Global Change in Agricultural Flash Drought over the 21st Century

Agricultural flash droughts are high-impact phenomena, characterized by rapid soil moisture dry down. The ensuing dry conditions can persist for weeks to months, with detrimental effects on natural ecosystems and crop cultivation. Increases in the frequency of these rare events in a future warmer climate would have significant societal impact. This study uses an ensemble of 10 Coupled Model Intercomparison Project(CMIP) models to investigate the projected change in agricultural flash drought during the 21st century. Comparison across geographical regions and climatic zones indicates that individual events are preceded by anomalously low relative humidity and precipitation, with long-term trends governed by changes in temperature, relative humidity, and soil moisture. As a result of these processes, the frequency of both upperlevel and root-zone flash drought is projected to more than double in the mid-and high latitudes over the 21st century, with hot spots developing in the temperate regions of Europe, and humid regions of South America, Europe, and southern Africa.

Characterization and Propagation of Historical and Projected Droughts in the Umatilla River Basin, Oregon, USA

Climate change is expected to have long-term impacts on drought and wildfire risks in Oregon as summers continue to become warmer and drier. This paper investigates the projected changes in drought characteristics and drought propagation in the Umatilla River Basin in northeastern Oregon for mid-century(2030–2059) and late-century(2070–2099) climate scenarios. Drought characteristics for projected climates were determined using downscaled CMIP5 climate datasets from ten climate models and Soil and Water Assessment Tool to simulate effects on hydrologic processes. Short-term(three months) drought characteristics(frequency, duration, and severity) were analyzed using four drought indices, including the Standardized Precipitation Index(SPI-3), Standardized Precipitation-Evapotranspiration Index(SPEI-3), Standardized Streamflow Index(SSI-3), and the Standardized Soil Moisture Index(SSMI-3). Results indicate that short-term meteorological droughts are projected to become more prevalent, with up to a 20% increase in the frequency of SPI-3drought events. Short-term hydrological droughts are projected to become more frequent(average increase of 11% in frequency of SSI-3 drought events), more severe, and longer in duration(average increase of 8% for short-term droughts).Similarly, short-term agricultural droughts are projected to become more frequent(average increase of 28% in frequency of SSMI-3 drought events) but slightly shorter in duration(average decrease of 4%) in the future. Historically, drought propagation time from meteorological to hydrological drought is shorter than from meteorological to agricultural drought in most sub-basins. For the projected climate scenarios, the decrease in drought propagation time will likely stress the timing and capacity of water supply in the basin for irrigation and other uses.

Assessing the Performance of CMIP6 Models in Simulating Droughts across Global Drylands

Both the attribution of historical change and future projections of droughts rely heavily on climate modeling. However,reasonable drought simulations have remained a challenge, and the related performances of the current state-of-the-art Coupled Model Intercomparison Project phase 6(CMIP6) models remain unknown. Here, both the strengths and weaknesses of CMIP6 models in simulating droughts and corresponding hydrothermal conditions in drylands are assessed.While the general patterns of simulated meteorological elements in drylands resemble the observations, the annual precipitation is overestimated by ~33%(with a model spread of 2.3%–77.2%), along with an underestimation of potential evapotranspiration(PET) by ~32%(17.5%–47.2%). The water deficit condition, measured by the difference between precipitation and PET, is 50%(29.1%–71.7%) weaker than observations. The CMIP6 models show weaknesses in capturing the climate mean drought characteristics in drylands, particularly with the occurrence and duration largely underestimated in the hyperarid Afro-Asian areas. Nonetheless, the drought-associated meteorological anomalies, including reduced precipitation, warmer temperatures, higher evaporative demand, and increased water deficit conditions, are reasonably reproduced. The simulated magnitude of precipitation(water deficit) associated with dryland droughts is overestimated by 28%(24%) compared to observations. The observed increasing trends in drought fractional area,occurrence, and corresponding meteorological anomalies during 1980–2014 are reasonably reproduced. Still, the increase in drought characteristics, associated precipitation and water deficit are obviously underestimated after the late 1990s,especially for mild and moderate droughts, indicative of a weaker response of dryland drought changes to global warming in CMIP6 models. Our results suggest that it is imperative to employ bias correction approaches in drought-related studies over drylands by using CMIP6 outputs.

