of B. utilis in areas near the treeline. Across different seasons, sites, and years, the most negative pre-dawn twig water potentials (Ψ PD ) and mid-day twig water potentials (Ψ MD ) were - 0.81 and - 1.24 MPa, respectively. The highest seasonal change (ΔΨ) in twig water potential (Ψ twig ) was in the post-monsoon season. Osmotic potential at full turgor (Ψπ 100 ) declined by - 0.66 MPa and osmotic potential at zero turgor (Ψπ 0 ) declined by - 1.07 MPa. The highest leaf conductance (gw) of 380.26 mmol m -2  s -1 was measured in the afternoon. During the initiation of flowering, Ψ PD of the twig was - 0.72 MPa and gradually rose to - 0.17 MPa by the end of the flowering period. This study provides key insight into the Ψ dynamics, leaf conductance, and phenology of B. utilis, highlighting its adaptation to changing environmental conditions and the need for effective management strategies to ensure the resilience and conservation of this Critically Endangered species., (© 2024. The Author(s).)

Climate change is raising concerns about how forests will respond to extreme droughts, heat waves and their co-occurrence. In this greenhouse study, we tested how carbon and water relations relate to seedling growth and mortality of northeastern US trees during and after extreme drought, warming, and combined drought and warming. We compared the response of our focal species red spruce (Picea rubens Sarg.) with a common associate (paper birch, Betula papyrifera Marsh.) and a species expected to increase abundance in this region with climate change (northern red oak, Quercus rubra L.). We tracked growth and mortality, photosynthesis and water use of 216 seedlings of these species through a treatment and a recovery year. Each red spruce seedling was planted in containers either alone or with another seedling to simulate potential competition, and the seedlings were exposed to combinations of drought (irrigated, 15-d 'short' or 30-d 'long') and temperature (ambient or 16 days at +3.5 °C daily maximum) treatments. We found dominant effects of the drought reducing photosynthesis, midday water potential, and growth of spruce and birch, but that oak showed considerable resistance to drought stress. The effects of planting seedlings together were moderate and likely due to competition for limited water. Despite high temperatures reducing photosynthesis for all species, the warming imposed in this study minorly impacted growth only for oak in the recovery year. Overall, we found that the diverse water-use strategies employed by the species in our study related to their growth and recovery following drought stress. This study provides physiological evidence to support the prediction that native species to this region like red spruce and paper birch are susceptible to future climate extremes that may favor other species like northern red oak, leading to potential impacts on tree community dynamics under climate change., (© The Author(s) 2024. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

Tropical montane evergreen broad-leaved forests cover the majority of forest areas and have high carbon storage in Xishuangbanna, southwest China. However, stem radial growth dynamics and their correlations with climate factors have never been analyzed in this forest type. By combining bi-weekly microcoring and high-resolution dendrometer measurements, we monitored xylogenesis and stem radius variations of the deciduous species Betula alnoides Buch.-Ham. ex D. Don and the evergreen species Schima wallichii (DC.) Korth. We analyzed the relationships between weekly climate variables prior to sampling and the enlarging zone width or wall-thickening zone width, as well as weekly radial increments and climate factors during two consecutive years (2020 to 2021) showing contrasting hydrothermal conditions in the pre-monsoon season. In the year 2020, which was characterized by a warmer and drier pre-monsoon season, the onset of xylogenesis and radial increments of B. alnoides and S. wallichii were delayed by three months and one month, respectively, compared with the year 2021. In 2020, xylem formation and radial increments were significantly reduced for B. alnoides, but not for S. wallichii. The thickness of enlarging zone and wall-thickening zone in S. wallichii were positively correlated with relative humidity, and minimum and mean air temperature, but were negatively correlated with vapor pressure deficit during 2020 to 2021. The radial increments of both species showed significant positive correlations with precipitation and relative humidity, and negative correlations with vapor pressure deficit and maximum air temperature during two years. Our findings reveal that drier pre-monsoon conditions strongly delay growth initiation and reduce stem radial growth, providing deep insights to understand tree growth and carbon sequestration potential in tropical forests under a predicted increase in frequent drought events., (© The Author(s) 2024. Published by Oxford University Press.)

The temperature sensitivities of photosynthesis and respiration remain a key uncertainty in predicting how forests will respond to climate warming. We grew seedlings of four temperate tree species, including Betula platyphylla, Fraxinus mandshurica, Juglans mandshurica and Tilia amurensis, at three temperature regimes (ambient, +2 °C, and +4 °C in daytime air temperature). We investigated net photosynthesis (Anet25), maximum rate of RuBP-carboxylation (Vcmax25) and RuBP-regeneration (Jmax25), stomatal conductance (gs25), mesophyll conductance (gm25), and leaf respiration (Rleaf) in dark (Rdark25) and in light (Rlight25) at 25 °C in all species. Additionally, we examined the temperature sensitivities of Anet, Vcmax, Jmax, Rdark and Rlight in F. mandshurica. Our findings showed that the warming-induced decreases in Anet25, Vcmax25 and Jmax25 were more prevalent in the late-successional species T. amurensis. Warming had negative impacts on gs25 in all species. Overall, Anet25 was positively correlated with Vcmax25 and Jmax25 across all growth temperatures. However, a positive correlation between Anet25 and gs25 was observed only under warming conditions, and gs25 was negatively associated with vapor pressure deficit. This implies that the vapor pressure deficit-induced decrease in gs25 was responsible for the decline in Anet25 at higher temperatures. The optimum temperature of Anet in F. mandshurica increased by 0.59 °C per 1.0 °C rise in growth temperature. While +2 °C elevated the thermal optima of Jmax, it did not affect the other temperature sensitivity parameters of Vcmax and Jmax. Rdark25 was not affected by warming in any species, and Rlight25 was stimulated in T. amurensis. The temperature response curves of Rdark and Rlight in F. mandshurica were not altered by warming, implying a lack of thermal acclimation. The ratios of Rdark25 and Rlight25 to Anet25 and Vcmax25 in T. amurensis increased with warming. These results suggest that Anet and Rleaf did not acclimate to warming synchronously in these temperate tree species., (© The Author(s) 2024. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

Polyploidization plays a crucial role in plant breeding and genetic improvement. Although the phenomenon of polyploidization affecting the area and number of plant epidermal pavement cells is well described, the underlying mechanism behind this phenomenon is still largely unknown. In this study, we found that the leaves of autotetraploid birch (Betula pendula) stopped cell division earlier and had a larger cell area. In addition, compared to diploids, tetraploids have a smaller stomatal density and fewer stomatal numbers. Genome-wide DNA methylation analysis revealed no significant difference in global DNA methylation levels between diploids and tetraploids. A total of 9154 differential methylation regions (DMRs) were identified between diploids and tetraploids, with CHH-type DMRs accounting for 91.73% of all types of DMRs. Further research has found that there are a total of 2105 differentially methylated genes (DMEGs) with CHH-type DMRs in birch. The GO functional enrichment results of DMEGs showed that differentially methylated genes were mainly involved in terms such as cellular process and metabolic process. The analysis of differentially methylated genes and differentially expressed genes suggests that hyper-methylation in the promoter region may inhibit the gene expression level of BpCYCD3;2 in tetraploids. To investigate the function of BpCYCD3;2 in birch, we obtained overexpression and repressed expression lines of BpCYCD3;2 through genetic transformation. The morphogenesis of both BpCYCD3;2-OE and BpCYCD3;2-RE lines was not affected. However, low expression of BpCYCD3;2 can lead to inhibition of cell division in leaves, and this inhibition of cell proliferation can be compensated for by an increase in cell size. Additionally, we found that the number and density of stomata in the BpCYCD3;2-RE lines were significantly reduced, consistent with the tetraploid. These data indicate that changes in cell division ability and stomatal changes in tetraploid birch can be partially attributed to low expression of the BpCYCD3;2 gene, which may be related to hyper-methylation in its promoter region. These results will provide new insights into the mechanism by which polyploidization affects plant development., Competing Interests: Declaration of Competing Interest The authors declare no conflicts of interest., (Copyright © 2024 Elsevier B.V. All rights reserved.)

