• Energy Research

Abstract Trees comprising high mountain forests have different requirements for site conditions (such as the water supply), thus current climate warming leads to varying reactions of upper forest boundaries depending on the site conditions and ecophysiological features of species. Positive reactions to an increasing heat supply during vegetative season may be hindered for drought-sensitive species by a water deficit in a cold environment, particularly during late winter or early spring. We investigated the radial growth of dark coniferous forest species Siberian stone pine (Pinus sibirica Du Tour) and Siberian fir (Abies sibirica Ledeb.) growing on slopes of different orientation (south-west, east, and north) near the upper forest boundary in an area undergoing fast climate warming: the Western Sayan Mountains (South Siberia, Russia), near a massive water reservoir. Correlations of tree-ring width chronologies with moving 21-day temperature series were used to more precisely determine the timing of temperature influence; an analysis of extreme and optimal years and multifactor regression modeling were applied to assess the most favorable/unfavorable thermal conditions in the study area and to estimate the tree growth reaction to the current climatic trends, respectively. Despite relatively low variation in growth (standard deviation Temperature increase in the study area during the last decades have occured about five times faster in the early spring (being enhanced by the reservoir) than in summer. This combination of spring and summer warming leads to an increase in P. sibirica radial growth on the northern slope and the stable growth of both species on sunlit slopes, i.e. providing a tentatively optimistic assessment of the dark coniferous forests’ near future in the region.

The response of vegetation to climate change is of special interest in regions where rapid warming is coupled with moisture deficit. This raises the question of the limits in plants’ acclimation ability and the consequent shifts of the vegetation cover. Radial growth dynamics and climatic response were studied in Scots pine (Pinus sylvestris L.), Siberian larch (Larix sibirica Ledeb.), and silver birch (Betula pendula Roth.) in the forest-steppe, and for Siberian elm (Ulmus pumila L.) in the steppe of South Siberia, as indicators of vegetation state and dynamics. Climate–growth relationships were analyzed by the following two approaches: (1) correlations between tree-ring width chronologies and short-term moving climatic series, and (2) optimization of the parameters of the Vaganov–Shashkin tree growth simulation model to assess the ecophysiological characteristics of species. Regional warming was accompanied by a slower increase of the average moisture deficit, but not in the severity of droughts. In the forest-steppe, the trees demonstrated stable growth and responded to the May–July climate. In the steppe, elm was limited by moisture deficit in May–beginning of June, during the peak water deficit. The forest-steppe stands were apparently acclimated successfully to the current climatic trends. It seems that elm was able to counter the water deficit, likely through its capacity to regulate transpiration by the stomatal morphology and xylem structure, using most of the stem as a water reservoir; earlier onset; and high growth rate, and these physiological traits may provide advantages to this species, leading to its expansion in steppes.

A joint analysis of dendrochronological and genomic data was performed to identify genetic mechanisms of adaptation and assess the adaptive genetic potential of Siberian stone pine (Pinus sibirica Du Tour) populations. The data obtained are necessary for predicting the effect of climate change and mitigating its negative consequences. Presented are the results of an association analysis of the variation of 84,853 genetic markers (single nucleotide polymorphisms—SNPs) obtained by double digest restriction-site associated DNA sequencing (ddRADseq) and 110 individual phenotypic traits, including dendrophenotypes based on the dynamics of tree-ring widths (TRWs) of 234 individual trees in six natural populations of Siberian stone pine, which have a history of extreme climatic stresses (e.g., droughts) and outbreaks of defoliators (e.g., pine sawfly [Neodiprion sertifer Geoff.]). The genetic structure of studied populations was relatively weak; samples are poorly differentiated and belong to genetically similar populations. Genotype–dendrophenotype associations were analyzed using three different approaches and corresponding models: General Linear Model (GLM), Bayesian Sparse Linear Mixed Model (BSLMM), and Bayesian-information and Linkage-disequilibrium Iteratively Nested Keyway (BLINK), respectively. Thirty SNPs were detected by at least two different approaches, and two SNPs by all three. In addition, three SNPs associated with mean values of recovery dendrophenotype (Rc) averaged across multiple years of climatic stresses were also found by all three methods. The sequences containing these SNPs were annotated using genome annotation of a very closely related species, whitebark pine (P. albicaulis Engelm.). We found that most of the SNPs with supposedly adaptive variation were located in intergenic regions. Three dendrophenotype-associated SNPs were located within the 10 Kbp regions and one in the intron of the genes encoding proteins that play a crucial role in ensuring the integrity of the plant’s genetic information, particularly under environmental stress conditions that can induce DNA damage. In addition, we found a correlation of individual heterozygosity with some dendrophenotypes. Heterosis was observed in most of these statistically significant cases; signs of homeostasis were also detected. Although most of the identified SNPs were not assigned to a particular gene, their high polymorphism and association with adaptive traits likely indicate high adaptive potential that can facilitate adaptation of Siberian stone pine populations to the climatic stresses and climate change.

