CN106591352A - 杀虫蛋白组合及其管理昆虫抗性的方法 - Google Patents
杀虫蛋白组合及其管理昆虫抗性的方法 Download PDFInfo
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Abstract
本发明涉及一种杀虫蛋白组合及其管理昆虫抗性的方法,所述管理昆虫抗性的方法包括将亚洲玉米螟至少与Cry2Ab蛋白和Cry1Ab蛋白接触。本发明通过利用两种杀虫蛋白Cry1Ab蛋白和Cry2Ab蛋白有效延缓或防止亚洲玉米螟产生抗性,从而实现对亚洲玉米螟的控制或防治,使植物更大限度的获得保护并稳定产量。
Description
技术领域
本发明涉及一种杀虫蛋白组合及其管理昆虫抗性的方法,特别是涉及组合使用Cry1Ab蛋白和Cry2Ab蛋白以管理亚洲玉米螟抗性的方法。
背景技术
昆虫害虫使全球农作物生产遭受着巨大的经济损失,并且农民每年都会面临由于虫害造成产量损失的威胁。亚洲玉米螟(Ostrinia furnacalis)俗名钻心虫,属鳞翅目螟蛾科,是我国玉米生产中的主要害虫,该类昆虫取食玉米叶片,蛀入玉米主茎或果穗内,降低光合作用,影响养分运输,还会导致各种次级病害的产生,致使玉米减产降质。近年来,随着气候条件的变化、耕作制度的改变、玉米种植密度的加大、肥水条件的提高,亚洲玉米螟的危害日益加重。
通过转化Bt(Bacillus thuringiensis)杀虫蛋白基因产生昆虫抗性植物的能力给现代农业带来了革命,并提高了杀虫蛋白及其基因的重要性和价值。已有数种Bt蛋白用于产生昆虫抗性的转基因植物中,包括Cry1Ab蛋白、Cry1Ac蛋白、Cry1F蛋白、Cry2Ab蛋白、Cry3Bb蛋白和Vip3A蛋白等。然而随着转基因作物的推广应用,昆虫在持续的选择压力下将进化出针对转基因植物中表达的Bt蛋白的抗性,这样的抗性一旦产生并且不能被有效控制的话,无疑将限制含有Bt蛋白的转基因作物品种的商业价值。只有实施合理的抗性管理策略,才能使这一现代化技术成果得以持久的利用。
生产上为了减少昆虫抗性的产生主要采用以下几种抗性管理策略:
1)田间设置庇护所。设置庇护所的目的在于保持一定比例非抗性的等位基因,以延缓耐受高剂量抗虫蛋白的昆虫的产生,但此种策略会使得农民在实际操作上较为繁琐,并且在一定程度上降低总产量。
2)提高单一抗虫蛋白的使用剂量,此种策略可持续性较差,昆虫在持续的选择压力下将对单一蛋白产生更高的抗性。
3)不同抗虫蛋白交替使用或共使用。鉴于Bt蛋白特异地结合敏感昆虫的受体,因此叠加使用抗虫蛋白需评估昆虫对不同抗虫蛋白是否存在交互抗性,即是否共享或竞争结合位点,具有较高的不确定性,故至今尚无关于Cry2Ab蛋白针对亚洲玉米螟与Cry1Ab蛋白是否具有交互抗性的相关报道。
发明内容
本发明的目的是提供一种杀虫蛋白组合及其管理昆虫抗性的方法,不仅首次提出了通过Cry2Ab蛋白和Cry1Ab蛋白组合控制抗性亚洲玉米螟的方法,而且有效延缓了亚洲玉米螟对单一蛋白产生抗性。
为实现上述目的,本发明提供了一种管理昆虫抗性的方法,所述方法包括将亚洲玉米螟至少与Cry2Ab蛋白和Cry1Ab蛋白接触。
进一步地,所述Cry2Ab蛋白和Cry1Ab蛋白存在于至少产生所述Cry2Ab蛋白和Cry1Ab蛋白的细菌或转基因植物中,所述亚洲玉米螟通过摄食所述细菌或所述转基因植物的组织至少与所述Cry2Ab蛋白和Cry1Ab蛋白接触,接触后所述亚洲玉米螟生长受到抑制和/或导致死亡,以实现管理亚洲玉米螟的抗性。
更进一步地,将对Cry1Ab蛋白产生抗性的亚洲玉米螟至少与所述Cry2Ab蛋白接触,所述Cry2Ab蛋白存在于至少产生所述Cry2Ab蛋白的细菌或转基因植物中,所述对Cry1Ab蛋白产生抗性的亚洲玉米螟通过摄食所述细菌或所述转基因植物的组织至少与所述Cry2Ab蛋白接触,接触后所述对Cry1Ab蛋白产生抗性的亚洲玉米螟生长受到抑制和/或导致死亡,以实现管理所述对Cry1Ab蛋白产生抗性的亚洲玉米螟的抗性。
再进一步地,将对Cry2Ab蛋白产生抗性的亚洲玉米螟至少与所述Cry1Ab蛋白接触,所述Cry1Ab蛋白存在于至少产生所述Cry1Ab蛋白的细菌或转基因植物中,所述对Cry2Ab蛋白产生抗性的亚洲玉米螟通过摄食所述细菌或所述转基因植物的组织至少与所述Cry1Ab蛋白接触,接触后所述对Cry2Ab蛋白产生抗性的亚洲玉米螟生长受到抑制和/或导致死亡,以实现管理所述对Cry2Ab蛋白产生抗性的亚洲玉米螟的抗性。
在上述技术方案中,所述转基因植物处于任意生育期。
进一步地,所述转基因植物的组织为根、叶片、茎秆、果实、雄穗、雌穗、花药或花丝。
更进一步地,所述对亚洲玉米螟危害植物的控制不因种植地点和/或种植时间的改变而改变。
可选择地,所述植物为玉米、小麦、高粱、谷子、水稻或大豆。
在上述技术方案中,所述Cry2Ab蛋白的氨基酸序列具有SEQ ID NO:1所示的氨基酸序列。
进一步地,所述Cry2Ab蛋白的氨基酸序列具有SEQ ID NO:1所示的氨基酸序列。
优选地,所述Cry1Ab蛋白的氨基酸序列具有SEQ ID NO:2所示的氨基酸序列。
进一步地,所述Cry1Ab蛋白的核苷酸序列具有SEQ ID NO:4所示的核苷酸序列。
为实现上述目的,本发明还提供了一种控制亚洲玉米螟的方法,所述方法包括将亚洲玉米螟至少与Cry2Ab蛋白和Cry1Ab蛋白接触,从而实现对亚洲玉米螟的控制。
进一步地,所述Cry2Ab蛋白和Cry1Ab蛋白存在于至少产生所述Cry2Ab蛋白和Cry1Ab蛋白的细菌或转基因植物中,所述亚洲玉米螟通过摄食所述细菌或所述转基因植物的组织至少与所述Cry2Ab蛋白和Cry1Ab蛋白接触,接触后所述亚洲玉米螟生长受到抑制和/或导致死亡,以实现对亚洲玉米螟危害植物的控制。
更进一步地,将对Cry1Ab蛋白产生抗性的亚洲玉米螟至少与所述Cry2Ab蛋白接触,所述Cry2Ab蛋白存在于至少产生所述Cry2Ab蛋白的细菌或转基因植物中,所述对Cry1Ab蛋白产生抗性的亚洲玉米螟通过摄食所述细菌或所述转基因植物的组织至少与所述Cry2Ab蛋白接触,接触后所述对Cry1Ab蛋白产生抗性的亚洲玉米螟生长受到抑制和/或导致死亡,以实现对所述对Cry1Ab蛋白产生抗性的亚洲玉米螟危害植物的控制。
再进一步地,将对Cry2Ab蛋白产生抗性的亚洲玉米螟至少与所述Cry1Ab蛋白接触,所述Cry1Ab蛋白存在于至少产生所述Cry1Ab蛋白的细菌或转基因植物中,所述对Cry2Ab蛋白产生抗性的亚洲玉米螟通过摄食所述细菌或所述转基因植物的组织至少与所述Cry1Ab蛋白接触,接触后所述对Cry2Ab蛋白产生抗性的亚洲玉米螟生长受到抑制和/或导致死亡,以实现对所述对Cry2Ab蛋白产生抗性的亚洲玉米螟危害植物的控制。
在上述技术方案中,所述转基因植物处于任意生育期。
进一步地,所述转基因植物的组织为根、叶片、茎秆、果实、雄穗、雌穗、花药或花丝。
优选地,所述对亚洲玉米螟危害植物的控制不因种植地点和/或种植时间的改变而改变。
可选择地,所述植物为玉米、小麦、高粱、谷子、水稻或大豆。
在上述技术方案中,所述Cry2Ab蛋白的氨基酸序列具有SEQ ID NO:1所示的氨基酸序列。
进一步地,所述Cry2Ab蛋白的核苷酸序列具有SEQ ID NO:3所示的核苷酸序列。
优选地,所述Cry1Ab蛋白的氨基酸序列具有SEQ ID NO:2所示的氨基酸序列。
进一步地,所述Cry1Ab蛋白的核苷酸序列具有SEQ ID NO:4所示的核苷酸序列。
为实现上述目的,本发明提供了一种Cry2Ab蛋白和Cry1Ab蛋白组合使用以防止或延缓亚洲玉米螟群体对Cry1Ab蛋白或Cry2Ab蛋白产生抗性的用途。
为实现上述目的,本发明还提供了一种Cry2Ab蛋白和Cry1Ab蛋白组合使用以控制对Cry1Ab蛋白或Cry2Ab蛋白产生抗性的亚洲玉米螟群体的用途。
本发明中所述的“接触”,是指昆虫和/或害虫触碰、停留和/或摄食植物、植物器官、植物组织或植物细胞,所述植物、植物器官、植物组织或植物细胞既可以是其体内表达杀虫蛋白,还可以是所述植物、植物器官、植物组织或植物细胞的表面具有杀虫蛋白和/或具有产生杀虫蛋白的微生物。
本发明所述的“控制”和/或“防治”是指亚洲玉米螟害虫至少与Cry2Ab蛋白和Cry1Ab蛋白接触,接触后亚洲玉米螟害虫生长受到抑制和/或导致死亡。进一步地,亚洲玉米螟害虫通过摄食植物组织至少与Cry2Ab蛋白和Cry1Ab蛋白接触,接触后全部或部分亚洲玉米螟害虫生长受到抑制和/或导致死亡。抑制是指亚致死,即尚未致死但能引起生长发育、行为、生理、生化和组织等方面的某种效应,如生长发育缓慢和/或停止。同时,植物在形态上应是正常的,且可在常规方法下培养以用于产物的消耗和/或生成。此外,含有编码Cry2Ab蛋白和Cry1Ab蛋白的多核苷酸序列的控制亚洲玉米螟害虫的植物和/或植物种子,在人工接种亚洲玉米螟害虫和/或亚洲玉米螟害虫自然发生危害的条件下,与非转基因的野生型植株相比具有减弱的植物损伤,具体表现包括但不限于改善的茎秆抗性、和/或提高的籽粒重量、和/或增产等。Cry2Ab蛋白和Cry1Ab蛋白对亚洲玉米螟的“控制”和/或“防治”作用是可以独立存在的,不因其它可“控制”和/或“防治”亚洲玉米螟害虫的物质的存在而减弱和/或消失。具体地,转基因植物(含有编码Cry2Ab蛋白和Cry1Ab蛋白的多核苷酸序列)的任何组织同时和/或不同步地,存在和/或产生,Cry2Ab蛋白和Cry1Ab蛋白和/或可控制亚洲玉米螟害虫的另一种物质,则所述另一种物质的存在既不影响Cry2Ab蛋白和Cry1Ab蛋白对亚洲玉米螟的“控制”和/或“防治”作用,也不能导致所述“控制”和/或“防治”作用完全和/或部分由所述另一种物质实现,而与Cry2Ab蛋白和Cry1Ab蛋白无关。通常情况下,在大田,亚洲玉米螟害虫摄食植物组织的过程短暂且很难用肉眼观察到,因此,在人工接种亚洲玉米螟害虫和/或亚洲玉米螟害虫自然发生危害的条件下,如转基因植物(含有编码Cry2Ab蛋白和Cry1Ab蛋白的多核苷酸序列)的任何组织存在死亡的亚洲玉米螟害虫、和/或在其上停留生长受到抑制的亚洲玉米螟害虫、和/或与非转基因的野生型植株相比具有减弱的植物损伤,即为实现了本发明的方法和/或用途,即通过亚洲玉米螟害虫至少与Cry2Ab蛋白和Cry1Ab蛋白接触以实现控制亚洲玉米螟害虫的方法和/或用途。
术语“植物”包括整株植物、植物细胞、植物器官、植物原生质体、植物可以从中再生的植物细胞组织培养物、植物愈伤组织、植物丛(plant clumps)和植物或植物部分中完整的植物细胞,所述植物部分例如胚、花粉、胚珠、种子、叶、花、枝、果实、茎秆、根、根尖、花药等。
本发明中所述的植物、植物组织或植物细胞的基因组,是指植物、植物组织或植物细胞内的任何遗传物质,且包括细胞核和质体和线粒体基因组。
