Responses of the antioxidant defense system to drought stress in the leaves of Fargesia denudata seedlings,the staple food of the giant panda

The responses of the antioxidant defense system in plant species to drought stress are still relatively unknown. In order to further understand how the system responds to drought stress, the leaves of Fargesia denudata seedlings were investigated. Antioxidant enzyme activities, antioxidant contents, hydrogen peroxide (H₂O₂), superoxide anion (O ₂ ^·? ) and MDA contents in the seedling leaves were measured under well-watered (WW), moderate drought-stressed (MD), and severe drought-stressed (SD) treatments. Although drought stress significantly increased H₂O₂ and O ₂ ^·? levels in F. denudata leaves, only weak lipid peroxidation was observed. This is attributed to the higher superoxide dismutase (SOD), catalase (CAT), ascorbate peroxidase (APX), glutathione reductase (GR), monodehydroascorbate reductase (MDHAR), and dehydroascorbate reductase (DHAR) activities in F. denudata leaves during the entire drought period. Reduced and oxidized ascorbate (AsA and DHA) contents were almost not affected by drought except that DHA under SD showed an obvious increase on day 30. Furthermore, reduced glutathione (GSH) content under drought stress significantly decreased, while oxidized glutathione (GSSG) markedly increased under SD on days 30 and 45 as well as under MD on day 30; as a result, the ratio GSH/GSSG declined considerably. These results indicated that GSH was involved in scavenging H₂O₂ and O ₂ ^·? under drought stress and it was more sensitive to drought stress in scavenging H₂O₂ and O ₂ ^·? than AsA. As a result, a highly efficient antioxidant defense system in drought-stressed F. denudate leaves operated mainly through the synergistic functioning of SOD, CAT, APX, MDHAR, DHAR, GR, and GSH against oxidative damage.     

Effects of silicon on defense of wheat against oxidative stress under drought at different developmental stages

The effects of silicon application before sowing on the drought-induced oxidative stress and antioxidant defense in wheat (Triticum aestivum L.) were investigated. Drought stress was applied by withholding watering till sampling at booting or filling stage. Application of Si increased the water potential of drought-stressed plants at filling stage, whereas it did not at booting stage. The superoxide dismutase (SOD) activity was inhibited and peroxidase (POD) activity was enhanced by drought at booting stage, and no differences were observed due to the Si treatment. At filling stage, however, application of Si increased the SOD activity and decreased the POD activity of drought-stressed plants. The catalase (CAT) activity was slightly increased by drought only in the absence of Si and at booting stage. The activity of glutathione reductase (GR) was not greatly influenced. Application of Si did not change the contents of H₂O₂, total soluble protein and protein carbonyl of drought-stressed plants at booting stage, whereas at filling stage, it decreased the content of H₂O₂ and protein carbonyl and increased the content of total soluble protein. The content of thiobarbituric acid reactive substances (TBARS) and the activities of acid phospholipase (AP) and lipoxygenase (LOX) in drought-stressed plants were also decreased by application of Si at both stages.     

Physiological and proteomic analysis of rice (<Emphasis Type="Italic">Oryza sativa</Emphasis> L.) in flag leaf during flowering stage and milk stage under drought stress

Hybrid rice has the inferiority of early senescence during reproductive stage. Early senescence of leaves induced by drought causes a constraint on crop productivity. Hence, to better understand traits of drought responsive mechanism in hybrid rice flag leaves during sensitive reproductive stage, we performed physiological measurements and conducted 2-D electrophoresis to investigate proteomic profile of rice flag leaves at flowering and milk stage. 43 proteins showed significant changes in silver stained gels were identified by MS/MS at flowering stage and 54 proteins at milk stage. For flowering stage, inactive CO₂ assimilation and ATP synthesis as well abnormal floral development could be postulated from decreased proteins. Additionally, higher levels of defense-related proteins and antioxidases in drought-stressed leaves suggested active ROS scavenging system. For milk stage, both photosynthesis and CO₂ assimilation were impaired. Disrupted oxidative defense and proteolysis system indicated redox imbalance. Upregulation of NADP-MDH facilitated for CO₂ assimilation and antioxidant system. In conclusion, drought stress during reproductive stage impacted a lot on biological processes in rice leaves at protein level, especially energy metabolism and redox balance.     

