CN112746073A - 一种通过多基因编辑获得高抗白粉病黄瓜种质材料的方法 - Google Patents
一种通过多基因编辑获得高抗白粉病黄瓜种质材料的方法 Download PDFInfo
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Abstract
本发明提供一种通过多基因编辑获得高抗白粉病黄瓜种质材料的方法,使CsMLO1、CsMLO8及CsMLO11基因产生双突变或三突变或使CsMLO1基因、CsMLO8基因及CsMLO11基因编码的氨基酸共同产生双突变或三突变来提高黄瓜白粉病的抗性;与现有技术相比,本发明采用CsMLO1基因、CsMLO8基因及CsMLO11基因共同调控黄瓜白粉病抗性,可获得白粉病完全抗性材料;与现有技术相比,本发明采用CRISPR/Cas9多基因编辑技术实现CsMLO1基因、CsMLO8基因及CsMLO11基因的共同突变,所获得的种质材料为不含外源转基因片段的无转基因痕迹突变体材料,转基因安全性风险更小。
Description
技术领域
本发明属于生物技术和分子生物学领域,具体涉及通过多基因编辑获得高抗白粉病黄瓜种质材料的方法以及相关黄瓜白粉病隐性抗病蛋白及其编码基因的应用。
背景技术
黄瓜白粉病是一种专性寄生的真菌性病害。近年来,在设施栽培条件下,作物种植密度增高、氮肥施用量增大、高温高湿的环境使白粉病造成的危害日益严重,严重影响了黄瓜的品质和产量。目前黄瓜白粉病的防治依然主要依靠使用化学农药,容易造成农药残留和生态环境污染。随着人们对食品安全和环境保护要求的提高,抗病品种培育成为防治白粉病的首选方法。然而黄瓜白粉病抗性是由多基因位点控制的数量性状,传统育种很难在不引入连锁累赘的情况下有效整合多个抗病基因,因此市面上普遍缺乏优良的抗性品种。为此通过生物学的方法提高黄瓜对白粉病的抗性是今后防治白粉病的有效策略之一。目前黄瓜白粉病抗性基因资源相对匮乏,函需发掘黄瓜抗病基因资源。克隆到新的抗病基因,利用CRISPR/Cas9多基因编辑技术和转基因的方法同时定点敲除多个隐性抗病基因,具有重要的经济意义。
MLO基因最早在大麦中发现,此基因的突变可以使大麦产生持久、广谱、高效的白粉病抗性。mlo隐性突变介导的白粉病抗性随后被广泛应用于其它多种作物的抗病育种中(Kusch and Panstruga,2017)。MLO基因家族成员之间存在功能冗余性,且不同成员突变对白粉病抗性的贡献率不相同,有主效和微效抗病基因之分。在双子叶植物中MLO基因第V分支成员参与介导白粉病抗性(Kusch and Panstruga,2017;Zheng et al.,2016)。黄瓜全基因组范围内CsMLO家族成员共13个,分为5个进化分支,其中第V分支包括三个基因CsMLO1、CsMLO8和CsMLO11(Schouten et al.,2014)。其中CsMLO8定位在白粉病主效QTL位点pm5.1区间内(Berg et al.,2015;Nie et al.,2015a),研究发现在抗病材料中由于转座子插入导致该基因失活突变(Nie et al.,2015b)。利用重测序数据分析发现,CsMLO8功能缺失突变在多个白粉病抗性种质中普遍存在,且异源表达CsMLO8可以恢复番茄mlo突变体的感病性,这些证据表明CsMLO8是pm5.1位点的白粉病隐性抗病基因(Berg et al.,2015;Nie etal.,2015b)。然而CsMLO8失活仅产生下胚轴对白粉病的完全抗性,叶片仅为部分抗性(Berget al.,2015)。
虽然利用传统的杂交方法可以获得多基因突变体,但是构建三个以上的多突变或整合遗传距离较近的多基因突变,其筛选工作量非常庞大,而且容易因连锁累赘而引入不良性状。CRISPR/Cas9多基因编辑技术,通过在载体上装载Cas9元件和多个sgRNA元件,可以实现同时对多个基因定向编辑,对于整合多个基因位点进行分子育种具有重要的意义。另外,在完成靶位点编辑后,通过自交或与野生型杂交可以将Cas9等转基因载体元件剔除,从而得到不含外源片段插入的无转基因痕迹突变体材料。这种经基因修饰的材料与传统的转基因材料不同,只是修饰了物种自身的基因,不引入其他外源基因,转基因安全风险性较小。
发明内容
为解决上述技术问题,本发明提供黄瓜白粉病隐性抗病相关蛋白及其编码基因与应用。
具体技术方案如下:
一种提高黄瓜白粉病抗性的方法,其不同之处在于,使CsMLO1、CsMLO8及CsMLO11基因中的两个或三个共同产生突变;
或
使CsMLO1基因、CsMLO8基因及CsMLO11基因编码的氨基酸中两个或三个共同产生突变来提高黄瓜白粉病的抗性;
所述CsMLO1的基因序列如SEQ ID NO.1所示,所述CsMLO8的基因序列如SEQ IDNO.2所示,所述CsMLO11的基因序列如SEQ ID NO.3所示。
进一步,CsMLO1基因控制的蛋白质的氨基酸编码为SEQ ID NO.13所示,CsMLO8基因控制的蛋白质的氨基酸编码为SEQ ID NO.14所示,CsMLO11基因控制的蛋白质的氨基酸编码为SEQ ID NO.15所示。
与现有技术相比,本发明通过使CsMLO1基因、CsMLO8基因及CsMLO11基因产生双突变或三突变调控黄瓜白粉病抗性,可获得白粉病完全高抗性材料。
进一步,使CsMLO1及CsMLO8基因共同产生突变或使CsMLO1及CsMLO8基因编码的氨基酸共同产生突变或使CsMLO1、CsMLO8及CsMLO11三个基因共同产生突变或使CsMLO1、CsMLO8及CsMLO11基因编码的氨基酸共同产生突变。
采取上述进一步技术方案的有益效果在于:上述突变方式可以使黄瓜的白粉病抗性大幅度增加。
进一步,使CsMLO1、CsMLO8及CsMLO11基因共同产生突变或使CsMLO1、CsMLO8及CsMLO11基因编码的氨基酸共同产生突变。
采取上述进一步技术方案的有益效果在于:三个基因共同突变可以得到黄瓜全抗性植株,且白粉病抗性几乎达到100%。
进一步,本发明通过采用CRISPR/Cas9基因编辑进行基因敲除实现基因突变。
进一步,所述CsMLO1基因发生的突变在CsMLO1靶点片段区域中,所述CsMLO1靶点片段序列为SEQ ID NO.4所示,所述CsMLO8基因发生的突变在CsMLO8靶点片段区域中,所述CsMLO8靶点片段序列为SEQ ID NO.5所示,所述CsMLO11基因发生的突变在CsMLO11靶点片段区域中,所述CsMLO11靶点片段序列为SEQ ID NO.6所示。
采取上述进一步技术方案的有益效果在于:通过在上述片段上突变,进行基因敲除。
一种调控黄瓜白粉病抗性的重组载体,其不同之处在于,所述调控黄瓜白粉病抗性的重组载体包括选自所述CsMLO1基因的所述CsMLO1靶点片段,序列如SEQ ID NO.4所示,选自所述CsMLO8基因的CsMLO8靶点片段,序列如SEQ ID NO.5所示,选自所述CsMLO11基因的CsMLO11靶点片段,序列如SEQ ID NO.6及载体质粒,所述CsMLO1靶点片段、所述CsMLO8靶点片段及所述CsMLO11靶点片段连接在所述载体质粒上。
与现有技术相比,本发明提供了调控黄瓜白粉病抗性的重组载体敲除CsMLO1基因和/或CsMLO8基因和/或CsMLO11基因,得到白粉病抗性良好的双突变基因编辑植株、三突变体基因编辑植株。
进一步,所述CsMLO1靶点片段、所述CsMLO8靶点片段及所述CsMLO11靶点片段之间通过依次连接的gRNA scaffold及TRNA进行串联,所述gRNA scaffold的序列为:
GTTTTAGAGCTAGAAATAGCAAGTTAAAATAAGGCTAGTCCGTTATCAACTTGAAAAAGTGGCACCGAGTCGGTGC;
所述tRNA的序列为:
AACAAAGCACCAGTGGTCTAGTGGTAGAATAGTACCCTGCCACGGTACAGACCCGGGTTCGATTCCCGGCTGGTGCA。
对于多sgRNA的组装,可以采用同一个启动子驱动多个间隔串联的sgRNA,并“借用”生物体内的tRNA加工系统或内含子剪切系统获得独立的sgRNA。
进一步,所述载体质粒为pBCG403。
一种制备上述调控黄瓜白粉病抗性的重组载体的方法,其不同之处在于,所述制备调控黄瓜白粉病抗性的重组载体的方法包括:将扩增引物分别扩增出含所述CsMLO1靶点片段的基因片段、含所述CsMLO8靶点片段的基因片段及含所述CsMLO11靶点片段的基因片段,然后各基因片段与线性化的载体质粒连接后,转入大肠杆菌,经过抽提后得到所述调控黄瓜白粉病抗性的重组载体。
与现有技术相比,本发明的有益效果在于,通过上述方法制备出了用于调控黄瓜白粉病抗性的重组载体,可用于敲除CsMLO1和/或CsMLO8和/或CsMLO11基因。
