外源亚精胺提高黄瓜幼苗耐盐性的生理调节功能研究
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摘要
设施土壤次生盐渍化是目前国内外设施栽培中的主要土壤障碍因子,严重制约着设施蔬菜的可持续发展。设施土壤发生盐害时,对黄瓜植株生长、产量和品质造成一系列不良影响,阻碍了黄瓜设施栽培的优质高效生产。多胺是生物代谢过程中产生的一类具有强烈生物活性的低分子量脂肪族含氮碱,研究表明多胺在植物抗逆境胁迫中发挥着重要作用。然而,关于多胺代谢变化与蔬菜作物耐盐性的关系以及多胺发挥的生理调节功能至今尚未阐明。
本文以耐盐性具有明显差异的两个黄瓜(Cucumis sativus L.)品种(耐盐性较强的‘长春密刺'和耐盐性较弱的‘津春2号')为试材,采用营养液栽培,通过营养液中添加外源Spd和多胺代谢抑制剂(MGBG和o-phen),研究盐(50 mmol·L~(-1)NaCl)胁迫下黄瓜植株生长、多胺代谢、活性氧代谢、质子泵活性、氮代谢、渗透调节物质积累等方面的变化,探讨黄瓜叶片和根系中多胺代谢变化、细胞器结合态多胺含量变化与植株耐盐性的关系,并探讨外源Spd在提高黄瓜耐盐性中发挥的生理调节功能。主要研究结果如下:
盐胁迫下,黄瓜幼苗叶片和根系中多胺代谢发生明显变化:‘长春密刺'幼苗中ADC、ODC、SAMDC和DAO活性升高幅度明显大于‘津春2号',而PAO活性升高幅度明显小于‘津春2号',引起‘津春2号'(游离态Spd+游离态Spm)/游离态Put比值显著降低、游离态Put/总游离态多胺比值显著升高,而‘长春密刺'变化不明显,且‘津春2号'体内结合态和束缚态尤其是束缚态多胺含量增加幅度明显小于‘长春密刺'。盐胁迫下外源0.1 mmol·L~(-1)Spd处理进一步提高了黄瓜叶片和根系中SAMDC和DAO活性,降低了PAO活性,引起(游离态Spd+游离态Spm)/游离态Put比值升高以及游离态Put/总游离态多胺比值降低,也引起结合态和束缚态尤其是束缚态多胺含量的显著增加,同时明显缓解了盐胁迫对黄瓜幼苗生长的抑制。外源MGBG处理能显著降低盐胁迫下幼苗体内游离态Spd和Spm含量,增加游离态Put含量,同时加重盐胁迫对幼苗生长的抑制,而外源Spd和Spm能缓解MGBG的伤害作用,且以Spd的缓解效果更明显。外源o-phen处理能显著降低盐胁迫下幼苗体内束缚态多胺含量,同时加重盐胁迫对幼苗生长的抑制。表明黄瓜幼苗叶片和根系中游离态Put迅速向游离态Spd和Spm转化,进而提高(游离态Spd+游离态Spm)/游离态Put比值、降低游离态Put/总游离态多胺比值,以及游离态Put迅速向结合态和束缚态多胺转化,尤其是向束缚态多胺转化,进而提高束缚态多胺含量,有利于提高黄瓜幼苗的耐盐性。
盐胁迫下,黄瓜细胞器结合态多胺含量变化与植株耐盐性密切相关:盐胁迫下,‘长春密刺'叶绿体TGase活性、结合态Put和Spd含量显著增加,而‘津春2号'变化不明显;‘长春密刺'叶绿体抗氧化酶活性和抗氧化剂含量增加幅度大于‘津春2号',而叶绿体H_2O_2和MDA含量升高幅度、净光合速率降低幅度小于‘津春2号'。暗示了叶绿体结合态Put和Spd含量的升高有利于阻止盐胁迫对叶绿体的伤害。外源Spd显著提高了盐胁迫下黄瓜叶绿体TGase活性、结合态多胺含量、抗氧化酶活性、抗氧化剂含量和净光合速率,降低了H_2O_2和MDA含量。盐胁迫下,‘长春密刺'根系线粒体结合态Spd含量、线粒体抗氧化酶活性和抗氧化剂含量增加幅度大于‘津春2号',而线粒体H_2O_2和MDA含量升高幅度小于‘津春2号'。暗示了线粒体结合态Spd含量的升高有利于阻止盐胁迫对线粒体的伤害。外源Spd显著提高了盐胁迫下黄瓜根系线粒体结合态多胺含量、抗氧化酶活性和抗氧化剂含量,降低了H_2O_2和MDA含量升高幅度。盐胁迫下,‘长春密刺'根系质膜结合态Spd含量、质膜H~+-ATPase活性显著升高,而‘津春2号'变化不明显;外源Spd显著提高了质膜结合态Spd含量,同时显著提高了质膜H~+-ATPase活性。表明黄瓜根系质膜结合态Spd通过提高质膜H~+-ATPase活性从而增强幼苗耐盐性。盐胁迫下,‘长春密刺'根系液泡膜结合态Put和Spd含量、液泡膜H~+-ATPase、H~+-PPase活性下降幅度小于‘津春2号';外源Spd显著提高了液泡膜结合态Spd含量,同时显著提高了液泡膜H~+-ATPase活性。表明黄瓜根系液泡膜结合态Spd通过提高液泡膜H~+-ATPase活性从而增强幼苗耐盐性。