本发明中所述的“重组”是指通常不能在自然界中发现并且因此通过人工干预产生的DNA和/或蛋白和/或生物体的形式。这种人工干预可产生重组DNA分子和/或重组植物。所述“重组DNA分子”是通过人工组合两种在其它情况下是分离的序列区段而获得的,例如通过化学合成或通过遗传工程技术操作分离的核酸区段。进行核酸操作的技术是众所周知的。
本发明术语“原毒素”或“毒素”或“杀虫毒素”指在中肠中发生任何断裂之前编码杀虫蛋白的全长基因的最初翻译产物。本发明术语“毒素”或“最小毒性片段”应理解为杀虫蛋白例如Cry2Ab或Cry1Ab蛋白的部分,其可以通过胰蛋白酶消化或通过在(目标昆虫,例如亚洲玉米螟)中肠液中水解而获得、且仍具有杀昆虫活性。通常在SDS-PAGE凝胶上,Cry1毒素具有约60-65kD的分子量,Cry2A毒素具有约50-58kD的分子量。
在本发明中,Cry2Ab蛋白和Cry1Ab蛋白在一种转基因植物中表达。这种超过一种的杀虫毒素在同一株转基因植物中共同表达可以通过遗传工程使植物包含并表达所需的基因来实现。另外,一种植物(第1亲本)可以通过遗传工程操作表达Cry2Ab蛋白质,第二种植物(第2亲本)可以通过遗传工程操作表达Cry1Ab蛋白质。通过第1亲本和第2亲本杂交获得表达引入第1亲本和第2亲本的所有基因的后代植物。
在本发明中,产生所述Cry1Ab蛋白的转基因植物包括但不限于Mon810转基因玉米事件和/或包含Mon810转基因玉米事件的植物材料(如在US6713259B2所描述的)、Bt11转基因玉米事件和/或包含Bt11转基因玉米事件的植物材料(如在USDA APHIS非管制状态申请95-195-01p所描述的,其所包含的Cry1Ab蛋白的氨基酸序列如本发明SEQ ID NO:3所示)、Bt176转基因玉米事件和/或包含Bt176转基因玉米事件的植物材料(如在USDA APHIS非管制状态申请94-319-01p所描述的,其所包含的Cry1Ab蛋白的氨基酸序列如US5625136B2所描述的)、TT51转基因水稻事件和/或包含TT51转基因水稻事件的植物材料(如在CN100582223C和CN101302520B所描述的)、223F-S21转基因水稻品系和/或包含223F-S21转基因水稻品系的植物材料(如在CN103773759A所描述的)、Mon15985转基因棉花事件和/或包含Mon15985转基因棉花事件的植物材料(如在CN101413028B所描述的)、MON531转基因棉花事件和/或包含MON531转基因棉花事件的植物材料(如在USDA APHIS非管制状态申请00-342-01p所描述的)、COT67B转基因棉花事件和/或包含COT67B转基因棉花事件的植物材料(如在USDA APHIS非管制状态申请07-108-01p所描述的)或者3006-210-23转基因棉花事件和/或包含3006-210-23转基因棉花事件的植物材料(如在USDA APHIS非管制状态申请03-036-02p所描述的)。
在本发明中,产生所述Cry2Ab蛋白的转基因植物包括但不限于Mon89034转基因玉米事件和/或包含Mon89034转基因玉米事件的植物材料(如在CN101495635A所描述的)、MON87751转基因大豆事件和/或包含MON87751转基因大豆事件的植物材料(如在USDAAPHIS非管制状态申请13-337-01p所描述的)、或者Mon15985转基因棉花事件和/或包含Mon15985转基因棉花事件的植物材料(如在CN101413028B所描述的)。
本领域技术人员所熟知的,DNA典型的以双链形式存在。在这种排列中,一条链与另一条链互补,反之亦然。由于DNA在植物中复制产生了DNA的其它互补链。这样,本发明包括对序列表中示例的多核苷酸及其互补链的使用。本领域常使用的“编码链”指与反义链结合的链。为了在体内表达蛋白质,典型将DNA的一条链转录为一条mRNA的互补链,它作为模板翻译出蛋白质。mRNA实际上是从DNA的“反义”链转录的。“有义”或“编码”链有一系列密码子(密码子是三个核苷酸,一次读三个可以产生特定氨基酸),其可作为开放阅读框(ORF)阅读来形成目的蛋白质或肽。本发明还包括与示例的DNA有相当功能的RNA。
任何常规的核酸杂交或扩增方法都可以用于鉴定本发明杀虫基因的存在。核酸分子或其片段在一定情况下能够与其他核酸分子进行特异性杂交。本发明中,如果两个核酸分子能形成反平行的双链核酸结构,就可以说这两个核酸分子彼此间能够进行特异性杂交。如果两个核酸分子显示出完全的互补性,则称其中一个核酸分子是另一个核酸分子的“互补物”。本发明中,当一个核酸分子的每一个核苷酸都与另一个核酸分子的对应核苷酸互补时,则称这两个核酸分子显示出“完全互补性”。如果两个核酸分子能够以足够的稳定性相互杂交从而使它们在至少常规的“低度严格”条件下退火且彼此结合,则称这两个核酸分子为“最低程度互补”。类似地,如果两个核酸分子能够以足够的稳定性相互杂交从而使它们在常规的“高度严格”条件下退火且彼此结合,则称这两个核酸分子具有“互补性”。从完全互补性中偏离是可以允许的,只要这种偏离不完全阻止两个分子形成双链结构。为了使一个核酸分子能够作为引物或探针,仅需保证其在序列上具有充分的互补性,以使得在所采用的特定溶剂和盐浓度下能形成稳定的双链结构。
本发明中,基本同源的序列是一段核酸分子,该核酸分子在高度严格条件下能够和相匹配的另一段核酸分子的互补链发生特异性杂交。促进DNA杂交的适合的严格条件,例如,大约在45℃条件下用6.0×氯化钠/柠檬酸钠(SSC)处理,然后在50℃条件下用2.0×SSC洗涤,这些条件对本领域技术人员是公知的。例如,在洗涤步骤中的盐浓度可以选自低度严格条件的约2.0×SSC、50℃到高度严格条件的约0.2×SSC、50℃。此外,洗涤步骤中的温度条件可以从低度严格条件的室温约22℃,升高到高度严格条件的约65℃。温度条件和盐浓度可以都发生改变,也可以其中一个保持不变而另一个变量发生改变。优选地,本发明所述严格条件可为在6×SSC、0.5%SDS溶液中,在65℃下与SEQ ID NO:3和SEQ ID NO:4发生特异性杂交,然后用2×SSC、0.1%SDS和1×SSC、0.1%SDS各洗膜1次。
因此,具有抗虫活性并在严格条件下与本发明SEQ ID NO:3或SEQ ID NO:4杂交的序列包括在本发明中。这些序列与本发明序列至少大约40%-50%同源,大约60%、65%或70%同源,甚至至少大约75%、80%、85%、90%、91%、92%、93%、94%、95%、96%、97%、98%、99%或更大的序列同源性。
本发明中所述的基因和蛋白质不但包括特定的示例序列,还包括保存了所述特定示例的蛋白质的杀虫活性特征的部分和/片段(包括与全长蛋白质相比在内和/或末端缺失)、变体、突变体、取代物(有替代氨基酸的蛋白质)、嵌合体和融合蛋白。所述“变体”或“变异”是指编码同一蛋白或编码有杀虫活性的等价蛋白的核苷酸序列。所述“等价蛋白”是指与权利要求的蛋白具有相同或基本相同的抗鳞翅目昆虫害虫的生物活性的蛋白。
本发明中所述的DNA分子或蛋白序列的“片段”或“截短”是指涉及的原始DNA或蛋白序列(核苷酸或氨基酸)的一部分或其人工改造形式(例如适合植物表达的序列),前述序列的长度可存在变化,但长度足以确保(编码)蛋白质为昆虫毒素。
使用标准技术可以修饰基因和容易的构建基因变异体。例如,本领域熟知制造点突变的技术。又例如美国专利号5605793描述了在随机断裂后使用DNA重装配产生其它分子多样性的方法。可以使用商业化核酸内切酶制造全长基因的片段,并且可以按照标准程序使用核酸外切酶。例如,可以使用酶诸如Bal31或定点诱变从这些基因的末端系统地切除核苷酸。还可以使用多种限制性内切酶获取编码活性片段的基因。可以使用蛋白酶直接获得这些毒素的活性片段。
本发明可以从B.t.分离物和/或DNA文库衍生出等价蛋白和/或编码这些等价蛋白的基因。有多种方法获取本发明的杀虫蛋白。例如,可以使用本发明公开和要求保护的杀虫蛋白的抗体从蛋白质混合物鉴定和分离其它蛋白。特别地,抗体可能是由蛋白最恒定和与其它Bt蛋白最不同的蛋白部分引起的。然后可以通过免疫沉淀、酶联免疫吸附测定(ELISA)或western印迹方法使用这些抗体专一地鉴定有特征活性的等价蛋白。可使用本领域标准程序容易的制备本发明中公开的蛋白或等价蛋白或这类蛋白的片段的抗体。然后可以从微生物中获得编码这些蛋白的基因。
由于遗传密码子的丰余性,多种不同的DNA序列可以编码相同的氨基酸序列。产生这些编码相同或基本相同的蛋白的可替代DNA序列正在本领域技术人员的技术水平内。这些不同的DNA序列包括在本发明的范围内。所述“基本上相同的”序列是指有氨基酸取代、缺失、添加或插入但实质上不影响杀虫活性的序列,亦包括保留杀虫活性的片段。
本发明中氨基酸序列的取代、缺失或添加是本领域的常规技术,优选这种氨基酸变化为:小的特性改变,即不显著影响蛋白的折叠和/或活性的保守氨基酸取代;小的缺失,通常约1-30个氨基酸的缺失;小的氨基或羧基端延伸,例如氨基端延伸一个甲硫氨酸残基;小的连接肽,例如约20-25个残基长。
保守取代的实例是在下列氨基酸组内发生的取代:碱性氨基酸(如精氨酸、赖氨酸和组氨酸)、酸性氨基酸(如谷氨酸和天冬氨酸)、极性氨基酸(如谷氨酰胺、天冬酰胺)、疏水性氨基酸(如亮氨酸、异亮氨酸和缬氨酸)、芳香氨基酸(如苯丙氨酸、色氨酸和酪氨酸),以及小分子氨基酸(如甘氨酸、丙氨酸、丝氨酸、苏氨酸和甲硫氨酸)。通常不改变特定活性的那些氨基酸取代在本领域内是众所周知的,并且已由,例如,N.Neurath和R.L.Hill在1979年纽约学术出版社(Academic Press)出版的《Protein》中进行了描述。最常见的互换有Ala/Ser,Val/Ile,Asp/Glu,Thu/Ser,Ala/Thr,Ser/Asn,Ala/Val,Ser/Gly,Tyr/Phe,Ala/Pro,Lys/Arg,Asp/Asn,Leu/Ile,Leu/Val,Ala/Glu和Asp/Gly,以及它们相反的互换。
对于本领域的技术人员而言显而易见地,这种取代可以在对分子功能起重要作用的区域之外发生,而且仍产生活性多肽。对于由本发明的多肽,其活性必需的并因此选择不被取代的氨基酸残基,可以根据本领域已知的方法,如定点诱变或丙氨酸扫描诱变进行鉴定(如参见,Cunningham和Wells,1989,Science244:1081-1085)。后一技术是在分子中每一个带正电荷的残基处引入突变,检测所得突变分子的抗虫活性,从而确定对该分子活性而言重要的氨基酸残基。底物-酶相互作用位点也可以通过其三维结构的分析来测定,这种三维结构可由核磁共振分析、结晶学或光亲和标记等技术测定(参见,如de Vos等,1992,Science 255:306-312;Smith等,1992,J.Mol.Biol 224:899-904;Wlodaver等,1992,FEBSLetters 309:59-64)。
