CN117264977B - 一种黄瓜抗蚜调控基因CsRBOHD及其应用 - Google Patents
一种黄瓜抗蚜调控基因CsRBOHD及其应用 Download PDFInfo
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Abstract
本发明涉及一种黄瓜抗蚜调控基因CsRBOHD及其应用，其核苷酸序列如SEQ ID NO.1所示。黄瓜抗蚜调控基因CsRBOHD的表达蛋白，氨基酸序列如SEQ ID NO.2所示。含有所述的黄瓜抗蚜调控基因CsRBOHD的植物病毒载体pV190。所述载体沉默CsRBOHD基因，在植株中低表达。所述转入载体为pV190，使用BsaI将pV190酶切进行线性化处理再进行重组，得到重组载体。本发明以黄瓜主栽品种“Superina”为材料，通过RACE技术克隆了CsRBOHD基因，同时通过酶切、重组反应，农杆菌转化法，对黄瓜叶片进行注射，并进行非选择性定居实验、ROS相关酶活性测定以及DAB染色，证明其为抗蚜性的调控因子，在增强植物抗性、改良抗蚜性筛选方法和创制种质资源等黄瓜育种领域有重要应用价值。
Description
技术领域
本发明涉及一种黄瓜抗蚜调控基因CsRBOHD及其应用，属于植物基因工程技术领域。
背景技术
黄瓜（Cucumis sativus L.）是葫芦科甜瓜属一年生攀援草本植物，是我国重要的蔬菜作物之一，也是全球重要的经济蔬菜作物。其果实可提取多种游离氨基酸以及挥发油、葫芦素、黄瓜酶等，广泛应用于化妆品、食品及药用等行业，经济和保健价值极高。
蚜虫，是半翅目蚜总科的统称，目前已经发现的蚜虫总共有10个科约4400种，其中多数属于蚜科，广泛分布于世界各地，每年造成粮食作物减产5-20%，是全球十大经济害虫之一。
研究表明，蚜虫危害作物如：菊花、甜瓜、茄子、辣椒等，尤其与黄瓜生产期一致，在黄瓜幼嫩部位产生大量蜜露造成煤污病，影响植物光合作用，进而导致植物养分不足，同时还能传播病毒，导致植株萎蔫或死亡，在生产上危害十分严重。生产上常用手段是喷施农药，会出现“3R”问题，从基因方面考虑在根本上解决生产上的虫害问题。目前对蚜虫研究已经从传统手段到分子水平，但在黄瓜上的抗蚜性报道较少。
RBOHD（Respiratory burst oxidase homolog D）属于呼吸爆发氧化酶家族，是一种NADPH氧化酶。RBOHD在植物中参与调节氧化还原反应，产生活性氧（ROS，reactiveoxygen species）。活性氧在植物中起着双重作用，既可以作为信号分子参与植物的生长发育和逆境应答，也可以作为有害物质对细胞结构和功能造成损害。RBOHD通过将电子从NADPH转移到氧分子上，产生超氧阴离子（O2-），进而生成其他ROS，如过氧化氢（H2O2）。这些ROS在植物中参与多种生物学过程，如细胞壁增强、植物免疫反应、植物对逆境的应答等。
研究表明，RBOHD在黄瓜中的表达受到多种逆境因子的调控，如高盐、干旱、寒冷等。RBOHD的过表达可以增强植物对逆境的抵抗能力，而其缺失则会导致植物对逆境的敏感性增加。
因此，本研究基于RBOHD在植物面对抗胁迫中起着重要的调节作用，参与植物的生长发育和逆境应答，通过VIGS技术沉默黄瓜植株中CsRBOHD基因、黄瓜蚜虫抗性筛选实验以及相关酶活性测定和染色分析，证明CsRBOHD基因可调控黄瓜蚜虫抗性，对CsRBOHD的研究有助于深入了解植物对蚜虫抗性的应答机制，为创制抗蚜性种质资源提供分子工具，具有降低环境污染和经济成本的潜在价值，同时也对生产上解决蚜虫虫害问题具有重要应用价值。
发明内容
本发明的目的是为了解决背景技术中的技术问题，提出一种黄瓜抗蚜调控基因CsRBOHD及其应用。
为了实现上述发明目的，本发明采用的技术方案如下：
一种黄瓜抗蚜调控基因CsRBOHD，其特征在于，其核苷酸序列如SEQ ID NO.1所示。
黄瓜抗蚜调控基因CsRBOHD的表达蛋白，氨基酸序列如SEQ ID NO .2所示。
