CN109593738A - 秋葵类胡萝卜素生物合成和抗逆性相关蛋白AePDS及其编码基因与应用 - Google Patents
秋葵类胡萝卜素生物合成和抗逆性相关蛋白AePDS及其编码基因与应用 Download PDFInfo
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Classifications
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- C—CHEMISTRY; METALLURGY
- C12—BIOCHEMISTRY; BEER; SPIRITS; WINE; VINEGAR; MICROBIOLOGY; ENZYMOLOGY; MUTATION OR GENETIC ENGINEERING
- C12N—MICROORGANISMS OR ENZYMES; COMPOSITIONS THEREOF; PROPAGATING, PRESERVING, OR MAINTAINING MICROORGANISMS; MUTATION OR GENETIC ENGINEERING; CULTURE MEDIA
- C12N9/00—Enzymes; Proenzymes; Compositions thereof; Processes for preparing, activating, inhibiting, separating or purifying enzymes
- C12N9/0004—Oxidoreductases (1.)
- C12N9/001—Oxidoreductases (1.) acting on the CH-CH group of donors (1.3)
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- C—CHEMISTRY; METALLURGY
- C12—BIOCHEMISTRY; BEER; SPIRITS; WINE; VINEGAR; MICROBIOLOGY; ENZYMOLOGY; MUTATION OR GENETIC ENGINEERING
- C12N—MICROORGANISMS OR ENZYMES; COMPOSITIONS THEREOF; PROPAGATING, PRESERVING, OR MAINTAINING MICROORGANISMS; MUTATION OR GENETIC ENGINEERING; CULTURE MEDIA
- C12N15/00—Mutation or genetic engineering; DNA or RNA concerning genetic engineering, vectors, e.g. plasmids, or their isolation, preparation or purification; Use of hosts therefor
- C12N15/09—Recombinant DNA-technology
- C12N15/63—Introduction of foreign genetic material using vectors; Vectors; Use of hosts therefor; Regulation of expression
- C12N15/79—Vectors or expression systems specially adapted for eukaryotic hosts
- C12N15/82—Vectors or expression systems specially adapted for eukaryotic hosts for plant cells, e.g. plant artificial chromosomes (PACs)
- C12N15/8241—Phenotypically and genetically modified plants via recombinant DNA technology
- C12N15/8242—Phenotypically and genetically modified plants via recombinant DNA technology with non-agronomic quality (output) traits, e.g. for industrial processing; Value added, non-agronomic traits
- C12N15/8243—Phenotypically and genetically modified plants via recombinant DNA technology with non-agronomic quality (output) traits, e.g. for industrial processing; Value added, non-agronomic traits involving biosynthetic or metabolic pathways, i.e. metabolic engineering, e.g. nicotine, caffeine
- C12N15/825—Phenotypically and genetically modified plants via recombinant DNA technology with non-agronomic quality (output) traits, e.g. for industrial processing; Value added, non-agronomic traits involving biosynthetic or metabolic pathways, i.e. metabolic engineering, e.g. nicotine, caffeine involving pigment biosynthesis
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- C—CHEMISTRY; METALLURGY
- C12—BIOCHEMISTRY; BEER; SPIRITS; WINE; VINEGAR; MICROBIOLOGY; ENZYMOLOGY; MUTATION OR GENETIC ENGINEERING
- C12N—MICROORGANISMS OR ENZYMES; COMPOSITIONS THEREOF; PROPAGATING, PRESERVING, OR MAINTAINING MICROORGANISMS; MUTATION OR GENETIC ENGINEERING; CULTURE MEDIA
- C12N15/00—Mutation or genetic engineering; DNA or RNA concerning genetic engineering, vectors, e.g. plasmids, or their isolation, preparation or purification; Use of hosts therefor
- C12N15/09—Recombinant DNA-technology
- C12N15/63—Introduction of foreign genetic material using vectors; Vectors; Use of hosts therefor; Regulation of expression
- C12N15/79—Vectors or expression systems specially adapted for eukaryotic hosts
- C12N15/82—Vectors or expression systems specially adapted for eukaryotic hosts for plant cells, e.g. plant artificial chromosomes (PACs)
- C12N15/8241—Phenotypically and genetically modified plants via recombinant DNA technology
- C12N15/8261—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield
- C12N15/8271—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield for stress resistance, e.g. heavy metal resistance
- C12N15/8273—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield for stress resistance, e.g. heavy metal resistance for drought, cold, salt resistance
