Abiotic stress treatment reveals expansin like A gene OfEXLA1 improving salt and drought tolerance of Osmanthus fragrans by responding to abscisic acid

Sweet osmanthus(Osmanthus fragrans) is a having general approval aromatic tree in China that is widely applied to landscaping and gardening. However, the evergreen tree adaptability is limited by many environmental stresses. Currently, limited information is available regarding the genetic analysis and functional identification of expansin genes in response to abiotic stress in sweet osmanthus. In this study, a total of 29 expansin genes were identified and divided into four groups by genome-wide analysis from the sweet osmanthus genome. Transcriptome and quantitative Real-time PCR analysis showed that the cell wall-localized protein expansin-like A(OfEXLA1) gene was significantly induced by salt and drought treatment. Histochemical GUS staining of transgenic Arabidopsis lines in which GUS activity was driven with the OfEXLA1 promoter, GUS activity was significantly induced by salt, drought, and exogenous abscisic acid(ABA). In yeast, we found OfEXLA1overexpression significantly improved the population of cells compared with wild-type strains after NaCl and polyethylene glycol(PEG)treatment. Additionally, OfEXLA1 overexpression not only promoted plant growth, but also improved the salt and drought tolerance in Arabidopsis. To gain insight into the role of ABA signaling in the regulation of OfEXLA1 improving abiotic tolerance in sweet osmanthus, four differentially expressed ABA Insensitive 5(ABI5)-like genes(OfABL4, OfABL5, OfABL7, and OfABL8) were identified from transcriptome, and dualluciferase(dual-LUC) and yeast one hybrid(Y1H) assay showed that OfABL4 and OfABL5 might bind to OfEXLA1 promoter to accumulate the OfEXLA1 expression by responding to ABA signaling to improve abiotic tolerance in sweet osmanthus. These results provide the information for understanding the molecular functions of expansin-like A gene and molecular breeding of sweet osmanthus in future.

Rice ONAC016 promotes leaf senescence through abscisic acid signaling pathway involving OsNAP

Senescence-induced NAC(senNAC)TFs play a crucial role in senescence during the final stage of leaf development.In this study,we identified a rice senNAC,ONAC016,which functions as a positive regulator of leaf senescence.The expression of ONAC016 increased rapidly in rice leaves during the progression of dark-induced and natural senescence.The onac016-1 knockout mutant showed a delayed leaf yellowing phenotype,whereas the overexpression of ONAC016 accelerated leaf senescence.Notably,ONAC016 expression was upregulated by abscisic acid(ABA),and thus detached leaves of the onac016-1 mutant remained green much longer under ABA treatment.Quantitative RT-PCR analysis showed that ONAC016 upregulates the genes associated with chlorophyll degradation,senescence,and ABA signaling.Yeast one-hybrid and dual-luciferase assays revealed that ONAC016 binds directly to the promoter regions of OsNAP,a key gene involved in chlorophyll degradation and ABA-induced senescence.Taken together,these results suggest that ONAC016 plays an important role in promoting leaf senescence through the ABA signaling pathway involving OsNAP.

VPD modifies CO_(2)fertilization effect on tomato plants via abscisic acid and jasmonic acid signaling pathways

Atmospheric CO_(2)concentration is elevated globally,which has“CO_(2)fertilization effects”and potentially improves plant photosynthesis,yield,and productivity.Despite the beneficial effect of CO_(2)fertilization being modulated by vapor pressure deficit(VPD),the underlying mechanism is highly uncertain.In the present study,the potential roles of hormones in determining CO_(2)fertilization effects under contrasting high and low VPD conditions were investigated by integrated physiological and transcriptomic analyses.Beneficial CO_(2)fertilization effects were offset under high VPD conditions and were constrained by plant water stress and photosynthetic CO_(2)utilization.High VPD induced a large passive water driving force,which disrupted the water balance and consequently caused plant water deficit.Leaf water potential,turgor pressure,and hydraulic conductance declined under high VPD stress.The physiological evidence combined with transcriptomic analyses demonstrated that abscisic acid(ABA)and jasmonic acid(JA)potentially acted as drought-signaling molecules in response to high VPD stress.Increased foliar ABA and JA content triggered stomatal closure to prevent excessive water loss under high VPD stress,which simultaneously increased the diffusion resistance for CO_(2)uptake from atmosphere to leaf intercellular space.High VPD also significantly increased mesophyll resistance for CO_(2)transport from stomatal cavity to fixation site inside chloroplast.The chloroplast“sink”CO_(2)availability was constrained by stomatal and mesophyll resistance under high VPD stress,despite the atmospheric“source”CO_(2)concentration being elevated.Thus,ABA-and JA-mediated drought-resistant mechanisms potentially modified the beneficial effect of CO_(2)fertilization on photosynthesis,plant growth,and yield productivity.This study provides valuable information for improving the utilization efficiency of CO_(2)fertilization and a better understanding of the physiological processes.