含有所述的黄瓜抗蚜调控基因CsRBOHD的植物病毒载体pV190。
所述载体沉默CsRBOHD基因，在植株中低表达。
所述转入载体为pV190，使用BsaI将pV190酶切进行线性化处理再进行重组，得到重组载体。
一种调控黄瓜抗蚜性的方法，所述调控基因沉默的方法包括：
步骤S1：将所述的重组载体转入至农杆菌；
步骤S2：将转入所述重组载体的农杆菌注射黄瓜苗子叶。
所述的黄瓜抗蚜调控基因CsRBOHD在调控黄瓜蚜虫抗性中的应用，所述CsRBOHD基因与ROS水平相关。
VIGS技术沉默CsRBOHD基因，ROS水平降低，植物抗性减弱。
本发明方法先进科学，通过本发明，一种黄瓜抗蚜调控基因CsRBOHD，其核苷酸序列如SEQ ID NO .1所示。所述黄瓜抗蚜调控基因CsRBOHD的表达蛋白，其氨基酸序列如SEQID NO .2所示。含有所述的黄瓜抗蚜调控基因CsRBOHD的VIGS载体pV190。含有所述的黄瓜抗蚜调控基因CsRBOHD的宿主细胞。所述黄瓜抗蚜调控基因CsRBOHD在增强黄瓜在生产上抗蚜虫的应用。
有益效果：本发明以黄瓜主栽品种“Superina”为材料，通过RACE技术克隆了CsRBOHD基因，同时通过酶切、重组反应，农杆菌转化法，对黄瓜叶片进行注射，并进行非选择性定居实验、ROS相关酶活性测定以及DAB染色，证明其为抗蚜性的调控因子，在增强植物抗性、改良抗蚜性筛选方法和创制种质资源等黄瓜育种领域有重要应用价值。
附图说明
图1是富集蚜虫相关通路图；接种蚜虫后转录组差异表达基因（DEGs）参与的生物过程的GO富集（展示差异最显著的20个生物学过程），依据Fold Enrichment>1,genecounts>10。
图2是RBOHs基因家族进化树图；拟南芥和黄瓜RBOHs家族9个关键基因（CsRBOHD、CsRBOHB、CsRBOHFa、CsRBOHFc、CsRBOHE、CsRBOHFb、CsRBOHH、 CsRBOHJa、CsRBOHJb）的进化树。
图3是6h、24h、72h接种蚜虫后CsRBOHs家族的转录组数据柱状图；接种蚜虫6h、24h、72h后9个基因表达水平，72h CsRBOHD基因明显上调。
图4是植物病毒介导载体pV190重组载体的质粒图谱；
图5是VIGS技术成功沉默CsRBOHD；注射子叶3周，植株叶片发生白化，即实验技术成功，注射沉默CsRBOHD植株发生基因沉默功能。
图6是荧光定量验证基因沉默后CsRBOHD表达显著下降；未沉默基因植株和已沉默基因CsRBOHD植株的抗蚜性鉴定；定量数据分析CsRBOHD基因表达水平下降50%，基因沉默后植株蚜虫数量是对照的1.75倍。
图7是CsRBOHD沉默后黄瓜叶片DAB染色。
图8是对照和沉默CsRBOHD基因的植株抗蚜性鉴定。
具体实施方式
下面结合附图以及附图说明对本发明做进一步说明。
实施例
GO富集分析
通过转录组测序后的GO富集分析找到与氧化水平相关的通路，通过CsRBOHs家族基因（CsRBOHD、CsRBOHB、CsRBOHFa、CsRBOHFc、CsRBOHE、CsRBOHFb、CsRBOHJa、CsRBOHJb）在接虫前后的基因表达差异，绘制柱状图。
实施例
VIGS技术
1、 通过RACE技术克隆CsRBOHD基因
基于黄瓜CDS序列及其蛋白保守结构域，设计引物，以为黄瓜叶片总RNA为模板，通过反转录得到CsRBOHD的cDNA。
I .引物设计
CsRBOHD RACE引物：
正向引物：GATTCGACCGGACTAAATCTGC
反向引物：GTTAGCTGAAGCACTCATACTAATAATC
II .RACE反应及载体重组过程
(1) RNA提取及鉴定
用“Superina”材料200 mg叶片在液氮中迅速研磨成粉末，取30-150 mg样本用RNAEasy Fast Plant Tissue Kit 试剂盒得到RNA溶液浓度鉴定或者存放-80℃；
(2) 反转录
