低氧胁迫下γ-氨基丁酸对甜瓜幼苗多胺代谢的影响及比较蛋白质组分析
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摘要
甜瓜(Cucumis melo L.)属典型的低氧敏感性植物,常常由于土壤紧实、灌水过多以及无土栽培中营养液溶氧浓度低、根垫形成等原因造成根际氧不足,造成产量和品质降低,现已成为制约设施甜瓜生产大规模发展的决定性障碍因素。本研究以‘西域一号’甜瓜为试验材料,采用营养液水培法,研究了低氧胁迫下外源添加γ-氨基丁酸(GABA)对甜瓜幼苗多胺代谢的影响,并通过比较蛋白质组学分析幼苗低氧胁迫下蛋白质组代谢的差异,筛选出低氧胁迫和外源GABA诱导产生的特异表达蛋白,为GABA提高甜瓜的耐低氧性提供了坚实的理论依据。主要研究结果如下:
(1)低氧胁迫下甜瓜根系和叶片中多胺含量呈先上升后下降的趋势,并在处理第4天达最大值;其中以游离态多胺含量最高,其次是结合态多胺,束缚态多胺含量最低;不同形态多胺在叶片中含量均为Spd>Put>Spm,在根系含量中均为Put>Spd>Spm;叶片中不同形态Spd、Spm含量大于根系,而不同形态Put含量则小于根系。外源GABA可促进Put、Spd、Spm的合成。而添加抑制剂D-Arg处理多胺含量显著降低。
(2)低氧胁迫诱导甜瓜幼苗根系和叶片中ADC、ODC、SAMDC活性显著提高;但根系中提高幅度较大,且在处理第4天达最大值,此后迅速下降,而叶片中ADC和SAMDC活性达到最大值后维持稳定。外源GABA提高了根系及叶片中ADC、ODC及SAMDC活性。
(3)低氧胁迫导致根系和叶片游离态和细胞壁结合态DAO、PAO活性显著提高,其中根系细胞壁结合态PAO和DAO活性显著低于叶片,而游离态DAO、PAO活性显著低于叶片。外源添加GABA处理显著降低了低氧胁迫下甜瓜幼苗根系和叶片DAO和PAO的活性。
(4)筛选并建立了甜瓜根系蛋白分离分析体系,通过对三种甜瓜根系蛋白提取方法的分析结果表明:TCA/丙酮沉淀法和SDS/酚提法得到的蛋白总含量较低;SDS/酚提取法和Tris/酚抽提-甲醇/乙酸铵沉淀法在SDS和2-DE胶中分离得到的蛋白条带和蛋白点数较多;TCA/丙酮法对分子量较高的偏碱性区域的蛋白提取效果不好,双向电泳胶图中丢失了较多蛋白点。
(5)运用双向电泳技术对处理4天甜瓜幼苗根系总蛋白进行比较蛋白质组研究,结果表明,低氧胁迫处理以及外源GABA处理胶图中蛋白点都显著增加;其中正常通气条件下外源GABA处理与对照相比,双向电泳胶图之间存在54个差异蛋白点;低氧胁迫处理与对照相比双向电泳胶图之间存在62个差异蛋白点;低氧胁迫外源添加GABA处理与低氧胁迫处理相比双向电泳胶图之间存在45个差异蛋白点。从中选择13个差异极显著点进行二级质谱分析,鉴定得到一些与抗逆相关的蛋白,如谷胱甘肽转移酶、NADP氧化还原酶、磷酸甘油酸激酶等,但是有4个差异点却未能得到准确鉴定结果,有待甜瓜数据库完善后进一步鉴定。
Melon (Cucumis melo L.) is a typical hypoxic-sensitive plant. But the yield and quality of melon often reduced by lacking of oxygen in the rhizosphere according to soil compaction, irrigation excessive, low dissolved oxygen concentration of nutrient solution in soilless culture and the root mat formation, which has become the decisive obstacles to large-scale development of melon production. In this paper, hydroponic experiments were performed to investigate the effects of exogenousγ-aminobutyric acid (GABA) application on polyamine metabolism and comparative proteomics of root total proteins in seedlings of melon cultivar‘Xiyu No.1’under hypoxia stress, which clarified the possible mechanisms of GABA enhancing hypoxia-tolerance in melon through metabolomics and proteomics technology.
