CN110225974B - 使用cry14来控制线虫害虫 - Google Patents
使用cry14来控制线虫害虫 Download PDFInfo
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- CN110225974B CN110225974B CN201780082613.6A CN201780082613A CN110225974B CN 110225974 B CN110225974 B CN 110225974B CN 201780082613 A CN201780082613 A CN 201780082613A CN 110225974 B CN110225974 B CN 110225974B
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Abstract
提供了赋予细菌、植物、植物细胞、组织和种子杀线虫活性的组合物和方法。特别地,提供了杀死或控制线虫害虫种群,特别是短体线虫,例如,短尾短体线虫种群的方法。该方法包括使线虫害虫与杀虫有效量的多肽接触,所述多肽包含杀线虫毒素,特别是对短体线虫属物种线虫,例如短尾短体线虫有活性的杀线虫毒素。还包括通过表达本发明的毒素来增加植物产量的方法。
Description
相关申请的交叉引用
本申请要求2016年12月22日提交的美国临时申请序列号62/438,420的权益,其内容通过引用整体并入本文。
电子提交的序列表的参考
序列表的官方副本通过EFS-Web作为ASCII格式的序列表以电子方式提交,其文件名为“APA16-6020WOSEQLIST.txt”,创建于2017年10月3日,大小为40千字节,并与说明书同时提交。该ASCII格式文档中包含的序列表是说明书的一部分,并且其全部内容通过引用并入本文。
发明领域
本发明涉及分子生物学领域。提供了使用Cry14控制线虫害虫的方法。
背景技术
线虫是活跃的、柔韧的、细长的生物,生活在潮湿表面或液体环境,包括土壤中的水膜和其他生物体内的潮湿组织中。许多线虫物种已经进化成非常成功的植物和动物寄生虫,并且造成农业和牲畜的显著经济损失以及人类的发病率和死亡率(Whitehead(1998)Plant Nematode Control.CAB International,New York)。
据估计,基于估计所有主要作物12%的年平均损失,寄生线虫在园艺和农业产业每年在全世界花费超过780亿美元。例如,估计线虫每年导致全球大豆损失约32亿美元(Barker等人(1994)Plant and Soil Nematodes:Societal Impact and Focus for theFuture.The Committee on National Needs and Priorities inNematology.Cooperative State Research Service,US Department of Agricultureand Society of Nematologists)。在高种植面积的作物市场中,大豆和棉花中的线虫损害最大。然而,还有数十种其他作物遭受严重的线虫侵袭,包括马铃薯、胡椒、洋葱、柑橘、咖啡、甘蔗、温室观赏植物和高尔夫球场草坪草。
已知线虫影响作物和植物的产量、生长和健康。由幼虫和/或成虫线虫引起的寄主植物根部的生理变化可导致虫瘿的形成,这导致植物根部的维管***的破坏。根伸长可以完全停止,并且可以引起由减少的根系提供的水和营养物的供应不足,导致叶子萎黄和/或枯萎,以及生长发育迟缓,其中任何一种都可以导致低产量或死亡。此外,线虫可引起生理效应,导致植物根对细菌和/或真菌侵袭的易感性增加,包括植物本来能抵抗的细菌和/或真菌。这种攻击可能导致广泛的二次腐变和腐烂。
根病变线虫短尾短体线虫(Pratylenchus brachyurus)已成为越来越重要的大豆病原体。它具有广泛的寄主范围,广泛分布于热带和亚热带地区,特别是在巴西、非洲和美国南部。短尾短体线虫已成为巴西Cerrado地区棉花和大豆种植者的一个担忧,被认为是该地区大豆的主要线虫病原体。在大豆中,这种线虫可以将产量降低30%至50%,在沙质土壤上观察到更大的损害。目前还没有发现耐短尾短体线虫的大豆品种。虽然已经研究了几种大豆基因型用于对短尾短体线虫的抗性,并且鉴定了一些耐受性增加的品种,但由于这种线虫是多食性的并且缺乏与其宿主的密切相互作用,因此育种抗短尾短体线虫的抗性品种很难(Machado(2014)Current Agricultural Science and Technology 20:26-35;Antonio等人,(2012)Soil productivity losses in area infested by the nematoidof the root lesions in Vera,MT.在:Brazilian Congress of Soy,6,2012,Cuiabá.Abstracts.Londrina:Embrapa Soja,4pp;Rios等人,(2016)Ciência Rural 46:580–584;Lima等人,2017,在:Soybean-The Basis of Yield,Biomass and Productivity书中,第6章;Minobu Kasai编,ISBN 978-953-51-3118-2,Print ISBN 978-953-51-3117-5,InTech;Inomoto等人,(2011)de culturas sob/>central para controle defitonematoides:/>populacional,patogenicidade e estimativa deperdas.Tropical Plant Pathology 36:178-185))。
已经以几种形式提供了控制线虫感染的方法。在许多情况下已尝试过生物和培养控制方法,包括植物检疫。在某些商业栽培品种(例如大豆)中可获得对某些线虫的遗传抗性,但这些品种数量受到限制,并且具有所需农艺特征和抗性的栽培品种的可得性受到限制。此外,基于通过有性杂交的遗传重组的常规植物育种来生产抗线虫的商业品种是缓慢的过程并且经常由于缺乏适当的种质而进一步受到阻碍。
控制植物寄生线虫的化学方法对许多缺乏足够天然抗性的作物仍然是必不可少的。然而,化学试剂通常不具有选择性,并且一些化学试剂在施用药剂后的一段时间内对非靶标生物体发挥作用,有效地破坏有益微生物群体。化学试剂可能在环境中持续存在并且只是缓慢代谢。
因此,需要用于控制农业重要植物中的线虫种群的其他手段。
发明内容
提供了赋予植物、植物细胞、组织和种子杀线虫活性的组合物和方法。特别地,提供了用于杀死或控制线虫害虫种群的方法,特别是病变线虫,例如短体线虫属物种(Pratylenchus sp),例如短尾短体线虫种群。本发明进一步提供对根结线虫(Meloidogynespp大豆害虫线虫,包括但不限于南方根结线虫(Meloidogyne incognita),花生根结线虫(Meloidogyne arenaria),北方根结线虫(Meloidogyne hapla),或爪哇根结线虫(Meloidogyne javanica),或其任何组合),肾形线虫(Rotylenchulus reniformis)和Lance线虫(纽带线虫(Hoplolaimus spp),例如哥伦比亚纽带线虫(H.columbus),H.galeatus和大针纽带线虫(H.magnistylus))的控制。该方法包括使线虫害虫与杀虫有效量的多肽接触,所述多肽包含杀线虫毒素,特别是对短体线虫,例如短尾短体线虫,根结线虫,肾状线虫或Lance线虫有活性的杀线虫毒素。在各种实施方案中,杀线虫毒素包含SEQID NO:1或2的氨基酸序列,或其杀虫有效变体或片段。在一些实施方案中,用于保护植物或其细胞免受线虫害虫种群,特别是短体线虫,例如短尾短体线虫,根结线虫,肾状线虫或Lance线虫的方法,包括在植物或细胞中表达编码SEQ ID NO:1或2的核酸,或其变体或片段,其中所述核酸与能够指导核酸在植物细胞中表达的启动子有效连接。
还包括用于增加植物产量的方法,包括在田地生长植物或其种子,所述植物或其种子在其基因组中稳定掺入DNA构建体,所述DNA构建体包含与能够指导核酸在植物细胞中表达的启动子有效连接的核酸,其中核酸编码SEQ ID NO:1或2,或其杀虫有效变体或片段。
本发明的组合物和方法可用于生产具有增强的线虫,例如短体线虫,根结线虫,肾状线虫,或Lance线虫抗性或耐受性的生物。这些生物和包含该生物的组合物是农业目的所需的。
附图简述
图1.美国的短体线虫抗性温室测定
具有EE-GM4的良种大豆植物在美国温室测定中控制短尾短体线虫。将表达SEQ IDNO:2(“EE-GM4”)的大豆植物与其他良种大豆品系进行比较:一个SCN易感成熟组(MG)3品系(“THORNE”),一个MG3 SCN易感品系,一个MG 6.2 SCN易感品系和一个MG9 SCN易感品系(“Susc WT”显示这三个品系的平均值),一个MG3 SCN抗性品系(具有来自PI88788的rhg1抗性等位基因,“SCN Res(PI88788)”)和一个来自北京的具有rhg1和Rhg4 SCN抗性的MG 6.2SCN抗性品系(“SCN Res(Peking)”)。绘出了侵袭后30天根中短体线虫的平均数量(每个条目5株植物),也显示在品种间观察到的变异(如通常在温室测定中所见)。结果显示,对于EE-GM4品系,对短体线虫的控制率为~85%。具有天然SCN抗性的大豆品系(来自Peking或PI88788)不能控制短尾短体线虫。
图2.巴西的短体线虫抗性温室分析
具有EE-GM4(“EE-GM4”)的大豆植物显著减少大豆根中的短尾短体线虫。短尾短体线虫从巴西的当地田地中分离出来。EE-GM4植物(两个不同的美国良种品系(都是成熟组6.2,一个SCN易感和一个具有Peking SCN抗性(“EE-GM4”))和五个巴西大豆品系,具有有限的短体线虫控制(“巴西品系”),一个巴西品系,标记为短体线虫的低Rf(繁殖因子)(“BRS7380(低Rf)”),一个美国良种品系(成熟组6.2),其是SCN易感(“SCN Susc”)和一个具有Peking SCN抗性的MG 6.2的美国良种品系(“SCN Res(Peking)”)在巴西的温室试验中进行了短体线虫控制的评价。绘出了这些条目的平均值,也显示了各品种间观察到的变异(如通常在温室测定中所见)。一个巴西大豆品系(BRS 7380)显示出短体线虫减少约89%。EE-GM4品系对短体线虫的控制率为~79%。携带针对SCN的Peking天然抗性的大豆品系不能控制短尾短体线虫。
详述
本发明涉及调节生物体,特别是植物或植物细胞中的线虫抗性的方法。“抗性”是指线虫在摄入本发明的多肽时被杀死或与本发明的多肽接触时线虫的运动、摄食、繁殖或其他功能受损。控制植物或其种子中的植物寄生线虫种群将改善结瘤、发芽、根发育、出苗和健康,包括对疾病(包括细菌或真菌疾病)的抵抗或保护免于疾病,这是本文所公开和描述的方法的重要益处。因此,如本文所述的方法可用于控制线虫种群,特别是短体线虫种群,例如短尾短体线虫,根结线虫,肾状线虫或Lance线虫,其提供植物和/或种子的改善的总体植物健康、营养和/或改善的农艺益处。与线虫种群控制有关的任何益处,例如,线虫总数/面积的减少,线虫卵/面积的减少,或对植物的损害的减少,都可以是本发明的农艺益处。控制线虫种群的次要益处包括但不限于改善的根发育(例如,改善的根或根毛生长),提高的产量,更快的出苗,改善的植物胁迫管理,包括增加的胁迫耐受性和/或改善的应力恢复,增加的机械强度,改善的抗旱性,减少真菌疾病感染和改善植物健康。还可以获得任何这些益处的组合。
本发明的方法包括生物体转化或使用包含编码本发明杀线虫蛋白质的异源核苷酸序列的生物体。本文描述的方法可用于控制或杀死线虫害虫种群和用于产生具有针对线虫害虫的杀线虫活性的组合物。
“杀虫毒素”或“杀虫蛋白”或“杀线虫活性”或“杀线虫毒素”是指对一种或多种线虫害虫具有活性的毒素,包括但不限于短体线虫,包括艾氏短体线虫(Pratylenchusalleni),短尾短体线虫,咖啡短体线虫(Pratylenchus coffeae),刻痕短体线虫(Pratylenchus crenatus),Pratylenchus dulscus,伪根结线虫(Pratylenchus fallax),Pratylenchus flakkensis,古氏短体线虫(Pratylenchus goodeyi),六裂短体线虫(Pratylenchus hexincisus),卢斯短体线虫(Pratylenchus loosi),Pratylenchusminutus,Pratylenchus mulchandi,Pratylenchus musicola,落选短体线虫(Pratylenchus neglectus),穿刺短体线虫(Pratylenchus penetrans),伤残短体线虫(Pratylenchus pratensis),Pratylenchus reniformia,斯氏短体线虫(Pratylenchusscribneri),索氏短体线虫(Pratylenchus thornei),Pratylenchus vulnus和玉米短体线虫(Pratylenchus zeae)。杀线虫蛋白质包括从本文公开的全长核苷酸序列推导的氨基酸序列,以及由于使用替代的下游起始位点,或者由于产生具有杀线虫活性的较短蛋白质的加工而比全长序列短的氨基酸序列。加工可以在蛋白质表达的生物体中发生,或者在摄入蛋白质后在害虫中发生。
在具体的实施方案中,杀线虫蛋白包含Cry14蛋白。在各种实施方案中,Cry14蛋白是Cry14Aa1(GENBANK登录号AAA21516)或Cry14Ab1(GENBANK登录号KC156652)。在一些实施方案中,Cry14Aa1蛋白包括SEQ ID NO:1中所示的氨基酸序列,以及其变体和片段。在其他实施方案中,Cry14Ab1蛋白包括SEQ ID NO:2中所示的氨基酸序列,以及其变体和片段。编码SEQ ID NO:1的示例性核苷酸序列示于SEQ ID NO:3和5中。编码SEQ ID NO:2的示例性核苷酸序列示于SEQ ID NO:4和6中。
因此,本文提供了杀死或控制线虫害虫种群的方法,例如,一种短体线虫种群,例如,短尾短体线虫,根结线虫,肾状线虫或Lance线虫,包括使线虫害虫接触,或将线虫害虫暴露于包含本发明杀线虫毒素的组合物。在具体的实施方案中,杀线虫蛋白质包含SEQ IDNO:1或2中所示的Cry14蛋白质,以及其变体和片段。
分离的核酸分子,以及它们的变体和片段
