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Roots are the interfaces between biochar particles and growing plants. Biochar application may alter root growth and traits and thereby affect plant performance. However, a comprehensive understanding of the effects of biochar on root traits is lacking. We conducted a meta‐analysis with 2108 paired observations from 136 articles to evaluate the responses of root traits associated with 13 variables under biochar application. Overall, biochar application increased root biomass (+32%), root volume (+29%) and surface area (39%). The biochar‐induced increases in root length (+52%) and number of root tips (+17%) were much larger than the increase in root diameter (+9.9%); this result suggests that biochar application benefits root morphological development to alleviate plant nutrient and water deficiency rather than to maximize biomass accumulation. Biochar application did not change root N concentration but significantly increased root P concentration (+22%), particularly when combined with N fertilization. Biochar application also affected root‐associated microbes and significantly increased the number of root nodules (+25%). The responses of root traits to biochar application were generally greater in annual plants than in perennial plants and were affected by soil texture and pH values. Moreover, it appears that biochar production process (pyrolysis temperature and time) plays a more important role in regulating root growth than does biochar source. Together, findings obtained from this meta‐analysis may have significant implications for the future sustainable development of biochar management to improve plant growth and functioning. To future sustainable biochar management, a comprehensive meta‐analysis with 2108 paired observations from 136 articles was conducted to evaluate responses of root traits associated with 13 variables under biochar application. Overall, biochar application significantly improved root growth and morphology, in particular, with overall stronger stimulation on root length than root diameter, which would benefit root morphological development to alleviate plant nutrient and water deficiency rather than to maximize biomass accumulation. Moreover, the responses of root traits to biochar application were affected by fertilizer input, plant species, soil pH and texture, and biochar production process.

Global climate change is expected to further increase the frequency and severity of extreme events, such as high temperature/heat waves as well as drought in the future. Thus, how plant responds to high temperature and drought has become a key research topic. In this study, we extracted data from Web of Science Core Collections database, and synthesized plant responses to high temperature and drought based on bibliometric methods using software of R and VOSviewer. The results showed that a stabilized increasing trend of the publications (1199 papers) was found during the period of 2008 to 2014, and then showed a rapid increase (2583 papers) from year 2015 to 2021. Secondly, the top five dominant research fields of plant responses to high temperature and drought were Plant Science, Agroforestry Science, Environmental Science, Biochemistry, and Molecular Biology, respectively. The largest amount of published article has been found in the Frontiers in Plant Science journal, which has the highest global total citations and H-index. We also found that the journal of Plant Physiology has the highest local citations. From the most cited papers and references, the most important research focus was the improvement of crop yield and vegetation stress resistance. Furthermore, “drought” has been the most prominent keyword over the last 14 years, and more attention has been paid to “climate change” over the last 5 years. Under future climate change, how to regulate growth and development of food crops subjected to high temperature and drought stress may become a hotspot, and increasing research is critical to provide more insights into plant responses to high temperature and drought by linking plant above-below ground components. To summarize, this research will contribute to a comprehensive understanding of the past, present, and future research on plant responses to high temperature and drought.

Widespread tree mortality associated with drought has been observed on all forested continents and global change is expected to exacerbate vegetation vulnerability. Forest mortality has implications for future biosphere–atmosphere interactions of carbon, water and energy balance, and is poorly represented in dynamic vegetation models. Reducing uncertainty requires improved mortality projections founded on robust physiological processes. However, the proposed mechanisms of drought-induced mortality, including hydraulic failure and carbon starvation, are unresolved. A growing number of empirical studies have investigated these mechanisms, but data have not been consistently analysed across species and biomes using a standardized physiological framework. Here, we show that xylem hydraulic failure was ubiquitous across multiple tree taxa at drought-induced mortality. All species assessed had 60% or higher loss of xylem hydraulic conductivity, consistent with proposed theoretical and modelled survival thresholds. We found diverse responses in non-structural carbohydrate reserves at mortality, indicating that evidence supporting carbon starvation was not universal. Reduced non-structural carbohydrates were more common for gymnosperms than angiosperms, associated with xylem hydraulic vulnerability, and may have a role in reducing hydraulic function. Our finding that hydraulic failure at drought-induced mortality was persistent across species indicates that substantial improvement in vegetation modelling can be achieved using thresholds in hydraulic function. The mechanisms underlying drought-induced tree mortality are not fully resolved. Here, the authors show that, across multiple tree species, loss of xylem conductivity above 60% is associated with mortality, while carbon starvation is not universal.

