乙烯诱导伏令夏橙果实脱落过程中重要功能基因的克隆、表达与功能分析
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摘要
柑橘果实发育过程中若受到过度胁迫会导致大量落果,从而造成大幅度减产,特别是晚熟品种在越冬时常会遇到低温,导致采前落果,造成巨大损失。另外,目前柑橘的采收方式主要为人工采摘,成本很高,在美国甚至超过柑橘生产成本的总和。因此,能实现果实的可控脱落成为柑橘生产者和科研工作者追求的主要目标之一。
本研究利用柑橘基因组芯片(Citrus Genome Array)研究乙烯诱导柑橘果实脱落并探讨其分子机理。研究中使用浓度为20μL/L的乙烯分别处理带叶的伏令夏橙(Citrus sinensis cv. Valencia)果实4 h、12 h和24 h,然后提取果实离层RNA进行芯片检测。聚类分析结果表明,芯片数据具有较好的重复性。12条典型基因的Real Time-PCR分析结果也表明这些基因表达的变化趋势与芯片数据基本一致。对乙烯诱导的差异表达基因进行分析,发现乙烯处理3个时间点上均表达的基因有510条(P<0.01)。对差异表达基因进行功能分析,发现乙烯诱导下上调表达的主要有编码膜类蛋白、能量代谢相关蛋白、胁迫相关蛋白、转录因子、丝氨酸／苏氨酸激酶类、磷酸酶类、钙离子结合蛋白类、水解酶类、乙烯代谢途径相关蛋白、转移酶类、受体蛋白类等基因,下调表达的主要包括编码核糖体类蛋白、水解酶类、水通道蛋白、细胞色素P450、金属离子结合蛋白、光合系统相关蛋白、ABC转运蛋白、脂酶类等基因。进一步分析发现,编码乙烯代谢途径相关蛋白的基因、伤害诱导基因和编码低氧胁迫蛋白的基因之间存在着关联表达,因而筛选出ETR2、低氧应答基因和伤害诱导基因进行深入研究。主要结果如下：
克隆了乙烯诱导表达的低氧应答基因(CsHRP)并进行表达分析。采用RACE技术克隆了CsHRP的全长cDNA并测序分析。结果表明,CsHRP的cDNA全长为795 bp, ORF编码98个氨基酸。序列比对分析结果显示CsHRP与甜橙低氧应答基因的同源性达97%。对蛋白质二级结构的预测结果推测CsHRP可能与抗病有关。采用Real-time PCR方法检测了CsHRP在外源乙烯诱导下的表达情况,结果显示CsHRP在乙烯处理前期(4 h)高度表达,12 h后减低到原来的水平。
乙烯可诱导伤害诱导蛋白(Wound induced proteins, WIPs)的表达。本研究克隆了柑橘的4条伤害诱导基因CsWIP1、CsWIP2、CsWIP3和CsWIP4,序列分析结果表明这4个伤害诱导基因分为两类,其中CsWIP1、CsWIP2和CsWIP3聚为一类,而CsWIP4与前三者在结构上差异较大故聚为另一类。WIP蛋白在C-端有一个非常保守的α-螺旋,在N-端也有一个α-螺旋,CsWIP1、CsWIP2和CsWIP3在两个α-螺旋处都保守,而CsWIP4只在C-端α-螺旋处保守。Real-time PCR结果显示：CsWIP1仅在外源乙烯处理下表达,受乙烯诱导；CsWIP1无论是在伤害处理还是在乙烯处理条件下,仅在叶片中表达,说明CsWIP2的表达具有组织特异性;CsWIP3在乙烯或伤害处理的叶片和离层中均表达,说明CsWIP3的表达没有组织特异性；CsWIP4仅在伤害处理下表达,基本上不受乙烯调控。
ETR2是乙烯受体基因之一。为深入了解乙烯受体基因(ETR2)在伏令夏橙果实脱落过程中的作用,本研究克隆了ETR2基因并研究了其表达。通过RACE技术得到ETR2的cDNA序列为1800 bp。序列比对分析表明伏令夏橙ETR2与其它甜橙ETR2的同源性达98%。ETR2蛋白质包含了3个结构域,分别为GAF结构域、组氨酸蛋白激酶(HPK)同源二聚体结合区域和类似CheY结构域。通过Real-time PCR检测了ETR2在外源乙烯诱导4 h、12 h和24 h时的表达量。结果显示,ETR2在乙烯诱导4 h、12 h和24 h的表达量分别是对照的2倍、4倍和1倍。
为进一步验证CsHRP和CsWIP1在柑橘果实脱落中的作用,本研究以转基因技术分别培育了CsHRP和CsWIP1的过量表达和RNAi转基因柑橘苗。目前已获得转基因抗性芽,现正在对转基因植株进行鉴定。
Developing citrus fruit will absciss under severe stresses, resulting in yield loses. Low winter temperature frequently causes pre-harvest fruit dropping and heavy loses. Citrus fruit are harvested by hands, which is very costly. For example, the harvesting cost has even surpassed the total production cost recently in USA. Therefore, achieving controllable fruit abscission is one of the main objectives being pursued by citrus growers and researchers.
