枳抗逆基因的克隆及功能鉴定
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摘要
柑橘是我国南方重要的水果之一,其产业发展对促进地区经济具有重要作用。近年来柑橘产业虽然取得了巨大的发展,但常受到干旱、盐及低温等各种非生物逆境胁迫,因此,如何高效培育柑橘抗逆新品种成为当前育种学家亟需解决的课题。柑橘育种方法主要有常规育种、芽变选种和基因工程等,但常规育种存在雌雄性败育,珠心胚干扰,遗传高度杂合等问题,而芽变选种培育周期过长,随着生物技术水平的提高,特别是基因工程的发展,为柑橘的育种提供了一条新的途径。抗逆机理解析及重要基因克隆是基因工程的前题,是柑橘分子育种的关键。但目前柑橘抗性基因的克隆较少,鉴定出功能的基因更少,这严重阻碍了柑橘抗逆分子育种的步伐。从前人的研究得知,转录因子ABF、bHLH、ICE1及蛋白激酶MAPK在植物抗非生物胁迫中起重要作用,因此本研究从枳中分离、克隆得到ABF,MAPK、bHLH及ICE1基因,并对他们进行抗逆功能鉴定。其目的是为了深入解析抗性基因的分子机理,获得拥有自主知识产权的柑橘抗性基因资源。主要研究结果如下：
1.采用电子克隆从枳中克隆得到ABA响应的转录因子(PtrABF),它包含1347bp的开放阅读框,编码448个氨基酸,预测蛋白质分子量为48kD,等电点9.66。多序列比对显示PtrABF与其它物种的ABF有很高的同源性。PtrABF定位于细胞核,有转录激活活性,能与ABRE元件结合,表明它是一个转录因子。表达模式分析显示它能被脱水、低温及ABA诱导。烟草中超表达PtrABF,表明转基因植株比未转化植株的抗脱水和抗旱能力显著提高。脱水和干旱条件下转基因植株比未转化植株活性氧积累低,三种抗氧化酶的活性及基因表达量均较高。另外,在干旱处理前后,9个逆境相关基因的表达量在转基因烟草中都比非转基因植株高。所有结果显示,PtrABF转基因植株抗旱能力显著提高。
2.利用电子克隆及RACE技术从枳中分离克隆出PtrMAPK,该基因包含1128bp开放性阅读框,编码375个氨基酸,蛋白质分子量为43kD,等电点为5.51。生物信息学分析表明PtrMAPK包括11个保守的激酶结构域和一个磷酸化位点,亚细胞定位表明它定位于细胞核。PtrMAPK受干旱及低温的诱导但不受盐的诱导。与未转化植株相比,PtrMAPK转基因烟草超表达植株的抗脱水和抗旱能力明显提高,进一步研究表明,抗性增强与抗氧化酶活性提高相关。此外,转基因烟草在干旱处理前后,相关逆境响应基因的表达量明显上升。研究结果表明,PtrMAPK转基因植株具有较强的抗旱能力。
3.从枳中克隆出PtrbHLH基因,该基因全长1921bp,它包括1464bp的开放性阅读框,编码487个氨基酸,等电点为5.30,分子量预测为53.6kD。多序列比对表明其编码的氨基酸序列中含有保守的ZIP结构域、bHLH结构域、C末段区域及SUMO结合结构域。PtrbHLH定位于细胞核,具有转录激活活性。ExPASy网站中的Signal P软件预测表明PtrbHLH氨基酸序列中具有一个核定位信号(NLS)及一个bHLH DNA结合结构域；SMART预测PtrbHLH氨基酸序列中含有一个转录激活结构域。PtrbHLH受低温、脱水及盐的诱导。超表达PtrbHLH的转基因柠檬和烟草植株的抗寒能力提高,但其RNAi-PtrbHLH转入枳使其抗性下降。采用Affymetrix芯片杂交分析表明,PtrbHLH的转基因柠檬中有108个基因上调,其中包括与H202清除有关的POD基因。进一步研究表明,超表达PtrbHLH的转基因植株(烟草和柠檬)中POD活性强,在氧化胁迫下抗性也增强。但采用POD抑制剂处理转基因烟草后,H202清除能力及抗寒性减弱。这些结果表明,PtrbHLH是通过调控POD来增强植物的抗寒能力。
4.采用电子克隆和RACE等手段从枳中克隆出PtrICE1基因,该基因包含一个1680bp的开放阅读框,编码559个氨基酸,蛋白预测的分子量为61kD,等电点为5.27。生物信息学分析显示PtrICE1氨基酸序列与其它植物氨基酸同源性较高,有两个保守的bHLH结构域和一个SUMO结合结构域。PtrICE1定位于细胞核,有很强的转录激活活性以及能与顺式作用元件MYC结合,这些都说明它是一个转录因子。在低温、脱水、盐及脱落酸诱导下PtrICE1的转录水平都有很大的提高,表明PtrICE1能受这些非生物逆境的诱导。PtrICE1在CaM35S启动子下在烟草中的超表达分析显示,它的抗寒能力得到很大的提高。
Citrus is one of the most important fruit trees in the southern part of China and possesses a pivotal position in the regional economy promotion. Though tremendous progress for the citrus industry has been achieved in recent decades, the developments of quality and yield are impeded by various stresses, such as drought, salt and low temperature, making stress tolerance one of the important objectives for citrus breeding. Due to polyembryony and sterility of reproductive organs, it is difficult for traditional breeding to obtain stress-tolerant germplasm, which can be complemented by gene engineering as an alternative. But so far, few stress-related genes were reported. ABA responsive element binding factor (ABF), Mitogen-activated protein kinase (MAPK), bHLH and ICE1are four well-known stress-related genes. In this study, all these three genes were cloned from Poncirus trifoliate and transferred into either tobacco or Arabidopsis for functional analysis to dissect the corresponding mechanism of stress tolerance and expand the pool of stress-related genes in Citrus. Main results were as follows:
1. PtrABF was obtained via in silico cloning, containing an open reading frame (ORF) of1347bp and encoding448amino acids with protein molecular weight (Mw)48kD and isoelectric point (pI)9.66. Positioning in the nucleus, strong transcriptional activation and binding with ABRE, these data proved that PtrABF is a transcription factor (TF), which can be induced by dehydration, low temperature and ABA. PtrABF transgenic tobacco showed improved dehydration and drought tolerance. It may be due to lower accumulation of reactive oxygen species (ROS) since the expression of anti-oxidant enzymes in transgenic plants were higher, so were the activities. In additional,9stress-related genes were observed higher expression in transgenic tobacco before and after drought treatment.
2. In silico cloning and5'-RACE were applied to obtain the full length of PtrMAPK with an ORF of1128bp, Mw43kD, pI5.51. Bioinformatics analysis revealed that PtrMAPK possesses11conserved kinase domains and a phosphorylation site. And subcellular localization analysis showed that it is situated in nucleus. The expression of PtrMAPK was induced by drought and low temperature but not salt. Overexpressed PtrMAPK tobacco showed elevated tolerance to dehydration and drought, lower water loss and ROS accumulation, higher anti-oxidant enzyme activities. Meanwhile, the expressions of ROS-scavenging and stress-related genes were higher in transgenic tobacco before and after drought stress.
3. PtrbHLH was isolated using the same method as PtrMAPK, owning an ORF of1464bp, Mw53.6kD, pI5.30. It enjoyed high amino acid homology with bHLH of other species. The bHLH family of transcription factors is characterized by an acidic domain near the N terminus and a conserved near the C terminus. At the C terninus of the PtrbHLH there are one bHLH zipper domain and one SUMO conjugation motif. PtrHLH was localized in nuleus and has strong transcriptional activation which demonstrated that it is a TF. Its expression can be induced by low temperature, dehydration, salt and ABA.and the overexpression of PtrbHLH in tobacco resulted in cold tolerance. Overexpressing PtrbHLH in lemon enhanced its cold tolerance but the introduction of RNAi-PtrbHLH into Poncirus undermined its cold tolerance. Affymetrix chip data revealed that there were108up-regulated genes in the lemon overexpressing PtrbHLH, including POD gene which can scavenge H2O2. Further experiments using PtrbHLH-overexpressing transgenic materials (tobacco and lemon) showed that POD activities have been obviously elevated, so have the anti-oxidative ability. When challenged with POD inhibitor, the transgenic tobacco demonstrated lowered POD activity and cold tolerance. These results suggested that PtrbHLH may enhance cold tolerance via manipulating POD activity.
