Drought stress reduces the feeding preference of Nilaparvata lugens due to the accumulation of abscisic acid and callose deposition in rice

The incidence of drought stress in plants has been increasing due to global warming, and the phytohormone abscisic acid (ABA) induces the formation of physical barriers in plants, such as callose accumulation. The brown planthopper (BPH; Nilaparvata lugens Stål) occurs throughout Asia and feeds on rice, but the effects of drought stress on BPH feeding remain unclear. In this study, we observed changes in callose formation and ABA content in rice during drought stress. ABA content and the relative expression of ABA synthesis genes OsNCED3 and OsNCED5 were higher in drought-stressed rice than the non-stressed control. Similarly, the expression levels of callose synthesis genes and callose deposition were significantly higher in drought-stressed rice as compared to non-stressed plants, and this impacted BPH feeding. Our results indicated that rice resistance to BPH increased during drought stress due to the accumulation of callose and increasing ABA levels. Our findings provide a basis for understanding BPH feeding performance on rice during drought stress and offer novel insights relative to control during periods of water shortage.     

AM1 is a potential ABA substitute for drought tolerance as revealed by physiological and ultra-structural responses of oilseed rape

Abscisic acid (ABA) is an important signaling molecule for plants under drought tolerance. However, ABA itself has many limitations to be used in agriculture practically. Recently, AM1 (ABA-mimicking ligand) has been found to replace ABA. In this study, we have investigated AM1’s potential role for drought tolerance by growing two contrasting rapeseed (Brassica napus L.) genotypes: Qinyou 8 (drought sensitive) and Q2 (drought resistant) with exogenous ABA or AM1 application under well-watered and drought-stressed conditions. Results demonstrate that drought stress has hampered plant growth (relative height growth rate, plant biomass, leaf area), plant water status (leaf relative water content, root moisture content, leaf water potential), photosynthetic gas exchange attributes like net photosynthesis rate (Pn), stomatal conductance (Gs), intercellular CO₂ concentration (Ci), transpiration rate (E); chlorophyll fluorescence parameters like photosynthetic efficiency (Fv/Fm), effective quantum yield of PSII (Φ _PSII ), photochemical quenching coefficient (qL), electron transport rate (ETR) and chlorophyll content, especially for Qinyou 8 significantly compared to well-watered plants. Whereas increased root/shoot ratio (R/S), water use efficiency (WUE) and non-photochemical quenching (NPQ) was recorded in both genotypes under drought stress. On the other hand, exogenous ABA or AM1 treatment has regulated all the above parameters in a rational way to avoid drought stress. Chloroplast transmission electron microscope images, especially for Qinyou8, have revealed that oxidative stress induced by drought has blurred the grana thylakoids, increased the size or number of plastoglobules due to lipid peroxidation, and the presence of starch granules depict weak capacity to convert them into simple sugars for osmotic adjustment. However, intact grana thylakoid, few plastoglobules with no or very few starch granules were observed in the chloroplast from ABA- or AM1-treated plants under drought. More importantly, AM1-treated plants under drought stress have responded in an extremely similar way like ABA-treated ones. Finally, it is suggested that AM1 is a potential ABA substitute for plant drought tolerance.     

ABA Inducible Rice Protein Phosphatase 2C Confers ABA Insensitivity and Abiotic Stress Tolerance in Arabidopsis

Arabidopsis PP2C belonging to group A have been extensively worked out and known to negatively regulate ABA signaling. However, rice (Oryza sativa) orthologs of Arabidopsis group A PP2C are scarcely characterized functionally. We have identified a group A PP2C from rice (OsPP108), which is highly inducible under ABA, salt and drought stresses and localized predominantly in the nucleus. Genetic analysis revealed that Arabidopsis plants overexpressing OsPP108 are highly insensitive to ABA and tolerant to high salt and mannitol stresses during seed germination, root growth and overall seedling growth. At adult stage, OsPP108 overexpression leads to high tolerance to salt, mannitol and drought stresses with far better physiological parameters such as water loss, fresh weight, chlorophyll content and photosynthetic potential (Fv/Fm) in transgenic Arabidopsis plants. Expression profile of various stress marker genes in OsPP108 overexpressing plants revealed interplay of ABA dependent and independent pathway for abiotic stress tolerance. Overall, this study has identified a potential rice group A PP2C, which regulates ABA signaling negatively and abiotic stress signaling positively. Transgenic rice plants overexpressing this gene might provide an answer to the problem of low crop yield and productivity during adverse environmental conditions.     