Effect of Selenium on Ascorbate�CGlutathione Metabolism During PEG-induced Water Deficit in Trifolium repens L.

To elucidate the effect of selenium (Se) on the ascorbate?Cglutathione (ASC?CGSH) cycle under drought stress, the activities of antioxidant enzymes and the levels of molecules involved in ASC?CGSH metabolism were studied in Trifolium repens seedlings subjected to polyethylene glycol (PEG)-induced water deficit alone or combined with 5???M Na₂SeO₄. Compared to the control, H₂O₂, thiobarbituric acid reactive substances (TBARS), ascorbate (ASC), dehydroascorbate (DHA), and glutathione disulfide (GSSG) contents increased, whereas a constant content of glutathione (GSH) and decreases in ASC/DHA and GSH/GSSG ratios were observed in the presence of PEG. The activities of ascorbate peroxidase (APX), dehydroascorbate reductase (DHAR), and glutathione reductase (GR) were upregulated, except for monodehydroascorbate reductase (MDHAR) activity during PEG-induced water deficit. Se application decreased the contents of H₂O₂, TBARS, DHA, and GSSG, increased the levels of GSH and ASC, and inhibited the decreases of ASC/DHA and GSH/GSSG ratios. Although it did not affect APX activity significantly, Se addition improved the activities of MDHAR, DHAR, and GR. Furthermore, GR activity showed the highest increase followed by that of DHAR and MDHAR in decreasing order. These data indicated that fluctuations in ASC?CGSH metabolism resulting from Se may have a positive effect on drought stress mitigation, and the regulation in the ASC?CGSH cycle can be attributed mainly to GR and DHAR in PEG?+?Se-treated T. repens seedlings.     

中国沙棘HrANR基因及黄酮类累积与抗旱的关系

花青素还原酶(anthocyanidin reductase, ANR)是合成黄酮类物质的关键酶之一,为明确其编码基因结构及干旱胁迫下的表达模式和黄酮类物质含量及二者之间的相关性,该文从中国沙棘转录组数据中筛选获得1个ANR基因,命名为HrANR基因。采用生物信息学软件对基因序列及编码蛋白进行分析,并对不同胁迫下各组织中HrANR基因的表达量和叶中黄酮类化合物含量进行相关性分析。结果表明:(1)中国沙棘HrANR基因ORF为1 017 bp,编码338个氨基酸,为稳定的亲水性蛋白,其ANR同源蛋白具有明显的科属特性。(2)干旱胁迫下HrANR基因在中国沙棘根、茎、叶中均有表达,但表达趋势不同,其中在根中的表达呈先升高后降低再升高的趋势,在茎中呈持续下降的趋势,在叶中呈先升高后持续降低的趋势。(3)通过芦丁标准曲线获得不同胁迫程度下中国沙棘叶内黄酮类的含量,表明黄酮类含量呈先持续上升,随后略有下降,复水后上升至最高点的变化趋势,表明干旱胁迫初期叶黄酮类含量与干旱胁迫呈正相关,在严重胁迫下黄酮类含量与胁迫呈负相关。(4)叶和茎的HrANR基因表达量与黄酮类含量呈负相关(P_叶=-0.751,P_茎=-0.934),根中呈正相关(P_根=0.444)。综上表明,中国沙棘HrANR基因的表达及黄酮类含量变化与其抗旱性密切相关,其结果为中国沙棘抗旱机制的阐明提供了依据。     