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- C—CHEMISTRY; METALLURGY
- C12—BIOCHEMISTRY; BEER; SPIRITS; WINE; VINEGAR; MICROBIOLOGY; ENZYMOLOGY; MUTATION OR GENETIC ENGINEERING
- C12Y—ENZYMES
- C12Y103/00—Oxidoreductases acting on the CH-CH group of donors (1.3)
- C12Y103/99—Oxidoreductases acting on the CH-CH group of donors (1.3) with other acceptors (1.3.99)
- C12Y103/99031—Phytoene desaturase (lycopene-forming) (1.3.99.31)
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- Life Sciences & Earth Sciences (AREA)
- Genetics & Genomics (AREA)
- Engineering & Computer Science (AREA)
- Chemical & Material Sciences (AREA)
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- Plant Pathology (AREA)
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- Nutrition Science (AREA)
- Breeding Of Plants And Reproduction By Means Of Culturing (AREA)
Abstract
本发明公开了一种秋葵类胡萝卜素生物合成关键酶蛋白AePDS及其编码基因在提高植物类胡萝卜素含量和耐盐性、抗旱性中的应用。本发明提供的蛋白，是如下(a)或(b)：(a)由序列表中SEQ ID NO.2所示的氨基酸序列组成的蛋白质；(b)将序列表中SEQ ID NO.2所示的氨基酸残基序列经过一个或几个氨基酸残基的取代和/或缺失和/或添加且与植物抗逆性相关的由序列SEQ ID NO.2衍生的蛋白质。实验证明，将所述蛋白的编码基因导入拟南芥中，转基因拟南芥中类胡萝卜素含量显著提高，同时提高转基因拟南芥植株的耐盐性和抗旱性。因此，类胡萝卜素生物合成和耐盐抗旱相关蛋白AePDS及其编码基因在调控植物类胡萝卜素生物合成以及耐盐抗旱性中具有重要的理论意义和实用价值；本发明将在农业领域具有广阔的应用空间和市场前景。
Description
技术领域
本发明属于生物技术领域，具体涉及秋葵类胡萝卜素生物合成和耐盐抗旱相关蛋白AePDS及其编码基因与应用。
背景技术
秋葵含有丰富的蛋白质、游离氨基酸、类胡萝卜素、各类维生素和磷、铁、钾、钙等矿质元素及由果胶和多糖等组成的粘性物质，具有多种保健功能，深受广大消费者欢迎。在动物中，类胡萝卜素物质也起着尤为重要的作用，但动物自身不能合成类胡萝卜素，只能从日常饮食中摄取。研究表明，类胡萝卜素与人类的身体健康有着密切的关联，是人类饮食结构中不可缺少的营养物质，在猝灭自由基、增加人体免疫力、预防心脑血管疾病和预防癌症、眼睛保护等方面具有非常重要的作用。β-胡萝卜素是人类及动物体内维生素A合成的前体。叶黄素和玉米黄质是人体视网膜黄斑的主要构成成分，可以预防老年人群中由黄斑病变所引起的失明。叶黄素和玉米黄质还具有消除自由基、抗衰老、抗癌和抗心血管疾病等作用，对人体健康非常有意义。
高等植物中类胡萝卜素的合成起始于异戊二烯焦磷酸(IPP)途径。三个IPP分子和一个二甲基丙烯基焦磷酸(DMAPP)分子在牻牛儿基牻牛儿基焦磷酸(GGPP)合酶(GGPS)催化下缩合形成C20 GGPP。GGPP是多种物质生物合成的共同前体，是形成植物类胡萝卜素最直接的前体。2个GGPP分子在八氢番茄红素合成酶(PSY)作用下缩合形成一个C40八氢番茄红素。八氢番茄红素在八氢番茄红素脱氢酶(PDS)、ζ-胡萝卜素脱氢酶(ZDS)和类胡萝卜素异构酶(CRTISO)3个酶共同催化下形成番茄红素。番茄红素经过两次番茄红素β-环化酶(LCYB)环化作用生成β-胡萝卜素；番茄红素在LCYB和番茄红素ε-环化酶(LCYE)环化形成α-胡萝卜素。α-胡萝卜素经过一步β-胡萝卜素羟化酶(BCH)羟化反应生成β-隐黄质，再经过一步BCH羟化反应形成玉米黄质。α-胡萝卜素经过BCH和ε-胡萝卜素羟化酶(ECH)两步羟化反应生成叶黄素。
世界上存在大面积的盐渍化的土地。据统计，全世界共有8亿hm2盐碱地，在灌溉地区还有占耕地面积33％的次生盐渍化土地，土壤的盐溃化严重影响现代农业的发展。就我国而言，在全国18亿亩耕地中有近十分之一的次生盐溃化土地，另外还有2000万hm2盐碱荒地。一般来说，盐浓度在0.2％-0.5％会影响作物的生长，但是盐碱地的盐分大都在0.6％-10％。大面积的盐渍化土地的存在严重影响了粮食生产，成为限制农业生产的主要因素。
因此，克隆秋葵类胡萝卜素生物合成关键基因AePDS，利用基因工程技术提高植物类胡萝卜素含量，同时改良植物耐盐性和抗旱性，培育高类胡萝卜素含量的秋葵新品种是改良秋葵营养保健功效的重要途径。
发明内容
本发明的一个目的是提供一种与秋葵类胡萝卜素积累和抗逆性相关蛋白及其编码基因和应用。
本发明所提供的与秋葵类胡萝卜素合成和耐盐抗旱相关蛋白，名称为AePDS，来源于秋葵(Abelmoschus esculentus)，是如下(a)或(b)的蛋白质：
(a)由序列表中SEQ ID NO.2所示的氨基酸序列组成的蛋白质；
(b)将序列表中SEQ ID NO.2所示的氨基酸序列经过一个或几个氨基酸残基的取代和/或缺失和/或添加且与植物类胡萝卜素积累和抗逆性相关的由SEQ ID NO.2衍生的蛋白质。
所述的SEQ ID NO.2由561个氨基酸残基组成。
编码上述蛋白的基因也属于本发明的保护范围。
所述基因是如下(a1)或(a2)或(a3)或(a4)所示的DNA分子：
(a1)编码区包括序列表中如SEQ ID NO.1所示核苷酸序列的DNA分子；
(a2)核苷酸序列是序列表中如SEQ ID NO.1所示的DNA分子；
(a3)在严格条件下与(a1)或(a2)限定的DNA序列杂交且编码与植物类胡萝卜素积累和抗逆性相关蛋白的DNA分子；
(a4)与(a1)或(a2)限定的DNA序列至少具有70％、至少具有75％、至少具有80％、至少具有85％、至少具有90％、至少具有95％、至少具有96％、至少具有97％、至少具有98％或者至少具有99％同源性且编码与植物类胡萝卜素积累和抗逆性相关蛋白的DNA分子。
所述的SEQ ID NO.1由1686个碱基组成，其开放阅读框(ORF)为自5′末端起第1位-第1686位碱基，编码的氨基酸序列是序列表中SEQ ID NO.2所示的蛋白。
上述的严格条件可为用6×SSC，0.5％SDS的溶液，在65℃下杂交，然后用2×SSC，0.1％SDS和1×SSC，0.1％SDS各洗膜一次。
含有编码上述蛋白的DNA分子或上述基因的表达盒、重组表达载体、转基因细胞系或重组微生物也属于本发明的保护范围。
所述重组表达载体是在载体pCBGUS的多克隆位点间插入所述编码基因得到的重组表达载体；
所述载体pCBGUS是通过包括如下步骤的方法得到的：
(1)将pCAMBIA1301载体经过HindIII和EcoRI双酶切，回收载体大片段；
(2)将pBI121载体经过HindIII和EcoRI双酶切，回收包含gusA基因的片段；
(3)将步骤(1)中回收的载体大片段与步骤(2)中回收的包含gusA基因的片段连接，得到重组载体pCBGUS。
所述pCAMBIA1301载体购自CAMBIA公司；所述pBI121载体购自Clontech公司。
扩增上述DNA分子全长或其任一片段的引物对也属于本发明的保护范围。
所述引物对为如下所示：
GSP-1：5’-CAGACGTTCTACCTGCACCC-3’
GSP-2：5’-GTTTTCCCAGTCACGAC-3’
GSP-3：5’-AGCATTCGTGGGATCAAGTC-3’
GSP-4：5’-CTCACCACCCAGCGATTCAA-3’
GSP-5：5’-ATGAGTCTTTCTGGGAGTGTTTCT-3’
GSP-6：5’-TCAGTGTCTGCTTGCTCCAG-3’
如下(b1)或(b2)或(b3)或(b4)的应用也是本发明保护的范围：
(b1)上述蛋白质，或，上述基因，或，含有上述基因的表达盒、重组载体、重组微生物或转基因细胞系，在调控植物中类胡萝卜素含量中的应用；
(b2)上述蛋白质，或，上述基因，或，含有上述基因的表达盒、重组载体、重组微生物或转基因细胞系，培育类胡萝卜素含量改变的转基因植物中的应用；
(b3)上述蛋白质，或，上述基因，或，含有上述基因的表达盒、重组载体、重组微生物或转基因细胞系，在调控植物抗逆性中的应用；
(b4)上述蛋白质，或，上述基因，或，含有上述基因的表达盒、重组载体、重组微生物或转基因细胞系，在培育抗逆性改变的转基因植物中的应用。
上述的抗逆性改变具体为耐盐性和/或抗旱性提高；
上述的调控植物抗逆性具体为提高植物耐盐性和/或抗旱性。
