水分胁迫下外源脱落酸提高甘蔗抗旱性的机理研究
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摘要
甘蔗是中国第一大糖料作物,甘蔗糖占我国食糖总产量90%以上,而我国甘蔗种植面积80%以上为丘陵旱地,蔗区农业水利条件较差,降水不均衡,土壤保水能力差,每年都面临不同程度干旱威胁。选育抗旱性强、水分利用率高的甘蔗品种是我国甘蔗糖业可持续发展的重要保证,但由于甘蔗遗传背景复杂,通过常规杂交育种方法进行甘蔗遗传改良所需周期比较长,亟需其它辅助手段解决甘蔗抗旱性问题。本研究选用甘蔗品种桂糖21号(GT21),比较在进行水分胁迫和胁迫加喷施外源脱落酸(ABA)处理条件下,甘蔗叶片内相关生理生化变化及其与抗旱性的关系,运用基因芯片比较两处理条件下差异表达基因异同并筛选ABA诱导基因,对ABA合成及信号转导过程关键基因进行克隆及表达分析,主要研究内容和结果如下：
1.以GT21为材料,研究水分胁迫及胁迫加叶面喷施ABA对甘蔗内源ABA含量及相关生理生化指标的影响,结果表明,干旱及干旱加外施ABA条件下,甘蔗内源ABA含量水平上升,但干旱加ABA处理增幅更显著,甘蔗叶片相对含水量随着胁迫加剧而逐渐下降,外施ABA能提高甘蔗保水能力。水分胁迫下,甘蔗叶内脯氨酸、可溶性蛋白、H2O2、丙二醛(MDA)含量增加,ABA处理能使渗透调节物质含量处于更高水平,并降低MDA含量,缓解其积累给细胞带来的伤害。水分胁迫及ABA处理均能提高GT21叶片中IAA含量而降低GA含量。ABA处理能防止叶绿素降解并对干旱引起的最大光能转化效率(Fv/Fm)、最大光化学效率(Fv'/Fm')、光系统II实际量子效率(PS2)下降有明显缓解作用。在干旱条件下,H2O2积累伴随着抗氧化系统相关酶CAT、GPX、GR和APX活性提高,ABA处理能进一步提高上述酶活性而使活性氧清除能力得到增强,表明在干旱条件下,外施ABA处理能增强甘蔗抗氧化防护系统,从而提高其抗旱性。
2.以已发表的NCED蛋白序列保守区域为模板,设计简并引物,运用RT-PCR方法,从GT21中克隆得到NCED基因序列表达标签后,采用RACE-PCR技术,扩增出该基因全长cDNA序列并命名为SoNCED。序列分析表明SoNCED全长2521bp,包含长度为1827bp的开放阅读框,编码608个氨基酸,推导氨基酸序列分子量65.9kDa,等电点6.04,聚类分析表明其与玉米和高梁等同属C4植物的NCED进化关系较近。生物信息学分析表明,SoNCED的N末端有典型叶绿体转运肽结构,组成其基本二级结构的α-helices、β-strands、β-propeller、与铁离子结合起催化作用的组氨酸位点,α-helices结构中富含疏水氨基酸残基等,性质高度保守。实时荧光定量PCR分析结果表明,25%PEG-6000模拟干旱胁迫处理、对照加ABA及干旱加ABA处理下,GT21叶片及根系SoNCED都随着02-产生速率升高表达量显著上升,并与ABA含量增长具有协同效应,表明SoNCED参与甘蔗叶片及根系内源ABA合成调控。
3.应用安捷伦4x44K基因芯片研究水分胁迫及胁迫加叶面喷施ABA对甘蔗基因表达的影响,结果表明,ABA处理在第3、第5天差异表达基因数量均高于单施干旱处理；胁迫前期,两处理差异基因以上调表达为主,后期下调表达占多数。从ABA处理第3天的样品中筛选到231个ABA响应基因,与第5天各处理差异基因相比,其中71个基因只在ABA处理第3天表达,72个在所有处理中共表达,38个只在ABA处理第3、第5天共表达,50个只在干旱处理第5天表达。使用BlastX对上述231个基因进行功能注释,共获得有效注释信息基因173个,基因功能分类结果表明,功能未知的假定蛋白所占比例最高,约占总数10%；参与脂类代谢、信号转导,编码蛋白激酶和渗透调节物质基因所占比例次之,都约占9%；参与蛋白质代谢、ABA代谢基因约占总数的6%；与光合作用、碳水化合物代谢、转运蛋白相关的基因分别约占5%；其它差异表达基因参与的功能还有呼吸作用、转录因子、激素代谢、氧化胁迫、氮代谢、细胞代谢等。实时荧光定量PCR实验表明芯片实验结果是可靠和可重复的。
4.以芯片实验基因筛选结果为参考,应用RCAE技术扩增出与水稻中非酵解型蛋白激酶基因SAPK3同源性极高的甘蔗基因全长cDNA并命名为SoSnRK2.1。序列分析表明,该基因全长cDNA为1385bp,开放阅读框长为1002bp,编码333个氨基酸,并与其它物种SnRK2基因具有很高同源性,聚类分析显示该基因属于SnRK2家族第一亚家族。预测SoSnRK2.1分子量为37.8kDa,等电点为5.54,是亲水性非跨膜蛋白质,在第5-261位氨基酸之间存在蛋白激酶保守催化域,分别具有11、3和8个潜在丝氨酸、苏氨酸、络氨酸磷酸化位点,二级结构中以环状结构所占比例最大。实时荧光定量PCR结果显示该基因在甘蔗叶片中受ABA及水分胁迫诱导表达,与ABA含量有着协同增长关系,推测其在叶中起着ABA信号转导作用；根系中不同处理该基因表达整体呈下降趋势,推测与ABA信号转导通路冗余性相关。
Sugarcane is the most important plant for sugar, and cane sugar occupies over90%of sugar production in China. However, over80%of the crop is planted in the hilly unpland area, where the rainfall is uneven and irrigation is not available, and drought has become one of the major limiting factors for sugarcane production in China. Breeding sugarcane varieties with strong drought tolerance and high water use efficiency is one of the most important strategies for sustainable development of the sugar industry. Due to the complex genetic background of sugarcane, however, crop modification through conventional hybrid breeding program is time costly and lowly efficient, other supplementary approaches including chemical regulation are necessary for improving the sugarcane drought resistance in practice. The aims of this research were to compare the physiological changes in leaves between the treatments of drought and drought plus foliar application of abscisic acid (ABA) under water stress conditions using sugarcane variety GT21, furthermore, to sieve the ABA responsive genes with microarray, and to clone the key gene involved in ABA biosynthesis and signal transduction.