The main results were presented as follows:
1. Hypoxia stress induced a significant increase in polyamines contents, polyamines of free form had higher content but lower content of bound form in leaves and roots of melon seedlings, the content sequence of PAs in roots was Put>Spd>Spm, and in leaves was Spd>Put>Spm. The three forms Spd and Spm contents in leaves were higher than that in roots, but Put contents were lower. Exogenous GABA significantly increased the contents of Put, Spd and Spm. However, simultaneously applying GABA and VGB could significantly decreased the content of PAs under hypoxia stress.
2. Hypoxia stress induced a significant increase in activities of polyamine synthesis in melon seedlings, especially the change in roots, activities of polyamine synthesis in roots fallen sharply after reached maximum in 4 days, but maintained stable of ADC and SAMDC activities in leaves after reached maxmum. Compare with hypoxia treatment, exogenous GABA significantly increased the activities of ADC, ODC and SAMDC both in roots and in leaves.
3. Compared with normoxic treatment, the activities of PBS-extractable form and cell-wall-bound form polyamine oxidase enzymes were higher in melon roots and in leaves under hypoxia stress; The activities of cell-wall-bound form DAO and PAO in roots were lower than in leaves, while the activities of PBS-extractable form DAO and PAO were higher than in leaves. Compared with hypoxia stress treatment, exogenous GABA significantly decreased the act ivies of DAO and PAO.
4. We selected and established a system for separation and analysis the melon root protein. Three improved methods were adopted to extract total protein,and which were analyzed and compared with each other, the results showed that TCA/acetone precipitation and SDS/phenol extraction had lower protein contents, more bands and spots could be detected on SDS gels and 2-DE gels by SDS/phenol extraction and Tris/phenol extraction-methanol/ammonium acetate precipitation, while TCA/acetone precipitation got the minimum spots and lost much of basic proteins.
5. Comparative proteomics analysis of melon root total proteins have been preformed after 4 days treatment, we found proteomics of the four treatments in melon root had some obvious changes. Employing computer analysis, and the result indicated that valid protein spots in gels of hypoxia and hypoxia+GABA treatments were significantly increased. Between normoxic treatment and normoxic+GABA treatment, there were 54 differential protein spots. Compared with normoxic and hypoxia treatments, 62 significant differential protein spots were detected in the images. And 45 differential protein spots were detected in images of hypoxia treatment and hypoxia+GABA treatment. In this paper, 13 extremely significant differential protein spots have been chosen to identify by MALDI-TOF/TOF Mass Spectrometry, and the identified proteins include some plant anti-versus relative proteins such as glutathione transferase, ferrodoxin NADP oxidoreductase, cytosolic phosphoglycerate kinase and so on. But there were 4 spots have not high identification rate by mass spectrometry, and these proteins need further study when a prefect database has established.
(1)低氧胁迫下甜瓜根系和叶片中多胺含量呈先上升后下降的趋势,并在处理第4天达最大值;其中以游离态多胺含量最高,其次是结合态多胺,束缚态多胺含量最低;不同形态多胺在叶片中含量均为Spd>Put>Spm,在根系含量中均为Put>Spd>Spm;叶片中不同形态Spd、Spm含量大于根系,而不同形态Put含量则小于根系。外源GABA可促进Put、Spd、Spm的合成。而添加抑制剂D-Arg处理多胺含量显著降低。
(2)低氧胁迫诱导甜瓜幼苗根系和叶片中ADC、ODC、SAMDC活性显著提高;但根系中提高幅度较大,且在处理第4天达最大值,此后迅速下降,而叶片中ADC和SAMDC活性达到最大值后维持稳定。外源GABA提高了根系及叶片中ADC、ODC及SAMDC活性。
(3)低氧胁迫导致根系和叶片游离态和细胞壁结合态DAO、PAO活性显著提高,其中根系细胞壁结合态PAO和DAO活性显著低于叶片,而游离态DAO、PAO活性显著低于叶片。外源添加GABA处理显著降低了低氧胁迫下甜瓜幼苗根系和叶片DAO和PAO的活性。
(4)筛选并建立了甜瓜根系蛋白分离分析体系,通过对三种甜瓜根系蛋白提取方法的分析结果表明:TCA/丙酮沉淀法和SDS/酚提法得到的蛋白总含量较低;SDS/酚提取法和Tris/酚抽提-甲醇/乙酸铵沉淀法在SDS和2-DE胶中分离得到的蛋白条带和蛋白点数较多;TCA/丙酮法对分子量较高的偏碱性区域的蛋白提取效果不好,双向电泳胶图中丢失了较多蛋白点。
(5)运用双向电泳技术对处理4天甜瓜幼苗根系总蛋白进行比较蛋白质组研究,结果表明,低氧胁迫处理以及外源GABA处理胶图中蛋白点都显著增加;其中正常通气条件下外源GABA处理与对照相比,双向电泳胶图之间存在54个差异蛋白点;低氧胁迫处理与对照相比双向电泳胶图之间存在62个差异蛋白点;低氧胁迫外源添加GABA处理与低氧胁迫处理相比双向电泳胶图之间存在45个差异蛋白点。从中选择13个差异极显著点进行二级质谱分析,鉴定得到一些与抗逆相关的蛋白,如谷胱甘肽转移酶、NADP氧化还原酶、磷酸甘油酸激酶等,但是有4个差异点却未能得到准确鉴定结果,有待甜瓜数据库完善后进一步鉴定。