上述的调控植物中类胡萝卜素含量具体为提高植物中类胡萝卜素的含量；
上述的植物中类胡萝卜素含量改变具体为类胡萝卜素含量提高。
本发明的另一个目的是提供一种培育类胡萝卜素含量和/或抗逆性高的转基因植物的方法。
本发明所提供的培育类胡萝卜素含量和/或抗逆性高的转基因植物的方法，为提高目的植物中上述蛋白质的含量或活性，得到转基因植物；
所述转基因植物的类胡萝卜素含量和/或抗逆性高于所述目的植物。
上述提高目的植物中上述蛋白质的含量或活性为提高目的植物中编码上述蛋白质的DNA分子的表达，具体为将编码上述蛋白质的DNA分子导入目的植物中，得到转基因植物；上述导入方式为通过重组表达载体导入。
本发明第三个目的是提供一种培育类胡萝卜素含量和/或抗逆性高的植物的方法。
本发明提供的方法，为培育上述方法得到的转基因植物。
上述方法中，所述抗逆性为耐盐性和/或抗旱性。
上述的类胡萝卜素含量具体由HPLC方法测定叶片中的类胡萝卜素含量增加来体现。
上述的耐盐性由增加植物根长和鲜重、增加SOD活性、增加POD活性、增加ABA含量、增加脯氨酸含量、降低过氧化氢(H2O2)含量和降低丙二醛(MDA)含量体现。
上述的抗旱性由增加植物根长和鲜重、增加SOD活性、增加POD活性、增加ABA含量、增加脯氨酸含量、降低过氧化氢(H2O2)含量和降低丙二醛(MDA)含量体现。
上述的类胡萝卜素含量为叶黄素、玉米黄质、β-胡萝卜素、α-胡萝卜素、β-隐黄质和/或总类胡萝卜素含量。
上述的植物是如下(c1)至(c4)中的任一种：
(c1)双子叶植物；
(c2)单子叶植物；
(c3)十字花科植物；
(c4)拟南芥。
本发明的实验证明，本发明发现了AePDS蛋白及其编码基因，将该基因导入拟南芥中，得到过表达AePDS基因的拟南芥植株。测定转基因植株中类胡萝卜素含量，发现叶片中的α-胡萝卜素、叶黄素、β-胡萝卜素、β-隐黄质、玉米黄质和总类胡萝卜素含量显著高于野生型拟南芥植株；将转基因拟南芥植株进行盐、旱胁迫处理，发现过表达株系与野生型拟南芥相比，耐盐性和抗旱性增强，具体体现在增加SOD活性、POD活性、ABA含量、脯氨酸含量和降低H2O2含量、MDA含量。结果表明，AePDS基因及其所编码的蛋白在植物提高类胡萝卜素含量以及对抗高盐和干旱过程中起着重要的作用。本发明所提供的AePDS蛋白及其编码基因在提高植物类胡萝卜素含量以及耐盐抗旱性研究中具有重要的应用价值。本发明将在农业领域具有广阔的应用空间和市场前景。
附图说明
图1本发明秋葵AePDS基因植物表达载体简图。
图2本发明AePDS转基因拟南芥植株的PCR检测结果图。
图3本发明AePDS基因在过表达拟南芥株系和野生型拟南芥植株中的表达。
图4本发明中叶黄素(lutein)、玉米黄质(zeaxanthin)、β-隐黄质(β-cryptoxanthin)、α-胡萝卜素(α-carotene)和β-胡萝卜素(β-carotene)的标准曲线图。
图5本发明AePDS转基因拟南芥植株在200mMNaCl和25％PEG6000的MS培养基上的生长和生根情况，WT为野生型拟南芥植株，L3和L4为转基因拟南芥植株。
图6本发明AePDS转基因拟南芥植株耐盐性和抗旱性盆栽鉴定，WT为野生型拟南芥植株，L3和L4为转基因拟南芥植株。
图7本发明AePDS转基因拟南芥植株抗逆生理生化指标测定，WT为野生型拟南芥植株，L3和L4为转基因拟南芥植株。
具体实施方式
下面结合具体实施例对本发明作进一步阐述，但不限制本发明。
下述实施例中，所用的试验材料及其来源包括：
秋葵(Abelmoschus esculentus)品种台湾五福，由淮阴工学院生命科学与食品工程学院江苏省植物生产与加工实践教育中心实验室保存。
大肠杆菌(Escherichia coli)DH5α由淮阴工学院生命科学与食品工程学院江苏省植物生产与加工实践教育中心实验室保存。克隆载体PMD-18-Simple T、各类限制性内切酶、Taq聚合酶、连接酶、dNTP、10×PCR buffer和DNA marker购自宝生物工程大连有限公司。所有的化学试剂都从美国西格玛化学公司和上海国药化学试剂公司购买。
本发明中常规的分子生物学操作具体参见《分子克隆》【Molecular Cloning.2nded.Cold Spring Harbor Laboratory Press,1989】。
下述实施例中常规的基因操作参照分子克隆文献进行【Sambook J,Frets EF,Mannsdes T et al.In:Molecular Cloning.2nd ed.Cold Spring Harbor LaboratoryPress,1989】。
实施例1秋葵AePDS蛋白及其编码基因的获得
1.实验材料
参照王旭等(2014)【王旭，韩春乐，周亚楠，王春国，宋文芹，陈成彬.黄秋葵查尔酮合成酶基因AeCHS的克隆与表达分析.植物遗传资源学报，2014，15(3)：561-567】的方法，将秋葵品种台湾五福植物叶片材料取下，液氮速冻，-80℃保存。
2.叶片总RNA提取和纯化
取台湾五福叶片约2.0g，在液氮中研磨成粉状，加入10mL离心管，用Applygen植物RNA提取试剂盒(Applygen Technologies Inc，Beijing)提取叶片总RNA，试剂盒中包括：Plant RNA Reagent，植物组织裂解、分离RNA、去除植物多糖和多酚；Extraction Reagent，有机抽提去除蛋白质、DNA、多糖和多酚；Plant RNA Aid，去除植物多糖多酚和次生代谢产物。利用QIAGEN Oligotex Mini mRNA Kit(QIAGEN，GmbH，Germany)从总RNA中纯化mRNA。最后，取1μL于1.2％琼脂糖凝胶电泳检测其完整性，另取2μL稀释至500μL，用紫外分光光度计检测其质量(OD260)和纯度(OD260/OD280)，提取台湾五福叶片总RNA，经非变性胶琼脂糖凝胶电泳检测，28S和18S条带清晰，且二者亮度比值为1.5～2︰1，表明总RNA没有降解，纯化所得mRNA符合实验要求，可用于秋葵AePDS蛋白cDNA全长的克隆。
3.秋葵AePDS蛋白cDNA的全长克隆
以获得的AePDS基因的EST片段设计引物进行AePDS蛋白cDNA的全长克隆。
(1)3’-RACE
以台湾五福cDNA为模板，用AePDSEST正向引物GSP-1和反向引物GSP-2进行PCR反应。引物序列如下：
GSP-1：5’-CAGACGTTCTACCTGCACCC-3’
GSP-2：5’-GTTTTCCCAGTCACGAC-3’
PCR得到的3’RACE片段，回收后连接pMD19-T载体(购自北京六合通经贸有限公司，产品目录号为D102A)进行TA克隆，以BcaBESTTM Sequencing Primers/M13 Primers通用引物进行测序。
(2)5’-RACE
以台湾五福cDNA为模板，用AePDSEST正向引物GSP-3和反向引物GSP-4进行PCR反应。引物序列如下：
GSP-3：5’-AGCATTCGTGGGATCAAGTC-3’
GSP-4：5’-CTCACCACCCAGCGATTCAA-3’
PCR得到的5’RACE片段，回收后连接pMD19-T载体(购自北京六合通经贸有限公司，产品目录号为D102A)进行TA克隆，以BcaBESTTM Sequencing Primers/M13 Primers通用引物进行测序。
(3)PCR扩增AePDS蛋白cDNA的编码区
利用DNAMAN 7.0软件拼接候选的秋葵AePDS蛋白cDNA序列。进一步设计正向引物GSP-5和反向引物GSP-6进行PCR扩增AePDS蛋白cDNA的编码区。引物序列如下：
GSP-5：5’-ATGAGTCTTTCTGGGAGTGTTTCT-3’
GSP-6：5’-TCAGTGTCTGCTTGCTCCAG-3’
以台湾五福叶片总RNA经Oligo(dT)反转录为模板，用高保真的FastPfu酶，进行PCR扩增，PCR条件为95℃1min，随后95℃20s，53℃20s和72℃1min，进行40个循环，最后72℃延伸10min。琼脂糖凝胶电泳检测PCR扩增产物，获得1686bp长度的扩增片段。
综合上述步骤的结果，获得了目的cDNA序列，其核苷酸序列如序列表中序列SEQID NO.1所示。序列表中序列SEQ ID NO.1由1686个碱基组成，自5’端第1位-第1686位碱基为其开放阅读框，编码具有序列表中序列SEQ ID NO.2所示的氨基酸残基序列的蛋白质。序列表中序列SEQ ID NO.2由561个氨基酸残基组成。将该基因命名为AePDS，将其编码的蛋白命名为AePDS。
实施例2 AePDS基因过表达载体的构建
将实施例1中测序鉴定正确的含有序列表SEQ ID NO.1所示核苷酸的DNA片段用BamH I和Sac I进行双酶切，用1％琼脂糖凝胶回收DNA片段，通过T4DNA连接酶将回收的AePDS基因片段与含有双35S启动子的载体pCBGUS连接，酶切鉴定和序列分析测定获得了含有秋葵AePDS基因的重组植物表达载体pCAMBIA1301-AePDS。该表达载体还包含gusA报告基因和带内含子卡那霉素抗性标记基因，载体如图1所示。
实施例3 AePDS基因转化拟南芥
将实施例2构建的秋葵AePDS基因的植物表达载体pCAMBIA1301-AePDS用蘸花法转化拟南芥，具体方法如下：
