除虫菊CDS_CCI2基因的克隆、功能分析及遗传转化体系建立
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摘要
植物萜类物质在呼吸作用、光合作用、生长与发育等生理生化过程起着重要的作用。菊科植物中存在大量的萜类物质，并具有重要的药用价值。各种萜类次生物质的合成在细胞内主要有两种经典的途径：甲羟戊酸（MVA）途径和2-甲基-D-赤藻糖醇-4-磷酸（MEP）途径。
除虫菊酯，一种存在于除虫菊的天然萜类物质，具有高效的杀虫活性，由两种酸（菊酸和除虫菊酸）和三种酮醇（除虫菊酮醇、瓜叶菊醇和茉莉酮醇）经酯化作用形成结构六个不规则的单萜组分组成。除虫菊酯中的酸类物质是经MEP途径发生苯丙环反应生物合成，而酮醇是由起始物源于亚麻酸的羟脂（Oxylipin）途径合成。
目前除了固醇和类胡萝卜素外，对不规则类异戊二烯生物合成的分子机制的研究很少。2001年首次在除虫菊中发现菊酰焦磷酸合成酶（CDS），Rivera等证实CDS催化两分子DMAPP聚合成菊酸是生物合成除虫菊酯反应的第一步，但对其调控除虫菊酯生物合成的分子机制尚不明确。
因此，本研究从除虫菊花蕾中分离了除虫菊酯生物合成途径中的关键酶CDS_CCI2基因；通过亚细胞定位、时空表达模式、胁迫处理、转基因分析、VIGS技术等对CDS_CCI2基因功能进行了研究，初步建立了CDS_CCI2基因在除虫菊代谢中的调控模型；同时建立了除虫菊的高效遗传转化体系，为深入阐明除虫菊酯生物合成的分子机制奠定了重要的理论基础。主要研究结果如下：
1、以除虫菊cDNA为模板，通过PCR技术克隆得到除虫菊CDS_CCI2基因并测序验证，该基因包含1,188bp的开放阅读框，编码395个氨基酸（GenBank登录号No.HQ235057）。系统进化树分析表明，该基因属于除虫菊CDS家族。
2、构建CDS_CCI2基因与GFP报告基因相融合的瞬时表达载体，转化烟草细胞原生质体，在激光共聚焦显微镜下观察发现CDS_CCI2蛋白定位在质体中，从而参与除虫菊代谢调控过程。
3、采用荧光定量PCR方法对CDS_CCI2基因的组织表达模式进行了分析。CDS_CCI2基因在除虫菊根、茎、叶、花蕾、花中均有表达，其中在花蕾表达最强。
4、通过荧光定量PCR技术分析CDS_CCI2基因在不同的激素处理下表达特征，结果显示CDS_CCI2基因受到乙烯、茉莉酸甲酯（MeJA）以及脱落酸（ABA）正调控作用。
5、通过对CDS_CCI2转基因番茄的鉴定分析，CDS_CCI2超表达植株呈现矮化、果实变小、无籽等表型，并且改变了其内源代谢产物的水平，推测其原因是CDS_CCI2竞争消耗了底物菊酰焦磷酸，从而减少了脱落酸和赤霉素的生物合成。
6、通过VIGS技术在除虫菊中敲除CDS_CCI2基因的功能，结果表明CDS_CCI2基因沉默后，影响了叶绿素、类胡萝卜素以及除虫菊酯的生物合成。
7、以除虫菊幼嫩子叶为材料，利用根癌农杆菌（Agrobacterium tumefaciens）介导的叶盘法转化除虫菊，成功建立了除虫菊遗传转化体系。
Higher plants biosynthesize thousands of functional terpenoids via differentmetabolite pathways. These varied terpenoids play important roles in plant respiration,photosynthesis, regulation of growth and development. Irregular monoterpenes arecommon and important in the Asteraceae family. There are two classical metabolicpathways in different subcellular compartments, i.e. cytosolic mevalonate (MVA)pathway and2-C-methyl-D-derythritol-4-phosphate (MEP) pathway.
