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Forests globally are experiencing severe droughts, leading to significant reductions in growth, crown dieback and even tree mortality. The ability of forest ecosystems to acclimate to prolonged and repeated droughts is critical for their survival with ongoing climate change. In a five-year throughfall exclusion experiment, we investigated the long-term physiological and morphological acclimation of mature Norway spruce (Picea abies [L.] KARST.) and European beech (Fagus sylvatica L.) to repeated summer drought at the leaf, shoot and whole tree level. Throughout the drought period, spruce reduced their total water use by 70 % to only 4-9 L per day and tree, while beech was less affected with about 30 % reduction of water use. During the first two summers, spruce achieved this by closing their stomata by up to 80 %. Additionally, from the second drought summer onwards, spruce produced shorter shoots and needles, resulting in a stepwise reduction of total leaf area of over 50 % by the end of the experiment. Surprisingly, no premature leaf loss was observed. This reduction in leaf area allowed a gradual increase in stomatal conductance. After the five-year drought experiment, water consumption per leaf area was the same as in the controls, while the total water consumption of spruce was still reduced. In contrast, beech showed no significant reduction in whole-tree leaf area, but nevertheless reduced water use by up to 50 % by stomatal closure. If the restriction of transpiration by stomatal closure is sufficient to ensure survival of Norway spruce during the first drought summers, then the slow but steady reduction in leaf area will ensure successful acclimation of water use, leading to reduced physiological drought stress and long-term survival. Neighboring beech appeared to benefit from the water-saving strategy of spruce by using the excess water., Competing Interests: Declaration of competing interest The authors declare the following financial interests/personal relationships which may be considered as potential competing interests: Thorsten E.E. Grams, Hans Pretzsch reports financial support was provided by German Research Foundation. Benjamin D. Hesse reports financial support was provided by German Federal Environmental Foundation. Thorsten E.E. Grams reports financial support was provided by Bavarian State Ministry of the Environment and Consumer Protection. Thorsten E.E. Grams reports financial support was provided by Bavarian State Ministry of the Food, Agriculture and Forestry. If there are other authors, they 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.)

Northern temperate coniferous forests serve as crucial connectors between boreal and temperate forests, yet they are vulnerable to various stressors such as climate change and human activities. Severe drought poses a significant threat to plant species within these forests, prompting recent research into its impacts. However, many studies lack explicit definitions of post-disturbance vegetation processes and fail to identify potential interactions with disturbance factors, necessitating comprehensive discussions. This study examines the effects of drought on tree growth patterns of the main dominant species in northern temperate regions: Abies nephrolepis and Picea jezoensis, along with two commonly associated Betula ermanii, and Quercus mongolica. Additionally, new disturbance factors in forests inhabited by these species (A. nephrolepis and P. jezoensis) were evaluated based on community classification. The study sites were located in the Mt. Baekdu (Changbai) and South Korea regions, which are positioned at the southern limit of the phytogeographical patterns of target species. Results indicate that A. nephrolepis and P. jezoensis exhibit high levels of recovery and resilience, while B. ermanii and Q. mongolica demonstrate high resistance. Species-specific responses align with drought intensity, with resistance, recovery, and resilience decreasing notably with increasing pre-drought radial growth. South Korean forests, the invasion of the vine species Tripterygium regelii after the death of A. nephrolepis in the overstory vegetation threatens the regeneration of new trees. However, certain environmental factors, such as high rock exposure and dense overstory canopy, limit vine invasion. Based on the results, pre-drought radial growth emerges as a key determinant in how trees respond to drought. Additionally, the results suggest the potential for new disturbances to emerge in forest gaps due to overstory vegetation mortality induced by global warming. These findings contribute to a deeper understanding of increasing drought stress, aid in identifying climate refugia, and inform conservation priorities based on habitat characteristics., 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.)

Population genetics theory predicts a relationship between fitness, genetic diversity (H 0 ) and effective population size (N e ), which is often tested through heterozygosity-fitness correlations (HFCs). We tested whether population and individual fertility and heterozygosity are correlated in two endangered Mexican spruces (Picea martinezii and Picea mexicana) by combining genomic, demographic and reproductive data (seed development and germination traits). For both species, there was a positive correlation between population size and seed development traits, but not germination rate. Individual genome-wide heterozygosity and seed traits were only correlated in P. martinezii (general-effects HFC), and none of the candidate single nucleotide polymorphisms (SNPs) associated with individual fertility showed heterozygote advantage in any species (no local-effects HFC). We observed a single and recent (c. 30 thousand years ago (ka)) population decline for P. martinezii; the collapse of P. mexicana occurred in two phases separated by a long period of stability (c. 800 ka). Recruitment always contributed more to total population census than adult trees in P. mexicana, while this was only the case in the largest populations of P. martinezii. Equating fitness to either H 0 or N e , as traditionally proposed in conservation biology, might not always be adequate, as species-specific evolutionary factors can decouple the expected correlation between these parameters., (© 2024 The Author(s). New Phytologist © 2024 New Phytologist Foundation.)

The influence of tree age on the growth response of Picea likiangensis, a predominant timber species in southwestern China, to climatic factors has been under-researched. In this study, we examined the relationships between tree age and the response of P. likiangensis to climatic factors and extreme drought events using tree-ring samples procured from the southeastern edge of the Tibetan Plateau. The results revealed differential responses of the radial growth of P. likiangensis trees of varying ages to climatic factors and extreme drought events. Specifically, deficient water availability during the early growing season emerged as the principal factor constraining radial growth across all age classes. Young and middle-aged trees (<100 years) demonstrated greater responsiveness to water availability than did mature trees (>100 years). Mature trees, in contrast, demonstrated markedly greater resistance to extreme drought events than young and middle-aged trees. Comparative studies of individual trees across different ages revealed negligible differences in the response of young and middle-aged trees to climatic factors and extreme drought events. Given these responses, future forest management practices should prioritize young and middle-aged trees that are more affected by drought to maximize the ecological value of the species. According to the specific research objectives, sample collection processes should classify mature trees and young and middle-aged trees, to minimize the influence of tree age on the final findings of the study., 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 B.V. All rights reserved.)

Mixed-species forests are, for multiple reasons, promising options for forest management in Central Europe. However, the extent to which interspecific competition affects tree hydrological processes is not clear. High-resolution dendrometers capture subdaily variations in stem diameter; they can simultaneously monitor stem growth (irreversible changes in diameter) and water status (reversible changes) of individual trees. Using the information on water status, we aimed to assess potential effects of tree species mixture, expressed as local neighborhood identity, on night-time rehydration and water stress. We deployed 112 sensors in pure and mixed forest stands of European beech, Norway spruce and Douglas fir on four sites in the northwestern Germany, measuring stem diameter in 10-min intervals for a period of four years (2019-2022). In a mixture distribution model, we used environmental variables, namely soil matric potential, atmospheric vapor pressure deficit, temperature, precipitation and neighborhood identity to explain night-time rehydration, measured as the daily minimum tree water deficit (TWDmin). TWDmin was used as a daily indicator of water stress and the daily occurrence of sufficient water supply, allowing for stem growth (potential growth). We found that species and neighborhood identity affected night-time rehydration, but the impacts varied depending on soil water availability. While there was no effect at high water availability, increasing drought revealed species-specific patterns. Beech improved night-time rehydration in mixture with Douglas fir, but not in mixture with spruce. Douglas fir, however, only improved rehydration at a smaller share of beech in the neighborhood, while beech dominance tended to reverse this effect. Spruce was adversely affected when mixed with beech. At species level and under dry conditions, we found that night-time rehydration was reduced in all species, but beech had a greater capacity to rehydrate under high to moderate soil water availability than the conifers, even under high atmospheric water demand. Our study gives new insights into neighborhood effects on tree water status and highlights the importance of species-specific characteristics for tree-water relations in mixed-species forests. It shows that drought stress of European beech can be reduced by admixing Douglas fir, which may point towards a strategy to adapt beech stands to climate change., (© The Author(s) 2024. Published by Oxford University Press.)