本发明的一个方面涉及分离的、重组的或嵌合的核酸分子,其包含编码杀线虫蛋白和多肽或其生物活性部分的核苷酸序列,以及足以用作杂交探针以鉴定编码具有序列同源性区域的蛋白质的核酸分子的核酸分子。本文还包括能够在本文其他地方定义的严格条件下与本发明的核苷酸序列杂交的核苷酸序列。如本文所用,术语“核酸分子”旨在包括DNA分子(例如重组DNA、cDNA或基因组DNA)和RNA分子(例如mRNA)和使用核苷酸类似物产生的DNA或RNA的类似物。核酸分子可以是单链或双链的,但优选是双链DNA。术语“重组”包括已经相对于天然多核苷酸或多肽操作的多核苷酸或多肽,使得多核苷酸或多肽与天然存在的不同(例如,化学组成或结构)。如本文所用,“分离的核酸(序列/分子)”或“分离的DNA(序列/分子)”是指不再处于从其分离的天然环境中的核酸或DNA(序列/分子)。例如,另一种细菌宿主或植物基因组中的核酸序列,或与来自另一来源的DNA或核酸(序列/分子)融合的核酸或DNA(序列/分子),例如当包含在嵌合基因中在(异源)植物可表达的启动子的控制下时。本发明的任何核酸或DNA,包括任何引物,也可以是非天然存在的,例如这样的核酸或DNA,其序列与天然存在的序列相同,但具有标记(天然发生的对应物所缺少的),或与天然存在的核苷酸相比具有至少一个核苷酸添加或置换或至少一个内部核苷酸缺失的序列,或与天然存在的核酸或DNA或其片段具有低于100%(不相同)同一性的序列的序列,或具有由不同来源的核苷酸序列组成的序列的核酸或DNA,所述核苷酸序列在自然界中不一起出现(嵌合或杂合DNA),或具有不同于天然核酸或DNA或其片段的序列的人造合成的核酸或DNA。
本文使用的分离的、重组的或嵌合的核酸(或DNA)是指不再在其天然环境中的核酸(或DNA),例如在体外或在重组细菌或植物宿主细胞中。在一些实施方案中,分离的、重组的或嵌合的核酸不含衍生该核酸的生物的基因组DNA中核酸天然侧翼的序列(优选蛋白质编码序列)(即位于核酸的5'和3'末端的序列)。出于本发明的目的,当用于指核酸分子时,“分离的”不包括分离的染色体。例如,在各种实施方案中,分离的δ-内毒素编码核酸分子可含有少于约5kb,4kb,3kb,2kb,1kb,0.5kb或0.1kb的天然位于衍生该核酸的细胞的基因组DNA中的核酸分子侧翼的核苷酸序列。在各种实施方案中,基本上不含细胞物质的δ-内毒素蛋白包括具有少于约30%,20%,10%或5%(干重)的非δ-内毒素蛋白的蛋白质制剂(也是在本文中称为“污染蛋白质”)。在一些实施方案中,本发明的重组核酸包含相对于SEQ IDNO:3-6的一个或多个核苷酸取代,或其变体或片段。
编码本发明蛋白质的核苷酸序列包括SEQ ID NO:3-6所示的序列,及其变体,片段和互补序列。“互补序列”是指与给定核苷酸序列充分互补,使得它可以与给定的核苷酸序列杂交,从而形成稳定的双链体的核苷酸序列。由这些核苷酸序列编码的杀线虫蛋白质的相应氨基酸序列示于SEQ ID NO:1和2中。
作为编码杀线虫蛋白的这些核苷酸序列的片段的核酸分子也包括在本发明中。“片段”是指编码杀线虫蛋白的核苷酸序列的一部分。核苷酸序列的片段可以编码杀线虫蛋白的生物活性部分,或者它可以是可以使用下文公开的方法用作杂交探针或PCR引物的片段。根据预期用途,作为编码杀线虫蛋白的核苷酸序列的片段的核酸分子包含至少约50、100、200、300、400、500、600、700、800、900、1000、1100、1200、1300、1350、1400个连续核苷酸或直至编码本文公开的杀线虫蛋白的全长核苷酸序列中存在的核苷酸数目。“连续的”核苷酸是指彼此紧邻的核苷酸残基。本发明的核苷酸序列的片段将编码保留杀线虫蛋白的生物活性的蛋白质片段,并因此保留针对线虫害虫的杀虫活性。因此,还涵盖了本文公开的多肽的生物学活性片段。“保留活性”意指片段具有杀线虫蛋白的杀虫活性的至少约30%,至少约50%,至少约70%,80%,90%,95%或更高。测量杀线虫活性的方法是本领域熟知的,并且也在本文中描述。
编码本发明蛋白质的生物活性部分的杀线虫蛋白质的核苷酸序列的片段将编码至少约15、25、30、50、75、100、125、150、175、200、250、300,350、400、450、500、550、600、650、700、750、800、850、900、950、1000、1050、1100或1150个连续氨基酸,或高达本发明的全长杀线虫蛋白质的氨基酸总数。在一些实施方案中,片段是蛋白水解切割片段。例如,相对于SEQID NO:1或2,蛋白水解切割片段可具有至少约30个氨基酸,至少约40个氨基酸,至少约50个,至少约100个氨基酸,约120,约130,约140,约150或约160个氨基酸的N末端或C末端截短。在一些实施方案中,本文包括的片段来自C末端结晶结构域的去除,例如通过蛋白水解,或通过在编码序列中***终止密码子。在进一步的实施方案中,本文包括的片段包含N-末端截短,并且N-末端截短可以在截短的N-末端包含甲硫氨酸残基。
在各种实施方案中,本发明的核酸包含SEQ ID NO:3-6的简并核酸,其中所述简并核苷酸序列编码与SEQ ID NO:1或2相同的氨基酸序列。
本发明优选的杀线虫蛋白质由与SEQ ID NO:3-6的核苷酸序列充分相同的核苷酸序列编码,或者杀线虫蛋白质与SEQ ID NO:1或2中所示的氨基酸序列充分相同。“充分相同”是指使用本文所述的比对程序之一使用标准参数与参考序列相比,具有至少约60%或65%序列同一性,约70%或75%序列同一性,约80%或85%序列同一性,约90%,91%,92%,93%,94%,95%,96%,97%,98%,99%或更高的序列同一性的氨基酸或核苷酸序列。本领域技术人员将认识到,通过考虑密码子简并性、氨基酸相似性、读码框定位等,可以适当调整这些值以确定由两个核苷酸序列编码的蛋白质的相应同一性。
为了确定两个氨基酸序列或两个核酸的百分比同一性,比对序列以达到最佳比较目的。两个序列之间的百分比同一性是序列共有的相同位置数的函数(即,同一性百分比=相同位置的数目/位置总数(例如,重叠位置)×100)。在一个实施方案中,两个序列长度相同。在另一个实施方案中,在整个参考序列(即,本文公开的序列,如SEQ ID NO:1-6中的任一个)上计算同一性百分比。两个序列之间的同一性百分比可以使用与下面描述的那些类似的技术来确定,允许或不允许空位。在计算百分比同一性时,通常计算完全匹配。空位,即在比对中在一个序列中存在残基而在另一个序列中不存在的位置被认为是具有不相同残基的位置。
可以使用数学算法来确定两个序列之间的百分比同一性。用于比较两个序列的数学算法的非限制性实例是Karlin和Altschul(1990)Proc.Natl.Acad.Sci.USA87:2264的算法,如在Karlin and Altschul(1993)Proc.Natl.Acad.Sci.USA 90:5873-5877中修改。这种算法被并入Altschul等人(1990)J.Mol.Biol.215:403的BLASTN和BLASTX程序中。BLAST核苷酸搜索可以用BLASTN程序进行,得分=100,字长=12,以获得与本发明的杀虫样核酸分子同源的核苷酸序列。BLAST蛋白质搜索可以用BLASTX程序进行,得分=50,字长=3,以获得与本发明的杀线虫蛋白质分子同源的氨基酸序列。为了获得用于比较目的的空位比对,可以如Altschul等人(1997)Nucleic Acids Res.25:3389所述利用Gapped BLAST(BLAST 2.0中)。或者,PSI-Blast可用于执行迭代搜索,以检测分子之间的距离关系。见Altschul等人,(1997)同上。当使用BLAST,Gapped BLAST和PSI-Blast程序时,可以使用各个程序(例如BLASTX和BLASTN)的默认参数。也可以通过检查手动进行比对。
用于比较序列的数学算法的另一个非限制性实例是ClustalW算法(Higgins等人(1994)Nucleic Acids Res.22:4673-4680)。ClustalW比较序列并比对整个氨基酸或DNA序列,因此可以提供关于整个氨基酸序列的序列保守性的数据。ClustalW算法用于几种商业上可获得的DNA/氨基酸分析软件包,例如Vector NTI Program Suite(InvitrogenCorporation,Carlsbad,CA)的ALIGNX模块。在用ClustalW比对氨基酸序列后,可以评估氨基酸同一性百分比。用于分析ClustalW比对的软件程序的非限制性实例是GENEDOCTM。(Karl Nicholas)允许评估多种蛋白质之间的氨基酸(或DNA)相似性和同一性。用于比较序列的数学算法的另一个非限制性实例是Myers和Miller(1988)CABIOS 4:11-17的算法。这种算法被并入ALIGN程序(版本2.0),该程序是GCG Wisconsin GeneticsSoftware Package,Version 10(可从Accelrys,Inc.,9685 Scranton Rd.,San Diego,CA,USA获得)的一部分。当利用ALIGN程序比较氨基酸序列时,可以使用PAM120权重残基表,空位长度罚分为12,空位罚分为4。
除非另有说明,否则GAP版本10(其使用Needleman和Wunsch(1970)J.Mol.Biol.48(3):443-453的算法)将使用以下参数用于确定序列同一性或相似性:使用GAP权重50和长度权重3,以及nwsgapdna.cmp评分矩阵确定核苷酸序列的%同一性和%相似性;使用8的GAP重量和2的长度重量以及BLOSUM62评分程序确定氨基酸序列的%同一性或%相似性。也可以使用等同程序。“等同程序”是指任何序列比较程序,对于所讨论的任何两个序列,当与GAP第10版产生的相应比对相比时,产生具有相同核苷酸残基匹配和相同百分比序列同一性的比对。
本发明还包括变体核酸分子。编码杀线虫蛋白质的核苷酸序列的“变体”包括编码本文公开的杀线虫蛋白但由于遗传密码的简并性以及如上所述充分相同的那些而保守地不同的那些序列。可以使用众所周知的分子生物学技术鉴定天然存在的等位基因变体,例如聚合酶链反应(PCR)和如下所述的杂交技术。变体核苷酸序列还包括合成衍生的核苷酸序列,其例如通过使用定点诱变产生但仍编码如下所述的本发明中公开的杀线虫蛋白。本发明包括的变体蛋白质具有生物学活性,即它们继续具有天然蛋白质的所需生物活性,即对线虫害虫的杀虫活性。“保留活性”意指变体将具有天然蛋白质的杀虫活性的至少约30%,至少约50%,至少约70%,或至少约80%。测量针对线虫害虫的杀虫活性的方法是本领域熟知的并且在本文其他地方描述。
技术人员将进一步理解,可以通过突变本发明的核苷酸序列引入变化,从而导致编码的杀线虫蛋白的氨基酸序列的变化,而不改变蛋白质的生物活性。因此,可以通过将一个或多个核苷酸取代、添加或缺失引入本文公开的相应核苷酸序列中来产生变体分离的核酸分子,使得将一个或多个氨基酸取代、添加或缺失引入编码的蛋白质中。可以通过标准技术引入突变,例如定点诱变和PCR介导的诱变。这些变体核苷酸序列也包括在本发明中。
例如,可以在一个或多个预测的非必需氨基酸残基处进行保守氨基酸取代。“非必需”氨基酸残基是可以从杀线虫蛋白的野生型序列改变而不改变生物活性的残基,而生物活性需要“必需”氨基酸残基。“保守氨基酸取代”是其中氨基酸残基被具有相似侧链的氨基酸残基取代的氨基酸取代。本领域已经定义了具有相似侧链的氨基酸残基家族。这些家族包括具有碱性侧链的氨基酸(例如,赖氨酸,精氨酸,组氨酸),酸性侧链(例如天冬氨酸,谷氨酸),不带电的极性侧链(例如甘氨酸,天冬酰胺,谷氨酰胺,丝氨酸,苏氨酸,酪氨酸,半胱氨酸),非极性侧链(如丙氨酸,缬氨酸,亮氨酸,异亮氨酸,脯氨酸,苯丙氨酸,蛋氨酸,色氨酸),β-支链侧链(如苏氨酸,缬氨酸,异亮氨酸)和芳香侧链的氨基酸(如酪氨酸,苯丙氨酸,色氨酸,组氨酸)。
δ-内毒素通常具有五个保守序列结构域和三个保守结构域(参见,例如,de Maagd等人(2001)Trends Genetics 17:193-199)。第一个保守结构域由7个α螺旋组成,参与膜***和孔形成。结构域II由以希腊键构型排列的三个β-折叠组成,结构域III由“果冻卷”形成的两个反平行β-折叠组成(de Maagd等人,2001,同上)。结构域II和III参与受体识别和结合,因此被认为是毒素特异性的决定因素。
氨基酸取代可以在保留功能的非保守区域中进行。通常,对于保守氨基酸残基或存在于保守基序内的氨基酸残基不会进行这种取代,其中这些残基对蛋白质活性是必需的。保守并且可能对蛋白质活性必需的残基的实例包括,例如,在相似或相关毒素与本发明的序列的比对中包含的所有蛋白质之间相同的残基(例如,在同源蛋白质的比对中相同的残基)。保守但可以允许保守氨基酸取代并且仍然保留活性的残基的实例包括,例如,在相似或相关毒素与本发明的序列的比对中包含的所有蛋白质之间仅具有保守取代的残基(例如,在比对同源蛋白质中包含的所有蛋白质之间仅具有保守取代的残基)。然而,本领域技术人员将理解,功能变体可以在保守残基中具有较小的保守或非保守改变。
或者,可以通过沿着全部或部分编码序列随机引入突变,例如通过饱和诱变来制备变体核苷酸序列,并且可以筛选所得突变体赋予针对线虫害虫的杀虫活性以鉴定保留活性的突变体的能力。诱变后,可以重组表达编码的蛋白质,并且可以使用标准测定技术测定蛋白质的活性。
可以使用诸如PCR、杂交等方法鉴定相应的杀虫序列,这些序列与本发明的序列具有实质同一性(例如,在整个参考序列上具有至少约70%,至少约75%,80%,85%,90%,95%或更多的序列同一性并且具有或赋予针对线虫害虫的杀虫活性。参见,例如,Sambrook和Russell(2001)Molecular Cloning:A Laboratory Manual。(Cold Spring HarborLaboratory Press,Cold Spring Harbor,NY)和Innis等人,(1990)PCR Protocols:AGuideto Methods and Applications(Academic Press,NY)。
在杂交方法中,全部或部分杀虫核苷酸序列可用于筛选cDNA或基因组文库。构建此类cDNA和基因组文库的方法通常是本领域已知的,并且在Sambrook和Russell,2001,同上中公开。所谓的杂交探针可以是基因组DNA片段,cDNA片段,RNA片段或其他寡核苷酸,并且可以用可检测基团如32P或任何其他可检测标记物标记,例如其他放射性同位素、荧光化合物、酶或酶辅因子。用于杂交的探针可以通过基于本文公开的已知杀线虫蛋白编码核苷酸序列标记合成寡核苷酸来进行。还可以使用基于核苷酸序列或编码的氨基酸序列中的保守核苷酸或氨基酸残基设计的简并引物。探针通常包含在严格条件下与编码本发明的杀线虫蛋白质或其片段或变体的核苷酸序列的至少约12,至少约25,至少约50,75,100,125,150,175或200个连续核苷酸杂交的核苷酸序列区域。制备用于杂交的探针的方法通常是本领域已知的,并且公开在Sambrook和Russe l l,2001,同上,在此引入作为参考。
