Polyamine catabolism influences antioxidative defense mechanism in shoots and roots of five wheat genotypes under high temperature stress

Effect of high temperature stress on polyamine catabolism and antioxidant enzyme activity in relation to glutathione, ascorbate and proline accumulation was studied in five wheat (Triticum aestivum L.) genotypes (differently susceptible to temperature stress). High temperature significantly increased the activities of superoxide dismutase (SOD), catalase (CAT), guaiacol peroxidase (GPX), ascorbate peroxidase (APX), dehydroascorbate reductase (DHAR), glutathione reductase (GR) and glutathione S-transferase (GST) in shoots of all genotypes. Higher activities of GPX in C 306, C 273 and APX in PBW 550, PBW 343 and PBW 534 demonstrate their important role in scavenging H₂O₂. Conversely, high temperature stress led to a significant decline in SOD, CAT, APX and GPX activities of roots with a subsequent increase in diamine oxidase (DAO) and polyamine oxidase (PAO) activities especially in PBW 550 and PBW 343. The concentration of ascorbic acid declined with the imposition of heat stress, however, polyamines responded to high temperature stress by increasing spermidine and spermine levels and decreasing putrescine levels. After exposure to high temperature, proline accumulation was significantly decreased in roots and increased in shoots though maximum concentration was achieved in C 306 genotype. Apparently, the wheat seedlings respond to high temperature mediated increase in reactive oxygen species (ROS) production by altering antioxidative defense mechanism and polyamine catabolism though differentially in five wheat genotypes. Among five genotypes studied, C 306 and C 273 seem to be better protected against temperature stress. The results suggested that shoots were more resistant against the destructive effects of ROS as is indicated by low levels of thiobarbituric acid reactive substances under high temperature stress.     

Effect of salicylic and abscisic acid administered through detached tillers on antioxidant system in developing wheat grains under heat stress

The mechanism imparting thermotolerance by salicylic acid (SA) and abscisic acid (ABA) is still unresolved using either spraying technique or in vitro conditions. Alternative way of studying these effects under near in vivo conditions is through the use of liquid culturing technique. Effects of SA and ABA (100 μM) on antioxidative enzymes, antioxidants and lipid peroxidation were studied in detached tillers of three wheat (Triticum aestivum L.) cultivars PBW 343, C 306 (heat tolerant) and WH 542 (heat susceptible) cultured in a liquid medium. Ears were subjected to heat shock treatment (45°C for 2 h) and then maintained at 25°C for 5 days. Heat shock treatment resulted in increased peroxidase (POD) activity, while superoxide dismutase (SOD) and catalase (CAT) activities were reduced compared to control. The decrease in CAT activity was more significant in susceptible cultivar WH 542. Concomitantly, content of α-tocopherol and lipid peroxides increased in heat-treated wheat ears, whereas contents of total ascorbate level were reduced. Following treatment with SA and ABA, activities of all three antioxidative enzymes increased in correspondence with an increase in ascorbate and α-tocopherol content. Apparently, lipid peroxide content was reduced by SA in heat tolerant cultivars (PBW 343 and C 306) whereas in susceptible cultivar it was decreased by ABA. The up-regulation of the antioxidant system by SA and ABA possibly contributes to better tolerance against heat shock-induced oxidative damage in wheat grains.     

Nitric oxide alleviates oxidative damage induced by high temperature stress in wheat

