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Forests play a crucial role in global carbon cycling by absorbing and storing significant amounts of atmospheric carbon dioxide. Although boreal forests contribute to approximately 45% of the total forest carbon sink, tree growth and soil carbon sequestration are constrained by nutrient availability. Here, we examine if long-term nutrient input enhances tree productivity and whether this leads to carbon storage or whether stimulated microbial decomposition of organic matter limits soil carbon accumulation. Over six decades, nitrogen, phosphorus, and calcium were supplied to a Pinus sylvestris-dominated boreal forest. We found that nitrogen fertilization alone or together with calcium and/or phosphorus increased tree biomass production by 50% and soil carbon sequestration by 65% compared to unfertilized plots. However, the nonlinear relationship observed between tree productivity and soil carbon stock across treatments suggests microbial regulation. When phosphorus was co-applied with nitrogen, it acidified the soil, increased fungal biomass, altered microbial community composition, and enhanced biopolymer degradation capabilities. While no evidence of competition between ectomycorrhizal and saprotrophic fungi has been observed, key functional groups with the potential to reduce carbon stocks were identified. In contrast, when nitrogen was added without phosphorus, it increased soil carbon sequestration because microbial activity was likely limited by phosphorus availability. In conclusion, the addition of nitrogen to boreal forests may contribute to global warming mitigation, but this effect is context dependent., (© 2024 The Author(s). Global Change Biology published by John Wiley & Sons Ltd.)

The ability of plants to retain nitrogen (N) for a long period of time is critical to their N use efficiency, growth, and fitness, particularly in infertile environments. The mean residence time of leaf N (MRT L ) and its two determinants, leaf lifespan and N resorption efficiency (r N , the fraction of the total leaf N pool that is resorbed during leaf senescence), have been hypothesized to increase plastically with decreasing soil N fertility but this remains to be fully tested. To avoid confusion by random changes in these characteristics in a relatively narrow N fertility range, MRT L , leaf lifespan, and N resorption efficiency were measured in Quercus glauca over a broad N fertility range. In the high to moderate N fertility range, leaf lifespan and r N increased with decreasing N addition rate, and thus the MRT L increased. However, in the moderate to low N fertility range, leaf lifespan increased but r N decreased significantly, so MRT L decreased. The decrease in r N occurred because the senesced leaf N concentration was almost constant at the lower limit while the green leaf N concentration decreased in this range. The hump-shaped quadratic responses of MRT L and r N along the N fertility gradient suggest that incorrect conclusions about the response of these traits to N fertility variation may be drawn from experiments that include only a few fertility levels, and N recycling within leaf canopy alone cannot achieve efficient N use in infertile environments., (© 2024 The Author(s). Physiologia Plantarum published by John Wiley & Sons Ltd on behalf of Scandinavian Plant Physiology Society.)

Reserves of non-structural carbohydrates (NSC) stored in living cells are essential for drought tolerance of trees. However, little is known about the phenotypic plasticity of living storage compartments (SC) and their interactions with NSC reserves under changing water availability. Here, we examined adjustments of SC and NSC reserves in stems and roots of seedlings of two temperate tree species, Acer negundo L. and Betula pendula Roth., cultivated under different substrate water availability. We found that relative contents of soluble NSC, starch and total NSC increased with decreasing water availability in stems of both species, and similar tendencies were also observed in roots of A. negundo. In the roots of B. pendula, soluble NSC contents decreased along with the decreasing water availability, possibly due to phloem decoupling or NSC translocation to shoots. Despite the contrast in organ responses, NSC contents (namely starch) positively correlated with proportions of total organ SC. Individual types of SC showed markedly distinct plasticity upon decreasing water availability, suggesting that water availability changes the partitioning of organ storage capacity. We found an increasing contribution of parenchyma-rich bark to the total organ NSC storage capacity under decreasing water availability. However, xylem SC showed substantially greater plasticity than those in bark. Axial storage cells, namely living fibers in A. negundo, responded more sensitively to decreasing water availability than radial parenchyma. Our results demonstrate that drought-induced changes in carbon balance affect the organ storage capacity provided by living cells, whose proportions are sensitively coordinated along with changing NSC reserves., (© 2024 The Author(s). Physiologia Plantarum published by John Wiley & Sons Ltd on behalf of Scandinavian Plant Physiology Society.)

All ecosystems contain both sources and sinks for atmospheric carbon (C). A change in their balance of net and gross ecosystem carbon uptake, ecosystem-scale carbon use efficiency (CUE ECO ), is a change in their ability to buffer climate change. However, anthropogenic nitrogen (N) deposition is increasing N availability, potentially shifting terrestrial ecosystem stoichiometry towards phosphorus (P) limitation. Depending on how gross primary production (GPP, plants alone) and ecosystem respiration (R ECO , plants and heterotrophs) are limited by N, P or associated changes in other biogeochemical cycles, CUE ECO may change. Seasonally, CUE ECO also varies as the multiple processes that control GPP and respiration and their limitations shift in time. We worked in a Mediterranean tree-grass ecosystem (locally called 'dehesa') characterized by mild, wet winters and summer droughts. We examined CUE ECO from eddy covariance fluxes over 6 years under control, +N and + NP fertilized treatments on three timescales: annual, seasonal (determined by vegetation phenological phases) and 14-day aggregations. Finer aggregation allowed consideration of responses to specific patterns in vegetation activity and meteorological conditions. We predicted that CUE ECO should be increased by wetter conditions, and successively by N and NP fertilization. Milder and wetter years with proportionally longer growing seasons increased CUE ECO , as did N fertilization, regardless of whether P was added. Using a generalized additive model, whole ecosystem phenological status and water deficit indicators, which both varied with treatment, were the main determinants of 14-day differences in CUE ECO . The direction of water effects depended on the timescale considered and occurred alongside treatment-dependent water depletion. Overall, future regional trends of longer dry summers may push these systems towards lower CUE ECO ., (© 2024 The Author(s). Global Change Biology published by John Wiley & Sons Ltd.)

Plants express diverse nutrient use and acquisition traits, but it is unclear how trait combinations at the species level are constrained by phylogeny, trait coordination, or trade-offs in resource investment. One trait - nitrogen (N) fixation - is assumed to correlate with other traits and used to define plant functional groups, despite potential confounding effects of phylogeny. We quantified growth, carbon metabolism, fixation rate, root phosphatase activity (RPA), mycorrhizal colonization, and leaf and root morphology/chemistry across 22 species of fixing and nonfixing tropical Fabaceae trees under common conditions. Belowground trait variation was high even among closely related species, and most traits displayed a phylogenetic signal, including N-fixation rate and nodule biomass. Across species, we observed strong positive correlations between physiological traits such as RPA and root respiration. RPA increased ~ fourfold per unit increase in fixation, supporting the debated hypothesis that N-fixers 'trade' N for phosphatases to enhance phosphorus acquisition. Specific root length and root N differed between functional groups, though for other traits, apparent differences became nonsignificant after accounting for phylogenetic nonindependence. We conclude that evolutionary history, trait coordination, and fixation ability contribute to nutrient trait expression at the species level, and recommend explicitly considering phylogeny in analyses of functional groupings., (© 2024 The Author(s). New Phytologist © 2024 New Phytologist Foundation.)

Background: Leaf nitrogen (N) and phosphorus (P) resorption is a fundamental adaptation strategy for plant nutrient conservation. However, the relative roles that environmental factors and plant functional traits play in regulating N and P resorption remain largely unclear, and little is known about the underlying mechanism of plant functional traits affecting nutrient resorption. Here, we measured leaf N and P resorption and 13 plant functional traits of leaf, petiole, and twig for 101 representative broad-leaved tree species in our target subtropical transitional forests. We integrated these multiple functional traits into the plant economics spectrum (PES). We further explored whether and how elevation-related environmental factors and these functional traits collectively control leaf N and P resorption., Results: We found that deciduous and evergreen trees exhibited highly diversified PES strategies, tending to be acquisitive and conservative, respectively. The effects of PES, rather than of environmental factors, dominated leaf N and P resorption patterns along the elevational gradient. Specifically, the photosynthesis and nutrient recourse utilization axis positively affected N and P resorption for both deciduous and evergreen trees, whereas the structural and functional investment axis positively affected leaf N and P resorption for evergreen species only. Specific leaf area and green leaf nutrient concentrations were the most influential traits driving leaf N and P resorption., Conclusions: Our study simultaneously elucidated the relative contributions of environmental factors and plant functional traits to leaf N and P resorption by including more representative tree species than previous studies, expanding our understanding beyond the relatively well-studied tropical and temperate forests. We highlight that prioritizing the fundamental role of traits related to leaf resource capture and defense contributes to the monitoring and modeling of leaf nutrient resorption. Therefore, we need to integrate PES effects on leaf nutrient resorption into the current nutrient cycling model framework to better advance our general understanding of the consequences of shifting tree species composition for nutrient cycles across diverse forests., (© 2024. The Author(s).)

