荒漠植物H~+-PPase基因的系统发育分析及SaVP1和KcNHX1基因的功能鉴定
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摘要
非生物胁迫是导致作物减产甚至植株死亡的重要因素。在非生物胁迫因子中,高温、干旱和盐碱是最常见、最广泛的影响植物生长、发育、生殖及产量等各个方面的逆境因子。高温和干旱主要对植物造成氧化胁迫和渗透胁迫,而盐胁迫主要对植物造成离子毒害和渗透胁迫。所有的逆境因子最终都会通过破坏细胞结构、损害胞内的生理功能从而影响植株的生长和发育。如何让植物在高温、干旱、盐碱等逆境条件下完成生活史并保持相应的产量成为目前亟待解决的问题。本研究从几种荒漠植物中克隆了抗逆相关基因H+转运无机焦磷酸酶(H+-PPase)基因和液泡膜Na+/H+逆向转运蛋白基因(NHX),并对H+-PPase基因的系统发育进行了分析,同时对来自苦豆子的H+-PPase和花花柴的NHX基因功能进行了研究,取得的主要结果如下：
1、克隆了来自新疆塔里木盆地7种荒漠植物的H+-PPase基因,并对其系统发育进行了分析。
本研究从荒漠植物苦豆子(Sophora alopecuroides L.)、乌拉尔甘草(Glycyrriza Uralensis Fisch.)、胀果甘草(Glycyrrhiza inflata Bat)、盐穗木(Glycyrrhiza inflata Bat).盐地碱蓬(Suaeda salsa)、硬枝碱蓬(Suaeda rigida)和花花柴(Karelinia caspica)7种荒漠植物中克隆得到了6条H+-PPase基因全长和8条H+-PPase基因片段序列。通过与NCBI数据库中已鉴定的240余条H+-PPase基因比对发现,新获得的这些基因除了具有H+-PPase家族特有的保守结构域外,该基因家族可能还存在两个新的保守结构域。利用17条来自与本研究所选材料亲缘关系较近的植物Ⅰ型H+-PPase基因、8条来自藻类、细菌和原生动物的Ⅰ型H+-PPase基因及9条Ⅱ型H+-PPase基因与本研究新克隆的14条基因进行了系统发育分析。聚类结果显示新克隆的14条基因可能都属于Ⅰ型H+-PPase基因。通过对上述H+-PPase基因编码蛋白质的3D结构预测发现,高等植物的Ⅰ型基因都以二聚体的形式存在,而所有的Ⅱ型基因以及来自低等生物的Ⅰ型基因则以单体的形式存在。SaVP3蛋白也以单体的形式存在,并与其他H+-PPase基因的同源性较低,因此推测SaVP3可能属于Ⅱ型H+-PPase基因或一类新的H+-PPase基因。这可能是由不同物种之间的基因交流所致,或者SaVP3和SaVP1基因可能是一对旁系同源基因。H+-PPase基因在进化上的高度保守性表明其对生物体功能的重要性,同时也表明这类基因在生物进化过程中所受的环境选择压力较小；H+-PPase基因家族的这些特点这对进一步研究其进化起源、基因结构与功能的关系提供了重要的参考。
2.SaVP1基因不仅能增强转基因植株的耐旱耐盐性,还具有耐高温的功能。
将来自荒漠植物苦豆子的SaVP1基因在拟南芥中超表达,结果发现转基因植株对干旱、高盐及高温的耐受性都强于野生型。在干旱和盐处理下,转基因植株中SOD酶活增强、MDA的含量降低,使得转基因植株的抗逆性明显强于野生型。将苗期的拟南芥在30-38℃处理4天后转基因植株的存活率在85%以上,而野生型仅为33%；而将结实期的植株在33℃处理1周后,转基因植株的结实性为野生型的2倍。这可能是在高温条件下SaVP1的超量表达增加了植株体内可溶性糖的积累以及POD的酶活,降低了MDA的含量；转基因植株在高温下的结实性也高于野生型,可能与SaVP1基因的超表达能够维持高温条件下花器官中较高的IAA含量有关。IAA的测试结果也说明在高温条件下转基因植株的花器官中IAA含量几乎是野生型的2倍,外施IAA的结果表明IAA的补充确实能部分恢复高温条件下的结实性,说明高温引起的结实性下降确实与IAA含量降低有关,而SaVP1的超表达能增加花器官中IAA的含量。同时在高温条件下,转基因植株叶片的气孔开度也明显增加,而且与K+和Ca2+相关的转运体基因的表达也明显上调,这些结果与转SaVP1基因植株具有耐旱、耐盐及耐高温的表型相吻合。
3、KcNHX1基因具有广谱抗逆性功能。
从荒漠植物花花柴中克隆了液泡膜Na+/H’逆向转运蛋自基因(KcNHX1),并对其蛋白质结构进行了分析,发现KcNHX1蛋白具有10次跨膜结构,pI值为6.27,分子量约为60kD。和SaVP1基因相似,将该基因在拟南芥中超表达,结果.显示转基因植株对干旱、高盐及高温的耐受性都强于野生型。苗期高温处理的结果显示转基因植株的存活率在88%以上,而野生型仅为18%；结实期在33℃处理1周后,转基因植株的结实性为野生型的2倍。而且在高温条件下KcNHX1的超量表达增强了转基因植株POD的酶活、增加了可溶性糖的积累、且MDA的含量降低,增加了花器官中IAA的含量,转基因植株在高温下的结实性也高于野生型。外施IAA结果表明IAA的补充确实能部分恢复高温条件下的拟南芥的结实性,说明KcNHX1的超表达能增加花器官中IAA的含量。同时在高温条件下,转基因植株叶片的气孔开度明显增加,而且与K+和Ca2+相关的转运体及离子通道蛋白基因的表达也明显上调,说明KcNHX1基因具有耐高温、耐旱、耐盐的功能。
通过本研究中所选的两个来自荒漠植物的SaVP1和KcNHX1基因的功能分析,表明来自荒漠植物的抗逆相关基因不仅具有模式甜土植物相应基因的功能,可能还具有一些新的功能尚待发掘。这不仅拓展了抗逆基因的来源,也为荒漠植物资源的开发利用、植物抗逆性机理研究及作物的抗逆性育种提供了重要的参考。