Effect of sodium nitroprusside (SNP), a donor of nitric oxide (NO) was examined in two wheat (Triticum aestivum L.) cultivars, C 306 (heat tolerant) and PBW 550 (comparatively heat susceptible) to study the extent of oxidative injury and activities of antioxidant enzyme in relation to high temperature (HT) stress. HT stress resulted in a marked decrease in membrane thermostability (MTS) and 2, 3, 5-triphenyl tetrazolium chloride (TTC) cell viability whereas content of lipid peroxide increased in both the cultivars. The tolerant cultivar C 306 registered less damage to cellular membranes compared to PBW 550 under HT stress. Activities of antioxidant enzymes viz, superoxide dismutase, catalase, ascorbate peroxidase, guaicol peroxidase and glutathione reductase increased with HT in both the cultivars. Following treatment with SNP, activities of all antioxidant enzymes further increased in correspondence with an increase in MTS and TTC. Apparently, lipid peroxide content was reduced by SNP more in shoots of heat tolerant cultivar C 306 indicating better protection over roots under HT stress. The up-regulation of the antioxidant system by NO possibly contributed to better tolerance against HT induced oxidative damage in wheat.     

Effect of water deficit on carbohydrate status and enzymes of carbohydrate metabolism in seedlings of wheat cultivars

The effect of water deficit on carbohydrate status and enzymes of carbohydrate metabolism (alpha and beta amylases, sucrose phosphate synthase, sucrose synthase, acid and alkaline invertases) in wheat (Triticum aestivum L.) was investigated in the seedlings of drought-sensitive (PBW 343) and drought-tolerant (C 306) cultivars. The water deficit was induced by adding 6% mannitol (water potential -0.815 Mpa) in the growth medium. The water deficit reduced starch content in the shoots of tolerant seedlings as compared to the sensitive ones, but increased sucrose content in the shoots and roots of tolerant seedlings, indicating their protective role during stress conditions. It also decreased the alpha-amylase activity in the endosperm of seedlings of both the cultivars, but increased alpha and beta amylase activities in the shoots of tolerant ones. Sucrose phosphate synthase (SPS) activity showed a significant increase at 6 days of seedling growth (DSG) in the shoots of stressed seedlings of tolerant cultivar. However, SPS activity in the roots of stressed seedlings of sensitive cultivar was very low at 4 DSG and appeared significantly only at day 6. Sucrose synthase (SS) activity was lower in the shoots and roots of stressed seedlings of tolerant cultivar than sensitive ones at early stage of seedling growth. Higher acid invertase activity in the shoots of seedlings of tolerant cultivar appeared to be a unique characteristic of this cultivar for stress tolerance. Alkaline invertase activity, although affected under water deficit conditions, but was too low as compared to acid invertase activity to cause any significant affect on sucrose hydrolysis. In conclusion, higher sucrose content with high SPS and low acid invertase and SS activities in the roots under water deficit conditions could be responsible for drought tolerance of C 306.     

Identification and expression analysis of group 3 LEA family genes in sorghum [Sorghum bicolor (L.) Moench]

Sorghum with its remarkable adaptability to drought and high temperature provides a model system for grass genomics and resource for gene discovery especially for abiotic stress tolerance. Group 3 LEA genes from barley and rice have been shown to play crucial role in abiotic stress tolerance. Here, we present a genome-wide analysis of LEA3 genes in sorghum. We identified four genes encoding LEA3 proteins in the sorghum genome and further classified them into LEA3A and LEA3B subgroups based on the conservation of LEA3 specific motifs. Further, expression pattern of these genes were analyzed in seeds during development and vegetative tissues under abiotic stresses. SbLEA3A group genes showed expression at early stage of seed development and increased significantly at maturity, while SbLEA3B group genes expressed only in matured seeds. Expression of SbLEA3 genes in response to abiotic stresses such as soil moisture deficit (drought), osmotic, salt, and temperature stresses, and exogenous ABA treatments was also studied in the leaves of 2-weeks-old seedlings. ABA and drought induced the expression of all LEA3 genes, while cold and heat stress induced none of them. Promoter analysis revealed the presence of multiple ABRE core cis-elements and a few low temperature response (LTRE)/drought responsive (DRE) cis-elements. This study suggests non-redundant function of LEA3 genes in seed development and stress tolerance in sorghum.     

