CN110267975A - Axmi669和axmi991毒素基因及其使用方法 - Google Patents
Axmi669和axmi991毒素基因及其使用方法 Download PDFInfo
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- CN110267975A CN110267975A CN201780084071.6A CN201780084071A CN110267975A CN 110267975 A CN110267975 A CN 110267975A CN 201780084071 A CN201780084071 A CN 201780084071A CN 110267975 A CN110267975 A CN 110267975A
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Classifications
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- C—CHEMISTRY; METALLURGY
- C12—BIOCHEMISTRY; BEER; SPIRITS; WINE; VINEGAR; MICROBIOLOGY; ENZYMOLOGY; MUTATION OR GENETIC ENGINEERING
- C12N—MICROORGANISMS OR ENZYMES; COMPOSITIONS THEREOF; PROPAGATING, PRESERVING, OR MAINTAINING MICROORGANISMS; MUTATION OR GENETIC ENGINEERING; CULTURE MEDIA
- C12N15/00—Mutation or genetic engineering; DNA or RNA concerning genetic engineering, vectors, e.g. plasmids, or their isolation, preparation or purification; Use of hosts therefor
- C12N15/09—Recombinant DNA-technology
- C12N15/63—Introduction of foreign genetic material using vectors; Vectors; Use of hosts therefor; Regulation of expression
- C12N15/79—Vectors or expression systems specially adapted for eukaryotic hosts
- C12N15/82—Vectors or expression systems specially adapted for eukaryotic hosts for plant cells, e.g. plant artificial chromosomes (PACs)
- C12N15/8241—Phenotypically and genetically modified plants via recombinant DNA technology
- C12N15/8261—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield
- C12N15/8271—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield for stress resistance, e.g. heavy metal resistance
- C12N15/8279—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield for stress resistance, e.g. heavy metal resistance for biotic stress resistance, pathogen resistance, disease resistance
- C12N15/8286—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield for stress resistance, e.g. heavy metal resistance for biotic stress resistance, pathogen resistance, disease resistance for insect resistance
-
- C—CHEMISTRY; METALLURGY
- C07—ORGANIC CHEMISTRY
- C07K—PEPTIDES
- C07K14/00—Peptides having more than 20 amino acids; Gastrins; Somatostatins; Melanotropins; Derivatives thereof
- C07K14/195—Peptides having more than 20 amino acids; Gastrins; Somatostatins; Melanotropins; Derivatives thereof from bacteria
- C07K14/32—Peptides having more than 20 amino acids; Gastrins; Somatostatins; Melanotropins; Derivatives thereof from bacteria from Bacillus (G)
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Abstract
本发明提供赋予细菌、植物、植物细胞、组织和种子杀农害活性的组合物和方法。提供包含毒素多肽的编码序列的组合物。编码序列可用于DNA构建体或表达盒中,用于在植物和细菌中转化和表达。组合物还包括转化的细菌、植物、植物细胞、组织和种子。特别地,提供分离的毒素核酸分子。另外,包括多核苷酸相应的氨基酸序列,以及特异性结合那些氨基酸序列的抗体。特别地,本发明提供分离的核酸分子,其包含编码SEQ ID NO:2所示氨基酸序列的核苷酸序列,或SEQ ID NO:1所示的核苷酸序列,以及其变体和片段。
Description
相关申请的交叉引用
本申请要求2016年11月23日提交的美国临时申请系列号62/425,669和2016年11月23日提交的美国临时申请系列号62/425,729的权益,其内容通过引用整体并入本文。
参考电子提交的序列表
序列表的官方副本作为ASCII格式的序列表通过EFS-Web以电子方式提交,其文件名为“APA16-6008WOSEQLIST.txt”,创建于2017年10月3日,大小为30kb,并与说明书同时提交。该ASCII格式的文档中包含的序列表是说明书的一部分,并且通过引用整体并入本文。
技术领域
本发明涉及分子生物学领域。本发明提供了编码杀虫蛋白的新基因。这些蛋白和编码它们的核酸序列可用于制备杀虫制剂和生产转基因抗虫植物中。
背景技术
苏云金芽孢杆菌(Bacillus thuringiensis)是形成革兰氏阳性孢子的土壤细菌,特征在于其能够产生对某些目和种的昆虫特别有毒的结晶包涵体,但对植物和其他非靶标生物体无害。由于该原因,包括苏云金芽孢杆菌菌株或其杀虫蛋白的组合物可用作环境可接受的杀虫剂以控制农业农害或用于各种人或动物疾病的昆虫载体。
来自苏云金芽孢杆菌的晶体(Cry)蛋白(δ-内毒素)主要对鳞翅目(Lepidopteran)、半翅目(Hemipteran)、双翅目(Dipteran)和鞘翅目(Coleopteran)幼虫具有有效的杀虫活性。这些蛋白还显示出对膜翅目(Hymenoptera),同翅目(Homoptera)、虱毛目(Phthiraptera)、食毛目(Mallophaga)和壁虱目(Acari)害虫的活性,以及对其他无脊椎动物目,如线虫(Nemathelminthes)、扁虫(Plaiyhelminthes)和肉鞭虫(Sarcomastigorphora)的活性(Feitelson(1993)The Bacillus Thuringiensis familytree.In Advanced Engineered Pesticides,Marcel Dekker,Inc.,New York,N.Y.)。这些蛋白最初主要基于其杀虫活性被分类为CryI至CryV。主要类别是鳞翅目特异的(I)、鳞翅目和双翅目特异的(II)、鞘翅目特异的(III)、双翅目特异的(IV)和线虫特异的(V)和(VI)。这些蛋白进一步分为亚科;每个科中更高度相关的蛋白被指定了分类字母,如Cry1A、Cry1B、Cry1C等。每个分类中甚至更密切相关的蛋白被命名为如Cry1C1、Cry1C2等。