Pyrethrins are a natural insecticide biosynthesized by Tanacetum cinerariifolium (T.cinerariaefolium). Meanwhile, there are a set of six structurally close monoterpeneesters produced by two monoterpenic acids (pyrethric acid and chrysanthemic acid)with either of three ketone alcohols(cinerolone, jasmolone and pyrethrolone)esterification in pyrethrins. Acid moieties of pyrethrins are terpenoids synthesized witha cyclopropane ring via the MEP pathway (2-C-methyl-D-derythritol-4-phosphatepathway), while rethrolone moieties are synthesized from linolenic acid via the oxylipinpathway.
Up to now, little is known about the irregular isoprenoids synthesis at themolecular level except for the sterol and carotenoid pathways. Rivera et al. stated thatchrysanthemyl pyrophosphate in T. cinerariaefolium involved in the first reaction inpyrethrins synthesis by condensing two molecules of DMAPP with CDS. Even thoughCDS was firstly identified ten years ago, the molecular functions of T.cinerariaefoliumCDS have not yet been clearly clarified during pyrethrins biosynthesis.
In this study, we cloned a CDS_CCI2gene from T.cinerariaefolium cDNA, andinvestigated its subcellular localization, expression patterns in different tissues or undervarious stress treatments. Meanwhile, CDS_CCI2was overexpressed in tomato plantsand silenced by VIGS strategy in T.cinerariaefolium as an attempt to characterize itsbiological function. Furthermore, an efficient Agrobacterium tumefaciens-mediatedtransformation system in T.cinerariaefolium was established, which is fundamental totransgenic study on pyrethrin biosynthesis.
The main results and conclusions are as follows:
1. Chrysanthemyl diphosphate synthase (CDS) gene named CDS_CCI2(GenBankaccession No.HQ235057) was isolated from T.cinerariaefolium. The cDNA ofCDS_CCI2is in length with an1188bp open reading frame (ORF) encoding a polypeptide of395amino acids. CDS_CCI2was classed into CDS family byphylogenetic analysis of putative CDS_CCI2protein.
2. The subcellular localization of the CDS_CCI2protein was investigated byfusion to GFP under the control of the CaMV35S promoter, and was transientlyexpressed in tobacco protoplasts. Fluorescence microscopy analysis associated withimage overlapping techniques showed that CDS_CCI2was able to fully direct thefusion protein to the plastid, indicating that CDS_CCI2may have a regulatory functionin pyrethrin biosynthesis.
3. Quantitative RT-PCR was performed to determine the expression pattern ofCDS_CCI2gene in different tissues of T.cinerariaefolium. It revealed that CDS_CCI2transcripts were detected in all tested tissues (Roots, leaves, stems, flower buds, flowers)and the CDS_CCI2expression was utmost in flower buds as well.
4. The expression of CDS_CCI2gene was detected in T.cinerariaefolium seedlingsexposed to various abiotic stresses. The results revealed that CDS_CCI2transcriptswere up-regulated by abscisic acid (ABA), methyl jasmonate (MeJA) and ethreltreatment.
5. To investigate the physiological roles of CDS_CCI2in T.cinerariaefoliumdevelopment, the CDS_CCI2cDNA was overexpressed in tomato. Transgenic35S::CDS_CCI2lines displayed various morphological defects, such as dwarf growthhabit, reduced flower fertility, small fruit and parthenocarpy compared with wild-typeplants. The hypocotyls in35S::CDS_CCI2seedlings were shorter than those inwild-type seedlings. Overexpression of CDS_CCI2also altered the production ofendogenous secondary metabolites. Our data indicated that CDS_CCI2affected thebiosynthesis of GA and ABA.
6. To characterize CDS_CCI2functionally, a loss of function approach wasimplemented using the tobacco rattle virus (TRV)-mediated gene silencing (VIGS)strategy. The present study uncovered the role of CDS_CCI2in affecting chlorophyll,carotenoids and pyrethrins biosynthesis.