上述得到的RNA用Vazyme HiScriptⅢ 1st Strand cDNA Synthesis Kit（+gDNAwiper）试剂盒逆转录为从cDNA；
将以下成分加入到一个置于冰上的RNase-free的小离心管中：1μgTotal RNA（一般2μL）, 6μL RNase-free ddH2O补至8μL;轻轻混匀，短暂离心，65℃加热5min，迅速置于冰上骤冷，并在冰上静置2min。
在上一步的混合液加2μL 5×gDNA wiper Mix；用移液器轻轻吹打混匀，42℃加热2min；
将在上一步的混合液加入体系：2μL 10×RT Mix，HiScriptⅢEnzyme Mix，2μLOligo(dT）20VN, RNase-free ddH2O补至20μL；
轻轻混匀，短暂离心，37℃温育45min，85℃加热5sec；进入PCR步骤，或-20℃保存反应物，并在半年内使用；长期存放建议分装后放-70℃，cDNA应避免反复冻融。构建载体。
（3）克隆目的片段（PCR）：
PCR反应体系为：25μL 2×Phanta Max Buffer（含有Mg2+，终浓度2μM）, 1μL dNTPMix(10mM), 1μL模板DNA，2μL Forward primer，2μL Reverse primer，1μLPhanta MaxSuper-Fidelity DNA Polymerase，ddH2O补至50μL。
反应程序：95℃3min，95℃15sec，58℃15sec，72℃60sec，35cycles，72℃5min。
（4）质粒扩繁
在50mL离心管，加入10mL LB液体培养基，千分之一的Kan，20μL PV190菌液（LiuMei et al., 2020），37℃，200r过夜培养；一是在超净工作台内操作，用灭菌1.5mL离心管加200μL 70%甘油，800μL菌液，放-80℃进行保菌；二是采用TIANGEN质粒提取试剂盒，提质粒进行酶切反应。
采用上述方法提PDS（八氢番茄红素脱氢酶）质粒（Liu Mei et al., 2020）；八氢番茄红素脱氢酶（Phytoene desaturase，PDS）是类胡萝卜素合成途径中的一个关键酶，当PDS 表达受阻时，植物便丧失了类胡萝卜素的光保护作用，从而呈现出白化效应。常以此作为实验操作成功与否的参照。
（5）酶切（载体线性化）。
酶切体系为：10μL 10×NEB Buffer，1μg质粒，5μL BamHⅠ，ddH2O补至100μL，30℃温热30min。
产物纯化
上述产物进行凝胶电泳检测，切取较亮的条带，采用TIANGEN产物纯化试剂盒进行纯化操作。
（6）目的片段与载体重组反应
反应体系为：（0.04×416bp）目的片段，[（0.02×10000）/载体浓度]载体，2μLExnaseⅡ；
反应程序：37 ℃ 30min。
如图4所示：植物病毒介导载体pV190质粒图谱；
2.转化
I .大肠杆菌转化
将新鲜制备或-70 ℃冻存的大肠杆菌TOP10感受态细胞在冰上融化；将100 μL感受态细胞分为两管，取5 μL纯化片段与克隆载体的连接产物，加入到一管50μL感受态细胞中，并轻轻混匀，冰浴30 min左右；42 ℃水浴中热击45 sec，迅速置于冰上2 min；加入700μL LB液体培养基，37℃&200 rpm摇菌1 h；5000 rpm离心3min，吸掉上层670 μL LB液体培养基，混匀剩余菌液；将菌液涂抹于含有Kan的LB筛选培养板上，37℃倒置培养过夜。
II .阳性筛选及测序分析
(1)摇菌：从筛选培养板上挑选单菌落接种于1ml LB液体培养基中，37℃，200rpm，摇菌过夜；直 接以培养过夜的菌落为模板进行重组转化子的PCR检测。
(2)反应程序：
反应体系为：10 μL 2×Rapid Taq Master Mix，1 μL载体上游引物，1 μL目的基因下游引物，挑取单菌落，ddH2O补至20 μL.