In mountain ecosystems, plants are sensitive to climate changes, and an entire range of species distribution can be observed in a small area. Therefore, mountains are of great interest for climate-growth relationship analysis. In this study, the Siberian spruce’s (Picea obovata Ledeb.) radial growth and its climatic response were investigated in the Western Sayan Mountains, near the Sayano-Shushenskoe Reservoir. Sampling was performed at three sites along an elevational gradient: at the lower border of the species range, in the middle, and at the treeline. Divergence of growth trends between individual trees was observed at each site, with microsite landscape-soil conditions as the most probable driver of this phenomenon. Cluster analysis of individual tree-ring width series based on inter-serial correlation was carried out, resulting in two sub-set chronologies being developed for each site. These chronologies appear to have substantial differences in their climatic responses, mainly during the cold season. This response was not constant due to regional climatic change and the local influence of the nearby Sayano-Shushenskoe Reservoir. The main response of spruce to growing season conditions has a typical elevational pattern expected in mountains: impact of temperature shifts with elevation from positive to negative, and impact of precipitation shifts in the opposite direction. Chronologies of trees, growing under more severe micro-conditions, are very sensitive to temperature during September-April and to precipitation during October-December, and they record both inter-annual and long-term climatic variation. Consequently, it would be interesting to test if they indicate the Siberian High anticyclone, which is the main driver of these climatic factors.

Текст статьи не публикуется в открытом доступе в соответствии с политикой журнала. The genetic mechanisms underlying the relationship of individual heterozygosity (IndHet) with heterosis and homeostasis are not fully understood. Such an understanding, however, would have enormous value as it could be used to identify trees better adapted to environmental stress. Dendrochronology data, in particular the individual average radial increment growth of wood measured as the average tree ring width (AvTRW) and the variance of tree ring width (VarTRW) were used as proxies for heterosis (growth rate measured as AvTRW) and homeostasis (stability of the radial growth of individual trees measured as VarTRW), respectively. These traits were then used to test the hypothesis that IndHet can be used to predict heterosis and homeostasis of individual trees. Wood core and needle samples were collected from 100 trees of Siberian larch (Larix sibirica Ledeb.) across two populations located in Eastern Siberia. DNA samples were obtained from the needles of each individual tree and genotyped for eight highly polymorphic microsatellite loci. Then mean IndHet calculated based on the genotypes of eight loci for each tree was correlated with the statistical characteristics of the measured radial growth (AvTRW and VarTRW) and the individual standardized chronologies. The analysis did not reveal significant relationships between the studied parameters. In order to account for the strong dependence of the radial growth on tree age the age curves were examined. An original approach was employed to sort trees into groups based on the distance between these age curves. No relationship was found between these groups and the groups formed based on heterozygosity. However, further work with more genetic markers and increased sample sizes is needed to test this novel approach for estimating heterosis and homeostasis.

Dendrogenomics is a new interdisciplinary approach that allows joint analysis of dendrological and genomic data and opens up new ways to study the temporal dynamics of forest treelines, delineate spatial and temporal population structures, decipher individual tree responses to abiotic and biotic stresses, and evaluate the adaptive genetic potential of forest tree populations. These data are needed for the prediction of climate change effects and mitigation of the negative effects. We present here an association analysis of the variation of 27 individual tree traits, including adaptive dendrophenotypes reflecting the individual responses of trees to drought stress, such as the resistance (Rt), recovery (Rc), resilience (Rs), and relative resilience (RRs) indexes measured in 136 Siberian larch trees in 5 populations in the foothills of the Batenevsky Ridge (Kuznetsk Alatau, Republic of Khakassia, Russia), with variation of 9742 SNPs genotyped using ddRADseq in the same trees. The population structure of five closely located Siberian larch populations was relatively weak (FST = 0.018). We found that the level of individual heterozygosity positively correlated with the Rc and RR indices for the five studied drought periods and partly with the Rs indices for three drought periods. It seems that higher individual heterozygosity improves the adaptive capabilities of the tree. We also discovered a significant negative relationship between individual heterozygosity and the Rt index in four out of five periods, which means that growth slows down during droughts more in trees with higher individual heterozygosity and is likely associated with energy and internal resource reallocation toward more efficient water and energy usage and optimization of larch growth during drought years. We found 371 SNPs with potentially adaptive variations significantly associated with the variation of adaptive dendrophenotypes based on all three different methods of association analysis. Among them, 26 SNPs were located in genomic regions carrying functional genes: 21 in intergenic regions and 5 in gene-coding regions. Based on the obtained results, it can be assumed that these populations of Siberian larch have relatively high standing adaptive genetic variation and adaptive potential underlying the adaptations of larch to various climatic conditions.