进一步,所述制备调控黄瓜白粉病抗性的重组载体的方法还包括利用引物模板扩增载体连接片段。
进一步,扩增含所述CsMLO11靶点片段的基因片段的CsMLO11-F引物序列如SEQ IDNO.7所示;扩增出含所述CsMLO1靶点片段的基因片段的上游引物CsMLO1-R序列如SEQ IDNO.8所示,扩增出含所述CsMLO1靶点片段的基因片段的下游引物CsMLO1-F序列如SEQ IDNO.9所示;扩增出含所述CsMLO8靶点片段的基因片段的上游引物CsMLO8-R序列如SEQ IDNO.10所示,扩增出含所述CsMLO8靶点片段的的基因片段的下游引物CsMLO8-F序列如SEQID NO.11所示,扩增载体连接片段的VECTOR-R引物如SEQ ID NO.12所示。
进一步,所述CsMLO11-F引物与所述CsMLO1-R引物为第一引物对,所述CsMLO1-F引物与CsMLO8-R引物为第二引物对,所述CsMLO8-F引物与VECTOR-R为第三引物对,分别进行扩增。
一种得到高抗性白粉病黄瓜的方法,其不同之处在于,所述得到高抗性白粉病黄瓜的方法包括:
步骤S1:将上述重组载体转入农杆菌。
步骤S2:将转入所述重组载体的农杆菌浸染黄瓜子叶。
与现有技术相比,本发明通过向黄瓜子叶中转入重组载体,使CsMLO1基因、CsMLO8基因、CsMLO11基因发生突变,将上述基因敲除。
进一步,所述得到高白粉病抗性黄瓜的方法还包括:
步骤S3:将所述步骤S2中得到的黄瓜植株与未经过处理的黄瓜植株杂交后再自交。
采取上述进一步技术方案的有益效果:通过杂交后再自交可将敲除系统去除,获得不含外源基因的基因编辑植株。
进一步,所述步骤S3中,经编辑位点的基因检测筛选出自交后代中的双突变体和/或三突变体,作为白粉病高抗性黄瓜编辑植株。
进一步,所述步骤S3中,经编辑位点的基因检测筛选出自交后代中的三突变体,作为白粉病高抗性黄瓜编辑植株。
附图说明
图1为CsMLO1、CsMLO8、CsMLO11基因sgRNA表达元件组。
图2为基因编辑的三突变体黄瓜T2代靶标位点编辑情况。
图3为基因编辑的Csmlo1/8双突变体黄瓜T2代靶标位点编辑情况。
图4为在田间发现白粉病粉层后,每隔五天分别对野生型与Csmlo1/8/11三突变体、调查病情级别。
图5为在田间白粉病自然发病情况下,野生型植株与Csmlo1/8/11三突变体植株的叶片感病情况。
图6为在田间白粉病自然发病情况下,野生型植株与Csmlo1、Csmlo8单突变体及Csmlo1/8双突变体植株的叶片感病情况。
具体实施方式
以下结合附图对本发明的原理和特征进行描述,所举实例只用于解释本发明,并非用于限定本发明的范围。
实施例1
1.1 CRISPR/Cas9表达载体的构建及农杆菌转化
1.sgRNA的设计
用Geneious软件选择CsMLO1、CsMLO8、CsMLO11基因(Cucumber(Chinese Long)v3Genome)的外显子区域进行sgRNA的设计:sgRNA序列具有5′-(N)20-NGG-3′结构(N为A、G、C、T四种碱基中的任意一个),且GC含量在40%-70%之间。
黄瓜CsMLO1 sgRNA序列如SEQ ID NO.4所示、CsMLO8 sgRNA序列如SEQ ID NO.5所示、CsMLO11基因的sgRNA序列如SEQ ID NO.6所示。
2.引物设计和PCR扩增
(1)根据CsMLO1、CsMLO8、CsMLO11基因的sgRNA设计引物序列。引物序列如下所示:
引物名称 | 引物序列(5′-3′) |
CsMLO11-F | SEQ ID NO.7 |
CsMLO1-R | SEQ ID NO.8 |
CsMLO1-F | SEQ ID NO.9 |
CsMLO8-R | SEQ ID NO.10 |
CsMLO8-F | SEQ ID NO.11 |
VECTOR-R | SEQ ID NO.12 |
其中,CsMLO11-F、VECTOR-R载体连接引物是根据pBCG403载体用BsaⅠ酶切后产生的粘性末端来设计。
pBCG403载体是在PBSE401载体(Xing,Huili et al.,2014.A CRISPR/Cas9toolkit for multiplex genome editing in plants[J].BMC Plant Biology.2014:14:327)基础上加入洋丁香黄曲叶病毒(CmYLCV)启动子序列、tRNA加工酶基因、花椰菜病毒终止子序列后获得。上海生工公司合成洋丁香黄曲叶病毒(CmYLCV)启动子序列、tRNA加工酶基因、花椰菜病毒终止子序列,共计851bp,连接到HindIII酶切pBSE401载体,得到pBCG403载体。
(2)PCR扩增
以测序正确的pT-tRNA为模板,扩增出sgRNA(扩增引物中包含sgRNA序列)、gRNAScaffold、tRNA(pT-tRNA模板载体上包含gRNA Scaffold、tRNA序列)相连的序列。
其中,pT-tRNA在天根pLB零背景快速克隆试剂盒(QIAGEN,Lethal Based FastCloning Kit,VT205)的T载体基础上加入gRNA Scaffold(Xing,Huili et al.,2014.ACRISPR/Cas9 toolkit for multiplex genome editing in plants[J].BMC PlantBiology.2014:14:327)、tRNA(Xie,Kabin et al.,2015.Boosting CRISPR/Cas9multiplex editing capability with the endogenous tRNA-processing system[J].Proc Natl Acad Sci U S A 112,3570-3575.)序列获得。其中gRNA Scaffold、tRNA序列片段均为上海生工公司合成后连接到T载体上。
其中,CsMLO11-F与CsMLO1-R作为扩增时的第一引物对,CsMLO1-F与CsMLO8-R作为扩增时的第二引物对,CsMLO8-F与VECTOR-R作为扩增时的第三引物对,各引物对分别进行扩增,表中统一用CsMLO-F及CsMLO-R表示,片段长度为212bp、195bp、111bp。
①PCR反应体系与条件:
②琼脂糖凝胶电泳检测:
称取0.5g琼脂糖置于锥形瓶中,加入25mL1×TAE溶液,微波炉加热煮沸至琼脂糖全部融化。将内槽置于水平位置并放好梳子,将冷却到65℃左右的琼脂糖凝胶液倒入内槽玻璃板上。室温下静置直至凝胶完全凝固,垂直轻拔梳子即配制成2%琼脂糖凝胶。将凝胶和内槽放入电泳槽中,用10μL微量移液器将DL 2000 DNA Marker(作为对照)与上述PCR扩增产物加入胶板的小槽内。加样后通电进行电泳,电压设为100V。电泳完毕后取出凝胶,用荧光核酸凝胶染色溶液染色约20min,通过凝胶成像仪进行紫外显影,可观察到用三对引物扩增分别在212bp、195bp、111bp处有一条明亮的条带,说明已成功扩增出了目的片段。
③PCR产物纯化回收:
用PCR产物纯化试剂盒(QIAGEN,QIAquick PCR Purification Kit(50),28104)进行纯化回收。按照说明书操作如下:将PCR产物加入2mL离心管中,加5个PCR产物体积的Buffer PB,混合均匀。将吸附柱置于2mL离心管中,并将上步混合均匀的溶液加入吸附柱中,12000rpm离心1min,弃滤液。将吸附柱置回2mL离心管,加入750μL BufferPE于吸附柱中,12000rpm离心1min,弃滤液。将吸附柱置回2mL离心管,再次12000rpm离心1min,弃滤液。将吸附柱置于洁净的1.5mL离心管中,放置5min。在吸附柱中央加入30μL Buffer EB(加热至65℃以提高洗脱效率),室温静置1min,12000rpm离心1min。获得PCR液体回收产物。3个片段液体回收浓度分别为100ng/μL、113ng/μL、64ng/μL。
3.扩增产物的酶切连接
获得PCR扩增产物后建立如下的酶切-连接体系
第一引物对扩增得到PCR片段1,包括:对应扩增MLO11的sgRNA全长、gRNAScaffold、tRNA、MLO1 sgRNA一半;第二引物对得到扩增PCR片段2,包括:对应扩增MLO1sgRNA的一半、gRNA Scaffold、tRNA、MLO8 sgRNA的一半;第三引物对得到扩增PCR片段3,包括对应扩增MLO8 sgRNA的一半、gRNA Scaffold。通过3个片段搭桥获得由tRNA间隔开来串联起来的sgRNA序列,以此实现将分别靶向3个CsMLO基因的sgRNA串联在一起。