Melatonin Alleviates Abscisic Acid Deficiency Inhibition on Photosynthesis and Antioxidant Systems in Rice under Salt Stress

Melatonin and abscisic acid,as major plant hormones,play important roles in the physiological and biochemical activities of crops,but the interaction between the two under salt stress is not yet clear.This study investigated the endogenous levels of melatonin and abscisic acid in rice by using exogenous melatonin,abscisic acid,and their synthetic inhibitors,and examined their interactions under salt stress.The research results indicate that melatonin and abscisic acid can improve rice salt tolerance.Melatonin alleviated the salt sensitivity caused by abscisic acid deficiency,increased antioxidant enzyme activity and antioxidant content in rice treated with abscisic acid synth-esis inhibitors,and reduced total reactive oxygen species content and thiobarbituric acid reactive substance accu-mulation.Melatonin also increased the activity of key photosynthetic enzymes and the content of photosynthetic pigments,maintaining the parameters of photosynthetic gas exchange and chlorophyllfluorescence.In summary,melatonin alleviated the effects of abscisic acid deficiency on photosynthesis and antioxidant systems in rice and improved salt tolerance.This study is beneficial for expanding the understanding of melatonin regulation of crop salt tolerance.

Overexpression of PbrGA2ox1 enhances pear drought tolerance through the regulation of GA_(3)-inhibited reactive oxygen species detoxification and abscisic acid signaling

Drought stress is a devastating natural disaster driven by the continuing intensification of global warming,which seriously threatens the productivity and quality of several horticultural crops,including pear.Gibberellins(GAs)play crucial roles in plant growth,development,and responses to drought stress.Previous studies have shown significant reductions of GA levels in plants under drought stress;however,our understanding of the intrinsic regulation mechanisms of GA-mediated drought stress in pear remains very limited.Here,we show that drought stress can impair the accumulation of bioactive GAs(BGAs),and subsequently identified PbrGA2ox1 as a chloroplast-localized GA deactivation gene.This gene was significantly induced by drought stress and abscisic acid(ABA)treatment,but was suppressed by GA_(3)treatment.PbrGA2ox1-overexpressing transgenic tobacco plants(Nicotiana benthamiana)exhibited enhanced tolerance to dehydration and drought stresses,whereas knock-down of PbrGA2ox1 in pear(Pyrus betulaefolia)by virus-induced gene silencing led to elevated drought sensitivity.Transgenic plants were hypersensitive to ABA,and had a lower BGAs content,enhanced reactive oxygen species(ROS)scavenging ability,and augmented ABA accumulation and signaling under drought stress compared to wild-type plants.However,the opposite effects were observed with PbrGA2ox1 silencing in pear.Moreover,exogenous GA_(3)treatment aggravated the ROS toxic effect and restrained ABA synthesis and signaling,resulting in the compromised drought tolerance of pear.In summary,our results shed light on the mechanism by which BGAs are eliminated in pear leaves under drought stress,providing further insights into the mechanism regulating the effects of GA on the drought tolerance of plants.

Effects of Abscisic Acid on Cold Resistance in Digitaria sanguinalis(L.)Scop.

[Objectives]This study was conducted to detect the protective effect of abscisic acid on chilling injury of Digitaria sanguinalis(L.)Scop,and whether this effect is related to antioxidant enzymes and osmotic adjustment.[Methods]D.sanguinalis plants were sprayed with abscisic acid solution,and exposed to cold stress at 15℃for 3 d after one day and then at 5℃for 25 to 30 d in a growth chamber.The changes of plant osmotic potential under this treatment were detected.[Results]Under low temperature stress,the osmotic potential of plants in the abscisic acid treatment and the control increased,but the osmotic potential level of the abscisic acid treatment plants was lower.The SOD activity of plants in the ABA treatment and the control decreased under low temperature stress.Under low temperature stress,the activity of catalase and peroxidase in ABA-treated plants was higher than that in control plants.[Conclusions]This study provides a theoretical basis for the impact of abscisic acid on the physiological response of D.sanguinalis to cold injury.