Time-Course Studies on Accumulation of Hydrophilic Antioxidants to Different Stress Regimes Followed by Recovery in Contrasting Cultivars of Wheat

Under unpredictable climatic scenarios, drought is one of the major environmental constraints limiting plant growth and productivity in arid and semi-arid regions. Rapid recovery from drought is of paramount importance for the persistence and survival of different crops growing worldwide. The boiling soluble proteins, BSPs (proteins remaining soluble upon boiling in aqueous solution) forms an instrumental part of the response to water deficit conditions and might be of key importance for the survival of plants under unfavourable environmental conditions. These BSPs are typified by two unique properties: high hydrophilicity and high thermal stability. The main objective of the study was to determine drought-induced changes in the markers of oxidative stress along with modulation in the activity of the boiling soluble antioxidants in response to different stress regimes followed by re-watering in Triticum aestivum L. In this study, we determined the indices of oxidative stress (membrane injury index (MII) and lipid peroxidation in terms of malondialdehyde (MDA) content) and activities of boiling soluble antioxidant enzymes in seeds of sensitive and tolerant cultivars of wheat at different duration of stress (3, 6 and 10 days) followed by recovery (post stress harvest). Water content recorded a decline in the sensitive (PBW 343 and PBW 621) as well as tolerant (PBW 527 and PBW 175) cultivars in stress duration and cultivar dependent manner and this was reversed following re-watering in all the cultivars. Oxidative stress indicators also increased in all the cultivars at different stress intensities but this was reversed following re-watering in the tolerant cvs. PBW 175 and PBW 527. At 3 and 6 days, boiling soluble monodehydroascorbate reductase (BsMDAR), boiling soluble protein disulphide isomerase (BsPDI) activity increased in both the tolerant cvs. PBW 175 and PBW 527 whereas boiling soluble guaiacol peroxidase (BsGPX) increased in the sensitive cv. PBW 343. However, as the stress intensity increased to 10 days, BsMDAR, boiling soluble glutathione-S-transferase (BsGST) and BsGPX increased only in the tolerant cvs. PBW 175 and PBW 527, thus accentuating their cardinal roles in stress tolerance under harsh drought conditions. Upon re-watering the stress plants after 10 days, BsMDAR increased only in the tolerant cv. PBW 175. On the other hand, boiling soluble protein disulphide isomerase (BsPDI) increased in both the tolerant cv. PBW 175 and susceptible cv. PBW 343, but with a greater enhancement in the cv. PBW 175. Based upon our results, biochemical significance of the boiling soluble antioxidants in the cultivars of wheat differing in drought resistance during different stress intensities and recovery is discussed.     

Antioxidant Responses of Lentil to Cold and Drought Stress

The effects of cold and drought stress on antioxidant responses and growth parameters in shoots and roots of lentil (Lens culinaris M cv Sultan 1) seedlings were investigated. Ten-day-old hydroponically grown seedlings were subjected to drought and cold (4°C) stress for 5 days. The length and fresh weight of shoots decreased significantly under both stress conditions, contrary to the increase in these growth parameters for roots under the same conditions. The oxidative damage as generation of malondialdehyde and hydrogen peroxide, was markedly higher in shoots under cold. Both stress conditions caused a significant increase in malondialdehyde levels in root tissues. The increase in proline levels was more pronounced under cold stress in shoots and roots. The tested stress conditions had no significant effect on chlorophyll contents. Superoxide dismutase activity was differentially altered in shoot and root tissues under drought and cold stress. The catalase activity was higher in roots under drought stress. On the other hand, ascorbate peroxidase activity increased in root tissues under cold stress. The results indicate that improved tolerance to cold and drought stress in root and shoot tissues of lentil might be correlated to the increased capacity of antioxidative defense system.     