基于氨基酸序列同源性而非昆虫靶标特异性描述了Cry基因的命名法(Crickmore等人(1998)Microbiol.Mol.Biol.Rev.62:807-813)。在该分类中,为每种毒素指定唯一名称,其包括第一等级(阿拉伯数字)、第二等级(大写字母)、第三等级(小写字母)和第四等级(另一阿拉伯数字)。罗马数字被替换为第一等级中的阿拉伯数字。具有低于45%序列同一性的蛋白具有不同的第一等级,第二和第三等级的标准分别为78%和95%。
晶体蛋白在被昆虫中肠摄入和溶解之前不会表现出杀虫活性。摄入的原毒素被昆虫消化道中的蛋白酶水解成活性毒性分子(Hofte and Whiteley(1989)Microbiol.Rev.53:242-255)。该毒素与靶幼虫中肠中的顶端刷状缘受体结合,插入顶膜形成离子通道或孔,导致幼虫死亡。
δ-内毒素通常具有五个保守序列域和三个保守结构域(参见,例如,de Maagd等人(2001)Trends Genetics 17:193-199)。第一个保守的结构域由7个α螺旋组成,其参与膜插入和孔形成。结构域II由以Greek关键构型排列的三个β-片层组成,结构域III由“果冻卷”形式的两个反平行β-片层组成(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。
除了δ-内毒素外,还有其他几种已知的杀虫蛋白毒素。VIP1/VIP2毒素(参见,例如,美国专利5,770,696)是二元杀虫毒素,其通过据信涉及受体介导的胞吞作用及随后的细胞毒性的机制表现出对昆虫的强活性,类似于其他二元(“A/B”)毒素的作用模式。A/B毒素,如VIP、C2、CDT、CST或炭疽杆菌(B.Anthracis)水肿和致死毒素,最初通过特定的受体介导的单体形式“B”组分结合,与靶细胞相互作用。然后这些单体形成同源七聚体。然后“B”七聚体-受体复合物作为对接平台发挥作用,其随后结合并允许酶促“A”组分通过受体介导的胞吞作用转运到胞质中。一旦进入细胞的胞质,“A”组分通过例如G-肌动蛋白的ADP-核糖基化或增加细胞内的环AMP(cAMP)水平来抑制正常细胞功能。参见Barth等人(2004)Microbiol Mol Biol Rev 68:373--402。
基于苏云金芽孢杆菌的杀虫剂的大量使用已经在小菜蛾,Plutella xylostella的田间种群中产生抗性(Ferre和Van Rie(2002)Annu.Rev.Entomol.47:501-533)。最常见的抗性机制是减少毒素与其特定中肠受体的结合。这也可能赋予对具有相同受体的其他毒素的交叉抗性(Ferre和Van Rie(2002))。
考虑到昆虫可导致的破坏性以及控制昆虫农害导致的产率改善,因此不断需要发现新形式的杀虫毒素。
发明概述
本发明提供了向细菌、植物、植物细胞、组织和种子赋予杀农害活性的组合物和方法。组合物包括编码杀农害(pesticidal)和杀昆虫(insectidal)多肽序列的核酸分子、包含这些核酸分子的载体和包含载体的宿主细胞。组合物还包括杀农害多肽序列和针对所述多肽的抗体。核苷酸序列可用于DNA构建体或表达盒中,用于在生物体(包括微生物和植物)中转化和表达。核苷酸或氨基酸序列可以是已经设计用于在生物体(包括但不限于微生物或植物)中表达的合成序列。组合物还包括含有本发明核苷酸序列的细菌、植物、植物细胞、组织和种子。
特别地,本发明提供了编码杀农害蛋白的分离的、重组的和嵌合的核酸分子。另外,本发明涵盖相应于该杀农害蛋白的氨基酸序列。特别地,本发明提供了包含核苷酸序列的分离的、重组的或嵌合的核酸分子以及其生物活性变体和片段,其中所述核苷酸序列编码SEQ ID NO:2或4中所示的氨基酸序列,或所述核苷酸序列示于SEQ ID NO:1或3中。本发明还包括与本发明的核苷酸序列互补或与本发明的序列或其互补序列杂交的核苷酸序列。本发明还提供载体、宿主细胞、植物和种子,其包含本发明的核苷酸序列、或编码本发明的氨基酸序列的核苷酸序列、或其生物活性变体和片段。
本发明提供用于产生本发明的多肽、以及用于使用这些多肽控制或杀死鳞翅目、半翅目、鞘翅目、线虫或双翅目农害的方法。本发明还包括用于检测样品中本发明的核酸和多肽的方法和试剂盒。
本发明的组合物和方法可用于产生具有增强的农害抗性或耐性的生物体。这些生物体和包含该生物体的组合物是农业目的所期望的。本发明的组合物还可用于产生具有杀农害活性的改变或改善的蛋白,或用于检测产品或生物体中杀农害蛋白或核酸的存在。
发明详述
本发明涉及用于调节生物体,特别是植物或植物细胞的农害抗性或耐性的组合物和方法。“抗性”意指,农害(例如昆虫)在摄入本发明的多肽或与本发明的多肽进行其它接触后被杀死。“耐性”意指,农害的运动、摄食、繁殖或其他功能的损害或减少。该方法包括用编码本发明的杀农害蛋白的核苷酸序列转化生物体。特别地,本发明的核苷酸序列可用于制备具有杀农害活性的植物和微生物。因此,提供了转化的细菌、植物、植物细胞、植物组织和种子。本发明组合物可以是芽孢杆菌或其他物种的杀农害核酸和蛋白。序列可用于构建表达载体用于随后转化入感兴趣的生物体,用作分离其他同源(或部分同源)基因的探针,以及用于通过本领域已知的方法产生改变的杀农害蛋白,所述方法如结构域交换或DNA改组,例如与Cry1、Cry2和Cry9内毒素家族成员进行的结构域交换或DNA改组。本发明蛋白可用于控制或杀死鳞翅目、半翅目、鞘翅目、双翅目和线虫农害群,以及用于产生具有杀农害活性的组合物。
“杀农害毒素”或“杀农害蛋白”意指对一种或多种农害具有毒性活性的毒素,或与这种蛋白具有同源性的蛋白,所述农害包括但不限于鳞翅目、双翅目和鞘翅目、或线虫门的成员。杀农害蛋白已经从生物体中分离,所述生物体包括例如芽孢杆菌属(Bacillus sp.)、双酶梭状芽孢杆菌(Clostridium bifermentans)和丽金龟子类芽孢杆菌(Paenibacilluspopilliae)。杀农害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,和比全长序列短的氨基酸序列,其中序列短的原因是使用了替代的下游起始位点或是由于加工产生了具有杀农害活性的较短蛋白。加工可以发生在表达蛋白的生物体中,或者在摄入蛋白后的农害中。
杀农害蛋白包括δ-内毒素。δ-内毒素包括鉴定为cry1至cry72、cyt1和cyt2的蛋白,以及Cyt样毒素。目前有超过250种已知的δ-内毒素种类,具有广泛的特异性和毒性。有关扩展列表,参见Crickmore等人(1998),Microbiol.Mol.Biol.Rev.62:807-813,有关定期更新,参见Crickmore等人(2003)"Bacillus thuringiensis toxin nomenclature,"在www.biols.susx.ac.uk/Home/Neil_Crickmore/Bt/index。
因此,本文提供了新的分离的、重组的或嵌合的核苷酸序列,其赋予杀农害活性。这些核苷酸序列编码与已知的δ-内毒素或二元毒素同源的多肽。还提供了杀农害蛋白的氨基酸序列。由该基因翻译产生的蛋白允许细胞控制或杀死摄入该蛋白的农害。
分离的核酸分子、其变体和片段
本发明的一个方面涉及分离的、重组的或嵌合的核酸分子,其中所述核酸分子包含编码杀农害蛋白和多肽或其生物活性部分的核苷酸序列,或者所述核酸分子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”包括相对于天然多核苷酸或多肽已经进行操作的多核苷酸或多肽,所述操作使得该多核苷酸或多肽与天然存在的多核苷酸或多肽不同(例如,在化学组成或结构上不同)。在另一个实施方案中,“重组”多核苷酸不含在该多核苷酸所源自的生物体的基因组DNA中天然存在的内部序列(即内含子)。这种多核苷酸的典型例子是所谓的互补DNA(cDNA)。