Cadmium toxicity is reduced by nitric oxide in rice leaves

We evaluate the protective effect of nitric oxide (NO) against Cadmium (Cd) toxicity in rice leaves. Cd toxicity of rice leaves was determined by the decrease of chlorophyll and protein contents. CdCl₂ treatment resulted in (1) increase in Cd content, (2) induction of Cd toxicity, (3) increase in H₂O₂ and malondialdehyde (MDA) contents, (4) decrease in reduced form glutathione (GSH) and ascorbic acid (ASC) contents, and (5) increase in the specific activities of antioxidant enzymes (superoxide dismutase, glutathione reductase, ascorbate peroxidase, catalase, and peroxidase). NO donors [N-tert-butyl-α-phenylnitrone, 3-morpholinosydonimine, sodium nitroprusside (SNP), and ASC + NaNO₂] were effective in reducing CdCl₂-induced toxicity and CdCl₂-increased MDA content. SNP prevented CdCl₂-induced increase in the contents of H₂O₂ and MDA, decrease in the contents of GSH and ASC, and increase in the specific activities of antioxidant enzymes. SNP also prevented CdCl₂-induced accumulation of NH₄ ⁺, decrease in the activity of glutamine synthetase (GS), and increase in the specific activity of phenylalanine ammonia-lyase (PAL). The protective effect of SNP on CdCl₂-induced toxicity, CdCl₂-increased H₂O₂, NH₄ ⁺, and MDA contents, CdCl₂-decreased GSH and ASC, CdCl₂-increased specific activities of antioxidant enzymes and PAL, and CdCl₂-decreased activity of GS were reversed by 2-(4-carboxy-2-phenyl)-4,4,5,5-tetramethyl-imidazoline-1-oxyl-3-oxide, a NO scavenger, suggesting that protective effect by SNP is attributable to NO released. Reduction of CdCl₂-induced toxicity by NO in rice leaves is most likely mediated through its ability to scavenge active oxygen species including H₂O₂.     

H_2S对干旱胁迫下白三叶幼苗抗氧化防御能力的影响

以‘拉丁诺’白三叶(Trifolium repens cv.‘Ladino’)为试验材料,研究外源H2S处理对PEG6 000(聚乙二醇)模拟干旱胁迫下白三叶叶片相对含水量(RWC)、膜脂过氧化、活性氧成分、抗氧化酶、抗坏血酸-谷胱甘肽循环代谢和非酶抗氧化物质的影响,以揭示H_2S调控白三叶抗旱性的生理机制。结果显示:(1)0.2 mmol/L的外源NaHS(H_2S供体)能显著提高干旱胁迫下白三叶的叶片相对含水量,维持显著较低的电解质渗透率(EL)和丙二醛(MDA)含量。(2)与直接干旱胁迫相比,干旱胁迫下外源添加NaHS处理的白三叶叶片内超氧化物歧化酶(SOD)、过氧化物酶(POD)、过氧化氢酶(CAT)活性显著增强,抗坏血酸-谷胱甘肽循环代谢中关键酶抗坏血酸过氧化物酶(APX)、脱氢抗坏血酸还原酶(DHAR)、单脱水抗坏血酸还原酶(MDHAR)和谷胱甘肽还原酶(GR)活性及其抗氧化中间产物抗坏血酸(AsA)、谷胱甘肽(GSH)含量也显著提高。(3)叶片类黄酮、总酚和原花青素的含量在一定的胁迫时间范围内亦显著增加,并伴随着活性氧成分O_2~(-·)产生速率和H_2O_2水平降低。研究认为,外源H2S能通过促进干旱胁迫下白三叶体内的多重抗氧化防御能力来提高其幼苗的抗旱性。     

Antioxidative responses to different altitudes in leaves of alpine plant Polygonum viviparum in summer