The high prevalence of hay fever in Europe has raised concerns about the implications of climate change-induced higher temperatures on pollen production. Our study focuses on downy birch pollen production across Europe by analyzing 456 catkins during 2019-2021 in 37 International Phenological Gardens (IPG) spanning a large geographic gradient. As IPGs rely on genetically identical plants, we were able to reduce the effects of genetic variability. We studied the potential association with masting behavior and three model specifications based on mean and quantile regression to assess the impact of meteorology (e.g., temperature and precipitation) and atmospheric gases (e.g., ozone (O 3 ) and carbon-dioxide (CO 2 )) on pollen and catkin production, while controlling for tree age approximated by stem circumference. The results revealed a substantial geographic variability in mean pollen production, ranging from 1.9 to 2.5 million pollen grains per catkin. Regression analyses indicated that elevated average temperatures of the previous summer corresponded to increased pollen production, while higher O 3 levels led to a reduction. Additionally, catkins number was positively influenced by preceding summer's temperature and precipitation but negatively by O 3 levels. The investigation of quantile effects revealed that the impacts of mean temperature and O 3 levels from the previous summer varied throughout the conditional response distribution. We found that temperature predominantly affected trees characterized by a high pollen production. We therefore suggest that birches modulate their physiological processes to optimize pollen production under varying temperature regimes. In turn, O 3 levels negatively affected trees with pollen production levels exceeding the conditional median. We conclude that future temperature increase might exacerbate pollen production while other factors may modify (decrease in the case of O 3 and amplify for precipitation) this effect. Our comprehensive study sheds light on potential impacts of climate change on downy birch pollen production, which is crucial for birch reproduction and human health., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier Inc. All rights reserved.)

The effects of rising atmospheric CO2 concentrations (Ca) with climate warming on intrinsic water-use efficiency and radial growth in boreal forests are still poorly understood. We measured tree-ring cellulose δ13C, δ18O, and tree-ring width in Larix dahurica (larch) and Betula platyphylla (white birch), and analyzed their relationships with climate variables in a boreal permafrost region of northeast China over past 68 years covering a pre-warming period (1951-1984; base period) and a warm period (1985-2018; warm period). We found that white birch but not larch significantly increased their radial growth over the warm period. The increased intrinsic water-use efficiency in both species was mainly driven by elevated Ca but not climate warming. White birch but not larch showed significantly positive correlations between tree-ring δ13C, δ18O and summer maximum temperature as well as vapor pressure deficit in the warm period, suggesting a strong stomatal response in the broad-leaved birch to temperature changes. The climate warming-induced radial growth enhancement in white birch is primarily associated with a conservative water-use strategy. In contrast, larch exhibits a profligate water-use strategy. It implies an advantage for white birch over larch in the warming permafrost regions., (© The Author(s) 2024. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Adventitious root formation is a key step in vegetative propagation via cuttings. It is crucial for establishing birch plantations and preserve birch varieties. Although previous studies have highlighted role of WOX11 in controlling adventitious root formation, no such study has been conducted in birch. Understanding the mechanism of adventitious root formation is essential for improvement of rooting or survival rate using stem cuttings in birch. In this study, we cloned BpWOX11 and produced BpWOX11 overexpression (OE) transgenic lines using the Agrobacterium-mediated plant transformation. OE lines exhibited early initiated adventitious root formation, leading to increase the rooting rate of stem cuttings plants. RNA sequencing analysis revealed that OE lines induced the gene expression related to expansin and cell division pathway, as well as defense and stress response genes. These may be important factors for the BpWOX11 gene to promote adventitious root formation in birch cuttings. The results of this study will help to further understand the molecular mechanisms controlling the formation of adventitious roots in birch., (© 2023. The Author(s).)

Background: We studied UPBEAT1 (UPB1) which regulated superoxide radical / hydrogen peroxide ratio together with peroxidase (POD) activity and PAL genes expression under different ways of apical meristem development during the xylem structural elements' formation in unique woody plants B. pendula var. pendula with straight-grained wood and B. pendula var. carelica with figured wood. The differentiation process predominanced in straight-grained wood ( B. pendula var. pendula ) or proliferation - in the figured wood. The investigation was conducted in the radial row (cambial zone - differentiating xylem - mature xylem) during the active cambial growth period., Objective: The study aimed to study the xylogenesis processes occurring in the 16-year-old straight-grained silver birch ( Betula pendula Roth) and Karelian birch ( Betula pendula Roth var. carelica (Mercl.) Hämet-Ahti) with figured wood., Methods: Hydrogen peroxide and superoxide radical contents and peroxidase activity were determined spectrophotometrically. Gene expression for PAL family genes and the UPBEAT1 gene was assessed using qRT-PCR., Results: Principal component analysis has confirmed trees with straight-grained and figured wood to be different according to UPBEAT1-ROS-POD- PAL system functioning., Conclusion: The higher superoxide radical/hydrogen peroxide ratio in figured Karelian birch, along with UPBEAT1 transcription factor and PAL genes upregulation, distinguished it from straight-grained silver birch. This metabolic picture confirmed the shift of Karelian birch xylogenesis towards proliferation processes, accompanied by ROS and phenolic compounds' flow and POD activity., (Copyright© Bentham Science Publishers; For any queries, please email at epub@benthamscience.net.)