As major terrestrial carbon sinks, forests play an important role in mitigating climate change. The relationship between the seasonal uptake of carbon and its allocation to woody biomass remains poorly understood, leaving a significant gap in our capacity to predict carbon sequestration by forests. Here, we compare the intra-annual dynamics of carbon fluxes and wood formation across the Northern hemisphere, from carbon assimilation and the formation of non-structural carbon compounds to their incorporation in woody tissues. We show temporally coupled seasonal peaks of carbon assimilation (GPP) and wood cell differentiation, while the two processes are substantially decoupled during off-peak periods. Peaks of cambial activity occur substantially earlier compared to GPP, suggesting the buffer role of non-structural carbohydrates between the processes of carbon assimilation and allocation to wood. Our findings suggest that high-resolution seasonal data of ecosystem carbon fluxes, wood formation and the associated physiological processes may reduce uncertainties in carbon source-sink relationships at different spatial scales, from stand to ecosystem levels., (© 2024. The Author(s).)

Plantations actively participate in the global carbon cycle and play a significant role in mitigating global climate change. However, the influence of forest management strategies, especially planting density management, on the biomass carbon storage and production value of plantations for ensuring carbon sink benefits is still unclear. In this study, we estimated the carbon sequestration and economic value of Pinus massoniana plantations with various stand densities and rotation ages using a growth model method. The results revealed that with increasing stand age, low-density plantations at 2000 trees·ha -1 (358.80 m 3 ·ha -1 ), as well as high-density plantations at 4500 trees·ha -1 (359.10 m 3 ·ha -1 ), exhibited nearly identical standing volumes, which indicated that reduced inter-tree competition intensity favors the growth of larger trees during later stages of development. Furthermore, an increase in planting density led to a decrease in the average carbon sequestration rate, carbon sink, and number of trees during the rapid growth period, indicating that broader spacing between trees is favorable for biomass carbon accumulation. Further, extending the rotation period from 15 to 20 years or 25 years and reducing the optimal planting density from 3000 to 2000 trees·ha -1 increased the overall benefits of combined timber and carbon sink income by 2.14 and 3.13 times, respectively. The results highlighted that optimizing the planting density positively impacts the timber productivity and carbon sink storage of Pinus massoniana plantations and boosts the expected profits of forest managers. Thus, future afforestation initiatives must consider stand age and planting density management to shift from a scale-speed pattern to a quality-benefit design., (© 2024. The Author(s).)

Herbivorous insects alter biogeochemical cycling within forests, but the magnitude of these impacts, their global variation, and drivers of this variation remain poorly understood. To address this knowledge gap and help improve biogeochemical models, we established a global network of 74 plots within 40 mature, undisturbed broadleaved forests. We analyzed freshly senesced and green leaves for carbon, nitrogen, phosphorus and silica concentrations, foliar production and herbivory, and stand-level nutrient fluxes. We show more nutrient release by insect herbivores at non-outbreak levels in tropical forests than temperate and boreal forests, that these fluxes increase strongly with mean annual temperature, and that they exceed atmospheric deposition inputs in some localities. Thus, background levels of insect herbivory are sufficiently large to both alter ecosystem element cycling and influence terrestrial carbon cycling. Further, climate can affect interactions between natural populations of plants and herbivores with important consequences for global biogeochemical cycles across broadleaved forests., (© 2024. The Author(s).)

Understanding the mechanisms controlling forest carbon accumulation is crucial for predicting and mitigating future climate change. Yet, it remains unclear whether the dominance of ectomycorrhizal (EcM) trees influences the carbon accumulation of entire forests. In this study, we analyzed forest inventory data from over 4000 forest plots across Northeast China. We find that EcM tree dominance consistently exerts a positive effect on tree, soil, and forest carbon stocks. Moreover, we observe that these positive effects are more pronounced during unfavorable climate conditions, at lower tree species richness, and during early successional stages. This underscores the potential of increasing the dominance of native EcM tree species not only to enhance carbon stocks but also to bolster resilience against climate change in high-latitude forests. Here we show that forest managers can make informed decisions to optimize carbon accumulation by considering various factors such as mycorrhizal types, climate, successional stages, and species richness., (© 2024. The Author(s).)

Hypoallometric (slope<1) scaling between metabolic rate and body mass is often regarded as near-universal across organisms. However, there are compelling reasons to question hypoallometric scaling in woody plants, where metabolic rate is directly proportional to leaf area. This leaf area must provide carbon to the volume of the metabolically active sapwood (VMASW). Within populations of a species, variants in which VMASW increases per unit leaf area with height growth (e.g. ⅔ or ¾ scaling) would have proportionally less carbon for growth and reproduction as they grow taller. Therefore, selection should favor individuals in which, as they grow taller, leaf area scales isometrically with shoot VMASW (slope=1). Using tetrazolium staining, we measured total VMASW and total leaf area (LAtot) across 22 individuals of Ricinus communis and confirmed that leaf area scales isometrically with VMASW, and that VMASW is much smaller than total sapwood volume. With the potential of the LAtot-VMASW relationship to shape factors as diverse as the crown area-stem diameter relationship, conduit diameter scaling, reproductive output, and drought-induced mortality, our work indicates that the notion that sapwood increases per unit leaf area with height growth requires revision., (© The Author(s) 2024. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For commercial re-use, please contact reprints@oup.com for reprints and translation rights for reprints. All other permissions can be obtained through our RightsLink service via the Permissions link on the article page on our site—for further information please contact journals.permissions@oup.com.)

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

Nonstructural carbohydrates (NSC) are essential for tree growth and adaptation, yet our understanding of the seasonal storage and mobilization dynamics of whole-tree NSC is still limited, especially when tree functional types are involved. Here, Quercus acutissima Carruth. and Pinus massoniana Lamb, with distinct life-history traits (i.e. a deciduous broadleaf species vs an evergreen coniferous species), were studied to assess the size and seasonal fluctuations of organ and whole-tree NSC pools with a focus on comparing differences in carbon resource mobilization patterns between the two species. We sampled the organs (leaf, branch, stem and root) of the target trees repeatedly over four seasons of the year. Then, NSC concentrations in each organ were paired with biomass estimates from the allometric model to generate whole-tree NSC pools. The seasonal dynamics of the whole-tree NSC of Q. acutissima and P. massoniana reached the peak in autumn and summer, respectively. The starch pools of the two species were supplemented in the growing season while the soluble sugar pools were the largest in the dormant season. Seasonal dynamics of organ-level NSC concentrations and pools were affected by organ type and tree species, with above-ground organs generally increasing during the growing season and P. massoniana roots decreasing during the growing season. In addition, the whole-tree NSC pools of P. massoniana were larger but Q. acutissima showed larger seasonal fluctuations, indicating that larger storage was not associated with more pronounced seasonal fluctuations. We also found that the branch and root were the most dynamic organs of Q. acutissima and P. massoniana, respectively, and were the major suppliers of NSC to support tree growth activities. These results provide fundamental insights into the dynamics and mobilization patterns of NSC at the whole-tree level, and have important implications for investigating environmental adaptions of different tree functional types., (© 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.)

