龙眼种质资源遗传多样性分析及低温对石硖龙眼影响研究
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摘要
中国是龙眼主要起源地之一,存在许多性状优良的野生和实生资源,虽然目前对龙眼资源的遗传多样性有较多的研究报道,但主要集中于国内栽培品种之间,对国外品种及国内龙眼非栽培资源研究不多。为此,本研究收集了大量野生和实生的龙眼资源,分析其遗传多样性,探明其亲缘关系,为龙眼资源的开发利用提供依据。近年来,低温寒冻害造成龙眼大面积死亡的现象时有发生,给龙眼生产带来巨大损失。石硖龙眼是我国的主栽品种之一,但低温对石硖龙眼影响的研究深度不够。为了更进一步探讨低温对石硖龙眼的影响,本研究从生理生化和分子水平探讨低温胁迫对石硖龙眼的影响,为龙眼栽培和抗寒育种提供依据。主要研究结论如下：
1、龙眼资源遗传多样性分析
利用SCoT和ISSR两种分子标记技术对国内3个省区及越南、泰国的龙眼种质资源的遗传多样性进行了分析,发现非栽培资源与栽培品种之间的遗传距离较远,中国龙眼基因资源丰富,泰国和越南的基因型龙眼分别聚类于2个不同的遗传类且遗传多样性较低。龙荔和龙眼之间的亲缘关系较远。
2、低温胁迫对龙眼生理生化特性的影响
以龙眼幼苗为材料,研究低温胁迫对龙眼生理生化指标的影响。在低温胁迫期间,各保护酶类、脯氨酸和可溶性蛋白含量呈先上升再下降的趋势；渗透物质可溶性糖含量一直呈上升趋势；相对含水量下降,电导率在达到最大值后急剧下降,MDA含量在处理前期处于较高水平；低温胁迫8小时前,对光合作用的影响以气孔限制为主,之后以非气孔限制为主,上述所测定的指标与龙眼低温胁迫呈显著或极显著相关。经主成分分析表明,前3个主成分包括的抗寒评价指标(保护酶类、膜透性和叶绿素荧光参数)可以进行抗寒性分析,可作为龙眼抗寒性评价的依据。另外,通过综合分析表明,4℃低温处理12小时内,可作为龙眼幼苗抗寒锻炼的适宜条件。
3、龙眼响应低温胁迫的蛋白质组学分析
建立了龙眼叶片总蛋白双向电泳体系,从蛋白质水平研究龙眼幼苗对低温的响应机制。从2-DE图谱获得了700多个有效蛋白质点,表达量差异在2倍以上有45个差异蛋白点,其中有36.7%的蛋白上调表达,63.3%的蛋白被下调表达。质谱鉴定出31个蛋白,它们主要是参与光合作用、自由基清除、细胞生长和分裂、信号传导、蛋白加工和基础代谢的蛋白。
4、抗寒相关基因的克隆与表达分析
应用RT-PCR和RACE方法克隆了蛋白质组学分析中的光合放氧33kD蛋白、锌指蛋白、抗坏血酸过氧化物酶、景天庚酮-1,7-二磷酸酶、Rubisco活化酶、泛素蛋白、磷酸核酮糖激酶、谷氨酰胺合成酶、热激蛋白、胚胎发育晚期丰富蛋白、14-3-3蛋白、叶绿体铜锌超氧化物歧化酶、咖啡酰辅酶A甲基转移酶、碳酸酐酶、水通道蛋白、钙依赖型蛋白激酶蛋白基因cDNA全长,并同源克隆了叶绿素a/b结合蛋白1、叶绿素a/b结合蛋白2、叶绿素a/b结合蛋白4和CBF基因cDNA全长。其中10个基因成功在大肠杆菌中表达,获得相应外源蛋白,并对热激蛋白经Western-blot验证确认。利用实时荧光定量技术对获得的基因进行了低温胁迫下时空表达模式分析,结果显示所有的基因在低温下均发生变化,但各基因的变化规律不同,如磷酸核酮糖激酶、泛素蛋白、Rubisco活化酶、谷氨酰胺合成酶、14-3-3蛋白、光合放氧33kD蛋白、水通道蛋白、叶绿素a/b结合蛋白、碳酸酐酶等基因在低温胁迫前期的变化较大,可能与各基因参与低温胁迫时的功能有关。已获得的cDNA全长中有43.8%在mRNA水平和蛋白水平表达相一致,56.2%的基因在转录水平上表达并没有直接伴随在蛋白水平的表达。
5、抗寒相关基因的转基因初步研究
根据已获得的龙眼CBF、热激蛋白和泛素蛋白的ORF序列设计真核表达引物,构建三个基因的正反义真核表达载体,转化农杆菌EHA105,采用农杆菌介导法转入野生型烟草植株,获得部分转基因烟草,为进一步研究龙眼抗寒基因的功能奠定了基础。
Dimocarpus longana Lour. belongs to Family Sapindaceae. China is one of the main origin centres, and there are many wild and seeding longan germplasm resources. In the last decade, molecular markers in characterizing and analyzing genetic diversity of longan has been successfully demonstrated by a number of studies. While these studies focused on local varieties of longan and the foreign cultivars or wild resources were rarely studied. In this paper, more resources with different genetic background were collected to study the diversity and genetic relationships by ISSR and SCoT markers. In recent years, cold and chilling injury resulted in serious economic losses in longan industry.'Shixia' longan is a main cultivars in China. However, there is a lack of depth