(3)过夜培养：在含有Kan抗生素 LB固体培养基上划线，放在37 ℃培养箱，板子倒放，12h；
(4)反应程序：95℃3min，95℃15sec，55℃15sec，72℃60sec，35cycles，72 ℃5min。
(5)菌落PCR检测为阳性的菌液送擎科生物科技有限公司测序。
Ⅲ.保菌、提质粒
保菌：待划线板子长出菌，挑单菌落在1.5mL离心管含有1mL千分之一的Kan的LB液体培养基进行过夜培养，然后加70%甘油，混匀放-80℃冰箱保存；
采用TIANGEN试剂盒对菌液进行质粒提取，进行农杆菌转化或者放-20℃。
Ⅳ.GV3101农杆菌转化
以下操作置于冰上操作：将-70℃冻存的农杆菌GV3101感受态细胞在冰上融化，取1000ng纯化片段与载体的连接产物，加入到100μL感受态细胞中，轻轻摇动混匀，依次冰上5min，液氮5 min，37℃水浴5 min，冰浴5 min；加入700 μLLB液体培养基，28℃，200r，2-3h；
6000 rpm离心1min，去除大部分LB液体培养基，剩余100 μL用移液枪轻轻吸打混匀，用涂布棒在含有抗生素Kan+Rif的LB固体培养基上；倒放28℃培养箱2-3d；
Ⅵ.菌落PCR鉴定
(1)操作与上述大肠杆菌菌落PCR程序一致；
(2)筛选出的阳性克隆进行摇菌、保菌。
Ⅴ.注射黄瓜子叶：
上述获得的农杆菌经过两次摇菌，第一次摇菌体系：5mLLB液体培养基，千分之一抗生素（Kan+Rif），150 μL菌液，28℃，200r过夜培养；第二次摇菌体系：150mLLB液体培养基，千分之一抗生素（Kan+Rif），150μL第一次摇的菌液，28℃，200r过夜培养。
(1)配置重悬浮液：1L dH2O，0.02gAS（乙酰丁香酮）用1mL酒精溶解，0.4g半胱氨酸，5mLTween 20，用磁力搅拌器500r 1h至全部溶解成透明；将上述菌液倒入50mL离心管，4000rpm离心5min，倒掉上清，用悬浮液调OD值约0.6-0.8，28℃黑暗2-3 h；
(2)注射：
采用5 mL针管吸取悬浮液注射在黄瓜子叶。
17-21d，观察注射PDS的黄瓜植株白化，即操作成功。
实施列3
荧光定量技术：
通过荧光定量PCR技术探究CsRBOHD基因沉默后的表达量，基于CsRBOHD基因的外显子区域设计荧光定量PCR引物正向引物5’- TCGCCGTTCATAGCGTTCAT -3’和反向引物5 '-CGAAGAACAGAGGCACGGAA - 3’，基 于 A c t i n 基 因 设 计 内 参 引 物 正 向 引 物5’- GCTGGATTCTGGTGATGGTG-3’和反向引物5’-AGCAAGGTCCAAACGGAGAA-3’。利用SYBRPremix Ex Taq(Takara公司)和荧光定量PCR仪q225(酷博公司)，参照生产商说明书PCR体系和程序，进行荧光定量PCR获取达到荧光阈值的循环数，计算出沉默CsRBOHD基因后的相对表达量。
如图5所示：CsRBOHD基因沉默后植株白化和基因表达量；
实施例4：黄瓜叶片被蚜虫刺吸前后的 DAB染色
通过使用DAB染液验证活性氧（ROS）水平，基于过氧化氢是活性氧的一种，过氧化氢能与DAB（3,3-二氨基联苯胺四盐酸盐）迅速反应生成棕红色化合物，从而定位组织中的过氧化氢，反应ROS水平。
如图7所示：CsRBOHD基因沉默后黄瓜叶片DAB染色；
实施例
对未沉默CsRBOHD基因的“Superina”、沉默CsRBOHD基因的“Superina”进行10 d后的蚜虫繁殖数量统计进行抗蚜性鉴定。利用GraphPad软件，参照软件说明和作图类型，对各自选取6株长势一致的黄瓜幼苗，进行数据统计，分析沉默CsRBOHD基因后的抗性差异，结果表明CsRBOHD沉默后蚜虫数量增多。
如图8所示：未沉默CsRBOHD基因和沉默CsRBOHD基因的植株抗蚜性鉴定。
黄瓜抗蚜调控基因CsRBOHD核苷酸序列为：
atgagacccc acgaacctta ttccggcaac aattccgacg ccgagagcgt cagtagcgtc
agaaggggtg accggagagc tttcagcggt ccaatttctt cttccactac caagcctagg
aagaatgcta aattcgacct ctcttcttct tcttcctccc tcaaagccgc cgatgacgac
gacacctatg ttgaaatcac ccttgatatc cgtgacgact ctgtcgccgt tcatagcgtt
cataccgccg gtcccggtca ggatcccaac tctcttgaag accctgaact ttcccttctc
gctaaacgaa ctctcgagaa gaaatcgtcc tctttccgtg cctctgttct tcgttccact
tcttcccgta tcaaacaggt ctctcaagag cttaaacgct tcacctcact taaccgccgc
acctccactc gccgattcga ccggactaaa tctgctgcca cccatgctct caaggggatg