本文使用的分离的、重组的或嵌合的核酸(或DNA)是指不再存在于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在该核酸所源自的生物体的基因组DNA中天然位于该核酸侧翼的序列(即位于该核酸的5'和3'端的序列)(优选蛋白编码序列)。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小于约5kb、4kb、3kb、2kb、1kb、0.5kb或0.1kb的、在衍生该核酸的细胞的基因组DNA中天然位于该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(在本文中也称为“污染蛋白”)的蛋白制剂。在一些实施方案中,本发明的重组核酸相对于SEQ ID NO:1或3包含一个或多个核苷酸取代,或包含其变体或片段。
编码本发明蛋白的核苷酸序列包括SEQ ID NO:1或3所示的序列,及其变体、片段和互补序列。“互补”意指与给定核苷酸序列充分互补的核苷酸序列,使得其可以与给定的核苷酸序列杂交,从而形成稳定的双链体。由这些核苷酸序列编码的杀农害蛋白的相应氨基酸序列示于SEQ ID NO:2或4中。
作为编码杀农害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”意指编码杀农害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农害蛋白的生物活性部分,或者其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,作为编码杀农害蛋白的核苷酸序列的片段的核酸分子,可以包含编码本文公开的杀农害蛋白的全长核苷酸序列中至少约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农害蛋白的全长核苷酸序列中存在的核苷酸数目。“连续”核苷酸意指彼此紧邻的核苷酸残基。本发明核苷酸序列的片段将编码保留杀农害蛋白的生物活性并因此保留杀农害活性的蛋白片段。因此,本发明还涵盖本文公开的多肽的生物学活性片段。“保留活性”意指片段具有杀农害蛋白的杀农害活性的至少约30%、至少约50%、至少约70%、80%、90%、95%或更高。在一个实施方案中,杀农害活性是杀鞘翅目昆虫活性。在另一个实施方案中,杀农害活性是杀鳞翅目昆虫活性。在另一个实施方案中,杀农害活性是杀线虫活性。在另一个实施方案中,杀农害活性是杀双翅目昆虫活性。在另一个实施方案中,杀农害活性是杀半翅目昆虫活性。测量杀农害活性的方法是本领域熟知的。参见例如,Czapla and Lang(1990)J.Econ.Entomol.83:2480-2485;Andrews等人(1988)Biochem.J.252:199-206;Marrone等人(1985)J.of Economic Entomology 78:290-293;和美国专利号5,743,477,所有这些通过引用整体并入本文。
编码杀农害蛋白的核苷酸序列的片段,在编码本发明蛋白的生物活性部分时,将编码本发明的全长杀农害蛋白中至少约15、25、30、50、75、100、125、150、175、200、250、300、350、400、450个连续氨基酸,或直至本发明的全长杀农害蛋白中存在的氨基酸总数。在一些实施方案中,片段是蛋白水解切割的片段。例如,相对于SEQ ID NO:2或4,蛋白水解切割的片段可具有至少约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,本文所涵盖的片段通过去除C-末端结晶结构域而产生,例如通过蛋白水解或通过在编码序列中插入终止密码子而产生。
在各种实施方案中,本发明的核酸包含SEQ ID NO:1或3的简并核酸,其中所述简并核苷酸序列编码与SEQ ID NO:2或4相同的氨基酸序列。
本发明优选的杀农害蛋白由与SEQ ID NO:1或3的核苷酸序列充分相同的核苷酸序列编码,或者杀农害蛋白与SEQ ID NO:2或4中所示的氨基酸序列充分相同。“充分相同”意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、阅读框定位等,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的目的。两个序列之间的百分比同一性是序列共有的相同位置数目的函数(即,百分比同一性=相同位置数目/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:1-4的任一个)上,计算百分比同一性。可以使用与下面描述的那些技术类似的技术,在允许或不允许空位的情况下,确定两个序列之间的百分比同一性。在计算百分比同一性时,通常计数确切匹配。空位,即在比对中在一个序列中存在残基而另一个序列中不存在残基的位置,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.USA 87:2264中公开的,且在Karlin和Altschul(1993)Proc.Natl.Acad.Sci.USA 90:5873-5877中修改的算法。这种算法被并入Altschul等人(1990)J.Mol.Biol.215:403的BLASTN和BLASTX程序中。可以用BLASTN程序进行BLAST核苷酸搜索,其中得分=100,字长=12,以获得与本发明的杀农害样核酸分子同源的核苷酸序列。可以用BLASTX程序进行BLAST蛋白搜索,其中得分=50,字长=3,以获得与本发明的杀农害蛋白分子同源的氨基酸序列。为了获得用于比较目的的带空位比对,可以如Altschul等人(1997)Nucleic Acids Res.25:3389所述,利用Gapped BLAST(BLAST 2.0中)。或者,可使用PSI-Blast进行迭代搜索以检测分子之间的距离关系。参见Altschul等人(1997)同上引文。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,如Vector NTI Program Suite(InvitrogenCorporation,Carlsbad,CA)的ALIGNX模块。在用ClustalW比对氨基酸序列后,可以评估百分比氨基酸同一性。用于分析ClustalW比对的软件程序的非限制性实例是GENEDOCTM。GENEDOCTM(Karl Nicholas)允许评估多种蛋白之间的氨基酸(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。这种算法被合并到ALIGN程序中(版本2.0),其是GCG Wisconsin Genetics SoftwarePackage,版本10(可从Accelrys,Inc.,9685Scranton 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12,空位罚分4。
除非另有说明,否则使用Needleman和Wunsch(1970)J.Mol.Biol.48(3):443-453的算法的GAP版本10,将用于确定序列同一性或相似性,其中使用以下参数:对于核苷酸序列的%同一性和%相似性,使用GAP权重50和长度权重3,以及nwsgapdna.cmp评分矩阵;对于氨基酸序列的%同一性或%相似性,使用GAP权重8和长度权重2,以及BLOSUM62评分程序。也可以使用等同程序。“等同程序”意指,对于讨论中的任何两个序列,当与GAP版本10产生的相应比对比较时,可以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的任何序列比较程序。
本发明还包括变体核酸分子。编码杀农害蛋白的核苷酸序列的“变体”包括编码本文公开的杀农害蛋白但由于遗传密码的简并性而保守地不同的那些序列,以及如上所述充分相同的那些序列。
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,如聚合酶链反应(PCR)和杂交技术,如下所述。变体核苷酸序列还包括合成产生的核苷酸序列,其例如通过使用定点诱变来产生,但仍编码本发明公开的杀农害蛋白,如下所述。本发明包括的变体蛋白是生物活性的,即其继续具有天然蛋白的期望生物活性,即杀农害活性。“保留活性”意指变体具有天然蛋白的杀农害活性的至少约30%、至少约50%、至少约70%、或至少约80%。测量杀农害活性的方法是本领域熟知的。参见例如,Czapla和Lang(1990)J.Econ.Entomol.83:2480-2485;Andrews等人(1988)Biochem.J.252:199-206;Marrone等人(1985)J.of Economic Entomology 78:290-293;和美国专利号5,743,477,所有这些通过引用整体并入本文。
