梅花脱水素基因家族的克隆与功能分析
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摘要
梅花(Prunus mume)属于蔷薇科李属植物,在东亚地区深受人们喜爱。梅花原产于长江流域,如何提高其抗逆能力,尤其是抗寒能力,对扩大梅花的应用范围十分重要。脱水素(又称第二类胚胎发育晚期丰富蛋白)在植物抵抗非生物胁迫中起重要作用。但目前对梅花脱水素家族基因的研究还很少。本研究首先在‘藏梅’全基因组测序的基础上,对梅花胚胎发育晚期丰富蛋白基因家族进行了生物信息分析；然后利用RT-PCR法对其中脱水素基因家族进行了克隆和表达分析；最后分别在大肠杆菌和烟草中超表达了梅花脱水素基因,并进行了初步的功能分析。主要结果如下：
1.首次预测了梅花胚胎发育晚期丰富蛋白基因家族成员：梅花中共有30个胚胎发育晚期丰富蛋白基因,分布于7个连锁群上。5个梅花胚胎发育晚期丰富蛋白基因属于片段重复,12个属于串联重复。根据序列相似性,可以将这些基因分为8组(LEA_1, LEA_2, LEA_3, LEA_4, LEA_5, PvLEA18,脱水素和SMP),其中脱水素家族含有6个成员(PmLEA8,PmLEA9, PmLEA10, PmLEA19,PmLEA20和PmLEA29)。在梅花胚胎发育晚期丰富蛋白基因中总共发现10次基因转换,其中7次发生在脱水素基因之间。
2.梅花脱水素基因表达受胁迫诱导,并具有明显的组织特异性：从‘北京玉蝶’梅花中克隆了5个脱水素基因(PmLEA8, PmLEA10, PmLEA19, PmLEA20, PmLEA29),并利用TA克隆的方法构建其GATEWAY克隆载体。测序结果表明,除PmLEA8外,其余基因序列与‘藏梅’全基因组数据库中的序列基本一致。表达分析证明：除PmLEA29在五种器官中(根、茎、叶、花、果实)均有高水平表达外,其余4个梅花脱水素基因表达均表现一定的组织特异性。6种胁迫处理下(低温、高温、干旱、盐、ABA和SA处理)梅花脱水素基因表达水平均有所升高,但PmLEA29在ABA处理下表达水平略有降低。
3.原核表达梅花脱水素可以提高大肠杆菌的抗逆能力：通过LR反应将克隆载体上的梅花脱水素基因重组到原核表达载体pDEST15上,转化大肠杆菌BL21(DE3),经1mM的IPTG诱导,表达了带GST标签的梅花脱水素蛋白。对转化大肠杆菌进行抗逆性分析证明,原核表达梅花脱水素可以提高大肠杆菌抗渗透胁迫和抗反复冻融能力。但不同梅花脱水素的改善效果不同,其中表达PmLEA8的改善效果稍差。
4.超表达梅花脱水素可以提高转基因烟草的抗寒能力：通过LR反应构建梅花脱水素基因的植物表达载体pEG203-PmLEA,转化农杆菌EHA105.采用农杆菌介导的叶圆盘法转化烟草NC89,经抗生素筛选和PCR验证获得转梅花脱水素基因烟草。其中,对转PmLEA20基因的烟草株系进行抗寒性分析证明：超表达梅花脱水素基因PmLEA20可以提高烟草种子在低温胁迫下的相对发芽率；低温下转PmLEA20基因烟草叶片电解质渗透率也明显低于对照。
本研究首次对梅花脱水素基因家族进行了克隆和功能分析,为利用梅花脱水素基因进行植物抗性改良、培育抗逆新品种,积累了新的分子材料,奠定了理论基础。
Mei (Prunus mume) belongs to the Rosaceae family. It is one of the most populous flowers in East Asia. Mei originated in the Yangtze River of China. Cold-hardiness breeding is important for mei utilization in the northern part of China. It has been demonstrated that dehydrins (also called group2late embryogenesis abundant proteins) play important roles in plant desiccation tolerance. In this study, we first did bioinformatics analysis to identify dehydrins in mei, and then cloned five dehydrin genes from P. mume 'Beijingyudie' using RT-PCR method. At last, mei dehydrin genes were overexpressed in Escherichia coli and tobacco to analysis their function.