Mountain environmental stresses result in increased formation of hydrogen peroxide (H₂O₂) and accumulation of malondialdehyde (MDA) in leaves of Polygonum viviparum. The activities of several antioxidative system enzymes such as superoxide dismutase (SOD, EC 1.15.1.1), catalase (CAT, EC 1.11.1.6), peroxidase (POD, EC 1.11.1.7), glutathione reductase (GR, EC 1.6.4.2), dehydroascorbate reductase (DHAR, EC 1.8.5.1) and the contents of several non-enzymatic antioxidants such as reduced form of ascorbate (ASC), dehydroascorbate (DHA), reduced glutathione (GSH), and oxidized glutathione (GSSG) were investigated in leaves of P. viviparum, which were collected from three altitudes (2,200, 3,200, and 3,900 m) of Tianshan Mountain in China. The activities of these four antioxidative enzymes were accompanied by increases of H₂O₂ levels from 2,200 to 3,200 m. However, the activities of CAT and POD were decreased, whereas the activities of SOD and GR continually increased at 3,900 m. Analyses of isoforms of SOD, CAT, POD, and GR showed that the leaves of P. viviparum exposed different altitude conditions are capable of differentially altering the intensity. Additionally, two new isoforms of SOD were detected at 3900 m. A continual increase in the ASC, ASC to DHA ratio, GSH and GSH/[GSH + GSSG] ratio, and the activity of DHAR were observed in leaves of P. viviparum with the elevation of altitude. These results suggest that the higher contents of ASC, GSH as well as an increase in reduced redox state may be essential to antioxidation processes in the leaves of P. viviparum, whereas antioxidant enzymes system is a cofactor in the processes.     

Ascorbate-glutathione metabolism during PEG-induced water deficit in <Emphasis Type="Italic">Trifolium repens</Emphasis>

In order to elucidate the response of the ascorbate-glutathione (ASC-GSH) cycle to drought stress, the activities of antioxidant enzymes and the levels of molecules involved in the ASC-GSH metabolism were studied in Trifolium repens L. seedlings subjected to PEG-induced water deficit. Compared to the control, the contents of H₂O₂, thiobarbituric acid reactive substances (TBARS), ascorbate (ASC), dehydroascorbate (DHA), and glutathione disulfide (GSSG) increased in PEG-treated seedlings, whereas the glutathione (GSH) content kept constant during the drought period. Further more, the ASC/DHA and GSH/GSSG ratios decreased in the presence of PEG. Except for that of monodehydroascorbate reductase (MDHAR), the activities of ascorbate peroxidase (APX), dehydroascorbate reductase (DHAR), and glutathione reductase (GR) were up-regulated during water deficit, and the increases in APX and DHAR activities were much higher than those in GR activity. These data indicate that fluctuations in the ASC-GSH metabolism resulted from PEG treatment may have a positive effect on drought stress mitigation in T. repens.     

沙芥属植物活性氧清除系统对干旱胁迫的响应

以沙芥属植物沙芥和斧形沙芥幼苗为试材,采用盆栽控水干旱方法,分析其在不同干旱胁迫强度下根和叶的活性氧水平、抗氧化酶活性和抗氧化剂含量的变化,并利用隶属函数法和抗旱系数法综合评价沙芥和斧形沙芥的抗旱性。结果表明: （1）随着干旱胁迫程度的加剧,沙芥和斧形沙芥的根和叶中O^-·₂产生速率及·OH、H₂O₂、MDA含量总体呈逐渐升高的趋势,且干旱胁迫下沙芥比斧形沙芥产生了更多的ROS和MDA。（2）随着干旱胁迫程度的加剧,沙芥和斧形沙芥的根和叶中POD、APX、GST活性及叶中GR活性均先升高后降低,叶中的SOD活性以及根中GR、GPX活性均先降低后升高,根和叶中的CAT活性、叶中的GPX活性和根中SOD活性均逐渐升高;但根和叶中的SOD、POD、CAT活性在各干旱处理下均表现为斧形沙芥高于沙芥。（3）沙芥和斧形沙芥的根和叶中AsA含量随着干旱胁迫程度的加剧而先升高后降低,GSH含量逐渐升高,CAR含量逐渐降低,而VE含量在叶中逐渐升高,在根中却逐渐降低;但斧形沙芥比沙芥合成更多的AsA和GSH,其植物体内AsA GSH循环系统能清除更多的ROS。（4）沙芥和斧形沙芥的根和叶中总抗氧化能力（T AOC）均随着干旱加剧逐渐增强,且斧形沙芥的总抗氧化能力强于沙芥;活性氧清除系统的平均隶属度和综合抗旱系数显示,轻度干旱胁迫下沙芥抗旱性强于斧形沙芥,中度和重度干旱胁迫下斧形沙芥的抗旱性强于沙芥。研究认为,在干旱胁迫条件下,斧形沙芥根叶中ROS和MDA含量明显低于沙芥,而其大部分抗氧化酶活性和抗氧化剂含量高于沙芥,斧形沙芥植株体内抗氧化系统表现出更强的活性氧清除能力,从而表现出更强的抗旱性。     