Atmospheric nitrogen (N) deposition in forests can affect soil microbial growth and turnover directly through increasing N availability and indirectly through altering plant-derived carbon (C) availability for microbes. This impacts microbial residues (i.e., amino sugars), a major component of soil organic carbon (SOC). Previous studies in forests have so far focused on the impact of understory N addition on microbes and microbial residues, but the effect of N deposition through plant canopy, the major pathway of N deposition in nature, has not been explicitly explored. In this study, we investigated whether and how the quantities (25 and 50 kg N ha -1  year -1 ) and modes (canopy and understory) of N addition affect soil microbial residues in a temperate broadleaf forest under 10-year N additions. Our results showed that N addition enhanced the concentrations of soil amino sugars and microbial residual C (MRC) but not their relative contributions to SOC, and this effect on amino sugars and MRC was closely related to the quantities and modes of N addition. In the topsoil, high-N addition significantly increased the concentrations of amino sugars and MRC, regardless of the N addition mode. In the subsoil, only canopy N addition positively affected amino sugars and MRC, implying that the indirect pathway via plants plays a more important role. Neither canopy nor understory N addition significantly affected soil microbial biomass (as represented by phospholipid fatty acids), community composition and activity, suggesting that enhanced microbial residues under N deposition likely stem from increased microbial turnover. These findings indicate that understory N addition may underestimate the impact of N deposition on microbial residues and SOC, highlighting that the processes of canopy N uptake and plant-derived C availability to microbes should be taken into consideration when predicting the impact of N deposition on the C sequestration in temperate forests., (© 2024 John Wiley & Sons Ltd.)

Amazonia contains the most extensive tropical forests on Earth, but Amazon carbon sinks of atmospheric CO 2 are declining, as deforestation and climate-change-associated droughts 1-4 threaten to push these forests past a tipping point towards collapse 5-8 . Forests exhibit complex drought responses, indicating both resilience (photosynthetic greening) and vulnerability (browning and tree mortality), that are difficult to explain by climate variation alone 9-17 . Here we combine remotely sensed photosynthetic indices with ground-measured tree demography to identify mechanisms underlying drought resilience/vulnerability in different intact forest ecotopes 18,19 (defined by water-table depth, soil fertility and texture, and vegetation characteristics). In higher-fertility southern Amazonia, drought response was structured by water-table depth, with resilient greening in shallow-water-table forests (where greater water availability heightened response to excess sunlight), contrasting with vulnerability (browning and excess tree mortality) over deeper water tables. Notably, the resilience of shallow-water-table forest weakened as drought lengthened. By contrast, lower-fertility northern Amazonia, with slower-growing but hardier trees (or, alternatively, tall forests, with deep-rooted water access), supported more-drought-resilient forests independent of water-table depth. This functional biogeography of drought response provides a framework for conservation decisions and improved predictions of heterogeneous forest responses to future climate changes, warning that Amazonia's most productive forests are also at greatest risk, and that longer/more frequent droughts are undermining multiple ecohydrological strategies and capacities for Amazon forest resilience., (© 2024. The Author(s), under exclusive licence to Springer Nature Limited.)

Methane is an important greenhouse gas 1 , but the role of trees in the methane budget remains uncertain 2 . Although it has been shown that wetland and some upland trees can emit soil-derived methane at the stem base 3,4 , it has also been suggested that upland trees can serve as a net sink for atmospheric methane 5,6 . Here we examine in situ woody surface methane exchange of upland tropical, temperate and boreal forest trees. We find that methane uptake on woody surfaces, in particular at and above about 2 m above the forest floor, can dominate the net ecosystem contribution of trees, resulting in a net tree methane sink. Stable carbon isotope measurement of methane in woody surface chamber air and process-level investigations on extracted wood cores are consistent with methanotrophy, suggesting a microbially mediated drawdown of methane on and in tree woody surfaces and tissues. By applying terrestrial laser scanning-derived allometry to quantify global forest tree woody surface area, a preliminary first estimate suggests that trees may contribute 24.6-49.9 Tg of atmospheric methane uptake globally. Our findings indicate that the climate benefits of tropical and temperate forest protection and reforestation may be greater than previously assumed., (© 2024. The Author(s).)

The uptake of carbon dioxide (CO 2 ) by terrestrial ecosystems is critical for moderating climate change 1 . To provide a ground-based long-term assessment of the contribution of forests to terrestrial CO 2 uptake, we synthesized in situ forest data from boreal, temperate and tropical biomes spanning three decades. We found that the carbon sink in global forests was steady, at 3.6 ± 0.4 Pg C yr -1 in the 1990s and 2000s, and 3.5 ± 0.4 Pg C yr -1 in the 2010s. Despite this global stability, our analysis revealed some major biome-level changes. Carbon sinks have increased in temperate (+30 ± 5%) and tropical regrowth (+29 ± 8%) forests owing to increases in forest area, but they decreased in boreal (-36 ± 6%) and tropical intact (-31 ± 7%) forests, as a result of intensified disturbances and losses in intact forest area, respectively. Mass-balance studies indicate that the global land carbon sink has increased 2 , implying an increase in the non-forest-land carbon sink. The global forest sink is equivalent to almost half of fossil-fuel emissions (7.8 ± 0.4 Pg C yr -1 in 1990-2019). However, two-thirds of the benefit from the sink has been negated by tropical deforestation (2.2 ± 0.5 Pg C yr -1 in 1990-2019). Although the global forest sink has endured undiminished for three decades, despite regional variations, it could be weakened by ageing forests, continuing deforestation and further intensification of disturbance regimes 1 . To protect the carbon sink, land management policies are needed to limit deforestation, promote forest restoration and improve timber-harvesting practices 1,3 ., (© 2024. This is a U.S. Government work and not under copyright protection in the US; foreign copyright protection may apply.)

Ozone uptake through the stomata in tree leaves is an important process for improving air quality by urban trees. Stomatal conductance (g s ) is a key determinant of stomatal ozone uptake. The parameterization of g s models for estimating stomatal ozone uptake of trees has mainly been carried out using g s data measured in seedling leaves although the leaf traits may differ between mature trees and seedlings. In the present study, we compared stomatal ozone uptake estimated by g s models parameterised with data from mature trees and seedlings of Zelkova serrata. We measured g s in leaves of mature trees and seedlings of Z. serrata using a leaf porometer for 3-4 growing seasons. The Jarvis-type g s model was parameterised with data from mature trees and seedlings, separately. The maximum g s , and the functions of the seedling g s estimation model regarding the response to air temperature, vapour pressure deficit and atmospheric ozone concentration were the factors inducing lower stomatal ozone uptake. In contrast, the function of the seedling g s estimation model regarding the response to irradiance resulted in a higher estimated stomatal ozone uptake. The estimated stomatal ozone uptake for one growing season (April-September) by the seedling g s estimation model was 27% lower than that by the mature tree g s estimation model. These results indicate that leaf gas exchange traits of Z. serrata were different between mature trees and seedlings, and that estimating ozone uptake in mature tree leaves using a model based on seedling g s measurements results in an underestimation., 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 Inc. All rights reserved.)

Investigating natural processes in arsenic (As) polluted areas and plants that have naturally chosen to grow there pose practical restoration recommendations. This study aimed to assess long-term changes in natural As dendroremediation dynamics for Quercus robur L. and Tilia cordata Mill., tree species capable of growing in areas polluted by mining activities. We examined total As and its forms, as well as B, Ca, K, Mg, Na and P, in soil and trees over 5 years. We also characterized pH and EC of soil, examined proline content in tree organs, and calculated Bioconcentration Factor (BCF) and Translocation Factor (TF) for As. Initial As concentrations in soil were 37.0 mg kg⁻ 1 under Q. robur and 34.7 mg kg⁻ 1 under T. cordata, significantly decreasing after 5 years to 10.5 mg kg⁻ 1 and 9.51 mg kg⁻ 1 , respectively. This corresponds to pollution reduction of up to 71.8%. A notable decrease in As(III) and dimethylarsinic acid, along with increase in other organic As forms in soil, was observed. Additionally, concentrations of essential elements in soil, as well as its pH and EC, decreased over time. Both tree species accumulated substantial amounts of As in their organs, but the dynamics of this process were species-specific. During first 4 years, T. cordata accumulated more As and exhibited higher BCF, but in the 5th year, it was clearly surpassed by Q. robur. The highest TF was calculated for Q. robur in year 3, and for T. cordata in years 2 and 3. Generally, limited aboveground movement of As was indicated: BCF >1 were calculated for years 2 and 3, while TF were consistently <1. Proline content increased significantly in all organs, correlating with As, especially in Q. robur. In contrast, Q. robur leaves mapping revealed stable macroelement distributions, but clear variations were observed for T. cordata., which may suggest specific reaction to stress. These findings suggest that both species can effectively restore As-polluted areas, though with different dynamics. The selection of species for dendrorestoration should be based on whether the goal is faster remediation with lesser overall reduction (e.g. T. cordata) or slower remediation with ultimately greater pollution reduction (e.g. Q. robur)., 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 Ltd. All rights reserved.)

