菊花DREB同源基因的克隆和功能分析
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摘要
菊花(Dendranthema×morifolium)为菊科菊属宿根草本花卉,该属各种主要分布于亚洲温带和亚热带部分地区,观赏价值极高,位居世界四大切花之首,在人们生活和花卉生产中占有非常重要的地位.菊花具有良好的耐逆性具有很强的生命力和适应性,研究和探讨菊花的耐逆分子机制具有非常重要的意义.
各种环境因素,如干旱、高盐、激素和低/高温等非生物胁迫对植物的生长发育造成很大影响.DREB转录因子是一类在植物抵抗非生物胁迫中起到关键作用的转录因子家族.DREB转录因子特异结合DRE/CRT顺式作用元件,激活一系列靶基因的表达.这些靶基因编码脯氨酸、蔗糖、LEA蛋白、AQP蛋白以及果糖合成酶等.在转基因拟南芥中DREB1A的超表达将激活至少40个逆境应答基因的表达,增强植物对逆境的忍耐力.一个转录因子可同时激活多个功能基因的表达,导入或改良一个关键的转录因子,促使多个功能基因发挥作用,从而提高植株的综合品质,因此转录因子在植物分子育种中具有巨大的潜在价值和广阔的应用前景.本研究内容主要包括:一、从菊花中克隆了两个与低温、高盐等非生物胁迫相关的DREB家族转录因子同源基因,并对它们的功能进行了初步研究;二、利用12份野生菊和5份栽培菊材料进行了DmDREBb基因的核苷酸多态性的初步分析.
本研究利用RACE-PCR的方法从耐寒性强的栽培菊花‘红枫'中分离克隆了两个DREB同源基因,分别命名为DmDREBa和DmDREBb.研究表明,DmDREBa和DmDREBb基因全长分别为872bp和675bp,分别编码192和185个氨基酸的完整阅读框,推测两个蛋白的分子量分别为21.66和20.99 kDa,等电点分别为6.72和6.80.两个基因提交NCBI,登录号分别为EF490996和EF487535.以相同的引物对菊花基因组DNA进行了扩增,得到了DmDREBs基因组序列,大小与cDNA序列一致.表明该基因中不含有内含子。经过Alighment比对,证实两个基因与已经报道的其他作物的DREB基因有着很高的相似性,并且和DREB1类基因有着更高的相似性.BLAST比较分析发现,除了个别氨基酸差异之外,DmDREBa比DmDREBb还明显多了七个“MPGVIDS”氨基酸残基(153—159).DmDREBa与DmDREBb在氨基酸水平上的一致性达到88%,核苷酸水平上为90%。另外在二级结构预测中发现两个蛋白均含有三个β-折叠和一个a-螺旋,含有一个AP2结构域,并且也具有此结构域两端的标签序列“PKKRAGGKKFMETRHP”和“DSAWR”.这些都符合DREB1亚家族的特征.系统进化树分析也表明DmDREBa和DmDREBb与DREB1亚家族的相似性比较高,与辣椒和葡萄等木本的双子叶植物距离比较近.Southern杂交表明,在‘红枫'中存在着DmDREBa和DmDREBb基因的多个拷贝或者同源基因.组织表达谱分析发现,DmDREBa和DmDREBb基因在茎、叶中的表达量比较高,而根和花中则表达量低.在各种非生物胁迫处理下,DmDREBa基因更能响应高盐的诱导,而DmDREBb基因则是在4℃低温处理下较早的强烈表达.DmDREBa和DmDREBb基因响应PEG6000模拟的干旱和40℃高温胁迫时候,表达被强烈抑制.酵母单杂交实验证实DmDREBa和DmDREBb具有与DRE顺式作用元件相结合的能力,并且具有转录激活活性.亚细胞定位研究也显示两者都定位到了细胞核内,这也表明两个基因编码的蛋白是转录因子.