aagtttatta ctgccaaaac cggcggtggt ggatcctccg ccggttgggc tccggttgag
aagcgattcg atgaactcac cgcctctacc aacggactcc tcccttcttc cttgtttgga
cagtgcattg gaatgaacaa agaatcgaaa gatttcgccg gtgagctttt ccgtgcactt
gctcgtaggc gtaacattac tggtgattcc atcaataaag ctcaacttaa agagttctgg
gatcagatct ccgatgacag cttcgattcc aggctacaaa ctttctttga catggtcgat
acggatgctg atggaagaat cacagaagaa gaagtggaag agattattag tatgagtgct
tcagctaacc aactttccac aattcaaaaa caagcaaaag aatacgctgc cttgatcatg
gaagaactcg accctggaaa tgctgggtac atcatgatac aaaatttgga aactctgtta
ctacaagctc caaatcaatc agtaagagtg agtgacagcc gagttctaag tcaattacta
agccaaaaac tgaagccaac aaacgaaacc aaccccatca taagaacata cgacaagttc
ctgtactttg tagaagacaa ttggcagagg atttgggttt tattgctctg gctcggcatc
tgcgcgggtc tcttcgctta caaattcatc caataccgta acagagcagt gttcaacgtc
atgggttact gtgtttccat cgctaaagga ggagcagaaa ctctaaaatt caacatggct
ctaattttac tccccgtttg tcgtaacacc atcacatggc ttcgtaacaa aaccaaactg
ggcctcatcg tacctttcga tgacaatctg aatttccata aagttatagc agttggaatt
tctgttggag tgggattaca cgcaattgct catttggctt gcgatttccc tcggcttctt
cacgcaacag aggaagaata tgaaccactg aagcgattct ttggagagga acagcctgat
aattattggt ggtttgttaa aggagtggaa ggtgttacag gcattataat ggttgtatta
atggccattg ctttcacatt agccactcca tggtttagaa gaaacaaact caaagtaccc
aaacccttga agaaactcac tggctttaat gccttttggt actcccacca tctcttcgta
gccgtctaca ccctcctggt cgtccacggt atctacctct acctgaccaa ggaatggtat
aagaaaacga cgtggatgta tttggctgta ccagtattac tttacgggtg cgagagattg
ataagagcat tcagatcagg gatcaagcca gtgaagattc ttaaggtggc cgtttatcct
ggaaacgttc tggcattgca catgtctaag ccacatgggt tcaaatacaa gagtgggcag
tacatgtttg ttaactgcag agatgtttcc ccattcgagt ggcatccatt ttcaattact
tcggctccag aagataatta tttgagtgtt catattcgaa cacttggtga ttggacaagg
aaacttaagg atgtattctc agaggtttgt cagcctcctc aagctggtaa aagtggactt
cttagagctg aatttttgca aggaggagct cctaatccca agtttccaaa gatattgatt
gacggtccat atggggcacc agcacaagac tacaaaaaat acgacgtggt attactagtg
ggtttgggaa ttggggcgac accgatggtg agtatagtaa aagacataat cgacaacata
gaggagaaag aatcagaagc aaatgcagta gaaaatgggc aagggcatag cagtcgagga
ggaagcaagc acggaaaagg attccgaacg aaaaaggcat acttttactg ggtgacaaga
gagcaagggt cattcgaatg gtttaaaggg ataatgaacg aggtagcaga aatggacgag