Background: The mechanisms that control the accumulation of woody biomass are of great interest to the study. Invertase and sucrose synthase are enzymes that are vital for distributing carbon in various biosynthetic pathways. Karelian birch ( Betula pendula var. carelica ) is a form of silver birch ( B. pendula Roth) and is characterized by disruption of the differentiation of cambium derivatives towards both the xylem and phloem, which leads to a change in the proportion of the conducting tissues' structural elements and the figured wood formation. We researched the expression profiles of genes encoding sucrose-cleaving enzymes ( CWINV and SUS gene families) and genes encoding CVIF protein, which is responsible for the post-translational regulation of the cell wall invertase activity., Objective: In our study, 16-year-old common silver birch ( Betula pendula var. pendula ) and Karelian birch were used for sampling non-figured and figured trunk section tissues, respectively. Samples were selected for the research based on the radial vector: non-conductive, conductive phloem, cambial zone - differentiating xylem - mature xylem., Methods: The enzyme's activity was investigated by biochemical methods. RT-PCR method was used to determine the level of gene expression. Anatomical and morphological methods were used to determine the stage of differentiation of xylem cambial derivatives., Results: Our research revealed a shift in the composition of xylem components in figured Karelian birch, characterized by increased parenchymatization and reduced vessel quantity. In all studied trunk tissues of Karelian birch, compared with common silver birch, an increase in the expression of the CWINV gene family and the SUS3 gene and a decrease in the expression of SUS4 were shown., Conclusion: Therefore, the increase in parenchymatization in figured Karelian birch is linked to a shift in sucrose metabolism towards the apoplastic pathway, indicated by a higher cell wall invertase activity and gene expression. The expression of the SUS4 gene correlates with the decrease in xylem increments and vessel proportion. The research findings will enhance our understanding of how sucrose breaking enzymes regulate secondary growth in woody plants and aid in developing practical timber cultivation methods., (Copyright© Bentham Science Publishers; For any queries, please email at epub@benthamscience.net.)

Silver birch trees ( Betula pendula Roth) are a pioneering species in post-industrial habitats, and have been associated with an expansive breeding strategy and low habitat requirements. We conducted ecophysiological and dendroclimatological studies to check whether there are any features of which the modification enables birch trees to colonise extreme habitats successfully. We characterised the efficiency of the photosynthetic apparatus, the gas exchange, the content of pigments in leaves, and the growth (leaf thickness and tree-ring width) of birch trees on a post-coal mine heap, a post-smelter heap, and a reference site. Birch growth was limited mainly by temperature and water availability during summer, and the leaves of the birch growing on post-industrial heaps were significantly thicker than the reference leaves. Moreover, birch trees growing on heaps were characterised by a significantly higher content of flavonols and anthocyanins in leaves and higher non-photochemical quenching. In addition, birches growing on the post-coal mine heap accumulated a concentration of Mn in their leaves, which is highly toxic for most plant species. Increasing the thickness of leaves, and the content of flavonols and anthocyanins, as well as efficient non-photochemical quenching seem to be important features that improve the colonization of extreme habitats by birches.

Water and nutrient are two critical factors that limit plant growth to spatial-temporal extents. Tree root foraging behavior has not received adequate attention in heterogeneous soil environments in temperate forest under drought pressure. In this study, birch (Betula platyphylla) and larch (Larix olgensis) seedlings were raised in pots in a split-root system with artificially heterogeneous soil environments to study the root foraging response to drought. Potted space was split into two halves where substrates were mixed with fertilizers in 67.5 mg nitrogen (N) plant-1 (N-P2O5-K2O, 14-13-13) to both halves as to create a homogeneous condition. Otherwise, a rate of 135 mg N plant-1 of fertilizers was delivered to a random half to create a heterogeneous condition. Half of seedlings were fully sub-irrigated every three days with the other half received the drought treatment by being watered every six days. Both birch and larch seedlings showed greater net shoot growth and biomass increment in well-watered condition, while root morphology was promoted by drought. Both species placed more fine roots with higher root N concentration in nutrient-enriched patches. In the heterogeneous pattern, birch showed a higher foraging precision assessed by biomass and greater foraging plasticity assessed in morphology and physiology. In contrast, larch seedlings had higher root N concentration in the well-watered condition. Neither species showed a significant response of N utilization to the heterogeneous pattern, but both used more N when water supply was improved. Overall, birch is better at acclimating to heterogeneous soil conditions, but its ability to seize N was lower than larch when drought was alleviated., Competing Interests: The authors declare no conflict of interest. The commercial affiliation of Harbin Gloria Pharmaceuticals Ltd. (Harbin city, postal code 150040, China) does not alter authors’ adherence to PLOS ONE policies on sharing data and materials.

Numerous long-term, free-air plant growth facilities currently explore vegetation responses to the ongoing climate change in northern latitudes. Open top chamber (OTC) experiments as well as the experimental set-ups with active warming focus on many facets of plant growth and performance, but information on morphological alterations of plant cells is still scarce. Here we compare the effects of in-situ warming on leaf epidermal cell expansion in dwarf birch, Betula nana in Finland, Greenland, and Poland. The localities of the three in-situ warming experiments represent contrasting regions of B. nana distribution, with the sites in Finland and Greenland representing the current main distribution in low and high Arctic, respectively, and the continental site in Poland as a B. nana relict Holocene microrefugium. We quantified the epidermal cell lateral expansion by microscopic analysis of B. nana leaf cuticles. The leaves were produced in paired experimental treatment plots with either artificial warming or ambient temperature. At all localities, the leaves were collected in two years at the end of the growing season to facilitate between-site and within-site comparison. The measured parameters included the epidermal cell area and circumference, and using these, the degree of cell wall undulation was calculated as an Undulation Index (UI). We found enhanced leaf epidermal cell expansion under experimental warming, except for the extremely low temperature Greenland site where no significant difference occurred between the treatments. These results demonstrate a strong response of leaf growth at individual cell level to growing season temperature, but also suggest that in harsh conditions other environmental factors may limit this response. Our results provide evidence of the relevance of climate warming for plant leaf maturation and underpin the importance of studies covering large geographical scales., Competing Interests: The authors have declared that no competing interests exist.

Curly birch (Betula pendula Roth. var. carelica [Merklin] Hämet-Ahti) is a disappearing representative of the Betula genus facing a regeneration failure in a large part of its natural distribution in Europe. The unique long-term study of clonal replications originating in heterogeneous environments enabled the evaluation of long-term survival and phenotypic stability of progenies in seed orchard to assess the conservation and commercial potential of heterovegetative propagation. Seventy-eight geographic sources (95 clone origins) representing the south distribution edge in East-Central Europe were analysed for species variation in survival, growth form, bark colour, and stem quality of parent trees and their vegetative progeny, and the effects of four parental site origin characteristics. The survival rate was 73% after 28-33 years of growth. Retention of curly-grained wood was high, the curly-grained wood structure is heritable and thus clonally efficiently achievable (only 3.5% of grafted individuals showed no occurrence of figured wood structure). The phenotypic expression of curliness manifested on the trunks as bulges, stem growth forms (tree/shrub) and stem technical quality showed a lower degree of stability (coincidence) between the parent trees and heterovegatively propagated progenies. Despite this, the conservation potential of seed orchard is very high, especially when stabilization of the stem growth forms affecting the survival and commercial value of progenies can be probably achieved by a more careful selection of scions. Overall, heterovegetative orchards seem to be a very promising method for the long-term conservation of curly birch populations, which, in addition to their great biological and ecological value, have considerable commercial potential.

