CTV强毒分离物对柑橘基因表达的影响及其与寄主互作蛋白的鉴定
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摘要
柑橘是一类重要的经济果树,全球年均贸易额已经达到90亿美元。由柑橘衰退病毒(Citrus tristeza virus, CTV)引起的柑橘衰退病被公认为柑橘栽培中最具威胁的病毒病害。病毒的成功侵染是病毒与寄主相互作用的结果,其在复制增殖过程中产生的病毒蛋白、核酸等分子引起寄主正常生理代谢的改变,并最终导致症状的产生。CTV侵染后能够诱导寄主植物表现茎陷点、苗黄、衰退等明显症状。然而,对于症状表现过程中CTV与寄主互作的分子机制仍然了解较少。为了分析CTV在转录和蛋白质水平上对寄主代谢造成的影响,本研究以CTV强毒株系N21为研究对象,对CTV侵染过程中寄主基因的转录应答反应和部分基因的时空表达动态进行了分析,并对CTV与寄主间的互作蛋白进行了筛选。取得的主要研究结果如下：
1、以感染CTV-N21并表现茎陷点的墨西哥株檬以及健康株檬植株为材料,采用抑制差减杂交技术(SSH)构建了一个库容量为14000个克隆的差减文库。通过反向Southern杂交筛选,共得到807个差异表达EST片段。测序及同源比对分析结果表明：这些EST片段代表430个寄主基因。其中上调表达基因282个,下调表达基因148个。使用荧光定量PCR技术对随机挑选25个基因在感染CTV-N21墨西哥株檬植株中的相对表达量进行的定量分析结果表明：正向文库中14个基因上调表达倍数为1.2-10倍,而反向文库中11个基因的下调表达倍数为1.1-7倍。以差异表达1.5倍为选择标准,则正向和反向文库的阳性率均在80%以上。对这430个基因进行功能注释并依据分子功能将其分为9类。其中,主要代谢和蛋白质代谢所占比例最大,分别占基因总数的26%和15.1%。在拟南芥数据库Tair中对这430个基因进行比对后共找到351个拟南芥同源基因。蛋白质互作预测结果显示这些基因编码产物在拟南芥代谢中形成复杂的互作网络。所占比例最大的互作来源于能量代谢。
2、对6个防御以及能量代谢相关基因在墨西哥株檬不同部位叶片组织中随CTV侵染时间(17-177dpi)的相对表达量变化进行了分析。3个防御相关基因分别编码几丁质酶(acidic class Ⅱ chitinase)、奇果类蛋白(miraculin-like protein)和Rsdv抗性类似蛋白(rsdvl resistance like protein);3个能量代谢相关基因编码蛋白为：碳酸酐酶(carbonic anhydrase)、1,5-二磷酸核酮糖脱氢酶(ribulose-1,5-bisphosphate carboxylase/oxygenase, RuBisCO)以及光捕捉复合体蛋白Lhca4(light-harvesting complex Ⅰ protein Lhca4)。荧光定量PCR分析结果显示：这些基因的表达量随CTV-N21侵染时间的延长表现出较大的波动性,在上部和下部叶片组织中的表达也存在差异。几丁质酶和奇果类蛋白基因均在接种后97d达到上调表达最大值。但两者在不同部位叶片中的表达量均存在差异；Rsdv抗性类似蛋白编码基因在上部叶片中表现为下调表达且呈总表达量呈持续下降趋势,而在下部叶片中则在接种后137d后表现为持续上调表达。光捕捉复合体蛋白编码基因在上部和下部叶片中的诱导表达量总体均呈下降趋势,并在接种后137d表现为下调表达。碳酸酐酶和RuBisCO编码基因总体呈下调表达,两者在不同部位叶片中的表达量存在较大差异。
3、通过RT-rPCR和体外融合方法分别构建了墨西哥株檬和枳壳的cDNA质粒文库,并利用酵母双杂交技术对CTV与寄主互作蛋白进行了筛选。初步筛选得到与病毒蛋白p20和p23互作的3个寄主蛋白。其在拟南芥中的同源蛋白分别为：碳硫裂合酶(Carbon-sulfur lyases)、蛋白质磷酸化酶(Protein phosphatase)和同源结构域亮氨酸拉链家族蛋白(Homeodomain leucine zipper family IV protein, HD-ZIP)。 BiFC荧光观察结果显示：p20与HD-ZIP沿细胞壁能够产生明显的互作信号,并在部分区域大量积累呈亮斑状。显示其互作可能发生在胞间连丝或细胞质特殊区域。
Citrus is one of the most economically important fruit tree speciese in the world. Its annual global trade value has reached nine billion dolars. Citrus tristeza virus is the causal agent of citrus tristeza, which was considered as the most devastating virus disease in citrus industry. The successful infection of a plant virus depends on the interaction between the virus and its host. The accumulation of virus in the process of infection disturbs the normal metabolism of host and result in the symptom exhibition. The infection of CTV could cause distinct symptoms like stem pitting, seedling yellow and tristeza on its host. However, the mechanism underlying the symptom development induced by CTV is still poorly understood, In this research, the transcriptional changes caused by a CTV severe isolate named N21were evaluated by using SSH, and the expression profiles of some candidate genes associated with defense and energy metabolism were analysed during a time course of CTV infection in leaves at different development stage. The proteins involved in the interactions between CTV and its suscept and resistance hosts were also identified. The main results obtained in this research were list as follows:
1. The transcriptional changes of Mexican lime response to CTV-N21infection were identified by SSH and two differentially expressed gene libraries consisted of14000clones were constructed. Screened by reverse southern blot, a total of774clones,501from the forward library and273from the reverse library, showing significant differential hybridization signals were selected and sequenced. A search for homologous genes of the selected EST sequences was performed by using tool of BlastX in NCBI database and the result showed that the657ESTs represent430genes, of which282were from forward library and148were from reverse library. The relative expression levels of25genes,14from forward-and11from reverse-library, in CTV-N21infected lime plants to that in mock inoculated lime were analyzed by qRT-PCR. Results showed that the relative expression levels of14genes from the forward library were up-regulated for1.2-10fold,11genes from the reverse library were down-regulated for1.1-7fold. According to a criterion of fold change≥1.5for positives, the credibility of all differentially expressed ESTs identified by SSH was above80%. According to GO annotation of molecular function, these430genes could be classified into nine main functional categories. The largest gene set involved in primary metabolism (26.0%), and then in protein metabolism (15.1%).351genes of Arabidopsis were found homologous to the screened430genes by BlastX search and their encoding proteins showed a complicated interaction network. The largest set of interactions was involved in energy metabolism.
2. Expression profile of six genes in upper and lower leaves were analysed by real-time PCR during a time course of17to177dpi with CTV-N21inoculation. Three of these genes including carbonic anhydrase, ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBisCO) and light-harvesting complex Ⅰ protein Lhca4were functionally involved in energy metabolism and other three genes including acidic class Ⅱ chitinase, miraculin-like protein and rsdvl resistance like protein were involved in defense and response to stress. The expression of these genes fluctuated during the infection of CTV-N21and the expression profiles differed in upper and lower leaves. The two genes encoding miraculin-like protein and acidic class Ⅱ chitinase had a similar induced expression tendency. Both of their expression peaked at97dpi, but showed different induction levels in upper and lower leaves. The expression level of the gene encoding a homolog of rsdvl resistance like protein showed to be persistantly repressed after CTV-N21infection in upper leaves. However, its expression in lower leaves was showed to be induced at137dpi. The induction of light-harvesting complex Ⅰ protein Lhca4gene showed consistently reduced in both upper and lower leaves, and turned to be repressed at137dpi. The expression of genes encoding carbonic anhydrase and RuBisCO was totally reduced during the infection of CTV-N21and showed a different expression profile in upper and lower leaves.