in the academic research on the effect of low temperature stress on'Shixia' Longan currently. In order to understand furtherly cold-responsive mechanisms of'Shixia' Longan, the cold-responsive physiology and molecular mechanism of'Shixia'Longan under low temperature stress were studied. The results are benefit for the cultivation and breeding of longan. The main results obtained were as follows:
1. Genetic diversity analysis of longan. The genetic relationships among68longan cultivars from China, Vietnam and Thailand were analyzed based on SCoT and ISSR. The results showed that the genetic similarity coefficient was big difference among cultivars and non-cultivars. The UPGMA analysis indicated that all of them were distinguished in China, Vietnam and Thailand, respectively. In addition, there were obvious differences between Dimocarpus confinis and longan. Comparison and mixture analysis of the two molecular markers demonstrated that both SCoT and ISSR are efficient approaches for genetic diversity analysis of longan germplasm.
2. The physiological and biochemical characteristics of longan seedlings under low temperature stress. The physiological and biochemical indexes related to chill resistance in the leaves of longan were investigated under4℃low temperatures compared with the normal temperature25℃(control). The results showed that there were significant or obviously significant correlations between every indexe under chill tolerance of longan. The changes of enzyme activity increased early and then decreased, the soluble sugar content overall assumed the trend of escalation, The electrolyte leakage and the soluble protein content increased early and than decreased, water content of leaves continuous increased and the MDA content was at a high level in the early treatment under low temperature stress. The photosynthesis was affected mainly by stomatal limitation during0～8h under cold stress, and photosynthesis was affected mainly by non-stomatal limitation after8h cold stress. The relevance of each indicator and principal component analysis showed that the first three principal components including the protective enzyme, membrane permeability, Chl Ⅱ fluorescence characteristics can be synthesis evaluation for cold stress of longan. the comprehensive analysis of all physiological and biochemical indicators showed that cold stress for12h at4℃is suitable condition for cold resistant exercise of longan seedlings.