Drought-Tolerant Rice at Molecular Breeding Eras:An Emerging Reality

Rice(Oryza sativa L.)stands as the most significantly influential food crop in the developing world,with its total production and yield stability affected by environmental stress.Drought stress impacts about 45%of the world’s rice area,affecting plants at molecular,biochemical,physiological,and phenotypic levels.The conventional breeding method,predominantly employing single pedigree selection,has been widely utilized in breeding numerous drought-tolerant rice varieties since the Green Revolution.With rapid progress in plant molecular biology,hundreds of drought-tolerant QTLs/genes have been identified and tested in rice crops under both indoor and field conditions.Several genes have been introgressed into elite germplasm to develop commercially accepted drought-tolerant varieties,resulting in the development of several drought-tolerant rice varieties through marker-assisted selection and genetically engineered approaches.This review provides up-to-date information on proof-of-concept genes and breeding methods in the molecular breeding era,offering guidance for rice breeders to develop drought-tolerant rice varieties.

Influence of varied drought types on soil conservation service within the framework of climate change:insights from the Jinghe River Basin,China

Severe soil erosion and drought are the two main factors affecting the ecological security of the Loess Plateau,China.Investigating the influence of drought on soil conservation service is of great importance to regional environmental protection and sustainable development.However,there is little research on the coupling relationship between them.In this study,focusing on the Jinghe River Basin,China as a case study,we conducted a quantitative evaluation on meteorological,hydrological,and agricultural droughts(represented by the Standardized Precipitation Index(SPI),Standardized Runoff Index(SRI),and Standardized Soil Moisture Index(SSMI),respectively)using the Variable Infiltration Capacity(VIC)model,and quantified the soil conservation service using the Revised Universal Soil Loss Equation(RUSLE)in the historical period(2000-2019)and future period(2026-2060)under two Representative Concentration Pathways(RCPs)(RCP4.5 and RCP8.5).We further examined the influence of the three types of drought on soil conservation service at annual and seasonal scales.The NASA Earth Exchange Global Daily Downscaled Projections(NEX-GDDP)dataset was used to predict and model the hydrometeorological elements in the future period under the RCP4.5 and RCP8.5 scenarios.The results showed that in the historical period,annual-scale meteorological drought exhibited the highest intensity,while seasonal-scale drought was generally weakest in autumn and most severe in summer.Drought intensity of all three types of drought will increase over the next 40 years,with a greater increase under the RCP4.5 scenario than under the RCP8.5 scenario.Furthermore,the intra-annual variation in the drought intensity of the three types of drought becomes smaller under the two future scenarios relative to the historical period(2000-2019).Soil conservation service exhibits a distribution pattern characterized by high levels in the southwest and southeast and lower levels in the north,and this pattern has remained consistent both in the historical and future periods.Over the past 20 years,the intra-annual variation indicated peak soil conservation service in summer and lowest level in winter;the total soil conservation of the Jinghe River Basin displayed an upward trend,with the total soil conservation in 2019 being 1.14 times higher than that in 2000.The most substantial impact on soil conservation service arises from annual-scale meteorological drought,which remains consistent both in the historical and future periods.Additionally,at the seasonal scale,meteorological drought exerts the highest influence on soil conservation service in winter and autumn,particularly under the RCP4.5 and RCP8.5 scenarios.Compared to the historical period,the soil conservation service in the Jinghe River Basin will be significantly more affected by drought in the future period in terms of both the affected area and the magnitude of impact.This study conducted beneficial attempts to evaluate and predict the dynamic characteristics of watershed drought and soil conservation service,as well as the response of soil conservation service to different types of drought.Clarifying the interrelationship between the two is the foundation for achieving sustainable development in a relatively arid and severely eroded area such as the Jinghe River Basin.