Spd参与盐胁迫下黄瓜体内活性氧代谢的调节:外源Spd能进一步提高盐胁迫下黄瓜叶片和根系中SOD、POD和CAT等抗氧化酶活性,降低O_2~-产生速率、H_2O_2和MDA含量及电解质渗漏率。并且叶片和根系中束缚态Spd含量与SOD、POD和CAT活性间呈显著正相关。而外源o-phen能显著降低盐胁迫下幼苗体内束缚态Spd含量,同时降低抗氧化酶活性,提高活性氧水平和膜脂伤害程度。表明外源Spd通过提高盐胁迫下黄瓜幼苗内源多胺尤其是内源束缚态Spd含量,从而提高抗氧化酶活性,降低活性氧水平以及膜脂伤害。
Spd参与盐胁迫下黄瓜体内氮代谢的调节:外源Spd显著抑制了盐胁迫下黄瓜叶片和根系中DNA、RNA等核酸含量的降低;抑制了蛋白酶活性的升高,进而减缓了可溶性蛋白质含量的下降以及总游离氨基酸含量的升高;外源Spd促进了盐胁迫下黄瓜植株对NO_3~--N的吸收和转运,诱导叶片和根系中NR活性提高,促进NO_3~--N的还原,通过促进NH_4~+-N的同化,缓解过量NH_4~+-N积累造成的氨毒害,最终缓解盐胁迫引起的氮代谢紊乱。
Spd诱导盐胁迫下黄瓜体内有机渗透调节物质的积累:外源Spd显著提高了盐胁迫下黄瓜叶片和根系中吡咯啉-5-羧酸合成酶(P5CS)活性,但对脯氨酸脱氢酶(ProDH)活性无明显影响,通过促进脯氨酸合成进而引起脯氨酸含量增加;外源Spd对可溶性糖含量无明显影响。表明外源Spd通过诱导脯氨酸的积累进行渗透调节,提高植株水分含量,同时进一步缓解氨毒害和活性氧伤害,最终缓解幼苗生长抑制。
Soil secondary salinization in protected cultivation is one of the major soil barrier factors which are seriously restricting the sustainable development of vegetable crops in greenhouse.Cucumber is an important horticultural crop.When cucumber plants are subjected to soil secondary salinization,the plant growth,yield and quality of cucumber is reduced.Polyamines are ubiquitous low-molecular-weight aliphatic amines that are involved in regulation of plant growth and development.It was reported that polyamines were involved in the defense reaction of higher plants to various environmental stresses. However,the relationship between polyamine metabolism and salt tolerance of vegetables and the physiological regulation function of polyamines is still not well known.