1. An experiment was set up to investigate the interrelationship between drought induced ABA biosynthesis and antioxidative defense system, and to confer the farther role of foliar application of ABA in imparting drought tolerance to the sugarcane plant. The results showed that drought treatment enhanced the ABA concentration in leaf but it was significantly higher in combined ABA treatment, suggesting its biosynthesis was triggered in leaf by the ABA application. The relative water content was dramatically decreased by drought treatment but it could maintain higher in ABA treatment. Both drought and ABA treatment could result in an increase in proline, soluble protein, auxin, H2O2and malondialdehyde (MDA) content but the plants applied with ABA were found to resist the accumulation of MDA and gibberellin. ABA application decreased the degradation of chlorophyll, counteracted, at least in part, the decrease in maximal PS Ⅱ efficiency(Fv/Fm), antennae efficiency of PS Ⅱ (Fv/Fm) and quantum efficiency of PSⅡ (PS2). Overproduction of H2O2in plants treated with water stress was followed by higher activities of catalase (CAT), glutathione peroxidase (GPX), glutathione reductase (GR) and ascorbate peroxidase (APX), but this increase was further improved by the ABA treatment. The results clearly suggests that though the tolerant variety showed an enhanced protective system against drought conditions, the foliar application of ABA further improved its tolerance by continuously triggering the over expression of antioxidative defense system.
2. Degenerated and specific primers were designed from highly homologous regions of NCED gene sequences of different crop plants taken from NCBI database, a full-length cDNA encoding NCED gene was isolated and characterized from leaves of Saccharum officinarum L.(So) using RT-PCR and RACE-PCR. The isolated full-length SoNCED (2521bp) with1827bp ORF, encoded a peptide of608amino acids. The calculated molecular weight of the protein was65.9kDa with isoelectric point of6.04. Conserved domains prediction indicated a chloroplast-targeting peptide located at N-terminus of SoNCED protein. Phylogenetic tree, constructed by Neighbor-joining method indicated that deduced protein sequences of SoNCED shared high identity with the NCEDs reported from Zea mays and Sorghum bicolor which all belong to the C4plants. Sequence alignment revealed that the basic secondary structure including a-helices,[3-strands, P-propeller and His residues coordinating catalytic sites of SoNCED were highly conserved as in the NCEDs from other plants. Tissue specific expression analysis using Quantitative real-time PCR (qRT-PCR) showed a significant increase in SoNCED mRNA level and its correlation with O2-production rate and ABA accumulation in leaves and roots of sugarcane variety GT21when exposed to water stress. Further, the stimulation of SoNCED mRNA level, O2-production rate and ABA content after exogenous application of ABA proved its involvement in pathways providing tolerance to drought stress.
