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Given growing concerns about global climate change, it is critical to understand both historical and current shifts in the hydroclimate, particularly in regions critically entwined with global circulation. The Tibetan Plateau, the Earth's largest and highest plateau, is a nexus for global atmospheric processes, significantly influencing East Asian hydroclimate dynamics through the synergy of the Asian Monsoon and the Westerlies. Yet, understanding historical and recent hydroclimate fluctuations and their wide-ranging ecological and societal consequences remains challenging due to short instrumental observations and partly ambiguous proxy reconstructions. Here, we present a precisely-dated 3476-year precipitation reconstruction derived from tree-ring δ 18 O data on the Tibetan Plateau, representing one of the few multi-millennia-long annually-resolved terrestrial δ 18 O records to date. Our findings reveal that the 20 th  century drought extremes are severe within the past three millennia, and likely linked to the weakening of both the Asian Monsoon and Westerlies due to anthropogenic aerosol emissions. Additionally, our analyses identified three distinct stages (110 BC-AD 280, AD 330-770 and AD 950-1300) characterized by shifts toward arid hydroclimate conditions, corresponding to significant social unrest and dynasty collapses, which underscores the potential societal impacts of severe hydroclimatic shifts., Competing Interests: Competing interests: The authors declare no competing interests., (© 2025. The Author(s).)

Oxygen (δ 18 O) and hydrogen (δ 2 H) stable isotope ratios are tightly coupled in precipitation and, albeit damped, in leaf water, but are often decoupled in tree-ring cellulose. The environmental and physiological conditions in which this decoupling occurs are not yet well understood. We investigated the relationships between δ 18 O and δ 2 H and tree-ring width (TRW), tree crown volume, tree age and climate in silver fir and Douglas-fir and found substantial differences between δ 18 O and δ 2 H. Overall, δ 18 O-δ 2 H correlations were weak to absent but became significantly negative under high summer vapour pressure deficit (VPD). δ 18 O and δ 2 H had positive and negative nonlinear relationships with TRW, respectively, with clear relationships at the site and tree levels for silver fir and, to a lesser extent, for Douglas-fir. Age trends for silver fir were weakly negative in δ 18 O but positive in δ 2 H. Tree crown volume and δ 18 O or δ 2 H had no significant relationships. Most strikingly, δ 18 O strongly depended on spring climate (precipitation and VPD), whereas δ 2 H depended on summer climate (temperature and VPD) for both species. Our study shows that the δ 18 O-δ 2 H decoupling in tree-ring cellulose in two temperate conifer species could be highlighted by their contrasting relationships to climate and tree intrinsic variables (TRW, age)., (© 2024 The Author(s). Plant, Cell & Environment published by John Wiley & Sons Ltd.)

Increases in temperatures and atmospheric CO2 concentration influence the growth performance of trees worldwide. The direction and intensity of tree growth and physiological responses to changing climate do, however, vary according to environmental conditions. Here we present complex, long-term, tree-physiological responses to unprecedented temperature increase in East Asia. For this purpose, we studied radial growth and isotopic (δ13C and δ18O) variations using tree-ring data for the past 100 yr of dominant Quercus mongolica trees from the cool-temperate forests from Hallasan, South Korea. Overall, we found that tree stem basal area increment, intercellular CO2 concentration and intrinsic water-use efficiency significantly increased over the last century. We observed, however, short-term variability in the trends of these variables among four periods identified by change point analysis. In comparison, δ18O did not show significant changes over time, suggesting no major hydrological changes in this precipitation-rich area. The strength and direction of growth-climate relationships also varied during the past 100 yr. Basal area increment (BAI) did not show significant relationships with the climate over the 1924-1949 and 1975-1999 periods. However, over 1950-1974, BAI was negatively affected by both temperature and precipitation, while after 2000, a temperature stimulus was observed. Finally, over the past two decades, the increase in Q. mongolica tree growth accelerated and was associated with high spring-summer temperatures and atmospheric CO2 concentrations and decreasing intrinsic water-use efficiency, δ18O and vapour pressure deficit, suggesting that the photosynthetic rate continued increasing under no water limitations. Our results indicate that the performance of dominant trees of one of the most widely distributed species in East Asia has benefited from recent global changes, mainly over the past two decades. Such findings are essential for projections of forest dynamics and carbon sequestration under climate change., (© The Author(s) 2024. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