In the present study,two cultivars of cucumber(Cucumis sativus L.),cvs.Changchun mici and Jinchun No.2,were used.It has been reported that cv.Jinchun No.2 is salinity-sensitive as in most cucumber cultivars,whereas cv.Changchun mici is comparatively tolerant to high salinity.By using exogenous Spd and polyamine metabolism inhibitor,MGBG and o-phen,we investigated the effects of salt stress(50 mmol·L~(-1)NaCl) on plant growth,polyamine metabolism,reactive oxygen metabolism,proton pump activity, nitrogen metabolism,osmo-compatible solute accumulation.The objectives of the present study were to elucidate the relationship between the changes of the polyamine metabolism in leaves and roots of cucumber and salt tolerance,the relationship between the contents of organelle bound polyamines and salt tolerance,and the physiology regulation function of exogenous Spd in improving salt tolerance of cucumber plants.Main research results were as follows:
Salt stress disturbed the polyamine metabolism homeostasis in leaves and roots of cucumber:During exposure to salt stress,the activities of ADC,ODC,SAMDC and DAO increased significantly,which were greater in cv.Changchun mici than in cv.Jinchun No.2. Salt stress also caused a significant increase in PAO activity,which was more marked in cv. Jinchun No.2 than in cv.Changchun mici.As a result,the ratios of(free-Spd+free-Spm) /free-Put decreased and the ratios of free-Put/total free-polyamines increased significantly in cv.Jinchun No.2,but not in cv.Changchun mici.Moreover,the salt stress-induced the increase of conjugated and bound polyamines,especially bound polyamines,was greater in cv.Changchun mici than in cv.Jinchun No.2.Under salt stress,exogenous Spd increased SAMDC and DAO activities,but decreased PAO activity.As a result,exogenous Spd induced the increase of the ratios of(free-Spd+free-Spm)/free-Put and the decrease of the ratios of free-Put/total free-polyamines.Exogenous Spd also induced the increase of conjugated and bound polyamines,especially bound polyamines.In addition,exogenous Spd reversed the detrimental effects of salt stress on plant growth of cucumber.Compared to the NaCl treatment,exogenous MGBG decreased significantly the contents of free Spd and Spm,but increased significantly the contents of free Put,and aggravated the salt stress-induced plant growth inhibition.The inhibiting effect of MGBG could be reversed partly by the combination with exogenous Spd and Spm,and Spd was the more effective. Compared to the NaCl treatment,exogenous o-phen decreased significantly the contents of bound polyamines,and aggravated the salt stress-induced plant growth inhibition.These results indicate that the alleviation of salt injury could be achieved by increasing the conversion of free Put to free Spd and Spm,resulting in the increase of the ratios of(free Spd+free Spm)/free Put and the decrease of the ratios of free-Put/total free polyamines,and by increasing the conversion of free Put to soluble conjugated and insoluble bound polyamines,especially bound polyamines.
The contents of organelle bound polyamines plays important roles in improving salt tolerance of cucumber:During exposure to salt stress,the activity of TGase,the contents of bound Put and Spd in chloroplast of cucumber increased significantly in cv.Changchun mici,but not in cv.Jinchun No.2.The increase of antioxidant enzymes activities and the antioxidant contents was greater in cv.Changchun mici than in cv.Jinchun No.2,but the increase of H_2O_2 and MDA contents and the decrease of net photosynthetic rates were greater in cv.Jinchun No.2 than in cv.Changchun mici.These results indicate that the alleviation of chloroplast injury could be achieved by increasing the content of bound Put and Spd in chloroplast.Exogenous Spd increased the activities of TGase and antioxidant enzymes,the contents of bound polyamines and antioxidant,and net photosynthetic rates, and decreased the contents of H_2O_2 and MDA.During exposure to salt stress,the contents of bound Spd,the activities of antioxidant enzymes and the contents of antioxidant in mitochondria of cucumber roots increased greater in cv.Changchun mici than in cv. Jinchun No.2,but the increase of H_2O_2 and MDA contents was greater in cv.Jinchun No.2 than in cv.Changchun mici.These results indicate that the alleviation of mitochondria injury could be achieved by increasing the content of bound Spd in mitochondria. Exogenous Spd increased the activities of antioxidant enzymes,the contents of bound polyamines and the antioxidant,and decreased the contents of H_2O_2 and MDA.During exposure to salt stress,the content of plasma membrane bound Spd and the activity of H~+-ATPase in plasma membrane of cucumber roots increased significantly in cv. Changchun mici,but not in cv.Jinchun No.2.Exogenous Spd increased significantly the content of plasma membrane bound Spd and the activity of H~+-ATPase in plasma membrane of cucumber roots.These results indicate that plasma membrane bound Spd in roots might enhance the salt tolerance of cucumber plants via maintenance the activity of plasma membrane H~+-ATPase.During exposure to salt stress,the decrease of the content of tonoplast bound Put and Spd and the activities of H~+-ATPase and H~+-PPase in tonoplast of cucumber roots was greater in cv.Jinchun No.2 than in cv.Changchun mici.Exogenous Spd increased significantly the content of tonoplast bound Spd and the activities of H~+-ATPase and H~+-PPase in tonoplast of cucumber roots.These results indicate that tonoplast bound Spd in roots might enhance the salt tolerance of cucumber plants via maintenance the activity of tonoplast H~+-ATPase.
