CN110499318B - 棉花抗黄萎病相关基因GhDEK的应用 - Google Patents
棉花抗黄萎病相关基因GhDEK的应用 Download PDFInfo
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- CN110499318B CN110499318B CN201910834989.9A CN201910834989A CN110499318B CN 110499318 B CN110499318 B CN 110499318B CN 201910834989 A CN201910834989 A CN 201910834989A CN 110499318 B CN110499318 B CN 110499318B
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- verticillium wilt
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Abstract
本发明属于农业生物技术领域，具体涉及棉花抗黄萎病相关基因GhDEK的应用。本发明的棉花抗黄萎病相关蛋白GhDEK，其氨基酸序列如SEQ ID NO.1所示。本发明的基因GhDEK的表达与棉花抗黄萎病呈正相关，可作为基因工程选育棉花抗病品种的候选基因。
Description
技术领域
本发明属于农业生物技术领域，具体涉及棉花抗黄萎病相关基因GhDEK的应用。
背景技术
棉花黄萎病作为棉花第一大病害，素有棉花“癌症”之称，其流行爆发给世界棉花产业的稳定发展造成严重威胁。然而由于棉花黄萎病是一种土传病害，防治难度极大，目前尚无理想的防治措施。因此，通过基因育种工程选育抗病品种成为解决棉花黄萎病的重要技术手段。
病原菌与植物的侵染和抗侵染机制一直是研究热点，植物与病原物之间的PTI和ETI抗病机理，成为研究抗病育种的重要手段。在病原物胁迫下，植物体内激活一系列信号转导，以激发植物的抗性反应，以抵御病原菌的侵染。现有技术研究表明，植物中的蛋白质通过发生蛋白质修饰，涉及植物生长、发育和免疫等。
发明内容
本发明的目的在于提供棉花抗黄萎病相关基因GhDEK的应用。
本发明的再一目的在于提供一种提高棉花黄萎病抗性品种的抗病性的方法。
本发明提供的与抗黄萎病相关的DEK结构域染色质相关蛋白(DEK domain-containing chromatin associated protein)GhDEK的氨基酸序列如SEQ ID NO.1所示：
通过蛋白质修饰组检测，蛋白GhDEK的第327位的苏氨酸(Thr，T)和329位的丝氨酸(Ser，S)发生了磷酸化修饰，第273位的赖氨酸(Lys，K)发生了乙酰化修饰。
根据本发明具体实施方式的蛋白GhDEK的编码基因的CDS序列如SEQ ID NO.2所示：
ATGGCGACGGAAACCCTAGATGACAAGAAACCGGAGGAAGAGGAGGTGAAAGACAAAGAAAACGAAGAGGGAAGTAAGGAGGTTTTGGAGAAACAAATGGAAGTTGAGGAGAAGGAGAACGAGGAAGAAGAAGAAGAAGAAGATGAGGAAGAGAAGAGGGAGGAGGAAGAAGAAGAGAGTGAAGATGAAGGGACTAAAAAGGTCAAAGGTAGTAGTCGAAAGGGGAGTTCTAGGAAATCTGGTCGAGATTCGGCTGAGAAGAAAGAGCCAGTGACGCCTAGTAGTGATAGGCCTACAAGGGAAAGGAAAGTCGTAGAAAGGTATTCAGCTCCTTCTGTTGCAAGGTATTCCTCGTCTAAAACTCTGTCAATTGAAAAGGGTCGGGGTACTCAGCTTAAAGATATTCCCAATGTGGCTTTCAAGTTGTCAAAGAGAAAAGCTGATGATAATCTGCAGATGCTTCATATAATTCTCTTTGGAAAGAAAGCAAAGCCTCACAGTTTGAAGAGAAACATTGGTCAATTTTCAGGCTATGTTTGGGTTGAGAACGAGCAGGAAAAACAAAAAGCGAAAGTAAGGGAAAAAATTGACAAATGTGTTAAAGAAAAATTGGTTGATTTCTGTGATTTGCTGAATATTCCATTTATGAGGACCAGTGTAAGAAAGGAGGAAGTCACTGCCAAATTATTGGAATTTTTGGAATCTCCTCATGCCACTACAGACATTCTTCTTGCTGACAAGGAACAGAAGGGTAAAAAGCGTAAGGCTACACCAAGCAAAAACATTGGTTCTGCAGAGGCATTGGATACATCAGCCAAGAAGCGACGAAAAACACCCCAAGGTGGAGAAAAGCGCAAGCGTTCATCCAAAGCTGAGGAGGAGGAAGATGATGATAAAGTTGAATCCCCTGTTGCTAGAGATGATTCTCATGAAGATGATGCTGACACTGCACCAAAAGAAGTGAATGATGATGAGGAGACTAAATCAGAGGAGGAAGAAGAACCCAAGAAGTCAAGCAAAAAGGGCACTTCAAAAAAGGTTGCAACAGAGAGTCCGGAGTCAAAAAGCAAAGATAAATCCGAATCTGGAAAGAATCTCACCCCTGCAAAATCTAGCAAAAAATCTTCTGGATCAACTTCAAAACAAGATGCTAGTGATGGTGGTGGGACTTCTGGCTCTAAATCAAAGGGTTCTGCATCAAAGAAGCCTAAGGTTGAAAAGGAAAACTCTAAGGATGGATCCACCAAAGAGAAGCTTGCAGT CAAGAAGCAAACAAATAAGTCATCAGCAAAGGTTTCTGCTAAATCACAAGGTAAAAGCAAAAGTGGCAAGAAACCTGAGCCTAGTAGGGAAGAGATTCATGAAGTTGTCGTAGATATTCTAAAAAAAGTGGACTTCAACACTGTAAGTATTTTGATAATCTCAGGTACACACTTTGACCTGGATTTGATGCACAGAAAAGCCGAGGTGAAGGATATCATTACAGATGTGATAAATAACATGTCTGATGAGGATGAGGAAGGGGATGAAAGTGAGGAGAATGCTGATACAGGTGGGGGTGCTGATAAAGATGGTGATGGAGATGATGATGCCTAG
本发明提供了基因GhDEK在棉花抗黄萎病上的应用。在接种病原菌后，蛋白GhDEK的第327位的苏氨酸(Thr，T)和329位的丝氨酸(Ser，S)发生了磷酸化修饰，第273位的赖氨酸(Lys，K)发生了乙酰化修饰；蛋白GhDEK在棉花体内表达上调，表明在病原菌的胁迫下激活了基因GhDEK的表达，通过激素(茉莉酸、乙烯、水杨酸和过氧化氢)处理后，GhDEK的表达发生了改变，表明GhDEK受激素的调控。
根据本发明具体实施方式的提高棉花抗黄萎病品种的抗病性的方法，所述方法包括在棉花抗性品种中过表达棉花抗黄萎病相关基因GhDEK的步骤；其中，所述方法还包括使用乙烯促进抗黄萎病相关基因GhDEK表达的步骤。
本发明的有益效果：
本发明提供一种与棉花抗黄萎病相关的DEK结构域染色质相关蛋白GhDEK，棉花在接种病原菌后，其在棉花体内的部分位点发生了磷酸化和乙酰化修饰，蛋白GhDEK在棉花体内表达上调。
本发明构建了基因GhDEK的沉默载体，利用病毒介导的基因沉默技术，抑制了GhDEK在棉花中的表达。对基因沉默植株接种棉花黄萎病病原菌Vd080孢子悬浮液后，沉默植物表现更感病，通过机理研究发现，沉默植株中木质部和胼胝质的合成下降，ROS减少，部分防御基因下调表达，因此，GhDEK与棉花抗黄萎病呈正相关，GhDEK可作为棉花抗病育种的候选基因。
附图说明
图1显示GhDEK在抗/感病品种中的表达情况；
图2显示激素处理后GhDEK的抗病品种中的表达情况；
图3显示激素处理后GhDEK的感病品种中的表达情况；
图4显示基因沉默后的白化现象及发病情况；
图5显示基因沉默后植株木质部的积累情况；
图6显示基因沉默后植株叶片胼胝质的积累情况；
图7显示基因沉默后植株叶片活性氧的爆发情况；
图8显示病菌处理后植株中相关防御基因的相对表达量。
具体实施方式
实施例1棉花蛋白质修饰的检测
以抗病品种中植棉2号和感病品种冀棉11号为植物材料，接种棉花黄萎病病原菌Vd080后，进行蛋白质修饰的检测，分析发现基因ID号为Gh_D07G0492的蛋白同时发生了磷酸化和乙酰化修饰，根据序列比对结果，命名为GhDEK。
实施例2不同抗性棉花品种中GhDEK的表达差异
在蛭石沙土纸钵中种植棉花抗病品种中植棉2号和感病品种冀棉11号，对抗病品种中植棉2号和感病品种冀棉11号不同时间段提取根部RNA，对GhDEK的表达进行分析。根据GhDEK的基因序列和CDS序列设计其荧光定量的引物为：
GhDEK-F:GCTATGTTTGGGTTGAGA，
GhDEK-R:TAGTGGCATGAGGAGATT。
如图1所示，在抗病品种中GhDEK的表达量显著高于感病品种，表明GhDEK在提高抗病品种的抗病性上起作用。
实施例3激素处理后对棉花中GhDEK表达的影响