3. An Agilent4×44K microarray was applied to investigate the gene differential expression under water stress and foliar application ABA. The results showed that the number of differentially expressed genes was higher in the drought plus ABA treatment than drought treatment only. Large part of the genes was first induced then depressed following the aggravated water stress in both treatments. Total231genes were identified as ABA responsive from ABA treated samples at3rd day. Compared to the samples taken at5th day,71of the aforementioned genes were specifically expressed at3rd day in ABA treated plants,72genes were co-expressed in all treatments and days,38genes were co-expressed in ABA treatment, and50genes were specifically expressed at5th day in drought treated plants. Annotation results done by BlastX showed that173of the231genes could be validated. Function categories indicated that the hypothetical proteins have taken up about10%of the annotated genes, while genes functioned in lipid metabolism, signal transduction, protein kinase and osmoregulation have taken up about9%separately. Genes related to protein and hormone metabolism have taken up about6%separately.5%of the genes are related to photosynthesis, carbohydrate metabolism and transport separately. The remaining genes take part in respiration, transcription, oxidative stress, nitrogen metabolism, cellular metabolism and so on. The differential expression patterns of nine candidate genes were chose confirmed by qRT-PCR and the results showed that the expression changes of these candidates were generally consistent with the microarray results.
4. After annotation of the ABA responsive genes, one with highly homologous to the sucrose non-fermenting related protein kinase (SnRK2) gene SAPK3in Oryza sativa was selected for further analysis, and the cDNA full-length of this gene was amplified using RACE-PCR and named SoSnRK2.1. The isolated full-length of SoSnRK2.1was1385bp with1002bp ORF, encoding a peptide of333amino acids. Sequence analysis indicated that SoSnRK2.1showed high similarities with SnRK2from other plant species, and belong to the first subfamily when clustered with the SnRK2s taken from Arabidopsis thaliana using a Neighbor-joining method. The calculated molecular weight of the protein was37.8kDa with isoelectric point of5.54, and predicted as a hydrophilic protein without transmenbrane domains. There was a protein kinase conservative catalytic domain between the5-261amino acid sites, with11,3and8potential phosphorylation sites of serine, threonine and tyrosine separately, and the loop structure have taken the most part in the second structure prediction of SoSnRK2.1. Tissue specific expression analysis using qRT-PCR showed a significant increase in SoSnRK2.1mRNA level induced by foliar application of ABA and water stress, and correlated with ABA accumulation in leaves, which indicated the signal transduction function of this gene in leaves, while the expressions were suppressed with the accumulation of ABA content in roots, and this might be due to the redundancy of SnRK2genes in plants, but should be for further research.
1.以GT21为材料,研究水分胁迫及胁迫加叶面喷施ABA对甘蔗内源ABA含量及相关生理生化指标的影响,结果表明,干旱及干旱加外施ABA条件下,甘蔗内源ABA含量水平上升,但干旱加ABA处理增幅更显著,甘蔗叶片相对含水量随着胁迫加剧而逐渐下降,外施ABA能提高甘蔗保水能力。水分胁迫下,甘蔗叶内脯氨酸、可溶性蛋白、H2O2、丙二醛(MDA)含量增加,ABA处理能使渗透调节物质含量处于更高水平,并降低MDA含量,缓解其积累给细胞带来的伤害。水分胁迫及ABA处理均能提高GT21叶片中IAA含量而降低GA含量。ABA处理能防止叶绿素降解并对干旱引起的最大光能转化效率(Fv/Fm)、最大光化学效率(Fv'/Fm')、光系统II实际量子效率(PS2)下降有明显缓解作用。在干旱条件下,H2O2积累伴随着抗氧化系统相关酶CAT、GPX、GR和APX活性提高,ABA处理能进一步提高上述酶活性而使活性氧清除能力得到增强,表明在干旱条件下,外施ABA处理能增强甘蔗抗氧化防护系统,从而提高其抗旱性。