NaCl Facilitates Cell Wall Phosphorus Reutilization in Abscisic Acid Dependent Manner in Phosphorus Deficient Rice Root

Phosphorus(P) starvation in rice facilitates the reutilization of root cell wall P by enhancing the pectin content. NaCl modulates pectin content, however, it is still unknown whether NaCl is also involved in the process of pectin regulated cell wall P remobilization in rice under P starved conditions. In this study, we found that 10 mmol/L NaCl increased the shoot and root biomasses under P deficiency to a remarkable extent, in company with the elevated shoot and root soluble P contents in rice. Further analysis indicated that exogenous NaCl enhanced the root cell wall P mobilization by increasing the pectin methylesterase activity and uronic acid content in pectin suggesting the involvement of NaCl in the process of cell wall P reutilization in P starved rice roots. Additionally, exogenous NaCl up-regulated the expression of P transporter OsPT6, which was induced by P deficiency, suggesting that NaCl also facilitated the P translocation prominently from root to shoot in P starved rice. Moreover, exogenous abscisic acid(ABA) can reverse the NaCl-mediated mitigation under P deficiency, indicating the involvement of ABA in the NaCl regulated root cell wall P reutilization. Taken together, our results demonstrated that NaCl can activate the reutilization of root cell wall P in P starved rice, which is dependent on the ABA accumulation pathway.

Fiber-specific increase of carotenoid content promotes cotton fiber elongation by increasing abscisic acid and ethylene biosynthesis

Cotton fiber is a raw material for the global textile industry and fiber quality is essential to its industrial application.Carotenoids are plant secondary metabolites that may serve as dietary components,regulate light harvesting,and scavenge reactive oxygen species.Although carotenoids accumulate predominantly in rapidly elongating cotton fibers,their roles in cotton fiber development remain poorly understood.In this study,a fiber-specific promoter proSCFP was applied to drive the expression of GhOR1Del,a positive regulator of carotenoid accumulation,to upregulate the carotenoid level in cotton fiber in planta.Fiber length,strength,and fineness were increased in proSCFP:GhOR1Del transgenic cotton and abscisic acid(ABA)and ethylene contents were increased in elongating fibers.The ABA downstream regulator GhbZIP27a stimulated the expression of the ethylene synthase gene GhACO3 by binding to its promoter,suggesting that ABA promoted fiber elongation by increasing ethylene production.These findings suggest the involvement of carotenoids and ABA signaling in promoting cotton fiber elongation and provide a strategy for improving cotton fiber quality.

Identification and Expression Analysis of Abscisic Acid Signal Transduction Genes in Hemp Seeds

Abscisic acid(ABA)is involved in regulating diverse biological processes,but its signal transduction genes and roles in hemp seed germination are not well known.Here,the ABA signaling pathway members,PYL,PP2C and SnRK2 gene families,were identified from the hemp reference genome,including 7 CsPYL(pyrab-actin resistance1-like,ABA receptor),8 CsPP2CA(group A protein phosphatase 2c),and 7 CsSnRK2(sucrose nonfermenting1-related protein kinase 2).The content of ABA in hemp seeds in germination stage is lower than that in non-germination stage.Exogenous ABA(1 or 10μM)treatment had a significant regulatory effect on the selected PYL,PP2C,SnRK2 gene families.CsAHG3 and CsHAI1 were most significantly affected by exogenous ABA treatment.Yeast two-hybrid experiments were performed to reveal that CsPYL5,CsSnRK2.2,and CsSnRK2.3 could interact with CsPP2CA7 and demonstrate that this interaction was ABA-independent.Our results indicated that CsPYL5,CsSnRK2.2,CsSnRK2.3 and CsPP2CA7 might involve in the ABA signaling transduction pathway of hemp seeds during the hemp seed germination stages.This study suggested that novel genetic views can be brought into investigation of ABA signaling pathway in hemp seeds and lay the foundation for further exploration of the mechanism of hemp seed germination.

RNAi-mediated suppression of the abscisic acid catabolism gene Os ABA8ox1 increases abscisic acid content and tolerance to saline–alkaline stress in rice(Oryza sativa L.)