1. The bioinformatics analysis of LEA gene family in P. mume:30LEA genes were identified from mei through genome-wide analysis. The PmLEA genes are distributed on all mei chromosomes except chromosome3. Twelve (40%) and four PmLEA genes are arranged in tandem and segmental duplications, respectively. The VmLEA genes could be divided into eight groups (LEA_1, LEA_2, LEA_3, LEA_4, LEA_5, PvLEA18, dehydrin and seed maturation protein). Six PmLEA genes (PmLEA8, PmLEA9, PmLEA10, PmLEA19, PmLEA20, PmLEA29) were divided into dehydrin group. Ten gene conversion events were observed in four LEA groups, with most of them (70%) identified in dehydrin group.
2. Cloning and expression analyses of dehydrin genes in P. mume:Five dehydrin genes (PmLEAs) were cloned from P. mume'Beijingyudie'using RT-PCR method. The entry vectors for PmLEAs were constructed using a TA method. The sequence of PmLEA8from P. mume'Beijingyudie'was longer than that from'Zangmei'genome database. The expressions of most genes were different among five organs (root, stem, leaf, flower, and fruit) except PmLEA29. The expression of most dehydrin genes was up-regulated under six abiotic stress treatments (cold, hot, drought, salt, ABA and SA treatment), except PmLEA29under ABA treatment.
3. Mei dehydrins confer the tolerance of E. coli:The E. coli expression vectors for dehydrin genes were established by LR recombination reaction. The mei dehydrins were expressed in E. coli BL21(DE3) after1mM IPTG induction. The results of a spot assay demonstrated that the expression of most mei dehydrins conferred the tolerance to the E. coli recombinant for cold and osmotic stresses.
4. Transgenic analysis of dehydrin genes of P. mume:The dehydrin genes in entry vector was recombined into plant expression vector pEG203through LR reaction and then transferred into tobacco NC89through Agrobacterium mediated leaf disc transformation. Twelve PmLEA20transgenic plants were identified through Basta selection and PCR test. These T1lines were used for germination analysis under low temperature and the relative germination rates of transgenic lines were higher than that of wild-type. The amount of electrolyte leakage in wild-type tobacco seedlings was higher than that of transgenic lines.
The cloning and functional analyses of P. mume dehydrin genes pave the way for their future utilization in plant resistance breeding.
1.首次预测了梅花胚胎发育晚期丰富蛋白基因家族成员：梅花中共有30个胚胎发育晚期丰富蛋白基因,分布于7个连锁群上。5个梅花胚胎发育晚期丰富蛋白基因属于片段重复,12个属于串联重复。根据序列相似性,可以将这些基因分为8组(LEA_1, LEA_2, LEA_3, LEA_4, LEA_5, PvLEA18,脱水素和SMP),其中脱水素家族含有6个成员(PmLEA8,PmLEA9, PmLEA10, PmLEA19,PmLEA20和PmLEA29)。在梅花胚胎发育晚期丰富蛋白基因中总共发现10次基因转换,其中7次发生在脱水素基因之间。
2.梅花脱水素基因表达受胁迫诱导,并具有明显的组织特异性：从‘北京玉蝶’梅花中克隆了5个脱水素基因(PmLEA8, PmLEA10, PmLEA19, PmLEA20, PmLEA29),并利用TA克隆的方法构建其GATEWAY克隆载体。测序结果表明,除PmLEA8外,其余基因序列与‘藏梅’全基因组数据库中的序列基本一致。表达分析证明：除PmLEA29在五种器官中(根、茎、叶、花、果实)均有高水平表达外,其余4个梅花脱水素基因表达均表现一定的组织特异性。6种胁迫处理下(低温、高温、干旱、盐、ABA和SA处理)梅花脱水素基因表达水平均有所升高,但PmLEA29在ABA处理下表达水平略有降低。
3.原核表达梅花脱水素可以提高大肠杆菌的抗逆能力：通过LR反应将克隆载体上的梅花脱水素基因重组到原核表达载体pDEST15上,转化大肠杆菌BL21(DE3),经1mM的IPTG诱导,表达了带GST标签的梅花脱水素蛋白。对转化大肠杆菌进行抗逆性分析证明,原核表达梅花脱水素可以提高大肠杆菌抗渗透胁迫和抗反复冻融能力。但不同梅花脱水素的改善效果不同,其中表达PmLEA8的改善效果稍差。
4.超表达梅花脱水素可以提高转基因烟草的抗寒能力：通过LR反应构建梅花脱水素基因的植物表达载体pEG203-PmLEA,转化农杆菌EHA105.采用农杆菌介导的叶圆盘法转化烟草NC89,经抗生素筛选和PCR验证获得转梅花脱水素基因烟草。其中,对转PmLEA20基因的烟草株系进行抗寒性分析证明：超表达梅花脱水素基因PmLEA20可以提高烟草种子在低温胁迫下的相对发芽率；低温下转PmLEA20基因烟草叶片电解质渗透率也明显低于对照。
