Seed biopriming with drought tolerant isolates of Trichoderma harzianum promote growth and drought tolerance in Triticum aestivum

Green house study was aimed to investigate the effect of seed biopriming with drought tolerant isolates of Trichoderma harzianum, viz. Th 56, 69, 75, 82 and 89 on growth of wheat under drought stress and to explore the mechanism underlying plant water stress resilience in response to Trichoderma inoculation. Measurements of relative water content, osmotic potential, osmotic adjustment, leaf gas exchange, chlorophyll fluorescence and membrane stability index were performed. In addition, analysis of the phenolics, proline, lipid peroxidation and measurements of phenylalanine ammonia‐lyase activity were carried out. Seed biopriming enhanced drought tolerance of wheat as drought induced changes like stomatal conductance, net photosynthesis and chlorophyll fluorescence were delayed. Drought stress from 4 to 13 days of withholding water induced an increase in the concentration of stress induced metabolites in leaves, while Trichoderma colonisation caused decrease in proline, malondialdehyde (MDA) and hydrogen peroxide (H₂O₂), and an increase in total phenolics. A common factor that negatively affects plants under drought stress conditions is accumulation of toxic reactive oxygen species (ROS), and we tested the hypothesis that seed biopriming reduced damages resulting from accumulation of ROS in stressed plants. The enhanced redox state of colonised plants could be explained by higher l ‐phenylalanine ammonia‐lyase (PAL) activity in leaves after 13 days of drought stress in Trichoderma treated plants. Similar activity was induced in untreated plants in response to drought stress but to a lower extent in comparison to treated plants. Our results support the hypothesis that seed biopriming in wheat with drought tolerant T. harzianum strains increased root vigour besides performing the process of osmoregulation. It ameliorates drought stress by inducing physiological protection in plants against oxidative damage, due to enhanced capacity to scavenge ROS and increased level of PAL, a mechanism that is expected to augment tolerance to abiotic stresses.     

干旱胁迫与施氮对麻疯树幼苗渗透调节物质积累的影响

采用盆栽控水的方法,研究干旱胁迫(80％ FC、60％ FC、40％ FC和20％ FC)及施氮(N0 0 g·pot-1、N1 1.2 g·pot1、Nm3.6 g·pot-1和Nh6.0 g·pot-1)对麻疯树幼苗叶、茎和根部主要渗透调节物质积累的影响.结果表明:干旱胁迫条件下,麻疯树幼苗茎和根部的游离脯氨酸、可溶性蛋白和茎部可溶性糖大量积累,叶片中脯氨酸含量也随干旱胁迫程度的增加大幅度上升;Na+、Ca2+和Mg2+在麻疯树幼苗叶、茎和根中大量积累,而K+仅在茎中大量积累,叶片和根部K+含量略微上升.施氮对植株渗透调节物质的影响与干旱胁迫强度和施氮水平有关.在80％ FC和60％ FC水分条件下,增加N肥施用量能明显促进麻疯树幼苗各组分渗透调节物质的积累;在40％ FC水分条件下,Nh处理对渗透调节物质积累的促进作用减弱;而在20％ FC条件下,N1处理植株的渗透调节能力较高,Nm和Nh处理对植株渗透调节的促进作用不明显甚至转为抑制.     

Drought tolerance through over-expression of the expansin gene TaEXPB23 in transgenic tobacco

Expansins are proteins that are the key regulators of wall extension during plant growth. To investigate the role of TaEXPB23, a wheat expansin gene, we analyzed TaEXPB23 mRNA expression levels in response to water stress in wheat and examined the drought resistance of transgenic tobaccos over-expressing TaEXPB23. We found that the expression of TaEXPB23 corresponded to wheat coleoptile growth and the response to water stress. The results also indicated that the transgenic tobacco lines lost water more slowly than the wild-type (WT) plants under drought stress; their cells could sustain a more integrated structure under water stress than that of WT. Other physiological and biochemical parameters under water stress, such as electrolyte leakage, malondialdehyde (MDA) level, photosynthetic rate, F_v/F_m and ΦPSII, also suggested that the transgenic tobaccos were more drought resistant than WT plants.     