Saline–alkaline(SA) stress is characterized by high salinity and high alkalinity(high p H), which severely inhibit plant growth and cause huge losses in crop yields worldwide. Here we show that a moderate elevation of endogenous abscisic acid(ABA) levels by RNAi-mediated suppression of Os ABA8 ox1(Os ABA8 ox1-kd), a key ABA catabolic gene, significantly increased tolerance to SA stress in rice plants. We produced Os ABA8 ox1-kd lines in two different japonica cultivars, Dongdao 4 and Nipponbare. Compared with nontransgenic control plants(WT), the Os ABA8 ox1-kd seedlings accumulated 25.9%–55.7% higher levels of endogenous ABA and exhibited reduced plasmalemma injury, ROS accumulation and Na;/K;ratio, and higher survival rates, under hydroponic alkaline conditions simulated by 10, 15, and 20 mmol L-1 of Na;CO;. In pot trials using SA field soils of different alkali levels(p H 7.59, 8.86, and 9.29), Os ABA8 ox1–kd plants showed markedly higher seedling survival rates and more vigorous plant growth, resulting in significantly higher yield components including panicle number(85.7%–128.6%), spikelets per panicle(36.9%–61.9%), branches(153.9%–236.7%), 1000–kernel weight(20.0%–28.6%), and percentage of filled spikelets(96.6%–1340.8%) at harvest time. Under severe SA soil conditions(p H = 9.29, EC = 834.4 μS cm-1),Os ABA8 ox1-kd lines showed an 194.5%–1090.8% increase in grain yield per plant relative to WT plants.These results suggest that suppression of Os ABA8 ox1 to increase endogenous ABA levels provides a new molecular approach for improving rice yield in SA paddies.

Exogenous Application of Abscisic Acid (ABA)Enhances Chilling Tolerance in Seedlings ofNapier Grass (Pennisetum purpureum Schum.)

Napier grass, an important forage crop with potentials in multi-purpose applications, is widely grown throughout the tropics and subtropics. Low temperature severely limits its productivity and geographical distribution in temperate regions of the world. In this study, we investigated the effect of exogenous abscisic Acid(ABA) on chilling tolerance of napier grass(Pennisetum purpureum Schum.) seedlings. Seven-day-old napier grass seedlings were cultured in dd H_2O or ABA solution at different concentrations and exposed to 1 ℃ for different time durations. The chilling injury, membrane stability index(MSI) and proline content were estimated from leaf samples. The results showed that there was obvious morphological injury of leaf blighting and restrained growth for the seedlings under chilling stress, but this damage can be largely reduced(by 2/3) when the seedlings were treated by 100 μmol/L ABA in the culture solution, and that the application of exogenous ABA can help to maintain a good stability of leaf cell membrane as expressed by a high MSI value and a low level of proline in leaf cells. These results suggested that exogenous ABA can significantly alleviate chilling injury in napier grass seedlings by maintaining the stability of leaf cell membrane during chilling stress, and that the chilling tolerance was not ensured by a proline accumulation although a passive accumulation of proline was observed in the seedlings under chilling stress. Our results lay a preliminary foundation for future investigations on the molecular mechanisms of ABA induced chilling or freezing tolerance in napier grass.

Research and Applicationon Plant Hormone--Abscisic Acid, ABA

Abscisic Acid (ABA), along with ethylene, gibberellins, cytokinins and auxins, is regarded as five kinds of important plant hormone. ABA was first isolated from cotton bud by Addcott Ohhuma’s group in 1963. Until 1965, its plane structure was determined. It was formally named as Abscisic acid in "the International Conference of Plant Regulator" in 1967. Scientists all over the world have made a long-term unremitting effort

In Vivo Tissue-dependent Abscisic Acid Specific-binding Proteins with High Affinity in Cytosol of Developing Apple Fruits