Metal(loid) pollution of soils has important negative effects on the environment and human health. For the rehabilitation of these soils, some eco-innovative strategies, such as phytoremediation, could be chosen. This practice could establish a plant cover to reduce the toxicity of the pollutants and stabilize the soil, preventing soil erosion and water leaching; this technique is called phytoremediation. For this, plants need to be tolerant to the pollutants present; thus, phytoremediation can have better outcomes if endemic species of the polluted area are used. Finally, to further improve phytoremediation success, amendments can be applied to ameliorate soil conditions. Different amendments can be used, such as biochar, a good metal(loid) immobilizer, compost, a nutrient-rich product and iron sulfate, an efficient arsenic immobilizer. These amendments can either be applied alone or combined for further positive effects. In this context, a mesocosm experiment was performed to study the effects of three amendments, biochar, compost and iron sulfate, applied alone or combined to a former mine technosol, on the soil properties and the phytoremediation potential of two endemic species, Alnus sp. and Betula sp. Results showed that the different amendments reduced soil acidity and decreased metal(loid) mobility, thus improving plant growth. Both species were able to grow on the amended technosols, but alder seedlings had a much higher growth compared to birch seedlings. Finally, the combination of compost with biochar and/or iron sulfate and the establishment of endemic alder plants could be a solution to rehabilitate a former mine technosol.

Birches are generally known for their high genetic and morphological variability, which has resulted in the description of many species. Ojców birch was described in 1809 by Willibald Suibert Joseph Gottlieb Besser in Poland. Since then, several studies assessing its taxonomy were conducted. Today, various authors present Ojców birch at different taxonomic ranks. In Czechia, the Ojców birch is classified a critically endangered taxon and confirmed at one locality consisting of several tens of individuals. However, before a strategy for its conservation can be applied, we consider it necessary to assess the taxonomic position of the endangered Czech population and to evaluate its relationship to the original Polish population. This study aimed to evaluate the morphometric and genetic variability between populations of B. ×oycoviensis in Poland and the Czechia and their relationship to regional populations of B. pendula, one of the putative parental species of the Ojców birch. Altogether, 106 individuals were sampled, including the holotype of B. szaferi, the second putative parental species of B. ×oycoviensis, received from the herbarium of W. Szafer, which is deposited at the Institute of Botany in Kraków. Morphological analyses identified differences in leaves between B. ×oycoviensis and B. pendula. However, no significant differences were found in genome size between selected taxa/working units except for B. pendula sampled in Czechia. The identified difference of the Czech population of B. pendula is probably caused by population variability. Genetic variability between all the taxa under comparison, regardless of their origin, was also very low; only the benchmark taxa (B. nana and B. humilis) clearly differed from all samples analyzed. The results indicate minute morphological and negligible genetic variability between the Czech and Polish populations of B. ×oycoviensis. In light of our results, the classification of B. ×oycoviensis as B. pendula var. oycoviensis seems more accurate than all hitherto presented alternatives (e.g. B. ×oycoviensis as a separate species)., Competing Interests: The authors have declared that no competing interests exist.

In forest regeneration areas, alongside roads and railways, under electric power lines and above gas pipe lines, there is a need for regular sprout control. A biocontrol method against broadleaved sprouting with formulations including the decay fungus Chondrostereum purpureum (Pers. Ex Fr.) Pouzar has been shown to be effective. Yet, heavy rain during spreading of this fungal inoculum on freshly cut stumps may affect the efficacy of the treatment, i.e., stump mortality during the following years. Thus, we performed an experiment where freshly cut birch stump surfaces (Betula pendula Roth and Betula pubescens Ehrh.) were treated with fungal inoculum under heavy irrigation and without it. Furthermore, two different adjuvants which aimed to fix the fungal inoculum to freshly cut stumps during irrigation and to protect against solar radiation were tested. Our results revealed that the artificial rainstorm treatment caused a delay in the efficacy of C. purpureum, but after three growing seasons, there was no significant difference in the mortality of birch stumps treated under irrigation or without it (stump mortalities 74 and 86%, respectively). Adjuvants did not improve the efficacy in stumps treated under irrigation nor in those treated without irrigation. KEY POINTS: • Heavy rain delayed the sprout control efficacy of a fungus Chondrostereum purpureum. • Final efficacy of formulations was the same in wet and dry conditions. • No additional adjuvants are needed to improve formulations.

Climate warming is anticipated to make high latitude ecosystems stronger C sinks through increasing plant production. This effect might, however, be dampened by insect herbivores whose damage to plants at their background, non-outbreak densities may more than double under climate warming. Here, using an open-air warming experiment among Subarctic birch forest field layer vegetation, supplemented with birch plantlets, we show that a 2.3 °C air and 1.2 °C soil temperature increase can advance the growing season by 1-4 days, enhance soil N availability, leaf chlorophyll concentrations and plant growth up to 400%, 160% and 50% respectively, and lead up to 122% greater ecosystem CO 2 uptake potential. However, comparable positive effects are also found when insect herbivory is reduced, and the effect of warming on C sink potential is intensified under reduced herbivory. Our results confirm the expected warming-induced increase in high latitude plant growth and CO 2 uptake, but also reveal that herbivorous insects may significantly dampen the strengthening of the CO 2 sink under climate warming.

Tree architecture has evolved to support a top-heavy above-ground biomass, but this integral feature poses a weight-induced challenge to trunk stability. Maintaining an upright stem is expected to require vertical proprioception through feedback between sensing stem weight and responding with radial growth. Despite its apparent importance, the principle by which plant stems respond to vertical loading forces remains largely unknown. Here, by manipulating the stem weight of downy birch (Betula pubescens) trees, we show that cambial development is modulated systemically along the stem. We carried out a genetic study on the underlying regulation by combining an accelerated birch flowering program with a recessive mutation at the ELIMÄKI locus (EKI), which causes a mechanically defective response to weight stimulus resulting in stem collapse after just 3 months. We observed delayed wood morphogenesis in eki compared with WT, along with a more mechanically elastic cambial zone and radial compression of xylem cell size, indicating that rapid tissue differentiation is critical for cambial growth under mechanical stress. Furthermore, the touch-induced mechanosensory pathway was transcriptionally misregulated in eki, indicating that the ELIMÄKI locus is required to integrate the weight-growth feedback regulation. By studying this birch mutant, we were able to dissect vertical proprioception from the gravitropic response associated with reaction wood formation. Our study provides evidence for both local and systemic responses to mechanical stimuli during secondary plant development., Competing Interests: Declaration of Interests The authors declare no competing interest, (Copyright © 2019 Elsevier Ltd. All rights reserved.)

MADS-box genes encode transcription factors involved in the control of many important developmental processes, especially the flower development of angiosperms. Analysis on gene regulatory relationship between MADS-box genes is useful for understanding the molecular mechanism of flower development. In this study, we focused on the regulatory relationship between MADS-box transcription factors APETALA1 (AP1) and PISTILLATA(PI)/DEFICIENS (DEF) in birch. We found that BpPI was an authentic target gene of BpAP1, and BpAP1 activated the expression of BpPI via directly binding to the CArG box motif. Functional analysis of BpPI showed that overexpression of BpPI may delay flowering via restricting flowering activators, in which BpAP1 was significantly down-regulated. We further investigated the regulatory of BpAP1 by BpPI, and found that BpPI could directly bind to the promoter of BpAP1 to restrict BpAP1 expression. In addition, we also found that BpPI could interact with its hypothetical partner BpDEF to co-regulate BpAP1 in birch. Our results suggested that overexpression of BpPI may delay flowering via restricting flowering activators, and there is a negative feedback loop between BpAP1 and BpPI/BpDEF heterodimer in birch. Our results will bring new evidences for further analysis of the molecular mechanism of flower formation in plants that produced unisexual flowers., (Copyright © 2019 Elsevier B.V. All rights reserved.)