因此,与SEQ ID NO:1和/或SEQ ID NO:2所示的氨基酸序列具有一定同源性的氨基酸序列也包括在本发明中。这些序列与本发明序列类似性/相同性典型的大于78%,优选的大于85%,更优选的大于90%,甚至更优选的大于95%,并且可以大于99%。也可以根据更特定的相同性和/或类似性范围定义本发明的优选的多核苷酸和蛋白质。例如与本发明示例的序列有78%、79%、80%、81%、82%、83%、84%、85%、86%、87%、88%、89%、90%、91%、92%、93%、94%、95%、96%、97%、98%或99%的相同性和/或类似性。
本发明术语“交互抗性”是指昆虫的一个品系由于相同抗性机理或相似作用机理或类似化学结构,对于选择药剂以外的其它从未使用过的一种药剂或一类药剂也产生抗药性的现象。特别的,本发明中“交互抗性”指对选择的杀虫蛋白以外的其它从未使用过的一种杀虫蛋白或一类杀虫蛋白也产生抗性的现象。被选择用于昆虫抗性管理的蛋白质需要独立地发挥其杀虫效果,从而对一种蛋白质产生的抗性不会赋予对第二种蛋白质的抗性(即对于第二种蛋白质无交叉抗性或有较低的交叉抗性)。例如对“蛋白A”有抗性的昆虫群体对“蛋白B”敏感,则人们可以得出结论,蛋白A和蛋白B无交叉抗性并且它们的组合可有效延迟对单一蛋白A的抗性。
本发明术语“抗性比例”是指同一种杀虫蛋白在昆虫的Bt抗性群体相对Bt敏感群体的LC50值提高的倍数,即抗性比例=Bt抗性株LC50值/Bt敏感株LC50值。
本发明中所使用的,昆虫物种群体对杀昆虫蛋白和/或表达杀昆虫蛋白植物(该植物以前控制或杀死所述昆虫群体)“产生抗性”、“已经产生抗性”或“已成为抗性的”是指,与首次引入该杀虫蛋白和/或该植物时对相同昆虫物种造成的控制效果和/或由相同昆虫物种造成的该植物的产量损失水平相比,在该植物中检测到由该昆虫群体控制效果减弱和/或造成的重复的且显著的产量损失。
亚洲玉米螟(Ostriniafurnacalis)与棉铃虫(HelicoverpaarmigeraHubner)虽然同属鳞翅目,但是亚洲玉米螟与棉铃虫在生物学上是清晰的、截然不同的两个物种,至少存在以下主要区别:1)亚洲玉米螟属螟蛾科,棉铃虫属夜蛾科。2)食性不同。亚洲玉米螟最常危害玉米,可危害玉米植株地上的各个部位,包括叶片、雄穗、茎秆、穗柄、穗轴,对各地的春、夏、秋播玉米都有不同程度危害,尤以夏播玉米最重;棉铃虫是危害棉花蕾铃期的重要钻蛀性害虫,主要蛀食蕾、花、铃,也取食嫩叶。3)分布区域不同。亚洲玉米螟分布在中国东部及西南主要玉米、高粱产区;而棉铃虫广泛分布在中国及世界各地,中国棉区和蔬菜种植区均有发生,黄河流域棉区、长江流域棉区受害较重;近年来,新疆棉区也时有发生。4)形态特征不同。亚洲玉米螟与棉铃虫的卵、幼虫、蛹、成虫的形态特征均有较大差异。5)生长习性和发生规律不同。玉米螟的发生代数随纬度而有显著的差异:在中国,北纬45°以北1代,45°-40°2代,40°-30°3代,30°-25°4代,25°-20°5-6代。海拔越高,发生代数越少;在四川省一年发生2-4代,温度高、海拔低,发生代数较多,通常以老熟幼虫在玉米茎秆、穗轴内或高粱、向日葵的秸秆中越冬,次年4-5月化蛹,蛹经过10天左右羽化。成虫夜间活动,飞翔力强,有趋光性,寿命5~10天,喜欢在离地50厘米以上、生长较茂盛的玉米叶背面中脉两侧产卵,一个雌蛾可产卵350-700粒,卵期3-5天;玉米螟适合在高温、高湿条件下发育,冬季气温较高,天敌寄生量少,有利于玉米螟的繁殖,危害较重;卵期干旱,玉米叶片卷曲,卵块易从叶背面脱落而死亡,危害较轻。而棉铃虫发生的代数因年份因地区而异,在山东省莱州市每年发生4代,九月下旬成长幼虫陆续下树入土,在苗木附近或杂草下5-10cm深的土中化蛹越冬;立春气温回升15℃以上时开始羽化,4月下旬至5月上旬为羽化盛期,成虫出现第一代在6月中下旬,第二代在7月中下旬,第三代在8月中下旬至9月上旬至10月上旬尚有棉铃虫出现,成虫有趋光性,羽化后即在夜间闪配产卵,卵散产,较分散,一头雌蛾一生可产卵500-1000粒,最高可达2700粒,卵多产在叶背面,也有产在正面、顶芯、叶柄、嫩茎上或农作的、杂草等其它植物上;幼虫孵化后有取食卵壳习性,初孵幼虫有群集限食习性,二三头、三五头在叶片正面或背面,头向叶缘排列、自叶缘向内取食,结果叶片被吃光,只剩主脉和叶柄,或成网状枯萎,造成干叶;1-2龄幼虫沿柄下行至银杏苗顶芽处自一侧蛀食或沿顶芽处下蛀入嫩枝,造成顶梢或顶部簇生叶死亡,危害十分严重;3龄前的幼虫食量较少,较集中,随着幼虫生长而逐渐分散,进入4龄食量大增,可食光叶片,只剩叶柄;幼虫7-8月份为害最盛;棉铃虫有转移危害的习性,一只幼虫可危害多株苗木;各龄幼虫均有食掉蜕下旧皮留头壳的习性,给鉴别虫龄造成一定困难,虫龄不整齐;棉铃虫发生的最适宜温度为25-28℃,相对湿度70-90%;第二代、第三代为害最为严重,严重地片虫口密度达98头/百叶,虫株率60-70%,个别地片达100%,受害叶片达1/3以上,影响叶产量20%,质量下降至少1个等级,苗木生长量影响很大。
综合上述,亚洲玉米螟与棉铃虫虽然同属鳞翅目,但是仅在形态特征和为害习性上就存在诸多方面的不同,且二者亲缘关系较远,无法交配产生后代。因此,二者中肠上皮细胞膜上表面上与Bt毒素结合的受体也是不同的。
昆虫对Bt蛋白的抗性机制不是单一的,Heckel(1994)等分析了昆虫对Bt蛋白产生抗性的潜在机理,认为抗性的产生主要和以下因素有关:1)Bt杀虫蛋白的溶解性:原毒素不能溶解或溶解性降低;2)Bt原毒素的蛋白水解性:水解不充分或过度水解;3)Bt蛋白与细胞膜上受体的结合:Bt蛋白与受体的结合由于竞争性抑制而受阻、受体的一级结构或二级结构的修饰作用发生改变,导致Bt蛋白与受体的结合位点减少;4)细胞膜上空洞的形成:空洞形成受阻或阻塞;5)中肠上皮的修复作用;6)行为机理等。其中毒素与细胞膜上受体结合能力的改变是主要抗性机理。
当两种或以上不同的Bt蛋白在昆虫中共享结合位点时,它们不能够提供用于昆虫抗性管理目的的良好组合。实际情况下,针对不同的昆虫采取何种抗性管理策略具有较高的不确定性。氨基酸序列差异较大的两个Bt蛋白也可能在某一昆虫物种中以高亲和力结合共同的结合位点,例如Cry1Ab和Cry1F蛋白在菜蛾(Plutella xylostella)中。而且,已发现在一个昆虫物种中不具有共享的结合位点的两种蛋白可以在另一个昆虫物种中共享结合位点,例如Fiuza等人(1996)发现Cry1Ac和Cry1Ba蛋白在二化螟(Chilo suppressalis)中共享结合位点,而Ballester等人(1999)发现以上两种蛋白在菜蛾中结合不同的结合位点。
本发明中,所述Cry2Ab蛋白可以具有序列表中SEQ ID NO:1所示的氨基酸序列;所述Cry1Ab蛋白可以具有序列表中SEQ ID NO:2所示的氨基酸序列。除了包含Cry2Ab蛋白和Cry1Ab蛋白的编码区外,也可包含其他元件,例如编码转运肽的编码区或编码选择性标记的蛋白质,本发明提供的Cry2Ab蛋白和Cry1Ab蛋白的核苷酸序列可以通过常规手段形成构建体。
此外,包含编码本发明Cry2Ab蛋白和Cry1Ab蛋白的构建体在植物中还可以与至少一种编码除草剂抗性基因的蛋白质一起表达,所述除草剂抗性基因包括但不限于,草胺膦抗性基因(如bar基因、pat基因)、苯敌草抗性基因(如pmph基因)、草甘膦抗性基因(如EPSPS基因)、溴苯腈(bromoxynil)抗性基因、磺酰脲抗性基因、对除草剂茅草枯的抗性基因、对氨腈的抗性基因或谷氨酰胺合成酶抑制剂(如PPT)的抗性基因,从而获得既具有高杀虫活性、又具有除草剂抗性的转基因植物。
本发明中所述调控序列包括但不限于启动子、转运肽、终止子、增强子、前导序列、内含子以及其它可操作地连接到所述Cry2Ab蛋白或所述Cry1Ab蛋白的调节序列。
所述启动子为植物中可表达的启动子,所述的“植物中可表达的启动子”是指确保与其连接的编码序列在植物细胞内进行表达的启动子。植物中可表达的启动子可为组成型启动子。指导植物内组成型表达的启动子的示例包括但不限于,来源于花椰菜花叶病毒的35S启动子、玉米Ubi启动子、水稻GOS2基因的启动子等。备选地,植物中可表达的启动子可为组织特异的启动子,即该启动子在植物的一些组织内如在绿色组织中指导编码序列的表达水平高于植物的其他组织(可通过常规RNA试验进行测定),如PEP羧化酶启动子。备选地,植物中可表达的启动子可为创伤诱导启动子。创伤诱导启动子或指导创伤诱导的表达模式的启动子是指当植物经受机械或由昆虫啃食引起的创伤时,启动子调控下的编码序列的表达较正常生长条件下有显著提高。创伤诱导启动子的示例包括但不限于,马铃薯和西红柿的蛋白酶抑制基因(pinⅠ和pinⅡ)和玉米蛋白酶抑制基因(MPI)的启动子。
所述转运肽(又称分泌信号序列或导向序列)是指导转基因产物到特定的细胞器或细胞区室,对受体蛋白质来说,所述转运肽可以是异源的,例如,利用编码叶绿体转运肽序列靶向叶绿体,或者利用‘KDEL’保留序列靶向内质网,或者利用大麦植物凝集素基因的CTPP靶向液泡。
所述前导序列包含但不限于,小RNA病毒前导序列,如EMCV前导序列(脑心肌炎病毒5’非编码区);马铃薯Y病毒组前导序列,如MDMV(玉米矮缩花叶病毒)前导序列;人类免疫球蛋白质重链结合蛋白质(BiP);苜蓿花叶病毒的外壳蛋白质mRNA的不翻译前导序列(AMVRNA4);烟草花叶病毒(TMV)前导序列。
所述增强子包含但不限于,花椰菜花叶病毒(CaMV)增强子、玄参花叶病毒(FMV)增强子、康乃馨风化环病毒(CERV)增强子、木薯脉花叶病毒(CsVMV)增强子、紫茉莉花叶病毒(MMV)增强子、夜香树黄化曲叶病毒(CmYLCV)增强子、木尔坦棉花曲叶病毒(CLCuMV)、鸭跖草黄斑驳病毒(CoYMV)和花生褪绿线条花叶病毒(PCLSV)增强子。
对于单子叶植物应用而言,所述内含子包含但不限于,玉米hsp70内含子、玉米泛素内含子、Adh内含子1、蔗糖合酶内含子或水稻Act1内含子。对于双子叶植物应用而言,所述内含子包含但不限于,CAT-1内含子、pKANNIBAL内含子、PIV2内含子和“超级泛素”内含子。
所述终止子可以为在植物中起作用的适合多聚腺苷酸化信号序列,包括但不限于,来源于农杆菌(Agrobacterium tumefaciens)胭脂碱合成酶(NOS)基因的多聚腺苷酸化信号序列、来源于蛋白酶抑制剂Ⅱ(pinⅡ)基因的多聚腺苷酸化信号序列、来源于豌豆ssRUBISCO E9基因的多聚腺苷酸化信号序列和来源于α-微管蛋白(α-tubulin)基因的多聚腺苷酸化信号序列。