3. Proteomic analysis of longan seedlings under low temperature stress. The proteome changes in one-year-old longan tree leaves subjected to a temperature of4℃for24h were studied with comparative proteomic analysis. Changes in the total proteins of longan leaves were examined by two-dimensional electropherosis and mass spectroscopy. More than700protein spots were detected on each gel, and45protein spots which expression difference were more than two times were obtained by PDQuest software analysis. Among45protein spots63.3%protein spots were down-regulated, and36.7%protein spots were up-regulated.31differentially expressed proteins were identified by Mass spectrometry, They couid participate in photosynthesis, free radical scavenging, cell growth/division, signal transduction, protein processing, metabolism and unclassified.
4. Cold-responsive genes cloning and expression. The full-length cDNA sequences were cloned from longan leaves in the way of RT-PCR and RACE techniques. They were genes involving in Oxygen evolving enhancer protein, Phd/F-box containing protein, Ascorbate peroxidase, Sedoheptulose-1,7-bisphosphatase, Rubisco activase, SUMO1protein、Phosphoribulokinase, Glutamine synthetase, Heat shock protein HSP, Late embryogenesis abundant protein,14-3-3protein, Chloroplast copper/zinc-superoxide dismutase, Caffeoyl-CoA O-methyltransferase, Carbonic anhydrase, Aquaporin gene, Calcium-dependent protein kinase gene, Light Harvesting Chlorophyll a/b binding proteins, CRT/DREB Binding Factor. The ten genes out of them were expressed in E.coli, their exogenous proteins were obtained, and heat shock protein (HSP) was confirmed by western blotting. To investigate changes in genes expression at the mRNA levels, their expression patterns in response to chilling stress were studied with quantitative real time PCR. The results showed that gene expression level were changed under low temperature stress, but every gene expression trend is different. For example, the genes involving in Phosphoribulokinase, SUMO1protein,14-3-3protein, Oxygen evolving enhancer protein, etc. changed largely during early period under low temperature stress, and these changes may be related to the genes function under low temperature stress. But only43.8%genes exhibited consistent expression patterns between their mRNAs and protein levels at the same time, while56.2%genes expressed at mRNA levels and unexpressed at protein levels.
5. Genetic transformation of cold-responsive genes. The primers of CBF, HSP and SUMO genes of longan have been designed by integrity of ORF sequence, and binary antisense and sense expression vectors of three genes were constructed by inserting the target fragment in PBI121with a double35S promoter. The PBI121-CBF (-/+), PBI121-HSP (-/+) and PBI121-SUMO (-/+) were successfully introduced into strain EHA105by Agrobacterium-mediated and acquired some transgenic tobaccos plants. The results will lay the foundation for further studying of cold resistant genes character and their function of longan.
1、龙眼资源遗传多样性分析