Community-scale impacts of glyphosate-based herbicides on wetland plant communities and the magnitude of those impacts that should be considered biologically relevant are poorly understood. We contrast three different thresholds for setting biologically meaningful critical effect sizes for complex ANOVA study designs. We use each of the of the critical effect sizes to determine optimal α levels for assessment of how different concentrations of glyphosate-based herbicides affect wetland plant communities over two years of herbicide application (alone and in combination with agricultural fertilizers) and two subsequent years without herbicide (or fertilizer) application. The application of glyphosate-based herbicides was found to result in a decrease in macrophyte species richness, an increase in macrophyte species evenness, a decrease in macrophyte cover and a reduction in community similarity. There was little evidence that nutrient additions directly or indirectly affected plant community endpoints. The glyphosate effects were evident in the first year of herbicide application in 2009, and became more pronounced in the second year of herbicide application in 2010. However, when herbicides were not applied in 2011, recovery was observed in most endpoints, with the exception being species evenness, for which partial recovery was not observed until 2012. Optimal α levels differed among the three critical effect sizes for each ANOVA term and endpoint combination, however regardless of differences in α levels, conclusions were generally consistent across all critical effect sizes., (Copyright © 2019 Elsevier Inc. All rights reserved.)

Background: Plant architecture, which is mostly determined by shoot branching, plays an important role in plant growth and development. Thus, it is essential to explore the regulatory molecular mechanism of branching patterns based on the economic and ecological importance. In our previous work, a multiple-branches birch mutant br was identified from 19 CINNAMOYL-COENZYME A REDUCTASE 1 (CCR1)-overexpressed transgenic lines, and the expression patterns of differentially expressed genes in br were analyzed. In this study, we further explored some other characteristics of br, including plant architecture, wood properties, photosynthetic characteristics, and IAA and Zeatin contents. Meanwhile, the T-DNA insertion sites caused by the insertion of exogenous BpCCR1 in br were identified to explain the causes of the mutation phenotypes., Results: The mutant br exhibited slower growth, more abundant and weaker branches, and lower wood basic density and lignin content than BpCCR1 transgenic line (OE2) and wild type (WT). Compared to WT and OE2, br had high stomatal conductance (Gs), transpiration rate (Tr), but a low non-photochemical quenching coefficient (NPQ) and chlorophyll content. In addition, br displayed an equal IAA and Zeatin content ratio of main branches' apical buds to lateral branches' apical buds and high ratio of Zeatin to IAA content. Two T-DNA insertion sites caused by the insertion of exogenous BpCCR1 in br genome were found. On one site, chromosome 2 (Chr2), no known gene was detected on the flanking sequence. The other site was on Chr5, with an insertion of 388 bp T-DNA sequence, resulting in deletion of 107 bp 5' untranslated region (UTR) and 264 bp coding sequence (CDS) on CORONATINE INSENSITIVE 1 (BpCOII). In comparison with OE2 and WT, BpCOI1 was down-regulated in br, and the sensitivity of br to Methyl Jasmonate (MeJA) was abnormal., Conclusions: Plant architecture, wood properties, photosynthetic characteristics, and IAA and Zeatin contents in main and lateral branches' apical buds changed in br over the study's time period. One T-DNA insertion was identified on the first exon of BpCOI1, which resulted in the reduction of BpCOI1 expression and abnormal perception to MeJA in br. These mutation phenotypes might be associated with a partial loss of BpCOI1 in birch.

The CUP-SHAPED COTYLEDON 2 ( CUC2 ) gene, which is negatively regulated by microRNA164 ( miR164 ), has been specifically linked to the regulation of leaf margin serration and the maintenance of phyllotaxy in model plants. However, few studies have investigated these effects in woody plants. In this study, we integrated genomic, transcriptomic, and physiology approaches to explore the function of BpCUC2 gene in Betula pendula growth and development. Our results showed that Betula pendula plants overexpressing BpCUC2 , which is targeted by BpmiR164 , exhibit shortened internodes and abnormal leaf shapes. Subsequent analysis indicated that the short internodes of BpCUC2 overexpressed transgenic lines and were due to decreased epidermal cell size. Moreover, transcriptome analysis, yeast one-hybrid assays, and ChIP-PCR suggested that BpCUC2 directly binds to the LTRECOREATCOR15 (CCGAC), CAREOSREP1 (CAACTC), and BIHD1OS (TGTCA) motifs of a series of IAA-related and cyclin-related genes to regulate expression. These results may be useful to our understanding of the functional role and genetic regulation of BpCUC2 .

In many woody dicot plant species, colder temperatures correlate with a greater degree of leaf dissection and with larger and more abundant leaf teeth (the serrated edges along margins). The measurement of site-mean characteristics of leaf size and shape (physiognomy), including leaf dissection and tooth morphology, has been an important paleoclimate tool for over a century. These physiognomic-based climate proxies require that all woody dicot plants at a site, regardless of species, change their leaf shape rapidly and predictably in response to temperature. Here we experimentally test these assumptions by growing five woody species in growth cabinets under two temperatures (17 and 25°C). In keeping with global site-based patterns, plants tend to develop more dissected leaves with more abundant and larger leaf teeth in the cool treatment. Overall, this upholds the assumption that leaf shape responds in a particular direction to temperature change. The assumption that leaf shape variables respond to temperature in the same way regardless of species did not hold because the responses varied by species. Leaf physiognomic models for inferring paleoclimate should take into account these species-specific responses., Competing Interests: The authors have declared that no competing interests exist.