Abiotic stresses have profound influence on crop growth and production which result in yield decrease or even plant death. Among the abiotic stresses, heat, drought and salt are universal stressors and often have an adverse impacts on plant development, growth, reproduction and yield. Heat and drought stresses mainly impose an oxidative damage and osmotic stress, and salt always led to ion toxicity and osmotic stress, which can disrupt cellular structures and impair key physiological functions through metabolic rearrangements. How to make plant survive and keep relative yield in stress such as high temperature, drought and salinity, is a big challenge for plant biologist. In this work, H+-PPase and Na+/H+antiporter from some eremophyte plants were cloned. The phylogeny of H+-PPase was analyzed, and function of H+-PPase from Sophora alopecuroides(SaVP1) and Na+/H+antiporter from Karelinia caspica (KcNHX1) was evaluated. The major results were presented as following:
1. Phylogenetic analysis of the novel H+-PPases from eremophytes
H+-translocating inorganic pyrophosphatases (H+-PPase) were recognized as the original energy donors in the development of plants. A large number of researchers have shown that H+-PPase could be an early origin protein that participated in many important biochemical and physiological processes. In this study we cloned14novel sequences from7eremophytes:Sophora alopecuroid (Sa), Glycyrrhiza uralensis (Gu), Glycyrrhiza inflata (Gi), Suaeda salsa (Ss), Suaeda rigida (Sr), Halostachys caspica (Hc), and Karelinia caspia (Kc). These novel sequences included6ORFs and8fragments, and they were identified as H+-PPases based on the typical conserved domains. Besides the identified domains, the result of sequence alignment showed that there could be two novel conserved motifs. A phylogenetic tree was constructed, including the14novel H+-PPase amino acid sequences and the other34identified H+-PPase protein sequences representing plants, algae, protozoans and bacteria. It was shown that these48H+-PPases were classified into two groups:type I and type II H+-PPase. The novel14H+-PPases from7eremophytes were classified into the type I H+-PPase. The3D structures of these H+-PPase proteins were predicted, which suggested that all type I H+-PPases from higher plants and algae were homodimers, while other type I H+-PPases from bacteria and protozoans and all type II H+-PPases were monomers. The3D structures of these novel H+-PPases were homodimers except for SaVP3which was a monomer. The result of little diversity at the amino acid sequence of H+-PPase implied that these genes play a vital role in the development of the organism and have suffered little selection pressure. But it is possible that different evolutionary histories or lateral gene transfers of H+-PPase between different (micro) organisms exist based SaVP3, or this gene could represent a novel type of H+-PPase, or they could be paralogs between SaVP3and SaVP1in Sophora alopecuroid based their lower homology. This regular structure could provide important evidence for the evolutionary origin and study of the relationship between the structure and function among members of the H+-PPase family.