利用SCoT和ISSR两种分子标记技术对国内3个省区及越南、泰国的龙眼种质资源的遗传多样性进行了分析,发现非栽培资源与栽培品种之间的遗传距离较远,中国龙眼基因资源丰富,泰国和越南的基因型龙眼分别聚类于2个不同的遗传类且遗传多样性较低。龙荔和龙眼之间的亲缘关系较远。
2、低温胁迫对龙眼生理生化特性的影响
以龙眼幼苗为材料,研究低温胁迫对龙眼生理生化指标的影响。在低温胁迫期间,各保护酶类、脯氨酸和可溶性蛋白含量呈先上升再下降的趋势；渗透物质可溶性糖含量一直呈上升趋势；相对含水量下降,电导率在达到最大值后急剧下降,MDA含量在处理前期处于较高水平；低温胁迫8小时前,对光合作用的影响以气孔限制为主,之后以非气孔限制为主,上述所测定的指标与龙眼低温胁迫呈显著或极显著相关。经主成分分析表明,前3个主成分包括的抗寒评价指标(保护酶类、膜透性和叶绿素荧光参数)可以进行抗寒性分析,可作为龙眼抗寒性评价的依据。另外,通过综合分析表明,4℃低温处理12小时内,可作为龙眼幼苗抗寒锻炼的适宜条件。
3、龙眼响应低温胁迫的蛋白质组学分析
建立了龙眼叶片总蛋白双向电泳体系,从蛋白质水平研究龙眼幼苗对低温的响应机制。从2-DE图谱获得了700多个有效蛋白质点,表达量差异在2倍以上有45个差异蛋白点,其中有36.7%的蛋白上调表达,63.3%的蛋白被下调表达。质谱鉴定出31个蛋白,它们主要是参与光合作用、自由基清除、细胞生长和分裂、信号传导、蛋白加工和基础代谢的蛋白。
4、抗寒相关基因的克隆与表达分析
应用RT-PCR和RACE方法克隆了蛋白质组学分析中的光合放氧33kD蛋白、锌指蛋白、抗坏血酸过氧化物酶、景天庚酮-1,7-二磷酸酶、Rubisco活化酶、泛素蛋白、磷酸核酮糖激酶、谷氨酰胺合成酶、热激蛋白、胚胎发育晚期丰富蛋白、14-3-3蛋白、叶绿体铜锌超氧化物歧化酶、咖啡酰辅酶A甲基转移酶、碳酸酐酶、水通道蛋白、钙依赖型蛋白激酶蛋白基因cDNA全长,并同源克隆了叶绿素a/b结合蛋白1、叶绿素a/b结合蛋白2、叶绿素a/b结合蛋白4和CBF基因cDNA全长。其中10个基因成功在大肠杆菌中表达,获得相应外源蛋白,并对热激蛋白经Western-blot验证确认。利用实时荧光定量技术对获得的基因进行了低温胁迫下时空表达模式分析,结果显示所有的基因在低温下均发生变化,但各基因的变化规律不同,如磷酸核酮糖激酶、泛素蛋白、Rubisco活化酶、谷氨酰胺合成酶、14-3-3蛋白、光合放氧33kD蛋白、水通道蛋白、叶绿素a/b结合蛋白、碳酸酐酶等基因在低温胁迫前期的变化较大,可能与各基因参与低温胁迫时的功能有关。已获得的cDNA全长中有43.8%在mRNA水平和蛋白水平表达相一致,56.2%的基因在转录水平上表达并没有直接伴随在蛋白水平的表达。
5、抗寒相关基因的转基因初步研究
根据已获得的龙眼CBF、热激蛋白和泛素蛋白的ORF序列设计真核表达引物,构建三个基因的正反义真核表达载体,转化农杆菌EHA105,采用农杆菌介导法转入野生型烟草植株,获得部分转基因烟草,为进一步研究龙眼抗寒基因的功能奠定了基础。
Dimocarpus longana Lour. belongs to Family Sapindaceae. China is one of the main origin centres, and there are many wild and seeding longan germplasm resources. In the last decade, molecular markers in characterizing and analyzing genetic diversity of longan has been successfully demonstrated by a number of studies. While these studies focused on local varieties of longan and the foreign cultivars or wild resources were rarely studied. In this paper, more resources with different genetic background were collected to study the diversity and genetic relationships by ISSR and SCoT markers. In recent years, cold and chilling injury resulted in serious economic losses in longan industry.'Shixia' longan is a main cultivars in China. However, there is a lack of depth in the academic research on the effect of low temperature stress on'Shixia' Longan currently. In order to understand furtherly cold-responsive mechanisms of'Shixia' Longan, the cold-responsive physiology and molecular mechanism of'Shixia'Longan under low temperature stress were studied. The results are benefit for the cultivation and breeding of longan. The main results obtained were as follows:
1. Genetic diversity analysis of longan. The genetic relationships among68longan cultivars from China, Vietnam and Thailand were analyzed based on SCoT and ISSR. The results showed that the genetic similarity coefficient was big difference among cultivars and non-cultivars. The UPGMA analysis indicated that all of them were distinguished in China, Vietnam and Thailand, respectively. In addition, there were obvious differences between Dimocarpus confinis and longan. Comparison and mixture analysis of the two molecular markers demonstrated that both SCoT and ISSR are efficient approaches for genetic diversity analysis of longan germplasm.