为了进一步了解DmDREBs基因的功能,我们对DmDREBa基因进行了烟草转化的研究.研究利用PCR的方法鉴定出了43株转基因阳性植株.随机选取10株转基因植株有7椿雀RNA转录水平能够表达.Southern杂交检测表明,目的基因已经整合到烟草的基因组中,并且以1-3个拷贝形式随机插入.另外,研究将DmDREBa基因过量表达的烟草在4℃低温、干旱和高盐胁迫后,发现转基因阳性植株的抗性比野生型烟草都有所增强.丙二醛(MDA)测定后,发现导入35S:DmDREBa基因的烟草丙二醛含量为0.481×10~(-3)μmol/g,低于野生型烟草的丙二醛含量(0.619×10~(-3)μmol/g),也说明转基因烟草提高了抗低温的能力.失水率分析表明,转基因阳性植株蒸腾失水量低于对照.随之对叶片的表皮气孔观察发现,转基因阳性植株上下表皮的气孔密度与对照相比有所降低,经t值测验,转基因烟草和野生型烟草的上、下表皮气孔数的差异均达到显著水平.经过低温、干旱以及高盐的胁迫处理后,转基因阳性植株较对照而言都表现出良好的抗性.
利用DmDREBb基因对12份野生菊和5份栽培菊进行了SNP的分析.结果发现,研究中野生菊中SNP的主要突变形式为颠换,而栽培菊中则是转换.野生菊的核苷酸多样性(π=0.02368)高于栽培菊(π=0.00963).表明栽培菊在进化过程中受到了强大的人工选择的作用.DmDREBb基因编码区内Ka/Ks远远小于1,说明DmDREBb基因序列十分保守,现代菊花在进化的过程中受到了净化选择的作用。LD分析发现LD强度随着距离的延伸而逐渐减弱,野生菊减弱趋势比栽培菊明显,栽培菊减弱趋势比较平缓.从野生菊到栽培菊LD强度大大提高.说明,菊花在栽培驯化过程中LD结构发生了变化,LD结构的变化与驯化过程中有遗传多样性的丢失.
Chrysanthemum(Dendranthema×morifolium Ramat.) is one type of herbage plants in chrysanthemum genus of composite.With important ornamental value,chrysanthemum grows in temperate zone and subtropics in Asia.Chrysanthemum is the fist one in the four dominating cut flowers in the world and it is very important in the people's life and the production of flowers.Moreover,with strong vitality and adaptability,chrysanthemum has good tolerance to abiotic stresses.Therefore,the study on molecular mechanism about stresses in chrysanthemum is valuable.
Plants are affected by many abiotic stresses,such as drought,high salinity,low/high temperature and hormone and so on,in the process of growth and development.DREB transcription factors play the key roles in response to abiotic stresses.DREB transcription factors specifically bind the DRE/CRT cis-acting element and trigger the downstream target genes.The coding products of target genes were functional proteins and enzymes such as proline,sucrose,LEA,aquaporin(AQP) and praline synthetase et al.In transgenic Arabidopsis plants,overexpression of DREB1A can introduce at least 40 downstream genes and increase the tolerance to abiotic stresses.One transcription factor introduces many downstream genes and improves the resistance activity to stresses.It had very large potential value and widest foreground to study transcription factors in plant breeding.The thesis aims included:firstly,two DREB-like genes were isolated from chrysanthemum and the relate function were studied;secondly,some information was obtained by the SNP analysis in chrysanthemum using DmDREBb gene.