Improvement of heat and drought photosynthetic tolerance in wheat by overaccumulation of glycinebetaine

Within their natural habitat, crops are often subjected to drought and heat stress, which suppress crop growth and decrease crop production. Causing overaccumulation of glycinebetaine (GB) has been used to enhance the crop yield under stress. Here, we investigated the response of wheat (Triticum aestivum L.) photosynthesis to drought, heat stress and their combination with a transgenic wheat line (T6) overaccumulating GB and its wild-type (WT) Shi4185. Drought stress (DS) was imposed by controlling irrigation until the relative water content (RWC) of the flag leaves decreased to between 78 and 82%. Heat stress (HS) was applied by exposing wheat plants to 40°C for 4 h. A combination of drought and heat stress was applied by subjecting the drought-stressed plants to a heat stress as above. The results indicated that all stresses decreased photosynthesis, but the combination of drought and heat stress exacerbated the negative effects on photosynthesis more than exposure to drought or heat stress alone. Drought stress decreased the transpiration rate (Tr), stomatal conductance (Gs) and intercellular CO₂ concentration (Ci), while heat stress increased all of these; the deprivation of water was greater under drought stress than heat stress, but heat stress decreased the antioxidant enzyme activity to a greater extent. Overaccumulated GB could alleviate the decrease of photosynthesis caused by all stresses tested. These suggest that GB induces an increase of osmotic adjustments for drought tolerance, while its improvement of the antioxidative defense system including antioxidative enzymes and antioxidants may be more important for heat tolerance.     

24-表油菜素内酯预处理对干旱胁迫下葡萄幼苗抗氧化系统和渗透调节物质的影响

以酿酒葡萄‘雷司令’(Riesling)一年生营养袋扦插苗为材料,采用人工气候室水培试验,考察在聚乙二醇6000(PEG)模拟干旱条件下,不同浓度(0.05、0.10和0.20mg/L)24-表油菜素内酯(EBR)预处理对‘雷司令’幼苗活性氧、抗氧化物质、渗透调节物质含量和抗氧化酶活性的影响,以揭示EBR预处理对干旱胁迫下葡萄幼苗的抗旱机理。结果显示:(1)与正常生长(对照)相比,干旱胁迫显著提高葡萄幼苗叶片中超氧阴离子自由基(■)、过氧化氢(H_2O_2)和丙二醛(MDA)含量;与干旱胁迫处理(PEG)相比,不同浓度EBR预处理均可降低叶片中■、H_2O_2和MDA的含量。(2)与对照相比,PEG处理显著降低葡萄幼苗叶片的抗坏血酸(AsA)和还原型谷胱甘肽(GSH)含量;与PEG处理相比,各浓度EBR预处理均可显著提高葡萄叶片AsA与GSH的含量,且以0.10mg/LEBR处理效果最好。(3)随着干旱胁迫时间的延长,葡萄幼苗叶片中的超氧化物歧化酶(SOD)、过氧化氢酶(CAT)、过氧化物酶(POD)与抗坏血酸过氧化物酶(APX)活性均呈先上升后下降的变化趋势,而在正常生长条件下酶活性基本保持不变;EBR预处理的葡萄叶片SOD、CAT、POD和APX活性均始终高于同期PEG处理。(4)PEG处理条件下,渗透调节物质脯氨酸和可溶性蛋白的含量整体高于对照;与PEG处理相比,不同浓度EBR预处理在干旱胁迫中后期均能显著提高葡萄叶片中脯氨酸和可溶性蛋白含量。研究表明,在干旱胁迫下,外源EBR预处理能够提高葡萄叶片抗氧化系统酶活性和渗透调节物质含量,有效降低干旱胁迫诱导的活性氧过度积累及膜脂过氧化程度,提高葡萄幼苗的抗旱能力,且以0.10mg/L EBR处理效果最佳。     

Responsiveness of Durum Wheat to Mycorrhizal Inoculation Under Different Environmental Scenarios