Spermidine takes part in the regulation of reactive oxygen metabolism of cucumber plants under salt stress:Exogenous Spd elevated the activities of antioxidant enzymes, SOD,POD and CAT,suppressed O_2~(-·) and H_2O_2 production,decreased the MDA content and leakage of electrolytes.A positive correlation was found between the contents of bound Spd and the activities of SOD,POD and CAT.Compared to the NaCl treatment,exogenous o-phen decreased significantly the contents of bound Spd and the activities of antioxidant enzymes,and aggravated the salt stress-induced reactive oxygen production and membrane damage.These results indicate that exogenous Spd can enhance antioxidant enzyme activities and reduce reactive oxygen production and membrane damage by increasing the bound Spd content,resulting in the improvement of the salt tolerance of cucumber plants.
Spermidine takes part in the regulation of nitrogen metabolism of cucumber plants under salt stress:Exogenous Spd inhibited significantly the decrease of DNA and RNA. Exogenous Spd also inhibited the increase of proteinase activity,alleviated the decrease of soluble protein content and the increase of free amino acids.Exogenous Spd improved the absorption and translocation of NO_3~--N in cucumber plants under salt stress,induced the increase of NR activity,and improved the deoxidization of NO_3~--N.Exogenous Spd also improved the assimilation of NH_4~+-N and decreased the accumulation of NH_4~+-N.As a result,the homeostasis of nitrogen metabolism in cucumber plants was alleviated.
Spermidine induces the accumulation of osmo-compatible solute in cucumber plants under salt stress:Exogenous Spd induced the increase of P5CS activity but not ProDH, resulting in the significant increase of proline level in cucumber plants under salt stress. Exogenous Spd has no effect on soluble sugar content in cucumber plants under salt stress. These results indicate that exogenous Spd can maintain high water content,alleviate ammonia and reactive oxygen injury in salt stressed cucumber seedlings by increasing the internal proline content,resulting in the alleviation of plant growth.
本文以耐盐性具有明显差异的两个黄瓜(Cucumis sativus L.)品种(耐盐性较强的‘长春密刺'和耐盐性较弱的‘津春2号')为试材,采用营养液栽培,通过营养液中添加外源Spd和多胺代谢抑制剂(MGBG和o-phen),研究盐(50 mmol·L~(-1)NaCl)胁迫下黄瓜植株生长、多胺代谢、活性氧代谢、质子泵活性、氮代谢、渗透调节物质积累等方面的变化,探讨黄瓜叶片和根系中多胺代谢变化、细胞器结合态多胺含量变化与植株耐盐性的关系,并探讨外源Spd在提高黄瓜耐盐性中发挥的生理调节功能。主要研究结果如下:
盐胁迫下,黄瓜幼苗叶片和根系中多胺代谢发生明显变化:‘长春密刺'幼苗中ADC、ODC、SAMDC和DAO活性升高幅度明显大于‘津春2号',而PAO活性升高幅度明显小于‘津春2号',引起‘津春2号'(游离态Spd+游离态Spm)/游离态Put比值显著降低、游离态Put/总游离态多胺比值显著升高,而‘长春密刺'变化不明显,且‘津春2号'体内结合态和束缚态尤其是束缚态多胺含量增加幅度明显小于‘长春密刺'。盐胁迫下外源0.1 mmol·L~(-1)Spd处理进一步提高了黄瓜叶片和根系中SAMDC和DAO活性,降低了PAO活性,引起(游离态Spd+游离态Spm)/游离态Put比值升高以及游离态Put/总游离态多胺比值降低,也引起结合态和束缚态尤其是束缚态多胺含量的显著增加,同时明显缓解了盐胁迫对黄瓜幼苗生长的抑制。外源MGBG处理能显著降低盐胁迫下幼苗体内游离态Spd和Spm含量,增加游离态Put含量,同时加重盐胁迫对幼苗生长的抑制,而外源Spd和Spm能缓解MGBG的伤害作用,且以Spd的缓解效果更明显。外源o-phen处理能显著降低盐胁迫下幼苗体内束缚态多胺含量,同时加重盐胁迫对幼苗生长的抑制。表明黄瓜幼苗叶片和根系中游离态Put迅速向游离态Spd和Spm转化,进而提高(游离态Spd+游离态Spm)/游离态Put比值、降低游离态Put/总游离态多胺比值,以及游离态Put迅速向结合态和束缚态多胺转化,尤其是向束缚态多胺转化,进而提高束缚态多胺含量,有利于提高黄瓜幼苗的耐盐性。
盐胁迫下,黄瓜细胞器结合态多胺含量变化与植株耐盐性密切相关:盐胁迫下,‘长春密刺'叶绿体TGase活性、结合态Put和Spd含量显著增加,而‘津春2号'变化不明显;‘长春密刺'叶绿体抗氧化酶活性和抗氧化剂含量增加幅度大于‘津春2号',而叶绿体H_2O_2和MDA含量升高幅度、净光合速率降低幅度小于‘津春2号'。暗示了叶绿体结合态Put和Spd含量的升高有利于阻止盐胁迫对叶绿体的伤害。外源Spd显著提高了盐胁迫下黄瓜叶绿体TGase活性、结合态多胺含量、抗氧化酶活性、抗氧化剂含量和净光合速率,降低了H_2O_2和MDA含量。盐胁迫下,‘长春密刺'根系线粒体结合态Spd含量、线粒体抗氧化酶活性和抗氧化剂含量增加幅度大于‘津春2号',而线粒体H_2O_2和MDA含量升高幅度小于‘津春2号'。暗示了线粒体结合态Spd含量的升高有利于阻止盐胁迫对线粒体的伤害。外源Spd显著提高了盐胁迫下黄瓜根系线粒体结合态多胺含量、抗氧化酶活性和抗氧化剂含量,降低了H_2O_2和MDA含量升高幅度。盐胁迫下,‘长春密刺'根系质膜结合态Spd含量、质膜H~+-ATPase活性显著升高,而‘津春2号'变化不明显;外源Spd显著提高了质膜结合态Spd含量,同时显著提高了质膜H~+-ATPase活性。表明黄瓜根系质膜结合态Spd通过提高质膜H~+-ATPase活性从而增强幼苗耐盐性。盐胁迫下,‘长春密刺'根系液泡膜结合态Put和Spd含量、液泡膜H~+-ATPase、H~+-PPase活性下降幅度小于‘津春2号';外源Spd显著提高了液泡膜结合态Spd含量,同时显著提高了液泡膜H~+-ATPase活性。表明黄瓜根系液泡膜结合态Spd通过提高液泡膜H~+-ATPase活性从而增强幼苗耐盐性。
Spd参与盐胁迫下黄瓜体内活性氧代谢的调节:外源Spd能进一步提高盐胁迫下黄瓜叶片和根系中SOD、POD和CAT等抗氧化酶活性,降低O_2~-产生速率、H_2O_2和MDA含量及电解质渗漏率。并且叶片和根系中束缚态Spd含量与SOD、POD和CAT活性间呈显著正相关。而外源o-phen能显著降低盐胁迫下幼苗体内束缚态Spd含量,同时降低抗氧化酶活性,提高活性氧水平和膜脂伤害程度。表明外源Spd通过提高盐胁迫下黄瓜幼苗内源多胺尤其是内源束缚态Spd含量,从而提高抗氧化酶活性,降低活性氧水平以及膜脂伤害。
Spd参与盐胁迫下黄瓜体内氮代谢的调节:外源Spd显著抑制了盐胁迫下黄瓜叶片和根系中DNA、RNA等核酸含量的降低;抑制了蛋白酶活性的升高,进而减缓了可溶性蛋白质含量的下降以及总游离氨基酸含量的升高;外源Spd促进了盐胁迫下黄瓜植株对NO_3~--N的吸收和转运,诱导叶片和根系中NR活性提高,促进NO_3~--N的还原,通过促进NH_4~+-N的同化,缓解过量NH_4~+-N积累造成的氨毒害,最终缓解盐胁迫引起的氮代谢紊乱。
Spd诱导盐胁迫下黄瓜体内有机渗透调节物质的积累:外源Spd显著提高了盐胁迫下黄瓜叶片和根系中吡咯啉-5-羧酸合成酶(P5CS)活性,但对脯氨酸脱氢酶(ProDH)活性无明显影响,通过促进脯氨酸合成进而引起脯氨酸含量增加;外源Spd对可溶性糖含量无明显影响。表明外源Spd通过诱导脯氨酸的积累进行渗透调节,提高植株水分含量,同时进一步缓解氨毒害和活性氧伤害,最终缓解幼苗生长抑制。
Soil secondary salinization in protected cultivation is one of the major soil barrier factors which are seriously restricting the sustainable development of vegetable crops in greenhouse.Cucumber is an important horticultural crop.When cucumber plants are subjected to soil secondary salinization,the plant growth,yield and quality of cucumber is reduced.Polyamines are ubiquitous low-molecular-weight aliphatic amines that are involved in regulation of plant growth and development.It was reported that polyamines were involved in the defense reaction of higher plants to various environmental stresses. However,the relationship between polyamine metabolism and salt tolerance of vegetables and the physiological regulation function of polyamines is still not well known.