1.农杆菌的准备
(1)将pCAMBIA1301-AePDS用电击法转化根癌农杆菌EHA105菌株(Biovector Co.,LTD)菌株，得到含有pCAMBIA1301-AePDS的重组农杆菌，并涂布于含有卡那霉素抗性的平板筛选转化子。
(2)挑取农杆菌单菌接种于5mL LB液体培养基(利福平50μg/mL，氯霉素100μg/mL)中，28℃，250rpm培养20h。
(3)取1mL菌液转接入20-30mL LB液体培养基(利福平50μg/mL，氯霉素100μg/mL)中，28℃，250rpm培养约12h，测OD 600≈1.5。
(4)8000rpm，4℃，10min离心收集菌体，重悬于农杆菌转化渗透液(5％蔗糖，0.05％Silwet L-77)并稀释至OD 600≈0.8。
2.拟南芥蘸花法转化
(1)将拟南芥的花薹浸入上述侵染液中，轻轻搅动约10s后取出，全部转化完毕后，用保鲜袋罩住拟南芥，以保持湿润环境，水平放置，22℃避光培养，24h后去掉保鲜袋直立培养。
(2)初次转化4d后，可再进行一次转化，重复两次，总共转化三次，这样可以对花序上发育的不同时期的花蕾进行转化，提高转化效率。
(3)生长约两个月后，收集种子，4℃冰箱储存待用。
经过蘸花法转化的拟南芥生长约两个月后，正常开花结子。
实施例4 AePDS基因转基因拟南芥植株分子检测
1.转基因拟南芥种子的筛选
(1)称25-30mg种子放入1.5mL离心管。
(2)1mL 75％乙醇消毒1min(不停摇晃振荡)，8000rpm离心5s，去上清。
(3)加入1mL过滤后的漂白粉(2.5％，g/100mL)消毒15min(不停摇晃振荡，充分消毒)，8000rpm离心5s，去上清。
(4)无菌水洗涤3-4次。
(5)将种子均匀的播撒到1/2MS平板(潮霉素50μg/mL)上，Parafilm膜封口，4℃冰箱放置两天，22℃，16h光照培养10天。
(6)将抗性植株移栽到盆中培养，苗稍大后，进行GUS活性检测，选出阳性植株(T1)培养至开花结实，收集T1植株上所结T2种子，进一步筛选得到T3种子。
2.转基因拟南芥植株PCR检测
(1)试验方法
用CTAB法提取T3拟南芥转基因植株和野生型植株的基因组DNA。用常规方法进行PCR检测，所使用的AePDS基因引物为：Primer 1：5’-TGCTTGTCCTTTGCAGGTAGT-3’和Primer2：5’-TCTGCTTGCTCCAGTCAACTT-3’。
(2)试验结果
电泳检测扩增结果见图2(图2中，泳道M为Maker；泳道W：水；泳道P：阳性对照(重组质粒pCAMBIA1301-AePDS)；泳道WT：野生型拟南芥植株；泳道L1-L10：为转化pCAMBIA1301-AePDS的拟南芥转基因植株)。从图中可见，转化pCAMBIA1301-AePDS的拟南芥拟转基因植株和阳性对照扩增出1537bp的目标条带，表明AePDS基因已经整合到拟南芥的基因组中，并证明这些再生植株为转基因植株；野生型拟南芥植株没有扩增出1537bp的目标条带。转基因植株为后续功能分析。
3.转基因拟南芥植株qRT-PCR检测
(1)试验方法
将阳性T3代转AePDS拟南芥株系提取RNA，反转录得到cDNA，进行qRT-PCR，以未转化的野生型为对照。AtActin基因为内参：AtActin-F：5’-GCACCCTGTTCTTCTTACCGA-3’和AtActin-R：5’-AGTAAGGTCACGTCCAGCAAGG-3’；AePDS引物序列为：AePDS-F：5’-GCGTACCTGATCGTGTGACT-3’和AePDS-R：5’-GGGTTGCCATCCAAGAATGC-3’。
(2)试验结果
结果如图3所示，WT为野生型拟南芥植株，L1、L2、L3、L4、L5、L6、L7、L8、L9、L10均为阳性T3代转AePDS拟南芥，表明AePDS在转基因拟南芥植株中有不同程度的表达。
实施例5高效液相色谱法测定AePDS基因转基因拟南芥植株叶片类胡萝卜素含量
1.标样的准备
(a)β-胡萝卜素(β-carotene)、玉米黄素(zeaxanthin)和叶黄素(lutein)标样购自sigma公司，商品号分别为C4582-10MG、14681-1、95507；β-隐黄质(β-cryptoxanthin)标样购自北京华美互利生化公司，商品号为0317S。
(b)α-胡萝卜素(α-carotene)标样：由于α-carotene标样极易降解，没有进行商业化标样，必须自行提取。具体方法如下：
(1)将胡萝卜丁放入食物料理机中打碎。
(2)将打碎的胡萝卜浆状物导入盛有5g硅藻土的大研钵中，混匀。
(3)加入适量的预冷丙酮用力研磨，将胡萝卜中的胡萝卜素萃取到丙酮中。
(4)将研磨液倒入磨砂漏斗中进行抽真空，黄色液体会被抽到三角瓶中，磨砂漏斗中的干燥物质用勺取出，再放入到大研钵中，加适量预冷丙酮再次用力研磨。重复5-6次，直到研磨液的颜色变为无色。
(5)将三角瓶中的金黄色液体(胡萝卜素萃取液)分若干次倒入分液漏斗中，每倒入一次胡萝卜素萃取液后，均倒入300mL的ddH2O，静置稍许，将下层透明液层排出到废液瓶中。
(6)胡萝卜素萃取液经过若干次ddH2O洗涤后，最后一次用饱和食盐水洗涤，以彻底除去所形成的乳状物层，同样排出下层的透明废液。
(7)用吸水纸擦干分液漏斗的出水孔。将金黄色的石油醚层(高纯度胡萝卜素萃取液)排出倒入一个干燥的锥形瓶中；加入适量无水硫酸钠(Na2SO4)干燥(至无水NaSO4结晶呈分散状态)，摇晃，吸取萃取液中的残留水分。
(8)将处理后的金黄色液体倒入一干燥的球形瓶中，再加入100mL 10％KOH(g/100ml)的甲醇溶液皂化和0.1％(g/100mL)二丁基羟基甲苯。
(9)将球形瓶接入旋转蒸发仪，旋转蒸发仪的另一接口接真空泵，真空蒸干液相石油醚，将有机相浓缩至大约5mL，之后用胶头滴管吸取石油醚(<5mL)以洗净球形瓶壁上的橙色固态物质，胶头滴管反复吹打橙色液相，使固态物质充分溶解，以获得更高纯度的胡萝卜素萃取液，工作时间约50min。
(10)柱填料硅藻土和氧化镁(1︰1)(质量比)于110℃活化4h，在干燥器里冷却、干燥。准备一个铁架台，将玻璃层析柱放置在铁架台上并固定。在玻璃层析柱下方接好一个带口三角瓶，三角瓶通过皮管连接真空泵。
(11)填柱：将高温激活的硅藻土氧化镁混合物小心倒入玻璃层析柱内，不时用柱塞棒小心压实，填柱约20cm左右，确保柱面水平；再在柱面上方加入细化无水硫酸钠层1cm，之后塞入一层1.5cm左右的脱脂棉层，保证硫酸钠层和脱脂棉层的相平。再次压实后，打开真空泵，抽真空1h。
(12)石油醚过柱：抽真空1小时后，不要关闭真空泵，沿柱壁加入石油醚润柱，调整真空泵，使流速为每秒2-3滴，并用柱塞棒保持固态面的水平，最终使石油醚润洗整个层析柱。
(13)用滴管将浓缩后的提取物小心转入层析柱中，至样品层进入到接近无水Na2SO4层时，再把冲洗圆底烧瓶的石油醚相转入层析柱中，过柱过程中要保证石油醚相始终高于无水Na2SO4层。
(14)层析工作过程中，一定要保证固相上方的石油醚液相的存在，石油醚一旦不足必须立即补充(层析柱需要避光，可以使用锡箔纸包裹)。
(15)α-胡萝卜素是第一个流出层析柱的物质，所以当分层的α-胡萝卜素层即将流出层析柱时，暂时关闭真空泵，更换新的带口三角瓶，以盛接α-胡萝卜素，连接好后，打开真空泵，金黄色的α-胡萝卜素会流出进入到三角瓶内(三角瓶同样需要避光)。
(16)将获得的金黄色液体倒入玻璃螺口离心管中保存，详细注明标样提取时间，parafilm严格密封，锡箔纸避光，-80℃冰箱直立保存。
(17)取出少量用N2气吹干(剩余的大量提取物密封、避光保存于-70℃，备用)，用1mL V乙腈︰V甲醇︰V二氯甲烷＝45︰20︰35的溶剂溶解。
(18)用1mL一次性注射器使溶质完全溶解后，通过0.22μL的过滤器转移至2mL棕色进样小瓶中，50μL进样检测提取样品的纯度。
2.标准品的配置以及标准曲线的绘制
将提取并检测的α-胡萝卜素(α-carotene)直接用于混合标样的制备；叶黄素(Lutein)标样用无水乙醇和乙醚溶解；玉米黄素(Zeaxanthin)用丙酮溶解；β-隐黄质(β-cryptoxanthin)标样用乙醚和石油醚溶解；β-胡萝卜素(β-catotene)标样则用石油醚进行溶解。将标准样品溶解后各取100μl至小容量瓶(5mL)定容，用紫外-可见光分光光度计在各成分特定波长下测定其吸光值。根据公式(1)计算浓度，公式(2)校对公式(1)计算所得浓度，用公式(3)配置成50mL混合标样。混合标样需经氮气浓缩后，用石油醚定容。分别取1mL、2mL、3mL和5mL混合标样并分别重复3次共45mL建立标准曲线，标样测定值满足表1所列各成分所在的浓度范围内即可进行标准曲线的绘制。混合标样中叶黄素(lutein)、玉米黄素(zeaxanthin)、β-隐黄质(β-cryptoxanthin)、α-胡萝卜素(α-carotene)和β-胡萝卜素(β-carotene)的标准曲线如图4所示。
式中OD为吸光值；为吸光系数；
校对浓度(μg/mL)＝浓度c×纯度(％) (2)
纯度(％)＝标样HPLC峰面积/HPLC峰总面积×100
a＝(50×b)/c (3)
式中50为混合标样总体积50mL；a为加入标样量(μg/mL)
b为浓度范围中值(μg/mL)；c为校对浓度(μg/mL)