Stem respiration constitutes a substantial proportion of autotrophic respiration in forested ecosystems, but its drivers across different spatial scales and land-use gradients remain poorly understood. This study quantifies and examines the impact of logging disturbance on stem CO 2 efflux (EA) in Malaysian Borneo. EA was quantified at tree- and stand-level in nine 1-ha plots over a logging gradient from heavily logged to old-growth using the static chamber method. Tree-level results showed higher EA per unit stem area in logged vs old-growth plots (37.0 ± 1.1 vs 26.92 ± 1.14 g C m -2  month -1 ). However, at stand-level, there was no difference in EA between logged and old-growth plots (6.7 ± 1.1 vs 6.0 ± 0.7 Mg C ha -1  yr -1 ) due to greater stem surface area in old-growth plots. Allocation to growth respiration and carbon use efficiency was significantly higher in logged plots. Variation in EA at both tree- and stand-level was driven by tree size, growth and differences in investment strategies between the forest types. These results reflect different resource allocation strategies and priorities, with a priority for growth in response to increased light availability in logged plots, while old-growth plots prioritise maintenance and cell structure., (© 2024 The Author(s). New Phytologist © 2024 New Phytologist Foundation.)

In this study, we assessed the impact of varied water deficit irrigation frequencies (T1: 2.5 L/4 days; T2: 5 L/8 days; CK: 5 L/4 days) on Zitian Seedless grapes from veraison to post-ripening. Notably, total soluble solids increased during on-tree storage compared to at maturity, while total anthocyanin content decreased, particularly in CK (60.16%), T1 (62.35%), and less in T2 (50.54%). Glucose and fructose levels increased significantly in T1 and T2, more so in T2, but slightly declined in CK. Tartaric acid content increased by 41.42% in T2. Moreover, compared to regular irrigation, water deficit treatments enhanced phenolic metabolites and volatile compounds, including chlorogenic acid, various flavonoids, viniferin, hexanal, 2-nonenal, 2-hexen-1-ol, (E)-, 3-hydroxy-dodecanoic acid, and 1-hexanol, etc. Overall, the T2 treatment outperformed T1 and CK in maintaining grape quality. This study reveals that combining on-tree storage with water deficit irrigation not only improves grape quality but also water efficiency., 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 Ltd. All rights reserved.)

To adapt to the complex belowground environment, plants make trade-offs between root resource acquisition and defence ability. This includes forming partnerships with different types of root associating microorganisms, such as arbuscular mycorrhizal and ectomycorrhizal fungi. These trade-offs, by mediating root chemistry, exert legacy effects on nutrient release during decomposition, which may, in turn, affect the ability of new roots to re-acquire resources, thereby generating a feedback loop. However, the linkages at the basis of this potential feedback loop remain largely unquantified. Here, we propose a trait-based root 'acquisition-defence-decomposition' conceptual framework and test the strength of relevant linkages across 90 angiosperm tree species. We show that, at the plant species level, the root-fungal symbiosis gradient within the root economics space, root chemical defence (condensed tannins), and root decomposition rate are closely linked, providing support to this framework. Beyond the dichotomy between arbuscular mycorrhizal-dominated versus ectomycorrhizal-dominated systems, we suggest a continuous shift in feedback loops, from 'high arbuscular mycorrhizal symbiosis-low defence-fast decomposition-inorganic nutrition' by evolutionarily ancient taxa to 'high ectomycorrhizal symbiosis-high defence-slow decomposition-organic nutrition' by more modern taxa. This 'acquisition-defence-decomposition' framework provides a foundation for testable hypotheses on multidimensional linkages between species' belowground strategies and ecosystem nutrient cycling in an evolutionary context., (© 2024. The Author(s).)

Deforestation of Atlantic Forest has caused prolonged drought events in the last decades. The need for reforestation is growing, and the development of native seedlings that are more tolerant to drought stress is necessary. A biotechnological tool that improves plant tolerance is the use of plant growth-promoting bacteria (PGPB) as inoculants. Two species of PGPB were inoculated in drought-stressed seedlings of two neotropical tree species that have been used in environmental restoration programs: Cecropia pachystachya and Cariniana estrellensis. Biometrical, physiological, and metabolomic parameters from carbon and nitrogen pathways were evaluated. We found that the PGPB positively influenced photosynthesis and growth parameters in both trees under drought. The enzymes activities, the tricarboxylic acid cycle intermediates, the amino acids, and protein contents were also influenced by the PGPB treatments. The results allowed us to find the specific composition of secondary metabolites of each plant species. This study provides evidence that there is not a single mechanism involved in drought tolerance and that the inoculation with PGPB promotes a broad-spectrum tolerance response in Neotropical trees. The inoculation with PGPB appears as an important strategy to improve drought tolerance in Atlantic Forest native trees and enhance environmental restoration programs' success. MAIN CONCLUSION: The association with plant growth-promoting bacteria improved the tolerance to drought in Neotropical trees through biochemical, physiological, and biometrical parameters. This can enhance the success of forest restoration programs., (© 2024. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Forest trees adopt effective strategies to optimize nitrogen (N) use through internal N recycling. In the context of more recurrent environmental stresses due to climate change, the question remains of whether increased frequency of drought or defoliation threatens this internal N recycling strategy. We submitted 8-year-old beech trees to 2 years of either severe drought (Dro) or manual defoliation (Def) to create a state of N starvation. At the end of the second year before leaf senescence, we labeled the foliage of the Dro and Def trees, as well as that of control (Co) trees, with 15N-urea. Leaf N resorption, winter tree N storage (total N, 15N, amino acids, soluble proteins) and N remobilization in spring were evaluated for the three treatments. Defoliation and drought did not significantly impact foliar N resorption or N concentrations in organs in winter. Total N amounts in Def tree remained close to those in Co tree, but winter N was stored more in the branches than in the trunk and roots. Total N amount in Dro trees was drastically reduced (-55%), especially at the trunk level, but soluble protein concentrations increased in the trunk and fine roots compared with Co trees. During spring, 15N was mobilized from the trunk, branches and twigs of both Co and Def trees to support leaf growth. It was only provided through twig 15N remobilization in the Dro trees, thus resulting in extremely reduced Dro leaf N amounts. Our results suggest that stress-induced changes occur in N metabolism but with varying severity depending on the constraints: within-tree 15N transport and storage strategy changed in response to defoliation, whereas a soil water deficit induced a drastic reduction of the N amounts in all the tree organs. Consequently, N dysfunction could be involved in drought-induced beech tree mortality under the future climate., (© The Author(s) 2024. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

Improving the availability of soil phosphorus (P) and promoting tree growth through tree species selection and assembly are the critical issue. We conducted an afforestation experiment following randomized block experimental design with 1, 2, 4, and 6 tree species richness in south subtropics, including Pinus massoniana , Mytilaria laosensis , Erythrophleum fordii , Castanopsis hystrix , Michelia macclurei , Manglietia glauca , Aquilaria sinensis , and Dalbergia odorifera . We measured the bioavailable P components (CaCl 2 -P, citrate-P, enzyme-P and HCl-P) and examined the effects of different tree species assembly on bioavailable P components and tree growth. The results showed that, compared with non-nitrogen-fixing tree species, the mixing of nitrogen-fixing tree species ( E. fordii and D. odorifera ) effectively increased the contents of soil water, total nitrogen, total phosphorus, and microbial biomass P (MBP). The assembly of specific tree species improved the accumulation of bioavailable P. Mixing of nitrogen-fixing tree species significantly increased CaCl 2 -P content by 46.2% to 160.3%, the enzyme-P content produced by microbial mineralization by 69.3% to 688.2%, and HCl-P by 31.5% to 81.3%, increased MBP by 81.8% to 149.4%, and microbial biomass N (MBN) by 88.1% to 160.6%, respectively. Redundancy and correlation analysis results showed that MBP, available P, total phosphorus, L-leucine aminopeptidase, cellobiose, acid phosphatase, MBN and soil organic carbon were key factors driving the variation of rhizosphere soil bioavailable P. Mixing of nitrogen-fixing tree species increased enzyme-P and citrate-P, and the availability of which were positively correlated to tree basal area. In this study, mixing of nitrogen-fixing tree species increased the rhizosphere soil bioavailable P content, which facilitates tree growth.