In this study, we applied thermal dissipation probe technology to examine sap flow in various directions (east, south, west, and north) and at different depths (0-2, 2-4, 4-6 cm) within the stem of natural Picea mongolica trees in the eastern of Otindag Sandy Land to provide a scientific basis for accurately quantifying water consumption of P. mongolica forests through transpiration and to enhance the understanding of water relations. The results showed that the diurnal variation of sap flow in different directions displayed a unimodal curve, with the sap flow sequence being south>east>west>north. The sap flow at different sapwood depths exhibited an obvious unimodal curve, with a significant decrease as sapwood depth increased. Compared with that calculated from the mean sap flux density in four directions (23.57 kg·d -1 ), water consumption calculated using the mean value in south-east, east-west, south-west, north-east, north-south, and north-west was overestimated by 10.2%, 5.5%, 14.5%, and underestimated by 12.3%, 8.2%, 9.8%, respectively. The water consumption calculated using the values from the east, south, and west was overestimated by 6.1%, 14.4%, and 15.4%, respectively, and underestimated by 30.7% in the north. In addition, compared with the water consumption calculated from the mean value in three sapwood depths (48.51 kg·d -1 ), that calculated using sap flux density at sapwood depths of 0-2, 2-4, and 4-6 cm were overestimated by 18.8%, underestimated by 1.7%, and underestimated by 62.9%, respectively. These results indicated that sap flow of P. mongolica had significant azimuthal and radial variations, which considerably influence the estimation of tree water consumption. Installing probes at 0-2 cm simultaneously in both the north and east of the trunk could effectively reduce the estimation error of whole-tree water consumption by 4.2%. This approach enabled the accurate quantification of water consumption of individual P. mongolica trees in sandy areas, thereby improving the precision of transpiration water consumption estimates scaling up from individual level to stand level.

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.)

Photosynthetic acclimation to both warming and elevated CO 2 of boreal trees remains a key uncertainty in modelling the response of photosynthesis to future climates. We investigated the impact of increased growth temperature and elevated CO 2 on photosynthetic capacity (V cmax and J max ) in mature trees of two North American boreal conifers, tamarack and black spruce. We show that V cmax and J max at a standard temperature of 25°C did not change with warming, while V cmax and J max at their thermal optima (T opt ) and growth temperature (T g ) increased. Moreover, V cmax and J max at either 25°C, T opt or T g decreased with elevated CO 2 . The J max /V cmax ratio decreased with warming when assessed at both T opt and T g but did not significantly vary at 25°C. The J max /V cmax increased with elevated CO 2 at either reference temperature. We found no significant interaction between warming and elevated CO 2 on all traits. If this lack of interaction between warming and elevated CO 2 on the V cmax , J max and J max /V cmax ratio is a general trend, it would have significant implications for improving photosynthesis representation in vegetation models. However, future research is required to investigate the widespread nature of this response in a larger number of species and biomes., (© 2024 The Author(s). Plant, Cell & Environment published by John Wiley & Sons Ltd.)

The photosynthetic acclimation of boreal evergreen conifers is controlled by regulatory and photoprotective mechanisms that allow conifers to cope with extreme environmental changes. However, the underlying dynamics of photosystem II (PSII) and photosystem I (PSI) remain unresolved. Here, we investigated the dynamics of PSII and PSI during the spring recovery of photosynthesis in Pinus sylvestris and Picea abies using a combination of chlorophyll a fluorescence, P700 difference absorbance measurements, and quantification of key thylakoid protein abundances. In particular, we derived a new set of PSI quantum yield equations, correcting for the effects of PSI photoinhibition. Using the corrected equations, we found that the seasonal dynamics of PSII and PSI photochemical yields remained largely in balance, despite substantial seasonal changes in the stoichiometry of PSII and PSI core complexes driven by PSI photoinhibition. Similarly, the previously reported seasonal up-regulation of cyclic electron flow was no longer evident, after accounting for PSI photoinhibition. Overall, our results emphasize the importance of considering the dynamics of PSII and PSI to elucidate the seasonal acclimation of photosynthesis in overwintering evergreens. Beyond the scope of conifers, our corrected PSI quantum yields expand the toolkit for future studies aimed at elucidating the dynamic regulation of PSI., (© The Author(s) 2024. Published by Oxford University Press on behalf of the Society for Experimental Biology.)

The analysis of genetic variation underlying local adaptation in natural populations, together with the response to different external stimuli, is currently a hot topic in forest sciences, with the aim of identifying genetic markers controlling key phenotypic traits of interest for their inclusion in restoration and breeding programs. In Europe, one of the main tree species is Norway spruce (Picea abies (L.) H.Karst.). Using the MassARRAY® platform, 568 trees from North Rhine-Westphalia (Germany) were genotyped with 94 single nucleotide polymorphisms (SNPs) related to circadian and growth rhythms, and to stress response. The association analysis of the selected markers with health status and elevation was performed using three different methods, and those identified by at least two of these were considered as high confidence associated SNPs. While just five markers showed a weak association with health condition, 32 SNPs were correlated with elevation, six of which were considered as high confidence associated SNPs, as indicated by at least two different association methods. Among these genes, thioredoxin and pseudo response regulator 1 (PRR1) are involved in redox homeostasis and ROS detoxification, APETALA2-like 3 (AP2L3), a transcription factor, is involved in seasonal apical growth, and a RPS2-like is a disease resistance gene. The function of some of these genes in controlling light-dependent reactions and metabolic processes suggests signatures of adaptation to local photoperiod and the synchronization of the circadian rhythm. This work provides new insights into the genetic basis of local adaptation over a shallow elevation gradient in Norway spruce., (© 2024 The Authors. Plant Biology published by John Wiley & Sons Ltd on behalf of German Society for Plant Sciences, Royal Botanical Society of the Netherlands.)

Background: Variations in hydraulic conductivity may arise from species-specific differences in the anatomical structure and function of the xylem, reflecting a spectrum of plant strategies along a slow-fast resource economy continuum. Spruce (Picea spp.), a widely distributed and highly adaptable tree species, is crucial in preventing soil erosion and enabling climate regulation. However, a comprehensive understanding of the variability in anatomical traits of stems and their underlying drivers in the Picea genus is currently lacking especially in a common garden., Results: We assessed 19 stem economic properties and hydraulic characteristics of 17 Picea species grown in a common garden in Tianshui, Gansu Province, China. Significant interspecific differences in growth and anatomical characteristics were observed among the species. Specifically, xylem hydraulic conductivity (K s ) and hydraulic diameter exhibited a significant negative correlation with the thickness to span ratio (TSR), cell wall ratio, and tracheid density and a significant positive correlation with fiber length, and size of the radial tracheid. PCA revealed that the first two axes accounted for 64.40% of the variance, with PC1 reflecting the trade-off between hydraulic efficiency and mechanical support and PC2 representing the trade-off between high embolism resistance and strong pit flexibility. Regression analysis and structural equation modelling further confirmed that tracheid size positively influenced K s , whereas the traits DWT, D&#95;r, and TSR have influenced K s indirectly. All traits failed to show significant phylogenetic associations. Pearson's correlation analysis demonstrated strong correlations between most traits and longitude, with the notable influence of the mean temperature during the driest quarter, annual precipitation, precipitation during the wettest quarter, and aridity index., Conclusions: Our results showed that xylem anatomical traits demonstrated considerable variability across phylogenies, consistent with the pattern of parallel sympatric radiation evolution and global diversity in spruce. By integrating the anatomical structure of the stem xylem as well as environmental factors of origin and evolutionary relationships, our findings provide novel insights into the ecological adaptations of the Picea genus., (© 2024. The Author(s).)

Modeling and simulating the growth of the branching of tree species remains a challenge. With existing approaches, we can reconstruct or rebuild the branching architectures of real tree species, but the simulation of the growth process remains unresolved. First, we present a tree growth model to generate branching architectures that resemble real tree species. Secondly, we use a quantitative morphometric approach to infer the shape similarity of the generated simulations and real tree species. Within a functional-structural plant model, we implement a set of biological parameters that affect the branching architecture of trees. By modifying the parameter values, we aim to generate basic shapes of spruce, pine, oak and poplar. Tree shapes are compared using geometric morphometrics of landmarks that capture crown and stem outline shapes. Five biological parameters, namely xylem flow, shedding rate, proprioception, gravitysense and lightsense, most influenced the generated tree branching patterns. Adjusting these five parameters resulted in the different tree shapes of spruce, pine, oak, and poplar. The largest effect was attributed to gravity, as phenotypic responses to this effect resulted in different growth directions of gymnosperm and angiosperm branching architectures. Since we were able to obtain branching architectures that resemble real tree species by adjusting only a few biological parameters, our model is extendable to other tree species. Furthermore, the model will also allow the simulation of structural tree-environment interactions. Our simplifying approach to shape comparison between tree species, landmark geometric morphometrics, showed that even the crown-trunk outlines capture species differences based on their contrasting branching architectures., (© The Author(s) 2024. Published by Oxford University Press.)