Flowering is a crucial process for plants that is under complex genetic control. AP1 acts as an organizer and a switch for the transition from vegetative to reproductive growth. In our previous study, we found that overexpression of BpAP1 significantly promoted the formation of male inflorescence in birch (Betula platyphlla × B. pendula). In this study, we aimed at investigating the molecular regulatory mechanism of BpAP1 during the process of male inflorescence formation in birch. We found that overexpression of BpAP1 affected the expression of many flowering-related genes, and had significant effect on B class MADS-box genes. A BpAP1-mediated gene regulatory network was constructed and B class gene BpDEF was finally predicted as a key target gene of BpAP1. Chromatin immunoprecipitation results indicated that BpAP1 could directly regulate BpDEF during the process of male inflorescence formation. Yeast one-hybrid assays and its validation in tobacco results suggested that BpAP1 regulated BpDEF via binding to a consensus DNA sequence known as CArG box. Gene function analysis of BpDEF indicated that BpDEF may function in sex-determination, and in particular specify the identity of male inflorescence in birch. Our results provide valuable clues for our understanding of the molecular mechanism of BpAP1 during the process of male inflorescence formation in birch., (© The Author(s) 2019. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Tree bark is a highly specialized array of tissues that plays important roles in plant protection and development. Bark tissues develop from two lateral meristems; the phellogen (cork cambium) produces the outermost stem-environment barrier called the periderm, while the vascular cambium contributes with phloem tissues. Although bark is diverse in terms of tissues, functions and species, it remains understudied at higher resolution. We dissected the stem of silver birch (Betula pendula) into eight major tissue types, and characterized these by a combined transcriptomics and metabolomics approach. We further analyzed the varying bark types within the Betulaceae family. The two meristems had a distinct contribution to the stem transcriptomic landscape. Furthermore, inter- and intraspecies analyses illustrated the unique molecular profile of the phellem. We identified multiple tissue-specific metabolic pathways, such as the mevalonate/betulin biosynthesis pathway, that displayed differential evolution within the Betulaceae. A detailed analysis of suberin and betulin biosynthesis pathways identified a set of underlying regulators and highlighted the important role of local, small-scale gene duplication events in the evolution of metabolic pathways. This work reveals the transcriptome and metabolic diversity among bark tissues and provides insights to its development and evolution, as well as its biotechnological applications., (© 2019 The Authors. New Phytologist © 2019 New Phytologist Trust.)

The retrogressive succession is an important driver for dynamics of soil organic carbon (SOC) and total nitrogen (TN). We studied the quantitative distribution and synergistic accumulation characteristics of soil organic carbon and nitrogen in the primary broadleaved Korean pine (KP) forest and Betula platyphylla (BP) secondary forest in Changbai Mountain through paired plot approach. Further, we analyzed the changes of carbon pool and carbon sink effect in temperate forest soil caused by secondary succession and their carbon-nitrogen coupling mechanism. The results showed that the BP forest accumulated more organic carbon and nitrogen in the surface and subsurface soil (0-20 cm) than the KP forest, with relatively low soil C/N. Compared with KP forest, soil organic carbon storage in BP forest (0-20 cm) was higher by 14.7 t·hm -2 , equivalent to a soil carbon sink gain of 29.4 g·m -2 ·a -1 . SOC and TN concentrations were positively correlated in each soil layer of all forest types, causing a co-accumulative relationship between SOC and TN. The coefficient of determination (R 2 ) between SOC and TN in the upper soil layers of BP forest was significantly higher than that of the KP forest, indicating that SOC accumulation under the relatively N-rich BP forest was more dependent on the accumulation of organic nitrogen. In the upper soil layers (0-10 cm) where organic matter concentrated, there was no significant difference in light fraction organic carbon and nitrogen stock between the two forest types, whereas the content, stock, and allocation percentage of heavy fraction organic carbon and nitrogen of BP forest were all significantly higher than that of the KP forest, with an average increment of 8.5 t·hm -2 in heavy fraction organic carbon stock. Those results indicated that the increase of soil organic carbon and nitrogen during secondary succession was mainly due to the increases of soil organic carbon and nitrogen pools in mineral-bound stability. The carbon-nitrogen coupling mechanisms in litter decomposition and soil organic matter formation was an important driving mechanisms underlying the changes of soil organic carbon and nitrogen pools during secondary succession.

Air temperatures and precipitation are predicted to increase in the future, especially at high latitudes and particularly so during winter. In contrast to air temperatures, changes in soil temperatures are more difficult to predict, as the fate of the insulating snow cover is crucial in this respect. Soil conditions can also be affected by rain-on-snow events and warm spells during winter, resulting in freeze-thaw cycles, compacted snow, ice encasement and local flooding. These adverse conditions during winter could counteract the otherwise positive effects of climate change on forest growth and productivity. For studying the effects of different winter and snow conditions on young Downy birch (Betula pubescens Ehrh.) seedlings, we carried out a laboratory experiment with birch seedlings subjected to four different winter scenarios: snow covering the seedlings (SNOW), compressed snow and ice encasement (ICE), flooded and frozen soil (FLOOD) and no snow at all (NO SNOW). After the winter treatments we simulated a spring and early summer period of 9.5 weeks, and monitored the growth by measuring shoot and root biomass of the seedlings, and starch and soluble sugar concentrations. We also assessed the stress experienced by the seedlings by measuring leaf chlorophyll fluorescence and gas exchange. Although no difference in mortality was observed between the treatments, the seedlings in the SNOW and ICE treatments had significantly higher shoot and root biomass compared with those in the FLOOD and NO SNOW treatments. We found higher starch concentrations in roots of the seedlings in the SNOW and ICE treatments, compared with those in the FLOOD and NO SNOW treatments, although photosynthesis did not differ. Our results suggest a malfunction of carbohydrate distribution in the seedlings of the FLOOD and NO SNOW treatments, probably resulting from decreased sinks. The results underline the importance of an insulating and protecting snow cover for small tree seedlings, and that future winters with changed snow pattern might affect the growth of tree seedlings and thus possibly species composition and forest productivity., (© The Author(s) 2018. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Due to the urban heat island effect, the time of plant pollination might markedly vary within the area of a city. However, existing pollen forecasts do not reflect the spatial variations in the pollen release time within a heterogeneous urban environment. The main objective of this study was to model the spatial pattern of flowering onset (and thus the moment of pollen release) in silver birch (Betula pendula Roth.) in Poznań (Western Poland) using land surface temperature (LST) data and in situ phenological observations. The onset of silver birch flowering was observed at 34 urban and rural sites (973 trees) in Poznań from 2012 to 2014. Forty-four thermal variables were retrieved from MODerate Resolution Imaging Spectroradiometer (MODIS) data. To predict the spatio-temporal distribution of B. pendula flowering onset dates in a city, the ordinary and partial least squares, support vector machine and random forest regression models were applied. The models' performance was examined by an internal repeated k-fold cross-validation and external validation with archival phenological data (2010). Birch flowering began significantly earlier in the urban sites compared to the rural sites (from -1.4 days in 2013, to -4.1 days in 2012). The maximum March LST difference between the urban and rural sites reached 2.4 °C in 2013 and 4.5 °C in 2012. The random forest model performed best at validation stage, i.e. the root mean square error between the predicted and observed onset dates was 1.461 days, and the determination coefficient was 0.829. A calibrated model for predicting the timing of flowering in a heterogeneous city area is an important step in developing a fine-scale forecasting system that can directly estimate pollen exposure in places where allergy sufferers live. Importantly, by incorporating only pre-flowering thermal data into the model, location-specific allergy forecasts can be delivered to the public before the actual flowering time., (Copyright © 2018 Elsevier B.V. All rights reserved.)