Preface to the Special Issue on Causes, Impacts, and Predictability of Droughts for the Past, Present, and Future

Drought is a recurring dry condition with below-normal precipitation and is often associated with warm temperatures or heatwaves. A drought event can develop slowly over several weeks or suddenly within days, commonly under abnormal atmospheric conditions(e.g., quasi-stationary high-pressure systems), and can persist for weeks, months, or even years.

Natural variation of GmFNSII-2 contributes to drought resistance by modulating enzyme activity in soybean

As an essential crop that provides vegetable oil and protein,soybean(Glycine max(L.)Merr.)is widely planted all over the world.However,the scarcity of water resources worldwide has seriously impacted on the quality and yield of soybean.To address this,exploring excellent genes for improving drought resistance in soybean is crucial.In this study,we identified natural variations of GmFNSII-2(flavone synthase II)significantly affect the drought resistance of soybeans.Through sequence analysis of GmFNSII-2 in 632 cultivated and 44 wild soybeans nine haplotypes were identified.The full-length allele GmFNSII-2^(C),but not the truncated allele GmFNSII-2^(A) possessing a nonsense nucleotide variation,increased enzyme activity.Further research found that GmDREB3,known to increase soybean drought resistance,bound to the promoter region of GmFNSII-2^(C).GmDREB3 positively regulated the expression of GmFNSII-2^(C),increased flavone synthase abundance and improved the drought resistance.Furthermore,a singlebase mutation in the GmFNSII-2^(C) promoter generated an additional drought response element(CCCCT),which had stronger interaction strength with GmDREB3 and increased its transcriptional activity under drought conditions.The frequency of drought-resistant soybean varieties with Hap 1(Pro:GmFNSII-2^(C))has increased,suggesting that this haplotype may be selected during soybean breeding.In summary,GmFNSII-2^(C) could be used for molecular breeding of drought-tolerant soybean.

Alternative Splicing of OsCYL4 Controls Drought Resistance via Regulating Water Loss and Reactive Oxygen Species-Scavenging in Rice

A rice cyclase gene,OsCYL4b,identified as an alternative splice variant of the cyclase gene OsCYL4a,is involved in the regulation of drought stress and oxidative response.Compared with OsCYL4a,OsCYL4b lacks the second exon,which is located in the conserved motif 3,and may be a functionally important site.Our results suggested that OsCYL4b was responsive to multiple abiotic stresses,and was localized to both the cytoplasm and plasma membrane.The overexpression of OsCYL4b resulted in significantly enhanced drought and osmotic stress tolerance,reduced water loss,and increased abscisic acid(ABA)content compared with the wild type(WT).

The BEL1-like transcription factor GhBLH5-A05 participates in cotton response to drought stress

Drought stress impairs crop growth and development.BEL1-like family transcription factors may be involved in plant response to drought stress,but little is known of the molecular mechanism by which these proteins regulate plant response and defense to drought stress.Here we show that the BEL1-like transcription factor GhBLH5-A05 functions in cotton(Gossypium hirsutum)response and defense to drought stress.Expression of GhBLH5-A05 in cotton was induced by drought stress.Overexpression of GhBLH5-A05 in both Arabidopsis and cotton increased drought tolerance,whereas silencing GhBLH5-A05 in cotton resulted in elevated sensitivity to drought stress.GhBLH5-A05 binds to cis elements in the promoters of GhRD20-A09 and GhDREB2C-D05 to activate the expression of these genes.GhBLH5-A05 interacted with the KNOX transcription factor GhKNAT6-A03.Co-expression of GhBLH5-A05 and GhKNAT6-A03 increased the transcription of GhRD20-A09 and GhDREB2C-D05.We conclude that GhBLH5-A05 acts as a regulatory factor with GhKNAT6-A03 functioning in cotton response to drought stress by activating the expression of the drought-responsive genes GhRD20-A09 and GhDREB2C-D05.