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,从而导致编码的杀农害蛋白的氨基酸序列的变化,而不改变蛋白的生物活性。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,以将一个或多个氨基酸取代、添加或缺失引入编码的蛋白中,来产生分离的核酸分子变体。可以通过标准技术引入突变,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”氨基酸残基是可以在杀农害蛋白的野生型序列中改变而不改变生物活性的残基,而“必需”氨基酸残基是生物活性所需的。“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(例如赖氨酸、精氨酸、组氨酸)、酸性侧链(例如天冬氨酸、谷氨酸)、不带电荷的极性侧链(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(例如丙氨酸、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(例如苏氨酸、缬氨酸、异亮氨酸)和芳香侧链(如酪氨酸、苯丙氨酸、色氨酸、组氨酸)的氨基酸。
δ-内毒素通常具有五个保守序列域和三个保守结构域(参见例如,de Maagd等人(2001)Trends Genetics 17:193-199)。第一保守结构域由7个α螺旋组成,参与膜插入和孔形成。结构域II由以Greek关键构型排列的三个β-片层组成,结构域III由“果冻卷”形式的两个反平行β-片层组成(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。
可以在保留功能的非保守区域中进行氨基酸取代。通常,不会对保守氨基酸残基或存在于保守基序内的氨基酸残基(其中这些残基对蛋白活性是必需的)进行这种取代。保守并且可能是对蛋白活性必需的残基的实例包括,例如,对于在本发明的序列与相似或相关毒素的比对中包含的所有蛋白,在这些蛋白之间相同的残基(例如,同源蛋白比对中相同的残基)。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,对于在本发明的序列与相似或相关毒素的比对中包含的所有蛋白,在这些蛋白之间仅具有保守取代的残基(例如,对于在同源蛋白的比对中包含的所有蛋白,在这些蛋白之间仅具有保守取代的残基)。然而,本领域技术人员将理解,功能变体可以在这些保守残基中具有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来制备变体核苷酸序列,如通过饱和诱变,并且可以筛选所得突变体赋予杀农害活性的能力,以鉴定保留活性的突变体。诱变后,可以重组表达编码的蛋白,并且可以使用标准测定技术测定蛋白的活性。
可以使用如PCR、杂交等方法鉴定相应的杀农害序列,这样的序列与本发明的序列具有实质同一性(例如,在整个参考序列上至少约70%、至少约75%、80%、85%、90%、95%或更高序列同一性)并具有或赋予杀农害活性。参见,例如,Sambrook and Russell(2001)Molecular Cloning:A Laboratory Manual(Cold Spring Harbor Laboratoiy Press,Cold Spring Harbor,NY)and Innis等人(1990)PCR Protocols:A Guide to Methods andApplications(Academic Press,NY)。
在杂交方法中,杀农害核苷酸序列的全部或部分可用于筛选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,同上引文中公开。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标记,或用任何其他可检测标记物,如其他放射性同位素、荧光化合物、酶或酶辅因子标记。可以基于本文公开的编码已知杀农害蛋白的核苷酸序列,通过标记合成寡核苷酸,来制备用于杂交的探针。还可以使用简并引物,所述简并引物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来设计。探针通常包含核苷酸序列区域,该区域在严格条件下与编码本发明杀农害蛋白的核苷酸序列或其片段或变体的至少约12、至少约25、至少约50、75、100、125、150、175或200个连续核苷酸杂交。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Russell,2001,同上引文中,在此引入作为参考。
例如,本文公开的完整杀农害序列或其一个或多个部分可用作能够与相应的杀农害蛋白样序列和信使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,所述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扩增来自所选生物体或样品的相应杀农害序列。该技术可用于从期望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。杂交技术包括铺板DNA文库的杂交筛选(噬斑或菌落;参见例如,Sambrook等人(1989)MolecularCloning:A Laboratory Manual(第2版,Cold Spring Harbor Laboratory Press,ColdSpring Harbor,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核苷酸序列的全部或部分杂交的核苷酸序列(例如,至少约300个核苷酸、至少约400、至少约500、1000、1200、1500、2000、2500、3000、3500、或直至本文公开的核苷酸序列的全长)。可以在严格条件下进行这些序列的杂交。“严格条件”或“严格杂交条件”是指,在该条件下,与其他序列相比,探针与其靶序列以可检测的更大程度杂交(例如,至少比背景高2倍)。严格条件是序列依赖性的,并且在不同情况下有所不同。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,严格条件是以下的条件:其中盐浓度小于约1.5M Na离子,通常约0.01-1.0MNa离子浓度(或其他盐),pH7.0至8.3,且对于短探针(例如,10至50个核苷酸),温度为至少约30℃,对于长探针(例如,大于50个核苷酸),温度为至少约60℃。加入去稳定剂如甲酰胺也可以实现严格条件。示例性的低严格条件包括在37℃用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50至55℃在1X至2X SSC(20X SSC=3.0MNaCl/0.3M柠檬酸三钠)中洗涤。示例性的中等严格条件包括在37℃在40至45%甲酰胺、1.0M NaCl、1%SDS中杂交,以及在55至60℃在0.5X至1X SSC中洗涤。示例性的高严格条件包括在37℃在50%甲酰胺、1M NaCl、1%SDS中杂交,以及在60至65℃在0.1X SSC中洗涤。任选地,洗涤缓冲液可包含约0.1%至约1%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,关键因子是最终洗涤溶液的离子强度和温度。对于DNA-DNA杂合体,Tm可以从Meinkoth和Wahl(1984)Anal.Biochem.138:267-284的公式中估计:Tm=81.5℃+16.6(log M)+0.41(%GC)-0.61(%form)-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,%form是杂交溶液中甲酰胺的百分比,L是杂交体的长度(以碱基对计)。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)。