To analysis the molecular mechanism of DREB in chrysanthemum,two DREB-like genes were isolated from chrysanthemum cv.'Hong Feng' using degenerate PCR and RACE methods in this study.These cDNAs were 872bp and 675bp in length containing a single open reading frame encoding 191,185 amino acids with predicted molecular masses of 21.66 and 20.99 kDa,and the pI were 6.72 and 6.80,respectively.Both sequences had been deposited to the GenBank database under accession numbers EF490996 and EF487535 for DmDREBa and DmDREBb,respectively.The genome sequences were obtained by the same primers with cDNAs and the lengths of both genes were also same with cDNA sequences.The results indicated that there was no intron in the two chrysanthemum DREB-like genes.The sequence alignment analysis showed that DmDREBs genes had high similarity with other DREB genes and had higher similarity with DREB1 genes.Blast analysis suggested high similarity between DmDREBa and DmDREBb with the overall identities 88%at amino acid level and 90%at nucleotide acids except for one obvious gap of MPGVIDS(152—159) in amino acids level.The DmDREBs proteins contained a conserved DNA-binding domain of 57 amino acids that presented in a large family of plants.DmDREBs genes revealed a typical DREB1-type nuclear localization signal(NLS) consensus PKKRAGGKKFMETRHP before the EREBP/AP2 domain and a DSAWR sequence after the domain,and that domain existed in all DREB1-type proteins from various species.Threeβ-sheets and an amphipathic a-helix were also found in the EREBP/AP2 domain of DmDREBa and DmDREBb,which may be responsible for DNA-binding activity.Thus,sequence analysis indicated that DmDREBa and DmDREBb were putative DREB1 homologs.Phylogenic analysis suggested that DmDREBs were attributable to the DREB1 subgroups comparing to DREB2 transcription factors. Interestingly,DmDREBs shared the highest similar to Capsicum annuum CaCBF1A (AY368482) and Vitis vinifera VvCBF1(AY390372)at amino acid level,two species that are chilling sensitive.Furthermore,Vitis vinifera is one of woody plants.Southern blot analysis was used to determine the approximate number of the DREB-like genes in chrysanthemum.Probing with the DmDREBa sequence indicated that chrysanthemum contains multiple DREB-like genes/alleles.To investigate the expression patterns of DmDREBs in chrysanthemum,the expression of the two genes at different organs was analyzed.Both genes were highly expressed in leaves and stems,but with low expression in roots or flowers at 15℃.The expression levels of DmDREBa and DmDREBb were different in several stresses.Under cold stress(4℃),DmDREBb was expressed highly at 0.5h,but DmDREBa was highly expressed in 24h under salt condition.The expression level of DmDREBa was also reduced during the whole process but a little expression was observed after 24h with 100μM ABA treatment.However,the expression level of DmDREBb was variable under 100μM ABA condition.The mRNA level of DmDREBa was inhibited significantly under PEG6000 and hot condition(40℃).The expression of DmDREBb was continued only 0.5h,and then was also inhibited strongly.These two proteins were found to have transcriptional activity and had the DRE-binding capacity as shown by yeast one-hybrid system and were localized to the nucleus of cells.These studies indicated that the two proteins were DREB transcription factors.
The DmDREBa gent was overexpressed in tobacco and 43 transgenic plants were obtained by PCR method.And southern blot analysis indicated that 1-3 copies of DmDREBa were randomly inserted.The tolerance of transgenic tobacco plants was increased comparing to the wild tobacco plants under low temperature,drought and high salinity conditions.The content of MDA in wild tobacco plants was 0.619×10~(-3)μmol/g which was more than 0.481×10~(-3)μmol/g in transgenic plants,which indicated the tolerance ability to low temperature of transgenic plants was increased.The lost water of transgenic tobacco was less than wild plants after natural transpiration.Further,the density of the stoma in the epidermis was observed and found that the numbers of stoma in upper or lower epidermis of transgenic plants were reduced comparing to wild tobacco.The result of t test suggested that the differences between the upper and lower epidermis of transgenic plants and wild type were significant.The transgenic tobaccos had better character than the wild plants under low temperature,drought and high salinity conditions.
At last the SNP research was carried out in chrysanthemum cultivars and wild plants using DmDREBb gene.The main mutations were transversions in chrysanthemum wild plants with more mutations of transitions in chrysanthemum cultivars.The results showed that the diversity of SNP in wild plants(π=0.02368) was higher than that in chrysanthemum cultivars(π=0.00963).The results indicated that artificial selection affected the evolution of chrysanthemum cultivars.The value of Ka/Ks was less than 1 suggested that the nucleotide acids sequence was very conservative and purifying selection affected the evolution of chrysanthemum cultivars.The analysis of LD was suggested that the strength of LD was decreased with the extrending of distance.The decreased trend of chrysanthemum wild plants was very evidence,but the trend was gentle in chrysanthemum cultivars.The structure of LD was changed during the domestication of chrysanthemum cultivars that affected diversity drop.