例如,本文公开的整个杀线虫序列或其一个或多个部分可用作能够与相应的杀线虫蛋白样序列和信使RNA特异性杂交的探针。为了在各种条件下实现特异性杂交,此类探针包括独特的序列,优选长度至少约10个核苷酸,或长度至少约20个核苷酸。此类探针可用于通过PCR扩增来自所选生物或样品的相应杀虫序列。该技术可用于从所需生物体中分离另外的编码序列或用作诊断测定法以确定生物体中编码序列的存在。杂交技术包括杂交筛选铺板DNA文库(噬斑或菌落;参见,例如,Sambrook等人(1989)Molecular Cloning:ALaboratory Manual(第2版,Cold Spring Harbor Laboratory Press,Cold SpringHarbor,New York)。
因此,本发明包括用于杂交的探针,以及能够与本发明的全部或部分核苷酸序列杂交的核苷酸序列(例如,至少约300个核苷酸,至少约400,至少约400,450,500,1000,1200,1500,2000,2500,3000,3500,或高达本文公开的核苷酸序列的全长)。这些序列的杂交可以在严格条件下进行。“严格条件”或“严格杂交条件”是指探针与其靶序列杂交的可检测程度大于其他序列(例如,至少比背景高2倍)的条件。严格条件是序列依赖性的,并且在不同情况下会有所不同。通过控制杂交和/或洗涤条件的严格性,可以鉴定与探针100%互补的靶序列(同源探测)。或者,可以调节严格条件以允许序列中的一些错配,从而检测较低程度的相似性(异源探测)。通常,探针长度小于约1000个核苷酸,优选长度小于500个核苷酸。
通常,严格条件是盐浓度小于约1.5M Na离子,通常约0.01-1.0M Na离子浓度(或其它盐),pH7.0至8.3,并且温度对于短探针(例如,10至50个核苷酸)至少约30℃和对于长探针(例如,大于50个核苷酸)至少约60℃的那些条件。加入去稳定剂如甲酰胺也可以达到严格的条件。示例性的低严格条件包括在37℃下与30至35%甲酰胺,1M NaCl,1%SDS(十二烷基硫酸钠)的缓冲溶液杂交,并在1X至2X SSC中(20X SSC=3.0M NaCl/0.3M柠檬酸三钠)在50至55℃下洗涤。示例性的中等严格条件包括在37℃下在40至45%甲酰胺,1.0M NaCl,1%SDS中杂交,以及在55至60℃下在0.5X至1X SSC中洗涤。示例性的高严格条件包括在50%甲酰胺,1M NaCl,1%SDS中于37℃杂交,以及在0.1X SSC中在60至65℃下洗涤。任选地,洗涤缓冲液可包含约0.1%至约1%的SDS。杂交持续时间通常小于约24小时,通常约4至约12小时。
特异性通常是杂交后洗涤的函数,关键因素是最终洗涤溶液的离子强度和温度。对于DNA-DNA杂交体,Tm可以从Meinkoth和Wahl(1984)Anal.Biochem.138:267-284的等式近似:Tm=81.5℃+16.6(log M)+0.41(%GC)-0.61(%形式)-500/L;其中M是单价阳离子的摩尔浓度,%GC是DNA中鸟苷和胞嘧啶核苷酸的百分比,%形式是杂交溶液中甲酰胺的百分比,L是碱基对中杂合体的长度。Tm是50%的互补靶序列与完全匹配的探针杂交的温度(在确定的离子强度和pH下)。每1%的不匹配,Tm降低约1℃;因此,可以调节Tm、杂交和/或洗涤条件以与具有所需同一性的序列杂交。例如,如果寻找具有≥90%同一性的序列,则Tm可以降低10℃。通常,严格条件选择为在确定的离子强度和pH下比特定序列及其互补体的热解链温度(Tm)低约5℃。然而,严格的严格条件可以利用在低于热解链温度(Tm)1,2,3或4℃下杂交和/或洗涤;中等严格条件可以利用在比热解链温度(Tm)低6,7,8,9或10℃下杂交和/或洗涤;低严格条件可以利用在比热解链温度(Tm)低11,12,13,14,15或20℃下杂交和/或洗涤。使用该等式、杂交和洗涤组合物以及所需的Tm,普通技术人员将理解,固有地描述了杂交和/或洗涤溶液严格性的变化。如果所需的错配程度导致Tm低于45℃(水溶液)或32℃(甲酰胺溶液),则优选增加SSC浓度以便可以使用更高的温度。核酸杂交的广泛指南见于Tijssen(1993)Laboratory Techniques in Biochemistry and Molecular Biology-Hybridization with Nucleic Acid Probes,Part I,Chapter 2(Elsevier,New York),和Ausubel等编辑,(1995)Current Protocols in Molecular Biology,第2章(GreenePublishing and Wiley-Interscience,New York)。见Sambrook等人,(1989)MolecularCloning:ALaboratory Manual(第2版,Cold Spring Harbor Laboratory Press,ColdSpring Harbor,New York)。
分离的蛋白质及其变体和片段
杀线虫蛋白质也包括在本发明内。“杀线虫蛋白质”是指具有SEQ ID NO:1或2所示氨基酸序列的蛋白质。还提供了片段、生物活性部分及其变体,并且可用于实施本发明的方法。“分离的蛋白质”或“重组蛋白质”用于指不再处于其天然环境,例如体外或重组细菌或植物宿主细胞中的蛋白质。在一些实施方案中,重组蛋白质是SEQ ID NO:1或2的变体,其中变体相对于SEQ ID NO:1或2包含至少一个氨基酸取代、缺失或***。
“片段”或“生物学活性部分”包括多肽片段,其包含与SEQ ID NO:1或2中所示的氨基酸序列充分相同的氨基酸序列,并且显示出针对线虫害虫的杀虫活性。杀线虫蛋白的生物活性部分可以是多肽,例如,长度为10,25,50,100,150,200,250,300,350,400,450,500,550,600,650,700,750,800,850,900,950,1000,1050,1100,1150,1200,1250,1300,1350或更多氨基酸。这些生物活性部分可以通过重组技术制备并评估针对线虫害虫的杀虫活性。测量针对线虫害虫的杀虫活性的方法是本领域熟知的(参见,例如,美国专利申请公开号US20160066584)并在本文其他地方描述。如本文所用,片段包含SEQ ID NO:1或2的至少8个连续氨基酸。然而,本发明包括其他片段,例如蛋白质中任何长度大于约10,20,30,50,100,150,200,250,300,350,400,450,500,550,600,650,700,750,800,850,900,950,1000,1050,1100,1150,1200,1250,1300,1350或更多个氨基酸的片段。
“变体”是指与SEQIDNO:1或2中任一个的氨基酸序列具有至少约60%,65%,约70%,75%,约80%,85%,约90%,91%,92%,93%,94%,95%,96%,97%,98%或99%同一性的氨基酸序列的蛋白质或多肽。变体还包括由在严格条件下,与SEQ ID NO:3-6的核酸分子或其互补序列杂交的核酸分子编码的多肽。变体包括由于诱变而在氨基酸序列上不同的多肽。本发明包括的变体蛋白质具有生物学活性,即它们继续具有天然蛋白质的所需生物活性,即保留针对线虫害虫的杀虫活性。在一些实施方案中,变体相对于天然蛋白质具有改善的活性。测量针对线虫害虫的杀虫活性的方法是本领域熟知的(参见,例如,美国专利申请公开号US 20160066584)并在本文其他地方描述。
细菌基因通常在开放读码框的起点附近具有多个甲硫氨酸起始密码子。通常,在这些起始密码子中的一个或多个处的翻译起始将导致功能性蛋白质的产生。这些起始密码子可包括ATG密码子。但是,芽孢杆菌等细菌还将密码子GTG识别为起始密码子,并且在GTG密码子处起始翻译的蛋白质在第一个氨基酸处含有甲硫氨酸。在极少数情况下,细菌***中的翻译可以在TTG密码子处起始,尽管在这种情况下TTG编码甲硫氨酸。此外,通常不先验地确定这些密码子中的哪一种在细菌中天然使用。因此,应理解,使用一种备选的甲硫氨酸密码子也可导致杀线虫蛋白的产生。这些杀线虫蛋白质包括在本发明中,并可用于本发明的方法中。应当理解,当在植物中表达时,有必要将备选的起始密码子改变为ATG以进行适当的翻译。
在本发明的各种实施方案中,杀线虫蛋白包括从本文公开的全长核苷酸序列推导的氨基酸序列,和由于使用备选的下游起始位点而短于全长序列的氨基酸序列。
还包括针对本发明多肽或其变体或片段的抗体。产生抗体的方法是本领域熟知的(参见,例如,Harlow和Lane(1988)Antibodies:A Laboratory Manual,Cold SpringHarbor Laboratory,Cold Spring Harbor,NY;美国专利No.4,196,265)。
因此,本发明的一个方面涉及特异性结合本发明的一种或多种蛋白质或肽分子及其同源物、融合物或片段的抗体、单链抗原结合分子或其他蛋白质。在特别优选的实施方案中,抗体特异性结合具有SEQ ID NO:1或2所示氨基酸序列的蛋白质或其片段。在另一个实施方案中,抗体特异性结合融合蛋白,所述融合蛋白包含选自SEQ ID NO:1或2所示氨基酸序列的氨基酸序列或其片段。在各种实施方案中,特异性结合本发明蛋白质或包含本发明蛋白质的融合蛋白质的抗体是非天然存在的抗体。
本发明的抗体可用于定量或定性检测本发明的蛋白质或肽分子,或用于检测蛋白质的翻译后修饰。如本文所用,如果这种结合不被非相关分子的存在竞争性抑制,则称抗体或肽“特异性结合”本发明的蛋白质或肽分子。
本发明的抗体可以包含在可用于检测本发明的蛋白质或肽分子的试剂盒中。本发明进一步包括检测本发明的蛋白质或肽分子(特别是由SEQ ID NO:1或2中所示的氨基酸序列编码的蛋白质,包括能够特异性结合本发明抗体的变体或其片段)的方法,包括使样品与本发明的抗体接触并确定样品是否含有本发明的蛋白质或肽分子。利用抗体检测目的蛋白质或肽的方法是本领域已知的。
改变的或改进的变体
已经认识到,杀线虫蛋白质的DNA序列可以通过各种方法改变,并且这些改变可以导致编码具有与本发明的杀线虫蛋白质编码的氨基酸序列不同的氨基酸序列的蛋白质的DNA序列。该蛋白质可以以各种方式改变,包括SEQ ID NO:1或2的一个或多个氨基酸的氨基酸取代、缺失、截短和***,包括多达约2,约3,约4,约5,约6,约7,约8,约9,约10,约15,约20,约25,约30,约35,约40,约45,约50,约55,约60,约65,约70,约75,约80,约85,约90,约100,约105,约110,约115,约120,约125,约130,约135,约140,约145,约150,约155或更多个氨基酸取代、缺失或***。用于这种操作的方法通常是本领域已知的。例如,杀线虫蛋白的氨基酸序列变体可以通过DNA中的突变来制备。这也可以通过几种形式的诱变和/或定向进化中的一种来完成。在一些方面,氨基酸序列中编码的变化基本上不会影响蛋白质的功能。这些变体将具有针对线虫害虫的所需杀虫活性。然而,应理解,通过在本发明的组合物上使用这些技术,可以改善杀线虫蛋白质赋予针对线虫害虫的杀虫活性的能力。例如,可以在宿主细胞中表达杀线虫蛋白,其在DNA复制过程中表现出高的碱基错误掺入率,例如XL-1 Red(Stratagene,La Jolla,CA)。在这些菌株中繁殖后,可以分离DNA(例如通过制备质粒DNA,或通过PCR扩增并将得到的PCR片段克隆到载体中),在非诱变菌株中培养杀线虫蛋白突变,并鉴定具有针对线虫害虫的杀虫活性的突变的基因,例如通过进行测定以测试针对线虫害虫的杀虫活性。此类测定可包括使植物与一种或多种害虫接触并确定植物存活和/或导致害虫死亡的能力。参见,例如,美国专利申请公开No.US 20160066584)。
或者,可以在氨基或羧基末端对许多蛋白质的蛋白质序列进行改变而基本上不影响活性。这可以包括由现代分子方法(例如PCR)引入的***、缺失或改变,包括PCR扩增,其通过在PCR扩增中使用的寡核苷酸中包含氨基酸编码序列来改变或延伸蛋白质编码序列。或者,添加的蛋白质序列可包括完整的蛋白质编码序列,例如本领域常用于产生蛋白质融合的序列。此类融合蛋白通常用于(1)增加目的蛋白质的表达(2)引入结合结构域,酶活性或表位以促进蛋白质纯化、蛋白质检测或本领域已知的其他实验用途(3)靶向蛋白质分泌或翻译成亚细胞细胞器,例如革兰氏阴性细菌的周质空间,或真核细胞的内质网,后者通常导致蛋白质的糖基化。
本发明的变体核苷酸和氨基酸序列还包括衍生自诱变和重组方法如DNA改组的序列。通过这种方法,可以使用一种或多种不同的杀线虫蛋白编码区来产生具有所需特性的新杀线虫蛋白。以这种方式,重组多核苷酸文库由包含序列区的相关序列多核苷酸群产生,所述序列区具有基本的序列同一性并且可以在体外或体内同源重组。例如,使用这种方法,可以在本发明的杀虫基因和其他已知的杀虫基因之间改组编码目的结构域的序列基序,以获得编码具有改善的感兴趣性质(例如增加的杀虫剂活性)的蛋白质的新基因。这种DNA改组的策略在本领域中是已知的。参见,例如,Stemmer(1994)Proc.Natl.Acad.Sci.USA 91:10747-10751;Stemmer(1994)Nature 370:389-391;Crameri等人,(1997)NatureBiotech.15:436-438;Moore等人,(1997)J.Mol.Biol.272:336-347;Zhang等人,(1997)Proc.Natl.Acad.Sci.USA 94:4504-4509;Crameri等人,(1998)Nature 391:288-291;和美国专利号5,605,793和5,837,458。
结构域交换或改组是产生改变的杀线虫蛋白质的另一种机制。可以在杀线虫蛋白质之间交换结构域,导致具有改善的针对线虫害虫或目标谱的杀虫活性的杂交或嵌合毒素。产生重组蛋白质并测试它们针对线虫害虫的杀虫活性的方法是本领域熟知的(参见,例如,Naimov等人(2001)Appl.Environ.Microbiol.67:5328-5330;de Maagd等人,(1996)Appl.Environ.Microbiol.62:1537-1543;Ge等人,(1991)J.Biol.Chem.266:17954-17958;Schnepf等人,(1990)J.Biol.Chem.265:20923-20930;Rang等人,91999)Appl.Environ.Microbiol.65:2918-2925)。