2.以已发表的NCED蛋白序列保守区域为模板,设计简并引物,运用RT-PCR方法,从GT21中克隆得到NCED基因序列表达标签后,采用RACE-PCR技术,扩增出该基因全长cDNA序列并命名为SoNCED。序列分析表明SoNCED全长2521bp,包含长度为1827bp的开放阅读框,编码608个氨基酸,推导氨基酸序列分子量65.9kDa,等电点6.04,聚类分析表明其与玉米和高梁等同属C4植物的NCED进化关系较近。生物信息学分析表明,SoNCED的N末端有典型叶绿体转运肽结构,组成其基本二级结构的α-helices、β-strands、β-propeller、与铁离子结合起催化作用的组氨酸位点,α-helices结构中富含疏水氨基酸残基等,性质高度保守。实时荧光定量PCR分析结果表明,25%PEG-6000模拟干旱胁迫处理、对照加ABA及干旱加ABA处理下,GT21叶片及根系SoNCED都随着02-产生速率升高表达量显著上升,并与ABA含量增长具有协同效应,表明SoNCED参与甘蔗叶片及根系内源ABA合成调控。
3.应用安捷伦4x44K基因芯片研究水分胁迫及胁迫加叶面喷施ABA对甘蔗基因表达的影响,结果表明,ABA处理在第3、第5天差异表达基因数量均高于单施干旱处理；胁迫前期,两处理差异基因以上调表达为主,后期下调表达占多数。从ABA处理第3天的样品中筛选到231个ABA响应基因,与第5天各处理差异基因相比,其中71个基因只在ABA处理第3天表达,72个在所有处理中共表达,38个只在ABA处理第3、第5天共表达,50个只在干旱处理第5天表达。使用BlastX对上述231个基因进行功能注释,共获得有效注释信息基因173个,基因功能分类结果表明,功能未知的假定蛋白所占比例最高,约占总数10%；参与脂类代谢、信号转导,编码蛋白激酶和渗透调节物质基因所占比例次之,都约占9%；参与蛋白质代谢、ABA代谢基因约占总数的6%；与光合作用、碳水化合物代谢、转运蛋白相关的基因分别约占5%；其它差异表达基因参与的功能还有呼吸作用、转录因子、激素代谢、氧化胁迫、氮代谢、细胞代谢等。实时荧光定量PCR实验表明芯片实验结果是可靠和可重复的。
4.以芯片实验基因筛选结果为参考,应用RCAE技术扩增出与水稻中非酵解型蛋白激酶基因SAPK3同源性极高的甘蔗基因全长cDNA并命名为SoSnRK2.1。序列分析表明,该基因全长cDNA为1385bp,开放阅读框长为1002bp,编码333个氨基酸,并与其它物种SnRK2基因具有很高同源性,聚类分析显示该基因属于SnRK2家族第一亚家族。预测SoSnRK2.1分子量为37.8kDa,等电点为5.54,是亲水性非跨膜蛋白质,在第5-261位氨基酸之间存在蛋白激酶保守催化域,分别具有11、3和8个潜在丝氨酸、苏氨酸、络氨酸磷酸化位点,二级结构中以环状结构所占比例最大。实时荧光定量PCR结果显示该基因在甘蔗叶片中受ABA及水分胁迫诱导表达,与ABA含量有着协同增长关系,推测其在叶中起着ABA信号转导作用；根系中不同处理该基因表达整体呈下降趋势,推测与ABA信号转导通路冗余性相关。
Sugarcane is the most important plant for sugar, and cane sugar occupies over90%of sugar production in China. However, over80%of the crop is planted in the hilly unpland area, where the rainfall is uneven and irrigation is not available, and drought has become one of the major limiting factors for sugarcane production in China. Breeding sugarcane varieties with strong drought tolerance and high water use efficiency is one of the most important strategies for sustainable development of the sugar industry. Due to the complex genetic background of sugarcane, however, crop modification through conventional hybrid breeding program is time costly and lowly efficient, other supplementary approaches including chemical regulation are necessary for improving the sugarcane drought resistance in practice. The aims of this research were to compare the physiological changes in leaves between the treatments of drought and drought plus foliar application of abscisic acid (ABA) under water stress conditions using sugarcane variety GT21, furthermore, to sieve the ABA responsive genes with microarray, and to clone the key gene involved in ABA biosynthesis and signal transduction.
1. An experiment was set up to investigate the interrelationship between drought induced ABA biosynthesis and antioxidative defense system, and to confer the farther role of foliar application of ABA in imparting drought tolerance to the sugarcane plant. The results showed that drought treatment enhanced the ABA concentration in leaf but it was significantly higher in combined ABA treatment, suggesting its biosynthesis was triggered in leaf by the ABA application. The relative water content was dramatically decreased by drought treatment but it could maintain higher in ABA treatment. Both drought and ABA treatment could result in an increase in proline, soluble protein, auxin, H2O2and malondialdehyde (MDA) content but the plants applied with ABA were found to resist the accumulation of MDA and gibberellin. ABA application decreased the degradation of chlorophyll, counteracted, at least in part, the decrease in maximal PS Ⅱ efficiency(Fv/Fm), antennae efficiency of PS Ⅱ (Fv/Fm) and quantum efficiency of PSⅡ (PS2). Overproduction of H2O2in plants treated with water stress was followed by higher activities of catalase (CAT), glutathione peroxidase (GPX), glutathione reductase (GR) and ascorbate peroxidase (APX), but this increase was further improved by the ABA treatment. The results clearly suggests that though the tolerant variety showed an enhanced protective system against drought conditions, the foliar application of ABA further improved its tolerance by continuously triggering the over expression of antioxidative defense system.