各种环境因素,如干旱、高盐、激素和低/高温等非生物胁迫对植物的生长发育造成很大影响.DREB转录因子是一类在植物抵抗非生物胁迫中起到关键作用的转录因子家族.DREB转录因子特异结合DRE/CRT顺式作用元件,激活一系列靶基因的表达.这些靶基因编码脯氨酸、蔗糖、LEA蛋白、AQP蛋白以及果糖合成酶等.在转基因拟南芥中DREB1A的超表达将激活至少40个逆境应答基因的表达,增强植物对逆境的忍耐力.一个转录因子可同时激活多个功能基因的表达,导入或改良一个关键的转录因子,促使多个功能基因发挥作用,从而提高植株的综合品质,因此转录因子在植物分子育种中具有巨大的潜在价值和广阔的应用前景.本研究内容主要包括:一、从菊花中克隆了两个与低温、高盐等非生物胁迫相关的DREB家族转录因子同源基因,并对它们的功能进行了初步研究;二、利用12份野生菊和5份栽培菊材料进行了DmDREBb基因的核苷酸多态性的初步分析.
本研究利用RACE-PCR的方法从耐寒性强的栽培菊花‘红枫'中分离克隆了两个DREB同源基因,分别命名为DmDREBa和DmDREBb.研究表明,DmDREBa和DmDREBb基因全长分别为872bp和675bp,分别编码192和185个氨基酸的完整阅读框,推测两个蛋白的分子量分别为21.66和20.99 kDa,等电点分别为6.72和6.80.两个基因提交NCBI,登录号分别为EF490996和EF487535.以相同的引物对菊花基因组DNA进行了扩增,得到了DmDREBs基因组序列,大小与cDNA序列一致.表明该基因中不含有内含子。经过Alighment比对,证实两个基因与已经报道的其他作物的DREB基因有着很高的相似性,并且和DREB1类基因有着更高的相似性.BLAST比较分析发现,除了个别氨基酸差异之外,DmDREBa比DmDREBb还明显多了七个“MPGVIDS”氨基酸残基(153—159).DmDREBa与DmDREBb在氨基酸水平上的一致性达到88%,核苷酸水平上为90%。另外在二级结构预测中发现两个蛋白均含有三个β-折叠和一个a-螺旋,含有一个AP2结构域,并且也具有此结构域两端的标签序列“PKKRAGGKKFMETRHP”和“DSAWR”.这些都符合DREB1亚家族的特征.系统进化树分析也表明DmDREBa和DmDREBb与DREB1亚家族的相似性比较高,与辣椒和葡萄等木本的双子叶植物距离比较近.Southern杂交表明,在‘红枫'中存在着DmDREBa和DmDREBb基因的多个拷贝或者同源基因.组织表达谱分析发现,DmDREBa和DmDREBb基因在茎、叶中的表达量比较高,而根和花中则表达量低.在各种非生物胁迫处理下,DmDREBa基因更能响应高盐的诱导,而DmDREBb基因则是在4℃低温处理下较早的强烈表达.DmDREBa和DmDREBb基因响应PEG6000模拟的干旱和40℃高温胁迫时候,表达被强烈抑制.酵母单杂交实验证实DmDREBa和DmDREBb具有与DRE顺式作用元件相结合的能力,并且具有转录激活活性.亚细胞定位研究也显示两者都定位到了细胞核内,这也表明两个基因编码的蛋白是转录因子.