4. PtrICE1was isolated same as PtrMAPK, owning an ORF of1680bp, Mw61kD, pI5.27. It enjoyed high amino acid homology with ICE1of other species. Two conserved bHLH motif and one SUMO binding domain were present in PtrICE1. Localized in nuleus, strong transcriptional activation and binding to cis element MYC demonstrated that it is a TF. Its expression can be induced by low temperature, dehydration, salt and ABA. And the overexpression of PtrICE1in tobacco resulted in cold tolerance.
1.采用电子克隆从枳中克隆得到ABA响应的转录因子(PtrABF),它包含1347bp的开放阅读框,编码448个氨基酸,预测蛋白质分子量为48kD,等电点9.66。多序列比对显示PtrABF与其它物种的ABF有很高的同源性。PtrABF定位于细胞核,有转录激活活性,能与ABRE元件结合,表明它是一个转录因子。表达模式分析显示它能被脱水、低温及ABA诱导。烟草中超表达PtrABF,表明转基因植株比未转化植株的抗脱水和抗旱能力显著提高。脱水和干旱条件下转基因植株比未转化植株活性氧积累低,三种抗氧化酶的活性及基因表达量均较高。另外,在干旱处理前后,9个逆境相关基因的表达量在转基因烟草中都比非转基因植株高。所有结果显示,PtrABF转基因植株抗旱能力显著提高。
2.利用电子克隆及RACE技术从枳中分离克隆出PtrMAPK,该基因包含1128bp开放性阅读框,编码375个氨基酸,蛋白质分子量为43kD,等电点为5.51。生物信息学分析表明PtrMAPK包括11个保守的激酶结构域和一个磷酸化位点,亚细胞定位表明它定位于细胞核。PtrMAPK受干旱及低温的诱导但不受盐的诱导。与未转化植株相比,PtrMAPK转基因烟草超表达植株的抗脱水和抗旱能力明显提高,进一步研究表明,抗性增强与抗氧化酶活性提高相关。此外,转基因烟草在干旱处理前后,相关逆境响应基因的表达量明显上升。研究结果表明,PtrMAPK转基因植株具有较强的抗旱能力。
3.从枳中克隆出PtrbHLH基因,该基因全长1921bp,它包括1464bp的开放性阅读框,编码487个氨基酸,等电点为5.30,分子量预测为53.6kD。多序列比对表明其编码的氨基酸序列中含有保守的ZIP结构域、bHLH结构域、C末段区域及SUMO结合结构域。PtrbHLH定位于细胞核,具有转录激活活性。ExPASy网站中的Signal P软件预测表明PtrbHLH氨基酸序列中具有一个核定位信号(NLS)及一个bHLH DNA结合结构域；SMART预测PtrbHLH氨基酸序列中含有一个转录激活结构域。PtrbHLH受低温、脱水及盐的诱导。超表达PtrbHLH的转基因柠檬和烟草植株的抗寒能力提高,但其RNAi-PtrbHLH转入枳使其抗性下降。采用Affymetrix芯片杂交分析表明,PtrbHLH的转基因柠檬中有108个基因上调,其中包括与H202清除有关的POD基因。进一步研究表明,超表达PtrbHLH的转基因植株(烟草和柠檬)中POD活性强,在氧化胁迫下抗性也增强。但采用POD抑制剂处理转基因烟草后,H202清除能力及抗寒性减弱。这些结果表明,PtrbHLH是通过调控POD来增强植物的抗寒能力。
4.采用电子克隆和RACE等手段从枳中克隆出PtrICE1基因,该基因包含一个1680bp的开放阅读框,编码559个氨基酸,蛋白预测的分子量为61kD,等电点为5.27。生物信息学分析显示PtrICE1氨基酸序列与其它植物氨基酸同源性较高,有两个保守的bHLH结构域和一个SUMO结合结构域。PtrICE1定位于细胞核,有很强的转录激活活性以及能与顺式作用元件MYC结合,这些都说明它是一个转录因子。在低温、脱水、盐及脱落酸诱导下PtrICE1的转录水平都有很大的提高,表明PtrICE1能受这些非生物逆境的诱导。PtrICE1在CaM35S启动子下在烟草中的超表达分析显示,它的抗寒能力得到很大的提高。
Citrus is one of the most important fruit trees in the southern part of China and possesses a pivotal position in the regional economy promotion. Though tremendous progress for the citrus industry has been achieved in recent decades, the developments of quality and yield are impeded by various stresses, such as drought, salt and low temperature, making stress tolerance one of the important objectives for citrus breeding. Due to polyembryony and sterility of reproductive organs, it is difficult for traditional breeding to obtain stress-tolerant germplasm, which can be complemented by gene engineering as an alternative. But so far, few stress-related genes were reported. ABA responsive element binding factor (ABF), Mitogen-activated protein kinase (MAPK), bHLH and ICE1are four well-known stress-related genes. In this study, all these three genes were cloned from Poncirus trifoliate and transferred into either tobacco or Arabidopsis for functional analysis to dissect the corresponding mechanism of stress tolerance and expand the pool of stress-related genes in Citrus. Main results were as follows:
1. PtrABF was obtained via in silico cloning, containing an open reading frame (ORF) of1347bp and encoding448amino acids with protein molecular weight (Mw)48kD and isoelectric point (pI)9.66. Positioning in the nucleus, strong transcriptional activation and binding with ABRE, these data proved that PtrABF is a transcription factor (TF), which can be induced by dehydration, low temperature and ABA. PtrABF transgenic tobacco showed improved dehydration and drought tolerance. It may be due to lower accumulation of reactive oxygen species (ROS) since the expression of anti-oxidant enzymes in transgenic plants were higher, so were the activities. In additional,9stress-related genes were observed higher expression in transgenic tobacco before and after drought treatment.
2. In silico cloning and5'-RACE were applied to obtain the full length of PtrMAPK with an ORF of1128bp, Mw43kD, pI5.51. Bioinformatics analysis revealed that PtrMAPK possesses11conserved kinase domains and a phosphorylation site. And subcellular localization analysis showed that it is situated in nucleus. The expression of PtrMAPK was induced by drought and low temperature but not salt. Overexpressed PtrMAPK tobacco showed elevated tolerance to dehydration and drought, lower water loss and ROS accumulation, higher anti-oxidant enzyme activities. Meanwhile, the expressions of ROS-scavenging and stress-related genes were higher in transgenic tobacco before and after drought stress.
3. PtrbHLH was isolated using the same method as PtrMAPK, owning an ORF of1464bp, Mw53.6kD, pI5.30. It enjoyed high amino acid homology with bHLH of other species. The bHLH family of transcription factors is characterized by an acidic domain near the N terminus and a conserved near the C terminus. At the C terninus of the PtrbHLH there are one bHLH zipper domain and one SUMO conjugation motif. PtrHLH was localized in nuleus and has strong transcriptional activation which demonstrated that it is a TF. Its expression can be induced by low temperature, dehydration, salt and ABA.and the overexpression of PtrbHLH in tobacco resulted in cold tolerance. Overexpressing PtrbHLH in lemon enhanced its cold tolerance but the introduction of RNAi-PtrbHLH into Poncirus undermined its cold tolerance. Affymetrix chip data revealed that there were108up-regulated genes in the lemon overexpressing PtrbHLH, including POD gene which can scavenge H2O2. Further experiments using PtrbHLH-overexpressing transgenic materials (tobacco and lemon) showed that POD activities have been obviously elevated, so have the anti-oxidative ability. When challenged with POD inhibitor, the transgenic tobacco demonstrated lowered POD activity and cold tolerance. These results suggested that PtrbHLH may enhance cold tolerance via manipulating POD activity.
4. PtrICE1was isolated same as PtrMAPK, owning an ORF of1680bp, Mw61kD, pI5.27. It enjoyed high amino acid homology with ICE1of other species. Two conserved bHLH motif and one SUMO binding domain were present in PtrICE1. Localized in nuleus, strong transcriptional activation and binding to cis element MYC demonstrated that it is a TF. Its expression can be induced by low temperature, dehydration, salt and ABA. And the overexpression of PtrICE1in tobacco resulted in cold tolerance.
引文
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