在另一个实施方案中,可以使用易错PCR,寡核苷酸定向诱变,装配PCR,性PCR诱变,体内诱变,盒式诱变,递归整体诱变,指数整体诱变,位点特异性诱变,基因重装配,基因位点饱和诱变,置换诱变,合成连接重装配(SLR),重组,递归序列重组,硫代磷酸酯修饰的DNA诱变,含尿嘧啶的模板诱变,缺口双链体诱变,点错配修复诱变,修复缺陷型宿主菌株诱变,化学诱变,放射性诱变,缺失诱变,限制性选择诱变,限制性纯化诱变,人工基因合成,整体诱变,嵌合核酸多聚体产生等中的一种或多种来获得变体核苷酸和/或氨基酸序列。
载体
本发明的杀虫序列可以在表达盒中提供,用于在感兴趣的宿主细胞例如,植物细胞或微生物中表达。“植物表达盒”是指能够导致从植物细胞中的开放读码框表达蛋白质的DNA构建体。通常这些含有启动子和编码序列。通常,此类构建体还含有3'非翻译区。此类构建体可含有“信号序列”或“前导序列”以促进肽向某些细胞内结构(例如叶绿体(或其他质体),内质网或高尔基体)的共翻译或翻译后转运。
“信号序列”是指已知或怀疑导致跨细胞膜的共翻译或翻译后肽转运的序列。在真核生物中,这通常涉及向高尔基体中分泌,其中一些产生糖基化。细菌的杀虫毒素通常作为原毒素合成,其在靶害虫的肠中被质子激活(Chang(1987)Methods Enzymol.153:507-516)。在本发明的一些实施方案中,信号序列位于天然序列中,或者可以源自本发明的序列。“前导序列”意指任何序列,其在翻译时产生足以触发肽链共翻译转移至亚细胞细胞器的氨基酸序列。因此,这包括通过进入内质网,通向液泡,包括叶绿体的质体,线粒体等靶向转运和/或糖基化的前导序列。因此,本文进一步提供了包含本发明的氨基酸序列的多肽,其与异源前导序列或信号序列有效连接。
“植物转化载体”是指有效转化植物细胞所必需的DNA分子。这种分子可以由一个或多个植物表达盒组成,并且可以组织成一个以上的“载体”DNA分子。例如,二元载体是植物转化载体,其利用两个非连续DNA载体编码用于转化植物细胞的所有必需的顺式和反式作用功能(Hellens和Mullineaux(2000)Trends in Plant Science 5:446-451)。“载体”是指设计用于在不同宿主细胞之间转移的核酸构建体。“表达载体”是指具有在外来细胞中掺入、整合和表达异源DNA序列或片段的能力的载体。盒将包括与本发明的序列有效连接的5'和/或3'调节序列。“有效连接”是指启动子和第二序列之间的功能性连接,其中启动子序列起始并介导对应于第二序列的DNA序列的转录。通常,有效连接的是指连接的核酸序列是连续的,并且在必要时连接两个蛋白质编码区,是连续的并且在相同的读码框中。在一些实施方案中,核苷酸序列与能够指导所述核苷酸序列在宿主细胞(例如微生物宿主细胞或植物宿主细胞)中表达的异源启动子有效连接。盒可以另外含有至少一个另外的基因,以共转化到生物体中。或者,可以在多个表达盒上提供另外的基因。
在各种实施方案中,本发明的核苷酸序列与异源启动子,例如植物启动子有效连接。“启动子”是指发挥指导下游编码序列转录功能的核酸序列。启动子与其他转录和翻译调节核酸序列(也称为“控制序列”)一起是表达目的DNA序列所必需的。
这种表达盒具有多个限制性位点,用于***杀虫序列,使其处于调节区的转录调节之下。
表达盒将包括在5'-3'方向的转录,转录和翻译起始区(即启动子),本发明的DNA序列,以及在植物中功能性的翻译和转录终止区(即终止区)。启动子对植物宿主和/或本发明的DNA序列可以是天然的或类似的,或外来的或异源的。另外,启动子可以是天然序列或可选地是合成序列。当启动子与植物宿主是“天然的”或“同源的”时,意指启动子存在于引入启动子的天然植物中。当启动子对于本发明的DNA序列是“外来的”或“异源的”时,意指启动子不是本发明的有效连接的DNA序列的天然或天然存在的启动子。启动子可以是诱导型或组成型的。它可以是天然存在的,可以由各种天然存在的启动子的部分组成,或者可以是部分或完全合成的。启动子设计的指导由启动子结构的研究提供,例如Harley和Reynolds(1987)Nucleic Acids Res.15:2343-2361。而且,可以优化启动子相对于转录起始的位置。参见,例如,Roberts等人,(1979)Proc.Natl.Acad.Sci.USA,76:760-764。用于植物的许多合适的启动子是本领域熟知的。
例如,用于植物的合适的组成型启动子包括:来自植物病毒的启动子,例如花生褪绿条纹花椰菜病毒(PC1SV)启动子(美国专利号5,850,019);来自花椰菜花叶病毒(CaMV)的35S启动子(Odell等人(1985)Nature313:810-812);Kay等人(1987)Science 236:1299-1302描述的35S启动子;小球藻病毒甲基转移酶基因的启动子(美国专利号5,563,328)和来自玄参花叶病毒(FMV)的全长转录启动子(美国专利号5,378,619);来自水稻肌动蛋白等基因的启动子(McElroy等人(1990)Plant Cell 2:163-171和美国专利5,641,876);遍在蛋白(Christensen等人(1989)Plant Mol.Biol.12:619-632和Christensen等人(1992)PlantMol.Biol.18:675-689)和Grefen等人(2010)Plant J,64:355-365;pEMU(Last等人(1991)Theor.Appl.Genet.81:581-588);MAS(Velten等人(1984)EMBO J.3:2723-2730和美国专利5,510,474);玉米H3组蛋白(Lepetit等人(1992)Mol.Gen.Genet.231:276-285和Atanassova等人(1992)Plant J.2(3):291-300);欧洲油菜ALS3(PCT申请WO97/41228);植物核酮糖-双羧化酶/加氧酶(RuBisCO)小亚基基因;圆环病毒(AU 689 311)或木薯叶脉花叶病毒(CsVMV,US 7,053,205);来自大豆的启动子(在WO/2014/150449中描述的Pbdc6或Pbdc7或在美国专利号7393948和美国专利号8395021中描述的遍在蛋白3启动子);和各种农杆菌基因的启动子(参见美国专利号4,771,002;5,102,796;5,182,200;和5,428,147)。
适用于植物的诱导型启动子包括:来自ACE1***的启动子,其响应铜(Mett等人(1993)PNAS 90:4567-4571);玉米In2基因的启动子,其响应于苯磺酰胺除草剂安全剂(Hershey等人(1991)Mol.Gen.Genetics 227:229-237和Gatz等人(1994)Mol.Gen.Genetics 243:32-38);来自Tn10的Tet阻遏物的启动子(Gatz等人(1991)Mol.Gen.Genet.227:229-237)。用于植物的另一种诱导型启动子是响应植物通常不响应的诱导剂的启动子。这种类型的示例性诱导型启动子是来自类固醇激素基因的诱导型启动子,其转录活性由糖皮质激素激素诱导(Schena等(1991)Proc.Natl.Acad.Sci.USA 88:10421)或最近应用嵌合转录激活因子XVE,用于由***激活的基于***受体的诱导型植物表达***(Zuo等人(2000)Plant J.,24:265-273)。用于植物的其他诱导型启动子描述于EP 332104,PCT WO 93/21334和PCT WO 97/06269中,其通过引用整体并入本文。也可以使用由其他启动子的部分以及部分或全部合成启动子组成的启动子。参见,例如,Ni等人(1995)Plant J.7:661-676和PCT WO 95/14098描述了用于植物的此类启动子。
在本发明的一个实施方案中,对植物的特定区域或组织特异的启动子序列可用于表达本发明的杀线虫蛋白质,例如对种子特异的启动子(Datla,R.等人,1997,Biotechnology Ann.Rev.3,269-296),尤其是napin启动子(EP 255 378 A1),菜豆蛋白启动子,谷蛋白启动子,向日葵蛋白(helianthinin)启动子(WO92/17580),白蛋白启动子(WO98/45460),油质蛋白启动子(WO98/45461),SAT1启动子或SAT3启动子(PCT/US98/06978)。
还可以使用诱导型启动子,其有利地选自苯丙氨酸氨裂解酶(PAL),HMG-CoA还原酶(HMG),几丁质酶,葡聚糖酶,蛋白酶抑制剂(PI),PR1家族基因,胭脂氨酸合酶(nos)和vspB启动子(US 5 670 349,表3),HMG2启动子(US 5 670 349),苹果β-半乳糖苷酶(ABG1)启动子和苹果氨基环丙烷羧酸合酶(ACC合酶)启动子(WO98/45445)。多个启动子可用于本发明的构建体中,包括连续使用。
启动子可以包括或被修饰以包括一个或多个增强子元件。在一些实施方案中,启动子可包括多个增强子元件。与不包含它们的启动子相比,含有增强子元件的启动子提供更高水平的转录。用于植物的合适增强子元件包括PC1SV增强子元件(美国专利号5,850,019),CaMV 35S增强子元件(美国专利号5,106,739和5,164,316)和FMV增强子元件(Maiti等人(1997)Transgenic Res.6:143-156);申请WO87/07644中描述的烟草花叶病毒(TMV)的翻译激活因子,或例如,Carrington&Freed 1990,J.Virol.64:1590-1597中描述的烟草蚀刻病毒(TEV)的翻译激活因子,或内含子,例如玉米的adh1内含子或稻肌动蛋白的内含子1。还参见PCT WO96/23898,WO2012/021794,WO2012/021797,WO2011/084370和WO2011/028914。
通常,此类构建体可含有5'和3'非翻译区。此类构建体可含有“信号序列”或“前导序列”以促进目的肽的共翻译或翻译后转运至某些细胞内结构,例如叶绿体(或其他质体),内质网或高尔基体,或被分泌。例如,可以将构建体改造成含有信号肽以促进肽向内质网的转移。“信号序列”是指已知或怀疑导致跨细胞膜的共翻译或翻译后肽转运的序列。在真核生物中,这通常涉及向高尔基体中的分泌,其中一些产生糖基化。“前导序列”意指任何序列,当翻译时,其产生足以触发肽链共转移至亚细胞细胞器的氨基酸序列。因此,这包括通过进入内质网,通向液泡,包括叶绿体的质体,线粒体等靶向转运和/或糖基化的前导序列。也可以优选将植物表达盒改造成含有内含子,使得内含子的mRNA加工是表达所必需的。
“3'非翻译区”是指位于编码序列下游的多核苷酸。聚腺苷酸化信号序列和编码能够影响多腺苷酸束向mRNA前体3'末端添加的调节信号的其他序列是3'非翻译区。“5'非翻译区”是指位于编码序列上游的多核苷酸。
其他上游或下游非翻译元件包括增强子。增强子是用于增加启动子区域表达的多核苷酸。增强子是本领域公知的,包括但不限于SV40增强子区和35S增强子元件。
终止区可以是转录起始区天然的,可以是与目标有效连接的DNA序列天然的,可以是植物宿主天然的,或者可以来自另一个来源(即,对于启动子、感兴趣的DNA序列、植物宿主或其任何组合来说是外源的或异源的)。方便的终止区可从根癌农杆菌(A.tumefaciens)的Ti质粒获得,例如章鱼碱合酶和胭脂碱合酶终止区。也参见Guerineau等人,(1991)Mol.Gen.Genet.262:141-144;Proudfoot(1991)Cell 64:671-674;Sanfacon等人,(1991)Genes Dev.5:141-149;Mogen等人,(1990)Plant Cell 2:1261-1272;Munroe等人,(1990)Gene 91:151-158;Ballas等人,(1989)Nucleic Acids Res.17:7891-7903;和Joshi等人,(1987)Nucleic Acid Res.15:9627-9639。
在适当的情况下,可以优化基因以增加转化的宿主细胞(合成的DNA序列)中的表达。也就是说,可以使用宿主细胞优选的密码子合成基因以改善表达,或者可以使用宿主偏好的密码子使用频率的密码子合成基因。合成DNA序列的开放读码框在细胞中的表达导致产生本发明的多肽。合成DNA序列可用于简单地去除不需要的限制性内切核酸酶位点,促进DNA克隆策略,改变或消除任何潜在的密码子偏倚,改变或改善GC含量,去除或改变备选读码框,和/或改变或去除可能存在于天然DNA序列中的内含子/外显子剪接识别位点,多腺苷酸化位点,Shine-Delgarno序列,不需要的启动子元件等。通常,基因的GC含量将增加。参见,例如,Campbell和Gowri(1990)Plant Physiol.92:1-11讨论了宿主偏好的密码子使用。本领域可用于合成植物优选基因的方法。参见,例如,美国专利号5,380,831和5,436,391,美国专利公开号20090137409和Murray等人,(1989)Nucleic Acids Res.17:477-498,在此引入作为参考。
也可以利用合成的DNA序列引入DNA序列的其他改进,例如引入内含子序列,产生表达为与细胞器靶向序列的蛋白质融合的DNA序列,例如叶绿体转运肽,质外体/液泡靶向肽,或导致所得肽保留在内质网中的肽序列。因此,在一个实施方案中,杀线虫蛋白靶向叶绿体用于表达。以这种方式,当杀线虫蛋白不直接***叶绿体中时,表达盒将另外含有编码转运肽的核酸,以将杀线虫蛋白导向叶绿体。此类转运肽在本领域中是已知的。参见,例如,VonHeijne等人,PlantMol.Biol.Rep.9:104-126;Clark等人,(1989)J.Biol.Chem.264:17544-17550;Della-Cioppa等人,(1987)Plant Physiol.84:965-968;Romer等人,(1993)Biochem.Biophys.Res.Commun.196:1414-1421;和Shah等人,(1986)Science233:478-481。
可以优化靶向叶绿体的杀虫基因以在叶绿体中表达,以解释植物细胞核和该细胞器之间密码子使用的差异。以这种方式,可以使用叶绿体优选密码子合成目标核酸。参见,例如,美国专利号5,380,831,其通过引用并入到本文中。
植物转化
本发明的方法包括将核苷酸构建体引入植物中。“引入”旨在以使得构建体进入植物细胞内部的方式向植物提供核苷酸构建体。本发明的方法不需要使用用于将核苷酸构建体引入植物的特定方法,只需要使核苷酸构建体进入植物的至少一个细胞的内部。将核苷酸构建体引入植物的方法是本领域已知的,包括但不限于稳定转化方法、瞬时转化方法和病毒介导的方法。