对于每1%的错配,Tm降低约1℃;因此,可以调节Tm、杂交和/或洗涤条件以与具有期望同一性的序列杂交。例如,如果寻找具有>90%同一性的序列,则Tm可以降低10℃。通常,严格条件选定为在确定的离子强度和pH下比具体序列与其互补序列的热熔点(Tm)低约5℃。然而,非常严格条件可以在比热熔点(Tm)低1、2、3或4℃的温度杂交和/或洗涤;中等严格条件可以在比热熔点(Tm)低6、7、8、9或10℃的温度杂交和/或洗涤;低严格条件可以在比热熔点(Tm)低11、12、13、14、15或20℃的温度杂交和/或洗涤。普通技术人员将理解,使用该公式、杂交和洗涤组合物以及期望的Tm,内在地描述了杂交和/或洗涤溶液的严格性的变化。如果期望的错配程度导致Tm小于45℃(水性溶液)或32℃(甲酰胺溶液),优选提高SSC浓度,以便可以使用更高的温度。关于核酸杂交的大量指导参见Tijssen(1993)Laboratory Techniques in Biochemistry and Molecular Biology--Hybridization with Nucleic Acid Probes,Part I,Chapter 2(Elsevier,New York);和Ausubel等人编辑(1995)Current Protocols in Molecular Biology,第2章(GreenePublishing and Wiley-Interscience,New York)。参见Sambrook等人(1989)MolecularCloning:A Laboratory Manual(第2版,Cold Spring Harbor Laboratory Press,ColdSpring Harbor,New York)。
分离的蛋白及其变体和片段
杀农害蛋白也包括在本发明内。“杀农害蛋白”意指具有SEQ ID NO:2或4所示氨基酸序列的蛋白。还提供了其片段、生物活性部分和变体,并且其片段、生物活性部分和变体可用于实施本发明的方法。“分离蛋白”或“重组蛋白”用于指不再处于其天然环境中的蛋白,例如在体外或重组细菌或植物宿主细胞中。在一些实施方案中,重组蛋白是SEQ ID NO:2或4的变体,其中变体相对于SEQ ID NO:2或4包含至少一个氨基酸取代、缺失或插入。
“片段”或“生物活性部分”包括多肽片段,其包含与SEQ ID NO:2或4中所示的氨基酸序列充分相同的氨基酸序列,并且显示出杀农害活性。杀农害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或更多个氨基酸。这些生物活性部分可以通过重组技术制备并评估杀农害活性。测量杀农害活性的方法是本领域熟知的。参见,例如,Czapla and Lang(1990)J.Econ.Entomol.83:2480-2485;Andrews等人(1988)Biochem.J.252:199-206;Marrone等人(1985)J.of EconomicEntomology 78:290-293;和美国专利号5,743,477,所有这些通过引用整体并入本文。如本文所用,片段包含SEQ ID NO:2或4的至少8个连续氨基酸。然而,本发明包括其他片段,例如蛋白中长度大于约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或更长氨基酸的任何片段。
“变体”意指具有与SEQ ID NO:2或4的氨基酸序列至少约60%、65%、约70%、75%、约80%、85%、约90%、91%、92%、93%、94%、95%、96%、97%、98%或99%相同的氨基酸序列的蛋白或多肽。变体还包括由在严格条件下与SEQ ID NO:1或3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。本发明涵盖的变体蛋白是生物活性的,即它们继续具有所期望的天然蛋白的生物活性,即保留杀农害活性。在一些实施方案中,变体相对于天然蛋白具有改善的活性。测量杀农害活性的方法是本领域熟知的。参见,例如,Czapla and Lang(1990)J.Econ.Entomol.83:2480-2485;Andrews等人(1988)Biochem.J.252:199-206;Marrone等人(1985)J of EconomicEntomology 78:290-293;和美国专利号5,743,477,所有这些通过引用整体并入本文。
细菌基因,如本发明的axmi基因,十分经常地在开放阅读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产生功能性蛋白。这些起始密码子可包括ATG密码子。但是,细菌如芽孢杆菌还将密码子GTG识别为起始密码子,并且在GTG密码子处起始翻译的蛋白在第一个氨基酸处含有甲硫氨酸。在极少数情况下,细菌系统中的翻译可以在TTG密码子处起始,但在这种情况下TTG编码甲硫氨酸。此外,常不能推理确定这些密码子中的哪一个在细菌中天然使用。因此,应理解,使用这些替代的甲硫氨酸密码子之一也可导致产生杀农害蛋白。这些杀农害蛋白包括在本发明中,并可用于本发明的方法中。应当理解,当在植物中表达时,必须将替代性起始密码子改变为ATG以便于正确翻译。
在本发明的各种实施方案中,杀农害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,和由于使用替代的下游起始位点而比全长序列短的氨基酸序列。因此,本发明的核苷酸序列和/或包含本发明核苷酸序列的载体、宿主细胞和植物(以及制备和使用本发明核苷酸序列的方法)可包含编码对应于SEQ ID NO:2或4的氨基酸序列的核苷酸序列。
本发明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;美国专利号4,196,265)。
因此,本发明的一个方面涉及抗体、单链抗原结合分子或其他蛋白,其特异性结合本发明的一种或多种蛋白或肽分子、及其同源物、融合物或片段。在特别优选的实施方案中,抗体特异性结合具有SEQ ID NO:2或4所示氨基酸序列的蛋白或其片段。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:2或4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或包含本发明蛋白的融合蛋白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或肽分子,或用于检测蛋白的翻译后修饰。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或肽分子(特别是由SEQ ID NO:2或4中所示的氨基酸序列编码的蛋白,包括能够特异性结合本发明抗体的其变体或片段)的方法,该方法包括使样品与本发明的抗体接触和确定样品是否含有本发明的蛋白或肽分子。利用抗体检测感兴趣蛋白或肽的方法是本领域已知的。
改变或改善的变体
可以认识到,可以通过各种方法改变杀农害蛋白的DNA序列,并且这些改变可以导致DNA序列编码蛋白,所述蛋白具有与本发明的杀农害蛋白编码的氨基酸序列不同的氨基酸序列。可以以各种方式改变该蛋白,包括SEQ ID NO:2或4的一个或多个氨基酸的氨基酸取代、缺失、截短和插入,包括多至约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、或更多个氨基酸的取代、缺失或插入。用于这种操作的方法通常是本领域已知的。例如,可以通过DNA中的突变制备杀农害蛋白的氨基酸序列变体。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的功能。这些变体将具有期望的杀农害活性。然而,应理解,通过在本发明的组合物上使用这些技术,可以改善杀农害蛋白赋予杀农害活性的能力。例如,可以在宿主细胞中表达杀农害蛋白,所述宿主细胞在DNA复制过程中表现出高的碱基错误并入率,如XL-1Red(Stratagene,LaJolla,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),在非致变菌株中培养杀农害蛋白突变,并鉴定具有杀农害活性的突变基因,例如通过进行测定法以测试杀农害活性。通常,将蛋白混合并用于摄食测定法(feeding assay)。参见,例如Marrone等人(1985)J.of Economic Entomology 78:290-293。此类测定法可包括使植物与一种或多种农害接触并确定植物存活和/或导致农害死亡的能力。导致毒性增加的突变的实例见于Schnepf等人(1998)Microbiol.Mol.Biol.Rev.62:775-806。