3. Moxican lime and C. trifoliata plasmid cDNA libraries were constructed by RT-rPCR and in vitro in fusion technology. The protein-protein interactions between virus and host were identified by cotransformation of the virus bait and library plasmid. The results indicated that CTV viral protein p20and p23may interact with three host proteins, which were homology to carbon-sulfur lyases, protein phosphatase, homeodomain leucine zipper family IV protein (HD-ZIP) in Arabidopsis. p20and HD-ZIP were shown interacted in BiFC analysis in Agrobacterium-infiltrated tobacco (Nicotiana benthamiana) leaves. The fluorescence signals resulted from their interaction accumulated and showed some bright spots along the cell wall, which implied that they may interact at cell wall and the bright spots may located at plasmodesmata.
1、以感染CTV-N21并表现茎陷点的墨西哥株檬以及健康株檬植株为材料,采用抑制差减杂交技术(SSH)构建了一个库容量为14000个克隆的差减文库。通过反向Southern杂交筛选,共得到807个差异表达EST片段。测序及同源比对分析结果表明：这些EST片段代表430个寄主基因。其中上调表达基因282个,下调表达基因148个。使用荧光定量PCR技术对随机挑选25个基因在感染CTV-N21墨西哥株檬植株中的相对表达量进行的定量分析结果表明：正向文库中14个基因上调表达倍数为1.2-10倍,而反向文库中11个基因的下调表达倍数为1.1-7倍。以差异表达1.5倍为选择标准,则正向和反向文库的阳性率均在80%以上。对这430个基因进行功能注释并依据分子功能将其分为9类。其中,主要代谢和蛋白质代谢所占比例最大,分别占基因总数的26%和15.1%。在拟南芥数据库Tair中对这430个基因进行比对后共找到351个拟南芥同源基因。蛋白质互作预测结果显示这些基因编码产物在拟南芥代谢中形成复杂的互作网络。所占比例最大的互作来源于能量代谢。
2、对6个防御以及能量代谢相关基因在墨西哥株檬不同部位叶片组织中随CTV侵染时间(17-177dpi)的相对表达量变化进行了分析。3个防御相关基因分别编码几丁质酶(acidic class Ⅱ chitinase)、奇果类蛋白(miraculin-like protein)和Rsdv抗性类似蛋白(rsdvl resistance like protein);3个能量代谢相关基因编码蛋白为：碳酸酐酶(carbonic anhydrase)、1,5-二磷酸核酮糖脱氢酶(ribulose-1,5-bisphosphate carboxylase/oxygenase, RuBisCO)以及光捕捉复合体蛋白Lhca4(light-harvesting complex Ⅰ protein Lhca4)。荧光定量PCR分析结果显示：这些基因的表达量随CTV-N21侵染时间的延长表现出较大的波动性,在上部和下部叶片组织中的表达也存在差异。几丁质酶和奇果类蛋白基因均在接种后97d达到上调表达最大值。但两者在不同部位叶片中的表达量均存在差异；Rsdv抗性类似蛋白编码基因在上部叶片中表现为下调表达且呈总表达量呈持续下降趋势,而在下部叶片中则在接种后137d后表现为持续上调表达。光捕捉复合体蛋白编码基因在上部和下部叶片中的诱导表达量总体均呈下降趋势,并在接种后137d表现为下调表达。碳酸酐酶和RuBisCO编码基因总体呈下调表达,两者在不同部位叶片中的表达量存在较大差异。
3、通过RT-rPCR和体外融合方法分别构建了墨西哥株檬和枳壳的cDNA质粒文库,并利用酵母双杂交技术对CTV与寄主互作蛋白进行了筛选。初步筛选得到与病毒蛋白p20和p23互作的3个寄主蛋白。其在拟南芥中的同源蛋白分别为：碳硫裂合酶(Carbon-sulfur lyases)、蛋白质磷酸化酶(Protein phosphatase)和同源结构域亮氨酸拉链家族蛋白(Homeodomain leucine zipper family IV protein, HD-ZIP)。 BiFC荧光观察结果显示：p20与HD-ZIP沿细胞壁能够产生明显的互作信号,并在部分区域大量积累呈亮斑状。显示其互作可能发生在胞间连丝或细胞质特殊区域。
Citrus is one of the most economically important fruit tree speciese in the world. Its annual global trade value has reached nine billion dolars. Citrus tristeza virus is the causal agent of citrus tristeza, which was considered as the most devastating virus disease in citrus industry. The successful infection of a plant virus depends on the interaction between the virus and its host. The accumulation of virus in the process of infection disturbs the normal metabolism of host and result in the symptom exhibition. The infection of CTV could cause distinct symptoms like stem pitting, seedling yellow and tristeza on its host. However, the mechanism underlying the symptom development induced by CTV is still poorly understood, In this research, the transcriptional changes caused by a CTV severe isolate named N21were evaluated by using SSH, and the expression profiles of some candidate genes associated with defense and energy metabolism were analysed during a time course of CTV infection in leaves at different development stage. The proteins involved in the interactions between CTV and its suscept and resistance hosts were also identified. The main results obtained in this research were list as follows:
1. The transcriptional changes of Mexican lime response to CTV-N21infection were identified by SSH and two differentially expressed gene libraries consisted of14000clones were constructed. Screened by reverse southern blot, a total of774clones,501from the forward library and273from the reverse library, showing significant differential hybridization signals were selected and sequenced. A search for homologous genes of the selected EST sequences was performed by using tool of BlastX in NCBI database and the result showed that the657ESTs represent430genes, of which282were from forward library and148were from reverse library. The relative expression levels of25genes,14from forward-and11from reverse-library, in CTV-N21infected lime plants to that in mock inoculated lime were analyzed by qRT-PCR. Results showed that the relative expression levels of14genes from the forward library were up-regulated for1.2-10fold,11genes from the reverse library were down-regulated for1.1-7fold. According to a criterion of fold change≥1.5for positives, the credibility of all differentially expressed ESTs identified by SSH was above80%. According to GO annotation of molecular function, these430genes could be classified into nine main functional categories. The largest gene set involved in primary metabolism (26.0%), and then in protein metabolism (15.1%).351genes of Arabidopsis were found homologous to the screened430genes by BlastX search and their encoding proteins showed a complicated interaction network. The largest set of interactions was involved in energy metabolism.
2. Expression profile of six genes in upper and lower leaves were analysed by real-time PCR during a time course of17to177dpi with CTV-N21inoculation. Three of these genes including carbonic anhydrase, ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBisCO) and light-harvesting complex Ⅰ protein Lhca4were functionally involved in energy metabolism and other three genes including acidic class Ⅱ chitinase, miraculin-like protein and rsdvl resistance like protein were involved in defense and response to stress. The expression of these genes fluctuated during the infection of CTV-N21and the expression profiles differed in upper and lower leaves. The two genes encoding miraculin-like protein and acidic class Ⅱ chitinase had a similar induced expression tendency. Both of their expression peaked at97dpi, but showed different induction levels in upper and lower leaves. The expression level of the gene encoding a homolog of rsdvl resistance like protein showed to be persistantly repressed after CTV-N21infection in upper leaves. However, its expression in lower leaves was showed to be induced at137dpi. The induction of light-harvesting complex Ⅰ protein Lhca4gene showed consistently reduced in both upper and lower leaves, and turned to be repressed at137dpi. The expression of genes encoding carbonic anhydrase and RuBisCO was totally reduced during the infection of CTV-N21and showed a different expression profile in upper and lower leaves.
3. Moxican lime and C. trifoliata plasmid cDNA libraries were constructed by RT-rPCR and in vitro in fusion technology. The protein-protein interactions between virus and host were identified by cotransformation of the virus bait and library plasmid. The results indicated that CTV viral protein p20and p23may interact with three host proteins, which were homology to carbon-sulfur lyases, protein phosphatase, homeodomain leucine zipper family IV protein (HD-ZIP) in Arabidopsis. p20and HD-ZIP were shown interacted in BiFC analysis in Agrobacterium-infiltrated tobacco (Nicotiana benthamiana) leaves. The fluorescence signals resulted from their interaction accumulated and showed some bright spots along the cell wall, which implied that they may interact at cell wall and the bright spots may located at plasmodesmata.
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