“植物”是指整株植物,植物器官(例如叶、茎、根等),种子,植物细胞,繁殖体,胚和它们的后代。植物细胞可以是分化的或未分化的(例如愈伤组织,悬浮培养细胞,原生质体,叶细胞,根细胞,韧皮部细胞,花粉)。
“转基因植物”或“转化植物”或“稳定转化的”植物或细胞或组织是指将外源核酸序列或DNA片段掺入或整合到植物细胞中的植物。这些核酸序列包括外源的或未存在于未转化的植物细胞中的核酸序列,以及可以是内源的或存在于未转化的植物细胞中的核酸序列。“异源”通常是指对细胞或其存在的天然基因组的一部分不是内源的核酸序列,并且已经通过感染,转染,显微注射,电穿孔,显微喷射等添加到细胞中。
本发明的转基因植物表达一种或多种本文公开的新毒素序列。在一些实施方案中,本发明的蛋白质或核苷酸序列有利地在植物中与编码蛋白质或RNA的其他基因组合,所述蛋白质或RNA赋予这些植物有用的农艺性质。在编码赋予转化植物有用农艺性质的蛋白质或RNA的基因中,可以提及编码赋予一种或多种除草剂耐受性的蛋白质的DNA序列,以及赋予某些昆虫耐受性的其他物质,赋予对某些疾病耐受性的那些,编码提供线虫或昆虫控制的RNA的DNA等。这些基因特别描述于公开的PCT专利申请WO91/02071和WO95/06128以及美国专利7,923,602和美国专利申请公开号20100166723中,其各自通过引用整体并入本文。在各种实施方案中,转基因植物还包含一种或多种用于昆虫抗性的其他基因(例如,Cry1,例如Cry1A,Cry1B,Cry1C,Cry1D,Cry1E和Cry1F家族的成员;Cry2,例如Cry2A家族的成员;Cry9,例如Cry9A,Cry9B,Cry9C,Cry9D,Cry9E和Cry9F家族的成员;等等)。本领域技术人员将理解,转基因植物可包含赋予目的农学性状的任何基因。
在编码赋予转化植物细胞和植物某些除草剂耐受性的蛋白质的DNA序列中,可以提及WO2009/152359中描述的bar或PAT基因或天蓝色链霉菌(Streptomyces coelicolor)基因,其赋予对草铵膦除草剂的耐受性,编码赋予对具有EPSPS作为靶标的除草剂例如草甘膦及其盐的耐受性合适的EPSPS的基因(US 4,535,060,US 4,769,061,US 5,094,945,US4,940,835,US 5,188,642,US 4,971,908,US 5,145,783,US 5,310,667,US 5,312,910,US5,627,061,US 5,633,435),编码草甘膦-n-乙酰转移酶的基因(例如,US 8,222,489,US 8,088,972,US 8,044,261,US 8,021,857,US 8,008,547,US 7,999,152,US 7,998,703,US7,863,503,US 7,714,188,US 7,709,702,US 7,666,644,US 7,666,643,US 7,531,339,US7,527,955和US 7,405,074),编码草甘膦氧化还原酶的基因(例如,US 5,463,175),或编码HPPD抑制剂耐受蛋白的基因(例如,描述于WO 2004/055191,WO 199638567,US 6791014,WO2011/068567,WO2011/076345,WO2011/085221,WO2011/094205,WO2011/068567,WO2011/094199,WO2011/094205,WO2011/145015,WO2012/056401和WO2014/043435的HPPD抑制剂耐受性基因)。
在编码赋予对具有EPSPS作为靶标的除草剂的耐受性的合适的EPSPS的DNA序列中,更特别地提及编码植物EPSPS的基因,特别是玉米EPSPS,特别是包含两个突变,特别是氨基酸位置102处的突变和氨基酸位置106处的突变的玉米EPSPS(WO2004/074443),并且其描述于专利申请US6566587,下文称为双突变体玉米EPSPS或2mEPSPS,或编码分离自农杆菌的EPSPS的基因,其描述于美国专利5,633,435的序列ID No.2和序列ID No.3,也称为CP4。
在编码赋予对具有EPSPS作为靶标的除草剂的耐受性的合适的EPSPS的DNA序列中,更具体地提及编码来自球形节杆菌的EPSPS GRG23的基因,以及突变体GRG23 ACE1,GRG23 ACE2,或GRG23 ACE3,特别是WO2008/100353中描述的GRG23的突变体或变体,例如WO2008/100353中的SEQ ID No.29的GRG23(ace3)R173K。
在编码EPSPS的DNA序列的情况下,更具体地是编码上述基因的序列,编码这些酶的序列有利地在编码转运肽的序列之前,特别是美国专利5,510,471或5,633,448中描述的“优化的转运肽”。
可以与本发明的核酸序列组合的示例性除草剂耐受性性状还包括至少一种ALS(乙酰乳酸合酶)抑制剂(WO2007/024782);突变的拟南芥ALS/AHAS基因(美国专利6,855,533);编码通过代谢赋予对2,4-D(2,4-二氯苯氧基乙酸)的耐受性的2,4-D-单加氧酶的基因(美国专利6,153,401);编码通过代谢赋予对麦草畏(3,6-二氯-2-甲氧基苯甲酸)的耐受性的麦草畏单加氧酶的基因(US 2008/0119361和US 2008/0120739)。
在各种实施方案中,本发明的核酸与一种或多种除草剂耐受基因堆叠,所述除草剂耐受基因包括一种或多种HPPD抑制剂除草剂耐受基因,和/或一种或多种耐受草甘膦和/或草铵膦的基因。
在编码涉及昆虫耐受性的蛋白质的DNA序列中,更具体地提及文献中广泛描述的Bt蛋白质,并且是本领域技术人员公知的。还将提及从细菌如发光杆菌(Photorhabdus)(WO97/17432和WO98/08932)中提取的蛋白质。
在编码赋予昆虫耐受性的新特性的感兴趣的蛋白质的这些DNA序列中,更具体地提及文献中广泛描述的Bt Cry或VIP蛋白质,并且是本领域技术人员公知的。这些包括Cry1F蛋白或衍生自Cry1F蛋白的杂合体(例如,US 6,326,169中描述的杂合Cry1A-Cry1F蛋白;US 6,281,016;US 6,218,188,或其毒性片段),Cry1A型蛋白或其毒性片段,优选如EP451878中所述的Cry1Ac蛋白或衍生自Cry1Ac蛋白的杂合体(例如,US5,880,275中描述的杂合Cry1Ab-Cry1Ac蛋白)或Cry1Ab或Bt2蛋白或其杀虫片段,如WO2002/057664中所述的Cry2Ae,Cry2Af或Cry2Ag蛋白或其毒性片段,WO 2007/140256(SEQ ID No.7)中描述的Cry1A.105蛋白或其毒性片段,NCBI登录号ABG20428的VIP3Aa19蛋白,NCBI登录号ABG20429的VIP3Aa20蛋白(WO2007/142840中的SEQ ID No.2),在COT202或COT203棉事件(分别为WO2005/054479和WO2005/054480)中产生的VIP3A蛋白,如WO2001/47952中所述的Cry蛋白,如Estruch等人(1996),Proc Natl Acad Sci US A.28;93(11):5389-94和US 6,291,156所述的VIP3Aa蛋白或其毒性片段,来自致病杆菌(Xenorhabdus)(如WO98/50427中所述),沙雷氏菌(Serratia)(特别是来自嗜虫沙雷菌(S.entomophila))或发光杆菌种菌株的杀虫蛋白,如WO98/08932中所述的来自发光杆菌的Tc-蛋白质(例如,Waterfield等人,2001,ApplEnviron Microbiol.67(11):5017-24;Ffrench-Constant和Bowen,2000,Cell Mol LifeSci;57(5):828-33)。此外,在一些(1-10,优选1-5)氨基酸上与任何上述序列,特别是其毒性片段的序列不同,或与转运肽,例如,质体转运肽或其他蛋白质或肽融合的任何一种这些蛋白质的任何变体或突变体包括在本文中。
在各种实施方案中,本发明的核酸可以在植物中与一种或多种赋予所需性状的基因组合,所述所需性状为例如除草剂耐受性,昆虫耐受性,耐旱性,线虫控制,水利用效率,氮利用效率,改善的营养价值,抗病性,改善的光合作用,改善的纤维品质,胁迫耐受性,改善的繁殖等。
可以在相同物种的植物中与本发明的基因组合(例如,通过杂交或通过将含有另一转基因事件的植物与本发明的嵌合基因重新转化)的特别有用的转基因事件,包括事件BPS-CV127-9(大豆,除草剂耐受性,保藏号为NCIMB No.41603,描述于WO2010/080829);事件DAS21606-3/1606(大豆,除草剂耐受性,保藏为PTA-11028,描述于WO2012/033794),事件DAS-44406-6/pDAB8264.44.06.1(大豆,除草剂耐受性,保藏为PTA-11336,描述在WO2012/075426中),事件DAS-14536-7/pDAB8291.45.36.2(大豆,除草剂耐受性,保藏为PTA-11335,描述于WO2012/075429),事件DAS68416(大豆,除草剂耐受性,保藏为ATCC PTA-10442,描述于WO2011/066384或WO2011/066360);事件DP-305423-1(大豆,品质性状,未保藏,描述于US-A 2008-312082或WO2008/054747);事件DP-356043-5(大豆,除草剂耐受性,保藏为ATCCPTA-8287,描述于US-A 2010-0184079或WO2008/002872);事件FG72(大豆,除草剂耐受性,保藏为PTA-11041,描述于WO2011/063413),事件LL27(大豆,除草剂耐受性,保藏为NCIMB41658,描述于WO2006/108674或US-A 2008-320616);事件LL55(大豆,除草剂耐受性,保藏为NCIMB 41660,描述于WO 2006/108675或US-A 2008-196127);事件MON87701(大豆,昆虫控制,保藏为ATCC PTA-8194,描述于US-A 2009-130071或WO2009/064652);事件MON87705(大豆,品质性状-除草剂耐受性,保藏为ATCC PTA-9241,描述于US-A 2010-0080887或WO2010/037016);事件MON87708(大豆,除草剂耐受性,保藏为ATCC PTA-9670,描述于WO2011/034704);事件MON87712(大豆,产量,保藏为PTA-10296,描述于WO2012/051199),事件MON87754(大豆,品质性状,保藏为ATCC PTA-9385,描述于WO2010/024976);事件MON87769(大豆,品质性状,保藏为ATCC PTA-8911,描述于US-A 2011-0067141或WO2009/102873);事件MON89788(大豆,除草剂耐受性,保藏为ATCC PTA-6708,描述于US-A2006-282915或WO2006/130436);事件SYHT0H2/SYN-000H2-5(大豆,除草剂耐受性,保藏为PTA-11226,描述于WO2012/082548),任选地与事件EE-GM1/LL27或事件EE-GM2/LL55(WO2011/063413A2)堆叠的事件EE-GM3/FG72(大豆,除草剂耐受性,ATCC登录号PTA-11041);事件DAS-68416-4(大豆,除草剂耐受性,ATCC登录号PTA-10442,WO2011/066360A1);事件DAS-68416-4(大豆,除草剂耐受性,ATCC登录号PTA-10442,WO2011/066384A1);事件DAS-21606-3(大豆,除草剂耐受性,ATCC登录号PTA-11028,WO2012/033794A2);事件MON-87712-4(大豆,品质性状,ATCC登录号:PTA-10296,WO2012/051199A2);事件DAS-44406-6(大豆,堆积的除草剂耐受性,ATCC登录号PTA-11336,WO2012/075426A1);事件DAS-14536-7(大豆,堆积的除草剂耐受性,ATCC登录号:PTA-11335,WO2012/075429A1);事件SYN-000H2-5(大豆,除草剂耐受性,ATCC登录号PTA-11226,WO2012/082548A2);事件8264.44.06.1(大豆,堆积的除草剂耐受性,登录号PTA-11336,WO2012075426A2);事件8291.45.36.2(大豆,堆积的除草剂耐受性,登录号PTA-11335,WO2012075429A2);事件SYHT0H2(大豆,ATCC登录号PTA-11226,WO2012/082548A2);事件pDAB8264.42.32.1(大豆,堆积的除草剂耐受性,ATCC登录号PTA-11993,WO2013/010094A1)。
此外,本文提供了用于生产与另一SCN抗性基因座/基因组合的、包含编码SEQ IDNO:1或2的核苷酸序列的大豆植物或种子的方法,例如通过组合包含编码SEQ ID NO:1或2的核苷酸序列的大豆植物或种子与在相同大豆植物/种子中发生的另一SCN抗性基因座/基因,并且种植包含编码SEQ ID NO:1或2的核苷酸序列和所述其他SCN抗性基因座/基因的种子。在一个实施方案中,本发明的植物、细胞或种子含有在大豆中出现的一种或多种其他SCN抗性基因座/基因,以在本发明的大豆植物,细胞或种子中获得不同SCN抗性来源的组合。已知几种大豆SCN抗性基因座或基因,并且这些的一种或多种可以与包含SEQ ID NO:1或2的植物在相同的植物、细胞或种子中组合,例如来自抗性源PI 88788,PI 548402(Peking),PI 437654(Hartwig或)的任一种SCN抗性基因座/基因,或它们的任何组合,或一种或多种天然SCN抗性基因座/基因rhg1,rhg1-b,rhg2,rhg3,Rhg4,Rhg5,qSCN11,cqSCN-003,cqSCN-005,cqSCN-006,cqSCN-007,或任何一种大豆染色体1,3,4,5,6,7,8,9,10,11,12,13,14,15,16,17,18,19或20中鉴定的任一个SCN抗性基因座,或其任何组合(Kim等人2016,Theor.Appl.Genet.129(12):2295-2311;Kim和Diers 2013,CropScience 53:775-785;Kazi等人2010,Theor.Appl.Gen.120(3):633-644;Glover等人2004,Crop Science 44(3):936-941;www.soybase.org;Concibido等人2004,Crop Science 44:1121-1131;Webb等人1995,Theor.Appl.Genet.91:574-581)。此外,在一个实施方案中,本发明的植物或种子与从SCN抗性来源PI 548316,PI 567305,PI 437654,PI 90763,PI404198B,PI 88788,PI468916,PI 567516C,PI 209332,PI 438489B,PI 89772,Peking,PI548402,PI 404198A,PI 561389B,PI 629013,PI 507471,PI 633736,PI 507354,PI404166,PI 437655,PI 467312,PI 567328,PI 22897,或PI 494182中的任一种获得的大豆中的一个或多个SCN抗性基因座组合。