4.转化大肠杆菌
在超净工作台中取5μL连接体系加入100μL大肠杆菌感受态DH5α中,轻弹混匀。冰浴5min,放入42℃水浴锅中热激90s,结束后立刻冰浴3min。加入1mLLB溶液,混匀;37℃、220rpm摇床培养30min。在固体LB+kana平板上均匀涂布约200μL转化菌。平板封口倒置于37℃培养箱中培养1d。待长出单菌落后,挑取单克隆菌斑接入LB+Kana液体培养基中,37℃、220rpm摇床培养30min。将单克隆菌液送擎科生物技术公司测序并用geneious软件进行序列比对,比对正确即说明重组表达载体构建成功(图1)。
实施例2
2.1农杆菌的转化
将擎科公司返回的测序正确的表达载体质粒转入农杆菌EHA105中,为后续黄瓜遗传转化侵染实验作准备。在超净工作台中取2μL质粒,加入100μL的EHA105农杆菌感受态细胞中,轻弹混匀。混匀后冰浴5min,放入液氮中冷冻1min,后迅速置于37℃水浴锅中水浴5min。加入800μL LB培养基,于28℃、220rpm摇床上培养2h。12000rpm离心1min以浓缩菌液,加入200μL LB液体培养基重悬,混匀,将其均匀涂在LB+Kana的固体培养基中,28℃暗培养2d。挑出农杆菌单克隆菌斑用LB+kana+rif液体摇菌12h,将菌液置于-80℃超低温冰箱中保存备用。
2.2黄瓜稳定遗传转化
取CU2黄瓜种质(Hu,Bowen et al.,2017.Engineering Non-transgenicGynoecious Cucumber Using an Improved Transformation Protocol and OptimizedCRISPR/Cas9 System[J].Molecular Plant.2017:1575–1578)于55℃温水中浸种半小时,除去种皮。在超净工作台中先用75%的酒精清洗30s,再用0.3%的次氯酸钠溶液浸泡15min,最后用无菌水冲洗5次。将消毒好的种子转移至种子萌发培养基M0上。于28度培养箱中暗培养24h,即可切取外植体。
吸取10μL农杆菌于30mLLB(含Kana、rif)培养基中28℃、220rpm摇床培养过夜。次日当菌液生长至OD600值为0.4–0.8时,6000rpm离心8min来收集农杆菌,并用M1液体培养基(即不加植物凝胶的M1培养基)重悬菌体并稀释到OD600为0.2。取已萌发的种子,在超净台中切去远端部分子叶并去掉胚轴,同时将两片子叶分开,即得到外植体。将切好的外植体放入重悬菌液中,在功率为100W的水浴超声波中超声30s。将超声后的子叶外植体和重悬液加入20mL注射器的针筒中,向前推动活塞直至10mL刻度位置处,用橡胶皮塞密封住注射器的头部针孔,向后方拔动活塞芯杆使活塞停留在20mL刻度处,保持1.5min后使活塞回到10mL刻度位置。之后再重复施加真空负压一次。侵染结束后,将外植体放到滤纸上吸干并将其转移到垫一层滤纸的共培养培养基M1上。于23℃,黑暗条件下共培养4天。在荧光体式显微镜下观察GFP的发光情况,评估侵染效率。
用无菌水清洗共培养后的外植体7-8遍,用灭过菌的吸水纸吸干表面附着的液体,将外植体斜插于恢复培养基M2上,恢复培养7天后更换到分化培养基M3上,两个星期后继代一次。在体式荧光显微镜下,挑选含有GFP荧光芽的子叶。继代到生根培养基M4的组培瓶中培养,直至生根。将生根良好的转基因阳性苗移栽到灭过菌的基质中,于光照培养箱中炼苗和蹲苗一个月,之后在人工气候室中培养直至开花。由于田间栽培管理原因,未能收到T1代自交瓜;所以用转基因T0代和野生型杂交收获的种子作为T1代,T1代自交留种分离得到T2代非转基因三突变体(pBCG403载体中含有独立的GFP表达框,利用GFP荧光辅助从T1代自交留种后代中筛选到无转基因痕迹的突变体)。
上述培养基中,M0培养基成分:MS 4.43g/L+蔗糖30g/L+植物凝胶3g/L+6BA 2mg/L+ABA 1mg/L,pH为5.6-5.8;
M1培养基成分:MS 4.43g/L+蔗糖30g/L+植物凝胶3g/L+6BA 2mg/L+ABA 1mg/L+AS80mg/L+MES 2.5mM,pH为5.6-5.8;
M2培养基成分:MS 4.43g/L+蔗糖30g/L+Agar 8g/L+6BA 2mg/L+ABA 1mg/L+TMT200mg/L,pH为5.6-5.8;
M3培养基成分:MS 4.43g/L+蔗糖30g/L+植物凝胶3g/L+6BA 2mg/L+ABA 1mg/L+PPT 2mg/L+TMT 200mg/L,pH为5.6-5.8;
M4培养基成分:MS4.43g/L+蔗糖30g/L+Agar 8g/L+TMT 200mg/L。
实施例3、基因编辑黄瓜植株DNA的提取与检测
3.1、基因编辑黄瓜植株DNA的提取
采用CTAB法提取实施例2中的T2代转基因黄瓜叶片DNA。具体包括以下步骤:取少量的黄瓜叶片约0.2-0.5g于2mL离心管中,加入干净的钢珠和1mL2%CTAB溶液,用磨样机研磨至粉末状(50Hz,30s)。将研磨后的样品置于65℃中水浴1h。期间倒转离心管数次。25℃,12000rpm离心10min。吸取上清液(约800μL)到新的2mL离心管中。在通风橱中加入等体积的氯仿/异戊醇(24:1)并颠倒混匀,12000rpm(4℃)离心15min。小心吸取上清液(约600μL),加入到已加入两倍体积(约1.2mL)-20℃预冷的无水乙醇的2mL离心管中,充分静置至白色团状沉淀出现。4℃冷冻离心机中10000g离心2min,弃上清液。用70%的乙醇冲洗两次后,再用无水乙醇冲洗一次,用枪头洗净液体,最后放在超净工作台上吹干。加入约50μLddH2O溶解DNA,37℃孵育1h。ddH2O溶解好的DNA放置于-20℃的冰箱中备用。
3.2、阳性单株基因编辑位点检测
以提出的转基因植株DNA为模板,分别利用引物MLO1-cexu-F、MLO1-cexu-R,MLO8-cexu-F、MLO8-cexu-R,MLO11-cexu-F、MLO11-cexu-R对包含CsMLO1、CsMLO8、CsMLO11基因三个靶标位点的区域进行扩增。挑选12个植株DNA样品PCR扩增产物送测,通过二代测序方法检测转基因黄瓜编辑类型。用Geneious软件对测序序列进行分析,发现CsMLO1、CsMLO8、CsMLO11基因被成功编辑,功能缺失的等位基因型相对于野生型表现为隐性(图2)。检测CsMLO1、CsMLO8、CsMLO11基因靶位点编辑效率所设引物序列如下所示:
引物名称 | 引物序列(5′-3′) |
MLO1-cexu-F | GCAAGGACCAATAACGGAGATATG |
MLO1-cexu-R | GGGCACCATCTTTGTACATTTGGACATG |
MLO8-cexu-F | GGTAGGAGGCGCATCTAGTCAC |
MLO8-cexu-R | CAGGCCTTCCATTTGCTCATCTAC |
MLO11-cexu-F | CCTACGCGTTCTAAAGTCAAAGTCATC |
MLO11-cexu-R | CCAGAGCTTCGTAGAGAGCTCG |
3.3筛选基因编辑植株
根据基因编辑位点的检测结果挑选双突变及三突变体,得到Csmlo1/8双突变体,Csmlo1/11双突变体、Csmlo8/11双突变体及Csmlo1/8/11三突变体。
对比例1
采取实施例1的CRISPR/Cas9基因敲除系统,基因敲除系统的转入、植株选育过程均与三突变体相同,不同的是本对比例经编辑位点检测,在T2代分离了仅CsMLO1靶点片段编辑的植株,得到Csmlo1单突变体。
对比例2
采取实施例1的CRISPR/Cas9基因敲除系统,基因敲除系统的转入、植株选育过程均与三突变体相同,不同的是本对比例经编辑位点检测,在T2代分离了仅CsMLO8靶点片段编辑的植株,得到Csmlo8单突变体。
实施例4、基因编辑黄瓜白粉病抗性鉴定
为了确定基因编辑黄瓜的白粉病抗性,我们将野生型和Csmlo1/8/11三突变体、Csmlo1单突变体、Csmlo8单突变体、Csmlo1/8的双突变体,随机种植在大棚中,在田间白粉病自然发病的情况下,对野生型和Csmlo1/8/11三突变体进行了病情指数调查。病情指数调查具体如下:在大棚中随机挑选4个调查样点,每个样点调查3株,野生型、三突变体、Csmlo1单突变体、Csmlo8单突变体、Csmlo1/8的双突变体分别调查8株;待发现白粉病粉层后每隔五天调查1次,每次分为上、中、下三个部位各取3片叶片进行调查并记录病情级别。病情级别的划定为6个级别(0,1,3,5,7,9)。0,叶片干净,没有病斑发生;1,少量细小模糊的白粉斑,病斑面积占叶片面积小于5%;3,白粉层薄,病斑面积占叶片面积大于5%且小于30%;5,白粉层厚,病斑面积占叶片面积大于30%且小于50%;7,白粉层厚,病斑面积占叶片面积大于50%且小于70%;9,白粉层厚,病斑面积占叶片面积大于70%且小于90%。调查并记录野生型和三突变体病情级别后用以下公式计算病情指数:病情指数=[Σ(各级病叶数×各级代表值)/调查总叶数×最高级代表值]×100。