在蛭石沙土纸钵中种植棉花抗病品种中植棉2号和感病品种冀棉11号，伤根后接种Vd080孢子悬浮液，不同时间段提取根部RNA。用0.5mM的过氧化氢(H2O2)、0.1mM的水杨酸(SA)，0.15mM的茉莉酸甲酯(JA)和1mM的乙烯(ET)喷施到叶面滴水为止，不同时间段提取根部RNA。根据GhDEK的基因序列和CDS序列设计其荧光定量的引物为：
GhDEK-F:GCTATGTTTGGGTTGAGA，
GhDEK-R:TAGTGGCATGAGGAGATT。
检测激素处理后GhDEK的表达情况。
如图2、3所示，在激素处理后，抗病品种的GhDEK对H2O2、SA和JA敏感，感病品种的GhDEK对SA和H2O2敏感，表明GhDEK的表达受激素的调控。
实施例4利用病毒介导的基因沉默技术(VIGS)研究GhDEK的功能
4.1棉花中基因GhDEK的沉默
设计GhDEK沉默载体的引物：
DEK-VIGS-F:GCTCTAGAAGGCTATGTTTGGGTT，
DEK-VIGS-R:GGGGTACCTTTACCCTTCTGTTCC。
以抗病品种中植棉2号的cDNA为模板扩增沉默片段并转化pYL-156载体，并转化大肠杆菌DH5α感受态细胞，测序验证正确后，提取pYL-156-GhDEK质粒，并转化农杆菌GV3101感受态，菌落PCR验证正确后，扩大培养，以pYL-156空载为对照，以pYL-156-PDS为正对照(PDS基因被沉默后，叶片表现出白化现象)，与辅助质粒pYL-192为混合静置后，用无针头注射器注射中植棉2号棉花叶片。注射处理后暗培养24h，置于正常光照下22℃培养。
待正对照出现白化表型时，如图4所示，利用荧光定量PCR检测沉默植株中GhDEK的表达量，选取沉默效果较好的植株进行进一步试验。
4.2沉默植株的抗病性研究
选取沉默效果较好的植株，待一片真叶初现时，接种10mL浓度为2×107CFU/mL大丽轮枝菌孢子液，置于25℃温室中正常光照生长。接菌后不同时间段取样提取沉默植株RNA用于检测防御相关基因的表达。在接菌3d时，检测棉花茎秆木质部和叶片胼胝质的积累，检测棉花叶片活性氧的爆发。15d时检测叶片细胞坏死情况。20d时，调查植株发病情况。
如图4所示，统计病情后计算得沉默植株的病情指数为60.52±3.37，病株率为100％，而对照的病情指数为21.92±2.68，病株率为83.74±2.56％，沉默植株与对照之间的病情指数和病株率均存在极显著差异。
将棉花幼苗茎秆用间苯三酚染色，浓硫酸孵育后，在正视显微镜下观察，如图5所示，沉默植株的木质部的积累显著低于非沉默植株。
利用苯胺蓝染色棉花叶片，紫外激发光下检视。如图6所示，沉默植株的胼胝质的积累量低于对照。
如图7所示，活性氧爆发的研究同样显示在沉默植物的叶片中褐色沉淀更少，说明活性氧爆发更弱。
接种病菌后的不同时间段沉默植株中的防御酶基因或防御酶代谢基因，检测相关基因的表达量变化情况如图8所示。
表1棉花防御相关基因的RT-qPCR引物
沉默植株中，苯丙氨酸解氨酶(GhPAL)、肉桂酸-4-羟基化酶(GhC4H1)、过氧化物酶(GhPOD)和多酚氧化酶(GhPPO)的表达量在接种病原菌后具有不同程度的降低。
棉花中过敏反应的标识基因GhHSR203J、GhHIN1和GhHSR203J在整个检测时期的表达水平均低于对照，表明GhDEK的沉默导致了活性氧的减少。GhPR3是乙烯(ET)信号通路的标识基因，在沉默植株中的表达均显著低于对照，表明ET在GhDEK抗病中起正调控作用。GhJaZ1是JA信号通路的标识基因，其表达在沉默植株中上调，表明JA与GhDEK是负调控作用。GhPR1是水杨酸信号通路的标识基因，沉默植株中的表达量高于对照，水杨酸在GhDEK的抗病调控中起负调控。GhNOA1是一氧化氮通路基因，在沉默植株中的表达量均显著低于对照，表明GhDEK与一氧化氮通路相关。因此，GhDEK在棉花抗黄萎病中具有重要的正调控作用，其可作为棉花抗黄萎病育种的重要候选基因。
实施例5拟南芥中过表达GhDEK研究的功能
以中植棉2号的cDNA为模板，克隆GhDEK的CDS序列，从而构建pCAMBIA2300-GhDEK的过表达载体，通过花盘转化技术，使拟南芥过表达DEK蛋白，通过连续3代抗性筛选及分子验证，获得阳性转基因苗。
栽培转基因拟南芥和野生型拟南芥，接种大丽轮枝菌病原菌，待发病后评价病情。结果显示，转基因拟南芥的病情指数为19.23±1.12，病株率为15.36±0.35％；野生型拟南芥的病情指数为56.78±2.18，病株率为25.32±1.02％，转基因拟南芥表现出明显的抗黄萎病性。
序列表
<110> 中国农业科学院棉花研究所
<120> 棉花抗黄萎病相关基因GhDEK的应用
<160> 2
<170> SIPOSequenceListing 1.0
<210> 1
<211> 531
<212> PRT
<213> 棉花(Malvaceae Gossypium)