OsASR5 enhances drought tolerance through a stomatal closure pathway associated with ABA and H2O2 signalling in rice

Drought is one of the major abiotic stresses that directly implicate plant growth and crop productivity. Although many genes in response to drought stress have been identified, genetic improvement to drought resistance especially in food crops is showing relatively slow progress worldwide. Here, we reported the isolation of abscisic acid, stress and ripening (ASR) genes from upland rice variety, IRAT109 (Oryza sativa L. ssp. japonica), and demonstrated that overexpression of OsASR5 enhanced osmotic tolerance in Escherichia coli and drought tolerance in Arabidopsis and rice by regulating leaf water status under drought stress conditions. Moreover, overexpression of OsASR5 in rice increased endogenous ABA level and showed hypersensitive to exogenous ABA treatment at both germination and postgermination stages. The production of H₂O₂, a second messenger for the induction of stomatal closure in response to ABA, was activated in overexpression plants under drought stress conditions, consequently, increased stomatal closure and decreased stomatal conductance. In contrast, the loss‐of‐function mutant, osasr5, showed sensitivity to drought stress with lower relative water content under drought stress conditions. Further studies demonstrated that OsASR5 functioned as chaperone‐like protein and interacted with stress‐related HSP40 and 2OG‐Fe (II) oxygenase domain containing proteins in yeast and plants. Taken together, we suggest that OsASR5 plays multiple roles in response to drought stress by regulating ABA biosynthesis, promoting stomatal closure, as well as acting as chaperone‐like protein that possibly prevents drought stress‐related proteins from inactivation.     

Chloroplast protein PLGG1 is involved in abscisic acid-regulated lateral root development and stomatal movement in Arabidopsis

The plant hormone abscisic acid (ABA) plays a crucial role in root architecture; however, the molecular mechanism of ABA-regulated lateral root (LR) growth is not well known. We screened an Arabidopsis thaliana mutant with LR growth that was sensitive to ABA from a T-DNA insertion mutant library, which was an allelic mutant of plgg1-1, termed plgg1-2. PLGG1 encodes a chloroplast protein that transports plastidic glycolate and glycerate. The length and number of LRs at the root-hypocotyl junction of plgg1-1 and plgg1-2 were significantly impaired under exogenous ABA treatment, and the transgenic plant complementary lines of plgg1-2 restored LR growth in response to ABA. In addition, we found that PLGG1 is involved in other major ABA responses, including ABA-inhibited seed germination, ABA-mediated stomatal movement, and drought tolerance. These findings open new perspectives on elucidating the mechanism of ABA response, and provide clues for analysing the functions of chloroplast proteins in regulating root growth.     

Time course analysis of ABA and non-ionic osmotic stress-induced changes in water status,chlorophyll fluorescence and osmotic adjustment in Arabidopsis thaliana wild-type (Columbia) and ABA-deficient mutant (aba2)

In the present study, we investigated time course changes of water status including relative water content (RWC), leaf osmotic potential (Ψ_Π), stomatal conductance (g_s), proline (Pro), chlorophyll fluorescence (F_v/F_m) and total chlorophyll content in the Arabidopsis thaliana under PEG-induced drought stress after exogenous ABA treatment. To a better explanation for the role of ABA in the water status of A. thaliana to drought stress, wild-type (Columbia) and ABA-deficient mutant (aba2) of A. thaliana were used in the present study. Moreover, three weeks old Arabidopsis seedlings were applied exogenously with 50 μM ABA and exposed to drought stress induced by 40% PEG8000 (−0.73 MPa) for 6 h, 12 h and 24 h (hours). Our findings indicate that RWC of wild-type and aba2 started to decrease in the first 12 h and 6 h of PEG-induced drought stress, respectively. However, exogenous treatment of 50 μM ABA increased their RWC under drought stress. On the other hand, while Ψ_Π of both genotypes started to decrease in the first 6 h of drought stress, these declines in Ψ_Π were prevented by ABA treatment under stress throughout the experiment; it was more pronounced in aba2 at 24 h. While the highest increase in g_s was obtained in aba2 after 24 h stress, ABA-induced highest decrease in g_s was obtained in the same genotype during 12 h, as compared to PEG-treated group alone. On the other hand, Pro content increased in all treatment groups of ABA-deficient mutant aba2 at 12 h and 24 h. However, Pro content in ABA + PEG treated aba2 plants was higher than in PEG- and ABA-treated plants alone at the end of the 24 h. Drought stress decreased F_v/F_m and total chlorophyll contents of both genotypes while 50 μM ABA alleviated these reductions during drought stress, as compared to PEG stressed plants. On the other hand, 50 μM ABA treatment alone did not create any remarkable effect on F_v/F_m and total chlorophyll contents.These findings indicate that exogenous ABA showed an alleviative effect against damage of drought stress on relative water content, osmotic potential, stomatal conductance, proline, chlorophyll fluorescence and total chlorophyll content of both genotypes during 24 h of drought stress treatment.     