Antioxidant system and photosynthetic characteristics responses to short-term PEG-induced drought stress in cucumber seedling leaves

The effect of short-term drought stress on the water content, antioxidant system and photosynthetic characteristics was investigated using cucumber (Cucumis sativus L.) seedlings. The results indicated that polyethylene glycol induced water stress reduced water content in shoots of cucumber seedling after treatment of 36 hours, and caused obvious reductions in net photosynthetic rate, stomatal conductance, intercellular CO₂ concentration and transpiration of leaves. In addition, water stress significantly reduced the photosynthetic pigment content and inhibited photochemical activity, including actual photochemical efficiency, maximal quantum yield of photosystem II photochemistry and coefficient for photochemical quenching. Meanwhile non-photochemical quenching increased. As responses to drought stress, significant increases in electrolyte leakage, malondialdehyde, superoxide anion and hydrogen peroxide levels were detected in leaves. The superoxide dismutases, catalase, glutathione reductase and dehydroascorbate reductase activities, protein, ascorbate and glutathione content, all decreased and peroxidases activity increased, while ascorbate peroxidase and monodehydroascorbate reductase activities exhibited different trend under different degree of water stress. Therefore, it can be concluded that water stress strongly disrupted the normal metabolism of leaves and restrained water absorption.     

Putrescine modulates antioxidant defense response in wheat under high temperature stress

Effects of putrescine (Put) on responses of wheat (Triticum aestivum) seedlings or detached tillers at mid-milky stage to high temperature (HT) stress were investigated. The heat tolerant cv. PBW 343 exhibited higher content of antioxidants and activities of antioxidative enzymes, while lower content of lipid peroxides as compared to the heat-sensitive cv. HD 2329. HT elevated peroxidase (POX) and superoxide dismutase (SOD) activities, while diamine oxidase (DAO) and polyamine oxidase (PAO) activities were reduced in roots, shoots and developing grains. Application of Put under HT further enhanced POX and SOD activities along with increased content of ascorbate and tocophereol in grains. Invariably POX and SOD revealed higher activities in roots while CAT, DAO and PAO activities were higher in shoots. The content of lipid peroxides was increased in roots and shoots of HT stressed seedlings but less in Put-treated cv. PBW 343.     

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.     

Stromal and thylakoid‐bound ascorbate peroxidases in NaCl‐treated wheat

Salinity is one of the most widespread causes of plant growth inhibition and reduction of agricultural productivity. Activated oxygen species seem to be implicated in the injury mechanism induced by salt. In our work, the effect of NaCl was studied in chloroplasts from seedlings of two cultivars of Triticum durum Desf., differently sensitive to drought and to heavy metals (cv. Ofanto more tolerant than cv. Adamello). The seedlings were grown for 9 days in Hoagland's no. 2 solution, to which increasing concentrations of NaCl (0, 50 and 100 m M ) were added. The results indicate an involvement of activated oxygen species in the mechanism of cellular toxicity of NaCl and the capability of the seedlings to induce their antioxidant defences. In chloroplasts of both cultivars, NaCl treatment induced a significant increase in ascorbate, even if cv. Ofanto showed the higher response. Ascorbate peroxidases appear to behave differently in the two cultivars in response to NaCl‐induced oxidative stress of chloroplasts. In seedlings of Adamello treated with 50 m M NaCl, a sudden increase in thylakoid‐bound ascorbate peroxidase (tAPX) activity and a decrease in that of stromal ascorbate peroxidase (sAPX) occurred. Differently, in Ofanto, a slight increase of both sAPX and tAPX was induced by NaCl. In both cultivars, three well‐resolved bands of sAPX were found. In chloroplasts of both cultivars, monodehydroascorbate reductase (MDHAR) and dehydroascorbate reductase (DHAR) behaved similarly, even if in Adamello a greater decrease of DHAR was shown. The experimental evidence suggested a more marked effect of NaCl stress on Adamello as compared with Ofanto.     