本发明中所述“有效连接”表示核酸序列的联结,所述联结使得一条序列可提供对相连序列来说需要的功能。在本发明中所述“有效连接”可以为将启动子与感兴趣的序列相连,使得该感兴趣的序列的转录受到该启动子控制和调控。当感兴趣的序列编码蛋白并且想要获得该蛋白的表达时“有效连接”表示:启动子与所述序列相连,相连的方式使得得到的转录物高效翻译。如果启动子与编码序列的连接是转录物融合并且想要实现编码的蛋白的表达时,制造这样的连接,使得得到的转录物中第一翻译起始密码子是编码序列的起始密码子。备选地,如果启动子与编码序列的连接是翻译融合并且想要实现编码的蛋白的表达时,制造这样的连接,使得5’非翻译序列中含有的第一翻译起始密码子与启动子相连结,并且连接方式使得得到的翻译产物与编码想要的蛋白的翻译开放读码框的关系是符合读码框的。可以“有效连接”的核酸序列包括但不限于:提供基因表达功能的序列(即基因表达元件,例如启动子、5’非翻译区域、内含子、蛋白编码区域、3’非翻译区域、聚腺苷化位点和/或转录终止子)、提供DNA转移和/或整合功能的序列(即T-DNA边界序列、位点特异性重组酶识别位点、整合酶识别位点)、提供选择性功能的序列(即抗生素抗性标记物、生物合成基因)、提供可计分标记物功能的序列、体外或体内协助序列操作的序列(即多接头序列、位点特异性重组序列)和提供复制功能的序列(即细菌的复制起点、自主复制序列、着丝粒序列)。
本发明中所述的“杀虫”或“抗虫”是指对农作物害虫是有毒的,从而实现“控制”和/或“防治”农作物害虫。优选地,所述“杀虫”或“抗虫”是指杀死农作物害虫。更具体地,目标昆虫是亚洲玉米螟害虫。
本发明中的植物,特别是玉米,在其基因组中含有外源DNA,所述外源DNA包含编码Cry1Ab蛋白和Cry2Ab蛋白的核苷酸序列,亚洲玉米螟害虫通过摄食植物组织与Cry1Ab蛋白和Cry2Ab蛋白接触,接触后亚洲玉米螟害虫生长受到抑制和/或导致死亡,同时亚洲玉米螟会延缓产生对于Cry1Ab蛋白的抗性。抑制是指致死或亚致死。同时,植物在形态上应是正常的,且可在常规方法下培养以用于产物的消耗和/或生成。此外,该植物可基本消除对化学或生物杀虫剂的需要。
本发明中,将外源DNA导入植物,如将编码所述Cry2Ab蛋白和/或所述Cry1Ab蛋白的基因或表达盒或重组载体导入植物细胞,常规的转化方法包括但不限于,农杆菌介导的转化、微量发射轰击、直接将DNA摄入原生质体、电穿孔或晶须硅介导的DNA导入。
本发明提供了一种杀虫蛋白组合及其管理昆虫抗性的方法,具有以下优点:
1、延缓抗性。昆虫在持续的选择压力下,会对单一Bt蛋白产生抗性,本发明通过利用两种杀虫蛋白Cry1Ab蛋白和Cry2Ab蛋白有效延缓或防止亚洲玉米螟产生抗性,从而实现对亚洲玉米螟的控制或防治。
2、有效控制抗性害虫。针对已经对Cry1Ab蛋白产生抗性的亚洲玉米螟施用Cry2Ab蛋白可有效控制抗性亚洲玉米螟为害植物,从而使植物更大限度的获得保护并稳定产量。
3、本发明是使Cry1Ab蛋白和Cry2Ab蛋白在植物体内进行表达,且只需种植能够该转基因植物即可,而不需要采用其它措施,从而节省了大量人力、物力和财力,同时效果稳定、彻底。
下面通过附图和实施例,对本发明的技术方案做进一步的详细描述。
附图说明
图1为本发明杀虫蛋白组合及其管理昆虫抗性的方法的含有Cry2Ab核苷酸序列的重组表达载体Cry2Ab-pET30的结构示意图;
图2为本发明杀虫蛋白组合及其管理昆虫抗性的方法的饲喂Cry1Ab蛋白处理的人工饲料的ACB-BtS和ACB-AbR的幼虫校正死亡率示意图;
图3为本发明杀虫蛋白组合及其管理昆虫抗性的方法的饲喂Cry2Ab蛋白处理的人工饲料的ACB-BtS和ACB-AbR的幼虫校正死亡率示意图;
图4为本发明杀虫蛋白组合及其管理昆虫抗性的方法的饲喂Cry1Ab蛋白处理的人工饲料的ACB-BtS和ACB-AbR的幼虫生长抑制率示意图;
图5为本发明杀虫蛋白组合及其管理昆虫抗性的方法的饲喂Cry2Ab蛋白处理的人工饲料的ACB-BtS和ACB-AbR的幼虫生长抑制率示意图。
具体实施方式
下面通过具体实施例进一步说明本发明杀虫蛋白组合及其管理昆虫抗性的方法的技术方案。
第一实施例、Cry2Ab蛋白的表达、纯化及活化
1、含有Cry2Ab核苷酸序列的重组表达载体的构建
Cry2Ab杀虫蛋白质的氨基酸序列(634个氨基酸),如序列表中SEQ ID NO:1所示;根据大肠杆菌密码子的偏好性对Cry2Ab杀虫蛋白质的核苷酸序列进行优化,获得编码相应于所述Cry2Ab杀虫蛋白质的氨基酸序列的Cry2Ab核苷酸序列(1905个核苷酸),如序列表中SEQ ID NO:3所示。
所述Cry2Ab核苷酸序列(如序列表中SEQ ID NO:3所示)由南京金斯瑞生物科技有限公司合成,且由其构建的重组表达载体Cry2Ab-pET30的结构示意图如图1所示(其中,Kan表示卡那霉素抗性基因;f1表示噬菌体f1的复制起点;Cry2Ab为Cry2Ab核苷酸序列(SEQ IDNO:3);Lac I表示操纵子;ori表示复制起点)。
然后将重组表达载体Cry2Ab-pET30用热激方法转化大肠杆菌T1感受态细胞(Transgen,Beijing,China,CAT:CD501),其热激条件为:50μl大肠杆菌T1感受态细胞、10μl质粒DNA(重组表达载体Cry2Ab-pET30),42℃水浴30秒;37℃振荡培养1小时(100rpm转速下摇床摇动),在表面涂有IPTG(异丙基硫代-β-D-半乳糖苷)和X-gal(5-溴-4-氯-3-吲哚-β-D-半乳糖苷)的氨苄青霉素(100mg/L)的LB平板(胰蛋白胨10g/L、酵母提取物5g/L、NaCl10g/L、琼脂15g/L,用NaOH调pH至7.5)上生长过夜。挑取白色菌落,在LB液体培养基(胰蛋白胨10g/L、酵母提取物5g/L、NaCl 10g/L、氨苄青霉素100mg/L,用NaOH调pH至7.5)中于温度37℃条件下培养过夜。碱法提取其质粒:将菌液在12000rpm转速下离心1min,去上清液,沉淀菌体用100μl冰预冷的溶液I(25mM Tris-HCl、10mM EDTA(乙二胺四乙酸)、50mM葡萄糖,pH8.0)悬浮;加入150μl新配制的溶液II(0.2M NaOH、1%SDS(十二烷基硫酸钠)),将管子颠倒4次,混合,置冰上3-5min;加入150μl冰冷的溶液III(4M醋酸钾、2M醋酸),立即充分混匀,冰上放置5-10min;于温度4℃、转速12000rpm条件下离心5min,在上清液中加入2倍体积无水乙醇,混匀后室温放置5min;于温度4℃、转速12000rpm条件下离心5min,弃上清液,沉淀用浓度(V/V)为70%的乙醇洗涤后晾干;加入30μl含RNase(20μg/ml)的TE(10mM Tris-HCl,1mM EDTA,pH8.0)溶解沉淀;于温度37℃下水浴30min,消化RNA;于温度-20℃保存备用。
提取的质粒经KpnI和HindIII酶切鉴定后,对阳性克隆进行测序验证,结果表明重组表达载体Cry2Ab-pET30中插入的所述Cry2Ab核苷酸序列为序列表中SEQ ID NO:3所示的核苷酸序列,即Cry2Ab核苷酸序列正确插入。
2、Cry2Ab蛋白的表达
将测序正确的重组表达载体Cry2Ab-pET30转化到大肠杆菌表达宿主BL21(DE3)(购自北京天根生化科技有限公司)中,具体转化方法为:取50μL冰上融化的BL21(DE3)感受态细胞,加入质粒DNA(重组表达载体Cry2Ab-pET30)并轻轻混匀,于冰上静置25min;于温度42℃下水浴90s,然后迅速放回冰上并静置2min;向离心管中加入800μL不含抗生素的LB液体培养基(胰蛋白胨10g/L、酵母提取物5g/L、NaCl 10g/L,用NaOH调pH至7.5),混匀后在温度37℃、转速150rpm条件下复苏60min;接着在转速4000rpm条件下离心1min,收集菌体,留取100μL左右上清轻轻吹打后重悬并涂布在含有50μg/mL卡那霉素的LB平板(胰蛋白胨10g/L、酵母提取物5g/L、NaCl 10g/L、琼脂15g/L,用NaOH调pH至7.5)上生长过夜。获得表达菌株后,按照下述步骤进行诱导表达:
步骤121、挑取一个含有重组表达载体Cry2Ab-pET30的阳性单克隆,在5mL的LB液体培养基(胰蛋白胨10g/L、酵母提取物5g/L、NaCl 10g/L、卡那霉素50μg/mL,用NaOH调pH至7.5)中于温度37℃条件下震荡培养,使其OD600值达到0.5-0.6;
步骤122、取0.5mL上述菌液于转速12000rpm条件下离心10min,分别取上清和沉淀作为未诱导表达的阴性对照;
步骤123、在剩余的4.5mL上述菌液中加入异丙基硫代半乳糖苷(IPTG)至终浓度为1mM,于转速150rpm、温度16℃条件下继续诱导培养20h;
步骤124、将继续诱导后的菌液于转速12000rpm条件下离心10min后收集菌体,并加入10mM Tris(pH 8.0)悬浮;
步骤125、超声波破碎悬浮后的菌体,于转速12000rpm条件下离心10min,分别收集上清和沉淀并进行SDS-PAGE蛋白电泳检测(参考《蛋白质电泳实验技术(第二版)》)Cry2Ab蛋白的表达,检测结果表明:上清和沉淀中均有Cry2Ab蛋白表达(72KDa)。
3、Cry2Ab蛋白的纯化
根据上述步骤121-125大量诱导表达Cry2Ab蛋白,将收集到的粗蛋白过镍柱纯化(His-Trap,FFGE Healthcare),具体步骤如下:
步骤131、将上述步骤124中收集的菌体溶于结合缓冲液(50mM NaH2PO4、500mMNaCl、20mM咪唑,pH 8.0)中,再置于冰上超声破碎,于转速12000rpm条件下离心30min后收集上清,将上清过0.45uM滤膜除杂;
步骤132、用所述结合缓冲液平衡镍柱,洗5个柱体积,流速为2mL/min;
步骤133、将过滤后的上清上样,流速为0.5mL/min;
步骤134、用所述结合缓冲液再洗5个柱体积,流速为2mL/min,使Cry2Ab蛋白充分挂柱;
步骤135、用所述结合缓冲液继续冲洗,洗去杂蛋白,流速为2mL/min;
步骤136、用洗脱缓冲液(50mM NaH2PO4、500mM NaCl、500mM咪唑,pH 8.0)进行梯度洗脱(依次为:浓度百分比为10%的所述洗脱缓冲液洗脱3个柱体积,浓度百分比为20%的所述洗脱缓冲液洗脱3个柱体积,浓度百分比为40%的所述洗脱缓冲液洗脱3个柱体积,浓度百分比为100%的所述洗脱缓冲液洗脱5个柱体积),流速为2mL/min,分别收集洗脱峰和洗脱液,经过SDS-PAGE蛋白电泳检测后获得纯化后的Cry2Ab蛋白。
4、Cry2Ab蛋白的活化
将上述纯化后的Cry2Ab蛋白溶于50mM碳酸钠缓冲液(pH 10.0)中;将胰蛋白酶配制成浓度为1mg/mL的水溶液,并按100:1(Cry2Ab蛋白:胰蛋白酶)的质量比向Cry2Ab蛋白中加入胰蛋白酶水溶液;混匀后在温度37℃下消化1-3小时。对活化后的Cry2Ab蛋白进行SDS-PAGE蛋白电泳检测,结果表明获得了酶切活化后的Cry2Ab蛋白。
用于下述实验的Cry1Ab蛋白购买自北京乐士宁科技有限公司。