本研究首次对梅花脱水素基因家族进行了克隆和功能分析,为利用梅花脱水素基因进行植物抗性改良、培育抗逆新品种,积累了新的分子材料,奠定了理论基础。
Mei (Prunus mume) belongs to the Rosaceae family. It is one of the most populous flowers in East Asia. Mei originated in the Yangtze River of China. Cold-hardiness breeding is important for mei utilization in the northern part of China. It has been demonstrated that dehydrins (also called group2late embryogenesis abundant proteins) play important roles in plant desiccation tolerance. In this study, we first did bioinformatics analysis to identify dehydrins in mei, and then cloned five dehydrin genes from P. mume 'Beijingyudie' using RT-PCR method. At last, mei dehydrin genes were overexpressed in Escherichia coli and tobacco to analysis their function.
1. The bioinformatics analysis of LEA gene family in P. mume:30LEA genes were identified from mei through genome-wide analysis. The PmLEA genes are distributed on all mei chromosomes except chromosome3. Twelve (40%) and four PmLEA genes are arranged in tandem and segmental duplications, respectively. The VmLEA genes could be divided into eight groups (LEA_1, LEA_2, LEA_3, LEA_4, LEA_5, PvLEA18, dehydrin and seed maturation protein). Six PmLEA genes (PmLEA8, PmLEA9, PmLEA10, PmLEA19, PmLEA20, PmLEA29) were divided into dehydrin group. Ten gene conversion events were observed in four LEA groups, with most of them (70%) identified in dehydrin group.
2. Cloning and expression analyses of dehydrin genes in P. mume:Five dehydrin genes (PmLEAs) were cloned from P. mume'Beijingyudie'using RT-PCR method. The entry vectors for PmLEAs were constructed using a TA method. The sequence of PmLEA8from P. mume'Beijingyudie'was longer than that from'Zangmei'genome database. The expressions of most genes were different among five organs (root, stem, leaf, flower, and fruit) except PmLEA29. The expression of most dehydrin genes was up-regulated under six abiotic stress treatments (cold, hot, drought, salt, ABA and SA treatment), except PmLEA29under ABA treatment.
3. Mei dehydrins confer the tolerance of E. coli:The E. coli expression vectors for dehydrin genes were established by LR recombination reaction. The mei dehydrins were expressed in E. coli BL21(DE3) after1mM IPTG induction. The results of a spot assay demonstrated that the expression of most mei dehydrins conferred the tolerance to the E. coli recombinant for cold and osmotic stresses.
4. Transgenic analysis of dehydrin genes of P. mume:The dehydrin genes in entry vector was recombined into plant expression vector pEG203through LR reaction and then transferred into tobacco NC89through Agrobacterium mediated leaf disc transformation. Twelve PmLEA20transgenic plants were identified through Basta selection and PCR test. These T1lines were used for germination analysis under low temperature and the relative germination rates of transgenic lines were higher than that of wild-type. The amount of electrolyte leakage in wild-type tobacco seedlings was higher than that of transgenic lines.
The cloning and functional analyses of P. mume dehydrin genes pave the way for their future utilization in plant resistance breeding.
引文
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