耐盐砧木嫁接调控H_2O_2清除系统提高黄瓜幼苗耐盐性的机制
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摘要
近年来,随着我国设施园艺的迅速发展,土壤次生盐渍化问题日趋加剧,已成为限制我国设施蔬菜产业发展的主要障碍。黄瓜(Cucumis sativus L.)是设施栽培中的主要蔬菜作物,对盐胁迫非常敏感。前人研究表明耐盐砧木嫁接能够提高黄瓜耐盐性,但其研究主要围绕嫁接对Na+和C1-吸收运转的调控机理,而关于嫁接对盐胁迫诱导的氧化伤害调控研究相对较少。本研究从组织水平、叶绿体水平及转录水平上研究嫁接黄瓜H202清除系统对盐胁迫的响应,阐明了盐胁迫下耐盐砧木嫁接对H202清除系统的调控机制。利用基因型不同的黄瓜接穗与南瓜砧木嫁接,以及将黄瓜与南瓜进行正反嫁接试验,揭示盐胁迫下耐盐砧木对提高黄瓜耐盐性起决定作用。通过研究嫁接植株根系中ABA含量及其信号途径中相关分子(H2O2、NO和MAPK级联)的变化,探索了耐盐砧木根系中ABA在调控黄瓜耐盐性中可能的信号途径。所取得主要结果如下：
1.以盐敏感型黄瓜品种津春2号为材料,将预喷施H202抑制剂(DPI,15μ.M)和清除剂(DMTU,5mM)的植株进行0和75 mM NaCl胁迫。研究了下黄瓜幼苗中H202含量变化及其与盐抑制黄瓜生长的关系。结果显示,预喷施H202抑制剂和清除剂降低了NaCl胁迫下黄瓜幼苗中H202和MDA含量,增加了其根系和地上部干重。相关性分析表明H202含量与根系和地上部干重呈显著负相关。此结果表明,盐胁迫诱导H202积累是盐胁迫抑制黄瓜生长的重要原因。
2.以津春2号黄瓜和耐盐砧木超级拳王(Cucurbita moschata Duch.)分别作砧木,以津春2号黄瓜作接穗进行嫁接。研究了黄瓜自根苗、黄瓜自嫁苗和耐盐砧木嫁接苗在0和75 mM NaCl胁迫下的生长与抗氧化系统的变化。结果表明,NaCl胁迫下耐盐砧木嫁接黄瓜植株具有较高的CAT、APX、GR和MDAR酶活性、GSH/GSSG比值和铁还原／总抗氧化能力,以此减少了黄瓜组织中H202和MDA的积累,缓解了盐胁迫对黄瓜幼苗生长的抑制。
3.研究了0、50、100 mM NaCl胁迫下黄瓜自嫁苗和耐盐砧木嫁接苗叶绿体中AsA-GSH循环及其超微结构的变化。结果显示,50、100 mM NaCl胁迫下砧木嫁接苗叶绿体中AsA-GSH循环相关酶APX、GR和DHAR活性与AsA/DHA比值,以及100 mM NaCl胁迫下MDAR活性、GSH含量及GSH/GSSG比值均显著高于自嫁苗,导致砧木嫁接苗叶绿体中H202含量较低。随盐胁迫浓度增加,自嫁苗叶绿体结构遭受破坏,尤其是在100 mM NaCl处理下,其叶绿体基粒与基质片层叠垛变薄、变模糊甚至片层断裂,而砧木嫁接苗叶绿体结构受NaCl伤害较轻。砧木嫁接使盐胁迫黄瓜叶绿体中具有较强的H202清除能力,以此保护了叶绿体超微结构和改善了黄瓜幼苗的生长与光合特性。
4.研究了0和75mM NaCl胁迫下黄瓜自嫁苗和耐盐砧木嫁接苗叶片中H2O2含量与重要抗氧化酶(CAT.APX.GR和MDAR)活性的动态变化,以及NaCl处理120 h时嫁接黄瓜中抗氧化基因mRNA的表达。结果显示,NaCl胁迫24 h至120h,自嫁苗叶片中H2O2含量逐渐增加,72 h时达到峰值,而砧木嫁接苗中H2O2含量增加较缓慢,且始终低于自嫁苗,这与其在NaCl处理48 h至120 h始终具有较高的保护酶(CAT.APX.GR和MDAR)活性有关。NaCl胁迫下自嫁苗叶片中CAT基因和根系中cAPX基因,以及嫁接苗叶片中cAPX基因mRNA表达量的变化均与其相应酶活性的变化趋势相一致,说明这些酶的活性受到了其转录水平上的调控。
5.分别以盐敏感型和耐盐型黄瓜品种津春2号与津育1号为接穗,以盐敏感型和耐盐型南瓜品种黑籽南瓜(Cucurbita ficifolia B.)和超级拳王为砧木。研究了不同接穗和砧木基因型对嫁接黄瓜生长和耐盐性的影响。结果表明：100 mM NaCl胁迫下,以超级拳王为砧木的嫁接苗盐害指数较低,植株生长受抑制较轻。相关性分析表明这与其较低的叶片Na+含量和根系H2O2含量,及较高的根系SOD.POD和CAT活性有关。此结果表明,与接穗相比,耐盐砧木对嫁接黄瓜的耐盐性起决定作用。
6.将盐敏感型黄瓜津春2号和耐盐型南瓜超级拳王进行正反嫁接获得4种嫁接组合(接穗/砧木),包括黄瓜/黄瓜、南瓜/南瓜、黄瓜/南瓜、南瓜/黄瓜。研究了0和75 mM NaCl胁迫对嫁接植株根系中H2O2和MDA含量、重要抗氧化酶活性、ABA含量及其信号途径中相关分子(H202.NO和MAPK级联)的影响。结果显示,NaCl胁迫下以南瓜为砧木嫁接植株根系中ABA含量增加幅度较大,信号调节基因MAPK1和MAPK6表达显著上调。此结果表明NaCl胁迫诱导了耐盐砧木根系中ABA积累及其介导的MAPK级联信号反应。这使得耐盐砧木嫁接黄瓜根系中具有较高的H202清除酶(CAT.APX.GR和MDAR)活性用以控制根系中H2O2与MDA含量在较低水平。
总之,本研究结果表明,盐胁迫诱导H2O2产生是盐胁迫抑制黄瓜幼苗生长的重要原因。耐盐砧木嫁接能够调控黄瓜幼苗组织和叶绿体中H2O2清除能力,从而提高其耐盐性。与接穗相比,盐胁迫下耐盐砧木对提高黄瓜耐盐性起决定作用。这可能是盐胁迫下耐盐砧木根系中ABA积累及其介导的MAPK级联信号调控的结果。