MYB proteins play important roles in the regulation of plant growth, development, and stress responses. Overexpression of BplMYB46 from Betula platyphylla improved plant salt and osmotic tolerances. In the present study, the interaction of eight avian myeloblastosis viral oncogene homolog (MYB) transcription factors with BplMYB46 was investigated using the yeast two-hybrid system, which showed that BplMYB46 could form homodimers and heterodimers with BplMYB6, BplMYB8, BplMYB11, BplMYB12, and BplMYB13. Relative beta-glucuronidase activity and chromatin immunoprecipitation assays showed that the interaction between BplMYB46 and the five MYBs increased the binding of BplMYB46 to the MYBCORE motif. A subcellular localization study showed that these MYBs were all located in the nucleus. Real-time fluorescence quantitative PCR results indicated that the expressions of BplMYB46 and the five MYB genes could be induced by salt and osmotic stress, and the BplMYB46 and BplMYB13 exhibited the most similar expression patterns. BplMYB46 and BplMYB13 co-overexpression in tobacco using transient transformation technology improved tobacco's tolerance to salt and osmotic stresses compared with overexpressing BplMYB13 or BplMYB46 alone. Taken together, these results demonstrated that BplMYB46 could interact with five other MYBs to form heterodimers that activate the transcription of target genes via an enhanced binding ability to the MYBCORE motif to mediate reactive oxygen species scavenging in response to salt and osmotic stresses.

The research aim was to assess the effects of the plant hormone abscisic acid (ABA) and the growth regulator paclobutrazol (PBZ) on root system development during the in vitro culture of different birch and aspen genotypes. The studied genotypes involved two aspen (Populus tremula and Populus tremuloides × P. tremula) and two silver birch (Betula pendula) trees, with one of the birches characterized by its inability to root in vitro. For experiments, apical shoot segments were cultured on nutrient medium enriched with either ABA or PBZ. Additionally, the analysis of the endogenous hormones in shoots developed on hormone-free medium was conducted by high-performance liquid chromatography. The endogenous concentration of auxin indole-3-acetic acid was much higher in the aspens than that in the birches, while the highest concentration of ABA was found in the root-forming birch. The culturing of this birch genotype on medium enriched with ABA resulted in an increased root length and a higher number of lateral roots without any negative effect on either shoot growth or adventitious root (AR) formation, although these two processes were largely inhibited by ABA in the aspens. Meanwhile, PBZ promoted AR formation in both aspen and birch cultures but impaired secondary root formation and shoot growth in birches. These results suggest the use of ABA for the in vitro rooting of birches and PBZ for the rooting of aspens., (© 2018 Scandinavian Plant Physiology Society.)

A greenhouse experiment was conducted to study the potential of intercropping Arundo donax with Broussonetia papyrifera or Morus alba to remediate heavy metal-contaminated soil. The results showed that intercropping the herbaceous plant A. donax with woody plants B. papyrifera or M. alba was beneficial for plant growth on heavy metal-contaminated soil. This can effectively enhance the comprehensive enrichment capacity of heavy metals and improve soil enzyme activities. The photosynthetic pigment contents in the leaves of A. donax, B. papyrifera , and M. alba decreased along with remediation time under monoculture treatment for each plant. However, compared with 90 d cultivation, the chlorophyll-a and carotenoid contents in B. papyrifera leaves and chlorophyll-b and carotenoid contents in M. alba leaves under intercropping treatment after 270 d cultivation were only slightly changed. Furthermore, chlorophyll-a, chlorophyll-b, and carotenoid contents in M. alba leaves under intercropping treatment were significantly ( P <0.05) increased by 99.1%, 177.1%, and 119.9%, respectively, compared with monoculture-treated M. alba , and the total biomass of M. alba increased significantly ( P <0.05) by 26.1%. Compared with monoculture-treated A. donax , the total accumulation amounts of Pb and Zn in the shoots of combined plants was significantly ( P <0.05) enhanced by 171% and 124% under intercropping treatment of A. donax with B. papyrifera . Compared with monoculture-treated M. alba and A. donax , the total accumulation amounts of As and Pb in the shoots of intercropped plants were significantly ( P <0.05) enhanced by 150% and 76.5%, respectively, under intercropping treatment of A. donax with M. alba . Moreover, the fractions of As, Cd, Pb, and Zn in contaminated soil slightly changed under intercropping treatment of A. donax with B. papyrifera or M. alba , and soil urease, acid phosphatase, and total phosphatase activity was superior to part of the monoculture treatments after 270 d cultivation. The results further suggested that intercropping A. donax with B. papyrifera or M. alba could be effectively used for heavy metal-contaminated soil remediation, while simultaneously improving the biological quality in contaminated soil.

A pot experiment was conducted to investigate the effects of nitrogen (N) addition (0, 20, 40 g N m-2 year-1, N0, N20, N40, respectively) on the growth, and biomass accumulation and allocation of coniferous and deciduous (Picea asperata Mast. and Betula albosinensis Burk.) seedlings under a range of soil moisture limitation (40%, 50%, 60%, 80% and 100% of field capacity, FC). At 100% FC, growth of shade-tolerant P. asperata increased with N supply, while that of shade-intolerant B. albosinensis reached a maximum at N20, declining somewhat thereafter. At 60% FC and lower moisture content, water availability limited the growth of P. asperata seedlings, while N availability became progressively limiting to growth with moisture increasing above 60% FC. The transition from principally water-limited response to N-limited response in B. albosinensis occurred at lower moisture content. For P. asperata, these patterns reflected the responses of roots, consistent with changes in root/shoot biomass. For B. albosinensis the response reflected changes in shoot dimensions and root biomass fraction, the latter decreasing with size and foliar [N]. We are not aware of another study demonstrating such differences in the shape of the growth responses of seedlings of differing potential growth rate, across a range in belowground resource supply. The responses of leaf photosynthesis (as well as photosynthetic water and N-use efficiencies) were consistent with the observed growth response of P. asperata to water and N availability, but not of B. albosinensis, suggesting that leaf area dynamics (not measured) dominated the response of this species. Betula albosinensis, a fast-growing species, has a relative narrow range of soil water and N availability for maximum growth, achieved by preferential allocation to the shoot as resources meet the requirements at moderate N and water supply. In contrast, P. asperata increases shoot biomass progressively with increasing resources up to moderate water supply, preferentially growing more roots when resources are not limiting, suggesting that its capacity to produce shoot biomass may reach a biological limit at moderate levels of resource supply.

As a photoreceptor specifically for UV-B light, UVR8 gene plays an important role in the photomorphogenesis and developmental growth of plants. In this research, we isolated the UVR8 gene from birch, named BpUVR8 (AHY02156). BpUVR8 overexpression rescued the uvr8 mutant phenotype using functional complementation assay of BpUVR8 in Arabidopsis uvr8 mutants, which showed that the function of UVR8 is conserved between Arabidopsis and birch. The expression analysis of BpUVR8 indicated that this gene is expressed in various tissues, but its expression levels in leaves are higher than in other organs. Moreover, abiotic stress factors, such as UV-B, salinity, and abscisic acid (ABA) can induce the expression of BpUVR8 gene. Interestingly, the analysis of promoter activity indicated that BpUVR8 promoter not only has the promoting activity but can also respond to the induction of abiotic stress and ABA signal. So, we analyzed its function in ABA response via transgenic UVR8 overexpression in Arabidopsis. The BpUVR8 enhances the susceptibility to ABA, which indicates that BpUVR8 is regulated by ABA and can inhibit seed germination. The root length of 20-day-old 35S::BpUVR8/WT transgenic plants was 18% reduced as compared to the wild-type under the ABA treatment. The membrane of the BpUVR8-overexpressing in Arabidopsis thaliana was the most damaged after ABA treatment and 35S::BpUVR8/WT transgenic plant was more sensitive to ABA than the wild type. These results showed that BpUVR8 is a positive regulator in the ABA signal transduction pathway. In the presence of low dose of UV-B, the sensitivity of wild-type and 35S::BpUVR8/WT plants to ABA was reduced. Moreover, BpUVR8 regulates the expression of a subset of ABA-responsive genes, both in Arabidopsis and Betula platyphylla, under the ABA treatment. Our data provide evidence that BpUVR8 is a positive regulator in the UV-B-induced photomorphogenesis in plants. Moreover, we propose from this research that BpUVR8 might have an important role in integrating plant growth and ABA signaling pathway., (Copyright © 2018 Elsevier B.V. All rights reserved.)