第二实施例、Bt敏感型亚洲玉米螟株系及亚洲玉米螟抗性株系(Cry1Ab蛋白)的生物测定
1、供试昆虫来源
供试昆虫为亚洲玉米螟(Ostrinia furnacalis)的Bt敏感株系(下文以ACB-BtS代表亚洲玉米螟Bt敏感株系)和亚洲玉米螟(Ostrinia furnacalis)的Bt抗性株系(下文以ACB-AbR代表亚洲玉米螟Bt抗性株系Ab0.8,即以Cry1Ab浓度为0.8μg/g的饲料继代培育的亚洲玉米螟Bt抗性株系),上述两种株系的昆虫均来自于中国农业科学院植物保护研究所。
2、亚洲玉米螟生物测定方法
针对每种Bt蛋白使用不同浓度进行生物测定,测定Cry1Ab蛋白的浓度范围为0μg/g至50μg/g,测定Cry2Ab蛋白的浓度范围为0μg/g至100μg/g。Bt蛋白溶液的制备方式为:将Cry1Ab蛋白与蒸馏水(或缓冲液)按照一定比例混合制成浓度分别为0μg/g、0.01μg/g、0.05μg/g、0.1μg/g、0.5μg/g、1μg/g、5μg/g、10μg/g和50μg/g的Cry1Ab蛋白溶液。将Cry2Ab蛋白与蒸馏水(或缓冲液)按照一定比例混合制成浓度分别为0μg/g、0.01μg/g、0.05μg/g、0.1μg/g、0.5μg/g、1μg/g、5μg/g、10μg/g、50μg/g和100μg/g的Cry2Ab蛋白溶液。将不同浓度的Bt蛋白溶液(Cry1Ab蛋白溶液或Cry2Ab蛋白溶液)与人工饲料(人工饲料来自于中国农业科学院植物保护研究所)按1.5:1分别混合成混合饲料,将混合饲料分散放置于48孔的生物测定板中,每个孔中放入约0.5g混合饲料,将与Bt蛋白溶液(Cry1Ab蛋白溶液或Cry2Ab蛋白溶液)等量的蒸馏水(或缓冲液)处理的人工饲料作为空白对照。在生物测定板每个孔的混合饲料表面上接种1只亚洲玉米螟的新生幼虫(存活<24h),用封口膜覆盖孔。将生物测定板在温度28℃、相对湿度80%、光周期(光/暗)16:8的条件下放置6天,接种后第7天开始记录幼虫死亡率和幼虫生长抑制率。昆虫株系与Bt蛋白溶液(Cry1Ab蛋白溶液或Cry2Ab蛋白溶液)浓度的组合每组重复3次,每个重复包括48只幼虫。
幼虫死亡率以“实际死亡率”(下文简称为死亡率)体现,计算死亡率需考虑实际死亡的幼虫和不显示体重显著增加(<0.1mg/幼虫)的存活幼虫。亚洲玉米螟的死亡率用如下的公式计算:死亡率(%)=100×[死亡幼虫数目+不显示体重显著增加(<0.1mg/幼虫)的存活幼虫数目]/测试昆虫的总数。每个幼虫的死亡率根据饲喂空白对照的幼虫死亡率进行校正。然后对经过校正的剂量和死亡率数据进行机率分析,应用POLO统计软件确定导致50%(LC50)死亡率的Bt蛋白浓度(Cry1Ab蛋白溶液或Cry2Ab蛋白溶液)和相应的95%置信区间。通过将ACB-AbR的LC50值除以ACB-BtS的LC50值计算抗性比例。
饲喂混合饲料的亚洲玉米螟的幼虫生长抑制率用以下公式计算:幼虫生长抑制(%)=100×(用空白对照饲喂的幼虫体重-用混合饲料饲喂的幼虫体重)/(用空白对照饲喂的幼虫体重)。如果没有体重显著增加(<0.1mg/幼虫)的幼虫,则指定该重复为100%的幼虫生长抑制。生长抑制数据用双向ANOVA进行分析,以昆虫株系和Bt蛋白溶液浓度作为两个主要因子。使用LSMEANS检验确定α=0.05水平的处理差异。
3、饲喂混合饲料的昆虫株系的幼虫死亡率及生长抑制率测定结果
饲喂Cry1Ab蛋白处理的混合饲料的ACB-BtS和ACB-AbR的幼虫死亡率均随着Cry1Ab蛋白浓度的增加而提高,且在同一浓度下ACB-BtS的幼虫死亡率显著高于ACB-AbR。如表1和图2所示,基于ACB-BtS的幼虫死亡率计算的LC50值为0.11μg/g;基于ACB-AbR的幼虫死亡率计算的LC50值16.25μg/g,因此ACB-BtS和ACB-AbR的LC50值的差异是极显著的,两者对于Cry1Ab蛋白的抗性差异也是极显著的,即Cry1Ab蛋白对于ACB-AbR具有较弱的杀虫活性。
饲喂Cry2Ab蛋白处理的混合饲料的ACB-BtS和ACB-AbR的幼虫死亡率均随着Cry2Ab蛋白浓度的增加而提高,且在同一浓度下ACB-BtS的幼虫死亡率高于ACB-AbR。如表1和图3所示,基于ACB-BtS的幼虫死亡率计算的LC50值为1.23μg/g;基于ACB-AbR的幼虫死亡率计算的LC50值3.57μg/g,因此ACB-BtS和ACB-AbR的LC50值的差异是显著的,但Cry2Ab蛋白对于ACB-BtS和ACB-AbR均具有杀虫活性。
如图4所示,横坐标为所用Bt蛋白浓度(μg/g)的对数值,纵坐标为对应亚洲玉米螟的生长抑制率,饲喂Cry1Ab蛋白处理的混合饲料的ACB-BtS和ACB-AbR的幼虫生长抑制率均随着Cry1Ab蛋白浓度的增加而提高,且在同一浓度下ACB-BtS的幼虫生长抑制率显著高于ACB-AbR;达到100%生长抑制率时,ACB-BtS所需Cry1Ab蛋白的浓度为约1μg/g,ACB-AbR所需Cry1Ab蛋白的浓度为约100μg/g,表明Cry1Ab蛋白对于ACB-AbR具有较弱的抑制活性。
如图5所示,饲喂Cry2Ab蛋白处理的混合饲料的ACB-BtS和ACB-AbR的幼虫生长抑制率均随着Cry2Ab蛋白浓度的增加而提高,且在同一浓度下ACB-BtS的幼虫生长抑制率高于ACB-AbR;达到100%生长抑制率时,ACB-BtS所需Cry2Ab蛋白的浓度为约50μg/g,ACB-AbR所需Cry2Ab蛋白的浓度为约50μg/g,表明Cry2Ab蛋白对于ACB-BtS和ACB-AbR具有相当的抑制活性。
表1:Bt蛋白对ACB-BtS和ACB-AbR新生幼虫的毒性结果
上述结果表明,ACB-AbR对Cry1Ab蛋白具有148倍抗性(即抗性比例为148),且Cry2Ab蛋白对于ACB-BtS和ACB-AbR均具有较好的杀虫活性,故ACB-AbR显示出对Cry2Ab蛋白较低的交互抗性,抗性比例显著降低为2.9。由此可见,Cry2Ab蛋白与Cry1Ab蛋白的交互抗性较低,Cry2Ab蛋白与Cry1Ab蛋白组合应用可以延缓或推迟亚洲玉米螟对Cry1Ab蛋白产生抗性,同时Cry2Ab蛋白可以有效管理亚洲玉米螟物种对Cry1Ab蛋白产生的抗性。
第三实施例、Cry2Ab蛋白与Cry1Ab蛋白在生产抗虫转基因植物中的用途
通过第二实施例的实验结果可知,Cry2Ab蛋白与Cry1Ab蛋白预期可以用于在转基因植物例如玉米植物中组合表达以延缓或防止亚洲玉米螟对该植物产生抗性。
第一种方法为相继转化:其中对已转化了第一基因(例如Cry1Ab基因)的植物进行再转化,以引入第二基因(例如Cry2Ab基因)。该相继转化优选地使用两个不同的选择标记基因,例如卡那霉素抗性基因和赋予对草铵膦除草剂的抗性的膦丝菌素乙酰转移酶基因(例如pat或bar基因)。
第二种方法为共转化方法:编码Cry1Ab蛋白的核苷酸序列在植物中与编码Cry2Ab蛋白的核苷酸序列一起表达,可以通过利用与各自基因连接的选择标记,整体筛选包含两个选择基因的植物。
第三种方法为独立的转化事件,将两个杀虫蛋白基因各自单独地转移至不同植物的基因组中,随后可以通过杂交将其组合在单独的植物中,并且可以使用DNA标记技术来选择包含这些不同基因的植物。
综上所述,本发明通过利用两种杀虫蛋白Cry1Ab蛋白和Cry2Ab蛋白有效延缓或防止亚洲玉米螟产生抗性,从而实现对亚洲玉米螟的控制或防治,使植物更大限度的获得保护并稳定产量;同时本发明使Cry1Ab蛋白和Cry2Ab蛋白在植物体内进行表达,且只需种植能够该转基因植物即可,而不需要采用其它措施,从而节省了大量人力、物力和财力,同时效果稳定、彻底。
最后所应说明的是,以上实施例仅用以说明本发明的技术方案而非限制,尽管参照较佳实施例对本发明进行了详细说明,本领域的普通技术人员应当理解,可以对本发明的技术方案进行修改或者等同替换,而不脱离本发明技术方案的精神和范围。
SEQUENCE LISTING
<110> 北京大北农科技集团股份有限公司
北京大北农生物技术有限公司
<120> 杀虫蛋白组合及其管理昆虫抗性的方法
<130> DBNBC115
<160> 4
<170> PatentIn version 3.5
<210> 1
<211> 634
<212> PRT
<213> Artificial Sequence
<220>
<223> Cry2Ab氨基酸序列
<400> 1
Met Asp Asn Ser Val Leu Asn Ser Gly Arg Thr Thr Ile Cys Asp Ala
1 5 10 15
Tyr Asn Val Ala Ala His Asp Pro Phe Ser Phe Gln His Lys Ser Leu
20 25 30
Asp Thr Val Gln Lys Glu Trp Thr Glu Trp Lys Lys Asn Asn His Ser
35 40 45
Leu Tyr Leu Asp Pro Ile Val Gly Thr Val Ala Ser Phe Leu Leu Lys
50 55 60
Lys Val Gly Ser Leu Val Gly Lys Arg Ile Leu Ser Glu Leu Arg Asn
65 70 75 80
Leu Ile Phe Pro Ser Gly Ser Thr Asn Leu Met Gln Asp Ile Leu Arg
85 90 95
Glu Thr Glu Lys Phe Leu Asn Gln Arg Leu Asn Thr Asp Thr Leu Ala
100 105 110
Arg Val Asn Ala Glu Leu Thr Gly Leu Gln Ala Asn Val Glu Glu Phe
115 120 125
Asn Arg Gln Val Asp Asn Phe Leu Asn Pro Asn Arg Asn Ala Val Pro
130 135 140
Leu Ser Ile Thr Ser Ser Val Asn Thr Met Gln Gln Leu Phe Leu Asn
145 150 155 160
Arg Leu Pro Gln Phe Gln Met Gln Gly Tyr Gln Leu Leu Leu Leu Pro
165 170 175
Leu Phe Ala Gln Ala Ala Asn Leu His Leu Ser Phe Ile Arg Asp Val
180 185 190
Ile Leu Asn Ala Asp Glu Trp Gly Ile Ser Ala Ala Thr Leu Arg Thr
195 200 205
Tyr Arg Asp Tyr Leu Lys Asn Tyr Thr Arg Asp Tyr Ser Asn Tyr Cys
210 215 220