Disentangling the factors influencing the climate sensitivity of trees is crucial to understanding the susceptibility of forests to climate change. Reducing tree-to-tree competition and mixing tree species are two strategies often promoted to reduce the drought sensitivity of trees, but it is unclear how effective these measures are in different ecosystems. Here, we studied the growth and physiological responses to climate and severe droughts of silver fir and Douglas-fir growing in pure and mixed conditions at three sites in Switzerland. We used tree-ring width data and carbon (δ13C), oxygen (δ18O) and hydrogen (δ2H) stable isotope ratios from tree-ring cellulose to gain novel information on water relations and the physiology of trees in response to drought and how tree species mixture and competition modulate these responses. We found significant differences in isotope ratios between trees growing in pure and mixed conditions for the two species, although these differences varied between sites, e.g. trees growing in mixed conditions had higher δ13C values and tree-ring width than trees growing in pure conditions for two of the sites. For both species, differences between trees in pure and mixed conditions regarding their sensitivity to temperature, precipitation, climatic water balance and vapor pressure deficit were minor. Furthermore, trees growing in pure and mixed conditions showed similar responses of tree-ring width and isotope ratios to the past severe droughts of 2003, 2015 and 2018. Competition had only a significantly negative effect on δ13C of silver fir, which may suggest a decrease in photosynthesis due to higher competition for light and nutrients. Our study highlights that tree species mixture may have only moderate effects on the radial growth and physiological responses of silver fir and Douglas-fir to climatic conditions and that site condition effects may dominate over mixture effects., (© The Author(s) 2024. Published by Oxford University Press.)

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

Understanding how trees prioritize carbon gain at the cost of drought vulnerability under severe drought conditions is crucial for predicting which genetic groups and individuals will be resilient to future climate conditions. In this study, we investigated variations in growth, tree-ring anatomy as well as carbon and oxygen isotope ratios to assess the sensitivity and the xylem formation process in response to an episode of severe drought in 29 mature white spruce (Picea glauca [Moench] Voss) families grown in a common garden trial. During the drought episode, the majority of families displayed decreased growth and exhibited either sustained or increased intrinsic water-use efficiency (iWUE), which was largely influenced by reduced stomatal conductance as revealed by the dual carbon‑oxygen isotope approach. Different water-use strategies were detected within white spruce populations in response to drought conditions. Our results revealed intraspecific variation in the prevailing physiological mechanisms underlying drought response within and among populations of Picea glauca. The presence of different genetic groups reflecting diverse water-use strategies within this largely-distributed conifer is likely to lessen the negative effects of drought and decrease the overall forest ecosystems' sensitivity to it., Competing Interests: Declaration of competing interest The authors have no conflicts of interest to declare that are relevant to the content of this article., (Copyright © 2024 The Author(s). Published by Elsevier B.V. All rights reserved.)

The oxygen isotope composition (δ 18 O) of tree-ring cellulose is used to evaluate tree physiological responses to climate, but their interpretation is still limited due to the complexity of the isotope fractionation pathways. We assessed the relative contribution of seasonal needle and xylem water δ 18 O variations to the intra-annual tree-ring cellulose δ 18 O signature of larch trees at two sites with contrasting soil water availability in the Swiss Alps. We combined biweekly δ 18 O measurements of soil water, needle water, and twig xylem water with intra-annual δ 18 O measurements of tree-ring cellulose, xylogenesis analysis, and mechanistic and structural equation modeling. Intra-annual cellulose δ 18 O values resembled source water δ 18 O mean levels better than needle water δ 18 O. Large parts of the rings were formed under high proportional exchange with unenriched xylem water (p ex ). Maximum p ex values were achieved in August and imprinted on sections at 50-75% of the ring. High p ex values were associated with periods of high atmospheric evaporative demand (VPD). While VPD governed needle water δ 18 O variability, we estimated a limited Péclet effect at both sites. Due to a variable p ex , source water has a strong influence over large parts of the intra-annual tree-ring cellulose δ 18 O variations, potentially masking signals coming from needle-level processes., (© 2023 The Authors. New Phytologist © 2023 New Phytologist Foundation.)