The in vivo highly tissue-dependent abscisic acid (ABA) specific-binding sites localized in cytosol were identified and characterized in the flesh of developing apple ( Malus pumila L. cv. Starkrimon) fruits. ABA binding activity was scarcely detectable in the microsomes and the cytosolic fraction isolated from the freshly harvested fruits via an in vitro ABA binding incubation of the subcellular fractions. If, however, instead that the subcellular fractions were in vitro incubated in H-3-ABA binding medium, the flesh tissue discs were directly in vivo incubated in H-3-ABA binding medium, a high ABA binding activity to the cytosolic fraction isolated from these tissue discs was detected. The in vivo ABA binding capacity of the cytosolic fraction was lost if the tissue discs had been pretreated with boiling water, indicating that the ABA binding needs a living state of tissue. The in vivo tissue-dependent binding sites were shown to possess protein nature with both active serine residua and thiol-group of cysteine residua in their functional binding center. The ABA binding of the in vivo tissue-dependent ABA binding sites to the cytosolic fraction was shown to be saturable, reversible, and of high affinity. The scatchard plotting gave evidence of two different classes of ABA binding proteins, one with a higher affinity ( Kd = 2.9 nmol/L) and the other with lower affinity ( Kd = 71.4 nmol/L). Phaseic acid, 2-trans-4-trans-ABA or cis-trans-(-)-ABA had substantially no affinity to the binding proteins, indicating their stereo-specificity to bind physiologically active ABA. The time course, pH- and temperature-dependence of the in vivo tissue-dependent binding proteins were determined. It is hypothesized that the detected ABA-binding proteins may be putative ABA-receptors that mediate ABA signals during fruit development.

Effects of Exogenous Abscisic Acid and Water Stress on the Growth Response of Subterranean Clover of Different Genotypes

The response of subterranean clover ( Trifolium subterraneum L.) to different abscisic acid (ABA) concentrations (10 -6 , 10 -5 , 10 -4 mol/L) in nutrient solution and to water stress were studied in growth room. Six cultivars of contrasting yield capacity were compared. Plants remained growing in Hoagland solution or pots until at least four full_developed leaves appeared. The ABA was then applied and the fresh weight, leaf number and length of the largest root were measured at 1, 4, 7 and 11 d after ABA treatment. The above parameters were also measured at 15 d under water stress. In all the tested genotypes ABA caused similar reduction in these growth parameters, as well as a significant decrease of leaf water potential which was dependent on ABA concentration. The average growth reduction after 11 d under 10 -4 mol/L ABA coincided with the range of these crops under water stress in pot experiments. On average of the different genotypes, leaf number, area of full_developed leaf and the dry weight per plant decreased by about 50% whereas the root/shoot ratio increased by 80%.The genotype variation and ranking for this treatment were rather similar to the same genotypes in pot experiments. The genotypes, Clare, Nuba and Seaton Park, showed the best results under both control and ABA treated conditions and water stress conditions. The similarity between the response to ABA in nutrient solution and to water stress opens the possibility to use this approach as a way to quantify the drought resistance of subterranean clover genotypes.

Exogenous Application of Abscisic Acid or Gibberellin Acid Has Different Effects on Starch Granule Size Distribution in Grains of Wheat

Granule size distribution of wheat starch is an important characteristic that can affect its chemical composition and the functionality of wheat products. Two high-yield winter wheat cultivars were used to evaluate the effects of the application of exogenous ABA or GA during the reproductive phase of the initial grain filling on starch granule size distribution and starch components in grains at maturity. The results indicated that a bimodal curve was found in the volume and surface area distribution of grain starch granules, and a unimodal curve was observed for the number distribution under all treatments. The exogenous ABA resulted in a significant increase in the proportions （both by volume and by surface area） of B-type （〈9.9 Ixm in diameter） starch granules, with a reduction in those of A-type （〉9.9 ~tm） starch granules, while, the exogenous GA3 led to converse effects on size distribution of those starch granules. The exogenous ABA also increased starch, amylose and amylopectin contents at maturity but significantly reduced the ratio of amylose to amylopectin. Application of GA3 significantly reduced starch content, amylopectin content but increased the ratio of amylose to amylopectin. The ratio of amylose to amylopectin showed a significant and negative relationship with the volume proportion of granules 〈9.9 μm, but was positively related to the volume proportion of granules 22.8-42.8 μm.

A novel impedance immunosensor based on O-phenylenediamine modified gold electrode to analyze abscisic acid

An impedance immunosensor based on O-phenylenediamine modified gold electrode for the determination of phytohormone abscisic acid(ABA) was proposed.The operating pH,absorption time,absorption temperature and concentration of anti-ABA antibody were investigated to optimize the analytical performance.The calibration curve for the determination of ABA was obtained from this impedance immunosensor under optimal conditions.The results showed that the detection limit at about 1 ng/mL in the range of 10-5000 ng/mL...