Responses of antioxidant defense system of <Emphasis Type="Italic">Helianthus annuus</Emphasis> to abscisic acid treatment under drought and waterlogging

The effect of abscisic acid (ABA) treatment on growth pigments and antioxidant defense system were investigated in seedlings of Helianthus annuus (cvs. Nantio F1 and Özdemirbey) subjected to drought and waterlogging stress. In addition, seedlings were sprayed with 10 M ABA three times every other day. Relative growth rate (RGR) was significantly reduced in both genotypes under drought stress, however, this growth inhibition was less in ABA-treated plants. Total chlorophyll content increased by drought stress in both genotypes. Ascorbate was not influenced by drought, while α-tocopherol increased in cv. Nantio F1. Ascorbate and α-tocopherol increased with drought stress in cv. Özdemirbey. ABA treatment decreased ascorbate and β-carotene contents while it increased α-tocopherol and xanthophylls contents under drought stress. The activity of superoxide dismutase (SOD) in both genotypes increased under drought stress-ABA combinations. Catalase (CAT) activity decreased under drought stress and drought-ABA combinations while it increased under waterlogging stress. Glutathione reductase (GR) activity decreased under drought stress but recovered with ABA treatment. The results suggested that ABA treatments have different effects on the components of antioxidant defense system in H. annuus genotypes and ABA may contribute drought-induced oxidative stress tolerance but not effects under waterlogging stress.     

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.     

Influence of silicon on antioxidant mechanisms and lipid peroxidation in chickpea (Cicer arietinum L.) cultivars under drought stress

Abstract

The effects of exogenous silicon (Si) on leaf relative water content (RWC), and the growth, Si concentrations, lipid peroxidation (MDA), lipoxygenase (LOX) activity, proline and H₂O₂ accumulation, non-enzymatic antioxidant activity (AA) and the activity of some antioxidant enzymes (superoxide dismutase, SOD; catalase, CAT; ascorbate peroxidase, APX) in shoots of ten chickpea cultivars grown under drought were investigated. Drought stress decreased the growth of all the cultivars while applied Si improved the growth at least five of the 10 chickpea cultivars. Silicon applied to the soil at 100 mg kg^?1 significantly increased Si concentrations of the cultivars and counteracted the deleterious effects of drought in 5 of the ten chickpea cultivars by increasing their RWC. In most cultivars tested H₂O₂, proline and MDA content and LOX activity were increased by drought whereas application of Si decreased their levels. APX activity was increased by drought but it was depressed by Si. In general, SOD and CAT activities of the cultivars were decreased by drought. Depending on cultivars, the CAT activity was decreased, and increased or unchanged in response to applied Si, while the SOD activity of the cultivars increased or unchanged by Si. The non-enzymatic antioxidant activity of the cultivars was also increased by Si. These observations implied an essential role for Si in minimizing drought stress-induced limitation of the growth and oxidative membrane damage in chickpea plants.     

Selenium Pretreatment Upregulates the Antioxidant Defense and Methylglyoxal Detoxification System and Confers Enhanced Tolerance to Drought Stress in Rapeseed Seedlings

In order to observe the possible regulatory role of selenium (Se) in relation to the changes in ascorbate (AsA) glutathione (GSH) levels and to the activities of antioxidant and glyoxalase pathway enzymes, rapeseed (Brassica napus) seedlings were grown in Petri dishes. A set of 10-day-old seedlings was pretreated with 25 μM Se (Sodium selenate) for 48 h. Two levels of drought stress (10% and 20% PEG) were imposed separately as well as on Se-pretreated seedlings, which were grown for another 48 h. Drought stress, at any level, caused a significant increase in GSH and glutathione disulfide (GSSG) content; however, the AsA content increased only under mild stress. The activity of ascorbate peroxidase (APX) was not affected by drought stress. The monodehydroascorbate reductase (MDHAR) and glutathione reductase (GR) activity increased only under mild stress (10% PEG). The activity of dehydroascorbate reductase (DHAR), glutathione S-transferase (GST), glutathione peroxidase (GPX), and glyoxalase I (Gly I) activity significantly increased under any level of drought stress, while catalase (CAT) and glyoxalase II (Gly II) activity decreased. A sharp increase in hydrogen peroxide (H₂O₂) and lipid peroxidation (MDA content) was induced by drought stress. On the other hand, Se-pretreated seedlings exposed to drought stress showed a rise in AsA and GSH content, maintained a high GSH/GSSG ratio, and evidenced increased activities of APX, DHAR, MDHAR, GR, GST, GPX, CAT, Gly I, and Gly II as compared with the drought-stressed plants without Se. These seedlings showed a concomitant decrease in GSSG content, H₂O₂, and the level of lipid peroxidation. The results indicate that the exogenous application of Se increased the tolerance of the plants to drought-induced oxidative damage by enhancing their antioxidant defense and methylglyoxal detoxification systems.     