如图4所示,发现随着时间的增长,田间野生型植株白粉病愈来愈严重,病情指数逐渐升高,Csmlo1及Csmlo8单基因敲除后,其发病情况相较于野生植株相差无几,而Csmlo1/8/11三突变体上下叶片几乎无白粉病病斑,获得了完全白粉病抗性,病情指数一直维持在较低的水平。显示只有CsMLO1、CsMLO8、CsMLO11基因共同突变后,才能使黄瓜对白粉病菌的抗病性大幅度增加。
其中,图5为三突变体与野生植株感病情况的对比,a、c为野生型植株;b、d为Csmlo1/8/11三突变体植株。可以看出野生型植株叶片白粉病较重,而Csmlo1/8/11三突变体植株叶片无白粉病病斑,图中a、b标尺为5cm;c、d标尺为2cm。
图6为Csmlo1单突变体、Csmlo8单突变体、Csmlo1/8双突变体与野生植株感病情况的对比。
综上所述,本发明提供了一种通过基因编辑手段创制具有白粉病高抗性黄瓜的方法,通过对CsMLO1、CsMLO8、CsMLO11基因的编辑,筛选出双突变体及三突变体,从实施例4效果检测可以发现,双突变体与三突变体相较于野生型与单突变体,其白粉病抗性大幅度提高。
不仅如此,三突变体在统计样本中,其抗性可以几乎达到100%。
以上所述仅为本发明的较佳实施例,并不用以限制本发明,凡在本发明的精神和原则之内,所作的任何修改、等同替换、改进等,均应包含在本发明的保护范围之内。
序列表
<110> 华中农业大学
<120> 一种通过多基因编辑获得高抗白粉病黄瓜种质材料的方法
<160> 15
<170> SIPOSequenceListing 1.0
<210> 1
<211> 1749
<212> DNA
<213> 黄瓜(cucumber)
<400> 1
atggcggggg cagccggtgg caagtcgctg gagcaaacac cgacatgggc cgttgccgtt 60
gtttgctttg ttttgctcgt catctctatt ttcatcgaat atagtctcca tcttatcgga 120
cattggctaa agaagagaca caaacgggcg ttgtttgaag cattagagaa gatcaaatca 180
gagcttatgt tattggggtt tatatcattg ctactaacgg tggggcaagg accaataacg 240
gagatatgta ttccacaaca tgtagctgca acgtggcatc catgtacaaa ggaaagagaa 300
gatgagatga acaaagaggt ggagaaatct gtggaacatt tgggtcttaa tcgccggaga 360
ctccttcatc tcctcggaaa tggtgaaagt ttccggcgga gtttggccgc tgcgggagga 420
gaggataaat gtgccgccaa gggtaaagct tcctttattt cagcagatgg aattcatcaa 480
cttcatatct tcatttttgt gttggctgtt tttcatgttt tgtattgtgt tctaacttat 540
gcgttggcta gagctaagat gaggagttgg aaaacatggg aaaaagagac caaaactgct 600
gaataccaat tctcacatga tccagagagg tttaggtttg caagagacac ctcatttggg 660
agaagacatt tgagcttttg gaccaaaaat cctgccttga tgtggatcgt ttgtttcttc 720
agacaatttg taagatctgt tccaaaagtt gattacttga cattaagaca tgggtttata 780
atggcacatt tagcacctca aagtcataca caatttgatt ttcaaaaata cattaataga 840
tcccttgaag aagacttcaa agttgttgtg ggaatcagcc caccaatttg gttctttgct 900
gttctatttc tcctctcaaa cactcacggt tggagggcgt atctatggct gccattcatc 960
ccactaatca ttttgctgtt gattggaaca aaattgcaag tgatcataac gaaaatggca 1020
ctaagaatac aagaaagagg tgaagtagtg aagggcgtgc cggtggtgga gcctggcgat 1080
gacctctttt ggtttaatcg acctcgcctt attctttatc tcatcaactt tgttctcttt 1140
caaaatgcct tccaagttgc cttctttgct tggacttggt atgagtttgg gttgaattct 1200
tgcttccatg agcatataga agatgtggtg atcagaattt ctatgggggt gcttgtacaa 1260
atcctttgca gttatgttac tcttcctctt tatgcactag tcactcagat gggttcaaca 1320
atgaagccaa ctatattcaa tgagagagtg gcagaggccc ttcgcaattg gtaccactcg 1380
gctcgaaagc acatcaaaca caaccgcggt tcggtcactc caatgtcgag ccgacccgcc 1440
accccgactc acagcatgtc acctgtccac cttctccgac actacaagag tgaagtcgat 1500
agcttccaca cctcaccgag aaggtcaccg ttcgacaccg atcgttggga caacgattcg 1560
ccctctccat ctcgccatgt tgatggttcg tcttcgtcac aaccccacgt tgagatggga 1620
ggttatgaaa aagatcccgt tgaatcaagt tcgtctcaag ttgatccggt tcaaccatct 1680
cgaaaccgca atcaacatga gattcatatt ggaggcccca aagacttttc atttgataga 1740
gttgaatga 1749
<210> 2
<211> 1725
<212> DNA
<213> 黄瓜(cucumber)
<400> 2
atggctgaat gtggaacaga gcagcgtact ttggaagata cctcaacttg ggctgttgcg 60
gttgtttgtt ttttcttggt tgttatttca atcttcattg aacatgtcat tcacctcact 120
ggaaagtggc tggagaaaag gcacaagcca gctcttgttg aagctctaga aaaggttaaa 180
gcagagctta tgctattggg attcatatcc ctacttctaa cgataggcca agatgctgtc 240
actcaaattt gtgtttcgaa agagcttgca gcaacttggc ttccctgtgc agcaagagct 300
aaaacaggag taaaagttgc gaagaacagt cgtcttagac ttcttgaatt tttagatcct 360
gactatggtt cgaggcgtat tttagcctcg aaaggagatg atgcatgcgc taagaggggc 420
caactcgctt tcgtgtcggc atatggaatc catcagctcc atattttcat cttcgtattg 480
gctgtcttcc atgtcctgta ctgcatcata actttggctt ttggcagaac aaagatgagc 540
aaatggaagg cctgggagga tgaaaccaag acaattgaat accagtacta taatgatcca 600