Effect of drought and ABA on growth, photosynthesis and antioxidant system of Cotinus coggygria seedlings under two different light conditions

We exposed seedlings of Cotinus coggygria var. cinerea to drought and exogenous abscisic acid (ABA) under two different light conditions. Two watering regimes (well-watered and drought), two exogenous ABA applications (no ABA and with ABA) and two light regimes (full sunlight and shade) were employed. Compared with well-watered treatment, drought treatment significantly reduced the relative growth rate, relative water content (RWC), net photosynthesis rate (A) and transpiration (E), but increased chlorophyll a (chla), carbon isotope (δ¹³C), endogenous ABA, malondialdehyde (MDA) and hydrogen peroxide (H₂O₂) contents, and guaiacol peroxidase (POD) and catalase (CAT) activities. There was an apparent alleviation of drought effects by shade, as indicated by the lower relative growth rate, and chlorophyll, MDA and H₂O₂ contents, and increases in indoleacetic acid (IAA) and reduced glutathione (GSH) contents. On the other hand, the exogenous ABA application under shade induced protective effects on drought-stressed seedlings, as visible in RWC, MDA, A, stomatal conductance (g_s), E, δ¹³C, ABA and IAA values. In all, our results suggest that seedlings of C. coggygria are more sensitive to drought under full-light than under shade.     

Photosynthetic characteristics and effects of exogenous glycine of Chorispora bungeana under drought stress

We investigated the photosynthetic characteristics of Chorispora bungeana under conditions of drought stress caused by different concentrations of polyethylene glycol-6000 (PEG; 0, 5, 20, and 40%) and various concentrations of exogenous glycine (0, 5, 10, and 20 mM) with 20% PEG. We showed that moderate and severe drought stress of PEG reduced the chlorophyll (Chl) content (both Chl a and b), maximal quantum yield of PSII photochemistry (F_v/F_m), actual photochemical efficiency of PSII in light (Y_II), and quantum yield of regulated energy dissipation (Y_NPQ), while Chl a/b and quantum yield of nonregulated energy dissipation (Y_NO) increased. The low and moderate drought stress increased Mg²⁺ and Fe³⁺ contents, while a decrease in Mg²⁺ and Fe³⁺ was found under severe drought stress. Compared to sole PEG stress, the addition of exogenous 10 mM glycine increased Chl, Mg²⁺ and Fe³⁺ contents, F_v/F_m, Y_II, and Y_NPQ, and reduced Y_NO. On the contrary, 20 mM glycine showed an opposite effect, except for Y_NO. Our results proved that Chl contents and fluorescence parameters are reliable indicators for drought tolerance of C. bungeana. We suggest that a proper glycine content can relieve the effect of drought stress on C. bungeana.     

Carotenoid deficiency impairs ABA and IAA biosynthesis and differentially affects drought and cold tolerance in rice

Plant responses to abiotic stresses are coordinated by arrays of growth and developmental programs. Phytohormones such as abscisic acid (ABA) and indole-3-acetic acid (IAA) play critical roles in developmental progresses and environmental responses through complex signalling networks. However, crosstalk between the two hormones at the biosynthesis level remains largely unknown. Here, we report that carotenoid-deficient mutants (phs1, phs2, phs3-1, phs4, and PDS-RNAi transgenic rice) were impaired in the biosynthesis of ABA and IAA. Under drought conditions, phs3-1 and PDS-RNAi transgenic rice showed larger stomata aperture and earlier wilting compared to the wild type at both seedling and panicle developmental stage. Interestingly, these carotenoid-deficient lines showed increased cold resistance, which was likely due to the combined effects of reduced IAA content, alleviated oxidative damage and decreased membrane penetrability. Furthermore, we found that IAA content was significantly declined in rice treated with fluridone (a carotenoid and ABA biosynthesis inhibitor), and expression of auxin synthesis and metabolism-related genes were altered in the fluridone-treated rice similar to that in the carotenoid-deficient mutants. In addition, exogenous IAA, but not ABA, could restore the dwarf phenotype of phs3-1 and PDS-RNAi transgenic rice. These results support a crosstalk between ABA and IAA at the biosynthesis level, and this crosstalk is involved in development and differentially affects drought and cold tolerance in rice.     