Drought-triggered repression of miR166 promotes drought tolerance in soybean

Drought stress limits agricultural productivity worldwide.Identifying and characterizing genetic components of drought stress-tolerance networks may improve crop resistance to drought stress.We show that the regulatory module formed by miR166 and its target gene,ATHB14-LIKE,functions in the regulation of drought tolerance in soybean(Glycine max).Drought stress represses the accumulation of miR166,leading to upregulation of its target genes.Optimal knockdown of miR166 in the stable transgenic line GmSTTM166 conferred drought tolerance without affecting yield.Expression of ABA signaling pathway genes was regulated by the miR166-mediated regulatory pathway,and ATHB14-LIKE directly activates some of these genes.There is a feedback regulation between ATHB14-LIKE and MIR166 genes,and ATHB14-LIKE inhibits MIR166 expression.These findings reveal that drought-triggered regulation of the miR166-mediated regulatory pathway increases plants drought resistance,providing new insights into drought stress regulatory network in soybean.

Leaf Morphology Genes SRL1 and RENL1 Co-Regulate Cellulose Synthesis and Affect Rice Drought Tolerance

The morphological development of rice(Oryza sativa L.)leaves is closely related to plant architecture,physiological activities,and resistance.However,it is unclear whether there is a co-regulatory relationship between the morphological development of leaves and adaptation to drought environment.In this study,a drought-sensitive,roll-enhanced,and narrow-leaf mutant(renl1)was induced from a semi-rolled leaf mutant(srl1)by ethyl methane sulfonate(EMS),which was obtained from Nipponbare(NPB)through EMS.Map-based cloning and functional validation showed that RENL1 encodes a cellulose synthase,allelic to NRL1/OsCLSD4.The RENL1 mutation resulted in reduced vascular bundles,vesicular cells,cellulose,and hemicellulose contents in cell walls,diminishing the water-holding capacity of leaves.In addition,the root system of the renl1 mutant was poorly developed and its ability to scavenge reactive oxygen species(ROS)was decreased,leading to an increase in ROS after drought stress.Meanwhile,genetic results showed that RENL1 and SRL1 synergistically regulated cell wall components.Our results revealed a theoretical basis for further elucidating the molecular regulation mechanism of cellulose on rice drought tolerance,and provided a new genetic resource for enhancing the synergistic regulation network of plant type and stress resistance,thereby realizing simultaneous improvement of multiple traits in rice.

Application of Polygonum minus Extract in Enhancing Drought Tolerance in Maize by Regulating Osmotic and Antioxidant System

Drought stress is a major factor affecting plant growth and crop yield production.Plant extracts as natural biostimulants hold great potential to strengthen plants to overcome drought impacts.To explore the effect of Polygonum minus extract(PME)in enhancing drought tolerance in plants,a study was set up in a glasshouse environment using 10 different treatment combinations.PME foliar application were designed in CRD and effects were closely observed related to the growth,physiology,and antioxidant system changes in maize(Zea mays L.)under well-watered and drought conditions.The seaweed extract(SWE)was used as a comparison.Plants subjected to drought stress exhibited a significant reduction in fresh weight,dry weight,relative water content(RWC),and soluble sugar,but they stimulated the phenolic,flavonoid,proline,glutathione(GSH),malondialdehyde(MDA)and antioxidant enzyme(catalase,CAT;peroxidase,POD;superoxide dismutase,SOD)activities.Foliar application of PME improved fresh and dry weight(FW:33.1%~41.4%;DW:48.0%~43.1%),chlorophyll content(Chl b:87.9%~100.76%),soluble sugar(23.6%~49.3%),and soluble protein(48.6%~56.9%)as well as antioxidant enzyme activities(CAT and POD)compared to CK under drought conditions.while decreasing the level of MDA.Notably,the mitigating effect of PME application with high concentration was more effective than those of SWE.Our study reveals that PME could alleviate drought stress by regulating osmoprotectant content and antioxidant defense system and can be used as an economical and environmentally friendly biostimulants for promoting maize growth under drought stress.