Plant meristems were previously thought to be sterile. Today, meristem-associated shoot endophytes are mainly reported as contaminants from plant tissue cultures, the number of observed species being very low. However, the few strains characterized have the capacity for infecting host cells and affecting plant growth and development. Here we studied the communities of endophytic bacteria in the buds of mountain birch (Betula pubescens ssp. czerepanovii (N. I. Orlova) Hämet-Ahti) exposed to winter moth (Operophtera brumata L.) herbivory, to identify differences between sprouts and branches of mature birch trees. Mountain birch of the high subarctic is cyclically exposed to winter moth and produces sprouts to generate new trees as a survival mechanism. The majority (54%) of operational taxonomic units belonged to Xanthomonadaceae and Pseudomonales of Proteobacteria. Most of the observed species were classified as Xanthomonas (28%). Sprout buds had the highest diversity, containing approximately three times more species, and significantly more (43%) Pseudomonas species than the mature trees (14%). Our results demonstrate that endophytic communities of buds are richer than previously thought. We suggest that the meristem-associated endophytes should be studied further for a possible role in sprouting and aiding regeneration of trees.

The alternative prey hypothesis predicts that the interaction between generalist predators and their main prey is a major driver of population dynamics of alternative prey species. In Fennoscandia, changes in climate and human land use are assumed to alter the dynamics of cyclic small rodents (main prey) and lead to increased densities and range expansion of an important generalist predator, the red fox Vulpes vulpes. In order to better understand the role of these potential changes in community structure on an alternative prey species, willow ptarmigan Lagopus lagopus, we analyzed nine years of population census data from SE Norway to investigate how community interactions affected their population dynamics. The ptarmigan populations showed no declining trend during the study period, and annual variations corresponded with marked periodic small rodent peaks and declines. Population growth and breeding success were highly correlated, and both demographic variables were influenced by an interaction between red fox and small rodents. Red foxes affected ptarmigan negatively only when small rodent abundance was low, which is in accordance with the alternative prey hypothesis. Our results confirm the important role of red fox predation in ptarmigan dynamics, and indicate that if small rodent cycles are disrupted, this may lead to decline in ptarmigan and other alternative prey species due to elevated predation pressure., Competing Interests: The authors have declared that no competing interests exist.

Worldwide tree mortality as induced by climate change presents a challenge to forest managers. To successfully manage vulnerable forests requires the capacity of regeneration to compensate for losses from tree mortality. We observed rapid regeneration and the growth release of young trees after warming-induced mortality in a David aspen-dominated (Populus davidiana) broad-leaved forest in Inner Mongolia, China, as based on individual tree measurements taken in 2012 and 2015 from a 6-ha permanent plot. Warming and drought stress killed large trees 10-15 m tall with a total number of 2881 trees during 2011-2012, and also thinned the upper crowns. David aspen recruitment increased 2 times during 2012-2015 and resulted in a high transition probability of David aspen replacing the same or other species, whereas the recruitment of Mongolian oak (Quercus mongolica) was much lower: it decreased from 2012 to 2015, indicating that rapid regeneration represented a regrowth phase for David aspen, and not succession to Mongolian oak. Further, we found that the recruitment density increased with canopy openness, thus implying that warming-induced mortality enhanced regeneration. Our results suggest that David aspen has a high regrowth ability to offset individual losses from warming-induced mortality. This important insight has implications for managing this vulnerable forest in the semi-arid region of northern China.

Global changes occurring under different environmental conditions have changed stand competition, as well as nutrient and light availability, which has resulted in changes in productivity. Therefore, in the present study, the characteristics of tree-ring width formation of the prevailing Lithuanian tree species, Norway spruce, Scots pine and silver and downy birch, and key factors resulting in their differences during the last 36-year period were investigated at forest sites located on poor mineral oligotrophic and on nutrient-rich organic mesoeutrophic soils. The aim of the study was as follows: first, to separately detect the maximum possible seasonal effect of three groups of variables - meteorology, acidifying pollutants and surface ozone on the stem basal area increment (BAI) of the evaluated tree species; second, to assess the significance of each group of variables affecting the BAI of these tree species integrally with the remaining groups of variables. Norway spruce was found to be well adapted to recent environmental changes, which makes it one of the most favourable tree species for silviculture in the northeastern part of Europe. The rapid increases recorded in growth intensity since 1980 were attributed to the increase in air temperature, precipitation amount, nitrogen deposition during the vegetative stage and reductions in SO 2 concentrations and S deposition. Scots pine demonstrated the highest level of resilience and capacity to adapt to recent global changes because its reaction to both negative and favourable environmental factors was best expressed. Silver and downy birch tree reactions to the effects of air concentrations of acidifying compounds, their deposition and surface ozone concentrations were the least expressed; however, a significant decline in growth intensity indicated that these tree species experienced a reduced resistance to recent changes in environmental conditions in the mature and over-mature age groups., (Copyright © 2017 Elsevier B.V. All rights reserved.)

Ectomycorrhizal (ECM) fungi can influence the establishment and performance of host species by increasing nutrient and water absorption. Therefore, understanding the response of ECM fungi to expected changes in the global climate is crucial for predicting potential changes in the composition and productivity of forests. While anthropogenic activity has, and will continue to, cause global temperature increases, few studies have investigated how increases in temperature will affect the community composition of ectomycorrhizal fungi. The effects of global warming are expected to be particularly strong at biome boundaries and in the northern latitudes. In the present study, we analyzed the effects of experimental manipulations of temperature and canopy structure (open vs. closed) on ectomycorrhizal fungi identified from roots of host seedlings through 454 pyrosequencing. The ecotonal boundary site selected for the study was between the southern boreal and temperate forests in northern Minnesota, USA, which is the southern limit range for Picea glauca and Betula papyrifera and the northern one for Pinus strobus and Quercus rubra. Manipulations that increased air and soil temperature by 1.7 and 3.4 °C above ambient temperatures, respectively, did not change ECM richness but did alter the composition of the ECM community in a manner dependent on host and canopy structure. The prediction that colonization of boreal tree species with ECM symbionts characteristic of temperate species would occur was not substantiated. Overall, only a small proportion of the ECM community appears to be strongly sensitive to warming.