Abscisic Acid Induces Rapid Reductions in Mesophyll Conductance to Carbon Dioxide

The rate of photosynthesis (A) of plants exposed to water deficit is a function of stomatal (g_s) and mesophyll (g_m) conductance determining the availability of CO₂ at the site of carboxylation within the chloroplast. Mesophyll conductance often represents the greatest impediment to photosynthetic uptake of CO₂, and a crucial determinant of the photosynthetic effects of drought. Abscisic acid (ABA) plays a fundamental role in signalling and co-ordination of plant responses to drought; however, the effect of ABA on g_m is not well-defined. Rose, cherry, olive and poplar were exposed to exogenous ABA and their leaf gas exchange parameters recorded over a four hour period. Application with ABA induced reductions in values of A, g_s and g_m in all four species. Reduced g_m occurred within one hour of ABA treatment in three of the four analysed species; indicating that the effect of ABA on g_m occurs on a shorter timescale than previously considered. These declines in g_m values associated with ABA were not the result of physical changes in leaf properties due to altered turgor affecting movement of CO₂, or caused by a reduction in the sub-stomatal concentration of CO₂ (C_i). Increased [ABA] likely induces biochemical changes in the properties of the interface between the sub-stomatal air-space and mesophyll layer through the actions of cooporins to regulate the transport of CO₂. The results of this study provide further evidence that g_m is highly responsive to fluctuations in the external environment, and stress signals such as ABA induce co-ordinated modifications of both g_s and g_m in the regulation of photosynthesis.     

The influence of ABA on water relation, photosynthesis parameters, and chlorophyll fluorescence under drought conditions in two maize hybrids with contrasting drought resistance

Drought is a major limitation of maize cultivation in Brazil. Agronomic and physiological practices have been considered to overcome this stress and consequently, increase grain production. The present study investigated the role of abscisic acid (ABA) application in some physiological parameters, in two hybrids with contrasting drought resistance (DKB 390 and BRS 1030 resistant and sensitive, respectively). Contrasting resistance to drought in these genotypes was determined in previous studies. Water deficit was imposed for 10 days at flowering stage, in association with the application of 100 μM abscisic acid on plant canopy. Evaluations of gas exchange, chlorophyll fluorescence, relative water content (RWC), and endogenous ABA content were performed during stress period and also at water recovery (recovery irrigation). A significant functional relationship was observed between RWC and the parameters of gas exchange and fluorescence. During water recovery, no differences were observed among the treatments. DKB 390 presented higher photosynthesis rate (P _n) and electron transport rate (ETR) under water stress, while BRS 1030 presented higher intercellular CO₂ concentration (C _i) and lower photochemical quenching (qP), non-photochemical quenching (NPQ), and lower F _v/F _m ratio. DBK 390 was more responsive to ABA application than BRS 1030, presenting higher endogenous ABA content in the first day of stress. DBK 390 with ABA application reduced the effect of water stress through maintenance of water status, an increase of photosynthetic parameters, and a decrease of decline in the functions of photosystem II during stress.     

Brassinolide alleviates the drought-induced adverse effects in barley by modulation of enzymatic antioxidants and ultrastructure

In the present study, possible role of exogenously applied brassinolide (BL) in alleviating the detrimental effects of polyethylene glycol (PEG)-inducted drought stress in barley was evaluated. Barley seedlings were subjected to two PEG levels (0 and 8% w/v) for 2 weeks followed by a foliar spray of BL (0, 0.01, 0.1 and 1 µM) to assess the changes in growth, gas exchange, chlorophyll contents, antioxidant enzyme activities and ultrastructural changes. PEG-induced drought considerably inhibited the barley growth in terms of shoot and root length, shoot, root fresh and dry weights, and chlorophyll contents. Moreover, extensive decrease by PEG-induced drought in gas exchange attributes i.e. net photosynthetic rate, transpiration rate, stomatal conductance and intercellular CO₂ were also recorded. However, exogenous application of BL remarkably improved the gas exchange attributes, chlorophyll contents and plant growth under either drought or control condition. Furthermore, foliar spray of BL decreased the production of malondialdehyde (MDA) and reactive oxygen species (ROS) in the leaves and roots by increasing enzymatic antioxidant activities under drought stress. The microscopic examination indicated that exogenous application of BL improved the cell structure and enabled a clean mesophyll cell having a well-developed chloroplast with thylakoid membranes of the drought stressed plants. On the basis of these findings, it can be concluded that exogenous application of BL could alleviate drought stress to barley by ameliorating photosynthetic parameters, improving antioxidants enzyme activities, reducing ultrastructural changes in leaf mesophyll and finally maintaining the plant growth.     