2. Degenerated and specific primers were designed from highly homologous regions of NCED gene sequences of different crop plants taken from NCBI database, a full-length cDNA encoding NCED gene was isolated and characterized from leaves of Saccharum officinarum L.(So) using RT-PCR and RACE-PCR. The isolated full-length SoNCED (2521bp) with1827bp ORF, encoded a peptide of608amino acids. The calculated molecular weight of the protein was65.9kDa with isoelectric point of6.04. Conserved domains prediction indicated a chloroplast-targeting peptide located at N-terminus of SoNCED protein. Phylogenetic tree, constructed by Neighbor-joining method indicated that deduced protein sequences of SoNCED shared high identity with the NCEDs reported from Zea mays and Sorghum bicolor which all belong to the C4plants. Sequence alignment revealed that the basic secondary structure including a-helices,[3-strands, P-propeller and His residues coordinating catalytic sites of SoNCED were highly conserved as in the NCEDs from other plants. Tissue specific expression analysis using Quantitative real-time PCR (qRT-PCR) showed a significant increase in SoNCED mRNA level and its correlation with O2-production rate and ABA accumulation in leaves and roots of sugarcane variety GT21when exposed to water stress. Further, the stimulation of SoNCED mRNA level, O2-production rate and ABA content after exogenous application of ABA proved its involvement in pathways providing tolerance to drought stress.
3. An Agilent4×44K microarray was applied to investigate the gene differential expression under water stress and foliar application ABA. The results showed that the number of differentially expressed genes was higher in the drought plus ABA treatment than drought treatment only. Large part of the genes was first induced then depressed following the aggravated water stress in both treatments. Total231genes were identified as ABA responsive from ABA treated samples at3rd day. Compared to the samples taken at5th day,71of the aforementioned genes were specifically expressed at3rd day in ABA treated plants,72genes were co-expressed in all treatments and days,38genes were co-expressed in ABA treatment, and50genes were specifically expressed at5th day in drought treated plants. Annotation results done by BlastX showed that173of the231genes could be validated. Function categories indicated that the hypothetical proteins have taken up about10%of the annotated genes, while genes functioned in lipid metabolism, signal transduction, protein kinase and osmoregulation have taken up about9%separately. Genes related to protein and hormone metabolism have taken up about6%separately.5%of the genes are related to photosynthesis, carbohydrate metabolism and transport separately. The remaining genes take part in respiration, transcription, oxidative stress, nitrogen metabolism, cellular metabolism and so on. The differential expression patterns of nine candidate genes were chose confirmed by qRT-PCR and the results showed that the expression changes of these candidates were generally consistent with the microarray results.
4. After annotation of the ABA responsive genes, one with highly homologous to the sucrose non-fermenting related protein kinase (SnRK2) gene SAPK3in Oryza sativa was selected for further analysis, and the cDNA full-length of this gene was amplified using RACE-PCR and named SoSnRK2.1. The isolated full-length of SoSnRK2.1was1385bp with1002bp ORF, encoding a peptide of333amino acids. Sequence analysis indicated that SoSnRK2.1showed high similarities with SnRK2from other plant species, and belong to the first subfamily when clustered with the SnRK2s taken from Arabidopsis thaliana using a Neighbor-joining method. The calculated molecular weight of the protein was37.8kDa with isoelectric point of5.54, and predicted as a hydrophilic protein without transmenbrane domains. There was a protein kinase conservative catalytic domain between the5-261amino acid sites, with11,3and8potential phosphorylation sites of serine, threonine and tyrosine separately, and the loop structure have taken the most part in the second structure prediction of SoSnRK2.1. Tissue specific expression analysis using qRT-PCR showed a significant increase in SoSnRK2.1mRNA level induced by foliar application of ABA and water stress, and correlated with ABA accumulation in leaves, which indicated the signal transduction function of this gene in leaves, while the expressions were suppressed with the accumulation of ABA content in roots, and this might be due to the redundancy of SnRK2genes in plants, but should be for further research.
引文
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