Stable isotope ratio analysis of tree rings has been widely and successfully applied in recent decades for climatic and environmental reconstructions. These studies were mostly conducted at an annual resolution, considering one measurement per tree ring, often focusing on latewood. However, much more information could be retrieved with high-resolution intra-annual isotope studies, based on the fact that the wood cells and the corresponding organic matter are continuously laid down during the growing season. Such studies are still relatively rare, but have a unique potential for reconstructing seasonal climate variations or short-term changes in physiological plant properties, like water-use efficiency. The reason for this research gap is mostly technical, as on the one hand sub-annual, manual splitting of rings is very tedious, while on the other hand automated laser ablation for high-resolution analyses is not yet well established and available. Here, we give an update on the current status of laser ablation research for analysis of the carbon isotope ratio (δ13C) of wood, describe an easy-to-use laser ablation system, its operation and discuss practical issues related to tree core preparation, including cellulose extraction. The results show that routine analysis with up to 100 laser shot-derived δ13C-values daily and good precision and accuracy (ca. 0.1‰) comparable to conventional combustion in an elemental analyzer are possible. Measurements on resin-extracted wood is recommended as most efficient, but laser ablation is also possible on cellulose extracted wood pieces. Considering the straightforward sample preparation, the technique is therefore ripe for wide-spread application. With this work, we hope to stimulate future progress in the promising field of high-resolution environmental reconstruction using laser ablation., (© The Author(s) 2022. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Recent experiments have underlined the potential of δ2H in tree-ring cellulose as a physiological indicator of shifts in autotrophic versus heterotrophic processes (i.e., the use of fresh versus stored non-structural carbohydrates). However, the impact of these processes has not yet been quantified under natural conditions. Defoliator outbreaks disrupt tree functioning and carbon assimilation, stimulating remobilization, therefore providing a unique opportunity to improve our understanding of changes in δ2H. By exploring a 700-year tree-ring isotope chronology from Switzerland, we assessed the impact of 79 larch budmoth (LBM, Zeiraphera griseana [Hübner]) outbreaks on the growth of its host tree species, Larix decidua [Mill]. The LBM outbreaks significantly altered the tree-ring isotopic signature, creating a 2H-enrichment and an 18O- and 13C-depletion. Changes in tree physiological functioning in outbreak years are shown by the decoupling of δ2H and δ18O (O-H relationship), in contrast to the positive correlation in non-outbreak years. Across the centuries, the O-H relationship in outbreak years was not significantly affected by temperature, indicating that non-climatic physiological processes dominate over climate in determining δ2H. We conclude that the combination of these isotopic parameters can serve as a metric for assessing changes in physiological mechanisms over time., (© The Author(s) 2023. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

Climate warming has profoundly altered the status of permafrost and has caused extensive permafrost degradation in the Northern Hemisphere. However, long-term observations investigating the hydrological dynamics of permafrost and its ecological effects on plant growth are lacking. Previous studies have reported tree-ring stable hydrogen isotope ratios of lignin methoxy groups (δ 2 H LM ) as an archive of hydrological signals. This study sampled tree-ring cores from a Larix gmelinii forest in Nanwenghe Forest Park, Northeastern China, and separately measured the tree-ring δ 2 H LM for earlywood and latewood from 1900 to 2020. Earlywood and latewood δ 2 H LM values, as well as the difference between them, showed no significant long-term trend from 1900 to 1987; however, they both exhibited significant increasing trends since 1988 at rates of 2.6 ‰ and 4.9 ‰ per decade, respectively. This variance changes the magnitude of the difference between the two chronologies and can be explained by the shift in source water δ 2 H values during tree growth. Based on a structural equation model analysis, when the influence of permafrost melting weakened due to permafrost degradation, the growing season temperature was better recorded in latewood δ 2 H LM through the effects of precipitation δ 2 H from July to September. Based on the environmental response of tree-ring δ 2 H LM in the permafrost region, permafrost degradation influences the source water δ 2 H values of trees, thereby affecting the expression of temperature signals in tree-ring δ 2 H LM . The novel results in this study provide a new perspective on permafrost degradation based on the dynamic responses of tree-ring δ 2 H LM to source water δ 2 H during permafrost degradation., 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 Elsevier B.V. All rights reserved.)

The impact of climate extremes on forest ecosystems is poorly understood but important for predicting carbon and water cycle feedbacks to climate. Some knowledge gaps still remain regarding how drought-related adjustments in intra-annual tree-ring characteristics directly impact tree carbon and water use. In this study we quantified the impact of an extreme summer drought on the water-use efficiency and carbon sequestration of four mature Norway spruce trees. We used detailed observations of wood formation (xylogenesis) and intra-annual tree-ring properties (quantitative wood anatomy and stable carbon isotopes) combined with physiological water-stress monitoring. During 41 d of tree water deficit, we observed an enrichment in 13 C but a reduction in cell enlargement and wall-thickening processes, which impacted the anatomical characteristics. These adjustments diminished carbon sequestration by 67% despite an 11% increase in water-use efficiency during drought. However, with the resumption of a positive hydric state in the stem, we observed a fast recovery of cell formation rates based on the accumulated assimilates produced during drought. Our findings enhance our understanding of carbon and water fluxes between the atmosphere and forest ecosystems, providing observational evidence on the tree intra-annual carbon sequestration and water-use efficiency dynamics to improve future generations of vegetation models., (© 2022 The Authors. New Phytologist © 2022 New Phytologist Foundation.)