Salicylic acid-induced physiological and biochemical changes in lemongrass varieties under water stress

Abstract

Salicylic acid (SA) treatment reduces the damaging action by water deficit on growth and accelerates a restoration of growth processes. The aim of the present work was to study the physiological and biochemical alteration induced by SA in lemongrass plants under stress conditions. Therefore, a pot culture experiment was conducted to test whether SA application at concentration of (10^?5 M) through foliar spray could protect lemongrass (Cymbopogon flexuosus Steud. Wats.) varieties (Neema and Krishna), subjected to drought stress on the basis of growth parameters and biochemical constituents, proline metabolism and quality attributes including citral content. The treatments were as follows: (i) 100% FC + 0 SA; (ii) 75% FC + 0 SA; (iii) 50% FC + 0 SA; (iv) 75% FC + 10^?5 M SA; and (v) 50% FC + 10^?5 M SA. The growth parameters were significantly reduced under the applied water stress levels; however, foliar application of salicylic acid (10^?5 M) improved the growth parameters in stress-affected plants. The plants under water stress exhibited a significant increase in activities of nitrate reductase and carbonic anhydrase, and electrolyte leakage, proline content, free amino acid and in PEP carboxylase activity. Content and yield of essential oil also significantly decreased in plants that faced water stress. Thus, it was concluded that variety Neema is the more tolerant variety as compared to Krishna on the basis of content and oil yield and well adapted to drought stress conditions.     

Antioxidant responses of wheat plants under stress

Currently, food security depends on the increased production of cereals such as wheat (Triticum aestivum L.), which is an important source of calories and protein for humans. However, cells of the crop have suffered from the accumulation of reactive oxygen species (ROS), which can cause severe oxidative damage to the plants, due to environmental stresses. ROS are toxic molecules found in various subcellular compartments. The equilibrium between the production and detoxification of ROS is sustained by enzymatic and nonenzymatic antioxidants. In the present review, we offer a brief summary of antioxidant defense and hydrogen peroxide (H₂O₂) signaling in wheat plants. Wheat plants increase antioxidant defense mechanisms under abiotic stresses, such as drought, cold, heat, salinity and UV-B radiation, to alleviate oxidative damage. Moreover, H₂O₂ signaling is an important factor contributing to stress tolerance in cereals.     

Drought acclimation reduces O2*- accumulation and lipid peroxidation in wheat seedlings

Abiotic stresses cause ROS accumulation, which is detrimental to plant growth. It is well known that acclimation of plants under mild or sub-lethal stress condition leads to development of resistance in plants to severe or lethal stress condition. The generation of ROS and subsequent oxidative damage during drought stress is well documented in the crop plants. However, the effect of drought acclimation treatment on ROS accumulation and lipid peroxidation has not been examined so far. In this study, the effect of water stress acclimation treatment on superoxide radical (O(2)(-z.rad;)) accumulation and membrane lipid peroxidation was studied in leaves and roots of wheat (Triticum aestivum) cv. C306. EPR quantification of superoxide radicals revealed that drought acclimation treatment led to 2-fold increase in superoxide radical accumulation in leaf and roots with no apparent membrane damage. However under subsequent severe water stress condition, the leaf and roots of non-acclimated plants accumulated significantly higher amount of superoxide radicals and showed higher membrane damage than that of acclimated plants. Thus, acclimation-induced restriction of superoxide radical accumulation is one of the cellular processes that confers enhanced water stress tolerance to the acclimated wheat seedlings.     

Response ofBromus inermis inoculated withGlomus fasciculatum to potassium fertilization and drought stress

Summary Bromus inermis Leyss. was grown in a 2×2×2 factorial design using different levels of mycorrhizal inoculation (inoculated and noninoculated), soil water stress (Ψ₁ or −0.8 MPa) and potassium (K) fertilization (0 or 150 ppm) as factors. Soil water stress and mycorrhizal inoculation significantly reduced plant top dry weight during the 18 week study. Chlamydospore production by the mycorrhizal symbiontGlomus fasciculatum (Thaxter sensu. Gerd.) Gerd. and Trappe was significantly reduced by soil water stress of −0.8 MPa. Potassium (K) fertilization did not significantly influence plant top growth or mycorrhizal colonization. However, foliar Ca and Mg were significantly lower in plants fertilized with K. Foliar Ca and Mg concentrations of P, K, N, Mn, Zn and Cu were significantly greater in drought stressed plants whereas Ca and Mg concentrations were significantly greater in well-watered plants.     