Understanding the physiological and biochemical responses of tree seedlings under extreme drought stress, along with recovery during rewatering, and potential intra-species differences, will allow us to more accurately predict forest responses under future climate change. Here, we selected seedlings from four provenances (AH (Anhui), JX (Jiangxi), HN (Hunan) and GX (Guangxi)) of Schima superba and carried out a simulated drought-rewatering experiment in a field-based rain-out shelter. Seedlings were progressively dried until they reached 50% and 88% loss of xylem hydraulic conductivity (PLC) (i.e. P 50 and P 88 ), respectively, before they were rehydrated and maintained at field capacity for 30 days. Leaf photosynthesis ( A sat ), water status, activity of superoxide dismutase (SOD), and proline (Pro) concentration were monitored and their associations were determined. Increasing drought significantly reduced A sat , relative water content (RWC) and SOD activity in all provenances, and Pro concentration was increased to improve water retention; all four provenances exhibited similar response patterns, associated with similar leaf ultrastructure at pre-drought. Upon rewatering, physiological and biochemical traits were restored to well-watered control values in P 50 -stressed seedlings. In P 88 -stressed seedlings, Pro was restored to control values, while SOD was not fully recovered. The recovery pattern differed partially among provenances. There was a progression of recovery following watering, with RWC firstly recovered, followed by SOD and Pro, and then A sat , but with significant associations among these traits. Collectively, the intra-specific differences of S. superba seedlings in recovery of physiology and biochemistry following rewatering highlight the need to consider variations within a given tree species coping with future more frequent drought stress.

Background Drought and nitrogen deposition are the major global change factors that alter forest dynamics by affecting tree growth and physiology. However, the impacts of increased nitrogen availability at pre-drought on trees remains poorly understood, and it remains unclear how these responses are coordinated. In this study, we conducted the fertilization-drought microcosm experiment using a widely distributed evergreen broadleaf tree species seedlings ( Schima superba ) in southern China. The experiment was conducted at 3 stages. First, four levels of N fertilization treatments (without N fertilization-NF, low N fertilization-LF, moderate N fertilization-MF, high N fertilization-HF) were applied for 60 days. Second, all seedlings were allowed to grow under four levels of N fertilization treatments for another 60 days to ensure that the N was absorbed by seedlings. Third, all seedling were subjected to three levels of sustained drought treatments for further 60 days. Traits related to growth and physiology were monitored. Results Our findings indicate that drought alone inhibited the growth and leaf photosynthetic rate of S. superba , while N fertilization alone stimulated the growth and leaf photosynthetic rate. Antecedent N fertilization alleviated the drought limitation on growth, due to the increased leaf photosynthetic rate ( A sat ) and instantaneous water use efficiency. Moderate N fertilization mitigated the negative effects of drought on A sat due to improved performance in stomatal conductance, leaf water potential and cell membrane permeability. Additionally, moderate N fertilization increased activities of antioxidant enzymes and osmoprotectants concentration under drought condition. Conclusions Overall, our findings suggest that increased N fertilization prior to drought can alleviate the negative effects of drought on growth and physiology, which is dependent on the magnitude of N fertilization and drought stress.

Eucalyptus species are widely planted for reforestation in subtropical China. However, the effects of Eucalyptus plantations on the regional water use remain poorly understood. In an age sequence of 2-, 4- and 6-year-old Eucalyptus plantations, the tree water use and soil evaporation were examined by linking model estimations and field observations. Results showed that annual evapotranspiration of each age sequence Eucalyptus plantations was 876.7, 944.1 and 1000.7 mm, respectively, accounting for 49.81%, 53.64% and 56.86% of the annual rainfall. In addition, annual soil evaporations of 2-, 4- and 6-year-old were 318.6, 336.1, and 248.7 mm of the respective Eucalyptus plantations. Our results demonstrated that Eucalyptus plantations would potentially reduce water availability due to high evapotranspiration in subtropical regions. Sustainable management strategies should be implemented to reduce water consumption in Eucalyptus plantations in the context of future climate change scenarios such as drought and warming.

Soybean adapts to phosphorus-deficient soils through three important phosphorus acquisition strategies, namely altered root conformation, exudation of carboxylic acids, and symbiosis with clumping mycorrhizal fungi. However, the trade-offs and regulatory mechanisms of these three phosphorus acquisition strategies in soybean have not been researched. In this study, we investigated the responses of ten different soybean varieties to low soil phosphorus availability by determining biomass, phosphorus accumulation, root morphology, exudation, and mycorrhizal colonization rate. Furthermore, the molecular regulatory mechanisms underlying root phosphorus acquisition strategies were examined among varieties with different low-phosphorus tolerance using transcriptome sequencing and weighted gene co-expression network analysis. The results showed that two types of phosphorus acquisition strategies-\"outsourcing\" and \"do-it-yourself\"-were employed by soybean varieties under low phosphorus availability. The \"do-it-yourself\" varieties, represented by QD11, Zh30, and Sd, obtained sufficient phosphorus by increasing their root surface area and secreting carboxylic acids. In contrast, the \"outsourcing\" varieties, represented by Zh301, Zh13, and Hc6, used increased symbiosis with mycorrhizae to obtain phosphorus owing to their large root diameters. Transcriptome analysis showed that the direction of acetyl-CoA metabolism could be the dividing line between the two strategies of soybean selection. ERF1 and WRKY1 may be involved in the regulation of phosphorus acquisition strategies for soybeans grown under low P environments. These findings will enhance our understanding of phosphorus acquisition strategies in soybeans. In addition, they will facilitate the development of breeding strategies that are more flexible to accommodate a variety of production scenarios in agriculture under low phosphorus environments.