表1标准溶液吸光系数和浓度范围
3.转基因拟南芥植株类胡萝卜素提取
拟南芥叶片：取移栽至营养土中2w的阳性T3代转AePDS拟南芥植株和野生型拟南芥植株的叶片，液氮速冻，然后磨成粉末状，各称取0.6g左右磨好的样品进行类胡萝卜素的提取。方法如下：
(1)对阳性T3代转AePDS拟南芥和野生型拟南芥植株样品进行磨样；
(2)称取0.6g磨好样品于25mL螺口玻璃离心管中，加入6mL 0.1％BHT(g/100ml)无水乙醇，涡旋20s；
(3)取出加120μL 80％(g/100ml)氢氧化钾溶液；
(4)涡旋20s，再放入85℃水浴5min；
(5)取出，再涡旋20s，再放入85℃水浴5min；
(6)取出后立即置于冰上，立即加3mL预冷ddH2O；
(7)加3mL正己烷，涡旋20s；
(8)2700rpm离心5min，用移液枪吸取上清至另一新的螺口玻璃离心管中；
(9)重复7,8步3次，将上清至另一新的螺口玻璃离心管中，终体积达到约12mL；
(10)在有上清液的新的螺口玻璃离心管中加3mL预冷ddH2O，涡旋，2700rpm离心5min；
(11)用移液枪吸取上层溶液(正己烷层)至新的尖底玻璃离心管中；
(12)再在水相的螺口管中加入3mL正己烷，涡旋，离心5min，再吸取上层正己烷层至新的尖底玻璃管中，重复2次；
(13)真空离心干燥总的正己烷；
(14)在干燥后的尖底玻璃离心管中加入1mL流动相，吸打混匀后通过0.22μm过滤器小心将液体加入棕色样品瓶中，用高效液相色谱法(HPLC)进行测定。
4.转基因拟南芥植株类胡萝卜素测定
(1)试验方法
利用高效液相色谱法测定(高效液相色谱仪为美国Agilent公司1200型)，方法如下：
(1)将上述得到的待测类胡萝卜素溶液在旋转蒸发器中集中收集，并用N2干燥。
(2)立即使类胡萝卜素溶解于1mL乙腈︰甲醇︰二氯甲烷(V︰V︰V)＝45︰20︰35的的溶剂中。
(3)将待测样品通过0.22μL注射器过虑后，直接将10μl样品到加入到色谱柱中。
(4)采用YMC C30色谱柱(250mm×4.6mm，5nm)，以乙腈︰甲醇︰二氯甲烷(V︰V︰V)＝75︰20︰5为流动相，流速1.8mL/min检测波长450nm处的吸收峰的变化。
(5)每个待测样品，重复测定3次。
(2)试验结果
根据上述建立的叶黄素(lutein)、玉米黄素(zeaxanthin)、β-隐黄质(β-cryptoxanthin)、α-胡萝卜素(α-carotene)和β-胡萝卜素(β-carotene)的标准曲线(图4)，分别测定转基因拟南芥叶片、转空载体对照拟南芥叶片和野生型对照拟南芥叶片的类胡萝卜素含量。其中总类胡萝卜素含量是五种类胡萝卜素含量之和。结果如表2所示，表2中L3、L4分别表示2个转基因拟南芥植株叶片的待测样品；CK代表转空载体对照拟南芥植株叶片；WT代表野生型拟南芥植株叶片。转空载体对照拟南芥植株叶片(CK)和野生型对照拟南芥植株叶片(WT)中的类胡萝卜素含量无显著差异；转基因拟南芥植株的叶黄素、玉米黄素、β-隐黄质、α-胡萝卜素、β-胡萝卜素、总类胡萝卜素含量与野生型拟南芥植株相比显著提高；叶黄素含量分别是野生型植株的1.35倍和1.21倍；玉米黄素含量分别是野生型植株的1.13倍和1.23倍；β-隐黄质含量分别是野生型植株的1.26倍和1.17倍；α-胡萝卜素含量分别是野生型植株的1.19倍和1.10倍；β-胡萝卜素含量分别是野生型植株的1.89倍和1.77倍；总类胡萝卜素含量分别是野生型植株的1.51倍和1.38倍；表明导入AePDS基因显著提高植物类胡萝卜素积累。
表2 AePDS转基因拟南芥植株叶片中类胡萝卜素含量
实施例6AePDS基因转基因拟南芥植株耐盐性和抗旱性鉴定
1.转基因植株耐盐性和抗旱性离体鉴定
(1)试验方法
将转基因拟南芥和野生型种子，消毒灭菌后播种继代培养于200mMNaCl和25％PEG6000(g/100mL)的1/2MS培养基上，胁迫培养2周后，观察拟南芥植株的生长状态和生根情况。
(2)试验结果
结果显示，转基因拟南芥植株的生长状态和生根情况显著优于野生型植株，转基因植株根长和植株鲜重均显著优于野生型植株(图5)，表明转基因植株的耐盐性和抗旱性较野生性有显著提高。
2.转基因植株耐盐性和抗旱性盆栽鉴定
(1)试验方法
将转基因拟南芥和野生型种子在1/2MS培养基上培养2周后，将植株移栽到盆中培养2周后，进行盐、干旱胁迫处理。用含有300mM NaCl的1/2霍格兰营养液每个2天灌溉1次，每次200mL，处理4周，观察植株生长情况并统计存活率；干旱处理6周后，观察植株生长情况并统计存活率。
(2)试验结果
结果显示，通过耐盐性和抗旱性盆栽鉴定，结果见图6，盐处理4周或干旱处理6周后，转基因植株的生长状态显著优于野生型植株，转基因植株的存活率显著高于野生性植株。表明过表达AePDS基因显著提高转基因拟南芥植株的耐盐性和抗旱性。
实施例7 AePDS基因转基因拟南芥植株抗性生理生化指标的测定
1.SOD活性测定
(1)试验方法
超氧化物歧化酶(SOD)活性可以作为植物抗逆性的一项生理生化指标。SOD的活性越低，植物遭受逆境伤害的程度越大。
测定方法参考文献【Feibing Wang,Weili Kong,Gary Wong,Lifeng Fu,RihePeng,Zhenjun Li,Quanhong Yao.AtMYB12regulates flavonoids accumulation andabiotic stress tolerance in transgenic Arabidopsis thaliana.MolecularGenetics and Genomics,2016,291:1545-1559】，检测拟南芥植株的SOD活性。拟南芥植株为上述盆栽鉴定中无胁迫处理2周的拟南芥植株、上述盆栽鉴定中盐处理1周的拟南芥植株和上述盆栽鉴定中干旱处理2周的拟南芥植株。实验需重复三次，结果取平均值。
(2)试验结果
实验结果见图7中A(Normal为空白对照，Salt stress为盐胁迫，Drought stress为干旱胁迫)。结果表明，转基因拟南芥L3植株和L4植株的SOD活性显著高于野生型拟南芥植株。
2.POD活性测定
(1)试验方法
过氧化物酶(POD)活性可以作为植物抗逆性的一项生理生化指标。POD的活性越低，植物遭受逆境伤害的程度越大。
测定方法参考文献【Feibing Wang,Weili Kong,Gary Wong,Lifeng Fu,RihePeng,Zhenjun Li,Quanhong Yao.AtMYB12 regulates flavonoids accumulationand abiotic stress tolerance in transgenic Arabidopsis thaliana.MolecularGenetics and Genomics,2016,291:1545-1559】，检测拟南芥植株的POD活性。拟南芥植株为上述盆栽鉴定中无胁迫处理2周的拟南芥植株、上述盆栽鉴定中盐处理1周的拟南芥植株和上述盆栽鉴定中干旱处理2周的拟南芥植株。实验需重复三次，结果取平均值。
(2)试验结果
实验结果见图7中B(Normal为空白对照，Salt stress为盐胁迫，Drought stress为干旱胁迫)。结果表明，转基因拟南芥L3植株和L4植株的POD活性显著高于野生型拟南芥植株。
3.ABA含量测定
(1)试验方法
ABA在植物逆境胁迫反应中具有重要作用。ABA可以提高植物的耐盐性，缓解盐分过多造成的渗透胁迫和离子胁迫，维持水分平衡，诱导植物渗透调剂物质脯氨酸大量积累，维持细胞膜结构的稳定性，提高保护性酶的活性。旱害胁迫时，ABA能明显减少叶片水分蒸发，降低叶片细胞膜透性，增加叶片细胞可溶性蛋白质含量，诱导生物膜系统保护酶形成，降低膜脂过氧化程度，增强抗氧化能力，提高植物的抗旱性。
测定方法参考文献【Feibing Wang,Weili Kong,Gary Wong,Lifeng Fu,RihePeng,Zhenjun Li,Quanhong Yao.AtMYB12 regulates flavonoids accumulationand abiotic stress tolerance in transgenic Arabidopsis thaliana.MolecularGenetics and Genomics,2016,291:1545-1559】，检测拟南芥植株的ABA含量。拟南芥植株为上述盆栽鉴定中无胁迫处理2周的拟南芥植株、上述盆栽鉴定中盐处理1周的拟南芥植株和上述盆栽鉴定中干旱处理2周的拟南芥植株。实验需重复三次，结果取平均值。。
(2)试验结果
实验结果见图7中C(Normal为空白对照，Salt stress为盐胁迫，Drought stress为干旱胁迫)。结果表明，转基因拟南芥L3植株和L4植株的ABA含量显著高于野生型拟南芥植株。
4.脯氨酸含量测定
(1)试验方法
植物在正常条件下，游离脯氨酸含量很低，但遇到盐、干旱等胁迫时，游离的氨基酸便会大量积累，并且积累指数和植物的抗逆性有关。因此，脯氨酸可以作为植物抗逆性的一项生化指标。