gcaagattta gatttgctag agatactacg tttggacgcc gacacttgag cttctggagt 660
cgtacaccaa tttccctctg gattgtttgt ttcttcagac agttctttgg atcagttacc 720
aaggttgatt acatgacact gagacatgga ttcattgttg cacatcttgc acccggaagt 780
gaagtaaaat ttgatttcca caaatacatt agcagatctc tggaagacga ctttaaagtt 840
gttgtgggga ttagtcccgc aatgtggcta tttgctgttc tcttcatcct aaccaataca 900
aatgggtggt attcatatct atggctgcct ttcatctcct taattataat tctattggtg 960
ggaacaaagc tccatgttat tataactcat atgggattga caattcaaga aaggggtcat 1020
gttgtgaagg gtgttcccgt cgttcagcct cgggatgacc tgttttggtt tggacgtcca 1080
caacttattc tcttcctgat ccactttgtt ctctttatga atgcatttca gcttgccttc 1140
tttgcttgga ccacatatgc atttaagtgg atgggttgtt tccatcagcg agttgaagat 1200
attgtcatca gactctcaat gggggttatc atacaagttc tctgcagtta tgtcacactc 1260
ccactctatg ctttggttac tcagatgggc tctaacatga gaccaaccat tttcaacgac 1320
cgagtggcga cggcattgaa gaactggcac cactcagcca agaagaacat gaagcagcac 1380
cgcaacccag acagtacctc accattctca agcaggccag ctactccaac tcacggcatg 1440
tctcctattc accttctgca caaacatcag catggcagca catctcccag gctatccgat 1500
gccgaacccg atcgttggga agagttgcct ccttcttcac accatagtag agccccccat 1560
catgataatc atcaagatca acaagaacaa tctgagacaa taattagaga acaggagatg 1620
acagttcaag gaccaagttc aagtgaaacc ggttccataa cacgtcctgc tcgccctcat 1680
caggaaatca ctaggactcc atcagacttc tcatttgcca aatga 1725
<210> 3
<211> 1782
<212> DNA
<213> 黄瓜(cucumber)
<400> 3
atggccggag gtggcgccgg aaggtccttg gaagagacgc cgacatgggc cgtcgccgcc 60
gtgtgctttg ttttggttct gatttctatt atcatcgaac acattctcca tctcatcgga 120
aagtggctaa agaagaaaca caaacgagct ctctacgaag ctctggagaa gattaaatca 180
gaactgatgc tgttgggatt catatcgctg ctgctgacgg tgggacaaag cctaatcaca 240
aatgtttgta taccacctga cgtggcagcc acgtggcatc catgtagtcc tcaaagagaa 300
gaagaattaa ctaaagaagc tgacctcgtc gattccgacc aaaatcgtcg aaaacttctc 360
gccctctccc atcacgtcaa cgccaccttc cgccgttccc tcgccgctgc cggtggtacc 420
gacaaatgtg ctgccaaggg taaagttcca tttgtatcgg aagggggtat tcatcagcta 480
catatattca tcttcgtact ggcagttttc catgttttgt attgtgtttt aactttagct 540
ttgggcaatg ccaagatgag aagttggaag tcatgggaaa aagagacaag aactgtggag 600
tatcaattct cacacgatcc ggaacggttt cgatttgcaa gagacacgtc atttgggaga 660
agacatttaa gcttttggac aaaatcccct ttcctcatat ggattgtttg tttcttcaga 720
caattcgtta ggtcggttcc aaaggttgat tacttgacct taagacatgg tttcgtcatg 780
gcacatctgg caccgcacag cgatcagaaa tttgactttc aaaaatacat aaaacgatct 840
cttgaagaag atttcaaggt ggtggtcagt atcagccctc cgatatggtt ctttgctgtc 900
ctcttcctac ttttcaacac ccacgggtgg agggcttatc tatggctacc ctttgttccg 960
ttaattatag tgttattggt ggggacaaag ttgcaagtga taataacgaa aatggcgctg 1020
aggatacaag aaagaggaga agtggtgaaa ggagtgccgg tggtagagcc aggggatgac 1080
cttttttggt tcaatcgccc tcgtcttatt ctttacctta tcaattttgt cctcttccag 1140
aatgcctttc agcttgcctt ttttgcttgg acttggaaag aatttgggat gaaatcttgt 1200
ttccatgagc acacagagga tttggtcatc agaataacaa tgggggttct cgttcaaatc 1260
ctttgcagtt atgtcacatt gccactttac gctctagtca cacagatggg ttcgacgatg 1320
aagcccacga ttttcaacga aagagtagcg acggcgttga gaaattggca ccacaccgct 1380
cgtaaacaca taaaacaaaa tcgtggctca atgacgccga tgtcgagccg ccctgcaacc 1440
ccctcccacc acttgtcacc cgtccacctc cttcgccact atcgaagcga attagatagc 1500
gttcatacgt ctcctagaag atccaatttc gacaccgatc agtgggaccc tgattcccct 1560
tccccttccc cttctcacca ctttcatcgt cgtccccatc ccggcgacgg ctccatttcc 1620
aaccatcacc gtgatgtgga ggccggggat cttgatgtcg atgttgaatc gcctcaaccc 1680
gaccgaacga cccagtcaat aaacccaaca aatattgagc accatgaaat tgacgtgggg 1740
tctaacgaat tctcattcga tagaagagtt gatagagtat aa 1782
<210> 4
<211> 20
<212> DNA
<213> 黄瓜(cucumber)
<400> 4
tttggccgct gcgggaggag 20
<210> 5
<211> 20
<212> DNA
<213> 黄瓜(cucumber)
<400> 5
atacgcctcg aaccatagtc 20
<210> 6
<211> 20
<212> DNA
<213> 黄瓜(cucumber)
<400> 6
cgtctcttcc aaggaccttc 20
<210> 7