To increase the resilience of forests to drought and other hazards, foresters are increasingly planting mixed stands. This requires knowledge about the drought response of tree species in pure and mixed-culture neighborhoods. In addition, drought frequently interacts with continued atmospheric nitrogen (N) deposition. To disentangle these factors for European beech, Norway spruce and Douglas fir, we conducted a replicated 3-factorial sapling growth experiment with three moisture levels, (high, medium, and low), two N levels (high and ambient), and pure and mixed-culture neighborhoods. We measured biomass, stomatal conductance (GS), shoot water potential (at predawn: ΨPD, midday, and turgor loss point: ΨTLP), branch xylem embolism resistance (Ψ50) and minimum epidermal conductance (Gmin). The three species differed most with respect to Gmin (10-fold higher in beech than in the conifers), hydroscape area (larger in beech), and the time elapsed to reach stomatal closure (TΨGS90) and ΨTLP (TTLP; shorter in beech), while Ψ50 and ΨTLP were remarkably similar. Neighborhood (pure vs mixed-culture) influenced biomass production, water status and hydraulic traits, notably GS (higher in Douglas fir, but lower in spruce and beech, in mixtures than pure culture), hydraulic safety margin (smaller for beech in mixtures), and TΨGS90 and TTLP (shorter for spruce in mixture). High N generally increased GS, but no consistent N effects on leaf water status and hydraulic traits were detected, suggesting that neighbor identity had a larger effect on plant water relations than N availability. We conclude that both tree neighborhood and N availability modulate the drought response of beech, spruce, and Douglas fir. Species mixing can alleviate the drought stress of some species, but often by disadvantaging other species. Thus, our study suggests that stabilizing and building resilience of production forests against a drier and warmer climate may depend primarily on the right species choice; species mixing can support the agenda., (© Crown copyright 2024.)

Using a unique 8-year data set (2010-2017) of phloem data, we studied the effect of temperature and precipitation on the phloem anatomy (conduit area, widths of ring, early and late phloem) and xylem-ring width in two coexisting temperate tree species, Picea abies and Fagus sylvatica, from three contrasting European temperate forest sites. Histometric analyses were performed on microcores taken from tree stems in autumn. We found high interannual variability and sensitivity of phloem anatomy and xylem-ring widths to precipitation and temperature; however, the responses were species- and site-specific. The contrasting response of xylem and phloem-ring widths of the same tree species to weather conditions was found at the two Slovenian sites generally well supplied with precipitation, while at the driest Czech site, the influence of weather factors on xylem and phloem ring widths was synchronised. Since widths of mean annual xylem and phloem increments were narrowest at the Czech site, this site is suggested to be most restrictive for the radial growth of both species. By influencing the seasonal patterns of xylem and phloem development, water availability appears to be the most important determinant of tissue- and species-specific responses to local weather conditions., (© 2024 The Authors. Plant, Cell & Environment published by John Wiley & Sons Ltd.)

The synthesis of a large body of evidence from field experiments suggests more diverse plant communities are more productive as well as more resistant to the effects of climatic extremes like drought. However, this view is strongly based on data from grasslands due to the limited empirical evidence from tree diversity experiments. Here we report on the relationship between tree diversity and productivity over 10 years in a field experiment established in 2005 that was then affected by the 2018 mega-drought in central Europe. Across a number of years, tree species diversity and productivity were significantly positively related; however, the slope switched to negative in the year of the drought. Net diversity effects increased through time, with complementarity effects making greater contributions to the net diversity effect than selection effects. Complementarity effects were clearly positive in three- and five-species mixtures before the drought (2012-2016) but were found to decrease in the year of the drought. Selection effects were clearly positive in 2016 and remained positive in the drought year 2018 in two-, three-, and five-species mixtures. The survival of Norway spruce (Picea abies) plummeted in response to the drought, and a negative relationship between species diversity and spruce survival was found. Taken together, our findings suggest that tree diversity per se may not buffer communities against the impacts of extreme drought and that tree species composition and the drought tolerance of tree species (i.e., species identity) will be important determinants of community productivity as the prevalence of drought increases., (© 2024 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Beneficial and negative effects of species interactions can strongly influence water fluxes in forest ecosystems. However, little is known about how trees dynamically adjust their water use when growing with interspecific neighbours. Therefore, we investigated the interaction effects between Fagus sylvatica (European beech) and Picea abies (Norway spruce) on water-use strategies and aboveground structural characteristics. We used continuous in situ isotope spectroscopy of xylem and soil water to investigate source water dynamics and root water uptake depths. Picea abies exhibited a reduced sun-exposed crown area in equally mixed compared with spruce-dominated sites, which was further correlated to a reduction in sap flow of -14.5 ± 8.2%. Contrarily, F. sylvatica trees showed +13.3 ± 33.3% higher water fluxes in equally mixed compared with beech-dominated forest sites. Although a significantly higher crown interference by neighbouring trees was observed, no correlation of water fluxes and crown structure was found. High time-resolved xylem δ2H values showed a large plasticity of tree water use (-74.1 to -28.5‰), reflecting the δ2H dynamics of soil and especially precipitation water sources. Fagus sylvatica in equally mixed sites shifted water uptake to deeper soil layers, while uptake of fresh precipitation was faster in beech-dominated sites. Our continuous in situ water stable isotope measurements traced root water uptake dynamics at unprecedented temporal resolution, indicating highly dynamic use of water sources in response to precipitation and to neighbouring species competition. Understanding this plasticity may be highly relevant in the context of increasing water scarcity and precipitation variability under climate change., (© The Author(s) 2023. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Continuous cover forestry (CCF) has gained interest as an alternative to even-aged management particularly on drained peatland forests. However, relatively little is known about the physiological response of suppressed trees when larger trees are removed as a part of CCF practices. Consequently, studies concentrating on process-level modeling of the response of trees to selection harvesting are also rare. Here, we compared, modeled and measured harvest response of previously suppressed Norway spruce (Picea abies) trees to a selection harvest. We quantified the harvest response by collecting Norway spruce tree-ring samples in a drained peatland forest site and measuring the change in stable carbon and oxygen isotopic ratios of wood formed during 2010-20, including five post-harvest years. The measured isotopic ratios were compared with ecosystem-level process model predictions for ${\kern0em }^{13}$C discrimination and ${\kern0em }^{18}$O leaf water enrichment. We found that the model predicted similar but lower harvest response than the measurements. Furthermore, accounting for mesophyll conductance was important for capturing the variation in ${\kern0em }^{13}$C discrimination. In addition, we performed sensitivity analysis on the model, which suggests that the modeled ${\kern0em }^{13}$C discrimination is sensitive to parameters related to CO2 transport through stomata to the mesophyll., (© The Author(s) 2023. Published by Oxford University Press.)

Conifers growing in temperate forests utilize sustained forms of thermal dissipation during winter to protect the photosynthetic apparatus from damage, which can be monitored via pronounced reductions in photochemical efficiency (Fv/Fm) during winter. Eastern white pine (Pinus strobus L.) and white spruce (Picea  glauca (Moench) Voss) are known to recover from winter stress at different rates, with pine recovering more slowly than spruce, suggesting different mechanisms for sustained dissipation in these species. Our objectives were to monitor pine and spruce throughout spring recovery in order to provide insights into key mechanisms for sustained dissipation in both species. We measured chlorophyll fluorescence, pigments, and abundance and phosphorylation status of key photosynthetic proteins. We found that both species rely on two forms of sustained dissipation involving retention of high amounts of antheraxanthin (A) + zeaxanthin (Z), one that is very slowly reversible and temperature independent and one that is more dynamic and occurs only on subzero days. Differences in protein abundance suggest that spruce, but not pine, likely upregulates cyclic or alternative pathways of electron transport involving the cytochrome b6f complex and photosystem I (PSI). Both species show an increased sustained phosphorylation of the D1 protein on subzero days, and spruce additionally shows dramatic increases in the sustained phosphorylation of light-harvesting complex II (LHCII) and other PSII core proteins on subzero days only, suggesting that a mechanism of sustained dissipation that is temperature dependent requires sustained phosphorylation of photosynthetic proteins in spruce, possibly allowing for direct energy transfer from PSII to PSI as a mechanism of photoprotection. The data suggest differences in strategy among conifers in mechanisms of sustained thermal dissipation in response to winter stress. Additionally, the flexible induction of sustained A + Z and phosphorylation of photosynthetic proteins in response to subzero temperatures during spring recovery seem to be important in providing photoprotection during transitional periods with high temperature fluctuation., (© The Author(s) 2023. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

In 2022, Europe emerged from eight of the hottest years on record, leading to significant spruce mortality across Europe. The particularly dry weather conditions of 2018 triggered an outbreak of bark beetles (Ips typographus), causing the loss of thousands of hectares of Norway spruce stands, including in Wallonia and North-eastern France. A methodology for detecting the health status of spruce was developed based on a dense time series of satellite imagery (Sentinel-2). The time series of satellite images allowed the modelling of the spectral response of healthy spruce forests over the seasons: a decrease in photosynthetic activity of the forest canopy causes deviations from this normal seasonal vegetation index trajectory. These anomalies are caused by a bark beetle attack and are detected automatically. The method leads in the production of an annual spruce health map of Wallonia and Grand-Est. The goal of this paper is to assess the damage caused by bark beetle using the resulting spruce health maps. A second objective was to compare the influence of basic variables on the mortality of spruce trees in these two regions. Lasted 6 years (2017-2022), bark beetle has destroyed 12.2% (23,674 ha) of the spruce area in Wallonia and Grand-Est of France. This study area is composed of three bioclimatic areas: Plains, Ardennes and Vosges, which have not been equally affected by bark beetle attacks. The plains were the most affected, with 50% of spruce forests destroyed, followed by the Ardennes, which lost 11.3% of its spruce stands. The Vosges was the least affected bioclimatic area, with 5.6% of spruce stands lost. For the most problematic sites, Norway spruce forestry should no longer be considered., (© 2024. The Author(s), under exclusive licence to Springer Nature Switzerland AG.)