Climate controls forest biomass production through direct effects on cambial activity and indirectly through interactions with CO 2 , air pollution, and nutrient availability. The atmospheric concentration of CO 2 , sulfur and nitrogen deposition can also exert a significant indirect control on wood formation since these factors influence the stomatal regulation of transpiration and carbon uptake, that is, intrinsic water use efficiency (iWUE). Here we provide 120-year long tree-ring time series of iWUE, stem growth, climatic and combined sulfur and nitrogen (SN) deposition trends for two common tree species, Pinus sylvestris (PISY) and Picea abies (PCAB), at their lower and upper distribution margins in Central Europe. The main goals were to explain iWUE trends using theoretical scenarios including climatic and SN deposition data, and to assess the contribution of climate and iWUE to the observed growth trends. Our results showed that after a notable increase in iWUE between the 1950s and 1980s, this positive trend subsequently slowed down. The substantial rise of iWUE since the 1950s resulted from a combination of an accelerated increase in atmospheric CO 2 concentrations (C a ) and a stable level of leaf intercellular CO 2 (C i ). The offset of observed iWUE values above the trajectory of a constant C i /C a scenario was explained by trends in SN deposition (all sites) together with the variation of drought conditions (low-elevation sites only). Increasing iWUE over the 20th and 21st centuries improved tree growth at low-elevation drought-sensitive sites. In contrast, at high-elevation PCAB sites, growth was mainly stimulated by recent warming. We propose that SN pollution should be considered in order to explain the steep increase in iWUE of conifers in the 20th century throughout Central Europe and other regions with a significant SN deposition history., Competing Interests: Declaration of competing interest The authors have no conflicts of interest to disclose and have all approved this submission., (Copyright © 2022 Elsevier B.V. All rights reserved.)

The intensity and frequency of droughts events are projected to increase in future with expected adverse effects for forests. Thus, information on the dynamics of tree water uptake from different soil layers during and after drought is crucial. We applied an in situ water isotopologue monitoring system to determine the oxygen isotope composition in soil and xylem water of European beech with a 2-h resolution together with measurements of soil water content, transpiration and tree water deficit. Using a Bayesian isotope mixing model, we inferred the relative and absolute contribution of water from four different soil layers to tree water use. Beech took up more than 50% of its water from the uppermost 5 cm soil layer at the beginning of the 2018 drought, but then reduced absolute water uptake from the drying topsoil by 84%. The trees were not able to quantitatively compensate for restricted topsoil water availability by additional uptake from deeper soil layers, which is related to the fine root depth distribution. Absolute water uptake from the topsoil was restored to pre-drought levels within 3 wk after rewetting. These uptake patterns help to explain both the drought sensitivity of beech and its high recovery potential after drought release., (© 2021 The Authors. New Phytologist © 2021 New Phytologist Foundation.)

The capacity of trees to release water from storage compartments into the transpiration stream can mitigate damage to hydraulic functioning. However, the location of these 'transient' water sources and also the pathways of water movement other than vertical through tree stems still remain poorly understood. We conducted an experiment on two tree species in a common garden in eastern Australia that naturally grow in regions of high (Eucalyptus tereticornis, 'Red Gum') and low (Eucalyptus sideroxylon, 'Ironbark') annual precipitation rates. Deuterium-enriched water (1350% label strength) was directly introduced into the transpiration stream of three trees per species for four consecutive days. Subsequently, the trees were felled, woody tissue samples were collected from different heights and azimuthal positions of the stems, and stable isotope ratios were determined on the water extracted from all samples. The presence/absence of the tracer along the radial and vertical stem axes in combination with xylem hydraulic properties inferred from sapflow, leaf and stem water potentials, wood moisture contents and anatomical sapwood characteristics elucidated species-specific patterns of short-term stem water storage and movement. The distribution of water isotopes at natural abundance among woody tissues indicated systematic differences with highest values of sapwood water and lower values in inner bark and heartwood. Presence of tracer in water of the inner bark highlighted the importance of this tissue as capacitor. Although injected at the northern side of stems, tracer was also discovered at the southern side, providing empirical evidence for circumferential flow in sapwood, particularly of Ironbark. Greater vertical water transport in Red Gum compared with more radial and circumferential water transport in Ironbark were associated with species-specific sapwood anatomy. Our study highlights the value of combining information from stable isotope tracers and wood anatomy to investigate patterns of water transport and storage of tall trees in situ., (© The Author(s) 2021. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)