2. Overexpression of the SaVPl enhanced tolerance not only to drought and salt, but also heat stress of Arabidopsis
The physiological role of a vacuolar H+-PPase (SaVP1) from an eremophyte Sophora alopecuroid was evaluated through overexpressing the gene in Arabidopsis thaliana. Overexpression of SaVP1in Arabidopsis enhanced tolerance to drought and salt stresses. Moreover, the SaVP1overexpressing plants showed enhanced tolerance to heat stress. At seedling stage, the survival rate was above85%for transgenic lines, and it was only33%in WT plants after heat stress. The fertility is twice in transgenic lines compared to WT plants after heat stress. It could be that SaVP1overexpression increased the accumulation of soluble sugar and chlorophyll, enhanced the POD activety, and decreased the MDA content in the transgenic plants. Simultaneously, SaVP1overexpression increased seed yield through maintaining higher content of IAA in flora organ than in wild type plants under high temperature. In addition, application of low concentration of IAA led to higher fertility of SaVP1overexpression plants than plants without IAA treatment and wild-type under high temperature. Bisides, It showed that overexpression of SaVP1resulted in several K+and Ca2+channel/transporters genes up-regulated. And the leaf stomatal aperture were increased in transgenic plants compared to wild type under high temperature stress. The results indicated that genes originated from eremophyte could be used as the candidate for enhancing heat, drought and salt tolerance of other plant species.
3. KcNHXl overexpressing confers multiple abiotic stress tolerance in Arabidopsis
The KcNHX1was cloned from eremophyte Karelinia caspia, its cDNA sequence contains1620bp, and codes539amino acid. The protein pI is6.27, the MW is about60kD with10putative transmemberane regions. Like SaVP1, the overexpression of KcNHX1in Arabidopsis enhanced tolerance to drought, salt and teat stress. At seedling stage, the survival rate was about88%for transgenic lines, and it was only18%in WT plants after heat stress. The fertility is higher in transgenic lines than in WT plants after heat stress. The overexpression of KcNHX1resulted in several K+and Ca2+channel/transporters genes up-regulated. Simultaneously, overexpression of KcNHXl maintained higher seed yield and IAA content in flora organ under high temperature than wild type plants. In addition, application of low concentration of IAA led to higher fertility of KcNHXl overexpression plants under high temperature. The results indicate that KcNHX1could be used as a candidate for enhancing drought, salt and heat tolerance of other plant species, too.
In this work, the results showed that genes related to tolerance might present some unknown function in addition to some identified function of corresponding genes in glycophyte through analyzing characterization of SaVP1and KcNHX1from eremophytes. Our research will expand the sources of stress-tolerance genes, and provide an important reference for the development and utilization of eremophytes, which would benefit crop breeding and the study of plant resistance mechanism.
1、克隆了来自新疆塔里木盆地7种荒漠植物的H+-PPase基因,并对其系统发育进行了分析。
本研究从荒漠植物苦豆子(Sophora alopecuroides L.)、乌拉尔甘草(Glycyrriza Uralensis Fisch.)、胀果甘草(Glycyrrhiza inflata Bat)、盐穗木(Glycyrrhiza inflata Bat).盐地碱蓬(Suaeda salsa)、硬枝碱蓬(Suaeda rigida)和花花柴(Karelinia caspica)7种荒漠植物中克隆得到了6条H+-PPase基因全长和8条H+-PPase基因片段序列。通过与NCBI数据库中已鉴定的240余条H+-PPase基因比对发现,新获得的这些基因除了具有H+-PPase家族特有的保守结构域外,该基因家族可能还存在两个新的保守结构域。利用17条来自与本研究所选材料亲缘关系较近的植物Ⅰ型H+-PPase基因、8条来自藻类、细菌和原生动物的Ⅰ型H+-PPase基因及9条Ⅱ型H+-PPase基因与本研究新克隆的14条基因进行了系统发育分析。聚类结果显示新克隆的14条基因可能都属于Ⅰ型H+-PPase基因。通过对上述H+-PPase基因编码蛋白质的3D结构预测发现,高等植物的Ⅰ型基因都以二聚体的形式存在,而所有的Ⅱ型基因以及来自低等生物的Ⅰ型基因则以单体的形式存在。SaVP3蛋白也以单体的形式存在,并与其他H+-PPase基因的同源性较低,因此推测SaVP3可能属于Ⅱ型H+-PPase基因或一类新的H+-PPase基因。这可能是由不同物种之间的基因交流所致,或者SaVP3和SaVP1基因可能是一对旁系同源基因。H+-PPase基因在进化上的高度保守性表明其对生物体功能的重要性,同时也表明这类基因在生物进化过程中所受的环境选择压力较小；H+-PPase基因家族的这些特点这对进一步研究其进化起源、基因结构与功能的关系提供了重要的参考。