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Glu Glu Glu Lys Arg Glu Glu Glu Glu Glu Glu Ser Glu Asp Glu Gly
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Gly Thr Gln Leu Lys Asp Ile Pro Asn Val Ala Phe Lys Leu Ser Lys
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Arg Lys Ala Asp Asp Asn Leu Gln Met Leu His Ile Ile Leu Phe Gly
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Lys Lys Ala Lys Pro His Ser Leu Lys Arg Asn Ile Gly Gln Phe Ser
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Gly Tyr Val Trp Val Glu Asn Glu Gln Glu Lys Gln Lys Ala Lys Val
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Arg Glu Lys Ile Asp Lys Cys Val Lys Glu Lys Leu Val Asp Phe Cys
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Asp Leu Leu Asn Ile Pro Phe Met Arg Thr Ser Val Arg Lys Glu Glu
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Val Thr Ala Lys Leu Leu Glu Phe Leu Glu Ser Pro His Ala Thr Thr
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Asp Ile Leu Leu Ala Asp Lys Glu Gln Lys Gly Lys Lys Arg Lys Ala
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atggcgacgg aaaccctaga tgacaagaaa ccggaggaag aggaggtgaa agacaaagaa 60
aacgaagagg gaagtaagga ggttttggag aaacaaatgg aagttgagga gaaggagaac 120
gaggaagaag aagaagaaga agatgaggaa gagaagaggg aggaggaaga agaagagagt 180
gaagatgaag ggactaaaaa ggtcaaaggt agtagtcgaa aggggagttc taggaaatct 240
ggtcgagatt cggctgagaa gaaagagcca gtgacgccta gtagtgatag gcctacaagg 300
gaaaggaaag tcgtagaaag gtattcagct ccttctgttg caaggtattc ctcgtctaaa 360
actctgtcaa ttgaaaaggg tcggggtact cagcttaaag atattcccaa tgtggctttc 420
aagttgtcaa agagaaaagc tgatgataat ctgcagatgc ttcatataat tctctttgga 480
aagaaagcaa agcctcacag tttgaagaga aacattggtc aattttcagg ctatgtttgg 540
gttgagaacg agcaggaaaa acaaaaagcg aaagtaaggg aaaaaattga caaatgtgtt 600
aaagaaaaat tggttgattt ctgtgatttg ctgaatattc catttatgag gaccagtgta 660
agaaaggagg aagtcactgc caaattattg gaatttttgg aatctcctca tgccactaca 720
gacattcttc ttgctgacaa ggaacagaag ggtaaaaagc gtaaggctac accaagcaaa 780
aacattggtt ctgcagaggc attggataca tcagccaaga agcgacgaaa aacaccccaa 840