Temperate forests are undergoing significant transformations due to the influence of climate change, including varying responses of different tree species to increasing temperature and drought severity. To comprehensively understand the full range of growth responses, representative datasets spanning extensive site and climatic gradients are essential. This study utilizes tree-ring data from 550 sites from the temperate forests of Czechia to assess growth trends of six dominant Central European tree species (European beech, Norway spruce, Scots pine, silver fir, sessile and pedunculate oak) over 1990-2014. By modeling mean growth series for each species and site, and employing principal component analysis, we identified the predominant growth trends. Over the study period, linear growth trends were evident across most sites (56% increasing, 32% decreasing, and 10% neutral). The proportion of sites with stationary positive trends increased from low toward high elevations, whereas the opposite was true for the stationary negative trends. Notably, within the middle range of their distribution (between 500 and 700 m a.s.l.), Norway spruce and European beech exhibited a mix of positive and negative growth trends. While Scots pine growth trends showed no clear elevation-based pattern, silver fir and oaks displayed consistent positive growth trends regardless of site elevation, indicating resilience to the ongoing warming. We demonstrate divergent growth trajectories across space and among species. These findings are particularly important as recent warming has triggered a gradual shift in the elevation range of optimal growth conditions for most tree species and has also led to a decoupling of growth trends between lowlands and mountain areas. As a result, further future shifts in the elevation range and changes in species diversity of European temperate forests can be expected., (© 2024 John Wiley & Sons Ltd.)

Climate change is rapidly altering weather patterns, resulting in shifts in climatic zones. The survival of trees in specific locations depends on their functional traits. Local populations exhibit trait adaptations that ensure their survival and accomplishment of growth and reproduction processes during the growing season. Studying these traits offers valuable insights into species responses to present and future environmental conditions, aiding the implementation of measures to ensure forest resilience and productivity. This study investigates the variability in functional traits among five black spruce (Picea mariana (Mill.) B.S.P.) provenances originating from a latitudinal gradient along the boreal forest, and planted in a common garden in Quebec, Canada. We examined differences in bud phenology, growth performance, lifetime first reproduction, and the impact of a late-frost event on tree growth and phenological adjustments. The findings revealed that trees from northern sites exhibit earlier budbreak, lower growth increments, and reach reproductive maturity earlier than those from southern sites. Late-frost damage affected growth performance, but no phenological adjustment was observed in the successive year. Local adaptation in the functional traits may lead to maladaptation of black spruce under future climate conditions or serve as a potent evolutionary force promoting rapid adaptation under changing environmental conditions., (© 2023. The Author(s).)

Tree line areas exhibited significant changes in response to climate change, including upward migration. Lower tree line dynamics are rarely studied, but as unique features in arid and semi-arid areas, they may influence forest distribution. Here, eight lower tree line plots in a Picea crassifolia Kom. (Qinghai spruce) forest in the arid and semi-arid Qilian Mountains of northwestern China were used to determine changes in tree line location and relationships with meteorological factors during 1968-2018. The results showed that the lower tree line descended by an average of 9.82 m during 1968 to 2018, and exhibited almost no change after 2008. The change in the lower tree line was significantly correlated with the annual average temperature (°C) and annual precipitation (mm) and may be affected by human activities. In the past 50 years, the lower tree line in arid areas exhibited a downward trend. Our findings indicate that the movement of the lower tree line is also an important aspect of climatic changes in coniferous forest distribution in arid and semi-arid mountains., Competing Interests: The authors have declared that no competing interests exist., (Copyright: © 2023 Fang et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

Abiotic factors such as water and nutrient availability can exert a dominant influence on the susceptibility of plants to various pathogens. Effects of abiotic environmental factors on phenolic compound concentrations in the plant tissue may represent one of the major underlying mechanisms, as these compounds are known to play a substantial role in plant resistance to pests. In particular, this applies to conifer trees, in which a large range of phenolic compounds are produced constitutively and/or induced by pathogen attack. We subjected Norway spruce saplings to water limitation and elevated nutrient supply over 2 years and subsequently controlled infection with the needle rust Chrysomyxa rhododendri (DC.) de Bary and analysed both constitutive and inducible phenolic compound concentrations in the needles as well as the degree of infection. Compared with the control group, both drought and fertilization profoundly modified the constitutive and pathogen-induced profiles of phenolic compounds, but had little impact on the total phenolic content. Fertilization predominantly affected the inducible phenolic response and led to higher infection rates by C. rhododendri. Drought stress, in contrast, mainly shaped the phenolic profiles in healthy plant parts and had no consequences on the plant susceptibility. The results show that specific abiotic effects on individual compounds seem to be decisive for the infection success of C. rhododendri, whereby the impaired induced response in saplings subjected to nutrient supplementation was most critical. Although drought effects were minor, they varied depending on the time and length of water limitation. The results indicate that prolonged drought periods in the future may not significantly alter the foliar defence of Norway spruce against C. rhododendri, but fertilization, often propagated to increase tree growth and forest productivity, can be counterproductive in areas with high pathogen pressure., (© The Author(s) 2023. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

Common-garden trials of forest trees provide phenotype data used to assess growth and local adaptation; this information is foundational to tree breeding programs, genecology, and gene conservation. As jurisdictions consider assisted migration strategies to match populations to suitable climates, in situ progeny and provenance trials provide experimental evidence of adaptive responses to climate change. We used drone technology, multispectral imaging, and digital aerial photogrammetry to quantify spectral traits related to stress, photosynthesis, and carotenoids, and structural traits describing crown height, size, and complexity at six climatically disparate common-garden trials of interior spruce (Picea engelmannii × glauca) in western Canada. Through principal component analysis, we identified key components of climate related to temperature, moisture, and elevational gradients. Phenotypic clines in remotely sensed traits were analyzed as trait correlations with provenance climate transfer distances along principal components (PCs). We used traits showing clinal variation to model best linear unbiased predictions for tree height (R 2  = .98-.99, root mean square error [RMSE] = 0.06-0.10 m) and diameter at breast height (DBH, R 2  = .71-.97, RMSE = 2.57-3.80 mm) and generated multivariate climate transfer functions with the model predictions. Significant (p < .05) clines were present for spectral traits at all sites along all PCs. Spectral traits showed stronger clinal variation than structural traits along temperature and elevational gradients and along moisture gradients at wet, coastal sites, but not at dry, interior sites. Spectral traits may capture patterns of local adaptation to temperature and montane growing seasons which are distinct from moisture-limited patterns in stem growth. This work demonstrates that multispectral indices improve the assessment of local adaptation and that spectral and structural traits from drone remote sensing produce reliable proxies for ground-measured height and DBH. This phenotyping framework contributes to the analysis of common-garden trials towards a mechanistic understanding of local adaptation to climate., (© 2023 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Lowland conifer forests dominated by black spruce (Picea mariana) and tamarack (Larix laricina) typically occur in peatlands in the boreal North American forest with near-surface water tables throughout the year. These forests are ecologically and economically important resources that may be impacted by climate change. However, information characterizing effects of forest disturbance, such as even-aged harvest on water table dynamics is needed to evaluate which forest tree species cover types are most hydrologically susceptible to even-aged harvest and changes in precipitation. We used a chronosequence approach to evaluate water table fluctuations and evapotranspiration across four stand age classes (<10, 15-30, 40-80, and >100-years old) and three distinct forest cover types (productive black spruce, stagnant black spruce, and tamarack) for a period of three years in Minnesota, USA. In general, there is limited evidence for elevated water tables in the younger age classes; the <10-year age class had no significant difference in mean weekly water table depth compared to the older age classes across all cover types. Estimated actual daily evapotranspiration (ET) generally agreed with the water table observations, with the exception of the tamarack cover type where ET was significantly lower in the <10-year age class. Productive black spruce sites that are 40-80-years old had higher evapotranspiration, and lower water table, possibly reflecting increased transpiration associated with the stem exclusion stage of stand development. Tamarack in the 40-80-year age class had higher water tables but no difference in ET compared to all other age classes, indicating that other external factors are driving higher water tables in that age class. To evaluate susceptibility to changing climate, we also assessed the sensitivity and response of water table dynamics to pronounced differences in growing season precipitation that occurred across study years. In general, tamarack forests are more sensitive to changes in precipitation compared to the two black spruce forest cover types. These findings can inform expected responses of site hydrology for a range of precipitation scenarios that may occur under future climate and be used by forest managers to evaluate hydrologic impacts of forest management activities across lowland conifer forest cover types., 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 © 2023. Published by Elsevier Ltd.)