Ile Asn Thr Tyr Gln Ser Ala Phe Lys Gly Leu Asn Thr Arg Leu His
225 230 235 240
Asp Met Leu Glu Phe Arg Thr Tyr Met Phe Leu Asn Val Phe Glu Tyr
245 250 255
Val Ser Ile Trp Ser Leu Phe Lys Tyr Gln Ser Leu Leu Val Ser Ser
260 265 270
Gly Ala Asn Leu Tyr Ala Ser Gly Ser Gly Pro Gln Gln Thr Gln Ser
275 280 285
Phe Thr Ser Gln Asp Trp Pro Phe Leu Tyr Ser Leu Phe Gln Val Asn
290 295 300
Ser Asn Tyr Val Leu Asn Gly Phe Ser Gly Ala Arg Leu Ser Asn Thr
305 310 315 320
Phe Pro Asn Ile Val Gly Leu Pro Gly Ser Thr Thr Thr His Ala Leu
325 330 335
Leu Ala Ala Arg Val Asn Tyr Ser Gly Gly Ile Ser Ser Gly Asp Ile
340 345 350
Gly Ala Ser Pro Phe Asn Gln Asn Phe Asn Cys Ser Thr Phe Leu Pro
355 360 365
Pro Leu Leu Thr Pro Phe Val Arg Ser Trp Leu Asp Ser Gly Ser Asp
370 375 380
Arg Glu Gly Val Ala Thr Val Thr Asn Trp Gln Thr Glu Ser Phe Glu
385 390 395 400
Thr Thr Leu Gly Leu Arg Ser Gly Ala Phe Thr Ala Arg Gly Asn Ser
405 410 415
Asn Tyr Phe Pro Asp Tyr Phe Ile Arg Asn Ile Ser Gly Val Pro Leu
420 425 430
Val Val Arg Asn Glu Asp Leu Arg Arg Pro Leu His Tyr Asn Glu Ile
435 440 445
Arg Asn Ile Ala Ser Pro Ser Gly Thr Pro Gly Gly Ala Arg Ala Tyr
450 455 460
Met Val Ser Val His Asn Arg Lys Asn Asn Ile His Ala Val His Glu
465 470 475 480
Asn Gly Ser Met Ile His Leu Ala Pro Asn Asp Tyr Thr Gly Phe Thr
485 490 495
Ile Ser Pro Ile His Ala Thr Gln Val Asn Asn Gln Thr Arg Thr Phe
500 505 510
Ile Ser Glu Lys Phe Gly Asn Gln Gly Asp Ser Leu Arg Phe Glu Gln
515 520 525
Asn Asn Thr Thr Ala Arg Tyr Thr Leu Arg Gly Asn Gly Asn Ser Tyr
530 535 540
Asn Leu Tyr Leu Arg Val Ser Ser Ile Gly Asn Ser Thr Ile Arg Val
545 550 555 560
Thr Ile Asn Gly Arg Val Tyr Thr Ala Thr Asn Val Asn Thr Thr Thr
565 570 575
Asn Asn Asp Gly Val Asn Asp Asn Gly Ala Arg Phe Ser Asp Ile Asn
580 585 590
Ile Gly Asn Val Val Ala Ser Ser Asn Ser Asp Val Pro Leu Asp Ile
595 600 605
Asn Val Thr Leu Asn Ser Gly Thr Gln Phe Asp Leu Met Asn Ile Met
610 615 620
Leu Val Pro Thr Asn Ile Ser Pro Leu Tyr
625 630
<210> 2
<211> 818
<212> PRT
<213> Artificial Sequence
<220>
<223> Cry1Ab氨基酸序列
<400> 2
Met Asp Asn Asn Pro Asn Ile Asn Glu Cys Ile Pro Tyr Asn Cys Leu
1 5 10 15
Ser Asn Pro Glu Val Glu Val Leu Gly Gly Glu Arg Ile Glu Thr Gly
20 25 30
Tyr Thr Pro Ile Asp Ile Ser Leu Ser Leu Thr Gln Phe Leu Leu Ser
35 40 45
Glu Phe Val Pro Gly Ala Gly Phe Val Leu Gly Leu Val Asp Ile Ile
50 55 60
Trp Gly Ile Phe Gly Pro Ser Gln Trp Asp Ala Phe Leu Val Gln Ile
65 70 75 80
Glu Gln Leu Ile Asn Gln Arg Ile Glu Glu Phe Ala Arg Asn Gln Ala
85 90 95
Ile Ser Arg Leu Glu Gly Leu Ser Asn Leu Tyr Gln Ile Tyr Ala Glu
100 105 110
Ser Phe Arg Glu Trp Glu Ala Asp Pro Thr Asn Pro Ala Leu Arg Glu
115 120 125
Glu Met Arg Ile Gln Phe Asn Asp Met Asn Ser Ala Leu Thr Thr Ala
130 135 140
Ile Pro Leu Phe Ala Val Gln Asn Tyr Gln Val Pro Leu Leu Ser Val
145 150 155 160
Tyr Val Gln Ala Ala Asn Leu His Leu Ser Val Leu Arg Asp Val Ser
165 170 175
Val Phe Gly Gln Arg Trp Gly Phe Asp Ala Ala Thr Ile Asn Ser Arg
180 185 190
Tyr Asn Asp Leu Thr Arg Leu Ile Gly Asn Tyr Thr Asp His Ala Val
195 200 205
Arg Trp Tyr Asn Thr Gly Leu Glu Arg Val Trp Gly Pro Asp Ser Arg
210 215 220
Asp Trp Ile Arg Tyr Asn Gln Phe Arg Arg Glu Leu Thr Leu Thr Val
225 230 235 240
Leu Asp Ile Val Ser Leu Phe Pro Asn Tyr Asp Ser Arg Thr Tyr Pro
245 250 255
Ile Arg Thr Val Ser Gln Leu Thr Arg Glu Ile Tyr Thr Asn Pro Val
260 265 270
Leu Glu Asn Phe Asp Gly Ser Phe Arg Gly Ser Ala Gln Gly Ile Glu
275 280 285
Gly Ser Ile Arg Ser Pro His Leu Met Asp Ile Leu Asn Ser Ile Thr
290 295 300
Ile Tyr Thr Asp Ala His Arg Gly Glu Tyr Tyr Trp Ser Gly His Gln
305 310 315 320
Ile Met Ala Ser Pro Val Gly Phe Ser Gly Pro Glu Phe Thr Phe Pro
325 330 335
Leu Tyr Gly Thr Met Gly Asn Ala Ala Pro Gln Gln Arg Ile Val Ala
340 345 350
Gln Leu Gly Gln Gly Val Tyr Arg Thr Leu Ser Ser Thr Leu Tyr Arg
355 360 365
Arg Pro Phe Asn Ile Gly Ile Asn Asn Gln Gln Leu Ser Val Leu Asp
370 375 380
Gly Thr Glu Phe Ala Tyr Gly Thr Ser Ser Asn Leu Pro Ser Ala Val
385 390 395 400
Tyr Arg Lys Ser Gly Thr Val Asp Ser Leu Asp Glu Ile Pro Pro Gln
405 410 415
Asn Asn Asn Val Pro Pro Arg Gln Gly Phe Ser His Arg Leu Ser His
420 425 430
Val Ser Met Phe Arg Ser Gly Phe Ser Asn Ser Ser Val Ser Ile Ile
435 440 445
Arg Ala Pro Met Phe Ser Trp Ile His Arg Ser Ala Glu Phe Asn Asn
450 455 460
Ile Ile Pro Ser Ser Gln Ile Thr Gln Ile Pro Leu Thr Lys Ser Thr
465 470 475 480
Asn Leu Gly Ser Gly Thr Ser Val Val Lys Gly Pro Gly Phe Thr Gly
485 490 495
Gly Asp Ile Leu Arg Arg Thr Ser Pro Gly Gln Ile Ser Thr Leu Arg
500 505 510
Val Asn Ile Thr Ala Pro Leu Ser Gln Arg Tyr Arg Val Arg Ile Arg
515 520 525
Tyr Ala Ser Thr Thr Asn Leu Gln Phe His Thr Ser Ile Asp Gly Arg