A greater understanding of how climate change will affect crop photosynthetic performance has been described as a target goal to improve yield potential. Other concomitant stressors can reduce the positive effect of elevated atmospheric CO₂ on wheat yield. Arbuscular mycorrhizal fungi (AMF) are symbiotic fungi predicted to be important in defining plant responses to rising atmospheric CO₂, but their role in response to global climatic change is still poorly understood. This study aimed to assess if increased atmospheric CO₂ interacting with drought can modify the effects of mycorrhizal symbiosis on flag leaf physiology in winter wheat. The study was performed in climate-controlled greenhouses with ambient (400 ppm, ACO₂) or elevated (700 ppm, ECO₂) CO₂ concentrations in the air. Within each greenhouse half of the plants were inoculated with Rhizophagus intraradices. When ear emergence began, half of the plants from each mycorrhizal and CO₂ treatment were subjected to terminal drought. At ACO₂ AMF improved the photochemistry efficiency of PSII compared with non-mycorrhizal plants, irrespective of irrigation regime. Mycorrhizal wheat accumulated more fructan than non-mycorrhizal plants under optimal irrigation. The level of proline in the flag leaf increased only in mycorrhizal wheat after applying drought. Mycorrhizal association avoided photosynthetic acclimation under ECO₂. However, nitrogen availability to flag leaves in mycorrhizal plants was lower under ECO₂ than at ACO₂. Results suggest that the mechanisms underlying the interactions between mycorrhizal association and atmospheric CO₂ concentration can be crucial for the benefits that this symbiosis can provide to wheat plants undergoing water deficit.     

Alleviation of drought stress and the physiological mechanisms in Citrus cultivar (Huangguogan) treated with methyl jasmonate

ABSTRACT

The role of exogenous methyl jasmonate (MeJA) in alleviating drought stress was investigated on Huangguogan. Except for intercellular CO₂ concentration, MeJA had little effect on net photosynthetic rate, stomatal conductance, and transpiration rate under drought stress. Compared with drought stress, MeJA significantly alleviated the decrease of chlorophyll content. However, chlorophyll a/b ratio was significantly increased. MeJA significantly increased proline and soluble sugar contents, significantly decreased the O₂ ^?· and H₂O₂ levels, and increased SOD and POD activities. In addition, the MDA content of drought stress was the highest of all treatments. MeJA significantly reduced MDA content in drought-stressed Huangguogan leaves. Although the Ascorbic acid (AsA) contents of 500 and 1000 mg L^?1 MeJA treatments were lower than that of 250 mg L^?1 MeJA, but all concentration of MeJA treatments delayed the decline of AsA content. Therefore, MeJA could induce drought stress tolerance by increasing the osmotic adjustment substances and antioxidant activities.     

玉米、花生及其间作茬口与施磷对冬小麦光合特性及产量的影响机制

该试验在玉米单作茬口、玉米-花生间作茬口(间作茬口)、花生单作茬口共3种茬口,以及0 kg P_2O_5·hm~(-2)(P_0)和180 kg P_2O_5·hm~(-2)(P_1) 2个磷水平下,研究了间作茬口与施磷对冬小麦分蘖、叶面积指数(LAI)、干物质积累、光合特性及产量的影响机制,为玉米花生间作与小麦-玉米复种轮作提供理论依据。结果表明:(1)间作茬口较玉米茬口显著提高了冬小麦有效分蘖数、LAI、净光合速率和干物质积累量,并提高了冬小麦旗叶的SPAD值、CO_2饱和点、光饱和点及最大净光合速率(P_(nmax))、表观量子效率(AQY)、羧化效率(CE)、最大羧化速率(V_(cmax))、最大RUBP再生的电子传递速率(J_(max))和最大磷酸丙糖利用速率(V_(TPU)),且CE、V_(cmax)、V_(TPU)的增幅均达到显著水平(P0.05),有效改善了冬小麦产量构成,显著提高籽粒产量(P0.05)。(2)间作茬口较花生茬口提高了冬小麦乳熟期的P_(nmax)、AQY、CE,增加了穗粒数和粒重,提高了产量。(3)与不施磷相比,施磷180 kg P_2O_5·hm~(-2)显著促进间作茬口冬小麦生长,显著提高冬小麦旗叶的SPAD值、P_(nmax)、AQY、CE、V_(cmax)、J_(max)、V_(TPU)和籽粒产量(P0.05)。研究发现,间作茬口较玉米茬口能有效增强冬小麦旗叶表观量子效率和CO_2羧化能力,显著提高小麦花后光合能力,促进冬小麦生长,从而增加穗粒数、粒重和籽粒产量,且间作茬口结合施磷180 kg P_2O_5·hm~(-2)效果更好。     