Melon (Cucumis melo L.) is a typical hypoxic-sensitive plant. But the yield and quality of melon often reduced by lacking of oxygen in the rhizosphere according to soil compaction, irrigation excessive, low dissolved oxygen concentration of nutrient solution in soilless culture and the root mat formation, which has become the decisive obstacles to large-scale development of melon production. In this paper, hydroponic experiments were performed to investigate the effects of exogenousγ-aminobutyric acid (GABA) application on polyamine metabolism and comparative proteomics of root total proteins in seedlings of melon cultivar‘Xiyu No.1’under hypoxia stress, which clarified the possible mechanisms of GABA enhancing hypoxia-tolerance in melon through metabolomics and proteomics technology.
The main results were presented as follows:
1. Hypoxia stress induced a significant increase in polyamines contents, polyamines of free form had higher content but lower content of bound form in leaves and roots of melon seedlings, the content sequence of PAs in roots was Put>Spd>Spm, and in leaves was Spd>Put>Spm. The three forms Spd and Spm contents in leaves were higher than that in roots, but Put contents were lower. Exogenous GABA significantly increased the contents of Put, Spd and Spm. However, simultaneously applying GABA and VGB could significantly decreased the content of PAs under hypoxia stress.
2. Hypoxia stress induced a significant increase in activities of polyamine synthesis in melon seedlings, especially the change in roots, activities of polyamine synthesis in roots fallen sharply after reached maximum in 4 days, but maintained stable of ADC and SAMDC activities in leaves after reached maxmum. Compare with hypoxia treatment, exogenous GABA significantly increased the activities of ADC, ODC and SAMDC both in roots and in leaves.
3. Compared with normoxic treatment, the activities of PBS-extractable form and cell-wall-bound form polyamine oxidase enzymes were higher in melon roots and in leaves under hypoxia stress; The activities of cell-wall-bound form DAO and PAO in roots were lower than in leaves, while the activities of PBS-extractable form DAO and PAO were higher than in leaves. Compared with hypoxia stress treatment, exogenous GABA significantly decreased the act ivies of DAO and PAO.
4. We selected and established a system for separation and analysis the melon root protein. Three improved methods were adopted to extract total protein,and which were analyzed and compared with each other, the results showed that TCA/acetone precipitation and SDS/phenol extraction had lower protein contents, more bands and spots could be detected on SDS gels and 2-DE gels by SDS/phenol extraction and Tris/phenol extraction-methanol/ammonium acetate precipitation, while TCA/acetone precipitation got the minimum spots and lost much of basic proteins.
5. Comparative proteomics analysis of melon root total proteins have been preformed after 4 days treatment, we found proteomics of the four treatments in melon root had some obvious changes. Employing computer analysis, and the result indicated that valid protein spots in gels of hypoxia and hypoxia+GABA treatments were significantly increased. Between normoxic treatment and normoxic+GABA treatment, there were 54 differential protein spots. Compared with normoxic and hypoxia treatments, 62 significant differential protein spots were detected in the images. And 45 differential protein spots were detected in images of hypoxia treatment and hypoxia+GABA treatment. In this paper, 13 extremely significant differential protein spots have been chosen to identify by MALDI-TOF/TOF Mass Spectrometry, and the identified proteins include some plant anti-versus relative proteins such as glutathione transferase, ferrodoxin NADP oxidoreductase, cytosolic phosphoglycerate kinase and so on. But there were 4 spots have not high identification rate by mass spectrometry, and these proteins need further study when a prefect database has established.
引文
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