7. Establishing an efficient Agrobacterium tumefaciens-mediated transformationsystem for T.cinerariaefolium is fundamental to transgenic study on pyrethrinsbiosynthesis. The results revealed that cotyledons from tissue culture were excellentexplants for transformation.
除虫菊酯，一种存在于除虫菊的天然萜类物质，具有高效的杀虫活性，由两种酸（菊酸和除虫菊酸）和三种酮醇（除虫菊酮醇、瓜叶菊醇和茉莉酮醇）经酯化作用形成结构六个不规则的单萜组分组成。除虫菊酯中的酸类物质是经MEP途径发生苯丙环反应生物合成，而酮醇是由起始物源于亚麻酸的羟脂（Oxylipin）途径合成。
目前除了固醇和类胡萝卜素外，对不规则类异戊二烯生物合成的分子机制的研究很少。2001年首次在除虫菊中发现菊酰焦磷酸合成酶（CDS），Rivera等证实CDS催化两分子DMAPP聚合成菊酸是生物合成除虫菊酯反应的第一步，但对其调控除虫菊酯生物合成的分子机制尚不明确。
因此，本研究从除虫菊花蕾中分离了除虫菊酯生物合成途径中的关键酶CDS_CCI2基因；通过亚细胞定位、时空表达模式、胁迫处理、转基因分析、VIGS技术等对CDS_CCI2基因功能进行了研究，初步建立了CDS_CCI2基因在除虫菊代谢中的调控模型；同时建立了除虫菊的高效遗传转化体系，为深入阐明除虫菊酯生物合成的分子机制奠定了重要的理论基础。主要研究结果如下：
1、以除虫菊cDNA为模板，通过PCR技术克隆得到除虫菊CDS_CCI2基因并测序验证，该基因包含1,188bp的开放阅读框，编码395个氨基酸（GenBank登录号No.HQ235057）。系统进化树分析表明，该基因属于除虫菊CDS家族。
2、构建CDS_CCI2基因与GFP报告基因相融合的瞬时表达载体，转化烟草细胞原生质体，在激光共聚焦显微镜下观察发现CDS_CCI2蛋白定位在质体中，从而参与除虫菊代谢调控过程。
3、采用荧光定量PCR方法对CDS_CCI2基因的组织表达模式进行了分析。CDS_CCI2基因在除虫菊根、茎、叶、花蕾、花中均有表达，其中在花蕾表达最强。
4、通过荧光定量PCR技术分析CDS_CCI2基因在不同的激素处理下表达特征，结果显示CDS_CCI2基因受到乙烯、茉莉酸甲酯（MeJA）以及脱落酸（ABA）正调控作用。
5、通过对CDS_CCI2转基因番茄的鉴定分析，CDS_CCI2超表达植株呈现矮化、果实变小、无籽等表型，并且改变了其内源代谢产物的水平，推测其原因是CDS_CCI2竞争消耗了底物菊酰焦磷酸，从而减少了脱落酸和赤霉素的生物合成。
6、通过VIGS技术在除虫菊中敲除CDS_CCI2基因的功能，结果表明CDS_CCI2基因沉默后，影响了叶绿素、类胡萝卜素以及除虫菊酯的生物合成。
7、以除虫菊幼嫩子叶为材料，利用根癌农杆菌（Agrobacterium tumefaciens）介导的叶盘法转化除虫菊，成功建立了除虫菊遗传转化体系。
Higher plants biosynthesize thousands of functional terpenoids via differentmetabolite pathways. These varied terpenoids play important roles in plant respiration,photosynthesis, regulation of growth and development. Irregular monoterpenes arecommon and important in the Asteraceae family. There are two classical metabolicpathways in different subcellular compartments, i.e. cytosolic mevalonate (MVA)pathway and2-C-methyl-D-derythritol-4-phosphate (MEP) pathway.