为了进一步了解DmDREBs基因的功能,我们对DmDREBa基因进行了烟草转化的研究.研究利用PCR的方法鉴定出了43株转基因阳性植株.随机选取10株转基因植株有7椿雀RNA转录水平能够表达.Southern杂交检测表明,目的基因已经整合到烟草的基因组中,并且以1-3个拷贝形式随机插入.另外,研究将DmDREBa基因过量表达的烟草在4℃低温、干旱和高盐胁迫后,发现转基因阳性植株的抗性比野生型烟草都有所增强.丙二醛(MDA)测定后,发现导入35S:DmDREBa基因的烟草丙二醛含量为0.481×10~(-3)μmol/g,低于野生型烟草的丙二醛含量(0.619×10~(-3)μmol/g),也说明转基因烟草提高了抗低温的能力.失水率分析表明,转基因阳性植株蒸腾失水量低于对照.随之对叶片的表皮气孔观察发现,转基因阳性植株上下表皮的气孔密度与对照相比有所降低,经t值测验,转基因烟草和野生型烟草的上、下表皮气孔数的差异均达到显著水平.经过低温、干旱以及高盐的胁迫处理后,转基因阳性植株较对照而言都表现出良好的抗性.
利用DmDREBb基因对12份野生菊和5份栽培菊进行了SNP的分析.结果发现,研究中野生菊中SNP的主要突变形式为颠换,而栽培菊中则是转换.野生菊的核苷酸多样性(π=0.02368)高于栽培菊(π=0.00963).表明栽培菊在进化过程中受到了强大的人工选择的作用.DmDREBb基因编码区内Ka/Ks远远小于1,说明DmDREBb基因序列十分保守,现代菊花在进化的过程中受到了净化选择的作用。LD分析发现LD强度随着距离的延伸而逐渐减弱,野生菊减弱趋势比栽培菊明显,栽培菊减弱趋势比较平缓.从野生菊到栽培菊LD强度大大提高.说明,菊花在栽培驯化过程中LD结构发生了变化,LD结构的变化与驯化过程中有遗传多样性的丢失.
Chrysanthemum(Dendranthema×morifolium Ramat.) is one type of herbage plants in chrysanthemum genus of composite.With important ornamental value,chrysanthemum grows in temperate zone and subtropics in Asia.Chrysanthemum is the fist one in the four dominating cut flowers in the world and it is very important in the people's life and the production of flowers.Moreover,with strong vitality and adaptability,chrysanthemum has good tolerance to abiotic stresses.Therefore,the study on molecular mechanism about stresses in chrysanthemum is valuable.
Plants are affected by many abiotic stresses,such as drought,high salinity,low/high temperature and hormone and so on,in the process of growth and development.DREB transcription factors play the key roles in response to abiotic stresses.DREB transcription factors specifically bind the DRE/CRT cis-acting element and trigger the downstream target genes.The coding products of target genes were functional proteins and enzymes such as proline,sucrose,LEA,aquaporin(AQP) and praline synthetase et al.In transgenic Arabidopsis plants,overexpression of DREB1A can introduce at least 40 downstream genes and increase the tolerance to abiotic stresses.One transcription factor introduces many downstream genes and improves the resistance activity to stresses.It had very large potential value and widest foreground to study transcription factors in plant breeding.The thesis aims included:firstly,two DREB-like genes were isolated from chrysanthemum and the relate function were studied;secondly,some information was obtained by the SNP analysis in chrysanthemum using DmDREBb gene.