Exogenous nitric oxide donor and arginine provide protection against short-term drought stress in wheat seedlings

Nitric oxide (NO) is an important plant signaling molecule that has a vital role in abiotic stress tolerance. In the present study, we assessed drought-induced (15 and 30% PEG, polyethylene glycol) damage in wheat (Triticum aestivum L. cv. Prodip) seedlings and mitigation by the synergistic effect of exogenous Arg (0.5 mM l-Arginine) and an NO donor (0.5 mM sodium nitroprusside, SNP). Drought stress sharply decreased the leaf relative water content (RWC) but markedly increased the proline (Pro) content in wheat seedlings. Drought stress caused overproduction of reactive oxygen species (ROS) and methylglyoxal (MG) due to the inefficiency of antioxidant enzymes, the glyoxalase system, and the ascorbate-glutathione pool. However, supplementation with the NO donor and Arg enhanced the antioxidant defense system (both non-enzymatic and enzymatic components) in drought-stressed seedlings. Application of the NO donor and Arg also enhanced the glyoxalase system and reduced the MG content by increasing the activities of the glyoxalase system enzymes (Gly I and Gly II), which restored the leaf RWC and further increased the Pro content under drought stress conditions. Exogenous NO donor and Arg application enhanced the endogenous NO content, which positively regulated the antioxidant system and reduced ROS production. Thus, the present study reveals the crucial roles of Arg and NO in enhancing drought stress tolerance in wheat seedlings by upgrading their water status and reducing oxidative stress and MG toxicity.     

拟南芥AtCIPK23基因对烟草抗旱能力的影响

为了探索拟南芥AtCIPK23基因对烟草耐旱能力的影响,对3个转AtCIPK23基因阳性纯合株系KA13、KA14和KA44与野生型烟草K326（对照）进行了自然干旱处理,测定离体叶片的失水速率、叶绿素含量、相对电导率、脯氨酸和可溶性糖含量,并分析了转基因及野生型材料对活性氧的清除能力,对活性氧清除基因NtSOD、NtCAT和NtAPX及干旱胁迫相关基因NtDREB、NtLEA5和NtCDPK2的表达量进行检测。结果表明：（1）转基因烟草离体叶片的失水速率明显低于K326;自然干旱7 d后,野生型K326出现了明显的干旱胁迫症状;干旱7 d进行复水后,转基因株系的复水存活率明显高于K326。（2）转基因株系中的叶绿素、脯氨酸及可溶性糖含量比K326显著提高,电导率则明显降低。（3）野生型烟草K326中H₂O₂的积累量明显高于3个转基因株系,转基因株系中ROS清除机制的3个关键基因NtSOD、NtCAT和NtAPX被诱导上调表达。（4）抗旱相关基因NtDREB、NtLEA5和NtCDPK2仅在转基因烟草中受干旱诱导。研究认为,AtCIPK23基因可能具有提高植物抗旱能力的功能。     

Sub-Surface Drip Irrigation in Associated with H₂O₂ Improved the Productivity of Maize under Clay-Rich Soil of Adana,Turkey