植物细胞的转化可以通过本领域已知的几种技术之一完成。可以修饰本发明的杀虫基因以获得或增强植物细胞中的表达。通常,表达这种蛋白质的构建体将含有驱动基因转录的启动子,以及允许转录终止和多腺苷酸化的3'非翻译区。这种构建体的组织在本领域中是众所周知的。在一些情况下,设计基因使得所得肽在植物细胞内分泌或以其他方式靶向可能是有用的。例如,可以将基因工程化为含有信号肽以促进肽向内质网的转移。也可以优选将植物表达盒设计成含有内含子,使得内含子的mRNA加工是表达所必需的。
通常将该“植物表达盒”***“植物转化载体”中。该植物转化载体可以包含实现植物转化所需的一种或多种DNA载体。例如,本领域的常见做法是利用由一个以上连续DNA区段组成的植物转化载体。这些载体在本领域通常称为“二元载体”。二元载体以及具有辅助质粒的载体最常用于农杆菌介导的转化,其中实现有效转化所需的DNA区段的大小和复杂性非常大,将功能分离到单独的DNA分子上是有利的。二元载体通常含有质粒载体,其含有T-DNA转移所需的顺式作用序列(例如左边界和右边界),可选择的标记物,其被工程化为能够在植物细胞中表达,和“目的基因”(一种工程化成能够在植物细胞中表达的基因,希望为所述植物细胞产生转基因植物)。该质粒载体上也存在细菌复制所需的序列。顺式作用序列以这样的方式排列,以允许有效转移到植物细胞中并在其中表达。例如,选择标记基因和杀虫基因位于左边界和右边界之间。第二质粒载体通常含有介导T-DNA从农杆菌转移到植物细胞的反式作用因子。该质粒通常含有毒力功能(Vir基因),其允许通过农杆菌感染植物细胞,并通过边界序列的切割和vir介导的DNA转移来转移DNA,如本领域所理解的(Hellens和Mullineaux(2000)Trend in Plant Science 5:446-451)。几种类型的农杆菌菌株(例如LBA4404,GV3101,EHA101,EHA105等)可用于植物转化。第二质粒载体不是通过其他方法,例如显微喷射,显微注射,电穿孔,聚乙二醇等转化植物所必需的。
通常,植物转化方法涉及将异源DNA转移到目标植物细胞(例如未成熟或成熟胚,悬浮培养物,未分化愈伤组织,原生质体等)中,然后应用最大阈值水平的适当选择(取决于选择标记基因)从一组未转化的细胞团中回收转化的植物细胞。通常将外植体转移到新鲜供应的相同培养基中并常规培养。随后,在置于补充有最大阈值水平的选择剂的再生培养基上后,将转化的细胞分化成枝条。然后将枝条转移到选择性生根培养基中以回收生根的枝条或小植物。然后转基因小植物生长成成熟植物并产生可育种子(例如Hiei等人(1994)The Plant Journal6:271-282;Ishida等人(1996)Nature Biotechnology14:745-750)。通常将外植体转移到新鲜供应的相同培养基中并常规培养。用于产生转基因植物的技术和方法的一般描述见Ayres和Park(1994)Critical Reviews in Plant Science 13:219-239和Bommineni和Jauhar(1997)Maydica 42:107-120。由于转化的材料含有许多细胞;转化的和未转化的细胞都存在于任何块的受试目标愈伤组织或组织或细胞组中。杀死未转化细胞并允许转化细胞增殖的能力导致转化的植物培养物。通常,去除未转化细胞的能力限制了转化植物细胞的快速恢复和转基因植物的成功产生。
转化方案以及将核苷酸序列引入植物的方案可以根据靶向转化的植物或植物细胞的类型,即单子叶植物或双子叶植物而变化。转基因植物的产生可以通过几种方法之一进行,包括但不限于显微注射,电穿孔,直接基因转移,通过农杆菌将异源DNA引入植物细胞(农杆菌介导的转化),用附着于颗粒的异源外来DNA轰击植物细胞,弹道粒子加速,气溶胶束转化(美国公开申请号20010026941;美国专利号4,945,050;国际公开号WO 91/00915;美国公开申请号2002015066),Lec1转化,以及各种其他非粒子直接介导的DNA转移方法。
转化叶绿体的方法是本领域已知的。参见,例如,Svab等人(1990)Proc.Natl.Acad.Sci.USA 87:8526-8530;Svab和Maliga(1993)Proc.Natl.Acad.Sci.USA90:913-917;Svab和Maliga(1993)EMBO J.12:601-606。该方法依赖于粒子枪递送含有选择标记的DNA并通过同源重组将DNA靶向质体基因组。另外,质体转化可以通过核编码和质体定向的RNA聚合酶的组织优选的表达反式激活沉默的质体携带的转基因来完成。McBride等人(1994)Proc.Natl.Acad.Sci.USA 91:7301-7305报道了这种***。
在将异源外源DNA整合到植物细胞中之后,然后在培养基中应用最大阈值水平的适当选择以杀死未转化的细胞,并通过定期转移到新鲜培养基中来分离和增殖从该选择处理中存活的推定转化的细胞。通过适当选择的连续传代和攻击,可以鉴定和增殖用质粒载体转化的细胞。然后可以使用分子和生物化学方法来确认整合的异源目的基因存在于转基因植物的基因组中。
已经转化的细胞可以按照常规方法生长成植物。参见,例如,McCormick等人,(1986)Plant Cell Reports 5:81-84。然后可以使这些植物生长,并用相同的转化菌株或不同菌株授粉,并鉴定具有所需表型特征的组成型表达的所得杂交体。可以生长两代或更多代以确保稳定维持和遗传所需表型特征的表达,然后收获种子以确保实现所需表型特征的表达。以这种方式,本发明提供了转化的种子(也称为“转基因种子”),其具有稳定地掺入其基因组中的本发明的核苷酸构建体,例如,本发明的表达盒。
植物转化评价
在将异源外源DNA引入植物细胞后,异源基因在植物基因组中的转化或整合通过各种方法证实,例如分析与整合基因相关的核酸、蛋白质和代谢物。
PCR分析是一种用于在移植到土壤之前的早期阶段筛选转化的细胞、组织或枝条中是否存在掺入的基因快速的方法(Sambrook和Russell(2001)Molecular Cloning:ALaboratory Manual.Cold Spring Harbor Laboratory Press,Cold春天港,纽约)。使用对目的基因或农杆菌载体背景等特异的寡核苷酸引物进行PCR。
植物转化可以通过基因组DNA的Southern印迹分析来确认(Sambrook和Russell,2001,同上)。通常,从转化体中提取总DNA,用合适的限制酶消化,在琼脂糖凝胶中分级分离并转移到硝酸纤维素膜或尼龙膜上。然后用例如放射性标记的32P靶DNA片段探测膜或“印迹”,以证实根据标准技术将导入的基因整合到植物基因组中(Sambrook和Russell,2001,同上)。
在Northern印迹分析中,从转化体的特定组织中分离RNA,在甲醛琼脂糖凝胶中分级分离,并根据本领域常规使用的标准方法印迹到尼龙滤膜上(Sambrook和Russell,2001,同上)。然后通过本领域已知的方法(Sambrook和Russell,2001,同上),通过将滤膜与源***虫基因的放射性探针杂交来测试由杀虫基因编码的RNA的表达。
可以对转基因植物进行蛋白质印迹,生物化学测定等,以通过标准程序(Sambrook和Russell,2001,同上)使用与存在于杀线虫蛋白上的一个或多个表位结合的抗体来确认由杀虫基因编码的蛋白质的存在。
植物中的杀虫活性
在本发明的另一个方面,可以产生表达杀线虫蛋白的转基因植物,所述杀线虫蛋白具有针对线虫害虫的杀虫活性。上述通过实施例描述的方法可用于产生转基因植物,但产生转基因植物细胞的方式对本发明并不重要。本领域已知或描述的方法,例如农杆菌介导的转化,生物射弹转化和非粒子介导的方法可以由实验者自行决定使用。表达杀线虫蛋白的植物可以通过本领域描述的常规方法分离,例如通过转化愈伤组织,筛选转化的愈伤组织和从这种转基因愈伤组织再生可育植物。在这样的过程中,可以使用任何基因作为选择标记,只要其在植物细胞中的表达赋予鉴定或选择转化细胞的能力。
已经开发了许多用于植物细胞的标记,例如对氯霉素,氨基糖苷G418,潮霉素等的抗性。编码参与叶绿体代谢的产物的其他基因也可用作选择标记。例如,可以特别使用对植物除草剂如草甘膦、溴苯腈或咪唑啉酮提供抗性的基因。已经报道了这些基因(Stalker等人(1985)J.Biol.Chem.263:6310-6314(溴苯腈抗性腈水解酶基因);和Sathasivan等人(1990)Nucl.Acids Res.18:2188(AHAS咪唑啉酮抗性基因)。另外,本文公开的基因可用作评估细菌或植物细胞转化的标记物。用于检测植物,植物器官(例如,叶,茎,根等),种子,植物细胞,繁殖体,胚胎或其后代中转基因的存在的方法在本领域中是熟知的。在一个实施方案中,通过测试针对线虫害虫的杀虫活性来检测转基因的存在。
可以测试表达杀线虫蛋白的可育植物对线虫害虫的杀虫活性,并且显示选择用于进一步育种的最佳活性的植物。用于测定害虫活性的方法是本领域可获得的。通常,将蛋白质混合并用于进食测定。参见,例如Marrone等人,(1985)J.of Economic Entomology 78:290-293。
本发明可用于转化任何植物物种,包括但不限于单子叶植物和双子叶植物。感兴趣的植物的实例包括但不限于玉米(玉米),高粱,小麦,向日葵,番茄,十字花科植物,辣椒,马铃薯,棉花,稻,大豆,甜菜,甘蔗,烟草,大麦和油菜,芸苔属植物,紫花苜蓿,黑麦,小米,红花,花生,甘薯,木薯,咖啡,椰子,菠萝,柑橘树,可可,茶,香蕉,鳄梨,无花果,番石榴,芒果,橄榄,木瓜,腰果,澳洲坚果,杏仁,燕麦,蔬菜,观赏植物和针叶树。
杀虫控制中的应用
使用包含本发明的核苷酸序列或其变体的菌株在害虫控制中或在其他生物体中用作杀虫剂的一般方法是本领域已知的。参见,例如美国专利No.5,039,523和EP0480762A2。
可以遗传改变微生物以含有编码SEQ ID NO:1或2的核苷酸序列,或其杀线虫活性变体或片段,并且蛋白质可以用于保护农作物和产品免受害虫侵害。在本发明的一个方面,用试剂处理产生毒素(杀虫剂)的生物体的整个(即未裂解的)细胞,当所述细胞施用于目标害虫环境时,所述试剂延长细胞中产生的毒素的活性。
或者,通过将杀虫基因引入细胞宿主中来产生杀虫剂。杀虫基因的表达直接或间接地导致农药的细胞内产生和维持。在本发明的一个方面,然后在将细胞应用于目标害虫的环境时延长细胞中产生的毒素活性的条件下处理这些细胞。所得产物保留了毒素的毒性。然后可以根据常规技术配制这些天然包封的杀虫剂,以应用于容纳目标害虫的环境,例如土壤,水和植物的叶子。参见例如EPA 0192319和其中引用的参考文献。或者,可以配制表达本发明基因的细胞,以允许将所得材料用作杀虫剂。
本发明的活性成分通常以组合物的形式施用,并且可以与其他化合物同时或相继施用于待处理的作物区域或植物。这些化合物可以是肥料,除草剂,冷冻保护剂,表面活性剂,洗涤剂,杀虫皂,休眠油,聚合物和/或定时释放或可生物降解的载体制剂,其允许在单次施用制剂后长期给药目标区域。如果需要,它们还可以是选择性除草剂,化学杀虫剂,杀病毒剂,杀微生物剂,杀螨剂,杀虫剂,杀真菌剂,杀细菌剂,杀线虫剂,杀软体动物剂或这些制剂中的几种的混合物,以及制剂领域中惯用的其它农业上可接受的载体,表面活性剂或促进应用的佐剂。合适的载体和佐剂可以是固体或液体,并且对应于制剂技术中通常采用的物质,例如,天然或再生矿物质,溶剂,分散剂,润湿剂,增粘剂,粘合剂或肥料。同样地,可以将制剂制成可食用的“诱饵”或制成害虫“诱捕器”,以允许由杀虫制剂的目标害虫摄食或摄取。
施用本发明的活性成分或本发明的农业化学组合物的方法包括叶施用、种子包衣和土壤施用,所述组合物含有至少一种本文公开为SEQ ID NO:1或2的杀线虫蛋白,或其杀线虫有效的变体或片段。施用次数和施用率取决于相应害虫的侵袭强度。
该组合物可以配制成粉末,粉剂,丸剂,颗粒剂,喷雾剂,乳剂,胶体剂,溶液剂等,并且可以通过干燥,冻干,匀浆,萃取,过滤,离心,沉降,或包含该多肽的细胞培养物的浓缩等常规方法制备。在含有至少一种这样的杀虫多肽的所有这些组合物中,多肽可以以约1%至约99%重量的浓度存在。
线虫害虫可以通过本发明的方法在给定区域中杀死或减少数量,或者可以预防性地应用于环境区域以防止易感害虫的侵袭。优选地,害虫摄入杀虫有效量的多肽或与杀虫有效量的多肽接触。“杀虫有效量”或“杀线虫有效量”是指能够使至少一种害虫死亡,或显著减少害虫生长、摄食或害虫或害虫感染的宿主植物的正常生理发育的杀虫剂或杀线虫剂的量。该量将根据以下因素而变化,例如,要控制的特定目标害虫,特定环境,位置,植物,作物或待处理的农业场所,环境条件,以及方法,速率,浓度,稳定性和杀线虫有效多肽组合物的施用量。制剂还可以在气候条件,环境考虑因素和/或施用频率和/或害虫侵袭的严重性方面变化。
所述杀虫组合物可以通过将细菌细胞、晶体和/或孢子悬浮液或分离的蛋白质组分与所需的农业上可接受的载体一起配制而制备。组合物可在给药前以适当的方式配制,例如冻干,冷冻干燥,干燥,或在含水载体,培养基或合适的稀释剂如盐水或其它缓冲液中。配制的组合物可以是粉剂或颗粒状物质,或油(植物或矿物质)中的悬浮液,或水或油/水乳液,或可湿性粉剂形式,或与适于农业应用的任何其他载体材料组合使用。合适的农业载体可以是固体或液体,并且是本领域熟知的。术语“农业上可接受的载体”包括通常用于杀虫剂制剂技术的所有佐剂,惰性组分,分散剂,表面活性剂,增粘剂,粘合剂等;这些是杀虫剂制剂领域的技术人员所熟知的。制剂可以与一种或多种固体或液体佐剂混合,并通过各种方法制备,例如,通过使用常规制剂技术将杀虫剂组合物与合适的佐剂均匀混合,共混和/或研磨。合适的制剂和施用方法描述于美国专利号6,468,523中,其通过引用并入本文。