测定方法参考文献【Feibing Wang,Weili Kong,Gary Wong,Lifeng Fu,RihePeng,Zhenjun Li,Quanhong Yao.AtMYB12 regulates flavonoids accumulationand abiotic stress tolerance in transgenic Arabidopsis thaliana.MolecularGenetics and Genomics,2016,291:1545-1559】，检测拟南芥植株的脯氨酸含量。拟南芥植株为上述盆栽鉴定中无胁迫处理2周的拟南芥植株、上述盆栽鉴定中盐处理1周的拟南芥植株和上述盆栽鉴定中干旱处理2周的拟南芥植株。实验需重复三次，结果取平均值。
(2)试验结果
实验结果见图7中D(Normal为空白对照，Salt stress为盐胁迫，Drought stress为干旱胁迫)。结果表明，转基因拟南芥L3植株和L4植株的脯氨酸含量显著高于野生型拟南芥植株。
5.H2O2含量测定
(1)试验方法
植物在逆境下或衰老时，由于体内活性氧代谢加强而使H2O2发生累积。H2O2可以直接或间接地氧化细胞内核酸，蛋白质等生物大分子，并使细胞膜遭受损害，从而加速细胞的衰老和解体。因此，H2O2的含量越高，植物遭受逆境伤害的程度越大。
测定方法参考文献【Feibing Wang,Weili Kong,Gary Wong,Lifeng Fu,RihePeng,Zhenjun Li,Quanhong Yao.AtMYB12 regulates flavonoids accumulationand abiotic stress tolerance in transgenic Arabidopsis thaliana.MolecularGenetics and Genomics,2016,291:1545-1559】，检测拟南芥植株的MDA含量。拟南芥植株为上述盆栽鉴定中无胁迫处理2周的拟南芥植株、上述盆栽鉴定中盐处理1周的拟南芥植株和上述盆栽鉴定中干旱处理2周的拟南芥植株。实验需重复三次，结果取平均值。
(2)试验结果
实验结果见图7中E(Normal为空白对照，Salt stress为盐胁迫，Drought stress为干旱胁迫)。结果表明，转基因拟南芥L3植株和L4植株的H2O2含量显著低于野生型拟南芥植株。
6.MDA含量测定
(1)试验方法
植物器官衰老或在逆境下遭受伤害，往往发生膜脂过氧化作用，丙二醛(MDA)是膜脂过氧化的最终分解产物，其含量可以反映植物遭受逆境伤害的程度，即MDA的含量越高，植物遭受逆境伤害的程度越大。
测定方法参考文献【Feibing Wang,Weili Kong,Gary Wong,Lifeng Fu,RihePeng,Zhenjun Li,Quanhong Yao.AtMYB12 regulates flavonoids accumulationand abiotic stress tolerance in transgenic Arabidopsis thaliana.MolecularGenetics and Genomics,2016,291:1545-1559】，检测拟南芥植株的MDA含量。拟南芥植株为上述盆栽鉴定中无胁迫处理2周的拟南芥植株、上述盆栽鉴定中盐处理1周的拟南芥植株和上述盆栽鉴定中干旱处理2周的拟南芥植株。实验需重复三次，结果取平均值。
(2)试验结果
实验结果见图7中F(Normal为空白对照，Salt stress为盐胁迫，Drought stress为干旱胁迫)。结果表明，转基因拟南芥L3植株和L4植株的MDA含量显著低于野生型拟南芥植株。
附：本发明所涉及到的核苷酸序列表：
<110>淮阴工学院
<120>秋葵类胡萝卜素生物合成和抗逆性相关蛋白AePDS及其编码基因与应用
<210>SEQ ID NO 1
<211>1686
<212>DNA
<213>秋葵(Abelmoschusesculentus)
<400>1
ATGAGTCTTTCTGGGAGTGTTTCTGCCTTGCACTTAAACTTCCAAAGAAACAGGATAAGCATGGGAAGTGTGTTGGCTTTTAAAAGTGGTGAATCCATGGGAAATTCCTTGAGAATTCCATTAAGAAAGAGGTCAAGTAAGGGTGCTTGTCCTTTGCAGGTAGTTTGCATAGATTATCCAAGGCCAGAGCTTGAGAATACTGTTAATTTTTTGGAGGCTGCTTCTCTGTCCGCTTCTTTTCGTTCTGCTCCCCGTCCACCTAAGCCATTGCAAATCATAATTGCTGGTGCAGGTTTGGCTGGTTTGTCAACTGCAAAGTATCTAGCGGATGCAGGTCATAAACCAATATTACTAGAAGCTAGAGATGTTCTAGGCGGAAAGGTGGCTGCATGGAAGGATGATGATGGAGATTGGTATGAGACAGGCCTACACATATTTTTCGGAGCCTACCCGAATGTGCAAAACTTGTTTGGAGAACTTGGCATCAATGATCGTCTGCAATGGAAGGAGCATTCTATGATATTTGCAATGCCAAATAAGCCTGGAGAATTCAGTCGTTTTGATTTTCCAGACGTTCTACCTGCACCCTTAAATGGGATATGGGCCATTTTGAAGAACAATGAAATGCTGACATGGCCTGAGAAAGTGAAATTTGCAATAGGACTCCTACCCGCAATGCTTGGTGGGCAACCTTATGTTGAGGCCCAGGATGGTTTATCTGTTAAAGAGTGGATGATAAAGCAGGGCGTACCTGATCGTGTGACTAAAGAGGTGTTTATTGCTATGTCAAAGGCTCTGAACTTCATTAACCCCGAGGAACTTTCAATGCAGTGTATATTGATTGCTTTGAATCGATTTCTTCAGGAAAAACATGGATCAAAGATGGCATTCTTGGATGGCAACCCTCCAGAGAGGCTTTGCATGCCAATCGTTAATCATATTGAATCGCTGGGTGGTGAGGTCAGGCTTAACTCACGAATAAAGAAAATAGAGCTCAATGATGATGGAACTGTGAAGAGTCTTCTCCTAATTAACGGCAATACAATTGAAGGAGATGCTTACGTAATTGCAACTCCTGTTGATGTCTTCAAGTTACTTTTGCCTGAAAACTGGAGAGATATTTCGTACTTCAAGAAACTAGAGAAATTAGTTGGAGTTCCGGTTATCAACGTTCACATCTGGTTTGATAGGAAATTGAAGAACACCTATGATCACCTACTCTTCAGCAGAAGTCCCCTTTTAAGTGTTTATGCCGACATGTCTGTAACATGCAAGGAATATTACAATCCGAACCAATCCATGTTGGAGTTAGTTTTTGCCCCTGCAGAAGAATGGATTGCACGAAGTGACTCTGAAATTATTGATGCTACAATGAAGGAACTTGCAAAGCTCTTCCCTGATGAAATATCTGCAGACCAGAGTAAAGCAAAAATCGTAAAGTACCATGTCGTTAAAACACCGAGATCTGTATATAAAACCGTTCCAAATTGTGAACCATGCCGTCCCTTGCAAAGATCTCCGGTACAGGGGTTCTATCTAGCAGGTGATTACACAAAGCAAAAGTATTTAGCTTCAATGGAAGGTGCTGTCCTCTCAGGGAAGCTTTGTGCACAGTCTATTGTACAGGATTATGAGTTGCTTTGTACTCTGGGACAAAGGAAGTTGACTGGAGCAAGCAGACACTGA
<210>SEQ ID NO 2
<211>561
<212>PRT
<213>秋葵(Abelmoschusesculentus)
<400>2
MSLSGSVSALHLNFQRNRISMGSVLAFKSGESMGNSLRIPLRKRSSKGACPLQVVCIDYPRPELENTVNFLEAASLSASFRSAPRPPKPLQIIIAGAGLAGLSTAKYLADAGHKPILLEARDVLGGKVAAWKDDDGDWYETGLHIFFGAYPNVQNLFGELGINDRLQWKEHSMIFAMPNKPGEFSRFDFPDVLPAPLNGIWAILKNNEMLTWPEKVKFAIGLLPAMLGGQPYVEAQDGLSVKEWMIKQGVPDRVTKEVFIAMSKALNFINPEELSMQCILIALNRFLQEKHGSKMAFLDGNPPERLCMPIVNHIESLGGEVRLNSRIKKIELNDDGTVKSLLLINGNTIEGDAYVIATPVDVFKLLLPENWRDISYFKKLEKLVGVPVINVHIWFDRKLKNTYDHLLFSRSPLLSVYADMSVTCKEYYNPNQSMLELVFAPAEEWIARSDSEIIDATMKELAKLFPDEISADQSKAKIVKYHVVKTPRSVYKTVPNCEPCRPLQRSPVQGFYLAGDYTKQKYLASMEGAVLSGKLCAQSIVQDYELLCTLGQRKLTGASRH
序列表
<110> 淮阴工学院
<120> 秋葵类胡萝卜素生物合成和抗逆性相关蛋白AePDS及其编码基因与应用