ggtggagaaa agcgcaagcg ttcatccaaa gctgaggagg aggaagatga tgataaagtt 900
gaatcccctg ttgctagaga tgattctcat gaagatgatg ctgacactgc accaaaagaa 960
gtgaatgatg atgaggagac taaatcagag gaggaagaag aacccaagaa gtcaagcaaa 1020
aagggcactt caaaaaaggt tgcaacagag agtccggagt caaaaagcaa agataaatcc 1080
gaatctggaa agaatctcac ccctgcaaaa tctagcaaaa aatcttctgg atcaacttca 1140
aaacaagatg ctagtgatgg tggtgggact tctggctcta aatcaaaggg ttctgcatca 1200
aagaagccta aggttgaaaa ggaaaactct aaggatggat ccaccaaaga gaagcttgca 1260
gtcaagaagc aaacaaataa gtcatcagca aaggtttctg ctaaatcaca aggtaaaagc 1320
aaaagtggca agaaacctga gcctagtagg gaagagattc atgaagttgt cgtagatatt 1380
ctaaaaaaag tggacttcaa cactgtaagt attttgataa tctcaggtac acactttgac 1440
ctggatttga tgcacagaaa agccgaggtg aaggatatca ttacagatgt gataaataac 1500
atgtctgatg aggatgagga aggggatgaa agtgaggaga atgctgatac aggtgggggt 1560
gctgataaag atggtgatgg agatgatgat gcctag 1596
Claims (4)
1.棉花抗黄萎病相关基因GhDEK在防治棉花黄萎病方面的应用，其中，所述棉花抗黄萎病相关基因GhDEK的核苷酸序列如SEQ ID NO.2所示。
2.棉花抗黄萎病相关基因GhDEK在提高棉花黄萎病抗病品种的抗病性方面的应用，其中，所述棉花抗黄萎病相关基因GhDEK的核苷酸序列如SEQ ID NO.2所示。
3.提高棉花黄萎病抗性品种的抗病性的方法，其特征在于，所述方法包括在棉花黄萎病抗性品种中过表达棉花抗黄萎病相关基因GhDEK的步骤，所述棉花抗黄萎病相关基因GhDEK的核苷酸序列如SEQ ID NO.2所示。
4.根据权利要求3所述的提高棉花黄萎病抗性品种的抗病性的方法，其特征在于，所述方法还包括使用乙烯促进抗黄萎病相关基因GhDEK表达的步骤。
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| CN107653258A (zh) * | 2017-11-16 | 2018-02-02 | 河南大学 | 棉花GhLecRK1基因在植物抗黄萎病中的应用 |
| CN109402147A (zh) * | 2018-11-02 | 2019-03-01 | 南京农业大学 | 抗棉花黄萎病的基因GbCYP86A1-1及其应用 |
| CN109706132A (zh) * | 2018-12-10 | 2019-05-03 | 中国农业科学院棉花研究所 | 棉花抗黄萎病相关蛋白GhMAPK13及其编码基因和应用 |
| CN109705201A (zh) * | 2018-12-10 | 2019-05-03 | 中国农业科学院棉花研究所 | 棉花抗黄萎病相关基因GhABC及其编码蛋白和应用 |
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| CN109402147A (zh) * | 2018-11-02 | 2019-03-01 | 南京农业大学 | 抗棉花黄萎病的基因GbCYP86A1-1及其应用 |
| CN109706132A (zh) * | 2018-12-10 | 2019-05-03 | 中国农业科学院棉花研究所 | 棉花抗黄萎病相关蛋白GhMAPK13及其编码基因和应用 |
| CN109705201A (zh) * | 2018-12-10 | 2019-05-03 | 中国农业科学院棉花研究所 | 棉花抗黄萎病相关基因GhABC及其编码蛋白和应用 |
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