2. The physiological and biochemical characteristics of longan seedlings under low temperature stress. The physiological and biochemical indexes related to chill resistance in the leaves of longan were investigated under4℃low temperatures compared with the normal temperature25℃(control). The results showed that there were significant or obviously significant correlations between every indexe under chill tolerance of longan. The changes of enzyme activity increased early and then decreased, the soluble sugar content overall assumed the trend of escalation, The electrolyte leakage and the soluble protein content increased early and than decreased, water content of leaves continuous increased and the MDA content was at a high level in the early treatment under low temperature stress. The photosynthesis was affected mainly by stomatal limitation during0～8h under cold stress, and photosynthesis was affected mainly by non-stomatal limitation after8h cold stress. The relevance of each indicator and principal component analysis showed that the first three principal components including the protective enzyme, membrane permeability, Chl Ⅱ fluorescence characteristics can be synthesis evaluation for cold stress of longan. the comprehensive analysis of all physiological and biochemical indicators showed that cold stress for12h at4℃is suitable condition for cold resistant exercise of longan seedlings.
3. Proteomic analysis of longan seedlings under low temperature stress. The proteome changes in one-year-old longan tree leaves subjected to a temperature of4℃for24h were studied with comparative proteomic analysis. Changes in the total proteins of longan leaves were examined by two-dimensional electropherosis and mass spectroscopy. More than700protein spots were detected on each gel, and45protein spots which expression difference were more than two times were obtained by PDQuest software analysis. Among45protein spots63.3%protein spots were down-regulated, and36.7%protein spots were up-regulated.31differentially expressed proteins were identified by Mass spectrometry, They couid participate in photosynthesis, free radical scavenging, cell growth/division, signal transduction, protein processing, metabolism and unclassified.
4. Cold-responsive genes cloning and expression. The full-length cDNA sequences were cloned from longan leaves in the way of RT-PCR and RACE techniques. They were genes involving in Oxygen evolving enhancer protein, Phd/F-box containing protein, Ascorbate peroxidase, Sedoheptulose-1,7-bisphosphatase, Rubisco activase, SUMO1protein、Phosphoribulokinase, Glutamine synthetase, Heat shock protein HSP, Late embryogenesis abundant protein,14-3-3protein, Chloroplast copper/zinc-superoxide dismutase, Caffeoyl-CoA O-methyltransferase, Carbonic anhydrase, Aquaporin gene, Calcium-dependent protein kinase gene, Light Harvesting Chlorophyll a/b binding proteins, CRT/DREB Binding Factor. The ten genes out of them were expressed in E.coli, their exogenous proteins were obtained, and heat shock protein (HSP) was confirmed by western blotting. To investigate changes in genes expression at the mRNA levels, their expression patterns in response to chilling stress were studied with quantitative real time PCR. The results showed that gene expression level were changed under low temperature stress, but every gene expression trend is different. For example, the genes involving in Phosphoribulokinase, SUMO1protein,14-3-3protein, Oxygen evolving enhancer protein, etc. changed largely during early period under low temperature stress, and these changes may be related to the genes function under low temperature stress. But only43.8%genes exhibited consistent expression patterns between their mRNAs and protein levels at the same time, while56.2%genes expressed at mRNA levels and unexpressed at protein levels.
5. Genetic transformation of cold-responsive genes. The primers of CBF, HSP and SUMO genes of longan have been designed by integrity of ORF sequence, and binary antisense and sense expression vectors of three genes were constructed by inserting the target fragment in PBI121with a double35S promoter. The PBI121-CBF (-/+), PBI121-HSP (-/+) and PBI121-SUMO (-/+) were successfully introduced into strain EHA105by Agrobacterium-mediated and acquired some transgenic tobaccos plants. The results will lay the foundation for further studying of cold resistant genes character and their function of longan.
引文
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