脱落酸和己糖激酶抑制剂对高表达C4-PEPC转基因稻苗耐旱性的影响

为了研究高表达转玉米C₄-磷酸烯醇式丙酮酸羧化酶（phosphoenolpyruvate carboxylase,PEPC）基因水稻（PC）的耐旱性机制,本研究以PC和未转基因野生型原种kitaake为材料,分别在光照和黑暗预处理24 h,其中光照处理中光强为600 μmol·m^-2·s^-1,预处理后稻苗再在15％聚乙二醇-6000模拟干旱胁迫下,同时使用药理学的方法,通过加入脱落酸和己糖激酶的专一性抑制剂100 μmol·L^-1去甲二氢愈创木酸和10 mmol·L^-1葡萄糖胺,观察两种水稻4~5叶期稻苗耐旱表现。结果发现,与WT水稻相比,在PEG-6000处理后,经过光预处理的PC水稻叶片相对含水量下降较少,始终比WT的高,而且丙二醛含量则较少,脯氨酸诱导增加,表现耐旱;而经过暗预处理后PC植株显著降低这个优势,表明光预处理有利于PC耐旱性的表现;黑暗预处理均显著下调了2供试材料植株叶片中可溶性糖的含量,而对可溶性蛋白的含量影响不显著;而光预处理后PC水稻叶片内可溶性糖含量比WT增加,而在黑暗预处理则PC的显著低于WT的,其中葡萄糖胺对光预处理下PC的可溶性糖含量的下调作用最显著;暗预处理逆转或消除了NO,H₂O₂和钙离子含量变化趋势,这些变化与暗预处理减少了两材料叶片蔗糖和葡萄糖含量变化同步;光暗预处理对两材料的PEPC酶活性的差异影响不大,表明外源玉米C₄-PEPC在水稻中是组成型表达。可见PC叶片可部分通过糖组分,参与内源糖介导ABA和HXK信号途径,缓解干旱处理对叶片的伤害,稳定光合能力。     

Silicon Alleviates Drought Stress of Rice Plants by Improving Plant Water Status,Photosynthesis and Mineral Nutrient Absorption

Drought is a major constraint for rice production in the rainfed lowlands in China. Silicon (Si) has been verified to play an important role in enhancing plant resistance to environmental stress. Two near-isogenic lines of rice (Oryza sativa L.), w-14 (drought susceptible) and w-20 (drought resistant), were selected to study the effects of exogenous Si application on the physiological traits and nutritional status of rice under drought stress. In wet conditions, Si supply had no effects on growth and physiological parameters of rice plants. Drought stress was found to reduce dry weight, root traits, water potential, photosynthetic parameters, basal quantum yield (F _v/F ₀), and maximum quantum efficiency of PSII photochemistry (F _v/F _m) in rice plants, while Si application significantly increased photosynthetic rate (Pr), transpiration rate (Tr), F _v/F ₀, and F _v/F _m of rice plants under drought stress. In addition, water stress increased K, Na, Ca, Mg, Fe content of rice plants, but Si treatment significantly reduced these nutrient level. These results suggested that silicon application was useful to increase drought resistance of rice through the enhancement of photochemical efficiency and adjustment of the mineral nutrient absorption in rice plants.     

Downregulation of OsPK1 Contributes to Oxidative Stress and the Variations in ABA/GA Balance in Rice

Abscisic acid (ABA) is a phytohormone that aids plants in coping with stress conditions. ABA and gibberellin (GA) are hormone partners that function via a complicated and antagonistic network. In ospk1, a dwarf rice mutant, the contents of ABA in the youngest leaf sheaths of 6-week-old seedlings and the uppermost internodes of heading stage plants were both increased, and the synthesis of bioactive GAs was suppressed, which may disharmonize ABA/GA balance. In ospk1, expression of three putative enzyme genes related to stress response was upregulated. A strong browning symptom was observed in the second internode (Int2, counted from the top) and part of panicles of ospk1 at the late productive phase. Furthermore, higher levels of H₂O₂ in flag leaf and Int2 were observed in ospk1 than those in wild type. These data suggest that ospk1 may undergo certain stress, especially oxidative stress. Here, we provide evidences that the downregulation of OsPK1 (a cytosolic pyruvate kinase) in ospk1 mutant results in variations in ABA/GA balance in rice and contributes to oxidative stress, which provide a new clue for understanding the connection of pyruvate kinase, ABA/GA balance, and oxidative stress in rice.