Prolonged drought and susceptibility to biotic stressors induced an extensive calamity in Norway spruce (Picea abies (L.) Karst.) and widespread crown defoliation in European beech (Fagus sylvatica L.) in Central Europe. For future management decisions, it is crucial to link changes in canopy cover to site conditions. However, current knowledge on the role of soil properties for drought-induced forest disturbance is limited due to the scarcity and low spatial resolution of soil information. We present a fine-scale assessment on the role of soil properties for forest disturbance in Norway spruce and European beech derived from optical remote sensing. A forest disturbance modeling framework based on Sentinel-2 time series was applied on 340 km 2 in low mountain ranges of Central Germany. Spatio-temporal information on forest disturbance was calculated at 10 m spatial resolution in the period 2019-2021 and intersected with high-resolution soil information (1:10,000) based on roughly 2850 soil profiles. We found distinct differences in disturbed area, depending on soil type, texture, stoniness, effective rooting depth and available water capacity (AWC). For spruce, we found a polynomial relationship between AWC (R 2  = 0.7) and disturbance, with highest disturbed area (65 %) for AWC between 90 and 160 mm. Interestingly, we found no evidence for generally higher disturbance on shallow soils, although stands on the deepest soils were significantly less affected. Noteworthy, sites affected first did not necessarily exhibit highest proportions of disturbed area post-drought, indicating recovery or adaptation. We conclude that site- and species-specific understanding of drought impacts benefits from a combination of remote sensing and fine-scale soil information. Since our approach revealed which sites were affected first and most, it qualifies for prioritizing in situ monitoring activities to most vulnerable stands in acute drought conditions as well as for developing long-term strategies for reforestation and site-specific risk assessment for precision forestry., 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 © 2023 Elsevier B.V. All rights reserved.)

It is assumed that the stimulatory effects of elevated CO2 concentration ([CO2]) on photosynthesis and growth may be substantially reduced by co-occurring environmental factors and the length of CO2 treatment. Here, we present the study exploring the interactive effects of three manipulated factors ([CO2], nitrogen supply and water availability) on physiological (gas-exchange and chlorophyll fluorescence), morphological and stoichiometric traits of Norway spruce (Picea abies) saplings after 2 and 3 years of the treatment under natural field conditions. Such multifactorial studies, going beyond two-way interactions, have received only limited attention until now. Our findings imply a significant reduction of [CO2]-enhanced rate of CO2 assimilation under reduced water availability which deepens with the severity of water depletion. Similarly, insufficient nitrogen availability leads to a down-regulation of photosynthesis under elevated [CO2] being particularly associated with reduced carboxylation efficiency of the Rubisco enzyme. Such adjustments in the photosynthesis machinery result in the stimulation of water-use efficiency under elevated [CO2] only when it is combined with a high nitrogen supply and reduced water availability. These findings indicate limited effects of elevated [CO2] on carbon uptake in temperate coniferous forests when combined with naturally low nitrogen availability and intensifying droughts during the summer periods. Such interactions have to be incorporated into the mechanistic models predicting changes in terrestrial carbon sequestration and forest growth in the future., (© The Author(s) 2023. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Mixed forest stands tend to be more resistant to drought than species-specific stands partially due to complementarity in root ecology and physiology. We asked whether complementary differences in the drought resistance of soil microbiomes might contribute to this phenomenon. We experimented on the effects of reduced soil moisture on bacterial and fungal community composition in species-specific (single species) and mixed-species root zones of Norway spruce and European beech forests in a 5-year-old throughfall-exclusion experiment and across seasonal (spring-summer-fall) and latitudinal moisture gradients. Bacteria were most responsive to changes in soil moisture, especially members of Rhizobiales, while fungi were largely unaffected, including ectomycorrhizal fungi (EMF). Community resistance was higher in spruce relative to beech root zones, corresponding with the proportions of drought-favored (more in spruce) and drought-sensitive bacterial taxa (more in beech). The spruce soil microbiome also exhibited greater resistance to seasonal changes between spring (wettest) and fall (driest). Mixed-species root zones contained a hybrid of beech- and spruce-associated microbiomes. Several bacterial populations exhibited either enhanced resistance or greater susceptibility to drought in mixed root zones. Overall, patterns in the relative abundances of soil bacteria closely tracked moisture in seasonal and latitudinal precipitation gradients and were more predictive of soil water content than other environmental variables. We conclude that complementary differences in the drought resistance of soil microbiomes can occur and the likeliest form of complementarity in mixed-root zones coincides with the enrichment of drought-tolerant bacteria associated with spruce and the sustenance of EMF by beech., 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 © 2023 Elsevier B.V. All rights reserved.)

As climate change progresses, the frequency and duration of drought stress events are increasing. While the mechanisms of drought acclimation of trees has received considerable attention in recent years, the recovery processes remain critically understudied. We used a unique throughfall exclusion experiment in a mature temperate mixed forest consisting of the more isohydric Norway spruce and more anisohydric European beech, to study the recovery and resilience after drought release. We hypothesized that pre-dawn water potential (ΨPD) of both species will increase within 1 day after watering, while the recovery of stomatal conductance (gs) and the reversal of osmoregulation will be significantly delayed in the more isohydric spruce. Furthermore, we hypothesized that the xylem sap flow density (udaily) will not fully recover within the growing season due to the strong drought impact. After 5 years of summer drought, trees showed significantly reduced ΨPD, udaily and increased osmoregulation in leaves, but only isohydric spruce displayed increased leaf abscisic acid concentrations. In line with our hypothesis, ΨPD and gs recovered within 1 day in beech. Conversely, isohydric spruce showed delayed increases in ΨPD and gs. The delay in recovery of spruce was partially related to the replenishment of the stem water reservoir, as indicated by the missing response of udaily at the crown base compared with DBH level upon watering. However, udaily fully recovered only in the next growing season for beech and was still reduced in spruce. Nevertheless, in both species, osmotic acclimations of leaves were reversed within several weeks. While both species displayed full resilience to drought stress in water-related physiology, the recovery time was in several cases, e.g., udaily, ΨPD and gs, shorter for beech than for spruce. With future increases in the frequency of drought events under ongoing climate change, tree species that recover more quickly will be favored., (© The Author(s) 2022. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

Against the backdrop of global change, the intensity, duration, and frequency of droughts are projected to increase and threaten forest ecosystems worldwide. Tree responses to drought are complex and likely to vary among species, drought characteristics, and site conditions. Here, we examined the drought response patterns of three major temperate tree species, s. fir (Abies alba), E. beech (Fagus sylvatica), and N. spruce (Picea abies), along an ecological gradient in the South - Central - East part of Germany that included a total of 37 sites with varying climatic and soil conditions. We relied on annual tree-ring data to assess the influence of different drought characteristics and (micro-) site conditions on components of tree resilience and to detect associated temporal changes. Our study revealed that nutrient regime, drought frequency, and hydraulic conditions in the previous and subsequent years were the main determinants of drought responses, with pronounced differences among species. Specifically, we found that (a) higher drought frequency was associated with higher resistance and resilience for N. spruce and E. beech; (b) more favorable climatic conditions in the two preceding and following years increased drought resilience and determined recovery potential of E. beech after extreme drought; (c) a site's nutrient regime, rather than micro-site differences in water availability, determined drought responses, with trees growing on sites with a balanced nutrient regime having a higher capacity to withstand extreme drought stress; (d) E. beech and N. spruce experienced a long-term decline in resilience. Our results indicate that trees under extreme drought stress benefit from a balanced nutrient supply and highlight the relevance of water availability immediately after droughts. Observed long-term trends confirm that N. spruce is suffering from persistent climatic changes, while s. fir is coping better. These findings might be especially relevant for monitoring, scenario analyses, and forest ecosystem management., Competing Interests: Declaration of competing interest None declared., (Copyright © 2023 The Authors. Published by Elsevier B.V. All rights reserved.)