In the present study,two cultivars of cucumber(Cucumis sativus L.),cvs.Changchun mici and Jinchun No.2,were used.It has been reported that cv.Jinchun No.2 is salinity-sensitive as in most cucumber cultivars,whereas cv.Changchun mici is comparatively tolerant to high salinity.By using exogenous Spd and polyamine metabolism inhibitor,MGBG and o-phen,we investigated the effects of salt stress(50 mmol·L~(-1)NaCl) on plant growth,polyamine metabolism,reactive oxygen metabolism,proton pump activity, nitrogen metabolism,osmo-compatible solute accumulation.The objectives of the present study were to elucidate the relationship between the changes of the polyamine metabolism in leaves and roots of cucumber and salt tolerance,the relationship between the contents of organelle bound polyamines and salt tolerance,and the physiology regulation function of exogenous Spd in improving salt tolerance of cucumber plants.Main research results were as follows:
Salt stress disturbed the polyamine metabolism homeostasis in leaves and roots of cucumber:During exposure to salt stress,the activities of ADC,ODC,SAMDC and DAO increased significantly,which were greater in cv.Changchun mici than in cv.Jinchun No.2. Salt stress also caused a significant increase in PAO activity,which was more marked in cv. Jinchun No.2 than in cv.Changchun mici.As a result,the ratios of(free-Spd+free-Spm) /free-Put decreased and the ratios of free-Put/total free-polyamines increased significantly in cv.Jinchun No.2,but not in cv.Changchun mici.Moreover,the salt stress-induced the increase of conjugated and bound polyamines,especially bound polyamines,was greater in cv.Changchun mici than in cv.Jinchun No.2.Under salt stress,exogenous Spd increased SAMDC and DAO activities,but decreased PAO activity.As a result,exogenous Spd induced the increase of the ratios of(free-Spd+free-Spm)/free-Put and the decrease of the ratios of free-Put/total free-polyamines.Exogenous Spd also induced the increase of conjugated and bound polyamines,especially bound polyamines.In addition,exogenous Spd reversed the detrimental effects of salt stress on plant growth of cucumber.Compared to the NaCl treatment,exogenous MGBG decreased significantly the contents of free Spd and Spm,but increased significantly the contents of free Put,and aggravated the salt stress-induced plant growth inhibition.The inhibiting effect of MGBG could be reversed partly by the combination with exogenous Spd and Spm,and Spd was the more effective. Compared to the NaCl treatment,exogenous o-phen decreased significantly the contents of bound polyamines,and aggravated the salt stress-induced plant growth inhibition.These results indicate that the alleviation of salt injury could be achieved by increasing the conversion of free Put to free Spd and Spm,resulting in the increase of the ratios of(free Spd+free Spm)/free Put and the decrease of the ratios of free-Put/total free polyamines,and by increasing the conversion of free Put to soluble conjugated and insoluble bound polyamines,especially bound polyamines.