This study used citrus genome array to probe ethylene-induced fruit abscission and and the underlying molecular mechanism. Ethylene at the concentration of 20μL/L was used to treat citrus mature fruit (Citrus sinensis cv. Valencia) for 4 h,12h and 24 h. Fruit abscission zones were collected and extracted for total RNA that was later analyzed by microarrays. Clustering analysis demonstrated that the replicability of the chip data were acceptable. qRT-PCR studies on 12 genes showed that their expression trends were similar to those showed by chip data. Results from analysis of chip data showed that there were 510 genes that were co-induced by ethylene at all the three sampling times (P＜0.01). Gene function assignment showed that the genes up-regulated by ethylene encode mostly membrane proteins, energy metabolism related proteins, stress-related proteins, transcription factors, serine/threonine kinases, phosphorylation enzymes, calcium-binding proteins, hydrolases, ethylene metabolic pathway proteins, transfer enzymes, and receptor proteins while those down-regulated encode ribosomal protein, hydrolytic enzymes, aquaporins, cytochrome P450, metal ion binding proteins, ABC transporters, lipase and so on. Further clustering showed that the genes encoding proteins involved in ethylene biosynthesis and signal transduction, in hypoxia stress and wounding responses were coordinately induced by ethylene. Therefore, ETR2, hypoxia responsive gene and wounding induced genes were selected for further study. The major results are as follows:
Hypoxia responsive gene CsHRP was cloned and analyzed for its expression. RACE was used to clone the full-lenth cDNA of the gene. The cloned cDNA was sequenced. Results showed that the full-length cDNA was 795 bp with an ORF encoding a peptide of 98 amino-acid residues. Sequence comparision showed that it shares 97%of similarity with a sweet orangen hypoxia responsive gene. Prediciton of the protein secondary structure suggested it might be related to plant resistance. Real-time PCR was used to quantify the gene's expression levels under exogenous ethylene treatment,and the results showed that that CsHRP was highly induced only at the early stage of ethylene treatment (4h), and treatment longer than 12h and 24h reversed its expression to the basal level.
Ethylene induced the expression of wounding-induced protein genes (WIPs). In this study 4 wounding-induced protein genes were cloned and designated as CsWIP1, CsWIP2, CsWIP3 and CsWIP4. Sequence comparison showed that they were strucuraly classified into two groups:group 1 contains CsWIP1, CsWIP2, CsWIP3 and group 2 contains only CsWIP4. WIPs generally contain two a-helices, a N-terninalα-helix and a C-ternimal a-helix. Simulation of these citrus wounding induced proteins demonstrated that CsWIP1, CsWIP2 and CsWIP3 are conservative in both helices while CsWIP4 is only conserved in its C-termianl helix. Real-time PCR results showed that CsWIP1 was expressed only under ethylene induction, indicating its ethylene dependence. CsWIP 2 was expressed only in leaves under both ethylene and wounding treatments, indicating its expression was tissue-specific. CsWIP 3 was universly expressed both in leaves and abscission zones either under ethylene treatment or under wounding treatment, indicating its expression was tissue-specific. CsWIP4 was only induced by wounding but not by ethylene.
ETR2 is a member of ethylene receptor gene family. To better understand the role of ETR2 in ethylene-induced fruit abscission, ETR2 was cloned and its expression was analyzed. Full-length cDNA of ETR2 was obtained by using RACE technology. The cloned cDNA was 1800bp in length. Sequence alignment analysis revealed that ETR2 of Valencia shares 98%homology with the same gene from other sweet orange(Citrus sinensis). Real-time PCR was used to detect the expression of ETR2 under exogenous ethylene treatment. The results showed that ETR2 was up-regulated by 2-folds and 4-folds after treatement with ethylene for 4h, and 12h, respectively, was reduced to its basal expression level after 24h of treatment.