Recent ecological research suggests that, in general, mixtures are more resistant to insect herbivores and pathogens than monocultures. However, we know little about mixtures with non-native trees, where enemy release could lead to patterns that differ from commonly observed relationships among native species. This becomes particularly relevant when considering that adaptation strategies to climate change increasingly promote a larger share of non-native tree species, such as North American Douglas fir in Central Europe. We studied leaf damage on European beech (Fagus sylvatica) saplings and mature trees across a wide range of site conditions in monocultures and mixtures with phylogenetically distant conifers native Norway spruce (Picea abies) and non-native Douglas fir (Pseudotsuga menziesii). We analyzed leaf herbivory and pathogen damage in relation to tree diversity and composition effects, as well as effects of environmental factors and plant characteristics. We observed lower sapling herbivory and tree sucking damage on beech in non-native Douglas fir mixtures than in beech monocultures, probably due to a lower herbivore diversity on Douglas fir trees, and higher pathogen damage on beech saplings in Norway spruce than Douglas fir mixtures, possibly because of higher canopy openness. Our findings suggest that for low diversity gradients, tree diversity effects on leaf damage can strongly depend on tree species composition, in addition to modifications caused by feeding guild and tree ontogeny. Moreover, we found that nutrient capacity modulated the effects of tree diversity, composition, and environmental factors, with different responses in sites with low or high nutrient capacity. The existence of contrasting diversity effects based on tree species composition provides important information on our understanding of the relationships between tree diversity and plant-herbivore interactions in light of non-native tree species introductions. Especially with recent Norway spruce die-off, the planting of Douglas fir as replacement is likely to strongly increase in Central Europe. Our findings suggest that mixtures with Douglas fir could benefit the survival or growth rates of beech saplings and mature trees due to lower leaf damage, emphasizing the need to clearly identify and compare the potential benefits and ecological trade-offs of non-native tree species in forest management under ongoing environmental change., (© 2022 The Authors. Ecological Applications published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

In a world of accelerating changes in environmental conditions driving tree growth, tradeoffs between tree growth rate and longevity could curtail the abundance of large old trees (LOTs), with potentially dire consequences for biodiversity and carbon storage. However, the influence of tree-level tradeoffs on forest structure at landscape scales will also depend on disturbances, which shape tree size and age distribution, and on whether LOTs can benefit from improved growing conditions due to climate warming. We analyzed temporal and spatial variation in radial growth patterns from ~5000 Norway spruce (Picea abies [L.] H. Karst) live and dead trees from the Western Carpathian primary spruce forest stands. We applied mixed-linear modeling to quantify the importance of LOT growth histories and stand dynamics (i.e., competition and disturbance factors) on lifespan. Finally, we assessed regional synchronization in radial growth variability over the 20th century, and modeled the effects of stand dynamics and climate on LOTs recent growth trends. Tree age varied considerably among forest stands, implying an important role of disturbance as an age constraint. Slow juvenile growth and longer period of suppressed growth prolonged tree lifespan, while increasing disturbance severity and shorter time since last disturbance decreased it. The highest age was not achieved only by trees with continuous slow growth, but those with slow juvenile growth followed by subsequent growth releases. Growth trend analysis demonstrated an increase in absolute growth rates in response to climate warming, with late summer temperatures driving the recent growth trend. Contrary to our expectation that LOTs would eventually exhibit declining growth rates, the oldest LOTs (>400 years) continuously increase growth throughout their lives, indicating a high phenotypic plasticity of LOTs for increasing biomass, and a strong carbon sink role of primary spruce forests under rising temperatures, intensifying droughts, and increasing bark beetle outbreaks., (© 2022 John Wiley & Sons Ltd.)

Ozone (O 3 ) enters plants through the stomata, passes into the intercellular air space and is decomposed in cell walls. Two factors that affect the O 3 level in the intercellular air space are the stomatal conductance and the concentration of biogenic volatile organic compounds (BVOCs). Stomatal conductance controls the O 3 flux into the air space and the intercellular BVOCs react with the O 3 . Therefore, the intercellular air space serves as a place where O 3 defense can occur, but it has received relatively little attention. This study aimed to explore potential plant-defense against O 3 in the intercellular air space by measuring the stomatal conductance and intercellular BVOC concentrations of Scots pine (Pinus sylvestris) and Norway spruce (Picea abies). Seedlings of both species were exposed to two levels of O 3 , ambient (15 nmol mol -1 ) and 80 nmol mol -1 (dropping to an ambient level at night), in plant growth chambers for five days in both spring and summer. We found that O 3 decreased stomatal conductance in both species and in both seasons, which can lower the O 3 flux into the intercellular air space. Intercellular BVOC concentrations were decreased in spring while increased in summer for both species in response to O 3 . This suggests that the BVOC protection in the intercellular air space is only of consequence in summer. These results demonstrate the potential for BVOCs to provide intercellular O 3 defense in both species, but with seasonal variation., 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 © 2022 The Authors. Published by Elsevier B.V. All rights reserved.)

Under climate change circumstances, increasing studies have reported the temporal instability of tree growth responses to climate, which poses a major challenge to linearly extrapolating past climate and future growth dynamics using tree-ring data. Space-for-time substitution (SFTS) is a potential solution to this problem that is widely used in the dendrochronology field to project past or future temporal growth response trajectories from contemporary spatial patterns. However, the projected accuracy of the SFTS in the climate effects on tree growth remains uncertain. Here, we empirically test the SFTS method by comparing the effect of spatial and temporal climate variations on climate responses of white spruce (Picea glauca), which has a transcontinental range in North America. We first applied a response surface regression model to capture the variations in growth responses along the spatial climate gradients. The results showed that the relationships between growth and June temperature varied along spatial climate gradients in a predictable way. And their relationships varied mainly along with local temperate condition. Then, the projected correlation coefficients between growth and climate using SFTS were compared against the observed. We found that the growth response changes caused by spatial versus temporal climate variations showed opposite trends. Moreover, the projected correlation coefficients using the SFTS were significantly lower than the observed. This finding suggests that applying the SFTS to project the growth response of white spruce might lead to an overestimation of the degree of tree maladaptation in future climate scenarios. And the overestimation is likely to get weaker from Alaska and Yukon Territory in the west to Quebec in the east. Although this is only a case study of the SFTS method for projecting tree growth response, our findings suggest that direct application of the SFTS method may not be applicable to all regions and all tree species., (© 2022 John Wiley & Sons Ltd.)

Despite recent advances in our understanding of drought impacts on tree functioning, we lack knowledge about the dynamic responses of mature trees to recurrent drought stress. At a subalpine forest site, we assessed the effects of three years of recurrent experimental summer drought on tree growth and water relations of Larix decidua Mill. and Picea abies (L. Karst.), two common European conifers representative for contrasting water-use strategies. We combined dendrometer and xylem sap flow measurements with analyses of xylem anatomy and non-structural carbohydrates and their carbon-isotope composition. Recurrent drought increased the effects of soil moisture limitation on growth and xylogenesis, and to a lesser extent on xylem sap flow. P. abies showed stronger growth responses to recurrent drought, reduced starch concentrations in branches and increased water-use efficiency when compared to L. decidua. Despite comparatively larger maximum tree water deficits than in P. abies, xylem formation of L. decidua was less affected by drought, suggesting a stronger capacity of rehydration or lower cambial turgor thresholds for growth. Our study shows that recurrent drought progressively increases impacts on mature trees of both species, which suggests that in a future climate increasing drought frequency could impose strong legacies on carbon and water dynamics of treeline species., (© 2022 The Authors. Plant, Cell & Environment published by John Wiley & Sons Ltd.)

Norway spruce (Picea abies (L.) Karst) trees are affected by ongoing climate change, including warming and exposure to phytotoxic levels of ozone. Non-volatile terpenoids and volatile terpenoids (biogenic organic volatile compounds, BVOCs) protect spruce against biotic and abiotic stresses. BVOCs also affect the atmosphere's oxidative capacity. Four-year-old Norway spruce were exposed to elevated ozone (EO) (1.4 × ambient) and warming (1.1 °C + ambient air) alone and in combination on an open-field exposure site in Central Finland. Net photosynthesis, needle terpenoid concentrations and BVOC emissions were measured four times during the experiment's second growing season: after bud opening in May, during the mid-growing season in June, and after needle maturation in August and September. Warming increased terpene concentrations in May due to advanced phenology and decreased them at the end of the growing season in matured current-year needles. Ozone enhanced these effects of warming on several compounds. Warming decreased concentrations of oxygenated sesquiterpenes in previous-year needles. Decreased emissions of oxygenated monoterpenes by warming and ozone alone in May were less prominent when ozone and warming were combined. A similar interactive treatment response in isoprene, camphene, tricyclene and α-pinene was observed in August when the temperature and ozone concentration was high. The results suggest long-term warming may reduce the terpenoid-based defence capacity of young spruce, but the defence capacity can be increased during the most sensitive growth phase (after bud break), and when high temperatures or ozone concentrations co-occur. Reduced BVOC emissions from young spruce may decrease the atmosphere's oxidative capacity in the warmer future, but the effect of EO may be marginal because less reactive minor compounds are affected., (© The Author(s) 2022. Published by Oxford University Press.)