530 535 540
Pro Ile Asn Gln Gly Asn Phe Ser Ala Thr Met Ser Ser Gly Ser Asn
545 550 555 560
Leu Gln Ser Gly Ser Phe Arg Thr Val Gly Phe Thr Thr Pro Phe Asn
565 570 575
Phe Ser Asn Gly Ser Ser Val Phe Thr Leu Ser Ala His Val Phe Asn
580 585 590
Ser Gly Asn Glu Val Tyr Ile Asp Arg Ile Glu Phe Val Pro Ala Glu
595 600 605
Val Thr Phe Glu Ala Glu Tyr Asp Leu Glu Arg Ala Gln Lys Ala Val
610 615 620
Asn Glu Leu Phe Thr Ser Ser Asn Gln Ile Gly Leu Lys Thr Asp Val
625 630 635 640
Thr Asp Tyr His Ile Asp Gln Val Ser Asn Leu Val Glu Cys Leu Ser
645 650 655
Asp Glu Phe Cys Leu Asp Glu Lys Lys Glu Leu Ser Glu Lys Val Lys
660 665 670
His Ala Lys Arg Leu Ser Asp Glu Arg Asn Leu Leu Gln Asp Pro Asn
675 680 685
Phe Arg Gly Ile Asn Arg Gln Leu Asp Arg Gly Trp Arg Gly Ser Thr
690 695 700
Asp Ile Thr Ile Gln Gly Gly Asp Asp Val Phe Lys Glu Asn Tyr Val
705 710 715 720
Thr Leu Leu Gly Thr Phe Asp Glu Cys Tyr Pro Thr Tyr Leu Tyr Gln
725 730 735
Lys Ile Asp Glu Ser Lys Leu Lys Ala Tyr Thr Arg Tyr Gln Leu Arg
740 745 750
Gly Tyr Ile Glu Asp Ser Gln Asp Leu Glu Ile Tyr Leu Ile Arg Tyr
755 760 765
Asn Ala Lys His Glu Thr Val Asn Val Pro Gly Thr Gly Ser Leu Trp
770 775 780
Pro Leu Ser Ala Pro Ser Pro Ile Gly Lys Cys Ala His His Ser His
785 790 795 800
His Phe Ser Leu Asp Ile Asp Val Gly Cys Thr Asp Leu Asn Glu Asp
805 810 815
Phe Arg
<210> 3
<211> 1905
<212> DNA
<213> Artificial Sequence
<220>
<223> Cry2Ab核苷酸序列
<400> 3
atggataact cagttctgaa ttcgggccgt accacgattt gcgatgccta taatgtcgcg 60
gcccatgacc cgtttagctt ccaacacaaa tctctggata ccgtgcagaa agaatggacg 120
gaatggaaga aaaacaatca tagcctgtac ctggacccga ttgtgggcac cgttgcgagc 180
tttctgctga agaaagtggg ctctctggtc ggtaaacgta ttctgagtga actgcgcaat 240
ctgatctttc cgtcaggttc gaccaacctg atgcaagata tcctgcgtga aacggaaaaa 300
ttcctgaacc agcgtctgaa taccgacacg ctggcgcgcg ttaacgccga actgaccggt 360
ctgcaagcaa acgtggaaga atttaatcgt caggttgata acttcctgaa cccgaatcgc 420
aacgctgtcc cgctgtctat taccagctct gtgaatacga tgcagcaact gtttctgaac 480
cgcctgccgc agttccaaat gcagggctat caactgctgc tgctgccgct gtttgcgcag 540
gcagctaatc tgcatctgag cttcattcgt gatgtgatcc tgaacgccga cgaatggggt 600
atttctgcgg ccaccctgcg tacgtatcgc gattacctga aaaattatac ccgcgactat 660
tccaactact gcatcaatac gtaccagtca gcatttaaag gcctgaatac ccgtctgcac 720
gatatgctgg aattccgcac ctatatgttt ctgaacgtgt tcgaatacgt tagcatttgg 780
tctctgttta aatatcagag cctgctggtt agctccggtg caaacctgta cgctagcggc 840
tctggtccgc agcaaaccca aagtttcacg tcccaggatt ggccgtttct gtattcactg 900
ttccaggtca attcgaacta cgtgctgaat ggctttagtg gtgcccgtct gtccaacacc 960
ttcccgaata tcgtgggtct gccgggttca accacgaccc acgcactgct ggcagctcgt 1020
gttaactatt cgggcggtat ttcatcgggc gatatcggtg cctcgccgtt taatcagaac 1080
ttcaattgta gcacctttct gccgccgctg ctgacgccgt tcgttcgtag ctggctggat 1140
agtggttccg accgtgaagg tgtcgcaacc gtgacgaatt ggcagaccga atcttttgaa 1200
acgaccctgg gcctgcgtag tggtgcattc accgctcgcg gcaactccaa ctacttcccg 1260
gattacttca tccgtaacat cagcggcgtg ccgctggttg tgcgcaacga agacctgcgt 1320
cgcccgctgc attataacga aattcgcaat atcgcctcac cgtcgggcac cccgggcggt 1380
gcacgtgcat acatggtttc tgtccacaac cgcaaaaaca atattcatgc ggtccacgaa 1440
aatggcagta tgatccatct ggccccgaac gattataccg gttttacgat ttccccgatc 1500
cacgcaaccc aagtgaacaa tcagacccgt acgtttattt cagaaaaatt cggcaatcag 1560
ggtgattcgc tgcgctttga acagaacaat acgaccgctc gttataccct gcgcggcaac 1620
ggtaatagtt ataacctgta cctgcgtgtt agctctatcg gcaattccac cattcgtgtt 1680
acgatcaacg gtcgcgtcta caccgcgacg aacgtgaata cgaccacgaa caatgatggc 1740
gtgaacgaca atggtgcacg ctttagcgat attaacatcg gcaatgtcgt ggctagttcc 1800
aatagcgatg ttccgctgga cattaacgtc accctgaatt ctggtacgca gtttgacctg 1860
atgaatatta tgctggtgcc gaccaacatc agcccgctgt attaa 1905
<210> 4
<211> 2457
<212> DNA
<213> Artificial Sequence
<220>
<223> Cry1Ab核苷酸序列
<400> 4
atggacaaca acccaaacat caacgagtgc atcccgtaca actgcctcag caaccctgag 60
gtcgaggtgc tcggcggtga gcgcatcgag accggttaca cccccatcga catctccctc 120
tccctcacgc agttcctgct cagcgagttc gtgccaggcg ctggcttcgt cctgggcctc 180
gtggacatca tctggggcat ctttggcccc tcccagtggg acgccttcct ggtgcaaatc 240
gagcagctca tcaaccagag gatcgaggag ttcgccagga accaggccat cagccgcctg 300
gagggcctca gcaacctcta ccaaatctac gctgagagct tccgcgagtg ggaggccgac 360
cccactaacc cagctctccg cgaggagatg cgcatccagt tcaacgacat gaacagcgcc 420
ctgaccaccg ccatcccact cttcgccgtc cagaactacc aagtcccgct cctgtccgtg 480
tacgtccagg ccgccaacct gcacctcagc gtgctgaggg acgtcagcgt gtttggccag 540
aggtggggct tcgacgccgc caccatcaac agccgctaca acgacctcac caggctgatc 600
ggcaactaca ccgaccacgc tgtccgctgg tacaacactg gcctggagcg cgtctggggc 660
cctgattcta gagactggat tcgctacaac cagttcaggc gcgagctgac cctcaccgtc 720
ctggacattg tgtccctctt cccgaactac gactcccgca cctacccgat ccgcaccgtg 780