2.SaVP1基因不仅能增强转基因植株的耐旱耐盐性,还具有耐高温的功能。
将来自荒漠植物苦豆子的SaVP1基因在拟南芥中超表达,结果发现转基因植株对干旱、高盐及高温的耐受性都强于野生型。在干旱和盐处理下,转基因植株中SOD酶活增强、MDA的含量降低,使得转基因植株的抗逆性明显强于野生型。将苗期的拟南芥在30-38℃处理4天后转基因植株的存活率在85%以上,而野生型仅为33%；而将结实期的植株在33℃处理1周后,转基因植株的结实性为野生型的2倍。这可能是在高温条件下SaVP1的超量表达增加了植株体内可溶性糖的积累以及POD的酶活,降低了MDA的含量；转基因植株在高温下的结实性也高于野生型,可能与SaVP1基因的超表达能够维持高温条件下花器官中较高的IAA含量有关。IAA的测试结果也说明在高温条件下转基因植株的花器官中IAA含量几乎是野生型的2倍,外施IAA的结果表明IAA的补充确实能部分恢复高温条件下的结实性,说明高温引起的结实性下降确实与IAA含量降低有关,而SaVP1的超表达能增加花器官中IAA的含量。同时在高温条件下,转基因植株叶片的气孔开度也明显增加,而且与K+和Ca2+相关的转运体基因的表达也明显上调,这些结果与转SaVP1基因植株具有耐旱、耐盐及耐高温的表型相吻合。
3、KcNHX1基因具有广谱抗逆性功能。
从荒漠植物花花柴中克隆了液泡膜Na+/H’逆向转运蛋自基因(KcNHX1),并对其蛋白质结构进行了分析,发现KcNHX1蛋白具有10次跨膜结构,pI值为6.27,分子量约为60kD。和SaVP1基因相似,将该基因在拟南芥中超表达,结果.显示转基因植株对干旱、高盐及高温的耐受性都强于野生型。苗期高温处理的结果显示转基因植株的存活率在88%以上,而野生型仅为18%；结实期在33℃处理1周后,转基因植株的结实性为野生型的2倍。而且在高温条件下KcNHX1的超量表达增强了转基因植株POD的酶活、增加了可溶性糖的积累、且MDA的含量降低,增加了花器官中IAA的含量,转基因植株在高温下的结实性也高于野生型。外施IAA结果表明IAA的补充确实能部分恢复高温条件下的拟南芥的结实性,说明KcNHX1的超表达能增加花器官中IAA的含量。同时在高温条件下,转基因植株叶片的气孔开度明显增加,而且与K+和Ca2+相关的转运体及离子通道蛋白基因的表达也明显上调,说明KcNHX1基因具有耐高温、耐旱、耐盐的功能。
通过本研究中所选的两个来自荒漠植物的SaVP1和KcNHX1基因的功能分析,表明来自荒漠植物的抗逆相关基因不仅具有模式甜土植物相应基因的功能,可能还具有一些新的功能尚待发掘。这不仅拓展了抗逆基因的来源,也为荒漠植物资源的开发利用、植物抗逆性机理研究及作物的抗逆性育种提供了重要的参考。
Abiotic stresses have profound influence on crop growth and production which result in yield decrease or even plant death. Among the abiotic stresses, heat, drought and salt are universal stressors and often have an adverse impacts on plant development, growth, reproduction and yield. Heat and drought stresses mainly impose an oxidative damage and osmotic stress, and salt always led to ion toxicity and osmotic stress, which can disrupt cellular structures and impair key physiological functions through metabolic rearrangements. How to make plant survive and keep relative yield in stress such as high temperature, drought and salinity, is a big challenge for plant biologist. In this work, H+-PPase and Na+/H+antiporter from some eremophyte plants were cloned. The phylogeny of H+-PPase was analyzed, and function of H+-PPase from Sophora alopecuroides(SaVP1) and Na+/H+antiporter from Karelinia caspica (KcNHX1) was evaluated. The major results were presented as following:
1. Phylogenetic analysis of the novel H+-PPases from eremophytes
H+-translocating inorganic pyrophosphatases (H+-PPase) were recognized as the original energy donors in the development of plants. A large number of researchers have shown that H+-PPase could be an early origin protein that participated in many important biochemical and physiological processes. In this study we cloned14novel sequences from7eremophytes:Sophora alopecuroid (Sa), Glycyrrhiza uralensis (Gu), Glycyrrhiza inflata (Gi), Suaeda salsa (Ss), Suaeda rigida (Sr), Halostachys caspica (Hc), and Karelinia caspia (Kc). These novel sequences included6ORFs and8fragments, and they were identified as H+-PPases based on the typical conserved domains. Besides the identified domains, the result of sequence alignment showed that there could be two novel conserved motifs. A phylogenetic tree was constructed, including the14novel H+-PPase amino acid sequences and the other34identified H+-PPase protein sequences representing plants, algae, protozoans and bacteria. It was shown that these48H+-PPases were classified into two groups:type I and type II H+-PPase. The novel14H+-PPases from7eremophytes were classified into the type I H+-PPase. The3D structures of these H+-PPase proteins were predicted, which suggested that all type I H+-PPases from higher plants and algae were homodimers, while other type I H+-PPases from bacteria and protozoans and all type II H+-PPases were monomers. The3D structures of these novel H+-PPases were homodimers except for SaVP3which was a monomer. The result of little diversity at the amino