agaggggtaa tcgagctgca caactactgc acgagcgtgt acgaagaagg tgacgctagg
tccgcattaa taacaatgct tcaatcatta caccatgcta agaacggagt ggatgtggta
tcagggacta gggtaaagtc acattttgcc aagccaaatt ggaggcaagt ttataaaaag
attaccctcc atcatccaga tacaaaagtt ggtgtgtttt actgtggaac accagtactt
acaaaggagc ttagtcaatt ggcatctgat ttcactcgta aaacatcaac taagtttgag
tttcataaag agaatttcta g
黄瓜抗蚜调控基因CsRBOHD的表达蛋白，氨基酸序列为：
Met Arg Pro His Glu Pro Tyr Ser Gly Asn Asn Ser Asp Ala Glu Ser
Val Ser Ser Val Arg Arg Gly Asp Arg Arg Ala Phe Ser Gly Pro Ile
Ser Ser Ser Thr Thr Lys Pro Arg Lys Asn Ala Lys Phe Asp Leu Ser
Ser Ser Ser Ser Ser Leu Lys Ala Ala Asp Asp Asp Asp Thr Tyr Val
Glu Ile Thr Leu Asp Ile Arg Asp Asp Ser Val Ala Val His Ser Val
His Thr Ala Gly Pro Gly Gln Asp Pro Asn Ser Leu Glu Asp Pro Glu
Leu Ser Leu Leu Ala Lys Arg Thr Leu Glu Lys Lys Ser Ser Ser Phe
Arg Ala Ser Val Leu Arg Ser Thr Ser Ser Arg Ile Lys Gln Val Ser
Gln Glu Leu Lys Arg Phe Thr Ser Leu Asn Arg Arg Thr Ser Thr Arg
Arg Phe Asp Arg Thr Lys Ser Ala Ala Thr His Ala Leu Lys Gly Met
Lys Phe Ile Thr Ala Lys Thr Gly Gly Gly Gly Ser Ser Ala Gly Trp
Ala Pro Val Glu Lys Arg Phe Asp Glu Leu Thr Ala Ser Thr Asn Gly
Leu Leu Pro Ser Ser Leu Phe Gly Gln Cys Ile Gly Met Asn Lys Glu
Ser Lys Asp Phe Ala Gly Glu Leu Phe Arg Ala Leu Ala Arg Arg Arg
Asn Ile Thr Gly Asp Ser Ile Asn Lys Ala Gln Leu Lys Glu Phe Trp
Asp Gln Ile Ser Asp Asp Ser Phe Asp Ser Arg Leu Gln Thr Phe Phe
Asp Met Val Asp Thr Asp Ala Asp Gly Arg Ile Thr Glu Glu Glu Val
Glu Glu Ile Ile Ser Met Ser Ala Ser Ala Asn Gln Leu Ser Thr Ile
Gln Lys Gln Ala Lys Glu Tyr Ala Ala Leu Ile Met Glu Glu Leu Asp
Pro Gly Asn Ala Gly Tyr Ile Met Ile Gln Asn Leu Glu Thr Leu Leu
Leu Gln Ala Pro Asn Gln Ser Val Arg Val Ser Asp Ser Arg Val Leu
Ser Gln Leu Leu Ser Gln Lys Leu Lys Pro Thr Asn Glu Thr Asn Pro
Ile Ile Arg Thr Tyr Asp Lys Phe Leu Tyr Phe Val Glu Asp Asn Trp
Gln Arg Ile Trp Val Leu Leu Leu Trp Leu Gly Ile Cys Ala Gly Leu
Phe Ala Tyr Lys Phe Ile Gln Tyr Arg Asn Arg Ala Val Phe Asn Val
Met Gly Tyr Cys Val Ser Ile Ala Lys Gly Gly Ala Glu Thr Leu Lys
Phe Asn Met Ala Leu Ile Leu Leu Pro Val Cys Arg Asn Thr Ile Thr
Trp Leu Arg Asn Lys Thr Lys Leu Gly Leu Ile Val Pro Phe Asp Asp