The composition of ecologically important moss-associated bacterial communities seems to be mainly driven by host species but may also be shaped by environmental conditions related with tree dominance. The moss phyllosphere has been studied in coniferous forests while broadleaf forests remain understudied. To determine if host species or environmental conditions defined by tree dominance drives the bacterial diversity in the moss phyllosphere, we used 16S rRNA gene amplicon sequencing to quantify changes in bacterial communities as a function of host species (Pleurozium schreberi and Ptilium crista-castrensis) and forest type (coniferous black spruce versus deciduous broadleaf trembling aspen) in eastern Canada. The overall composition of moss phyllosphere was defined by the interaction of both factors, though most of the bacterial phyla were determined by a strong effect of forest type. Bacterial α-diversity was highest in spruce forests, while there was greater turnover (β-diversity) and higher γ-diversity in aspen forests. Unexpectedly, Cyanobacteria were much more relatively abundant in aspen than in spruce forests, with the cyanobacteria family Nostocaceae differing the most between forest types. Our results advance the understanding of moss-associated microbial communities among coniferous and broadleaf deciduous forests, which are important with the increasing changes in tree dominance in the boreal system., (© 2022 Society for Applied Microbiology and John Wiley & Sons Ltd.)

Pathogenic diseases frequently occur in drought-stressed trees. However, their contribution to the process of drought-induced mortality is poorly understood. We combined drought and stem inoculation treatments to study the physiological processes leading to drought-induced mortality in Norway spruce (Picea abies) and Scots pine (Pinus sylvestris) saplings infected with Heterobasidion annosum s.s. We analysed the saplings' water status, gas exchange, nonstructural carbohydrates (NSCs) and defence responses, and how they related to mortality. Saplings were followed for two growing seasons, including an artificially induced 3-month dormancy period. The combined drought and pathogen treatment significantly increased spruce mortality; however, no interaction between these stressors was observed in pine, although individually each stressor caused mortality. Our results suggest that pathogen infection decreased carbon reserves in spruce, reducing the capacity of saplings to cope with drought, resulting in increased mortality rates. Defoliation, relative water content and the starch concentration of needles were predictors of mortality in both species under drought and pathogen infection. Infection and drought stress create conflicting needs for carbon to compartmentalize the pathogen and to avoid turgor loss, respectively. Heterobasidion annosum reduces the functional sapwood area and shifts NSC allocation patterns, reducing the capacity of trees to cope with drought., (© 2022 The Authors. Plant, Cell & Environment published by John Wiley & Sons Ltd.)

Moss plays an important role in boreal forest ecosystems as an understory bryophyte species. Clearcut harvesting is a common boreal forest regeneration method that can expose understory vegetation to abiotic stressors impeding their recovery following post-harvest conditions. Very little is known concerning how moss remodel their chloroplast lipidome to enhance photosynthetic performance for successful acclimation to light and water stress during boreal forest regeneration following clearcut harvesting. The chloroplast lipidome and photosynthetic performance of Sphagnum sp. and three feathermoss species (Pleurozium schreberi, Hylocomium splendens, and Ptilium crista-castrensis) from a boreal black spruce (Picea mariana) forest were assessed using liquid chromatography-mass spectrometry (LC-MS), photospectrometry, and light response curves. We observed an overall increase in monogalactosyldiacylglycerol (MGDG) and sulfoquinovosyldiacylglycerol (SQDG) and decrease in digalactosyldiacylglycerol (DGDG) and phosphatidylglycerol (PG). In addition, unsaturation of the chloroplast lipidome occurred concomitant with photoprotection by carotenoid pigments to enhance the efficiency and photosynthetic capacity in moss exposed to light and water stress following clearcut harvesting. This appears to be a successful acclimation strategy used by moss to circumvent light stress during boreal forest regeneration following clearcut harvesting. These findings could be of significance in the development of boreal forest management strategies following resource harvesting., (Copyright © 2022 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Recent climate projections predict a more rapid increase of winter temperature than summer and global temperature averages in temperate and cold environments. As there is relatively little experimental knowledge on the effect of winter warming on cambium phenology and stem growth in species growing in cold environments, the setting of manipulative experiments is considered of primary importance, and they can help to decipher the effect of reduced winter chilling and increased forcing temperatures on cambium reactivation, growth and xylem traits. In this study, localized stem heating was applied to investigate the effect of warming from the rest to the growth phase on cambium phenology, intra-annual stem growth dynamics and ring wood features in Picea abies (L.) H.Karst. We hypothesized that reduced winter chilling induces a postponed cambium dormancy release and decrease of stem growth, while high temperature during cell wall lignification determines an enrichment of latewood-like cells. The heating device was designed to maintain a +5 °C temperature delta with respect to air temperature, thus allowing an authentic scenario of warming. Continuous stem heating from the rest (November) to the growing phase determined, at the beginning of radial growth, a reduction of the number of cell layers in the cambium, higher number of cell layers in the wall thickening phase and an asynchronous stem radial growth when comparing heated and ambient saplings. Nevertheless, heating did not induce changes in the number of produced cell layers at the end of the growing season. The analyses of two-photon fluorescence images showed that woody rings formed during heating were enriched with latewood-like cells. Our results showed that an increase of 5 °C of temperature applied to the stem from the rest to growth might not influence, as generally reported, onset of cambial activity, but it could affect xylem morphology of Norway spruce in mountain environments., (© The Author(s) 2021. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Norway spruce (Picea abies) is an economically and ecologically important tree species that grows across northern and central Europe. Treating Norway spruce with jasmonate has long-lasting beneficial effects on tree resistance to damaging pests, such as the European spruce bark beetle Ips typographus and its fungal associates. The (epi)genetic mechanisms involved in such long-lasting jasmonate induced resistance (IR) have gained much recent interest but remain largely unknown. In this study, we treated 2-year-old spruce seedlings with methyl jasmonate (MeJA) and challenged them with the I. typographus vectored necrotrophic fungus Grosmannia penicillata. MeJA treatment reduced the extent of necrotic lesions in the bark 8 weeks after infection and thus elicited long-term IR against the fungus. The transcriptional response of spruce bark to MeJA treatment was analysed over a 4-week time course using mRNA-seq. This analysis provided evidence that MeJA treatment induced a transient upregulation of jasmonic acid, salicylic acid and ethylene biosynthesis genes and downstream signalling genes. Our data also suggests that defence-related genes are induced while genes related to growth are repressed by methyl jasmonate treatment. These results provide new clues about the potential underpinning mechanisms and costs associated with long-term MeJA-IR in Norway spruce., (© 2022 The Authors. Plant, Cell & Environment published by John Wiley & Sons Ltd.)

The boreal forest represents the terrestrial biome most heavily affected by climate change. However, no consensus exists regarding the impacts of these changes on the growth of tree species therein. Moreover, assessments of young tree responses in metrics transposable to forest management remain scarce. Here, we assessed the impacts of climate change on black spruce (Picea mariana [Miller] BSP) and jack pine (Pinus banksiana Lambert) growth, two dominant tree species in boreal forests of North America. Starting with a retrospective analysis including data from 2591 black spruces and 890 jack pines, we forecasted trends in 30-year height growth at the transitions from closed to open boreal coniferous forests in Québec, Canada. We considered three variables: (1) height growth, rarely used, but better-reflecting site potential than other growth proxies, (2) climate normals corresponding to the growth period of each stem, and (3) site type (as a function of texture, stoniness, and drainage), which can modify the effects of climate on tree growth. We found a positive effect of vapor pressure deficit on the growth of both species, although the effect on black spruce leveled off. For black spruce, temperatures had a positive effect on the height at 30 years, which was attenuated when and where climatic conditions became drier. Conversely, drought had a positive effect on height under cold conditions and a negative effect under warm conditions. Spruce growth was also better on mesic than on rocky and sub-hydric sites. For portions of the study areas with projected future climate within the calibration range, median height-change varied from 10 to 31% for black spruce and from 5 to 31% for jack pine, depending on the period and climate scenario. As projected increases are relatively small, they may not be sufficient to compensate for potential increases in future disturbances like forest fires., (© 2021 John Wiley & Sons Ltd.)