Maize being sub-tropical crop is sensitive to water deficit during the early growth stages; particularly clay-rich soil, due to the compaction of the soil. It is well-documented that potential sub-surface drip irrigation (SDI) (Full irrigation; SDIFull (100% field capacity (FC)), Deficit irrigation; SDIDeficit (70% FC)) improves water use efficiency, which leads to increased crop productivity; since it has a constraint that SDI excludes soil air around the root-zone during irrigation events, which alter the root function and crop performance. Additionally, in clayrich soils, the root system of plants generally suffers the limitation of oxygen, particularly the temporal hypoxia, and occasionally from root anoxia; while SDI system accomplishes with the aerating stream of irrigation in the rhizosphere could provide oxygen root environment. The oxygen can be introduced into the irrigation stream of SDI through two ways: the venturi principle, or by using solutions of hydrogen peroxide through the air injection system. Therefore, the application of hydrogen peroxide (H₂O₂; HP) can mitigate the adverse effect of soil compactness and also lead to improving the growth, yield and yield attributes of maize in clay-rich soil. Considering the burning issue, a field study was conducted in consecutive two seasons of 2017 and 2018; where hybrid maize was cultivated as a second crop, to evaluate the effect of liquid-injection of H₂O₂ (HP) into the irrigation stream of SDI on the performance of maize in a clay-rich soil field of Adana, Turkey. When soil water content decreased in 50% of available water, irrigation was performed. The amount of water applied to reach the soil water content to the field capacity is SDIFull (100% FC) and 70% FC of this water is SDIDeficit (70% FC). In the irrigation program, hydrogen peroxide (HP) was applied at intervals of 7 days on average according to available water with and without HP: SDIFull (100% FC) + 0 ppm HP with full SDI irrigation; SDIFull (100% FC) + 250 ppm HP with deficit SDI irrigation; SDIDeficit (70% FC) + 0 ppm HP, SDIDeficit (70% FC) + 250 ppm HP and SDIDeficit (70% FC) + 500 ppm HP. Deficit irrigation (SDIDeficit (70% FC)) program was started from tasseling stage and continued up to the physiological maturity stage with sub-soil drip irrigation. H₂O₂ was applied 3 times during the growing season. Two years’ results revealed that the liquid-injection of H₂O₂ into the irrigation stream of SDI improved the growth and yield-related attributes and grain yield of maize. Based on the obtained results, during the extreme climatic condition in the year 2017, SDIFull (100% FC) + 250 ppm HP was more effective than SDIFull (100% FC) + 0 ppm HP on all traits for relative to full irrigation. While, during the favourable climatic condition in the 2018 season, SDIFull (100% FC) + 250 ppm HP was more effective than full irrigation with SDIFull (100% FC) + 0 ppm HP for the grain yield, grains, and SPAD value. Accordingly, the most effective treatment was SDIFull (100% FC) + 250 ppm HP, as it gave the highest growth and yield-related attributes and grain yield of maize followed by SDIDeficit (70% FC) + 250 ppm HP. Therefore, SDIFull with 250 ppm H₂O₂ using as liquid-injection may be recommended to mitigate the adverse effect of soil compactness particularly water-deficit stress in clay-rich soil for the sustainability of maize production.     

Effects of drought stress on phosphorus and potassium uptake dynamics in summer maize (Zea mays) throughout the growth cycle

The effects of drought stress on the phosphorus (P) and potassium (K) uptake dynamics of summer maize (Zea mays L.) throughout the growth cycle were studied. Field trials were conducted under a completely randomized design with three field water capacity (FC) regimes: 75?% FC was well watered and considered to be the control, 55?% FC represented moderate stress (MS), and 35?% FC represented severe stress (SS). The water regimes were applied from the third leaf stage until maturity. Drought stress induced sharp decreases in total K and P uptake of maize organs at different developmental stages and, in particular, detrimentally affected the nutrient uptake capability of roots. SS caused more deleterious effect than MS on both total K and P uptake by plant organs. The results suggested maize plants differ in their ability to maintain nutrient uptake under drought stress, and it is highly dependent on the intensity and duration of drought stress and the developmental stage. The decrease in total K and P uptake caused by both MS and SS was accompanied by reduction in biomass production in drought-stressed tissues. The biomass allocation patterns in response to drought stress fluctuated strong mostly because of competitive changes in the shoot and roots at different stages, thus the root:shoot ratio increased at some stages and decreased at other stages. SS induced a dramatic reduction in the harvest index (HI), whereas MS slightly decreased HI. Thus, water limitation caused lower K and P uptake and HI.     

Alterations in Growth and Yield of Camelina Induced by Different Planting Densities under Water Deficit Stress

Camelina (Camelina sativa L.) is famous for its oil quality and unique fatty acid pattern. Growth and yield of crops reduced under water deficit conditions. Environmental threat such as drought or water deficit condition is the emerging problem which creates the negative impact on the growth of plants. Based upon the current situation a pot study was performed in rain out-shelter to explore the effect of different plant densities (15, 10 and 5 plants per pot) on growth and seed yield of two camelina genotypes under normal (100% WHC) and water deficit (60% WHC) conditions by using completely randomized design with factorial arrangement having three replicates. Results indicated that individual effects of plant densities and water deficit stress levels considerably influenced the growth and seed yield of camelina but interaction effects did not indicate any significant variation. Maximum values of leaf area index (LAI) and crop growth rate (CGR) were recorded in P3 treatment (15 plants per pot). However, maximum values of leaf area duration (LAD), net assimilation rate (NAR), yield and yield components were observed in the treatment P₁ (5 plants per pot). Water deficit condition (60% WHC) significantly minimized the growth, seed yield (0.82 g/m² ) and yield components of camelina genotypes. Both camelina genotypes (611 and 618) did not differ significantly under water deficit conditions.     