In recent years, with the rapid development of protected horticulture in China, soil secondary salinization has been more and more severe. It has become the main obstacle of limiting the development of vegetable production under protected horticulture. Cucumber (Cucumis sativus L.) is the main vegetables under the protected cultivation conditions, and is sensitive to salt stress. It has been reported that grafting with salt-tolerant rootstock can improve cucumber tolerance to salt stress, but these researches are mainly about the mechanism of grafting on regulating the Na+and Cl- uptake and transport, little study has been down on the research of grafting on regulating the salt-induced oxidative stress. In our present study, the mechanism of grafting with salt-tolerant rootstock on regulating the H2O2-scavenging system under salt stress were evaluated, by investigating the response of H2O2-scavenging system to salt stress at grafted cucumber organic, chloroplast, and transcript levels. The salt-tolerant rootstock was clarified to play a primary important effect in determining the salt tolerance of cucumber seedlings under salt stress, according to the experiments of grafting with different genotype cultivars of cucumber scions and pumpkin rootstocks, and reciprocal grafting between cucumber and pumpkin. We explored the possible signal pathway of ABA in salt-tolerant rootstock roots on regulating the salt tolerance of cucumber, by investigating the ABA content and some signal molecules involved in ABA signal pathway (H2O2, NO and MAPK cascades) in grafted plant roots. The main contents and results in the present study are as follows:
1. A salt-sensitive cultivar of cucumber (cv.'Jinchun No.2') was used. Plants pre-sprayed with H2O2 inhibitor (DPI,15μM) and scavenger (DMTU.5 mM) were exposed to 0 and 75 mM NaC1 stress. The changes in H2O2 content in cucumber seedlings and what this has to do with the salt-induced growth inhibition were investigated. The results showed that pre-spraying H2O2 inhibitor and scavenger decreased the H2O2 and MDA contents, while increased the root and shoot dry weights of cucumber seedling under NaCl stress. Correlation analysis showed that H2O2 content was negatively correlated with root and shoot dry weights. This result implied that the salt-induced H2O2 accumulation was an important reason for plant growth inhibition caused by salt stress.