Pyrethrins are a natural insecticide biosynthesized by Tanacetum cinerariifolium (T.cinerariaefolium). Meanwhile, there are a set of six structurally close monoterpeneesters produced by two monoterpenic acids (pyrethric acid and chrysanthemic acid)with either of three ketone alcohols(cinerolone, jasmolone and pyrethrolone)esterification in pyrethrins. Acid moieties of pyrethrins are terpenoids synthesized witha cyclopropane ring via the MEP pathway (2-C-methyl-D-derythritol-4-phosphatepathway), while rethrolone moieties are synthesized from linolenic acid via the oxylipinpathway.
Up to now, little is known about the irregular isoprenoids synthesis at themolecular level except for the sterol and carotenoid pathways. Rivera et al. stated thatchrysanthemyl pyrophosphate in T. cinerariaefolium involved in the first reaction inpyrethrins synthesis by condensing two molecules of DMAPP with CDS. Even thoughCDS was firstly identified ten years ago, the molecular functions of T.cinerariaefoliumCDS have not yet been clearly clarified during pyrethrins biosynthesis.
In this study, we cloned a CDS_CCI2gene from T.cinerariaefolium cDNA, andinvestigated its subcellular localization, expression patterns in different tissues or undervarious stress treatments. Meanwhile, CDS_CCI2was overexpressed in tomato plantsand silenced by VIGS strategy in T.cinerariaefolium as an attempt to characterize itsbiological function. Furthermore, an efficient Agrobacterium tumefaciens-mediatedtransformation system in T.cinerariaefolium was established, which is fundamental totransgenic study on pyrethrin biosynthesis.
The main results and conclusions are as follows:
1. Chrysanthemyl diphosphate synthase (CDS) gene named CDS_CCI2(GenBankaccession No.HQ235057) was isolated from T.cinerariaefolium. The cDNA ofCDS_CCI2is in length with an1188bp open reading frame (ORF) encoding a polypeptide of395amino acids. CDS_CCI2was classed into CDS family byphylogenetic analysis of putative CDS_CCI2protein.
2. The subcellular localization of the CDS_CCI2protein was investigated byfusion to GFP under the control of the CaMV35S promoter, and was transientlyexpressed in tobacco protoplasts. Fluorescence microscopy analysis associated withimage overlapping techniques showed that CDS_CCI2was able to fully direct thefusion protein to the plastid, indicating that CDS_CCI2may have a regulatory functionin pyrethrin biosynthesis.
3. Quantitative RT-PCR was performed to determine the expression pattern ofCDS_CCI2gene in different tissues of T.cinerariaefolium. It revealed that CDS_CCI2transcripts were detected in all tested tissues (Roots, leaves, stems, flower buds, flowers)and the CDS_CCI2expression was utmost in flower buds as well.
4. The expression of CDS_CCI2gene was detected in T.cinerariaefolium seedlingsexposed to various abiotic stresses. The results revealed that CDS_CCI2transcriptswere up-regulated by abscisic acid (ABA), methyl jasmonate (MeJA) and ethreltreatment.
5. To investigate the physiological roles of CDS_CCI2in T.cinerariaefoliumdevelopment, the CDS_CCI2cDNA was overexpressed in tomato. Transgenic35S::CDS_CCI2lines displayed various morphological defects, such as dwarf growthhabit, reduced flower fertility, small fruit and parthenocarpy compared with wild-typeplants. The hypocotyls in35S::CDS_CCI2seedlings were shorter than those inwild-type seedlings. Overexpression of CDS_CCI2also altered the production ofendogenous secondary metabolites. Our data indicated that CDS_CCI2affected thebiosynthesis of GA and ABA.
6. To characterize CDS_CCI2functionally, a loss of function approach wasimplemented using the tobacco rattle virus (TRV)-mediated gene silencing (VIGS)strategy. The present study uncovered the role of CDS_CCI2in affecting chlorophyll,carotenoids and pyrethrins biosynthesis.
7. Establishing an efficient Agrobacterium tumefaciens-mediated transformationsystem for T.cinerariaefolium is fundamental to transgenic study on pyrethrinsbiosynthesis. The results revealed that cotyledons from tissue culture were excellentexplants for transformation.
引文
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