The responses of the enzymes related with ascorbate–glutathione cycle during drought stress in apple leaves

To explore the significance of the ascorbate–glutathione cycle under drought stress, the leaves of 2-year-old potted apple (Malus domestica Borkh.) plants were used to investigate the changes of each component of the ascorbate–glutathione cycle as well as the gene expression of dehydroascorbate reductase (DHAR, EC 1.8.5.1), ascorbate peroxidase (APX, EC 1.11.1.11) and glutathione reductase (GR, EC 1.6.4.2) under drought stress. The results showed that the malondialdehyde (MDA) and H₂O₂ concentrations in apple leaves increased during drought stress and began to decrease after re-watering. The contents of total ascorbate, reduced ascorbic acid (AsA), total glutathione and glutathione (GSH) were obviously upregulated in apple leaves when the soil water content was 40–45%. With further increase of the drought level, the contents of the antioxidants and especially redox state of AsA and GSH declined. However, levels of them increased again after re-watering. Moreover, drought stress induced significant increase of the activities of enzymes such as APX, scavenging H₂O₂, and also of monodehydroascorbate reductase (MDHAR, EC 1.6.5.4), DHAR and GR used to regenerate AsA and GSH, especially when the soil water content was above 40–45%. During severe drought stress, activities of the enzymes were decreased and after re-watering increased again. Gene expression of cytoplasmic DHAR, cytoplasmic APX and cytoplasmic GR showed similar changes as the enzyme activities, respectively. The results suggest that the ascorbate–glutathione cycle is up-regulated in response to drought stress, but cannot be regulated at severe drought stress conditions.     

Salicylic acid pretreatment induces drought tolerance and delays leaf rolling by inducing antioxidant systems in maize genotypes

Salicylic acid (SA) acts as an endogenous signal molecule responsible for inducing abiotic stress tolerance in plants. In this study, the role of SA in improving drought tolerance in two maize cultivars (Zea mays L.) differing in their tolerance to drought was evaluated. The plants were regularly watered per pot and grown until the grain filling stage (R2) under a rainout shelter. At stage R2, parts of the plants were treated with SA, after which drought stress was applied. Leaf samples were harvested on the 10th and 17th days of the drought. Some antioxidant enzyme activity, such as the superoxide dismutase (SOD), catalase (CAT), ascorbate peroxidase (APX), glutathione reductase (GR), monodehydroascorbate reductase (MDHAR), dehydroascorbate reductase (DHAR), hydrogen peroxide (H₂O₂) and malondialdehyde (MDA) content, was measured during the drought period. Exogenous SA prevented water loss and delayed leaf rolling in comparison with control leaves in both cultivars. As a consequence of drought stress, lipid peroxidation, measured in terms of malondialdehyde content, was prevented by SA. SA pretreatment induced all antioxidant enzyme activities, and to a greater extent than the control leaves, during drought. SA also caused a reduction in the ascorbate (ASC) and glutathione (GSH) content in two maize cultivars. The H₂O₂ level was higher in SA pretreated plants than the controls in both cultivars. Pretreatment with SA further enhanced the activities of antioxidant enzymes and the concentrations of non-enzymatic antioxidants in the tolerant cultivar compared with the sensitive cultivar. Results suggested that exogenous SA could help reduce the adverse effects of drought stress and might have a key role in providing tolerance to stress by decreasing water loss and inducing the antioxidant system in plants with leaf rolling, an alternative drought protection mechanism.