Effects of Formulated Fertilizer Synergist on Abscisic Acid Accumulation, Proline Content and Photosynthetic Characteristics of Rice under Drought

To investigate the effects of formulated fertilizer synergist on the drought tolerance in rice, pot experiment was conducted to analyze the photosynthetic characteristics and the accumulation of abscisic acid （ABA） and proline in middle-season rice variety Peiliangyou 93. The synergist could improve the net photosynthetic rate, and coordination between the water loss and the CO2 absorption as well as reduce the harmful effect on photosynthetic process under drought conditions. Under drought, the ABA accumulated massively both in roots and leaves, while the ABA content in roots was far higher than that in leaves. The results indicate that synergist could increase the ABA accumulation, but reduce the proline accumulation in rice plant under drought.

K^+channels inhibited by hydrogen peroxide mediate abscisic acid signaling in Vicia guard cells

A number of studies show that environmental stress conditions increase abscisic acid (ABA) and hydrogen peroxide (H2O2) levels in plant cells. Despite this central role of ABA in altering stomatal aperture by regulating guard cell ion transport, little is known concerning the relationship between ABA and H2O2 in signal transduction leading to stomatal movement. Epidermal strip bioassay illustrated that ABA- inhibited stomatal opening and ABA-induced stomatal closure were abolished partly by externally added catalase (CAT) or diphenylene iodonium (DPl), which are a H2O2 scavenger and a NADPH oxidase inhibitor respectively. In contrast, internally added CAT or DPI nearly completely or partly reversed ABA-induced closure in half-stoma. Consistent with these results, whole-cell patch-clamp analysis showed that intracellular application of CAT or DPI partly abolished ABA-inhibited inward K+ current across the plasma membrane of guard cells. H2O2 mimicked ABA to inhibit inward K+ current, an effect which was reversed by the addition of ascorbic acid (Vc) in patch clamping micropipettes. These results suggested that H2O2 mediated ABA-induced stomatal movement by targeting inward K+ channels at plasma membrane.

Effects of Exogenous Trehalose on the Metabolism of Sugar and Abscisic Acid in Tomato Seedlings Under Salt Stress

Salt stress a ects the growth and development of plants, which results in a decrease in crop quality and yield. In this study, we used tomato seedlings treated with salt and trehalose as experimental materials and analyzed them using the technique for order preference by similarity to ideal solution analysis to select the optimal trehalose concentration for treatment. We also determined the contents of sugar and abscisic acid (ABA) and detected the expression of genes involved in the metabolism of sugar and ABA by quantitative real-time PCR. Results showed that the optimal trehalose concentration was 2 mmol/L for tomato seedlings under salt stress. Exogenous trehalose decreased the starch content and increased the soluble sugar con- tent by a ecting the expression of genes related to the metabolism of starch and soluble sugar. Exogenous trehalose altered the accumulation and distribution of sugar by inducing the upregulation of sugar transporter genes. Furthermore, trehalose increased the ABA content to induce salt stress response by regulating the expression of genes related to the synthesis and metabolism of ABA. In conclusion, trehalose can e ectively alleviate salt stress and enhance salt tolerance of tomato. These ndings provide a novel perspective and a better resource to investigate the salt tolerance mechanism and a new method for alleviating salt stress in tomato.

Abscisic acid and jasmonic acid are involved in drought priming-induced tolerance to drought in wheat

Drought stress is a limiting factor for wheat production and food security.Drought priming has been shown to increase drought tolerance in wheat.However,the underlying mechanisms are unknown.In the present study,the genes encoding the biosynthesis and metabolism of abscisic acid(ABA)and jasmonic acid(JA),as well as genes involved in the ABA and JA signaling pathways were up-regulated by drought priming.Endogenous concentrations of JA and ABA increased following drought priming.The interplay between JA and ABA in plant responses to drought priming was further investigated using inhibitors of ABA and JA biosynthesis.Application of fluridone(FLU)or nordihydroguaiaretic acid(NDGA)to primed plants resulted in lower chlorophyll-fluorescence parameters and activities of superoxide dismutase and glutathione reductase,and higher cell membrane damage,compared to primed plants(PD)under drought stress.NDGA+ABA,but not FLU+JA,restored priming-induced tolerance,as indicated by a finding of no significant difference from PD under drought stress.Under drought priming,NDGA induced the suppression of ABA accumulation,while FLU did not affect JA accumulation.These results were consistent with the expression of genes involved in the biosynthesis of ABA and JA.They suggest that ABA and JA are required for priming-induced drought tolerance in wheat,with JA acting upstream of ABA.