或者,可以在氨基或羧基末端对许多蛋白的蛋白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引入的插入、缺失或改变,所述方法如PCR,包括:通过在PCR扩增中使用的寡核苷酸中包含编码氨基酸的序列,通过PCR扩增来改变或延伸编码蛋白的序列。或者,添加的蛋白序列可包括完整的蛋白编码序列,如本领域常用于产生蛋白融合物的那些序列。此类融合蛋白通常用于(1)增加感兴趣蛋白的表达,(2)引入结合结构域、酶活性或表位以促进蛋白纯化、蛋白检测或本领域已知的其他实验用途,(3)使蛋白分泌或翻译靶向亚细胞器,如革兰氏阴性细菌的周质空间,或真核细胞的内质网,后者常导致蛋白的糖基化。
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农害蛋白编码区来产生具有期望特性的新的杀农害蛋白。以这种方式,由一组包含序列区的相关序列多核苷酸,产生重组多核苷酸文库,所述序列区具有实质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,可以在本发明的杀农害基因和其他已知的杀农害基因之间改组编码感兴趣结构域的序列基序,以获得编码具有改善的目的特性的蛋白的新基因,如增加的杀农害活性。这种DNA改组策略在本领域中是已知的。参见,例如,Stemmer(1994)Proc.Natl.Acad.Sci.USA91:10747-10751;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农害蛋白的另一种机制。可以在杀农害蛋白之间交换结构域,从而产生具有改善的杀农害活性或目标谱的杂合或嵌合毒素。产生重组蛋白并测试其杀农害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,可以使用下列方法中的一种或多种来获得变体核苷酸和/或氨基酸序列:易错PCR、寡核苷酸定向诱变、装配PCR、有性PCR诱变、体内诱变、盒诱变、递归整体诱变、指数整体诱变、位点特异性诱变、基因重排、基因位点饱和诱变、排列诱变(permutational mutagenesis)、合成连接重排(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(gapped duplex mutagenesis)、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。
载体
本发明的杀农害序列可以表达盒中提供,用于在目的宿主细胞例如,植物细胞或微生物中的表达。“植物表达盒”意指能够在植物细胞中导致从开放阅读框表达蛋白的DNA构建体。通常其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(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”意指已知或怀疑可以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,其通常涉及向高尔基体中的分泌,以及由此导致的一定糖基化。细菌的杀昆虫毒素常作为原毒素合成,其在目标农害的肠道中通过原蛋白水解(protolytically)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运至亚细胞器的氨基酸序列。因此,这包括通过进入内质网、进入液泡、质体(包括叶绿体、线粒体)等进行靶向转运和/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可操作地连接。
“植物转化载体”意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载体编码所有必需的顺式和反式作用功能,用于转化植物细胞(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并入、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可操作地连接的5'和/或3'调节序列。“可操作地连接”意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,可操作地连接意指连接的核酸序列是连续的,并且当需要连接两个蛋白编码区时,是连续的并且在相同的阅读框中。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(如微生物宿主细胞或植物宿主细胞)中表达的异源启动子可操作地连接。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(一个或多个)。
在各种实施方案中,本发明的核苷酸序列与能够指导核苷酸序列在细胞中(如在植物细胞或微生物中)表达的异源启动子可操作地连接。“启动子”是指用于指导下游编码序列转录的核酸序列。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。
这种表达盒提供有多个限制性位点,可以用于将杀农害序列插入在调节区的转录调节下。
表达盒在转录的5'-3'方向包括转录和翻译起始区(即启动子)、本发明的DNA序列、以及在植物中发挥功能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同源的,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。当启动子对植物宿主是“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可操作地连接的DNA序列的天然或自然存在的启动子。启动子可以是诱导型或组成型的。其可以是天然存在的,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。启动子设计的指导可以通过启动子结构的研究来提供,如Harleyand 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,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)Nature 313:810-812);Kay等人(1987)Science 236:1299-1302中描述的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);来自基因的启动子,如稻肌动蛋白(McElroy等人(1990)Plant Cell 2:163-171和美国专利5,641,876);泛素(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中描述的泛素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)。
用于植物的合适的诱导型启动子包括:来自ACE1系统的启动子,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诱导(Schena等人(1991)Proc.Natl.Acad.Sci.USA 88:10421),或最近应用的嵌合转录激活物XVE,用于由雌二醇激活的基于雌激素受体的诱导型植物表达系统(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EP332104、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和部分或全部合成的启动子的部分组成的启动子。参见,例如,Ni等人(1995)Plant J.7:661-676和PCT WO 95/14098,其描述了用于植物的这种启动子。