BackgroundThe prevalence of understory removal and anthropogenic nitrogen (N) deposition has significantly altered the ecological processes of forest ecosystems at both regional and global scales. However, it remains a pressing challenge to understand how N deposition and understory removal affect leaf nutrient dynamics, nutrient resorption, litter decomposition, and their linkages for better managing forest ecosystems under nutrient imbalances induced by N enrichment. To address this research gap, a field manipulation experiment was carried out in a subtropical Cunninghamia lanceolata plantation with four treatments including: control (CK), canopy N addition (CN), understory removal (UR), and canopy N addition plus understory removal (CN × UR). Green and senesced leaf N and phosphorus (P) concentrations, N and P resorption efficiencies, litter decomposition, and their correlations were measured.ResultsThe results revealed that the average N concentrations of green early and late leaves in UR were increased by 6.61 and 18.89% compared to CK. UR had the highest whereas CN had the lowest P concentrations in green leaves across the two sampling seasons. Following this, UR, leaf type, season, and their interactions significantly affected leaf N, P, and N:P (P < 0.05). The highest leaf N resorption (32.68%) and P resorption efficiencies (63.96%) were recorded in UR. Litter decomposition was significantly retarded in UR (P < 0.01) relative to CN. The regression analysis demonstrated that leaf nutrient status was significantly interconnected with leaf nutrient resorption efficiencies. In addition, leaf nutrient dynamics were strongly correlated with litter nutrients, indicating that both were coupled.ConclusionThese findings can deepen our knowledge of biogeochemical cycling and reveal contrasting nutrient-acquisition strategies on N and P limitation in response to UR and CN. Considering the P limitation, it is important to note that P was resorbed more efficiently, illustrating a remarkable nutrient preservation approach for nutrient-limitations. Resorption may be a crucial mechanism for keeping nutrients in these forests, so better understory management practices are required to prevent reliance on external nutrient pools. Overall, this study sheds meaningful insights into the ability of forest adaptation in response to global climatic change.

Subtropical tree species may experience severe drought stress due to variable rainfall under future climates. However, the capacity to restore hydraulic function post-drought might differ among co-occurring species with contrasting leaf habits (e.g., evergreen and deciduous) and have implications for future forest composition. Moreover, the links between hydraulic recovery and physiological and morphological traits related to water-carbon availability are still not well understood. Here, potted seedlings of six tree species (four evergreen and two deciduous) were grown outdoors under a rainout shelter. They grew under favorable water conditions until they were experimentally subjected to a soil water deficit leading to losses of ca. 50% of hydraulic conductivity, and then soils were re-watered to field capacity. Traits related to carbon and water relations were measured. There were differences in drought responses and recovery between species, but not as a function of evergreen or deciduous groups. Sapindus mukorossi exhibited the most rapid drought response, which was associated with a suite of physiological and morphological traits (larger plant size, the lowest hydraulic capacitance (Cbranch), higher minimum conductance (gmin) and lower HV (Huber value)). Upon re-watering, xylem water potential exhibited fast recovery in 1–3 days among species, while photosynthesis at saturating light (Asat) and stomatal conductance (gs) recovery lagged behind water potential recovery depending on species, with gs recovery being more delayed than Asat in most species. Furthermore, none of the six species exhibited significant hydraulic recovery during the 7 days re-watering period, indicating that xylem refilling was apparently limited; in addition, NSC availability had a minimal role in facilitating hydraulic recovery during this short-term period. Collectively, if water supply is limited by insignificant hydraulic recovery post-drought, the observed carbon assimilation recovery of seedlings may not be sustained over the longer term, potentially altering seedling regeneration and shifting forest species composition in subtropical China under climate change.

The deposition of reactive nitrogen (N) has substantially increased in subtropical regions due to human activities. However, the effects of long‐term N addition on the water‐use efficiency of subtropical forests are poorly understood. Here, we conducted an 11‐year experiment in a subtropical Cunninghamia lanceolate plantation with four N‐addition levels: N0, N1, N2, and N3 (equivalent to 0, 6, 12, and 24 g N m−2 year−1, respectively). A thermal dissipation probe system was used to calculate sap flow, and plant biomass carbon was assessed by field investigation. The whole‐plant water use and water‐use efficiency were estimated. In addition, the δ13C of tree rings was used to indicate the plant intrinsic water‐use efficiency. The results showed that N3 treatment significantly increased the annual sap flow velocity, especially in summer and winter. Annual water use, plant growth, and water‐use efficiency did not significantly differ among the N treatments, but water use tended to be higher in N3 treatment than in N0 treatment. Furthermore, the significant reduction of δ13C in N3 treatment than in N0 treatment supported the inference that N addition could increase water use. We conclude that long‐term addition of high levels (but not of low levels) of N increased whole‐plant water use in C. lanceolate plantations. Our findings indicate that N deposition accompanied by high temperature and drought events may negatively affect water balance in subtropical forests.