Physiology of medicinal and aromatic plants under drought stress

Drought poses a significant challenge,restricting the productivity of medicinal and aromatic plants.The strain induced by drought can impede vital processes like respiration and photosynthesis,affecting various aspects of plants’growth and metabolism.In response to this adversity,medicinal plants employ mechanisms such as morphological and structural adjustments,modulation of drought-resistant genes,and augmented synthesis of secondary metabolites and osmotic regulatory substances to alleviate the stress.Extreme water scarcity can lead to leaf wilting and may ultimately result in plant death.The cultivation and management of medicinal plants under stress conditions often differ from those of other crops.This is because the main goal with medicinal plants is not only to increase the yield of the above-ground parts but also to enhance the production of active ingredients such as essential oils.To elucidate these mechanisms of drought resistance in medicinal and aromatic plants,the current review provides a summary of recent literature encompassing studies on the morphology,physiology,and biochemistry of medicinal and aromatic plants under drought conditions.

More tree growth reduction due to consecutive drought and its legacy effect for a semiarid larch plantation in Northwest China

Extreme climate has increasingly led to negative impacts on forest ecosystems globally,especially in semiarid areas where forest ecosystems are more vulnerable.However,it is poorly understood how tree growth is affected by different drought events.In 2006–2009,the larch plantations in the semiarid areas of Northwest China were negatively affected by four consecutive dry years,which was a very rare phenomenon that may occur frequently under future climate warming.In this study,we analyzed the effect of these consecutive dry years on tree growth based on the data of the tree rings in the dominant layer of the forest canopy on a larch plantation.We found that the tree-ring width index(RWI)in dry years was lower than that in normal years,and it experienced a rapidly decreasing trend from 2006 to 2009(slope=-0.139 year^(-1),r=-0.94)due to water supply deficits in those dry years.Drought induced legacy effects of tree growth reduction,and consecutive dry years corresponded with greater growth reductions and legacy effects.Growth reductions and legacy effects were significantly stronger in the third and fourth consecutive dry years than that of single dry year(p<0.05),which might have been due to the cumulative stress caused by consecutive dry years.Our results showed that larch trees experienced greater tree growth reduction due to consecutive dry years and their legacy effect,and the trees had lower recovery rates after consecutive dry years.Our results highlight that consecutive dry years pose a new threat to plantations under climate warming,and thus,the effect of climate extremes on tree growth should be considered in growth models in semiarid areas.

Annual growth of Fagus orientalis is limited by spring drought conditions in Iran’s Golestan Province

Due to the lack of a uniform and accurate defi-nition of‘drought’,several indicators have been introduced based on different variables and methods,and the efficiency of each of these is determined according to their relationship with drought.The relationship between two drought indices,SPI(standardized precipitation index)and SPEI(standard-ized precipitation-evapotranspiration index)in different sea-sons was investigated using annual rings of 15 tree samples to determine the effect of drought on the growth of oriental beech(Fagus orientalis Lipsky)in the Hyrcanian forests of northern Iran.The different evapotranspiration calcula-tion methods were evaluated on SPEI efficiency based on Hargreaves-Samani,Thornthwaite,and Penman-Monteith methods using the step-by-step M5 decision tree regression method.The results show that SPEI based on the Penman-Monteith in a three-month time scale(spring)had similar temporal changes and a better relationship with annual tree rings(R^(2)=0.81)at a 0.05 significant level.Abrupt change and a decreasing trend in the time series of annual tree rings are similar to the variation in the SPEI based on the Penman-Monteith method.Factors affecting evapotranspiration,temperature,wind speed,and sunshine hours(used in the Penman-Monteith method),increased but precipitation decreased.Using non-linear modeling methods,SPEI based on Penman-Monteith best illustrated climate changes affecting tree growth.