提高植物产量的方法
提供了用于增加植物产量的方法。所述方法包括提供表达编码本文中公开的杀线虫多肽序列的多核苷酸的植物或植物细胞,并且在被线虫害虫侵袭(或易受线虫害虫侵袭)的田地使该植物或其种子生长,其中所述多肽对所述线虫害虫具有杀线虫活性。在一些实施方案中,本文所述Cry14多肽对短体线虫属物种(Pratylenchus spp.)具有杀线虫活性,并且所述田地被所述短体线虫侵袭。在各种实施方案中,短体线虫是短尾短体线虫。在另外的实施方案中,线虫是根结线虫,病变线虫或Lance线虫。如本文中所定义,植物的“产量”指由植物产生的生物质的质量和/或数量。“生物质”意指任何被测量的植物产物。生物质产生的增加是被测量植物产物的产量上的任意改善。增加植物产量具有几项商业应用。例如,增加植物叶片生物质可以增加用于人或动物消费的叶菜类的产量。另外,增加叶片生物质可以用于增加植物衍生的药用或工业产物的生产。产量的增加可以包含任何统计学上显著的增加,包括但不限于与不表达本文所述的杀虫蛋白的植物相比,产量增加至少1%、增加至少3%、增加至少5%、增加至少10%、增加至少20%、增加至少30%、至少50%、至少70%、至少100%或更多。在具体的方法中,植物产量由于表达本文中所公开杀线虫蛋白的植物对线虫害虫的抗性增强而增加。杀线虫蛋白的表达导致害虫侵袭或进食的能力降低。在各种实施方案中,与不表达本发明的杀线虫蛋白的植物相比,杀线虫蛋白的表达导致改善的根发育(例如,改善的根或根毛生长),改善的产量,更快的出苗,改善的植物胁迫管理,包括增加的胁迫耐受性和/或改善的胁迫恢复,增加的机械强度,改善的抗旱性,减少的真菌病感染,以及改善的植物健康。
还可以用一种或多种化学组合物对植物进行处理,所述的化学组合物包括一种或多种除草剂、杀虫剂或杀真菌剂。示例性化学组合物包括:水果/蔬菜除草剂:莠去津(Atrazine)、除草定(Bromacil)、敌草隆(Diuron)、草甘膦(Glyphosate)、利谷隆(Linuron)、嗪草酮(Metribuzin)、西玛津(Simazine)、氟乐灵(Trifluralin)、吡氟禾草灵(Fluazifop)、草铵膦(Glufosinate)、氯吡嘧磺隆(Halosulfuron Gowan)、百草枯(Paraquat)、炔草胺(Propyzamide)、稀禾定(Sethoxydim)、氟丙嘧草酯(Butafenacil)、氯吡嘧磺隆(Halosulfuron)、Indaziflam;水果/蔬菜杀虫剂:涕灭威(Aldicarb)、苏云金芽孢杆菌(Bacillus thuriengiensis)、甲萘威(Carbaryl)、克百威(Carbofuran)、毒死蜱(Chlorpyrifos)、氯氰菊酯(Cypermethrin)、溴氰菊酯(deltamethrin)、阿维菌素(Abamectin)、氟氯氰菊酯/β-氟氯氰菊酯(Cyfluthrin/Beta-Cyfluthrin)、顺式氰戊菊酯(Esfenvalerate)、高三氟氯氰菊酯(Lambda-cyhalothrin)、灭螨醌(Acequinocyl)、联苯肼酯(Bifenazate)、甲氧虫酰肼(Methoxyfenozide)、双苯氟脲(Novaluron)、环虫酰肼(Chromafenozide)、噻虫啉(Thiacloprid)、呋虫胺(Dinotefuran)、嘧螨酯(fluacrypyrim)、螺螨酯(spirodiclofen)、γ-氯氟氰菊酯(gama-cyhalothrin)、螺甲螨酯(spiromesifen)、艾克敌(spinosad)、氯虫苯甲酰胺(Rynaxypyr)、溴氰虫酰胺(Cyazypyr)、杀铃脲(Triflumuron)、螺虫乙酯(Spirotetramat)、吡虫啉(Imidacloprid)、氟虫双酰胺(Flubendiamide)、硫双威(Thiodicarb)、氰氟虫腙(Metaflumizone)、氟啶虫胺腈(Sulfoxaflor)、丁氟螨酯(Cyflumetofen)、Cyanopyrafen、可尼丁(Clothianidin)、噻虫嗪(Thiamethoxam)、Spinotoram、硫双威(Thiodicarb)、氟啶虫酰胺(Flonicamid)、甲硫威(Methiocarb)、氨基阿维菌素苯甲酸盐(Emamectin-benzoate)、二唑虫(Indoxacarb)、苯线磷(Fenamiphos)、吡丙醚(Pyriproxifen)、苯丁锡(Fenbutatin-oxid);水果/蔬菜杀真菌剂:唑嘧菌胺(Ametoctradin)、腈嘧菌酯(Azoxystrobin)、苯噻菌胺(Benthiavalicarb)、啶酰菌胺(Boscalid)、克菌丹(Captan)、多菌灵(Carbendazim)、百菌清(Chlorothalonil)、铜(Copper)、氰霜唑(Cyazofamid)、环氟菌胺(Cyflufenamid)、霜脲氰(Cymoxanil)、环丙唑醇(Cyproconazole)、环丙嘧啶(Cyprodinil)、苯醚甲环唑(Difenoconazole)、烯酰吗啉(Dimetomorph)、二噻农(Dithianon)、咪唑菌酮(Fenamidone)、环酰菌胺(Fenhexamid)、氟啶胺(Fluazinam)、咯菌腈(Fludioxonil)、氟吡菌胺(Fluopicolide)、氟吡菌酰胺(Fluopyram)、氟嘧菌酯(Fluoxastrobin)、氟唑菌酰胺(Fluxapyroxad)、灭菌丹(Folpet)、福賽得(Fosetyl)、异菌脲(Iprodione)、丙森锌(Iprovalicarb)、吡唑萘菌胺(Isopyrazam)、苯氧菊(Kresoxim-methyl)、代森锰锌(Mancozeb)、双炔酰菌胺(Mandipropamid)、甲霜灵/精甲霜灵(Metalaxyl/mefenoxam)、代森联(Metiram)、苯菌酮(Metrafenone)、腈菌唑(Myclobutanil)、戊菌唑(Penconazole)、吡噻菌胺(Penthiopyrad)、啶氧菌酯(Picoxystrobin)、霜霉威(Propamocarb)、丙环唑(Propiconazole)、甲代森锌(Propineb)、碘喹唑酮(Proquinazid)、丙硫菌唑(Prothioconazole)、吡唑醚菌酯(Pyraclostrobin)、嘧霉胺(Pyrimethanil)、苯氧喹啉(Quinoxyfen)、螺环菌胺(Spiroxamine)、硫磺(Sulphur)、戊唑醇(Tebuconazole)、硫芬酯(Thiophanate-methyl)、肟菌酯(Trifloxystrobin);
谷类植物除草剂:
2.4-D、酰嘧磺隆(Amidosulfuron)、溴苯腈(Bromoxynil)、唑草酮-E(Carfentrazone-E)、氯麦隆(Chlorotoluron)、氯磺隆(Chlorsulfuron)、炔草酸-P(Clodinafop-P)、二氯吡啶酸(Clopyralid)、麦草畏(Dicamba)、禾草灵酸-M(Diclofop-M)、吡氟草胺(Diflufenican)、恶禾草灵(Fenoxaprop)、双氟磺草胺(Florasulam)、氟酮磺隆-NA(Flucarbazone-NA)、氟噻草胺(Flufenacet)、氟啶嘧磺隆-M(Flupyrsulfuron-M)、氟草烟(Fluroxypyr)、呋草酮(Flurtamone)、草甘膦、碘甲磺隆(Iodosulfuron)、碘苯腈(Ioxynil)、异丙隆(Isoproturon)、MCPA、甲磺胺磺隆(Mesosulfuron)、甲磺隆(Metsulfuron)、二甲戊灵(Pendimethalin)、唑啉草酯(Pinoxaden)、丙苯磺隆(Propoxycarbazone)、苄草丹(Prosulfocarb)、甲氧磺草胺(Pyroxsulam)、磺酰磺隆(Sulfosulfuron)、噻磺隆(Thifensulfuron)、三甲苯草酮(Tralkoxydim)、醚苯磺隆(Triasulfuron)、苯磺隆(Tribenuron)、氟乐灵、三氟甲磺隆(Tritosulfuron);谷类植物杀 真菌剂:腈嘧菌酯、联苯吡菌胺(Bixafen)、啶酰菌胺、多菌灵、百菌清、环氟菌胺、环丙唑醇、嘧菌环胺、醚菌胺(Dimoxystrobin)、环氧康那唑(Epoxiconazole)、苯锈啶(Fenpropidin)、丁苯吗啉(Fenpropimorph)、氟吡菌酰胺、氟嘧菌酯、氟喹唑(Fluquinconazole)、氟唑菌酰胺、吡唑萘菌胺、醚菌酯、叶菌唑(Metconazole)、苯菌酮、吡噻菌胺、啶氧菌酯、咪鲜胺(Prochloraz)、丙环唑、碘喹唑酮、丙硫菌唑、吡唑醚菌酯、苯氧喹啉、螺环菌胺、戊唑醇、硫芬酯、肟菌酯;谷类植物杀虫剂:乐果(Dimethoate)、高三氟氯氰菊酯(λ-cyhalthrin)、溴氰菊酯、α-氯氰菊酯、β-氟氯氰菊酯、联苯菊酯(Bifenthrin)、吡虫啉、可尼丁、噻虫嗪、噻虫啉、啶虫脒(Acetamiprid)、呋虫胺(Dinetofuran)、Clorphyriphos、抗蚜威(Pirimicarb)、甲硫威、氟啶虫胺腈;玉米除草剂:莠去津、甲草胺(Alachlor)、溴苯腈、乙草胺(Acetochlor)、麦草畏、二氯吡啶酸、二甲吩草胺((S-)Dimeth enamid)、草铵膦、草甘膦、异唑草酮(Isoxaflutole)、精异丙甲草胺(S-Metolachlor)、甲基磺草酮(Mesotrione)、烟嘧磺隆(Nicosulfuron)、氟嘧磺隆(Primisulfuron)、玉嘧磺隆(Rimsulfuron)、磺草酮(Sulcotrione)、甲酰胺磺隆(Foramsulfuron)、苯吡唑草酮(Topramezone)、环磺酮(Tembotrione)、苯嘧磺草胺(Saflufenacil)、酮脲磺草吩(Thiencarbazone)、氟噻草胺、Pyroxasulfon;玉米杀虫剂:克百威、毒死蜱、联苯菊酯、溴氰虫酰胺、氟虫腈(Fipronil)、吡虫啉、高三氟氯氰菊酯、七氟菊酯(Tefluthrin)、特丁硫磷(Terbufos)、噻虫嗪、可尼丁、螺环菌胺、氟虫双酰胺、杀铃脲、氯虫苯甲酰胺、溴氰菊酯、硫双威、β-氟氯氰菊酯、氯氰菊酯、联苯菊酯、虱螨脲(Lufenuron)、丁基嘧啶磷(Tebupirimphos)、乙虫腈(Ethiprole)、溴氰虫酰胺、噻虫啉、啶虫脒、呋虫胺、阿维菌素(Avermectin);玉米杀真菌剂:嘧菌酯、联苯吡菌胺、啶酰菌胺、环丙唑醇、醚菌胺、环氧康那唑、种衣酯(Fenitropan)、氟吡菌酰胺、氟嘧菌酯、氟唑菌酰胺、吡唑萘菌胺、叶菌唑、吡噻菌胺、啶氧菌酯、戊唑醇、丙硫菌唑、吡唑醚菌酯、戊唑醇、肟菌酯;稻除草剂:丁草胺(Butachlor)、敌稗(Propanil)、四唑嘧磺隆(Azimsulfuron)、苄嘧磺隆(Bensulfuron)、氰氟草酯(Cyhalofop)、杀草隆(Daimuron)、四唑酰草胺(Fentrazamide)、咪唑磺隆(Imazosulfuron)、苯噻草胺(Mefenacet)、/>嗪草酮(Oxaziclomefone)、吡嘧磺隆(Pyrazosulfuron)、稗草畏(Pyributicarb)、二氯喹啉酸(Quinclorac)、禾草丹(Thiobencarb)、茚草酮(Indanofan)、氟噻草胺、四唑酰草胺、氯吡嘧磺隆、/>嗪草酮、双环磺草酮(Benzobicyclon)、环酯草醚(Pyriftalid)、五氟磺草胺(Penoxsulam)、双草醚(Bispy ribac)、丙炔/>草酮(Oxadiargyl)、乙氧磺隆(Ethoxysulfuron)、丙草胺(Pretilachlor)、甲基磺草酮、Tefuryltrione、/>草酮(Oxadiazone)、/>唑禾草灵、Pyrimisulfan;稻杀虫剂:二嗪农(Diazinon)、仲丁威(Fenobucarb)、丙硫克百威(Benfuracarb)、噻嗪酮(Buprofezin)、呋虫胺(Dinotefnran)、氟虫腈、吡虫啉、异丙威(Isoprocarb)、噻虫啉、环虫酰肼、可尼丁、乙虫腈、氟虫双酰胺、氯虫苯甲酰胺、溴氰菊酯、啶虫脒、噻虫嗪、溴氰虫酰胺、艾克敌、Spinotoram、氨基阿维菌素苯甲酸盐、氯氰菊酯、毒死蜱、醚菊酯(Etofenprox)、克百威、丙硫克百威、氟啶虫胺腈;稻杀真菌剂:嘧菌酯、多菌灵、环丙酰亚胺(Carpropamid)、双氯氰菌胺(Diclocymet)、苯醚甲环唑、敌瘟磷(Edifenphos)、嘧菌腙(Ferimzone)、庆大霉素(Gentamycin)、己唑醇(Hexaconazole)、恶霉灵(Hymexazol)、异稻瘟净(Iprobenfos,IBP)、稻瘟灵(Isoprothiolane)、异噻菌胺(Isotianil)、春雷霉素(Kasugamycin)、代森锰锌、苯氧菌胺(Metominostrobin)、肟醚菌胺(Orysastrobin)、戊菌隆(Pencycuron)、烯丙苯噻唑(probenazole)、戊唑醇、甲代森锌、咯喹酮(Pyroquilon)、戊唑醇、甲基硫菌灵、噻酰菌胺(Tiadinil)、三环唑(Tricyclazole)、三氟啶磺隆、有效霉素(Validamycin);棉花除草剂:敌草隆、伏草隆(Fluometuron)、MSMA、乙氧氟草醚(Oxyfluorfen)、扑草净(Prometryn)、氟乐灵、唑草酮、烯草酮(Clethodim)、吡氟禾草灵(Fluazifop-butyl)、草甘膦、氟草敏(Norflurazon)、二甲戊灵、嘧草硫醚(Pyrithiobac-sodium)、三氟啶磺隆、吡喃草酮(Tepraloxydim)、草铵膦、丙炔氟草胺(Flumioxazin)、噻苯隆(Thidiazuron);棉花杀虫剂:乙酰甲胺磷(Acephate)、涕灭威、毒死蜱、氯氰菊酯、溴氰菊酯、阿维菌素、啶虫脒、依马菌素苯甲酸盐(EmamectinBenzoate)、吡虫啉、/>二唑虫、高三氟氯氰菊酯、艾克敌、硫双威、γ-氯氟氰菊酯、螺甲螨酯、啶虫丙醚(Pyridalyl)、氟啶虫酰胺、氟虫双酰胺、杀铃脲、氯虫苯甲酰胺、β-氟氯氰菊酯、螺虫乙酯、可尼丁、噻虫嗪、噻虫啉、呋虫胺、氟虫双酰胺、溴氰虫酰胺、艾克敌、Spinotoram、γ氯氟氰菊酯、4-[[(6-氯吡啶-3-基)甲基](2,2-二氟乙基)氨基]呋喃-2(5H)-酮、硫双威、阿维菌素、氟啶虫酰胺、啶虫丙醚、螺甲螨酯、氟啶虫胺腈;棉花杀真菌剂:嘧菌酯、联苯吡菌胺、啶酰菌胺、多菌灵、百菌清、铜、环丙唑醇、苯醚甲环唑、醚菌胺、环氧康那唑、咪唑菌酮、氟啶胺、氟吡菌酰胺、氟嘧菌酯、氟唑菌酰胺、异菌脲、吡唑萘菌胺、异噻菌胺、代森锰锌、代森锰(Maneb)、苯氧菌胺、吡噻菌胺、啶氧菌酯、甲代森锌、丙硫菌唑、吡唑醚菌酯、五氯硝基苯(Quintozene)、戊唑醇、氟醚唑(Tetraconazole)、甲基硫菌灵、肟菌酯;大豆除草剂:甲草胺、灭草松(Bentazone)、氟乐灵、氯嘧磺隆(Chlorimuron-Ethyl)、氯酯磺草胺(Cloransulam-Methyl)、/>唑禾草灵、氟磺胺草醚(Fomesafen)、吡氟禾草灵、草甘膦、甲氧咪草烟(Imazamox)、咪唑喹啉酸(Imazaquin)、咪唑乙烟酸(Imazethapyr)、精异丙甲草胺、嗪草酮、二甲戊灵、吡喃草酮、草铵膦;大豆杀虫剂:高三氟氯氰菊酯、灭多威(Methomyl)、吡虫啉、可尼丁、噻虫嗪、噻虫啉、啶虫脒、呋虫胺、氟虫双酰胺、氯虫苯甲酰胺、溴氰虫酰胺、艾克敌、Spinotoram、氨基阿维菌素苯甲酸盐、氟虫腈、乙虫腈、溴氰菊酯、β-氟氯氰菊酯、γ和λ氯氟氰菊酯、4-[[(6-氯吡啶-3-基)甲基](2,2-二氟乙基)氨基]呋喃-2(5H)-酮、螺虫乙酯、Spinodiclofen、杀铃脲、氟啶虫酰胺、硫双威、β-氟氯氰菊酯;大豆杀真菌剂:嘧菌酯、联苯吡菌胺、啶酰菌胺、多菌灵、百菌清、铜、环丙唑醇、苯醚甲环唑、醚菌胺、环氧康那唑、氟啶胺、氟吡菌酰胺、氟嘧菌酯、粉唑醇(Flutriafol)、氟唑菌酰胺、吡唑萘菌胺、异菌脲、异噻菌胺、代森锰锌、代森锰、叶菌唑、苯氧菌胺、腈菌唑、吡噻菌胺、啶氧菌酯、戊唑醇、甲代森锌、丙硫菌唑、吡唑醚菌酯、戊唑醇、氟醚唑、甲基硫菌灵、肟菌酯;甜菜除草 