The contribution of litterfall nutrient return to the maintenance of soil carbon pool and nutrient cycling is a crucial aspect of forest ecosystem functioning. Taking 21 tree species in subtropical young plantations as subjects, we investigated the correlation between litterfall nutrient return characteristics and functional traits of leaf and root and. The results showed notable variations in litterfall production, standing crop, and nutrient return across all the examined tree species. Mytilaria laosensis exhibited the highest litterfall production (689.2 g·m -2 ·a -1 ) and standing crop (605.1 g·m -2 ), while Cryptomeria fortunei demonstrated the lowest litterfall production (36.0 g·m -2 ·a -1 ) and standing crop (10.0 g·m -2 ). The nitrogen and phosphorus return amounts of 21 species ranged from 3.0 to 48.3 kg·hm -2 and from 0.1 to 2.0 kg·hm -2 , respectively. Castanopsis fissa demonstrated the highest nitrogen return, while Liquidambar formosana exhibited the highest phosphorus return. C. fortunei had the lowest nitrogen and phosphorus return. Results of the stepwise regression analysis indicated that litterfall production exhibited a significant negative correlation with leaf nitrogen content and leaf dry matter content, and a significant positive correlation with fine root tissue density. Additionally, leaf nitrogen content, leaf dry matter content, and specific root length had a significant negative impact on standing crop. The structural equation modelling results indicated that leaf dry matter content had a direct or indirect negative effect on nitrogen return amount through the reduction of litterfall production. Conversely, fine root tissue density had a significant positive impact on nitrogen return amount by increasing litter leaf nitrogen content. Both leaf nitrogen content and leaf dry matter content had direct or indirect negative effects on phosphorus return amount through the reduction of litterfall production. In conclusion, the tree species with low leaf nitrogen content and dry matter content, as well as high fine root tissue density, was recommended for the establishment of plantations in the subtropical zone in order to enhance nutrient cycling through litter decomposition and improve soil fertility and forest productivity.

Plants adjust their growth and development in response to changing light caused by canopy shade. The molecular mechanisms underlying shade avoidance responses have been widely studied in Arabidopsis and annual crop species, yet the shade avoidance signalling in woody perennial trees remains poorly understood. Here, we first showed that PtophyB1/2 photoreceptors serve conserved roles in attenuating the shade avoidance syndrome (SAS) in poplars. Next, we conducted a systematic identification and characterization of eight PtoPIF genes in Populus tomentosa. Knocking out different PtoPIFs led to attenuated shade responses to varying extents, whereas overexpression of PtoPIFs, particularly PtoPIF3.1 and PtoPIF3.2, led to constitutive SAS phenotypes under normal light and enhanced SAS responses under simulated shade. Notably, our results revealed that distinct from Arabidopsis PIF4 and PIF5, which are major regulators of SAS, the Populus homologues PtoPIF4.1 and PtoPIF4.2 seem to play a minor role in controlling shade responses. Moreover, we showed that PtoPIF3.1/3.2 could directly activate the expression of the auxin biosynthetic gene PtoYUC8 in response to shade, suggesting a conserved PIF-YUC-auxin pathway in modulating SAS in tree. Overall, our study provides insights into shared and divergent functions of PtoPIF members in regulating various aspects of the SAS in Populus., (© 2024 John Wiley & Sons Ltd.)

Carbon influences the evolution and functioning of plants and their roots. Previous work examining a small number of commonly measured root traits has revealed a global multidimensionality of the resource economics traits in fine roots considering carbon as primary currency but without considering the diversity of carbon-related traits. To address this knowledge gap, we use data from 66 tree species from a tropical forest to illustrate that root economics space co-varies with a novel molecular-level traits space based on nuclear magnetic resonance. Thinner fine roots exhibit higher proportions of carbohydrates and lower diversity of molecular carbon than thicker roots. Mass-denser fine roots have more lignin and aromatic carbon compounds but less bioactive carbon compounds than lighter roots. Thus, the transition from thin to thick fine roots implies a shift in the root carbon economy from 'do-it-yourself' soil exploration to collaboration with mycorrhizal fungi, while the shift from light to dense fine roots emphasizes a shift from acquisitive to conservative root strategy. We reveal a previously undocumented role of molecular-level carbon traits that potentially undergird the multidimensional root economics space. This finding offers new molecular insight into the diversity of root form and function, which is fundamental to our understanding of plant evolution, species coexistence and adaptations to heterogeneous environments., (© 2024. The Author(s), under exclusive licence to Springer Nature Limited.)

Large trees in plantations generally produce more wood per unit of resource use than small trees. Two processes may account for this pattern: greater photosynthetic resource use efficiency or greater partitioning of carbon to wood production. We estimated gross primary production (GPP) at the individual scale by combining transpiration with photosynthetic water-use efficiency of Eucalyptus trees. Aboveground production fluxes were estimated using allometric equations and modeled respiration; total belowground carbon fluxes (TBCF) were estimated by subtracting aboveground fluxes from GPP. Partitioning was estimated by dividing component fluxes by GPP. Dominant trees produced almost three times as much wood as suppressed trees. They used 25 ± 10% (mean ± SD) of their photosynthates for wood production, whereas suppressed trees only used 12 ± 2%. By contrast, dominant trees used 27 ± 19% of their photosynthate belowground, whereas suppressed trees used 58 ± 5%. Intermediate trees lay between these extremes. Photosynthetic water-use efficiency of dominant trees was c. 13% greater than the efficiency of suppressed trees. Suppressed trees used more than twice as much of their photosynthate belowground and less than half as much aboveground compared with dominant trees. Differences in carbon partitioning were much greater than differences in GPP or photosynthetic water-use efficiency., (© 2024 The Authors. New Phytologist © 2024 New Phytologist Foundation.)

The capacity for terrestrial ecosystems to sequester additional carbon (C) with rising CO 2 concentrations depends on soil nutrient availability 1,2 . Previous evidence suggested that mature forests growing on phosphorus (P)-deprived soils had limited capacity to sequester extra biomass under elevated CO 2 (refs. 3-6 ), but uncertainty about ecosystem P cycling and its CO 2 response represents a crucial bottleneck for mechanistic prediction of the land C sink under climate change 7 . Here, by compiling the first comprehensive P budget for a P-limited mature forest exposed to elevated CO 2 , we show a high likelihood that P captured by soil microorganisms constrains ecosystem P recycling and availability for plant uptake. Trees used P efficiently, but microbial pre-emption of mineralized soil P seemed to limit the capacity of trees for increased P uptake and assimilation under elevated CO 2 and, therefore, their capacity to sequester extra C. Plant strategies to stimulate microbial P cycling and plant P uptake, such as increasing rhizosphere C release to soil, will probably be necessary for P-limited forests to increase C capture into new biomass. Our results identify the key mechanisms by which P availability limits CO 2 fertilization of tree growth and will guide the development of Earth system models to predict future long-term C storage., (© 2024. Crown.)

Physiological water stress induced by low root temperatures might contribute to species-specific climatic limits of tree distribution. We investigated the low temperature sensitivity of root water uptake and transport in seedlings of 16 European tree species which reach their natural upper elevation distribution limits at different distances to the alpine treeline. We used 2 H-H 2 O pulse-labelling to quantify the water uptake and transport velocity from roots to leaves in seedlings exposed to constant 15°C, 7°C or 2°C root temperature, but identical aboveground temperatures between 20°C and 25°C. In all species, low root temperatures reduced the water transport rate, accompanied by reduced stem water potentials and stomatal conductance. At 7°C root temperature, the relative water uptake rates among species correlated positively with the species-specific upper elevation limits, indicating an increasingly higher sensitivity to lower root zone temperatures, the lower a species' natural elevational distribution limit. Conversely, 2°C root temperature severely inhibited water uptake in all species, irrespective of the species' thermal elevational limits. We conclude that low temperature-induced hydraulic constraints contribute to the cold distribution limits of temperate tree species and are a potential physiological cause behind the low temperature limits of plant growth in general., (© 2024 The Authors. Plant, Cell & Environment published by John Wiley & Sons Ltd.)