Photosynthesis and antioxidative defense mechanisms in deciphering drought stress tolerance of crop plants

Crop plants are regularly exposed to an array of abiotic and biotic stresses, among them drought stress is a major environmental factor that shows adverse effects on plant growth and productivity. Because of this these factors are considered as hazardous for crop production. Drought stress elicits a plethora of responses in plants resulting in strict amendments in physiological, biochemical, and molecular processes. Photosynthesis is the most fundamental physiological process affected by drought due to a reduction in the CO₂ assimilation rate and disruption of primary photosynthetic reactions and pigments. Drought also expedites the generation of reactive oxygen species (ROS), triggering a cascade of antioxidative defense mechanisms, and affects many other metabolic processes as well as affecting gene expression. Details of the drought stress-induced changes, particularly in crop plants, are discussed in this review, with the major points: 1) leaf water potentials and water use efficiency in plants under drought stress; 2) increased production of ROS under drought leading to oxidative stress in plants and the role of ROS as signaling molecules; 3) molecular responses that lead to the enhanced expression of stress-inducible genes; 4) the decrease in photosynthesis leading to the decreased amount of assimilates, growth, and yield; 5) the antioxidant defense mechanisms comprising of enzymatic and non-enzymatic antioxidants and the other protective mechanisms; 6) progress made in identifying the drought stress tolerance mechanisms; 7) the production of transgenic crop plants with enhanced tolerance to drought stress.     

Combined effect of salicylic acid and potassium mitigates drought stress through the modulation of physio-biochemical attributes and key antioxidants in wheat

Improving physio-biochemical traits in wheat under drought stress conditions has received more research attention in recent years for better adaptability and higher yield. In this study, we explored the potential bio-physiological mechanisms underlying improved plant growth and water use efficiency in wheat following soil application of potassium (0 and 100 kg ha^?1) and seed primed salicylic acid (SA) (150 mg per L) and SA foliar application (100 mg per L) under drought stresses (100%, 60% and 30% FC). Two years' average data revealed that inducing drought stress resulted in a decrease in plant pigments content, growth traits, and plant water status however, the influence was substantially reduced with the combined application of K and SA under drought stress conditions. The SA foliar spray in combination with K had increased chlorophyll a (174% and 83%), chl b (130% and 192%), chl a + b (156% and 120), carotenoid (22% and 11%), proline contents (24% and 29%) leaf relative water content (24% and 29%) while reduced leaf WSD (17% and 20%), WRC (6% and 7%), and WUC (23% and 28%) under mild and severe drought stresses, respectively. The increase in grain yield by 41% and 37% with enhanced water use efficiency was obtained with combined foliar SA and K under mild and severe drought stress, respectively indicating its vital role in overcoming the deleterious effects of drought via regulation of osmotic and metabolic processes and stabilizes cell components. RDA analysis revealed that the studied traits were completely discriminated under severe stress than mild or no drought stress. A positive and significant association was found between plant pigments with seed yield whereas a negative and significant correlation existed between water leaf traits and plant pigments. It was concluded that both foliar SA and seed primed SA with K fertilization combat the adverse effects of drought and improved plant water status as well as growth and bio-physiological traits of wheat under drought stress conditions.     

Exogenous application of calcium chloride in wheat genotypes alleviates negative effect of drought stress by modulating antioxidant machinery and enhanced osmolyte accumulation