Drought events influence nutrient canopy exchanges and green leaf partitioning during senescence in a deciduous forest

The increase in the frequency and intensity of drought events expected in the coming decades in Western Europe may disturb forest biogeochemical cycles and create nutrient deficiencies in trees.One possible origin of nutrient deficiency is the disturbance of the partitioning of the green leaf pool during the leaf senescence period between resorption,foliar leaching and senesced leaves.However,the effects of drought events on this partitioning and the consequences for the maintenance of tree nutrition are poorly documented.An experiment in a beech forest in Meuse(France)was conducted to assess the effect of drought events on nutrient canopy exchanges and on the partitioning of the green leaf pool during the leaf senescence period.The aim was to identify potential nutritional consequences of droughts for trees.Monitoring nutrient dynamics,including resorption,chemistry of green and senesced leaves,foliar absorption and leaching in mature beech stands from 2012 to 2019 allowed us to compare the nutrient exchanges for three nondry and three dry years(i.e.,with an intense drought event during the growing season).During dry years,we observed a decrease by almost a third of the potassium(K)partitioning to resorption(i.e.resorption efficiency),thus reducing the K reserve in trees for the next growing season.This result suggests that with the increased drought frequency and intensity expected for the coming decades,there will be a risk of potassium deficiency in trees,as already observed in a rainfall exclusion experiment on the same study site.Reduced foliar leaching and higher parititioning to the senesced leaves for K and phosphorus(P)were also observed.In addition,a slight increase in nitrogen(N)resorption efficiency occurred during dry years which is more likely to improve tree nutrition.The calcium(Ca)negative resorption decreased,with no apparent consequence in our study site.Our results show that nutrient exchanges in the canopy and the partitioning of the green leaf pool can be modified by drought events,and may have consequences on tree nutrition.

Leading Pattern of Spring Drought Variability over East Asia and Associated Drivers

Drought events have become more frequent and intense over East Asia in recent decades,leading to huge socioeconomic impacts.Although the droughts have been studied extensively by cases or for individual regions,their leading variability and associated causes remain unclear.Based on the Standardized Precipitation Evapotranspiration Index(SPEI)and ERA5 reanalysis product from 1979 to 2020,this study evealuates the severity of spring droughts in East Asia and investigates their variations and associated drivers.The results indicate that North China and Mongolia have experienced remarkable trends toward dryness during spring in recent decades,while southwestern China has witnessed an opposite trend toward wetness.The first Empirical Orthogonal Function mode of SPEI variability reveals a similar seesawing pattern,with more severe dryness in northwestern China,Mongolia,North China,South Korea,and Japan but increased wetness in Southwestern China and southeast Asia.Further investigation reveals that the anomalously dry(wet)surface in North(Southwestern)China is significantly associated with anomalously high(low)temperature,less(more)precipitation,and reduced(increased)soil moisture during the previous winter and early spring,regulated by an anomalous anticyclone(cyclone)and thus reduced(increased)water vapor convergence.The spring dry-wet pattern in East Asia is also linked to cold sea surface temperature anomalies in the central-eastern Pacific.The findings of this study have important implications for improving the prediction of spring drought events in East Asia.

Grain Yield,Biomass Accumulation,and Leaf Photosynthetic Characteristics of Rice under Combined Salinity-Drought Stress