Forest ecosystems store large amounts of carbon and can be important sources, or sinks, of the atmospheric carbon dioxide that is contributing to global warming. Understanding the carbon storage potential of different forests and their response to management and disturbance events are fundamental to developing policies and scenarios to partially offset greenhouse gas emissions. Projections of live tree carbon accumulation are handled differently in different models, with inconsistent results. We developed growth-and-yield style models to predict stand-level live tree carbon density as a function of stand age in all vegetation types of the coastal Pacific region, US (California, Oregon, and Washington), from 7,523 national forest inventory plots. We incorporated site productivity and stockability within the Chapman-Richards equation and tested whether intensively managed private forests behaved differently from less managed public forests. We found that the best models incorporated stockability in the equation term controlling stand carrying capacity, and site productivity in the equation terms controlling the growth rate and shape of the curve. RMSEs ranged from 10 to 137 Mg C/ha for different vegetation types. There was not a significant effect of ownership over the standard industrial rotation length (~50 yrs) for the productive Douglas-fir/western hemlock zone, indicating that differences in stockability and productivity captured much of the variation attributed to management intensity. Our models suggest that doubling the rotation length on these intensively managed lands from 35 to 70 years would result in 2.35 times more live tree carbon stored on the landscape. These findings are at odds with some studies that have projected higher carbon densities with stand age for the same vegetation types, and have not found an increase in yields (on an annual basis) with longer rotations. We suspect that differences are primarily due to the application of yield curves developed from fully-stocked, undisturbed, single-species, "normal" stands without accounting for the substantial proportion of forests that don't meet those assumptions. The carbon accumulation curves developed here can be applied directly in growth-and-yield style projection models, and used to validate the predictions of ecophysiological, cohort, or single-tree style models being used to project carbon futures for forests in the region. Our approach may prove useful for developing robust models in other forest types., Competing Interests: The authors have declared that no competing interests exist., (Copyright: This is an open access article, free of all copyright, and may be freely reproduced, distributed, transmitted, modified, built upon, or otherwise used by anyone for any lawful purpose. The work is made available under the Creative Commons CC0 public domain dedication.)

The leaves of many trees emit volatile organic compounds (abbreviated as BVOCs), which protect them from various damages, such as herbivory, pathogens, and heat stress. For example, isoprene is highly volatile and is known to enhance the resistance to heat stress. In this study, we analyze the optimal seasonal schedule for producing isoprene in leaves to mitigate damage. We assume that photosynthetic rate, heat stress, and the stress-suppressing effect of isoprene may vary throughout the season. We seek the seasonal schedule of isoprene production that maximizes the total net photosynthesis using Pontryagin's maximum principle. The isoprene production rate is determined by the changing balance between the cost and benefit of enhanced leaf protection over time. If heat stress peaks in midsummer, isoprene production can reach its highest levels during the summer. However, if a large portion of leaves is lost due to heat stress in a short period, the optimal schedule involves peaking isoprene production after the peak of heat stress. Both high photosynthetic rate and high isoprene volatility in midsummer make the peak of isoprene production in spring. These results can be clearly understood by distinguishing immediate impacts and the impacts of future expectations., (© 2024. The Author(s).)

The air pollution remediation is naturally carried out by plants. Their overground parts called phyllosphere are a type of a natural filter on which pollutants can be adsorb. Moreover, microbial communities living in phyllosphere perform a variety of biochemical processes removing also chemical pollutants. As their pollution is nowadays a burning issue especially for highly developed countries, the development of effective and ecological technologies for air treatment are of the utmost importance. The use of phyllosphere bacteria in the process of air bioremediation is a promising technology. This article reviews the role of phyllospheric bacteria in air bioremediation processes especially linked with the moderate climate plants. Research results published so far indicate that phyllosphere bacteria are able to metabolize the air pollutants but their potential is strictly determined by plant-phyllospheric bacteria interaction. The European tree species most commonly used for this purpose are also presented. The collected information filled the gap in the practical use of tree species in air bioremediation in the moderate climate zone., 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 Ltd. All rights reserved.)

Decomposition of dead organic matter is fundamental to carbon (C) and nutrient cycling in terrestrial ecosystems, influencing C fluxes from the biosphere to the atmosphere. Theory predicts and evidence strongly supports that the availability of nitrogen (N) limits litter decomposition. Positive relationships between substrate N concentrations and decomposition have been embedded into ecosystem models. This decomposition paradigm, however, relies on data mostly from short-term studies analyzing controls on early-stage decomposition. We present evidence from three independent long-term decomposition investigations demonstrating that the positive N-decomposition relationship is reversed and becomes negative during later stages of decomposition. First, in a 10-y decomposition experiment across 62 woody species in a temperate forest, leaf litter with higher N concentrations exhibited faster initial decomposition rates but ended up a larger recalcitrant fraction decomposing at a near-zero rate. Second, in a 5-y N-enrichment experiment of two tree species, leaves with experimentally enriched N concentrations had faster decomposition initial rates but ultimately accumulated large slowly decomposing fractions. Measures of amino sugars on harvested litter in two experiments indicated that greater accumulation of microbial residues in N-rich substrates likely contributed to larger slowly decomposing fractions. Finally, a database of 437 measurements from 120 species in 45 boreal and temperate forest sites confirmed that higher N concentrations were associated with a larger slowly decomposing fraction. These results challenge the current treatment of interactions between N and decomposition in many ecosystems and Earth system models and suggest that even the best-supported short-term controls of biogeochemical processes might not predict long-term controls., Competing Interests: Competing interests statement:The authors declare no competing interest.

The oxygen and hydrogen isotopic composition (δ18O, δ2H) of plant tissues are key tools for the reconstruction of hydrological and plant physiological processes and may therefore be used to disentangle the reasons for tree mortality. However, how both elements respond to soil drought conditions before death has rarely been investigated. To test this, we performed a greenhouse study and determined predisposing fertilization and lethal soil drought effects on δ18O and δ2H values of organic matter in leaves and tree rings of living and dead saplings of five European tree species. For mechanistic insights, we additionally measured isotopic (i.e. δ18O and δ2H values of leaf and twig water), physiological (i.e. leaf water potential and gas-exchange) and metabolic traits (i.e. leaf and stem non-structural carbohydrate concentration, carbon-to-nitrogen ratios). Across all species, lethal soil drought generally caused a homogenous 2H-enrichment in leaf and tree-ring organic matter, but a low and heterogenous δ18O response in the same tissues. Unlike δ18O values, δ2H values of tree-ring organic matter were correlated with those of leaf and twig water and with plant physiological traits across treatments and species. The 2H-enrichment in plant organic matter also went along with a decrease in stem starch concentrations under soil drought compared with well-watered conditions. In contrast, the predisposing fertilization had generally no significant effect on any tested isotopic, physiological and metabolic traits. We propose that the 2H-enrichment in the dead trees is related to (i) the plant water isotopic composition, (ii) metabolic processes shaping leaf non-structural carbohydrates, (iii) the use of carbon reserves for growth and (iv) species-specific physiological adjustments. The homogenous stress imprint on δ2H but not on δ18O suggests that the former could be used as a proxy to reconstruct soil droughts and underlying processes of tree mortality., (© The Author(s) 2024. Published by Oxford University Press.)