Asn Leu Asn Phe His Lys Val Ile Ala Val Gly Ile Ser Val Gly Val
Gly Leu His Ala Ile Ala His Leu Ala Cys Asp Phe Pro Arg Leu Leu
His Ala Thr Glu Glu Glu Tyr Glu Pro Leu Lys Arg Phe Phe Gly Glu
Glu Gln Pro Asp Asn Tyr Trp Trp Phe Val Lys Gly Val Glu Gly Val
Thr Gly Ile Ile Met Val Val Leu Met Ala Ile Ala Phe Thr Leu Ala
Thr Pro Trp Phe Arg Arg Asn Lys Leu Lys Val Pro Lys Pro Leu Lys
Lys Leu Thr Gly Phe Asn Ala Phe Trp Tyr Ser His His Leu Phe Val
Ala Val Tyr Thr Leu Leu Val Val His Gly Ile Tyr Leu Tyr Leu Thr
Lys Glu Trp Tyr Lys Lys Thr Thr Trp Met Tyr Leu Ala Val Pro Val
Leu Leu Tyr Gly Cys Glu Arg Leu Ile Arg Ala Phe Arg Ser Gly Ile
Lys Pro Val Lys Ile Leu Lys Val Ala Val Tyr Pro Gly Asn Val Leu
Ala Leu His Met Ser Lys Pro His Gly Phe Lys Tyr Lys Ser Gly Gln
Tyr Met Phe Val Asn Cys Arg Asp Val Ser Pro Phe Glu Trp His Pro
Phe Ser Ile Thr Ser Ala Pro Glu Asp Asn Tyr Leu Ser Val His Ile
Arg Thr Leu Gly Asp Trp Thr Arg Lys Leu Lys Asp Val Phe Ser Glu
Val Cys Gln Pro Pro Gln Ala Gly Lys Ser Gly Leu Leu Arg Ala Glu
Phe Leu Gln Gly Gly Ala Pro Asn Pro Lys Phe Pro Lys Ile Leu Ile
Asp Gly Pro Tyr Gly Ala Pro Ala Gln Asp Tyr Lys Lys Tyr Asp Val
Val Leu Leu Val Gly Leu Gly Ile Gly Ala Thr Pro Met Val Ser Ile
Val Lys Asp Ile Ile Asp Asn Ile Glu Glu Lys Glu Ser Glu Ala Asn
Ala Val Glu Asn Gly Gln Gly His Ser Ser Arg Gly Gly Ser Lys His
Gly Lys Gly Phe Arg Thr Lys Lys Ala Tyr Phe Tyr Trp Val Thr Arg
Glu Gln Gly Ser Phe Glu Trp Phe Lys Gly Ile Met Asn Glu Val Ala
Glu Met Asp Glu Arg Gly Val Ile Glu Leu His Asn Tyr Cys Thr Ser
Val Tyr Glu Glu Gly Asp Ala Arg Ser Ala Leu Ile Thr Met Leu Gln
Ser Leu His His Ala Lys Asn Gly Val Asp Val Val Ser Gly Thr Arg
Val Lys Ser His Phe Ala Lys Pro Asn Trp Arg Gln Val Tyr Lys Lys
Ile Thr Leu His His Pro Asp Thr Lys Val Gly Val Phe Tyr Cys Gly
Thr Pro Val Leu Thr Lys Glu Leu Ser Gln Leu Ala Ser Asp Phe Thr
Arg Lys Thr Ser Thr Lys Phe Glu Phe His Lys Glu Asn Phe
Claims (2)
1.黄瓜抗蚜调控基因CsRBOHD在调控黄瓜蚜虫抗性中的应用，其特征在于，所述CsRBOHD基因与ROS水平相关；黄瓜抗蚜调控基因CsRBOHD，其核苷酸序列如SEQ ID NO.1所示；黄瓜抗蚜调控基因CsRBOHD的表达蛋白，氨基酸序列如SEQ ID NO .2所示。
2.根据权利要求1所述的应用，其特征在于，VIGS技术沉默CsRBOHD基因，ROS水平降低，黄瓜蚜虫抗性减弱。
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