剂:氯草敏(Chloridazon)、双苯胺灵(Desmedipham)、乙氧呋草黄(Ethofumesate)、甲双苯胺灵(Phenmedipham)、野麦畏(Triallate)、二氯吡啶酸、吡氟禾草灵、环草定(Lenacil)、苯嗪草酮(Metamitron)、氯甲喹啉酸(Quinmerac)、噻草酮(Cycloxydim)、氟胺磺隆(Triflusulfuron)、吡喃草酮、喹禾灵(Quizalofop);甜菜杀虫剂:吡虫啉、可尼丁、噻虫嗪、噻虫啉、啶虫脒、呋虫胺、溴氰菊酯、β-氟氯氰菊酯、γ/λ氯氟氰菊酯、4-[[(6-氯吡啶-3-基)甲基](2,2-二氟乙基)氨基]呋喃-2(5H)-酮、七氟菊酯、氯虫苯甲酰胺、Cyaxypyr、氟虫腈、克百威;卡诺拉油菜除草剂:二氯吡啶酸、禾草灵酸、吡氟禾草灵、草铵膦、草甘膦、吡唑草胺(Metazachlor)、氟乐灵、胺苯磺隆(EthaMetsulfuron)、氯甲喹啉酸、喹禾灵、烯草酮、吡喃草酮;卡诺拉油菜杀真菌剂:嘧菌酯、联苯吡菌胺、啶酰菌胺、多菌灵、环丙唑醇、嘧菌环胺、醚菌胺、环氧康那唑、氟啶胺、氟吡菌酰胺、氟嘧菌酯、氟硅唑(Flusilazole)、氟唑菌酰胺、异菌脲、吡唑萘菌胺、缩节胺(Mepiquat-chloride)、叶菌唑、苯氧菌胺、多效唑(Paclobutrazole)、吡噻菌胺、啶氧菌酯、咪鲜胺、丙硫菌唑、吡唑醚菌酯、戊唑醇、甲基硫菌灵、肟菌酯、乙烯菌核利(Vinclozolin);卡诺拉油菜杀虫剂:克百威、噻虫啉、溴氰菊酯、吡虫啉、可尼丁、噻虫嗪、啶虫脒、呋虫胺、β-氟氯氰菊酯、γ和λ氯氟氰菊酯、τ-氟胺氰菊酯(Fluvaleriate)、乙虫腈、艾克敌、Spinotoram、氟虫双酰胺、氯虫苯甲酰胺、溴氰虫酰胺、4-[[(6-氯吡啶-3-基)甲基](2,2-二氟乙基)氨基]呋喃-2(5H)-酮。
提供以下实施例是为了说明而不是为了限制。
实验实施例
实施例1.大豆中Cry14Aa1的表达
表达Cry14Aa1(SEQIDNO:3)的大豆事件是使用含有编码4-羟基苯丙酮酸双加氧酶蛋白(HPPD)抑制剂耐受性除草剂基因(描述于WO2014043435)和Cry14Aa1的基因的构建体,通过农杆菌介导的Thorne大豆植物转化而开发的。野生型Thorne大豆作为非线虫抗性对照。当在大豆植物中表达时,Cry14Aa1减少了与野生型植物相比在根中繁殖的短尾短体线虫的数量。三个独立的事件品系始终给出相同的结果,显示多次重新测试时线虫减少。在所有三个事件中,线虫的数量减少了60-85%。
实施例2.大豆中Cry14Ab1的表达
表达Cry14Ab1(SEQ ID NO:4)的EE-GM4大豆事件是使用含有编码4-羟基苯丙酮酸双加氧酶蛋白(HPPD)抑制剂耐受性除草剂基因和Cry14Ab1的基因的构建体(在WO2014043435中描述),通过农杆菌介导的转化Thorne大豆植物而开发的。野生型Thorne大豆作为非线虫抗性对照。当在大豆植物中表达时,Cry14Ab1减少了与野生型植物相比在根中繁殖的短尾短体线虫的数量。将未转化的Thorne和EE-GM4种子进行发芽并在温室中种植以检查对病灶线虫短尾短体线虫的控制。当2周龄时,将短尾短体线虫(#1500/植物,不同的发育阶段)施用于植物。施用后30天,从根中提取短体线虫并计数。将含有EE-GM4的植物根部中发现的线虫的平均数量与野生型Thorne植物根中发现的短体线虫的平均数量进行比较。与Thorne对照根相比,在含有EE-GM4的植物的根中发现平均少了约80-90%的短体线虫,表明通过大豆事件EE-GM4显著控制病灶线虫。
图1显示了在美国短尾短体线虫温室测定的结果,比较了5个良种大豆品系中的具有EE-GM4的良种品系(一个SCN易感(MG1),一个SCN抗性(PI88788,MG3),一个SCN易感(MG)6.2),一个SCN抗性(Peking,MG 6.2)和一个对SCN易感和SCN抗性的SCN易感(MG 9)的美国大豆品系。大豆植物生长在小锥形盆中并保持在温度变化在25-32℃之间的温室中。从南卡罗来纳州获得并在温室中增加的短尾短体线虫用于接种V2-V3发育阶段的植物。每株植物接种约1500个卵+成虫,每个条目有5株植物。在侵袭三十天后,从根中提取线虫和卵并计数。每个条目在两个独立的实验中进行。虽然SCN易感和SCN抗性的美国大豆品系未显示对短体线虫的控制,但具有EE-GM4的植物显示短体线虫的约85%控制。
图2显示了在巴西短尾短体线虫温室测定的结果,将具有EE-GM4的大豆植物与没有抗性的巴西大豆品系和1个低Rf品系,以及SCN易感和抗性植物比较。大豆品系在小锥形盆中生长并保持在温度变化在25-32℃之间的温室中。从巴西田地获得并在温室中增加的短尾短体线虫用于接种V2-V3发育阶段的植物。每株植物接种大约1000个卵+成虫,每个条目有5株植物。侵袭后三十天,从根中提取线虫和卵并计数。显示的结果来自单个实验。一个巴西大豆品系(BRS 7380),标记为短体线虫的低繁殖因子,显示出短体线虫减少了约89%。具有EE-GM4的植物给出对短体线虫的控制率为~97%。对SCN(rhg1+Rhg4)具有天然抗性的大豆品系不能控制短尾短体线虫。
此外,含有EE-GM4的植物可用于控制根结线虫(RKN),例如南方根结线虫。虽然南方根结线虫的种群不能很好地感染Thorne野生型大豆,但与未转化的Thorne植物相比,使用EE-GM4的Thorne植物显示出平均的RKN卵数/根质量进一步减少。
实施例3.用于植物表达的基因的载体
本发明的编码区与适当的启动子和终止子序列连接,用于在植物中表达。此类序列在本领域中是熟知的。产生和确认启动子-基因-终止子构建体的技术也是本领域熟知的。
在本发明的一个方面,设计并产生合成的DNA序列。这些合成序列相对于亲本序列具有改变的核苷酸序列,但编码与亲本序列基本相同的蛋白质。在一些实施方案中,合成DNA序列包含SEQ ID NO:3或4。
在本发明的另一方面,设计合成基因的修饰形式,使得所得肽靶向植物细胞器,例如内质网或质外体。已知导致融合蛋白靶向植物细胞器的肽序列是本领域已知的。例如,来自白羽扇豆Lupinus albus的酸性磷酸酶基因的N-末端区域(ID GI:14276838,Miller等人(2001)Plant Physiology 127:594-606)在本领域已知导致异源蛋白质的内质网靶向。如果得到的融合蛋白还含有在C末端包含肽N-末端-赖氨酸-天冬氨酸-谷氨酸-亮氨酸(即,“KDEL”基序,SEQ ID NO:7)的内质网保留序列,则融合蛋白将靶向内质网。如果融合蛋白在C末端缺乏内质网靶向序列,则蛋白质将靶向内质网,但最终将被隔绝在质外体中。
因此,该基因编码融合蛋白,该融合蛋白含有融合到本发明的氨基酸序列的N-末端的、来自白羽扇豆Lupinus albus的酸性磷酸酶基因的N-末端31个氨基酸(IDGI:14276838,Miller等,2001,同上),以及C-末端的KDEL(SEQ ID NO:7)序列。因此,预测所得蛋白质在植物细胞中表达后靶向植物内质网。
将上述植物表达盒与合适的植物选择标记组合以帮助选择转化的细胞和组织,并连接到植物转化载体中。这些可包括来自农杆菌介导的转化的二元载体或用于气溶胶或生物射弹转化的简单质粒载体。
在本发明中,包含编码Cry14A(SEQ ID NO:1或2)的合成基因的表达盒与水稻(Oryza sativa)的蔗糖合酶1基因的启动子区(Wang等人(1992)Plant MolecularBiology,19,881-885)或花椰菜花叶病毒35S转录物的启动子区域(Odell等人(1985)Nature 313,810-812)和矮牵牛的叶绿素a/b结合蛋白基因的前导序列(Harpster等人,(1988)Molecular and General Genetics 212,182-190)有效连接。表达盒还包含来自pTiT37的T-DNA的胭脂碱合酶基因的3'非翻译区(Depicker等人,(1982)Journal ofMolecular and Applied Genetics 1,561-573),其有效连接于Cry14序列的3'末端。
实施例4:大豆转化
使用本领域熟知的方法实现大豆转化,例如基本上使用Paz等人(2006),Plantcell Rep.25:206描述的方法,使用根癌农杆菌介导的转化大豆半种子外植体。使用环磺酮作为选择标记鉴定转化体。观察到绿芽的出现,并记录为对除草剂异唑草酮或环磺酮的耐受性的指标。耐受的转基因芽将显示与未用异/>唑草酮或环磺酮处理的野生型大豆芽相当的正常绿化,而用相同量的异/>唑草酮或环磺酮处理的野生型大豆芽将完全漂白。这表明HPPD蛋白的存在使得能够耐受HPPD抑制剂除草剂,如异/>唑草酮或环磺酮。
耐受的绿芽被转移到生根培养基或移植。在适应期后,将生根的小植株转移到温室中。然后用HPPD抑制剂除草剂喷洒含有转基因的植物,例如用100g AI/ha比率的环磺酮或用补充有硫酸铵甲酯菜籽油的300g AI/ha比率的甲基磺草酮。施用后十天,评估由施用除草剂引起的症状,并与在相同条件下在野生型植物上观察到的症状进行比较。
实施例5:棉花T0植物的建立和选择
使用本领域熟知的方法,特别是PCT专利公开WO 00/71733中描述的优选方法,实现棉花转化。将再生的植物转移到温室中。在适应期后,用HPPD抑制剂除草剂喷洒经充分生长的植物,例如相当于补充有硫酸铵和甲酯菜籽油的100或200gAI/ha的环磺酮。喷洒施用后7天,评估由于用除草剂处理引起的症状,并与在相同条件下进行相同处理的野生型棉花植物上观察到的症状进行比较。
说明书中提及的所有出版物和专利申请表示本发明所属领域的技术人员的技术水平。所有出版物和专利申请均通过引用并入本文,其程度如同每个单独的出版物或专利申请被具体和单独地指出为通过引用并入。
尽管为了清楚理解的目的已经通过说明和实施例详细地描述了前述发明,但显而易见的是,可以在所附权利要求的范围内实施某些改变和修改。
Claims (7)
1.一种控制短尾短体线虫(Pratylenchus brachyurus)害虫种群的方法,包括使所述种群与杀线虫有效量的多肽接触,所述多肽为SEQ ID NO:1或2的氨基酸序列,其中所述多肽具有针对所述短尾短体线虫害虫种群的杀线虫活性。
2.权利要求1所述的方法,其中所述多肽为SEQ ID NO:1所示氨基酸序列的多肽。
3.权利要求1所述的方法,其中所述多肽为SEQ ID NO:2所示氨基酸序列的多肽。
4.权利要求1-3任一项所述的方法,其中所述多肽在大豆植物中表达。
5.一种保护植物以对抗短尾短体线虫害虫的方法,包括在植物或其细胞中表达与启动子有效连接的核苷酸序列,所述启动子能够指导所述核苷酸序列在植物细胞中的表达,其中所述核苷酸序列选自:
a)SEQ ID NO:3或4中所示的核苷酸序列;和
b)编码多肽的核苷酸序列,所述多肽为SEQ ID NO:1或2的氨基酸序列,其中所述多肽具有针对所述短尾短体线虫害虫的杀线虫活性。
6.一种增加植物产量的方法,包括在田地生长植物或其种子,所述植物或其种子在其基因组中稳定地掺入DNA构建体,所述DNA构建体包含与启动子有效连接的核苷酸序列,所述启动子能够指导所述核苷酸序列在植物细胞中的表达,其中所述核苷酸序列选自:
a)SEQ ID NO:3或4中所示的核苷酸序列;和
b)编码多肽的核苷酸序列,所述多肽为SEQ ID NO:1或2的氨基酸序列,其中所述多肽具有针对短尾短体线虫害虫的杀线虫活性;
其中所述田地被短尾短体线虫害虫侵袭。
7.权利要求5或6所述的方法,其中所述植物还包含一种或多种编码一种或多种昆虫毒素的核苷酸序列。
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AU2017382305A1 (en) | 2019-07-18 |
MX2019007599A (es) | 2019-12-16 |
CN110225974A (zh) | 2019-09-10 |
EA201991547A1 (ru) | 2020-01-17 |
US20210212323A1 (en) | 2021-07-15 |
CL2019001753A1 (es) | 2019-09-13 |
BR112019012897A2 (pt) | 2020-01-07 |
IL267428A (en) | 2019-08-29 |
US20190364908A1 (en) | 2019-12-05 |
PH12019501447A1 (en) | 2020-03-16 |
JP2020504609A (ja) | 2020-02-13 |
EP3559241A1 (en) | 2019-10-30 |
WO2018119336A1 (en) | 2018-06-28 |
CO2019006843A2 (es) | 2019-10-09 |
CA3047582A1 (en) | 2018-06-28 |
KR20190095411A (ko) | 2019-08-14 |
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