Source-sink balance in plants determines carbon distribution, and altering it can impact carbon fixation, transport, and allocation. We aimed to investigate the effect of altered source-sink ratios on carbon fixation, transport, and distribution in 'Valencia' sweet orange (Citrus x sinensis) by various defoliation treatments (0%, 33%, 66%, and 83% leaf removal). Gas exchange parameters were measured on 0 and 10 days after defoliation using A/C i response curves, and leaf export was measured two days after defoliation using radioisotope tracer techniques. Greater defoliation increased the maximum rate of carboxylation (V cmax ), electron transport rate (J 1200 ), and triose-phosphate utilization rate (TPU). Leaf export was unaffected by defoliation but increased in leaves closer to the shoot apex. Basipetal translocation velocity in the trunk remained unaltered, indicating that more photosynthates remained in the shoot rather than being transported directly to the root sink. Defoliated plants initiated more new flush shoots but accumulated less shoot biomass per plant after 8 weeks. Carbon allocation to fine roots was smaller in defoliated plants, suggesting defoliation led to retention of carbohydrates in aboveground organs such as the trunk and other shoots from previous growing cycles. In conclusion, the low source-sink ratio increased carbon fixation without impacting individual leaf export in citrus. The results suggest that intermediate sinks such as the aboveground perennial organs play a role in mediating the translocation velocity. Further research is necessary to better understand the dynamics of source-sink regulation in citrus trees., (© 2024 Scandinavian Plant Physiology Society.)

Tracer injection has long been recognized as a valuable tool for delineating tree hydraulics and assessing water transport pathways. Recently, isotope tracers have emerged as innovative instruments for investigating tree hydraulics, providing new insights into tree water dynamics. Nevertheless, there is a critical need for further research to comprehensively grasp water movement and distribution within trees. A previously introduced technique for analyzing the isotopic ratio of water in wet tissues, offering millimeter-scale resolution for visualizing tracer movement, faces challenges due to its underdeveloped sample preparation techniques. In this study, we introduced an H 2 18 O tracer into S. gracilistyla samples, exclusively comprising indeterminate roots, stems, and leaves, cultivated through hydroponics and grown within the current year. Our objective was to assess the axial distribution of the tracer in the xylem. Additionally, we devised a novel method for preparing frozen wet tissue samples, enhancing the repeatability and success rate of experiments. The results demonstrated that all frozen wet tissue samples exhibited an average water loss rate of less than 0.6%. Isotopic analysis of these samples unveiled a consistent decline in tracer concentration with increasing height in all Salix specimens, with three out of five samples revealing a significant isotope gradient. Our findings affirm the efficacy and practicality of combining isotopic labeling with freezing, stabilization, and preparation techniques. Looking ahead, our isotopic labeling and analysis methods are poised to transcend woody plants, finding extensive applications in plant physiology and ecohydrology., (© 2024 Scandinavian Plant Physiology Society.)

With over one-third of terrestrial net primary productivity transferring to the litter layer annually, the carbon release from litter serves as a crucial valve in atmospheric carbon dioxide concentrations. However, few quantitative global projections of litter carbon release rate in response to climate change exist. Here, we combined a global foliar litter carbon release dataset (8973 samples) to generate spatially explicitly estimates of the response of their residence time (τ) to climate change. Results show a global mean litter carbon release rate ( k $$ k $$ ) of 0.69 year -1 (ranging from 0.09-5.6 year -1 ). Under future climate scenarios, global mean τ is projected to decrease by a mean of 2.7% (SSP 1-2.6) and 5.9% (SSP 5-8.5) during 2071-2100 period. Locally, the alleviation of temperature and moisture restrictions corresponded to obvious decreases in τ in cold and arid regions, respectively. In contract, τ in tropical humid broadleaf forests increased by 4.6% under SSP 5-8.5. Our findings highlight the vegetation type as a powerful proxy for explaining global patterns in foliar litter carbon release rates and the role of climate conditions in predicting responses of carbon release to climate change. Our observation-based estimates could refine carbon cycle parameterization, improving projections of carbon cycle-climate feedbacks., (© 2024 John Wiley & Sons Ltd.)

Manganese (Mn) deficiency is a widespread occurrence across different landscapes, including agricultural systems and managed forests, and causes interruptions in the normal metabolic functioning of plants. The microelement is well-characterized for its role in the oxygen-evolving complex in photosystem II and maintenance of photosynthetic structures. Mn is also required for a variety of enzymatic reactions in secondary metabolism, which play a crucial role in defense strategies for trees. Despite the strong relationship between Mn availability and the biosynthesis of defense-related compounds, there are few studies addressing how Mn deficiency can impact tree defense mechanisms and the ensuing ecological patterns and processes. Understanding this relationship and highlighting the potentially deleterious effects of Mn deficiency in trees can also inform silvicultural and management decisions to build more robust forests. In this review, we address this relationship, focusing on forest trees. We describe Mn availability in forest soils, characterize the known impacts of Mn deficiency in plant susceptibility, and discuss the relationship between Mn and defense-related compounds by secondary metabolite class. In our review, we find several lines of evidence that low Mn availability is linked with lowered or altered secondary metabolite activity. Additionally, we compile documented instances where Mn limitation has altered the defense capabilities of the host plant and propose potential ecological repercussions when studies are not available. Ultimately, this review aims to highlight the importance of untangling the effects of Mn limitation on the ecophysiology of plants, with a focus on forest trees in both managed and natural stands., 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 Masson SAS. All rights reserved.)

Polycyclic aromatic hydrocarbon (PAH) concentrations in the leaves and 1-year-old branches of three common tree species growing in a middle-sized city located in a moderate climate zone were estimated. For this purpose, PAH phytoaccumulation in Platanus × acerifolia, Celtis australis, and Tilia grandifolia species from highly urbanized, traffic congested, and highly PAH-contaminated streets was compared with trees from non-contaminated parks in the same urban core. The gathered data was used to define 17 PAH profiles, identify the main PAH pollution emission sources, and determine the organ and species specificity of PAHs accumulation. Due to the direct absorption of polluted air via stomata, the leaves accumulated up to 30% more PAHs compared to the 1-year-old branches. As expected, PAH concentrations were much higher in street trees, while heavy weight PAHs (with five and six rings) were accumulated in the highest concentrations. The highest foliar Σ 17 PAH concentrations were detected in street-grown C. australis, followed by P. acerifolia and T. grandifolia (502.68, 488.45, and 339.47 ng g -1 dry weight (DW), respectively). The same pattern was noted for Σ 17 PAHs in branches (414.89, 327.58, and 342.99 ng g -1 DW, respectively). Thus, T. grandifolia emerged as the least effective PAH sink as it accumulated up to ~ 40% less PAHs than P. acerifolia and C. australis leaves/branches. Among the 17 tracked PAHs, benzo[a]anthracene, benzo[a]pyrene, dibenzo[a,h]anthracene, and pyrene were found to have accumulated in the highest concentrations in all analyzed species irrespective of the site, and accounted for more than 50% of the total detected PAHs. Finally, a "black box" about species and organ specificity, as well as specific drivers that limit PAHs uptake capacity by trees, was opened, while this work provides insights into further PAH phytoremediation strategies., (© 2024. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Replanting broadleaved trees in monoculture conifer plantations has been shown to improve the ecological environment. However, not much is known about the distribution properties of soil phosphate-mobilizing bacteria (PMB) under different mixed plantings or how PMB affects biometabolism-driven phosphorus (P) bioavailability. The phoD and pqqC genes serve as molecular markers of PMB because they regulate the mobilization of organic (Po) and inorganic (Pi) P. Differences in soil bioavailable P concentration, phoD- and pqqC-harboring PMB communities, and their main regulators were analyzed using biologically-based P (BBP) and high-throughput sequencing approaches after combining coniferous trees (Pinus massoniana) and five individual broadleaved trees (Bretschneidera sinensis, Michelia maudiae, Cercidiphyllum japonicum, Manglietia conifera, and Camellia oleifera). The findings revealed that the contents of litter P, soil organic carbon (SOC), available Pi (CaCl 2 -P), and labile Po (Enzyme-P) were significantly higher in conifer-broadleaf mixed plantations than those in the monospecific Pinus massoniana plantations (PM), especially in the mixed stands with the introduction of Cercidiphyllum japonicum, Michelia maudiae, and Camellia oleifera. Conifer-broadleaf mixing had little effect on the abundance of phoD and pqqC genes but significantly altered species composition within the communities. Conifer-broadleaf mixing improved soil microbial habitat mainly by increasing the pH, increasing carbon source availability and nutrient content, decreasing exchangeable Fe 3+ and Al 3+ content, and decreasing the activation degrees of Fe and Al oxides in acidic soils. A small group of taxa (phoD: Bradyrhizobium, Tardiphaga, Nitratireductor, Mesorhizobium, Herbaspirillum, and Ralstonia; pqqC: Burkholderia, Variovorax, Bradyrhizobium, and Leptothrix) played a key role in the synthesis of P-related enzymes (e.g., alkaline phosphomonoesterase, ALP) and in lowering the levels of mineral-occluded (HCl-P) and chelated (Citrate-P) Pi. Overall, our findings highlight that mixing conifers and broadleaves could change the PMB communities that produce ALP and dissolve Pi to make P more bioavailable., 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 Ltd. All rights reserved.)