2. Cucumber plants (cv.'Jinchun No.2') were self-grafted or grafted onto salt-tolerant rootstock'Chaojiquanwang'(Cucurbita moschata Duch). The changes in plant growth and antioxidant defense system of non-grafted, self-grafted, and rootstock-grafted cucumber seedlings under 0 and 75 mM NaCl stress were investigated. The results showed that cucumber plants grafted onto salt-tolerant rootstock had higher activities of CAT, APX, GR, and MDAR, GSH/GSSG ratio, and the ferric reducing ability/antioxidant power, thereby decreasing the H2O2 and MDA contents in cucumber tissue. As a result, the salt-induced growth inhibition was alleviated.
3. Chloroplast ultrastructure and the AsA-GSH cycle system in chloroplasts of self-grafted and rootstock-grafted cucumber leaves under 0,50, and 100 mM NaC1 stress were investigated. The results showed that the activities of APX, GR, and DHAR involved in AsA-GSH cycle and AsA/DHA ratio in the chloroplasts of rootstock-grafted plants under 50 and 100 mM NaCl, as well as the MDAR activity, GSH content, and GSH/GSSG ratio under 100 mM NaCl were all significant higher than those in self-grafted plants. This resulted in the lower H2O2 content in the chloroplast of rootstock-grafted plants. With increasing levels of NaC1 concentration, the chloroplast structure of self-grafted plants was visibly damaged. Particularly under 100 mM NaCl, the stacks of grana and intergranal lamellae were thin and obscure, even fractured, whereas the damage in the chloroplast of rootstock-grafted plants were less severe. The results suggest that rootstock grafting enhances the H2O2-scavenging capacity of the AsA-GSH cycle in the chloroplasts under salt stress, thereby protecting the chloroplast structure and improving the photosynthetic performance and growth of cucumber seedlings.
4. Changes in the time course of H2O2 content and main antioxidant enzyme (CAT, APX, GR and MDAR) activities in the leaves of self-grafted and rootstock-grafted cucumber plants under 0 and 75 mM NaCl stress, as well as the mRNA levels of antioxidant enzyme genes in grafted cucumber plants under NaCl stress for 120 h were studied. The results showed that the H2O2 content in self-grafted leaves increased gradually during NaC1 stress from 24 h to 120 h, and showing a peak at 72 h, whereas the H2O2 content in rootstock-grafted increased more slowly and kept lower all the time. This could be associated with its higher antioxidant enzyme activities during NaCl stress from 48 h to 120 h. The changes in the mRNA level of CAT and cAPX in self-grafted plants leaves and roots, respectively, and the mRNA level of cAPX in rootstock-grafted plants leaves were consistent with the changes in their enzyme activity levels under NaCl stress. This result indicated that the activities of these enzymes were regulated at their mRNA levels.
5. Two cucumber cultivars (cv.'Jinchun No.2', a relative salt-sensitive cultivar, and 'Jinyu No.1', a relative salt-tolerant cultivar) were grafted onto two rootstocks ('Figleaf Gourd', a relative salt-sensitive cultivar, and'Chaojiquanwang', a relative salt-tolerant cultivar), respectively. The effects of scion and rootstock on plant growth and the antioxidant defense systems of cucumber were assessed. The results showed that the salt injury index of plants grafted onto Chaojiquanwang was lower and the growth reduction was less severe under 100 mM NaCl stress. Correlation analysis showed that this could be partially attributed to its lower leaf Na+content and root H2O2 content, as well as the higher activities of SOD, POD, and CAT in the roots. This result implied that the salt-tolerant rootstock played a primary important effect in determining the salt tolerance of grafted cucumber seedlings under salt stress compared with the scion genotypes.
6. A salt-sensitive cucumber cultivar (cv.'Jinchun No.2') and a salt-tolerant pumpkin cultivar ('Chaojiquanwang') were reciprocal grafted. Four types of grafted plants (scion/rootstock) were obtained, including cucumber/cucumber, pumpkin/pumpkin, cucumber/pumpkin, and pumpkin/cucumber. The effects of 0 and 75 mM NaC1 stress on H2O2 and MDA contents, antioxidant enzyme activities, ABA content and some signal molecules involved in ABA signal pathway (H2O2, NO and MAPK cascades) in grafted plant roots were investigated. The results showed that the ABA contents increased and the expression of signal regulatory genes such as MAPK1 and MAPK6 up-regulated significantly in the grafted plants roots under NaC1 stress when pumpkin was used as rootstock. The results implied that salinity induced the production of ABA and the response of MAPK cascades mediated by ABA in salt-tolerant rootstock roots. This resulted in the higher H2O2-scavenging enzyme activities in salt-tolerant rootstock roots for controlling the H2O2 and MDA contents at lower levels.