<160> 14
<170> SIPOSequenceListing 1.0
<210> 1
<211> 1686
<212> DNA
<213> 秋葵(Abelmoschusesculentus)
<400> 1
atgagtcttt ctgggagtgt ttctgccttg cacttaaact tccaaagaaa caggataagc 60
atgggaagtg tgttggcttt taaaagtggt gaatccatgg gaaattcctt gagaattcca 120
ttaagaaaga ggtcaagtaa gggtgcttgt cctttgcagg tagtttgcat agattatcca 180
aggccagagc ttgagaatac tgttaatttt ttggaggctg cttctctgtc cgcttctttt 240
cgttctgctc cccgtccacc taagccattg caaatcataa ttgctggtgc aggtttggct 300
ggtttgtcaa ctgcaaagta tctagcggat gcaggtcata aaccaatatt actagaagct 360
agagatgttc taggcggaaa ggtggctgca tggaaggatg atgatggaga ttggtatgag 420
acaggcctac acatattttt cggagcctac ccgaatgtgc aaaacttgtt tggagaactt 480
ggcatcaatg atcgtctgca atggaaggag cattctatga tatttgcaat gccaaataag 540
cctggagaat tcagtcgttt tgattttcca gacgttctac ctgcaccctt aaatgggata 600
tgggccattt tgaagaacaa tgaaatgctg acatggcctg agaaagtgaa atttgcaata 660
ggactcctac ccgcaatgct tggtgggcaa ccttatgttg aggcccagga tggtttatct 720
gttaaagagt ggatgataaa gcagggcgta cctgatcgtg tgactaaaga ggtgtttatt 780
gctatgtcaa aggctctgaa cttcattaac cccgaggaac tttcaatgca gtgtatattg 840
attgctttga atcgatttct tcaggaaaaa catggatcaa agatggcatt cttggatggc 900
aaccctccag agaggctttg catgccaatc gttaatcata ttgaatcgct gggtggtgag 960
gtcaggctta actcacgaat aaagaaaata gagctcaatg atgatggaac tgtgaagagt 1020
cttctcctaa ttaacggcaa tacaattgaa ggagatgctt acgtaattgc aactcctgtt 1080
gatgtcttca agttactttt gcctgaaaac tggagagata tttcgtactt caagaaacta 1140
gagaaattag ttggagttcc ggttatcaac gttcacatct ggtttgatag gaaattgaag 1200
aacacctatg atcacctact cttcagcaga agtccccttt taagtgttta tgccgacatg 1260
tctgtaacat gcaaggaata ttacaatccg aaccaatcca tgttggagtt agtttttgcc 1320
cctgcagaag aatggattgc acgaagtgac tctgaaatta ttgatgctac aatgaaggaa 1380
cttgcaaagc tcttccctga tgaaatatct gcagaccaga gtaaagcaaa aatcgtaaag 1440
taccatgtcg ttaaaacacc gagatctgta tataaaaccg ttccaaattg tgaaccatgc 1500
cgtcccttgc aaagatctcc ggtacagggg ttctatctag caggtgatta cacaaagcaa 1560
aagtatttag cttcaatgga aggtgctgtc ctctcaggga agctttgtgc acagtctatt 1620
gtacaggatt atgagttgct ttgtactctg ggacaaagga agttgactgg agcaagcaga 1680
cactga 1686
<210> 2
<211> 561
<212> PRT
<213> 秋葵(Abelmoschusesculentus)
<400> 2
Met Ser Leu Ser Gly Ser Val Ser Ala Leu His Leu Asn Phe Gln Arg
1 5 10 15
Asn Arg Ile Ser Met Gly Ser Val Leu Ala Phe Lys Ser Gly Glu Ser
20 25 30
Met Gly Asn Ser Leu Arg Ile Pro Leu Arg Lys Arg Ser Ser Lys Gly
35 40 45
Ala Cys Pro Leu Gln Val Val Cys Ile Asp Tyr Pro Arg Pro Glu Leu
50 55 60
Glu Asn Thr Val Asn Phe Leu Glu Ala Ala Ser Leu Ser Ala Ser Phe
65 70 75 80
Arg Ser Ala Pro Arg Pro Pro Lys Pro Leu Gln Ile Ile Ile Ala Gly
85 90 95
Ala Gly Leu Ala Gly Leu Ser Thr Ala Lys Tyr Leu Ala Asp Ala Gly
100 105 110
His Lys Pro Ile Leu Leu Glu Ala Arg Asp Val Leu Gly Gly Lys Val
115 120 125
Ala Ala Trp Lys Asp Asp Asp Gly Asp Trp Tyr Glu Thr Gly Leu His
130 135 140
Ile Phe Phe Gly Ala Tyr Pro Asn Val Gln Asn Leu Phe Gly Glu Leu
145 150 155 160
Gly Ile Asn Asp Arg Leu Gln Trp Lys Glu His Ser Met Ile Phe Ala
165 170 175
Met Pro Asn Lys Pro Gly Glu Phe Ser Arg Phe Asp Phe Pro Asp Val
180 185 190
Leu Pro Ala Pro Leu Asn Gly Ile Trp Ala Ile Leu Lys Asn Asn Glu
195 200 205
Met Leu Thr Trp Pro Glu Lys Val Lys Phe Ala Ile Gly Leu Leu Pro
210 215 220
Ala Met Leu Gly Gly Gln Pro Tyr Val Glu Ala Gln Asp Gly Leu Ser
225 230 235 240
Val Lys Glu Trp Met Ile Lys Gln Gly Val Pro Asp Arg Val Thr Lys
245 250 255
Glu Val Phe Ile Ala Met Ser Lys Ala Leu Asn Phe Ile Asn Pro Glu
260 265 270
Glu Leu Ser Met Gln Cys Ile Leu Ile Ala Leu Asn Arg Phe Leu Gln
275 280 285
Glu Lys His Gly Ser Lys Met Ala Phe Leu Asp Gly Asn Pro Pro Glu
290 295 300
Arg Leu Cys Met Pro Ile Val Asn His Ile Glu Ser Leu Gly Gly Glu
305 310 315 320
Val Arg Leu Asn Ser Arg Ile Lys Lys Ile Glu Leu Asn Asp Asp Gly
325 330 335
Thr Val Lys Ser Leu Leu Leu Ile Asn Gly Asn Thr Ile Glu Gly Asp
340 345 350
Ala Tyr Val Ile Ala Thr Pro Val Asp Val Phe Lys Leu Leu Leu Pro
355 360 365
Glu Asn Trp Arg Asp Ile Ser Tyr Phe Lys Lys Leu Glu Lys Leu Val
370 375 380