To further characterize the CsHRP and CsWIP1, transgenic plants over expressing CsHRP and CsWIP1 have been obtained. Their RNAi transgenic plants have been alsoe obtained. Characterization of the transgenic plants is under way.
本研究利用柑橘基因组芯片(Citrus Genome Array)研究乙烯诱导柑橘果实脱落并探讨其分子机理。研究中使用浓度为20μL/L的乙烯分别处理带叶的伏令夏橙(Citrus sinensis cv. Valencia)果实4 h、12 h和24 h,然后提取果实离层RNA进行芯片检测。聚类分析结果表明,芯片数据具有较好的重复性。12条典型基因的Real Time-PCR分析结果也表明这些基因表达的变化趋势与芯片数据基本一致。对乙烯诱导的差异表达基因进行分析,发现乙烯处理3个时间点上均表达的基因有510条(P<0.01)。对差异表达基因进行功能分析,发现乙烯诱导下上调表达的主要有编码膜类蛋白、能量代谢相关蛋白、胁迫相关蛋白、转录因子、丝氨酸／苏氨酸激酶类、磷酸酶类、钙离子结合蛋白类、水解酶类、乙烯代谢途径相关蛋白、转移酶类、受体蛋白类等基因,下调表达的主要包括编码核糖体类蛋白、水解酶类、水通道蛋白、细胞色素P450、金属离子结合蛋白、光合系统相关蛋白、ABC转运蛋白、脂酶类等基因。进一步分析发现,编码乙烯代谢途径相关蛋白的基因、伤害诱导基因和编码低氧胁迫蛋白的基因之间存在着关联表达,因而筛选出ETR2、低氧应答基因和伤害诱导基因进行深入研究。主要结果如下：
克隆了乙烯诱导表达的低氧应答基因(CsHRP)并进行表达分析。采用RACE技术克隆了CsHRP的全长cDNA并测序分析。结果表明,CsHRP的cDNA全长为795 bp, ORF编码98个氨基酸。序列比对分析结果显示CsHRP与甜橙低氧应答基因的同源性达97%。对蛋白质二级结构的预测结果推测CsHRP可能与抗病有关。采用Real-time PCR方法检测了CsHRP在外源乙烯诱导下的表达情况,结果显示CsHRP在乙烯处理前期(4 h)高度表达,12 h后减低到原来的水平。
乙烯可诱导伤害诱导蛋白(Wound induced proteins, WIPs)的表达。本研究克隆了柑橘的4条伤害诱导基因CsWIP1、CsWIP2、CsWIP3和CsWIP4,序列分析结果表明这4个伤害诱导基因分为两类,其中CsWIP1、CsWIP2和CsWIP3聚为一类,而CsWIP4与前三者在结构上差异较大故聚为另一类。WIP蛋白在C-端有一个非常保守的α-螺旋,在N-端也有一个α-螺旋,CsWIP1、CsWIP2和CsWIP3在两个α-螺旋处都保守,而CsWIP4只在C-端α-螺旋处保守。Real-time PCR结果显示：CsWIP1仅在外源乙烯处理下表达,受乙烯诱导；CsWIP1无论是在伤害处理还是在乙烯处理条件下,仅在叶片中表达,说明CsWIP2的表达具有组织特异性;CsWIP3在乙烯或伤害处理的叶片和离层中均表达,说明CsWIP3的表达没有组织特异性；CsWIP4仅在伤害处理下表达,基本上不受乙烯调控。
ETR2是乙烯受体基因之一。为深入了解乙烯受体基因(ETR2)在伏令夏橙果实脱落过程中的作用,本研究克隆了ETR2基因并研究了其表达。通过RACE技术得到ETR2的cDNA序列为1800 bp。序列比对分析表明伏令夏橙ETR2与其它甜橙ETR2的同源性达98%。ETR2蛋白质包含了3个结构域,分别为GAF结构域、组氨酸蛋白激酶(HPK)同源二聚体结合区域和类似CheY结构域。通过Real-time PCR检测了ETR2在外源乙烯诱导4 h、12 h和24 h时的表达量。结果显示,ETR2在乙烯诱导4 h、12 h和24 h的表达量分别是对照的2倍、4倍和1倍。
为进一步验证CsHRP和CsWIP1在柑橘果实脱落中的作用,本研究以转基因技术分别培育了CsHRP和CsWIP1的过量表达和RNAi转基因柑橘苗。目前已获得转基因抗性芽,现正在对转基因植株进行鉴定。
Developing citrus fruit will absciss under severe stresses, resulting in yield loses. Low winter temperature frequently causes pre-harvest fruit dropping and heavy loses. Citrus fruit are harvested by hands, which is very costly. For example, the harvesting cost has even surpassed the total production cost recently in USA. Therefore, achieving controllable fruit abscission is one of the main objectives being pursued by citrus growers and researchers.