<211> 60
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 7
tttggtctca tgcacgtctc ttccaaggac cttcgtttta gagctagaaa tagcaagtta 60
<210> 8
<211> 44
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 8
tttggtctca cccgcagcgg ccaaatgcac cagccgggaa tcga 44
<210> 9
<211> 45
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 9
tttggtctca cgggaggagg ttttagagct agaaatagca agtta 45
<210> 10
<211> 42
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 10
tttggtctca gttcgaggcg tattgcacca gccgggaatc ga 42
<210> 11
<211> 47
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 11
tttggtctca gaaccatagt cgttttagag ctagaaatag caagtta 47
<210> 12
<211> 32
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 12
tttggtctca tgttgcaccg actcggtgcc ac 32
<210> 13
<211> 582
<212> PRT
<213> 黄瓜(cucumber)
<400> 13
Met Ala Gly Ala Ala Gly Gly Lys Ser Leu Glu Gln Thr Pro Thr Trp
1 5 10 15
Ala Val Ala Val Val Cys Phe Val Leu Leu Val Ile Ser Ile Phe Ile
20 25 30
Glu Tyr Ser Leu His Leu Ile Gly His Trp Leu Lys Lys Arg His Lys
35 40 45
Arg Ala Leu Phe Glu Ala Leu Glu Lys Ile Lys Ser Glu Leu Met Leu
50 55 60
Leu Gly Phe Ile Ser Leu Leu Leu Thr Val Gly Gln Gly Pro Ile Thr
65 70 75 80
Glu Ile Cys Ile Pro Gln His Val Ala Ala Thr Trp His Pro Cys Thr
85 90 95
Lys Glu Arg Glu Asp Glu Met Asn Lys Glu Val Glu Lys Ser Val Glu
100 105 110
His Leu Gly Leu Asn Arg Arg Arg Leu Leu His Leu Leu Gly Asn Gly
115 120 125
Glu Ser Phe Arg Arg Ser Leu Ala Ala Ala Gly Gly Glu Asp Lys Cys
130 135 140
Ala Ala Lys Gly Lys Ala Ser Phe Ile Ser Ala Asp Gly Ile His Gln
145 150 155 160
Leu His Ile Phe Ile Phe Val Leu Ala Val Phe His Val Leu Tyr Cys
165 170 175
Val Leu Thr Tyr Ala Leu Ala Arg Ala Lys Met Arg Ser Trp Lys Thr
180 185 190
Trp Glu Lys Glu Thr Lys Thr Ala Glu Tyr Gln Phe Ser His Asp Pro
195 200 205
Glu Arg Phe Arg Phe Ala Arg Asp Thr Ser Phe Gly Arg Arg His Leu
210 215 220
Ser Phe Trp Thr Lys Asn Pro Ala Leu Met Trp Ile Val Cys Phe Phe
225 230 235 240
Arg Gln Phe Val Arg Ser Val Pro Lys Val Asp Tyr Leu Thr Leu Arg
245 250 255
His Gly Phe Ile Met Ala His Leu Ala Pro Gln Ser His Thr Gln Phe
260 265 270
Asp Phe Gln Lys Tyr Ile Asn Arg Ser Leu Glu Glu Asp Phe Lys Val
275 280 285
Val Val Gly Ile Ser Pro Pro Ile Trp Phe Phe Ala Val Leu Phe Leu
290 295 300
Leu Ser Asn Thr His Gly Trp Arg Ala Tyr Leu Trp Leu Pro Phe Ile
305 310 315 320
Pro Leu Ile Ile Leu Leu Leu Ile Gly Thr Lys Leu Gln Val Ile Ile
325 330 335
Thr Lys Met Ala Leu Arg Ile Gln Glu Arg Gly Glu Val Val Lys Gly
340 345 350
Val Pro Val Val Glu Pro Gly Asp Asp Leu Phe Trp Phe Asn Arg Pro
355 360 365
Arg Leu Ile Leu Tyr Leu Ile Asn Phe Val Leu Phe Gln Asn Ala Phe
370 375 380
Gln Val Ala Phe Phe Ala Trp Thr Trp Tyr Glu Phe Gly Leu Asn Ser
385 390 395 400
Cys Phe His Glu His Ile Glu Asp Val Val Ile Arg Ile Ser Met Gly
405 410 415
Val Leu Val Gln Ile Leu Cys Ser Tyr Val Thr Leu Pro Leu Tyr Ala
420 425 430
Leu Val Thr Gln Met Gly Ser Thr Met Lys Pro Thr Ile Phe Asn Glu
435 440 445
Arg Val Ala Glu Ala Leu Arg Asn Trp Tyr His Ser Ala Arg Lys His
450 455 460
Ile Lys His Asn Arg Gly Ser Val Thr Pro Met Ser Ser Arg Pro Ala
465 470 475 480
Thr Pro Thr His Ser Met Ser Pro Val His Leu Leu Arg His Tyr Lys
485 490 495
Ser Glu Val Asp Ser Phe His Thr Ser Pro Arg Arg Ser Pro Phe Asp
500 505 510
Thr Asp Arg Trp Asp Asn Asp Ser Pro Ser Pro Ser Arg His Val Asp
515 520 525
Gly Ser Ser Ser Ser Gln Pro His Val Glu Met Gly Gly Tyr Glu Lys
530 535 540
Asp Pro Val Glu Ser Ser Ser Ser Gln Val Asp Pro Val Gln Pro Ser
545 550 555 560
Arg Asn Arg Asn Gln His Glu Ile His Ile Gly Gly Pro Lys Asp Phe
565 570 575
Ser Phe Asp Arg Val Glu
580
<210> 14
<211> 574
<212> PRT
<213> 黄瓜(cucumber)
<400> 14
Met Ala Glu Cys Gly Thr Glu Gln Arg Thr Leu Glu Asp Thr Ser Thr
1 5 10 15