Transpiration is a significant part of water cycle in forest ecosystems, influenced by meteorological factors and potentially constrained by soil moisture. We used Granier-type thermal dissipation probes to monitor xylem sap flow dynamics of three tree species ( Quercus liaotungensis , Platycladus orientalis , and Robinia pseudoacacia ) in a semi-arid loess hilly region, and to continuously monitor the key meteorological factors and soil water content (SWC). We established the SWC thresholds delineating soil moisture-limited and -unlimited sap flow responses to transpiration drivers. The results showed that mean sap flux density ( J s ) of Q. liaotungensis and R. pseudoacacia was significantly higher during period with higher soil moisture compared to lower soil moisture, while the difference in J s for P. orientalis between the two periods was not significant. We used an exponential saturation function to fit the relationship between the J s of each tree species and the integrated transpiration variable (VT) which reflected solar radiation and vapor pressure deficit. The difference in the fitting curve parameters indicated that there were distinct response patterns between J s and VT under different soil moisture conditions. There was a threshold in soil moisture limitation on sap flow for each species, which was identified as 0.129 m 3 ·m -3 for Q. liaotungensis , 0.116 m 3 ·m -3 for P. orientalis , and 0.108 m 3 ·m -3 for R. pseudoacacia . Below the thresholds, J s was limited by soil moisture. Above these points, the normalized sensitivity index (NSI) for Q. liaotungensis and P. orientalis reached saturation, while that of R. pseudoacacia did not reach saturation but exhibited a significant reduction in moisture limitation. Among the three species, P. orientalis was the most capable of overcoming soil moisture constraints.

The natural abundance of stable carbon and nitrogen isotopes (δ 13 C and δ 15 N) in leaves can provide comprehensive information on the physiological and ecological processes of plants and has been widely used in ecological research. However, recent studies on leaf δ 13 C and δ 15 N have focused mainly on woody species, few studies have been conducted on herbs in different vegetation types, and their differences and driving factors are still unclear. In this study, we focused on the herbs in subalpine coniferous forests, alpine shrublands, and alpine mea-dows on the eastern Qinghai-Tibet Plateau, and investigated the differences in leaf δ 13 C and δ 15 N of herbs and the driving factors. The results showed that there were significant differences in leaf δ 13 C and δ 15 N values of herbs among different vegetation types, with the highest δ 13 C and δ 15 N values in alpine meadows, followed by alpine shrublands, and the lowest in subalpine coniferous forests. Using variation partitioning analysis, we revealed that differences in leaf δ 13 C and δ 15 N of herbs among various vegetation types were driven by both leaf functional traits and climate factors, with the contribution of leaf functional traits being relatively higher than that of climate factors. Hierarchical partitioning results indicated that mean annual temperature (MAT), chlorophyll content index, leaf nitrogen content per unit area (N area ), and leaf mass per area were the main drivers of leaf δ 13 C variations of herbs across different vegetation types, while the relative importance of N area and MAT for variation in leaf δ 15 N of herbs was much higher than those other variables. There was a strong coupling relationship between leaf δ 13 C and δ 15 N as indicated by the result of the ordinary least squares regression. Our findings could provide new insights into understanding the key drivers of leaf δ 13 C and δ 15 N variations in herbs across different vegetation types.

Non-structural carbohydrates (NSCs) are building blocks for biomass and fuel metabolic processes. However, it remains unclear how tropical forests mobilize, export, and transport NSCs to cope with extreme droughts. We combined drought manipulation and ecosystem 13CO2 pulse-labeling in an enclosed rainforest at Biosphere 2, assessed changes in NSCs, and traced newly assimilated carbohydrates in plant species with diverse hydraulic traits and canopy positions. We show that drought caused a depletion of leaf starch reserves and slowed export and transport of newly assimilated carbohydrates below ground. Drought effects were more pronounced in conservative canopy trees with limited supply of new photosynthates and relatively constant water status than in those with continual photosynthetic supply and deteriorated water status. We provide experimental evidence that local utilization, export, and transport of newly assimilated carbon are closely coupled with plant water use in canopy trees. We highlight that these processes are critical for understanding and predicting tree resistance and ecosystem fluxes in tropical forest under drought., (© The Author(s) 2024. Published by Oxford University Press on behalf of the Society for Experimental Biology.)

The loss of leaves and needles in tree crowns and tree mortality are increasing worldwide, mostly as a result of more frequent and severe drought stress. Scots pine (Pinus sylvestris L.) is a tree species that is strongly affected by these developments in many regions of Europe and Asia. So far, changes in metabolic pathways and metabolite profiles in needles and roots on the trajectory toward mortality are unknown, although they could contribute to a better understanding of the mortality mechanisms. Therefore, we linked long-term observations of canopy defoliation and tree mortality with the characterization of the primary metabolite profile in needles and fine roots of Scots pines from a forest site in the Swiss Rhone valley. Our results show that Scots pines are able to maintain metabolic homeostasis in needles over a wide range of canopy defoliation levels. However, there is a metabolic tipping point at around 80-85% needle loss. Above this threshold, many stress-related metabolites (particularly osmoprotectants, defense compounds and antioxidants) increase in the needles, whereas they decrease in the fine roots. If this defoliation tipping point is exceeded, the trees are very likely to die within a few years. The different patterns between needles and roots indicate that mainly belowground carbon starvation impairs key functions for tree survival and suggest that this is an important factor explaining the increasing mortality of Scots pines., (© The Author(s) 2024. Published by Oxford University Press.)

Many plants, especially trees, emit isoprene in a highly light- and temperature-dependent manner. The advantages for plants that emit, if any, have been difficult to determine. Direct effects on membranes have been disproven. New insights have been obtained by RNA sequencing, proteomic and metabolomic studies. We determined the responses of the phosphoproteome to exposure of Arabidopsis leaves to isoprene in the gas phase for either 1 or 5 h. Isoprene effects that were not apparent from RNA sequencing and other methods but were apparent in the phosphoproteome include effects on chloroplast movement proteins and membrane remodelling proteins. Several receptor kinases were found to have altered phosphorylation levels. To test whether potential isoprene receptors could be identified, we used molecular dynamics simulations to test for proteins that might have strong binding to isoprene and, therefore might act as receptors. Although many Arabidopsis proteins were found to have slightly higher binding affinities than a reference set of Homo sapiens proteins, no specific receptor kinase was found to have a very high binding affinity. The changes in chloroplast movement, photosynthesis capacity and so forth, found in this work, are consistent with isoprene responses being especially useful in the upper canopy of trees., (© 2023 The Authors. Plant, Cell & Environment published by John Wiley & Sons Ltd.)

Trees can effectively capture airborne particles and improve air quality. However, the specific response of phyllosphere microbiome (PMo) in different plant species to particulate matter (PM) and the heavy metals it contains are not yet fully understood. In this study, we investigated the impact of PM on the diversity and function of PMo in Loropetalum chinense and Osmanthus fragrans trees grown in industrial and clean zones with varying levels of PM pollution. Our findings revealed that leaf dust had a significant negative effect on microbial richness, with O. fragrans exhibiting higher microbial diversity than L. chinense. The dominant phylum of phyllosphere bacteria in all samples was Proteobacteria, and the dominant genera were Stenotrophomonas and Delftia. The relative abundance of these genera varied significantly among plant species and regions. Our results showed that PM had a significant impact on the community composition of PMo, with the presence of heavy metals exerting a greater effect than particle size. Moreover, the foliar microbial community of plants grown in industrial zones exhibited significantly higher metabolic functions related to stress resistance and disease resistance compared to plants in control zones. These findings highlight the structural and functional responses of PMo to PM and indicate their potential for enhancing plant adaptation to environmental stress., 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.)