Climate change in the conifer-dominated boreal forest is expected to lead to warmer but more dynamic winter air temperatures, reducing the depth and duration of snow cover and lowering winter soil temperatures. To gain insight into the mechanisms that have enabled conifers to dominate extreme cold environments, we performed genome-wide RNA-Seq analysis from needles and roots of non-dormant two-year Norway spruce (Picea abies (L.) H. Karst), and contrasted these response to herbaceous model Arabidopsis We show that the main transcriptional response of Norway spruce needles exposed to cold was delayed relative to Arabidopsis, and this delay was associated with slower development of freezing tolerance. Despite this difference in timing, Norway spruce principally utilizes early response transcription factors (TFs) belonging to the same gene families as Arabidopsis, indicating broad evolutionary conservation of cold response networks. In keeping with their different metabolic and developmental states, needles and root of Norway spruce showed contrasting results. Regulatory network analysis identified both conserved TFs with known roles in cold acclimation (e.g. homologs of ICE1, AKS3, and of the NAC and AP2/ERF superfamilies), but also a root-specific bHLH101 homolog, providing functional insights into cold stress response strategies in Norway spruce., (© 2021 The Authors. Plant, Cell & Environment published by John Wiley & Sons Ltd.)

Increasing seawater exposure is causing mortality of coastal forests, yet the physiological response associated with seawater-induced tree mortality, particularly in non-halophytes, is poorly understood. We investigated the shifts in carbon and nitrogen (N) metabolism of mature Sitka-spruce trees that were dying after an ecosystem-scale manipulation of tidal seawater exposure. Soil porewater salinity and foliar ion concentrations increased after seawater exposure and were strongly correlated with the percentage of live foliated crown (PLFC; e.g., crown 'greenness', a measure of progression to death). Co-occurring with decreasing PLFC was decreasing photosynthetic capacity, N-investment into photosynthesis, N-resorption efficiency and non-structural carbohydrate (soluble sugars and starch) concentrations, with the starch reserves depleted to near zero when PLFC dropped below 5%. Combined with declining PLFC, these changes subsequently decreased total carbon gain and thus exacerbated the carbon starvation process. This study suggests that an impairment in carbon and N metabolism during the mortality process after seawater exposure is associated with the process of carbon starvation, and provides critical knowledge necessary to predict sea-level rise impacts on coastal forests., (© The Author(s) 2021. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

Sea-level rise is one of the most critical challenges facing coastal ecosystems under climate change. Observations of elevated tree mortality in global coastal forests are increasing, but important knowledge gaps persist concerning the mechanism of salinity stress-induced nonhalophytic tree mortality. We monitored progressive mortality and associated gas exchange and hydraulic shifts in Sitka-spruce (Picea sitchensis) trees located within a salinity gradient under an ecosystem-scale change of seawater exposure in Washington State, USA. Percentage of live foliated crown (PLFC) decreased and tree mortality increased with increasing soil salinity during the study period. A strong reduction in gas exchange and xylem hydraulic conductivity (Ks) occurred during tree death, with an increase in the percentage loss of conductivity (PLC) and turgor loss point (πtlp). Hydraulic and osmotic shifts reflected that hydraulic function declined from seawater exposure, and dying trees were unable to support osmotic adjustment. Constrained gas exchange was strongly related to hydraulic damage at both stem and leaf levels. Significant correlations between foliar sodium (Na+) concentration and gas exchange and key hydraulic parameters (Ks, PLC, and πtlp) suggest that cellular injury related to the toxic effects of ion accumulation impacted the physiology of these dying trees. This study provides evidence of toxic effects on the cellular function that manifests in all aspects of plant functioning, leading to unfavourable osmotic and hydraulic conditions., (© American Society of Plant Biologists 2021. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

This study developed somatic embryogenesis protocols for Picea pungens (Engelm), an important ornamental species, including initiation, proliferation, maturation, germination, and acclimation. Somatic embryogenic tissues were induced from mature zygotic embryos of five families, with a frequency of [Formula: see text] 22% for each. Embryogenic tissues (ET) from 13 clones of three families were proliferated for one week, achieving an average rate of 179.1%. The ET of 38 clones of three families were cultured in maturation medium for six weeks; 188 mature embryos on average were counted per gram ET cultured, of which [Formula: see text] 81.1% appeared normal, and each clone developed at least 28 normally matured embryos. A total of 69.9% or more of cotyledonary somatic embryos germinated normally and developed into normal emblings. The experiment of transplanting the emblings into a greenhouse had an average survival rate of 68.5%. Considerable variation among and within families during initiation and proliferation was observed, but this variation decreased in the maturation and germination. Changing the concentration of plant growth regulator of the initiation medium did not significantly change the initiation frequency. We recommend incorporating these protocols into the current Picea pungens practical programs, although further research is essential to increase efficiencies and reduce cost., (© 2021. The Author(s).)

Picea mongolica, a native species with excellent industrial wood quality and strong sand-fixing capacity, may be utilized in construction of urban green spaces in arid areas in China. However, now the sustainability of the ecosystems where this species grows is at serious risk due to a lack of natural regeneration. In this study, we developed an efficient regeneration system and comprehensively analyzed various factors affecting somatic embryogenesis (SE) using zygotic embryos as explants. We identified the optimal plant growth regulators (PGRs) performance and the best donor trees (k81) for the generation of somatic embryos (SEMs). Additionally, we confirmed that the positive developmental window of SEMs initiation was at the end of July to early August, which is when zygotic embryos was at the late embryogeny. In this time period, specific transcripts associated with the regulation of epigenetic modifications, plant hormone-related genes, and embryonic development-related transcription factors play important roles for early SEMs initiation. These results may provide a valuable resource for vegetative propagation of Picea mongolica. Our results may help to establish a reliable protocol for plantlet regeneration, which may facilitate urban greening applications and industrialization in arid areas.

Fine roots (< 2 mm in diameter) play an important role in belowground ecosystem processes, and their physiological ecology is easily altered by nitrogen deposition. To better understand the response of physiological and ecological processes of fine roots to nitrogen deposition, a manipulation experiment was conducted to investigate the effects of exogenous nitrogen addition (control (0 kg ha -1  a -1 ), low (5 kg ha -1  a -1 ), moderate (10 kg ha -1  a -1 ), and high nitrogen (20 kg ha -1  a -1 )) on the biomass, morphological characteristics, chemical elements and nonstructural carbohydrates of fine roots in a Picea schrenkiana forest. We found that most fine roots were located in the 0-20 cm of soil layer across all nitrogen treatment groups (42.81-52.09% of the total biomass). Compared with the control, the biomass, specific root length and specific root area of the fine roots increased in the medium nitrogen treatment, whereas the fine roots biomass was lower in the high nitrogen treatment than in the other treatments. In fine roots, nitrogen addition promotes the absorption of nitrogen and phosphorus and their stoichiometric ratio, while reducing the content of nonstructural carbohydrates. The content of nonstructural carbohydrates in the small-diameter roots (< 1 mm in diamter) in each nitrogen treatment group was lower than that in the large-diameter roots. Correlation analysis showed that soil carbon and nitrogen were positively correlated with fine root biomass and specific root length and negatively correlated with the nonstructural carbohydrates. Our findings demonstrate that medium nitrogen addition is conducive to the development of fine root morphology, while excessive nitrogen can suppress the growth of root systems.

Norway spruce has a broad natural distribution range, which results in a substantial variety of its physiological and genetic variation. There are three distinct altitudinal ecotypes described in this tree species. The physiological optimum of each ecotype may be shifted due to ongoing climate change, especially in traits associated with water demand that might be crucial for adaptation. Dehydrins are proteins that help to mitigate the adverse effects of dehydration. Dehydrin gene expression patterns appeared to be a suitable marker for plant stress assessment. Genetically determined differences in response between individuals and populations were formerly studied, however, mainly in controlled conditions. We evaluated ecotypic variation in dehydrin gene expression in a clonal bank comprised of all three ecotypes. A genetic relationship among targeted trees was uncovered utilizing GBS (Genotyping by Sequencing) platform. We sampled 4-6 trees of each ecotype throughout 15 months period. Subsequently, we assessed the RNA expression of dehydrin genes by qRT-PCR. For this study, we deliberately selected dehydrins from different categories. Our findings detected significant differences among ecotypes in dehydrin expression. The association of recorded climatic variables and individual gene expression across the study period was evaluated and revealed, for certain genes, a correlation between dehydrin gene expression and precipitation, temperature, and day-length.