tcccaactga cccgcgaaat ctacaccaac cccgtcctgg agaacttcga cggtagcttc 840
aggggcagcg cccagggcat cgagggctcc atcaggagcc cacacctgat ggacatcctc 900
aacagcatca ctatctacac cgatgcccac cgcggcgagt actactggtc cggccaccag 960
atcatggcct ccccggtcgg cttcagcggc cccgagttta cctttcctct ctacggcacg 1020
atgggcaacg ccgctccaca acaacgcatc gtcgctcagc tgggccaggg cgtctaccgc 1080
accctgagct ccaccctgta ccgcaggccc ttcaacatcg gtatcaacaa ccagcagctg 1140
tccgtcctgg atggcactga gttcgcctac ggcacctcct ccaacctgcc ctccgctgtc 1200
taccgcaaga gcggcacggt ggattccctg gacgagatcc caccacagaa caacaatgtg 1260
ccccccaggc agggtttttc ccacaggctc agccacgtgt ccatgttccg ctccggcttc 1320
agcaactcgt ccgtgagcat catcagagct cctatgttct cctggattca tcgcagcgcg 1380
gagttcaaca atatcattcc gtcctcccaa atcacccaaa tccccctcac caagtccacc 1440
aacctgggca gcggcacctc cgtggtgaag ggcccaggct tcacgggcgg cgacatcctg 1500
cgcaggacct ccccgggcca gatcagcacc ctccgcgtca acatcaccgc tcccctgtcc 1560
cagaggtacc gcgtcaggat tcgctacgct agcaccacca acctgcaatt ccacacctcc 1620
atcgacggca ggccgatcaa tcagggtaac ttctccgcca ccatgtccag cggcagcaac 1680
ctccaatccg gcagcttccg caccgtgggt ttcaccaccc ccttcaactt ctccaacggc 1740
tccagcgttt tcaccctgag cgcccacgtg ttcaattccg gcaatgaggt gtacattgac 1800
cgcattgagt tcgtgccagc cgaggtcacc ttcgaagccg agtacgacct ggagagagcc 1860
cagaaggctg tcaatgagct cttcacgtcc agcaatcaga tcggcctgaa gaccgacgtc 1920
actgactacc acatcgacca agtctccaac ctcgtggagt gcctctccga tgagttctgc 1980
ctcgacgaga agaaggagct gtccgagaag gtgaagcatg ccaagcgtct cagcgacgag 2040
aggaatctcc tccaggaccc caatttccgc ggcatcaaca ggcagctcga ccgcggctgg 2100
cgcggcagca ccgacatcac gatccagggc ggcgacgatg tgttcaagga gaactacgtg 2160
actctcctgg gcactttcga cgagtgctac cctacctact tgtaccagaa gatcgatgag 2220
tccaagctca aggcttacac tcgctaccag ctccgcggct acatcgaaga cagccaagac 2280
ctcgagattt acctgatccg ctacaacgcc aagcacgaga ccgtcaacgt gcccggtact 2340
ggttccctct ggccgctgag cgcccccagc ccgatcggca agtgtgccca ccacagccac 2400
cacttctcct tggacatcga tgtgggctgc accgacctga acgaggactt tcggtag 2457
Claims (26)
1.一种管理昆虫抗性的方法,其特征在于,包括将亚洲玉米螟至少与Cry2Ab蛋白和Cry1Ab蛋白接触。
2.根据权利要求1所述管理昆虫抗性的方法,其特征在于,所述Cry2Ab蛋白和Cry1Ab蛋白存在于至少产生所述Cry2Ab蛋白和Cry1Ab蛋白的细菌或转基因植物中,所述亚洲玉米螟通过摄食所述细菌或所述转基因植物的组织至少与所述Cry2Ab蛋白和Cry1Ab蛋白接触,接触后所述亚洲玉米螟生长受到抑制和/或导致死亡,以实现管理亚洲玉米螟的抗性。
3.根据权利要求1所述管理昆虫抗性的方法,其特征在于,将对Cry1Ab蛋白产生抗性的亚洲玉米螟至少与所述Cry2Ab蛋白接触,所述Cry2Ab蛋白存在于至少产生所述Cry2Ab蛋白的细菌或转基因植物中,所述对Cry1Ab蛋白产生抗性的亚洲玉米螟通过摄食所述细菌或所述转基因植物的组织至少与所述Cry2Ab蛋白接触,接触后所述对Cry1Ab蛋白产生抗性的亚洲玉米螟生长受到抑制和/或导致死亡,以实现管理所述对Cry1Ab蛋白产生抗性的亚洲玉米螟的抗性。
4.根据权利要求1所述管理昆虫抗性的方法,其特征在于,将对Cry2Ab蛋白产生抗性的亚洲玉米螟至少与所述Cry1Ab蛋白接触,所述Cry1Ab蛋白存在于至少产生所述Cry1Ab蛋白的细菌或转基因植物中,所述对Cry2Ab蛋白产生抗性的亚洲玉米螟通过摄食所述细菌或所述转基因植物的组织至少与所述Cry1Ab蛋白接触,接触后所述对Cry2Ab蛋白产生抗性的亚洲玉米螟生长受到抑制和/或导致死亡,以实现管理所述对Cry2Ab蛋白产生抗性的亚洲玉米螟的抗性。
5.根据权利要求2-4任一项所述管理昆虫抗性的方法,其特征在于,所述转基因植物处于任意生育期。
6.根据权利要求2-4任一项所述管理昆虫抗性的方法,其特征在于,所述转基因植物的组织为根、叶片、茎秆、果实、雄穗、雌穗、花药或花丝。
7.根据权利要求2-4任一项所述管理昆虫抗性的方法,其特征在于,所述对亚洲玉米螟危害植物的控制不因种植地点和/或种植时间的改变而改变。
8.根据权利要求2-7任一项所述管理昆虫抗性的方法,其特征在于,所述植物为玉米、小麦、高粱、谷子、水稻或大豆。
9.根据权利要求1-8任一项所述管理昆虫抗性的方法,其特征在于,所述Cry2Ab蛋白的氨基酸序列具有SEQ ID NO:1所示的氨基酸序列。
10.根据权利要求9所述管理昆虫抗性的方法,其特征在于,所述Cry2Ab蛋白的核苷酸序列具有SEQ ID NO:3所示的核苷酸序列。
11.根据权利要求1-10任一项所述管理昆虫抗性的方法,其特征在于,所述Cry1Ab蛋白的氨基酸序列具有SEQ ID NO:2所示的氨基酸序列。
12.根据权利要求11所述管理昆虫抗性的方法,其特征在于,所述Cry1Ab蛋白的核苷酸序列具有SEQ ID NO:4所示的核苷酸序列。
13.一种控制亚洲玉米螟的方法,其特征在于,包括将亚洲玉米螟至少与Cry2Ab蛋白和Cry1Ab蛋白接触,从而实现对亚洲玉米螟的控制。
14.根据权利要求13所述控制亚洲玉米螟的方法,其特征在于,所述Cry2Ab蛋白和Cry1Ab蛋白存在于至少产生所述Cry2Ab蛋白和Cry1Ab蛋白的细菌或转基因植物中,所述亚洲玉米螟通过摄食所述细菌或所述转基因植物的组织至少与所述Cry2Ab蛋白和Cry1Ab蛋白接触,接触后所述亚洲玉米螟生长受到抑制和/或导致死亡,以实现对亚洲玉米螟危害植物的控制。
15.根据权利要求13所述控制亚洲玉米螟的方法,其特征在于,将对Cry1Ab蛋白产生抗性的亚洲玉米螟至少与所述Cry2Ab蛋白接触,所述Cry2Ab蛋白存在于至少产生所述Cry2Ab蛋白的细菌或转基因植物中,所述对Cry1Ab蛋白产生抗性的亚洲玉米螟通过摄食所述细菌或所述转基因植物的组织至少与所述Cry2Ab蛋白接触,接触后所述对Cry1Ab蛋白产生抗性的亚洲玉米螟生长受到抑制和/或导致死亡,以实现对所述对Cry1Ab蛋白产生抗性的亚洲玉米螟危害植物的控制。
16.根据权利要求13所述控制亚洲玉米螟的方法,其特征在于,将对Cry2Ab蛋白产生抗性的亚洲玉米螟至少与所述Cry1Ab蛋白接触,所述Cry1Ab蛋白存在于至少产生所述Cry1Ab蛋白的细菌或转基因植物中,所述对Cry2Ab蛋白产生抗性的亚洲玉米螟通过摄食所述细菌或所述转基因植物的组织至少与所述Cry1Ab蛋白接触,接触后所述对Cry2Ab蛋白产生抗性的亚洲玉米螟生长受到抑制和/或导致死亡,以实现对所述对Cry2Ab蛋白产生抗性的亚洲玉米螟危害植物的控制。
17.根据权利要求14-16任一项所述控制亚洲玉米螟的方法,其特征在于,所述转基因植物处于任意生育期。
18.根据权利要求14-16任一项所述控制亚洲玉米螟的方法,其特征在于,所述转基因植物的组织为根、叶片、茎秆、果实、雄穗、雌穗、花药或花丝。
19.根据权利要求14-16任一项所述控制亚洲玉米螟的方法,其特征在于,所述对亚洲玉米螟危害植物的控制不因种植地点和/或种植时间的改变而改变。
20.根据权利要求14-19任一项所述控制亚洲玉米螟的方法,其特征在于,所述植物为玉米、小麦、高粱、谷子、水稻或大豆。
21.根据权利要求13-20任一项所述控制亚洲玉米螟的方法,其特征在于,所述Cry2Ab蛋白的氨基酸序列具有SEQ ID NO:1所示的氨基酸序列。
22.根据权利要求21所述控制亚洲玉米螟的方法,其特征在于,所述Cry2Ab蛋白的核苷酸序列具有SEQ ID NO:3所示的核苷酸序列。
23.根据权利要求13-22任一项所述控制亚洲玉米螟的方法,其特征在于,所述Cry1Ab蛋白的氨基酸序列具有SEQ ID NO:2所示的氨基酸序列。
24.根据权利要求23所述控制亚洲玉米螟的方法,其特征在于,所述Cry1Ab蛋白的核苷酸序列具有SEQ ID NO:4所示的核苷酸序列。
25.一种Cry2Ab蛋白和Cry1Ab蛋白组合使用以防止或延缓亚洲玉米螟群体对Cry1Ab蛋白或Cry2Ab蛋白产生抗性的用途。
26.一种Cry2Ab蛋白和Cry1Ab蛋白组合使用以控制对Cry1Ab蛋白或Cry2Ab蛋白产生抗性的亚洲玉米螟群体的用途。
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