Gly Val Pro Val Ile Asn Val His Ile Trp Phe Asp Arg Lys Leu Lys
385 390 395 400
Asn Thr Tyr Asp His Leu Leu Phe Ser Arg Ser Pro Leu Leu Ser Val
405 410 415
Tyr Ala Asp Met Ser Val Thr Cys Lys Glu Tyr Tyr Asn Pro Asn Gln
420 425 430
Ser Met Leu Glu Leu Val Phe Ala Pro Ala Glu Glu Trp Ile Ala Arg
435 440 445
Ser Asp Ser Glu Ile Ile Asp Ala Thr Met Lys Glu Leu Ala Lys Leu
450 455 460
Phe Pro Asp Glu Ile Ser Ala Asp Gln Ser Lys Ala Lys Ile Val Lys
465 470 475 480
Tyr His Val Val Lys Thr Pro Arg Ser Val Tyr Lys Thr Val Pro Asn
485 490 495
Cys Glu Pro Cys Arg Pro Leu Gln Arg Ser Pro Val Gln Gly Phe Tyr
500 505 510
Leu Ala Gly Asp Tyr Thr Lys Gln Lys Tyr Leu Ala Ser Met Glu Gly
515 520 525
Ala Val Leu Ser Gly Lys Leu Cys Ala Gln Ser Ile Val Gln Asp Tyr
530 535 540
Glu Leu Leu Cys Thr Leu Gly Gln Arg Lys Leu Thr Gly Ala Ser Arg
545 550 555 560
His
<210> 3
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 3
cagacgttct acctgcaccc 20
<210> 4
<211> 17
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 4
gttttcccag tcacgac 17
<210> 5
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 5
agcattcgtg ggatcaagtc 20
<210> 6
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 6
ctcaccaccc agcgattcaa 20
<210> 7
<211> 24
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 7
atgagtcttt ctgggagtgt ttct 24
<210> 8
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 8
tcagtgtctg cttgctccag 20
<210> 9
<211> 21
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 9
tgcttgtcct ttgcaggtag t 21
<210> 10
<211> 21
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 10
tctgcttgct ccagtcaact t 21
<210> 11
<211> 21
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 11
gcaccctgtt cttcttaccg a 21
<210> 12
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 12
agtaaggtca cgtccagcaa gg 22
<210> 13
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 13
gcgtacctga tcgtgtgact 20
<210> 14
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 14
gggttgccat ccaagaatgc 20
Claims (10)
1.一种蛋白，是如下(a)或(b)：
(a)由序列表中SEQ ID NO.2所示的氨基酸序列组成的蛋白质；
(b)将序列表中SEQ ID NO.2所示的氨基酸残基序列经过一个或几个氨基酸残基的取代和/或缺失和/或添加且与植物类胡萝卜素积累和抗逆性相关的由SEQ ID NO.2衍生的蛋白质。
2.编码权利要求1所述蛋白的基因。
3.如权利要求2所述的基因，其特征在于：所述基因是如下(a1)-(a4)中任何一种的DNA分子：
(a1)编码区包括序列表中如SEQ ID NO.1所示核苷酸序列的DNA分子；
(a2)核苷酸序列是序列表中如SEQ ID NO.1所示的DNA分子；
(a3)在严格条件下与(a1)或(a2)限定的DNA序列杂交且编码与植物类胡萝卜素积累和抗逆性相关蛋白的DNA分子；
(a4)与(a1)或(a2)限定的DNA序列至少具有70％、至少具有75％、至少具有80％、至少具有85％、至少具有90％、至少具有95％、至少具有96％、至少具有97％、至少具有98％或者至少具有99％同源性且编码与植物类胡萝卜素积累和抗逆性相关蛋白的DNA分子。
4.含有权利要求2或3所述基因的表达盒、重组载体、重组微生物或转基因细胞系。
5.(b1)或(b2)或(b3)或(b4)的应用：
(b1)权利要求1所述蛋白质，或，权利要求2所述基因，或，含有权利要求2或3所述基因的表达盒、重组载体、重组微生物或转基因细胞系，在调控植物中类胡萝卜素含量中的应用；
(b2)权利要求1所述蛋白质，或，权利要求2所述基因，或，含有权利要求2或3所述基因的表达盒、重组载体、重组微生物或转基因细胞系，在培育类胡萝卜素含量改变的转基因植物中的应用；
(b3)权利要求1所述蛋白质，或，权利要求2所述基因，或，含有权利要求2或3所述基因的表达盒、重组载体、重组微生物或转基因细胞系，在调控植物抗逆性中的应用；
(b4)权利要求1所述蛋白质，或，权利要求2或3所述基因，或，含有权利要求2或3所述基因的表达盒、重组载体、重组微生物或转基因细胞系，在培育抗逆性改变的转基因植物中的应用。
6.一种培育类胡萝卜素含量和/或抗逆性高的转基因植物的方法，其特征在于：提高目的植物中权利要求1所述蛋白质的含量或活性，得到转基因植物；所述转基因植物的类胡萝卜素含量和/或抗逆性高于所述目的植物。
7.一种培育类胡萝卜素含量和/或抗逆性高的植物的方法，其特征在于：为培育权利要求6所述方法得到的转基因植物。
8.如权利要求1所述蛋白质，或，权利要求5所述的应用，或，权利要求6或7所述的方法，其特征在于：所述抗逆性为耐盐性和/或抗旱性。
9.如权利要求1或8所述的蛋白质，或，权利要求5或8所述的应用，或，权利要求6或7或8所述的方法，其特征在于：所述类胡萝卜素含量为叶黄素(lutein)、玉米黄质(zeaxanthin)、β-隐黄质(β-cryptoxanthin)、α-胡萝卜素(α-carotene)、β-胡萝卜素(β-carotene)和/或总类胡萝卜素(total carotenoids)含量。
10.如权利要求1、8或9所述蛋白质，或，权利要求5、8或9所述的应用，或，权利要求6至9任一所述的方法，其特征在于：所述植物是如下(c1)至(c4)中的任一种：
(c1)双子叶植物；
(c2)单子叶植物；
(c3)十字花科植物；
(c4)拟南芥。
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| CN116536283A (zh) * | 2022-12-31 | 2023-08-04 | 淮阴工学院 | 秋葵类胡萝卜素生物合成和抗逆性相关蛋白AePSY及其编码基因和应用 |
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| CN116445464A (zh) * | 2022-12-28 | 2023-07-18 | 淮阴工学院 | 一种提高植物类胡萝卜素积累和耐盐性相关蛋白AeLCYE及其编码基因与应用 |
| CN116536283A (zh) * | 2022-12-31 | 2023-08-04 | 淮阴工学院 | 秋葵类胡萝卜素生物合成和抗逆性相关蛋白AePSY及其编码基因和应用 |
| CN116640195A (zh) * | 2023-05-10 | 2023-08-25 | 西南大学 | 柑橘CsPIF4基因及其编码的蛋白与应用 |
| CN116640195B (zh) * | 2023-05-10 | 2024-03-12 | 西南大学 | 柑橘CsPIF4基因及其编码的蛋白与应用 |
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