In conclusion, the present study suggests that the salt-induced H2O2 production result in the plant growth inhibition caused by salt stress. Grafting with salt-tolerant rootstock can improve the salt tolerance of cucumber seedlings by regulating the H2O2-scavenging system in cucumber organic and chloroplast. The salt-tolerant rootstock plays a primary important effect in determining the salt tolerance of cucumber seedlings under salt stress compared with the scion genotypes. This could be attributed to the production of ABA and the regulation of ABA-mediated MAPK cascades in salt-tolerant rootstock roots under salt stress.
1.以盐敏感型黄瓜品种津春2号为材料,将预喷施H202抑制剂(DPI,15μ.M)和清除剂(DMTU,5mM)的植株进行0和75 mM NaCl胁迫。研究了下黄瓜幼苗中H202含量变化及其与盐抑制黄瓜生长的关系。结果显示,预喷施H202抑制剂和清除剂降低了NaCl胁迫下黄瓜幼苗中H202和MDA含量,增加了其根系和地上部干重。相关性分析表明H202含量与根系和地上部干重呈显著负相关。此结果表明,盐胁迫诱导H202积累是盐胁迫抑制黄瓜生长的重要原因。
2.以津春2号黄瓜和耐盐砧木超级拳王(Cucurbita moschata Duch.)分别作砧木,以津春2号黄瓜作接穗进行嫁接。研究了黄瓜自根苗、黄瓜自嫁苗和耐盐砧木嫁接苗在0和75 mM NaCl胁迫下的生长与抗氧化系统的变化。结果表明,NaCl胁迫下耐盐砧木嫁接黄瓜植株具有较高的CAT、APX、GR和MDAR酶活性、GSH/GSSG比值和铁还原／总抗氧化能力,以此减少了黄瓜组织中H202和MDA的积累,缓解了盐胁迫对黄瓜幼苗生长的抑制。
3.研究了0、50、100 mM NaCl胁迫下黄瓜自嫁苗和耐盐砧木嫁接苗叶绿体中AsA-GSH循环及其超微结构的变化。结果显示,50、100 mM NaCl胁迫下砧木嫁接苗叶绿体中AsA-GSH循环相关酶APX、GR和DHAR活性与AsA/DHA比值,以及100 mM NaCl胁迫下MDAR活性、GSH含量及GSH/GSSG比值均显著高于自嫁苗,导致砧木嫁接苗叶绿体中H202含量较低。随盐胁迫浓度增加,自嫁苗叶绿体结构遭受破坏,尤其是在100 mM NaCl处理下,其叶绿体基粒与基质片层叠垛变薄、变模糊甚至片层断裂,而砧木嫁接苗叶绿体结构受NaCl伤害较轻。砧木嫁接使盐胁迫黄瓜叶绿体中具有较强的H202清除能力,以此保护了叶绿体超微结构和改善了黄瓜幼苗的生长与光合特性。
4.研究了0和75mM NaCl胁迫下黄瓜自嫁苗和耐盐砧木嫁接苗叶片中H2O2含量与重要抗氧化酶(CAT.APX.GR和MDAR)活性的动态变化,以及NaCl处理120 h时嫁接黄瓜中抗氧化基因mRNA的表达。结果显示,NaCl胁迫24 h至120h,自嫁苗叶片中H2O2含量逐渐增加,72 h时达到峰值,而砧木嫁接苗中H2O2含量增加较缓慢,且始终低于自嫁苗,这与其在NaCl处理48 h至120 h始终具有较高的保护酶(CAT.APX.GR和MDAR)活性有关。NaCl胁迫下自嫁苗叶片中CAT基因和根系中cAPX基因,以及嫁接苗叶片中cAPX基因mRNA表达量的变化均与其相应酶活性的变化趋势相一致,说明这些酶的活性受到了其转录水平上的调控。
5.分别以盐敏感型和耐盐型黄瓜品种津春2号与津育1号为接穗,以盐敏感型和耐盐型南瓜品种黑籽南瓜(Cucurbita ficifolia B.)和超级拳王为砧木。研究了不同接穗和砧木基因型对嫁接黄瓜生长和耐盐性的影响。结果表明：100 mM NaCl胁迫下,以超级拳王为砧木的嫁接苗盐害指数较低,植株生长受抑制较轻。相关性分析表明这与其较低的叶片Na+含量和根系H2O2含量,及较高的根系SOD.POD和CAT活性有关。此结果表明,与接穗相比,耐盐砧木对嫁接黄瓜的耐盐性起决定作用。
6.将盐敏感型黄瓜津春2号和耐盐型南瓜超级拳王进行正反嫁接获得4种嫁接组合(接穗/砧木),包括黄瓜/黄瓜、南瓜/南瓜、黄瓜/南瓜、南瓜/黄瓜。研究了0和75 mM NaCl胁迫对嫁接植株根系中H2O2和MDA含量、重要抗氧化酶活性、ABA含量及其信号途径中相关分子(H202.NO和MAPK级联)的影响。结果显示,NaCl胁迫下以南瓜为砧木嫁接植株根系中ABA含量增加幅度较大,信号调节基因MAPK1和MAPK6表达显著上调。此结果表明NaCl胁迫诱导了耐盐砧木根系中ABA积累及其介导的MAPK级联信号反应。这使得耐盐砧木嫁接黄瓜根系中具有较高的H202清除酶(CAT.APX.GR和MDAR)活性用以控制根系中H2O2与MDA含量在较低水平。
总之,本研究结果表明,盐胁迫诱导H2O2产生是盐胁迫抑制黄瓜幼苗生长的重要原因。耐盐砧木嫁接能够调控黄瓜幼苗组织和叶绿体中H2O2清除能力,从而提高其耐盐性。与接穗相比,盐胁迫下耐盐砧木对提高黄瓜耐盐性起决定作用。这可能是盐胁迫下耐盐砧木根系中ABA积累及其介导的MAPK级联信号调控的结果。