在本发明的一个实施方案中,对植物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农害蛋白,如对种子特异的启动子(Datla,R等人,1997,BiotechnologyAnn.Rev.3,269-296),特别是油菜籽蛋白启动子(EP 255 378A1)、菜豆蛋白启动子、谷蛋白启动子、向日葵蛋白启动子(WO92/17580)、白蛋白启动子(WO98/45460)、油质蛋白启动子(W098/45461)、SAT1启动子或SAT3启动子(PCT/US98/06978)。
还可以使用的诱导型启动子有利地选自苯丙氨酸解氨酶(PAL)、HMG-CoA还原酶(HG)、几丁质酶、葡聚糖酶、蛋白酶抑制剂(PI)、PR1家族基因、胭脂碱合酶(nos)和vspB启动子(US 5 670 349,表3)、HMG2启动子(US 5 670349)、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,包括相继地使用。
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。与不包含增强子元件的启动子相比,含有增强子元件的启动子提供更高水平的转录。适用于植物的增强子元件包括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)的翻译激活子、或Carrington&Freed 1990,J.Virol.64:1590-1597中描述的烟草蚀刻病毒(TEV)的翻译激活子,或内含子,如玉米的adhl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,如叶绿体(或其他质体)、内质网或高尔基体,或促进目的肽的分泌。例如,可以工程化构建体使其含有信号肽以促进肽向内质网的转移。“信号序列”意指已知或怀疑可以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,其通常涉及向高尔基体的分泌,以及产生的一定糖基化。“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运至亚细胞器的氨基酸序列。因此,这包括通过进入内质网、进入液泡、质体(包括叶绿体、线粒体)等的靶向转运和/或糖基化的前导序列。还优选工程化植物表达盒以含有内含子,从而为了表达需要进行内含子的mRNA加工。
“3'非翻译区”意指位于编码序列下游的多核苷酸。能够影响多腺苷酸束向mRNA前体3'末端添加的聚腺苷酸化信号序列和其他编码调节信号的序列是3'非翻译区。“5'非翻译区”意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。
终止区可以相对于转录起始区是天然的,可以相对于可操作地连接的目的DNA序列是天然的,可以相对于植物宿主是天然的,或者可以来自另一个来源(即,对于启动子、目的DNA序列、植物宿主或其任何组合而言,是外来的或异源的)。方便的终止区可从根瘤农杆菌(A.Tumefaciens)的Ti质粒获得,如章鱼碱合酶和胭脂碱合酶终止区。还参见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;Balias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,可以优化一种或多种基因以增加在转化的宿主细胞中的表达(合成的DNA序列)。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,或者可以以宿主优选的密码子使用频率来使用密码子以合成基因。合成DNA序列的开放阅读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好、改变或改善GC含量、去除或改变替代阅读框,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、聚腺苷酸化位点、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讨论了宿主优选的密码子使用。用于合成植物优选基因的方法可在本领域获得。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)NucleicAcids Res.17:477-498,通过引用并入本文。
也可以利用合成的DNA序列向DNA序列导入其他改善,如导入内含子序列、产生DNA序列以表达为与细胞器靶向序列的蛋白融合物,细胞器靶向序列为例如叶绿体转运肽、质体/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,杀农害蛋白靶向叶绿体用于表达。以这种方式,当杀农害蛋白不直接插入叶绿体中时,表达盒将另外含有编码转运肽的核酸,以将杀农害蛋白导向叶绿体。此类转运肽在本领域中是已知的。参见,例如,Von Heijne等人(1991)Plant Mol.Biol.Rep.9:104-126;Clark等人(1989)JBiol.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)Science 233:478-481。
可以优化待靶向叶绿体的杀农害基因以用于在叶绿体中表达,以解决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的核酸。参见,例如,美国专利号5,380,831,其通过引用并入本文。
植物转化
本发明的方法包括将核苷酸构建体导入植物中。“导入”意指以使构建体进入植物细胞内部的方式向植物呈递核苷酸构建体。本发明的方法不需要使用将核苷酸构建体导入植物的特定方法,只需要使核苷酸构建体进入植物的至少一个细胞的内部。将核苷酸构建体导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”意指整株植物、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“转化的植物”或“稳定转化的”植物或细胞或组织是指,植物已经并入或整合了外源核酸序列或DNA片段到植物细胞中。这些核酸序列包括外源的或不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,核酸序列相对于该核酸序列所在的天然基因组的细胞或部分不是内源的,该核酸序列通过感染、转染、显微注射、电穿孔、微弹射(microprojection)等添加到该细胞中。
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,本发明的蛋白或核苷酸序列有利地在植物中与编码蛋白或RNA的其他基因组合,所述蛋白或RNA赋予这些植物有用的农艺特性。在编码赋予转化的植物有用的农艺特性的蛋白或RNA的基因中,可以提及编码如下蛋白和RNA的DNA序列:赋予对一种或多种除草剂耐性的蛋白,赋予对某些昆虫耐性的蛋白、赋予对某些疾病耐性的蛋白,可以提供线虫或昆虫控制的RNA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。
在编码赋予转化的植物细胞和植物对某些除草剂耐性的蛋白的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性;编码合适的EPSPS的基因,其赋予对具有EPSPS作为靶标的除草剂的耐性,如草甘膦及其盐(US 4,535,060、US 4,769,061、US 5,094,945、US 4,940,835、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-乙酰基转移酶的基因(例如,US 8,222,489、US8,088,972、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,HPPD抑制剂耐性基因,描述于WO 2004/055191、WO199638567、US 6791014、WO2011/068567、WO2011/076345、WO2011/085221、WO2011/094205、WO2011/068567、WO2011/094199、WO2011/094205、WO2011/145015、WO2012/056401和WO2014/043435)。