The contents of organelle bound polyamines plays important roles in improving salt tolerance of cucumber:During exposure to salt stress,the activity of TGase,the contents of bound Put and Spd in chloroplast of cucumber increased significantly in cv.Changchun mici,but not in cv.Jinchun No.2.The increase of antioxidant enzymes activities and the antioxidant contents was greater in cv.Changchun mici than in cv.Jinchun No.2,but the increase of H_2O_2 and MDA contents and the decrease of net photosynthetic rates were greater in cv.Jinchun No.2 than in cv.Changchun mici.These results indicate that the alleviation of chloroplast injury could be achieved by increasing the content of bound Put and Spd in chloroplast.Exogenous Spd increased the activities of TGase and antioxidant enzymes,the contents of bound polyamines and antioxidant,and net photosynthetic rates, and decreased the contents of H_2O_2 and MDA.During exposure to salt stress,the contents of bound Spd,the activities of antioxidant enzymes and the contents of antioxidant in mitochondria of cucumber roots increased greater in cv.Changchun mici than in cv. Jinchun No.2,but the increase of H_2O_2 and MDA contents was greater in cv.Jinchun No.2 than in cv.Changchun mici.These results indicate that the alleviation of mitochondria injury could be achieved by increasing the content of bound Spd in mitochondria. Exogenous Spd increased the activities of antioxidant enzymes,the contents of bound polyamines and the antioxidant,and decreased the contents of H_2O_2 and MDA.During exposure to salt stress,the content of plasma membrane bound Spd and the activity of H~+-ATPase in plasma membrane of cucumber roots increased significantly in cv. Changchun mici,but not in cv.Jinchun No.2.Exogenous Spd increased significantly the content of plasma membrane bound Spd and the activity of H~+-ATPase in plasma membrane of cucumber roots.These results indicate that plasma membrane bound Spd in roots might enhance the salt tolerance of cucumber plants via maintenance the activity of plasma membrane H~+-ATPase.During exposure to salt stress,the decrease of the content of tonoplast bound Put and Spd and the activities of H~+-ATPase and H~+-PPase in tonoplast of cucumber roots was greater in cv.Jinchun No.2 than in cv.Changchun mici.Exogenous Spd increased significantly the content of tonoplast bound Spd and the activities of H~+-ATPase and H~+-PPase in tonoplast of cucumber roots.These results indicate that tonoplast bound Spd in roots might enhance the salt tolerance of cucumber plants via maintenance the activity of tonoplast H~+-ATPase.
Spermidine takes part in the regulation of reactive oxygen metabolism of cucumber plants under salt stress:Exogenous Spd elevated the activities of antioxidant enzymes, SOD,POD and CAT,suppressed O_2~(-·) and H_2O_2 production,decreased the MDA content and leakage of electrolytes.A positive correlation was found between the contents of bound Spd and the activities of SOD,POD and CAT.Compared to the NaCl treatment,exogenous o-phen decreased significantly the contents of bound Spd and the activities of antioxidant enzymes,and aggravated the salt stress-induced reactive oxygen production and membrane damage.These results indicate that exogenous Spd can enhance antioxidant enzyme activities and reduce reactive oxygen production and membrane damage by increasing the bound Spd content,resulting in the improvement of the salt tolerance of cucumber plants.
Spermidine takes part in the regulation of nitrogen metabolism of cucumber plants under salt stress:Exogenous Spd inhibited significantly the decrease of DNA and RNA. Exogenous Spd also inhibited the increase of proteinase activity,alleviated the decrease of soluble protein content and the increase of free amino acids.Exogenous Spd improved the absorption and translocation of NO_3~--N in cucumber plants under salt stress,induced the increase of NR activity,and improved the deoxidization of NO_3~--N.Exogenous Spd also improved the assimilation of NH_4~+-N and decreased the accumulation of NH_4~+-N.As a result,the homeostasis of nitrogen metabolism in cucumber plants was alleviated.
Spermidine induces the accumulation of osmo-compatible solute in cucumber plants under salt stress:Exogenous Spd induced the increase of P5CS activity but not ProDH, resulting in the significant increase of proline level in cucumber plants under salt stress. Exogenous Spd has no effect on soluble sugar content in cucumber plants under salt stress. These results indicate that exogenous Spd can maintain high water content,alleviate ammonia and reactive oxygen injury in salt stressed cucumber seedlings by increasing the internal proline content,resulting in the alleviation of plant growth.
引文
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