In recent years, with the rapid development of protected horticulture in China, soil secondary salinization has been more and more severe. It has become the main obstacle of limiting the development of vegetable production under protected horticulture. Cucumber (Cucumis sativus L.) is the main vegetables under the protected cultivation conditions, and is sensitive to salt stress. It has been reported that grafting with salt-tolerant rootstock can improve cucumber tolerance to salt stress, but these researches are mainly about the mechanism of grafting on regulating the Na+and Cl- uptake and transport, little study has been down on the research of grafting on regulating the salt-induced oxidative stress. In our present study, the mechanism of grafting with salt-tolerant rootstock on regulating the H2O2-scavenging system under salt stress were evaluated, by investigating the response of H2O2-scavenging system to salt stress at grafted cucumber organic, chloroplast, and transcript levels. The salt-tolerant rootstock was clarified to play a primary important effect in determining the salt tolerance of cucumber seedlings under salt stress, according to the experiments of grafting with different genotype cultivars of cucumber scions and pumpkin rootstocks, and reciprocal grafting between cucumber and pumpkin. We explored the possible signal pathway of ABA in salt-tolerant rootstock roots on regulating the salt tolerance of cucumber, by investigating the ABA content and some signal molecules involved in ABA signal pathway (H2O2, NO and MAPK cascades) in grafted plant roots. The main contents and results in the present study are as follows:
1. A salt-sensitive cultivar of cucumber (cv.'Jinchun No.2') was used. Plants pre-sprayed with H2O2 inhibitor (DPI,15μM) and scavenger (DMTU.5 mM) were exposed to 0 and 75 mM NaC1 stress. The changes in H2O2 content in cucumber seedlings and what this has to do with the salt-induced growth inhibition were investigated. The results showed that pre-spraying H2O2 inhibitor and scavenger decreased the H2O2 and MDA contents, while increased the root and shoot dry weights of cucumber seedling under NaCl stress. Correlation analysis showed that H2O2 content was negatively correlated with root and shoot dry weights. This result implied that the salt-induced H2O2 accumulation was an important reason for plant growth inhibition caused by salt stress.
2. Cucumber plants (cv.'Jinchun No.2') were self-grafted or grafted onto salt-tolerant rootstock'Chaojiquanwang'(Cucurbita moschata Duch). The changes in plant growth and antioxidant defense system of non-grafted, self-grafted, and rootstock-grafted cucumber seedlings under 0 and 75 mM NaCl stress were investigated. The results showed that cucumber plants grafted onto salt-tolerant rootstock had higher activities of CAT, APX, GR, and MDAR, GSH/GSSG ratio, and the ferric reducing ability/antioxidant power, thereby decreasing the H2O2 and MDA contents in cucumber tissue. As a result, the salt-induced growth inhibition was alleviated.