To analysis the molecular mechanism of DREB in chrysanthemum,two DREB-like genes were isolated from chrysanthemum cv.'Hong Feng' using degenerate PCR and RACE methods in this study.These cDNAs were 872bp and 675bp in length containing a single open reading frame encoding 191,185 amino acids with predicted molecular masses of 21.66 and 20.99 kDa,and the pI were 6.72 and 6.80,respectively.Both sequences had been deposited to the GenBank database under accession numbers EF490996 and EF487535 for DmDREBa and DmDREBb,respectively.The genome sequences were obtained by the same primers with cDNAs and the lengths of both genes were also same with cDNA sequences.The results indicated that there was no intron in the two chrysanthemum DREB-like genes.The sequence alignment analysis showed that DmDREBs genes had high similarity with other DREB genes and had higher similarity with DREB1 genes.Blast analysis suggested high similarity between DmDREBa and DmDREBb with the overall identities 88%at amino acid level and 90%at nucleotide acids except for one obvious gap of MPGVIDS(152—159) in amino acids level.The DmDREBs proteins contained a conserved DNA-binding domain of 57 amino acids that presented in a large family of plants.DmDREBs genes revealed a typical DREB1-type nuclear localization signal(NLS) consensus PKKRAGGKKFMETRHP before the EREBP/AP2 domain and a DSAWR sequence after the domain,and that domain existed in all DREB1-type proteins from various species.Threeβ-sheets and an amphipathic a-helix were also found in the EREBP/AP2 domain of DmDREBa and DmDREBb,which may be responsible for DNA-binding activity.Thus,sequence analysis indicated that DmDREBa and DmDREBb were putative DREB1 homologs.Phylogenic analysis suggested that DmDREBs were attributable to the DREB1 subgroups comparing to DREB2 transcription factors. Interestingly,DmDREBs shared the highest similar to Capsicum annuum CaCBF1A (AY368482) and Vitis vinifera VvCBF1(AY390372)at amino acid level,two species that are chilling sensitive.Furthermore,Vitis vinifera is one of woody plants.Southern blot analysis was used to determine the approximate number of the DREB-like genes in chrysanthemum.Probing with the DmDREBa sequence indicated that chrysanthemum contains multiple DREB-like genes/alleles.To investigate the expression patterns of DmDREBs in chrysanthemum,the expression of the two genes at different organs was analyzed.Both genes were highly expressed in leaves and stems,but with low expression in roots or flowers at 15℃.The expression levels of DmDREBa and DmDREBb were different in several stresses.Under cold stress(4℃),DmDREBb was expressed highly at 0.5h,but DmDREBa was highly expressed in 24h under salt condition.The expression level of DmDREBa was also reduced during the whole process but a little expression was observed after 24h with 100μM ABA treatment.However,the expression level of DmDREBb was variable under 100μM ABA condition.The mRNA level of DmDREBa was inhibited significantly under PEG6000 and hot condition(40℃).The expression of DmDREBb was continued only 0.5h,and then was also inhibited strongly.These two proteins were found to have transcriptional activity and had the DRE-binding capacity as shown by yeast one-hybrid system and were localized to the nucleus of cells.These studies indicated that the two proteins were DREB transcription factors.
The DmDREBa gent was overexpressed in tobacco and 43 transgenic plants were obtained by PCR method.And southern blot analysis indicated that 1-3 copies of DmDREBa were randomly inserted.The tolerance of transgenic tobacco plants was increased comparing to the wild tobacco plants under low temperature,drought and high salinity conditions.The content of MDA in wild tobacco plants was 0.619×10~(-3)μmol/g which was more than 0.481×10~(-3)μmol/g in transgenic plants,which indicated the tolerance ability to low temperature of transgenic plants was increased.The lost water of transgenic tobacco was less than wild plants after natural transpiration.Further,the density of the stoma in the epidermis was observed and found that the numbers of stoma in upper or lower epidermis of transgenic plants were reduced comparing to wild tobacco.The result of t test suggested that the differences between the upper and lower epidermis of transgenic plants and wild type were significant.The transgenic tobaccos had better character than the wild plants under low temperature,drought and high salinity conditions.
At last the SNP research was carried out in chrysanthemum cultivars and wild plants using DmDREBb gene.The main mutations were transversions in chrysanthemum wild plants with more mutations of transitions in chrysanthemum cultivars.The results showed that the diversity of SNP in wild plants(π=0.02368) was higher than that in chrysanthemum cultivars(π=0.00963).The results indicated that artificial selection affected the evolution of chrysanthemum cultivars.The value of Ka/Ks was less than 1 suggested that the nucleotide acids sequence was very conservative and purifying selection affected the evolution of chrysanthemum cultivars.The analysis of LD was suggested that the strength of LD was decreased with the extrending of distance.The decreased trend of chrysanthemum wild plants was very evidence,but the trend was gentle in chrysanthemum cultivars.The structure of LD was changed during the domestication of chrysanthemum cultivars that affected diversity drop.
引文
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