Trp Ala Val Ala Val Val Cys Phe Phe Leu Val Val Ile Ser Ile Phe
20 25 30
Ile Glu His Val Ile His Leu Thr Gly Lys Trp Leu Glu Lys Arg His
35 40 45
Lys Pro Ala Leu Val Glu Ala Leu Glu Lys Val Lys Ala Glu Leu Met
50 55 60
Leu Leu Gly Phe Ile Ser Leu Leu Leu Thr Ile Gly Gln Asp Ala Val
65 70 75 80
Thr Gln Ile Cys Val Ser Lys Glu Leu Ala Ala Thr Trp Leu Pro Cys
85 90 95
Ala Ala Arg Ala Lys Thr Gly Val Lys Val Ala Lys Asn Ser Arg Leu
100 105 110
Arg Leu Leu Glu Phe Leu Asp Pro Asp Tyr Gly Ser Arg Arg Ile Leu
115 120 125
Ala Ser Lys Gly Asp Asp Ala Cys Ala Lys Arg Gly Gln Leu Ala Phe
130 135 140
Val Ser Ala Tyr Gly Ile His Gln Leu His Ile Phe Ile Phe Val Leu
145 150 155 160
Ala Val Phe His Val Leu Tyr Cys Ile Ile Thr Leu Ala Phe Gly Arg
165 170 175
Thr Lys Met Ser Lys Trp Lys Ala Trp Glu Asp Glu Thr Lys Thr Ile
180 185 190
Glu Tyr Gln Tyr Tyr Asn Asp Pro Ala Arg Phe Arg Phe Ala Arg Asp
195 200 205
Thr Thr Phe Gly Arg Arg His Leu Ser Phe Trp Ser Arg Thr Pro Ile
210 215 220
Ser Leu Trp Ile Val Cys Phe Phe Arg Gln Phe Phe Gly Ser Val Thr
225 230 235 240
Lys Val Asp Tyr Met Thr Leu Arg His Gly Phe Ile Val Ala His Leu
245 250 255
Ala Pro Gly Ser Glu Val Lys Phe Asp Phe His Lys Tyr Ile Ser Arg
260 265 270
Ser Leu Glu Asp Asp Phe Lys Val Val Val Gly Ile Ser Pro Ala Met
275 280 285
Trp Leu Phe Ala Val Leu Phe Ile Leu Thr Asn Thr Asn Gly Trp Tyr
290 295 300
Ser Tyr Leu Trp Leu Pro Phe Ile Ser Leu Ile Ile Ile Leu Leu Val
305 310 315 320
Gly Thr Lys Leu His Val Ile Ile Thr His Met Gly Leu Thr Ile Gln
325 330 335
Glu Arg Gly His Val Val Lys Gly Val Pro Val Val Gln Pro Arg Asp
340 345 350
Asp Leu Phe Trp Phe Gly Arg Pro Gln Leu Ile Leu Phe Leu Ile His
355 360 365
Phe Val Leu Phe Met Asn Ala Phe Gln Leu Ala Phe Phe Ala Trp Thr
370 375 380
Thr Tyr Ala Phe Lys Trp Met Gly Cys Phe His Gln Arg Val Glu Asp
385 390 395 400
Ile Val Ile Arg Leu Ser Met Gly Val Ile Ile Gln Val Leu Cys Ser
405 410 415
Tyr Val Thr Leu Pro Leu Tyr Ala Leu Val Thr Gln Met Gly Ser Asn
420 425 430
Met Arg Pro Thr Ile Phe Asn Asp Arg Val Ala Thr Ala Leu Lys Asn
435 440 445
Trp His His Ser Ala Lys Lys Asn Met Lys Gln His Arg Asn Pro Asp
450 455 460
Ser Thr Ser Pro Phe Ser Ser Arg Pro Ala Thr Pro Thr His Gly Met
465 470 475 480
Ser Pro Ile His Leu Leu His Lys His Gln His Gly Ser Thr Ser Pro
485 490 495
Arg Leu Ser Asp Ala Glu Pro Asp Arg Trp Glu Glu Leu Pro Pro Ser
500 505 510
Ser His His Ser Arg Ala Pro His His Asp Asn His Gln Asp Gln Gln
515 520 525
Glu Gln Ser Glu Thr Ile Ile Arg Glu Gln Glu Met Thr Val Gln Gly
530 535 540
Pro Ser Ser Ser Glu Thr Gly Ser Ile Thr Arg Pro Ala Arg Pro His
545 550 555 560
Gln Glu Ile Thr Arg Thr Pro Ser Asp Phe Ser Phe Ala Lys
565 570
<210> 15
<211> 593
<212> PRT
<213> 黄瓜(cucumber)
<400> 15
Met Ala Gly Gly Gly Ala Gly Arg Ser Leu Glu Glu Thr Pro Thr Trp
1 5 10 15
Ala Val Ala Ala Val Cys Phe Val Leu Val Leu Ile Ser Ile Ile Ile
20 25 30
Glu His Ile Leu His Leu Ile Gly Lys Trp Leu Lys Lys Lys His Lys
35 40 45
Arg Ala Leu Tyr Glu Ala Leu Glu Lys Ile Lys Ser Glu Leu Met Leu
50 55 60
Leu Gly Phe Ile Ser Leu Leu Leu Thr Val Gly Gln Ser Leu Ile Thr
65 70 75 80
Asn Val Cys Ile Pro Pro Asp Val Ala Ala Thr Trp His Pro Cys Ser
85 90 95
Pro Gln Arg Glu Glu Glu Leu Thr Lys Glu Ala Asp Leu Val Asp Ser
100 105 110
Asp Gln Asn Arg Arg Lys Leu Leu Ala Leu Ser His His Val Asn Ala
115 120 125
Thr Phe Arg Arg Ser Leu Ala Ala Ala Gly Gly Thr Asp Lys Cys Ala
130 135 140
Ala Lys Gly Lys Val Pro Phe Val Ser Glu Gly Gly Ile His Gln Leu
145 150 155 160
His Ile Phe Ile Phe Val Leu Ala Val Phe His Val Leu Tyr Cys Val
165 170 175
Leu Thr Leu Ala Leu Gly Asn Ala Lys Met Arg Ser Trp Lys Ser Trp
180 185 190
Glu Lys Glu Thr Arg Thr Val Glu Tyr Gln Phe Ser His Asp Pro Glu
195 200 205
Arg Phe Arg Phe Ala Arg Asp Thr Ser Phe Gly Arg Arg His Leu Ser