Plants use various mechanisms to cope with drought constraints at morphological, physiological, and biochemical level by means of different adaptive mechanisms. All organisms use a network of signal transduction pathways to control their metabolism and to adapt to the environment. Among these pathways, calcium (Ca²⁺) ions play an important role as a universal second messenger. Calcium has unique properties and universal ability to transmit diverse signals that trigger primary physiological actions in the cell in response to hormones, pathogens, and stress factors. Calcium plays a fundamental role in regulating the polar growth of cells and tissues and participates in plant adaptation to various stress factors. This study was conducted to examine the role of Ca²⁺ in ameliorating the adverse effect of drought stress responses in two contrasting wheat genotypes, HD 2733 (drought sensitive) and HD 2987 (drought tolerant), differing in their drought tolerance. The plants were treated with mannitol or Hoagland solution and then supplemented with CaCl₂ (10 mM). Measurements of seed germination, shoot growth, and chlorophyll content showed that calcium treatment increased all these factors in tolerant genotype (HD 2987) under induced stress condition. Drought stress reduced relative water content, osmolyte, and soluble sugar accumulation in both the genotypes, but CaCl₂ supplementation increased all the components under stress condition in HD 2987 as compared to HD 2733. The oxidative damage caused by induced stress was lower in HD 2987 compared to HD 2733 genotypes as assessed by their higher photosynthetic capacity and lower electrolyte leakage, malondialdehyde (MDA) content as well as H₂O₂ accumulation. Less accumulation of superoxide and H₂O₂ was also observed in HD 2987 genotype after CaCl₂ supplementation combined with mannitol treatment. In addition, the enhanced accumulation of calcium in the HD 2987 genotype is correlated with the higher activities of antioxidant enzymes than HD 2733 genotype under similar stress conditions. Our findings provide evidence of the protective role of exogenous calcium in conferring better tolerance against mannitol-induced drought stress in wheat genotypes, which could be useful as genetic stock to develop wheat tolerant varieties in breeding programs.     

Enhancing Drought Tolerance in Wheat through Improving MorphoPhysiological and Antioxidants Activities of Plants by the Supplementation of Foliar Silicon

The main objective of the research is to assess the role of foliar application of silicon (Si) for enhancing the survival ability of wheat under drought stress through improving its morphology, physicochemical and antioxidants activities. Treatments were five doses of Si at the rate of 2, 4, 6 and 8 mM and a control. After completion of seeds germination, pots were divided into four distinct groups at various field capacity (FC) levels, such as 100% FC (well-irrigated condition), 75% FC (slight water deficit), 50% FC (modest water deficit) and 25% FC (severe water deficit stress condition). Foliar application of Si at the rate of 2, 4, 6 and 8 mM and a control were given after 30 days of sowing at the tillering stage of wheat. Findings of the present investigation indicated that increasing the level of water deficit stress reduced the morphological parameters (such as root and shoot fresh and dry-biomass weight) and physico-biochemical events ((such as chlorophyll contents by estimating SPAD value), total free amino acid (TFAA), total soluble sugar (TSS), total soluble protein (TSP), total proline (TP), CAT (catalase), POD (peroxidase), SOD (superoxide dismutase) and APX (ascorbate peroxidase)) of wheat; while foliar application of Si at 6 mM at tillering stage enhanced the drought tolerance in wheat by increasing morphology and physiochemical characters under all levels of drought stress. Similarly, antioxidants activities in wheat also enhanced by the application of Si at 6 mM under normal as well as all drought stress levels. Therefore, it may be concluded that foliar application of Si at 6 mM at the tillering stage of wheat is an important indication for increasing the drought tolerance by improving the morphology, physico-biochemical and antioxidants activities in plants under deficit water (drought) conditions.     

iTRAQ‐based quantitative proteomic analysis reveals new metabolic pathways of wheat seedling growth under hydrogen peroxide stress

As an abundant ROS, hydrogen peroxide (H₂O₂) plays pivotal roles in plant growth and development. In this work, we conducted for the first time an iTRAQ‐based quantitative proteomic analysis of wheat seedling growth under different exogenous H₂O₂ treatments. The growth of seedlings and roots was significantly restrained by increased H₂O₂ concentration stress. Malondialdehyde, soluble sugar, and proline contents as well as peroxidase activity increased with increasing H₂O₂ levels. A total of 3 425 proteins were identified by iTRAQ, of which 157 showed differential expression and 44 were newly identified H₂O₂‐responsive proteins. H₂O₂‐responsive proteins were mainly involved in stress/defense/detoxification, signal transduction, and carbohydrate metabolism. It is clear that up‐regulated expression of signal transduction and stress/defence/detoxification‐related proteins under H₂O₂ stress, such as plasma membrane intrinsic protein 1, fasciclin‐like arabinogalactan protein, and superoxide dismutase, could contribute to H₂O₂ tolerance of wheat seedlings. Increased gluconeogenesis (phosphoenol‐pyruvate carboxykinase) and decreased pyruvate kinase proteins are potentially related to the higher H₂O₂ tolerance of wheat seedlings. A metabolic pathway of wheat seedling growth under H₂O₂ stress is presented.