This study used citrus genome array to probe ethylene-induced fruit abscission and and the underlying molecular mechanism. Ethylene at the concentration of 20μL/L was used to treat citrus mature fruit (Citrus sinensis cv. Valencia) for 4 h,12h and 24 h. Fruit abscission zones were collected and extracted for total RNA that was later analyzed by microarrays. Clustering analysis demonstrated that the replicability of the chip data were acceptable. qRT-PCR studies on 12 genes showed that their expression trends were similar to those showed by chip data. Results from analysis of chip data showed that there were 510 genes that were co-induced by ethylene at all the three sampling times (P＜0.01). Gene function assignment showed that the genes up-regulated by ethylene encode mostly membrane proteins, energy metabolism related proteins, stress-related proteins, transcription factors, serine/threonine kinases, phosphorylation enzymes, calcium-binding proteins, hydrolases, ethylene metabolic pathway proteins, transfer enzymes, and receptor proteins while those down-regulated encode ribosomal protein, hydrolytic enzymes, aquaporins, cytochrome P450, metal ion binding proteins, ABC transporters, lipase and so on. Further clustering showed that the genes encoding proteins involved in ethylene biosynthesis and signal transduction, in hypoxia stress and wounding responses were coordinately induced by ethylene. Therefore, ETR2, hypoxia responsive gene and wounding induced genes were selected for further study. The major results are as follows:
Hypoxia responsive gene CsHRP was cloned and analyzed for its expression. RACE was used to clone the full-lenth cDNA of the gene. The cloned cDNA was sequenced. Results showed that the full-length cDNA was 795 bp with an ORF encoding a peptide of 98 amino-acid residues. Sequence comparision showed that it shares 97%of similarity with a sweet orangen hypoxia responsive gene. Prediciton of the protein secondary structure suggested it might be related to plant resistance. Real-time PCR was used to quantify the gene's expression levels under exogenous ethylene treatment,and the results showed that that CsHRP was highly induced only at the early stage of ethylene treatment (4h), and treatment longer than 12h and 24h reversed its expression to the basal level.
Ethylene induced the expression of wounding-induced protein genes (WIPs). In this study 4 wounding-induced protein genes were cloned and designated as CsWIP1, CsWIP2, CsWIP3 and CsWIP4. Sequence comparison showed that they were strucuraly classified into two groups:group 1 contains CsWIP1, CsWIP2, CsWIP3 and group 2 contains only CsWIP4. WIPs generally contain two a-helices, a N-terninalα-helix and a C-ternimal a-helix. Simulation of these citrus wounding induced proteins demonstrated that CsWIP1, CsWIP2 and CsWIP3 are conservative in both helices while CsWIP4 is only conserved in its C-termianl helix. Real-time PCR results showed that CsWIP1 was expressed only under ethylene induction, indicating its ethylene dependence. CsWIP 2 was expressed only in leaves under both ethylene and wounding treatments, indicating its expression was tissue-specific. CsWIP 3 was universly expressed both in leaves and abscission zones either under ethylene treatment or under wounding treatment, indicating its expression was tissue-specific. CsWIP4 was only induced by wounding but not by ethylene.
ETR2 is a member of ethylene receptor gene family. To better understand the role of ETR2 in ethylene-induced fruit abscission, ETR2 was cloned and its expression was analyzed. Full-length cDNA of ETR2 was obtained by using RACE technology. The cloned cDNA was 1800bp in length. Sequence alignment analysis revealed that ETR2 of Valencia shares 98%homology with the same gene from other sweet orange(Citrus sinensis). Real-time PCR was used to detect the expression of ETR2 under exogenous ethylene treatment. The results showed that ETR2 was up-regulated by 2-folds and 4-folds after treatement with ethylene for 4h, and 12h, respectively, was reduced to its basal expression level after 24h of treatment.
To further characterize the CsHRP and CsWIP1, transgenic plants over expressing CsHRP and CsWIP1 have been obtained. Their RNAi transgenic plants have been alsoe obtained. Characterization of the transgenic plants is under way.
引文
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