210 215 220
Phe Trp Thr Lys Ser Pro Phe Leu Ile Trp Ile Val Cys Phe Phe Arg
225 230 235 240
Gln Phe Val Arg Ser Val Pro Lys Val Asp Tyr Leu Thr Leu Arg His
245 250 255
Gly Phe Val Met Ala His Leu Ala Pro His Ser Asp Gln Lys Phe Asp
260 265 270
Phe Gln Lys Tyr Ile Lys Arg Ser Leu Glu Glu Asp Phe Lys Val Val
275 280 285
Val Ser Ile Ser Pro Pro Ile Trp Phe Phe Ala Val Leu Phe Leu Leu
290 295 300
Phe Asn Thr His Gly Trp Arg Ala Tyr Leu Trp Leu Pro Phe Val Pro
305 310 315 320
Leu Ile Ile Val Leu Leu Val Gly Thr Lys Leu Gln Val Ile Ile Thr
325 330 335
Lys Met Ala Leu Arg Ile Gln Glu Arg Gly Glu Val Val Lys Gly Val
340 345 350
Pro Val Val Glu Pro Gly Asp Asp Leu Phe Trp Phe Asn Arg Pro Arg
355 360 365
Leu Ile Leu Tyr Leu Ile Asn Phe Val Leu Phe Gln Asn Ala Phe Gln
370 375 380
Leu Ala Phe Phe Ala Trp Thr Trp Lys Glu Phe Gly Met Lys Ser Cys
385 390 395 400
Phe His Glu His Thr Glu Asp Leu Val Ile Arg Ile Thr Met Gly Val
405 410 415
Leu Val Gln Ile Leu Cys Ser Tyr Val Thr Leu Pro Leu Tyr Ala Leu
420 425 430
Val Thr Gln Met Gly Ser Thr Met Lys Pro Thr Ile Phe Asn Glu Arg
435 440 445
Val Ala Thr Ala Leu Arg Asn Trp His His Thr Ala Arg Lys His Ile
450 455 460
Lys Gln Asn Arg Gly Ser Met Thr Pro Met Ser Ser Arg Pro Ala Thr
465 470 475 480
Pro Ser His His Leu Ser Pro Val His Leu Leu Arg His Tyr Arg Ser
485 490 495
Glu Leu Asp Ser Val His Thr Ser Pro Arg Arg Ser Asn Phe Asp Thr
500 505 510
Asp Gln Trp Asp Pro Asp Ser Pro Ser Pro Ser Pro Ser His His Phe
515 520 525
His Arg Arg Pro His Pro Gly Asp Gly Ser Ile Ser Asn His His Arg
530 535 540
Asp Val Glu Ala Gly Asp Leu Asp Val Asp Val Glu Ser Pro Gln Pro
545 550 555 560
Asp Arg Thr Thr Gln Ser Ile Asn Pro Thr Asn Ile Glu His His Glu
565 570 575
Ile Asp Val Gly Ser Asn Glu Phe Ser Phe Asp Arg Arg Val Asp Arg
580 585 590
Val
Claims (10)
1.一种提高黄瓜白粉病抗性的方法,其特征在于,使CsMLO1、CsMLO8及CsMLO11基因中的两个或三个共同产生突变;
或
使CsMLO1基因、CsMLO8基因及CsMLO11基因编码的氨基酸中的两个或三个共同产生突变来提高黄瓜白粉病的抗性;
所述CsMLO1的基因序列如SEQ ID NO.1所示,所述CsMLO8的基因序列如SEQ ID NO.2所示,所述CsMLO11的基因序列如SEQ ID NO.3所示。
2.根据权利要求1所述的提高黄瓜白粉病抗性的方法,其特征在于,所述CsMLO1基因发生的突变在CsMLO1靶点片段区域中,所述CsMLO1靶点片段序列为SEQ ID NO.4所示,所述CsMLO8基因发生的突变在CsMLO8靶点片段区域中,所述CsMLO8靶点片段序列为SEQ IDNO.5所示,所述CsMLO11基因发生的突变在CsMLO11靶点片段区域中,所述CsMLO11靶点片段序列为SEQ ID NO.6所示。
3.一种调控黄瓜白粉病抗性的重组载体,其特征在于,所述调控黄瓜白粉病抗性的重组载体包括选自所述CsMLO1基因的所述CsMLO1靶点片段,序列如SEQ ID NO.4所示,选自所述CsMLO8基因的CsMLO8靶点片段,序列如SEQ ID NO.5所示,选自所述CsMLO11基因的CsMLO11靶点片段,序列如SEQ ID NO.6及骨架载体质粒,所述CsMLO1靶点片段、所述CsMLO8靶点片段及所述CsMLO11靶点片段连接在所述骨架载体质粒上。
4.根据权利要求3所述的调控黄瓜白粉病抗性的重组载体,其特征在于,所述CsMLO1靶点片段、所述CsMLO8靶点片段及所述CsMLO11靶点片段之间通过依次连接的gRNA scaffold及tRNA进行间隔串联,所述gRNA scaffold的序列为:
GTTTTAGAGCTAGAAATAGCAAGTTAAAATAAGGCTAGTCCGTTATCAACTTGAAAAAGTGGCACCGAGTCGGTGC;
所述tRNA的序列为:
AACAAAGCACCAGTGGTCTAGTGGTAGAATAGTACCCTGCCACGG TACAGACCCGGGTTCGATTCCCGGCTGGTGCA。
5.一种制备权利要求3所述的调控黄瓜白粉病抗性的重组载体的方法,其特征在于,所述制备调控黄瓜白粉病抗性的重组载体的方法包括:将扩增引物扩增出含所述CsMLO1靶点片段的基因片段、含所述CsMLO8靶点片段的基因片段及含所述CsMLO11靶点片段的基因片段,然后各基因片段与线性化的载体质粒连接后,转入大肠杆菌,经过抽提后得到所述调控黄瓜白粉病抗性的重组载体。
6.根据权利要求5所述的制备调控黄瓜白粉病抗性的重组载体的方法,其特征在于,所述制备调控黄瓜白粉病抗性的重组载体的方法还包括利用引物扩增载体连接片段。
7.根据权利要求6所述的制备调控黄瓜白粉病抗性的重组载体的方法,其特征在于,扩增含所述CsMLO11靶点片段的基因片段的CsMLO11-F引物序列如SEQ ID NO.7所示;扩增出含所述CsMLO1靶点片段的基因片段的上游引物CsMLO1-R序列如SEQ ID NO.8所示,扩增出含所述CsMLO1靶点片段的基因片段的下游引物CsMLO1-F序列如SEQ ID NO.9所示;扩增出含所述CsMLO8靶点片段的基因片段的上游引物CsMLO8-R序列如SEQ ID NO.10所示,扩增出含所述CsMLO8靶点片段的的基因片段的下游引物CsMLO8-F序列如SEQ ID NO.11所示,扩增载体连接片段的VECTOR-R引物如SEQ ID NO.12所示。
8.根据权利要求6所述的制备调控黄瓜白粉病抗性的重组载体的方法,其特征在于,所述CsMLO11-F引物与所述CsMLO1-R引物为第一引物对,所述CsMLO1-F引物与CsMLO8-R引物为第二引物对,所述CsMLO8-F引物与VECTOR-R为第三引物对,分别进行扩增。
9.一种得到高白粉病抗性黄瓜的方法,其特征在于,所述得到高白粉病抗性黄瓜的方法包括:
步骤S1:将权利要求3所述的重组载体转入农杆菌。
步骤S2:将转入所述重组载体的农杆菌浸染黄瓜子叶。
10.根据权利要求9所述的得到高白粉病抗性黄瓜的方法,其特征在于,所述得到高白粉病抗性黄瓜的方法还包括:
步骤S3:将所述步骤S2中得到的黄瓜植株与未经过处理的黄瓜植株杂交后再自交。
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