acid sequence of H+-PPase implied that these genes play a vital role in the development of the organism and have suffered little selection pressure. But it is possible that different evolutionary histories or lateral gene transfers of H+-PPase between different (micro) organisms exist based SaVP3, or this gene could represent a novel type of H+-PPase, or they could be paralogs between SaVP3and SaVP1in Sophora alopecuroid based their lower homology. This regular structure could provide important evidence for the evolutionary origin and study of the relationship between the structure and function among members of the H+-PPase family.
2. Overexpression of the SaVPl enhanced tolerance not only to drought and salt, but also heat stress of Arabidopsis
The physiological role of a vacuolar H+-PPase (SaVP1) from an eremophyte Sophora alopecuroid was evaluated through overexpressing the gene in Arabidopsis thaliana. Overexpression of SaVP1in Arabidopsis enhanced tolerance to drought and salt stresses. Moreover, the SaVP1overexpressing plants showed enhanced tolerance to heat stress. At seedling stage, the survival rate was above85%for transgenic lines, and it was only33%in WT plants after heat stress. The fertility is twice in transgenic lines compared to WT plants after heat stress. It could be that SaVP1overexpression increased the accumulation of soluble sugar and chlorophyll, enhanced the POD activety, and decreased the MDA content in the transgenic plants. Simultaneously, SaVP1overexpression increased seed yield through maintaining higher content of IAA in flora organ than in wild type plants under high temperature. In addition, application of low concentration of IAA led to higher fertility of SaVP1overexpression plants than plants without IAA treatment and wild-type under high temperature. Bisides, It showed that overexpression of SaVP1resulted in several K+and Ca2+channel/transporters genes up-regulated. And the leaf stomatal aperture were increased in transgenic plants compared to wild type under high temperature stress. The results indicated that genes originated from eremophyte could be used as the candidate for enhancing heat, drought and salt tolerance of other plant species.
3. KcNHXl overexpressing confers multiple abiotic stress tolerance in Arabidopsis
The KcNHX1was cloned from eremophyte Karelinia caspia, its cDNA sequence contains1620bp, and codes539amino acid. The protein pI is6.27, the MW is about60kD with10putative transmemberane regions. Like SaVP1, the overexpression of KcNHX1in Arabidopsis enhanced tolerance to drought, salt and teat stress. At seedling stage, the survival rate was about88%for transgenic lines, and it was only18%in WT plants after heat stress. The fertility is higher in transgenic lines than in WT plants after heat stress. The overexpression of KcNHX1resulted in several K+and Ca2+channel/transporters genes up-regulated. Simultaneously, overexpression of KcNHXl maintained higher seed yield and IAA content in flora organ under high temperature than wild type plants. In addition, application of low concentration of IAA led to higher fertility of KcNHXl overexpression plants under high temperature. The results indicate that KcNHX1could be used as a candidate for enhancing drought, salt and heat tolerance of other plant species, too.
In this work, the results showed that genes related to tolerance might present some unknown function in addition to some identified function of corresponding genes in glycophyte through analyzing characterization of SaVP1and KcNHX1from eremophytes. Our research will expand the sources of stress-tolerance genes, and provide an important reference for the development and utilization of eremophytes, which would benefit crop breeding and the study of plant resistance mechanism.
引文
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