Simultaneous stresses of salinity and drought often coincide during rice-growing seasons in saline lands,primarily due to insufficient water resources and inadequate irrigation facilities.Consequently,combined salinity-drought stress poses a major threat to rice production.In this study,two salinity levels(NS,non-salinity;HS,high salinity)along with three drought treatments(CC,control condition;DJ,drought stress imposed at jointing;DH,drought stress imposed at heading)were performed to investigate their combined influences on leaf photosynthetic characteristics,biomass accumulation,and rice yield formation.Salinity,drought,and their combination led to a shortened growth period from heading to maturity,resulting in a reduced overall growth duration.Grain yield was reduced under both salinity and drought stress,with a more substantial reduction under the combined salinity-drought stress.The combined stress imposed at heading caused greater yield losses in rice compared with the stress imposed at jointing.Additionally,the combined salinity-drought stress induced greater decreases in shoot biomass accumulation from heading to maturity,as well as in shoot biomass and nonstructural carbohydrate(NSC)content in the stem at heading and maturity.However,it increased the harvest index and NSC remobilization reserve.Salinity and drought reduced the leaf area index and SPAD value of flag leaves and weakened the leaf photosynthetic characteristics as indicated by lower photosynthetic rates,transpiration rates,and stomatal conductance.These reductions were more pronounced under the combined stress.Salinity,drought,and especially their combination,decreased the activities of ascorbate peroxidase,catalase,and superoxide dismutase,while increasing the contents of malondialdehyde,hydrogen peroxide,and superoxide radical.Our results indicated a more significant yield loss in rice when subjected to combined salinity-drought stress.The individual and combined stresses of salinity and drought diminished antioxidant enzyme activities,inhibited leaf photosynthetic functions,accelerated leaf senescence,and subsequently lowered assimilate accumulation and grain yield.

Unexpected Diversity in Ecosystem Nutrient Responses to Experimental Drought in Temperate Grasslands

The responses of ecosystem nitrogen (N) and phosphorus (P) to drought are an important component of globalchange studies. However, previous studies were more often based on site-specific experiments, introducing a significantuncertainty to synthesis and site comparisons. We investigated the responses of vegetation and soil nutrientsto drought using a network experiment of temperate grasslands in Northern China. Drought treatment (66%reduction in growing season precipitation) was imposed by erecting rainout shelters, respectively, at the driest,intermediate, and wettest sites. We found that vegetation nutrient concentrations increased but soil nutrient concentrationsdecreased along the aridity gradient. Differential responses were observed under experimentaldrought among the three grassland sites. Specifically, the experimental drought did not change vegetation andsoil nutrient status at the driest site, while strongly reduced vegetation but increased soil nutrient concentrationsat the site with intermediate precipitation. On the contrary, experimental drought increased vegetation N concentrationsbut did not change vegetation P and soil nutrient concentrations at the wettest site. In general, the differentialeffects of drought on ecosystem nutrients were observed between manipulative and observationalexperiments as well as between sites. Our research findings suggest that conducting large-scale, consistent, andcontrolled network experiments is essential to accurately evaluate the effects of global climate change on terrestrialecosystem bio-geochemistry.

Improved observation of colonized roots reveals the regulation of arbuscule development and senescence by drought stress in the arbuscular mycorrhizae of citrus

Citrus is the typical mycorrhizal fruit tree species establishing symbiosis with arbuscular mycorrhizal (AM) fungi. However, arbuscule development and senescence in colonized citrus roots, especially in response to drought stress, remain unclear, which is mainly due to the difficulty in clearing and staining lignified roots with the conventional method. Here, we improved the observation of colonized roots of citrus plants with the sectioning method, which enabled the clear observation of AM fungal structures. Furthermore, we investigated the effects of one week of drought stress on arbuscule development and senescence with the sectioning method. Microscopy observations indicated that drought stress significantly decreased mycorrhizal colonization (F%and M%) although it did not affect plant growth performance. Fluorescence probes (WGA 488 and/or Nile red) revealed that drought stress inhibited arbuscule development by increasing the percentage of arbuscules at the early stage and decreasing the percentages of arbuscules at the midterm and mature stages. Meanwhile, drought stress accelerated arbuscule senescence, which was characterized by the increased accumulation of neutral lipids. Overall, the sectioning method developed in this study enables the in-depth investigation of arbuscule status, and drought stress can inhibit arbuscule development but accelerate arbuscule senescence in the colonized roots of citrus plants. This study paves the way to elaborately dissecting the arbuscule dynamics in the roots of fruit tree species in response to diverse abiotic stresses.