Antioxidant response of wheat roots to drought acclimation

Wheat (Triticum aestivum L.) seedlings of a drought-resistant cv. C306 were subjected to severe water deficit directly or through stress cycles of increasing intensity with intermittent recovery periods. The antioxidant defense in terms of redox metabolites and enzymes in root cells and mitochondria was examined in relation to membrane damage. Acclimated seedlings exhibited higher relative water content and were able to limit the accumulation of H₂O₂ and membrane damage during subsequent severe water stress conditions. This was due to systematic up-regulation of superoxide dismutase, ascorbate peroxidase (APX), catalase, peroxidases, and ascorbate–glutathione cycle components at both the whole cell level as well as in mitochondria. In contrast, direct exposure of severe water stress to non-acclimated seedlings caused greater water loss, excessive accumulation of H₂O₂ followed by elevated lipid peroxidation due to the poor antioxidant enzyme response particularly of APX, monodehydroascorbate reductase, dehydroascorbate reductase, glutathione reductase, and ascorbate–glutathione redox balance. Mitochondrial antioxidant defense was found to be better than the cellular defense in non-acclimated roots. Termination of stress followed by rewatering leads to a rapid enhancement in all the antioxidant defense components in non-acclimated roots, which suggested that the excess levels of H₂O₂ during severe water stress conditions might have inhibited or down-regulated the antioxidant enzymes. Hence, drought acclimation conferred enhanced tolerance toward oxidative stress in the root tissue of wheat seedlings due to both reactive oxygen species restriction and well-coordinated induction of antioxidant defense.     

黄土高原冰草叶片抗坏血酸和谷胱甘肽合成及循环代谢对干旱胁迫的生理响应

通过盆栽实验, 对干旱胁迫下黄土高原地区冰草(Agropyron cristatum)叶片的抗坏血酸和谷胱甘肽合成及循环代谢相关酶及物质含量进行了研究。结果表明: 冰草可以通过增强叶片的抗坏血酸和谷胱甘肽合成及循环代谢酶: 抗坏血酸过氧化物酶、谷胱甘肽还原酶、脱氢抗坏血酸还原酶、单脱氢抗坏血酸还原酶、L-半乳糖酸-1, 4-内酯脱氢酶和γ-谷氨酰半胱氨酸合成酶活性, 维持植物体内抗坏血酸和谷胱甘肽水平及氧化还原状态, 从而抵御干旱造成的氧化胁迫。但叶片抗坏血酸和谷胱甘肽合成及循环代谢对不同水平干旱胁迫的响应, 随胁迫时间的延长而不同。在胁迫24天以前, 严重干旱下叶片的抗坏血酸和谷胱甘肽合成及循环代谢增强较显著; 在胁迫24天后, 由于该胁迫下植物所遭受的氧化胁迫较为严重, 叶片中上述6种酶的活性均呈降低趋势。而在中度干旱下叶片抗坏血酸和谷胱甘肽合成及循环代谢相关的6种酶在整个胁迫过程中均保持较高的活性。这说明, 冰草能够长时间有效地抵御中度干旱所造成的氧化胁迫, 但只能在一定时间范围内有效地抵御严重干旱所造成的氧化胁迫, 胁迫时间延长则会降低其抵御严重干旱的能力。