3. Chloroplast ultrastructure and the AsA-GSH cycle system in chloroplasts of self-grafted and rootstock-grafted cucumber leaves under 0,50, and 100 mM NaC1 stress were investigated. The results showed that the activities of APX, GR, and DHAR involved in AsA-GSH cycle and AsA/DHA ratio in the chloroplasts of rootstock-grafted plants under 50 and 100 mM NaCl, as well as the MDAR activity, GSH content, and GSH/GSSG ratio under 100 mM NaCl were all significant higher than those in self-grafted plants. This resulted in the lower H2O2 content in the chloroplast of rootstock-grafted plants. With increasing levels of NaC1 concentration, the chloroplast structure of self-grafted plants was visibly damaged. Particularly under 100 mM NaCl, the stacks of grana and intergranal lamellae were thin and obscure, even fractured, whereas the damage in the chloroplast of rootstock-grafted plants were less severe. The results suggest that rootstock grafting enhances the H2O2-scavenging capacity of the AsA-GSH cycle in the chloroplasts under salt stress, thereby protecting the chloroplast structure and improving the photosynthetic performance and growth of cucumber seedlings.
4. Changes in the time course of H2O2 content and main antioxidant enzyme (CAT, APX, GR and MDAR) activities in the leaves of self-grafted and rootstock-grafted cucumber plants under 0 and 75 mM NaCl stress, as well as the mRNA levels of antioxidant enzyme genes in grafted cucumber plants under NaCl stress for 120 h were studied. The results showed that the H2O2 content in self-grafted leaves increased gradually during NaC1 stress from 24 h to 120 h, and showing a peak at 72 h, whereas the H2O2 content in rootstock-grafted increased more slowly and kept lower all the time. This could be associated with its higher antioxidant enzyme activities during NaCl stress from 48 h to 120 h. The changes in the mRNA level of CAT and cAPX in self-grafted plants leaves and roots, respectively, and the mRNA level of cAPX in rootstock-grafted plants leaves were consistent with the changes in their enzyme activity levels under NaCl stress. This result indicated that the activities of these enzymes were regulated at their mRNA levels.
5. Two cucumber cultivars (cv.'Jinchun No.2', a relative salt-sensitive cultivar, and 'Jinyu No.1', a relative salt-tolerant cultivar) were grafted onto two rootstocks ('Figleaf Gourd', a relative salt-sensitive cultivar, and'Chaojiquanwang', a relative salt-tolerant cultivar), respectively. The effects of scion and rootstock on plant growth and the antioxidant defense systems of cucumber were assessed. The results showed that the salt injury index of plants grafted onto Chaojiquanwang was lower and the growth reduction was less severe under 100 mM NaCl stress. Correlation analysis showed that this could be partially attributed to its lower leaf Na+content and root H2O2 content, as well as the higher activities of SOD, POD, and CAT in the roots. This result implied that the salt-tolerant rootstock played a primary important effect in determining the salt tolerance of grafted cucumber seedlings under salt stress compared with the scion genotypes.
6. A salt-sensitive cucumber cultivar (cv.'Jinchun No.2') and a salt-tolerant pumpkin cultivar ('Chaojiquanwang') were reciprocal grafted. Four types of grafted plants (scion/rootstock) were obtained, including cucumber/cucumber, pumpkin/pumpkin, cucumber/pumpkin, and pumpkin/cucumber. The effects of 0 and 75 mM NaC1 stress on H2O2 and MDA contents, antioxidant enzyme activities, ABA content and some signal molecules involved in ABA signal pathway (H2O2, NO and MAPK cascades) in grafted plant roots were investigated. The results showed that the ABA contents increased and the expression of signal regulatory genes such as MAPK1 and MAPK6 up-regulated significantly in the grafted plants roots under NaC1 stress when pumpkin was used as rootstock. The results implied that salinity induced the production of ABA and the response of MAPK cascades mediated by ABA in salt-tolerant rootstock roots. This resulted in the higher H2O2-scavenging enzyme activities in salt-tolerant rootstock roots for controlling the H2O2 and MDA contents at lower levels.
In conclusion, the present study suggests that the salt-induced H2O2 production result in the plant growth inhibition caused by salt stress. Grafting with salt-tolerant rootstock can improve the salt tolerance of cucumber seedlings by regulating the H2O2-scavenging system in cucumber organic and chloroplast. The salt-tolerant rootstock plays a primary important effect in determining the salt tolerance of cucumber seedlings under salt stress compared with the scion genotypes. This could be attributed to the production of ABA and the regulation of ABA-mediated MAPK cascades in salt-tolerant rootstock roots under salt stress.
引文
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