在编码合适的EPSPS(其赋予对具有EPSPS作为靶标的除草剂耐性)的DNA序列中,更特别地提及:编码植物EPSPS的基因,特别是编码玉米EPSPS,特别是包含两个突变的玉米EPSPS,特别是包含氨基酸位置102的突变和氨基酸位置106的突变(WO2004/074443),其描述于专利申请US 6566587,下文称为双突变体玉米EPSPS或2mEPSPS;或编码从农杆菌(Agrobacterium)分离的EPSPS的基因,其在美国专利5,633,435中描述为序列ID No.2和序列ID No.3,也称为CP4。
在编码赋予对具有EPSPS作为靶标的除草剂耐性的合适EPSPS的DNA序列中,更具体地提及如下基因,所述基因编码来自球形节杆菌(Arthrobacter globiformis)的EPSPSGRG23,以及突变体GRG23 ACE1、GRG23 ACE2、或GRG23ACE3,特别是WO2008/100353中描述的GRG23的突变体或变体,如WO2008/100353中SEQ ID No.29的GRG23(ace3)R173K。
在编码EPSPS、更具体地编码上述基因的DNA序列的情况下,编码这些酶的序列有利地之前具有编码转运肽的序列,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性性状还可以包括: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)耐性的2,4-D-单加氧酶的基因(美国专利6,153,401);和编码通过代谢赋予对麦草畏(3,6-二氯-2-甲氧基苯甲酸)耐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,本发明的核酸可以与一种或多种除草剂耐性基因堆叠,所述除草剂耐性基因包括一种或多种HPPD抑制剂除草剂耐性基因,和/或一种或多种耐受草甘膦和/或草铵膦的基因。
在编码与昆虫耐性特性相关的蛋白的DNA序列中,更具体地提及文献中广泛描述的并且是本领域技术人员公知的Bt蛋白。还可以提及从细菌如光杆状细菌(Photorhabdus)中提取的蛋白(WO97/17432和WO98/08932)。
在这些编码赋予对昆虫的新耐性特性的目的蛋白的DNA序列中,更具体地提及在文献中广泛描述并且本领域技术人员公知的Bt Cry或VIP蛋白。这些蛋白包括Cry1F蛋白或衍生自Cry1F蛋白的杂合体(例如,US6,326,169;US6,281,016;US6,218,188中描述的杂合Cry1A-Cry1F蛋白;或其毒性片段)、Cry1A型蛋白或其毒性片段,优选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EP451878中描述的Cry1Ab或Bt2蛋白或其杀昆虫片段,WO2002/057664中描述的Cry2Ae、Cry2Af或Cry2Ag蛋白或其毒性片段、WO2007/140256中描述的Cry1A.105蛋白(SEQ ID No.7)或其毒性片段、NCBI登录号ABG20428的VIP3Aal9蛋白、NCBI登录号ABG20429的VIP3Aa20蛋白(WO2007/142840中的SEQ ID No.2)、在COT202或COT203棉花事件中产生的VIP3A蛋白(分别为WO2005/054479和WO2005/054480)、如在WO2001/47952中描述的Cry蛋白、如Estruch等人(1996),Proc Natl Acad Sci U S A.28;93(11):5389-94和US 6,291,156中所述的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S.entomophila)或光杆状物种菌株的杀昆虫蛋白,如如WO98/08932中所述的来自光杆状菌的Tc-蛋白(例如,Waterfield等人,2001,Appl EnvironMicrobiol.67(11):5017-24;Ffrench-Constant and Bowen,2000,Cell Mol Life Sci.;57(5):828-33)。此外,本文包括这些蛋白之任何一种的任何变体或突变体,其在一些(1-10个,优选1-5个)氨基酸上与任何上述序列不同,特别是其毒性片段的序列,或与转运肽如质体转运肽或其他蛋白或肽融合的序列。
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期望性状的基因组合,如除草剂耐性、昆虫耐性、耐旱性、线虫控制、水利用效率、氮利用效率、改善的营养值、疾病抗性、改善的光合作用、改善的纤维质量、胁迫耐性、改善的繁殖等。
可以与本发明的基因在相同物种植物中组合的特别有用的转基因事件(例如,通过交叉育种(crossing)或通过用本发明的嵌合基因再转化含有另一转基因事件的植物),包括事件531/PV-GHBK04(棉花,昆虫控制,描述于WO2002/040677中);事件1143-14A(棉花,昆虫控制,未保藏,描述于WO2006/128569中);事件1143-51B(棉花,昆虫控制,未保藏,描述于WO2006/128570中);事件1445(棉花,除草剂耐性,未保藏,描述于US-A 2002-120964或WO2002/034946中);事件17053(稻,除草剂耐性,保藏为PTA-9843,描述于WO2010/117737中);事件17314(稻,除草剂耐性,保藏为PTA-9844,描述于WO2010/117735中);事件281-24-236(棉花,昆虫控制-除草剂耐性,保藏为PTA-6233,描述于WO2005/103266或US-A 2005-216969中);事件3006-210-23(棉花,昆虫控制-除草剂耐性,保藏为PTA-6233,描述于US-A2007-143876或WO2005/103266中);事件3272(玉米,品质性状,保藏为PTA-9972,描述于WO2006/098952或US-A 2006-230473中);事件33391(小麦,除草剂耐性,保藏为PTA-2347,描述于WO2002/027004中);事件40416(玉米,昆虫控制-除草剂耐性,保藏为ATCC PTA-11158,描述于WO11/075593中);事件43A47(玉米,昆虫控制-除草剂耐性,保藏为ATCC PTA-11509,描述于WO2011/075595中);事件5307(玉米,昆虫控制,保藏为ATCC PTA-9561,描述于WO2010/077816中);事件ASR-368(翦股颖,除草剂耐性,保藏为ATCC PTA-4816,描述于US-A 2006-162007或WO2004/053062中);事件B16(玉米,除草剂耐性,未保藏,描述于US-A2003-126634中);事件BPS-CV127-9(大豆,除草剂耐性,保藏为NCIMB No.41603,描述于WO2010/080829中);事件BLR1(油菜,雄性不育恢复,保藏为NCIMB 41193,描述于WO2005/074671中);事件CE43-67B(棉花,昆虫控制,保藏为DSM ACC2724,描述于US-A 2009-217423或WO2006/128573中);事件CE44-69D(棉花,昆虫控制,未保藏,描述于US-A 2010-0024077中);事件CE44-69D(棉花,昆虫控制,未保藏,描述于WO2006/128571中);事件CE46-02A(棉花,昆虫控制,未保藏,描述于WO2006/128572中);事件COT102(棉花,昆虫控制,未保藏,描述于US-A 2006-130175或WO2004/039986中);事件COT202(棉花,昆虫控制,未保藏,描述于US-A 2007-067868或WO2005/054479中);事件COT203(棉花,昆虫控制,未保藏,描述于WO2005/054480中);事件DAS21606-3/1606(大豆,除草剂耐性,保藏为PTA-11028,描述于WO2012/033794中);事件DAS40278(玉米,除草剂耐性,保藏为ATCC PTA-10244,描述于WO2011/022469中);事件DAS-44406-6/pDAB8264.44.06.1(大豆,除草剂耐性,保藏为PTA-11366,描述于WO2012/075426中);事件DAS-14536-7/pDAB8291.45.36.2(大豆,除草剂耐性,保藏为PTA-11335,描述于WO2012/075429中);事件DAS-59122-7(玉米,昆虫控制-除草剂耐性,保藏为ATCC PTA11384,描述于US-A 2006-070139中);事件DAS-59132(玉米,昆虫控制-除草剂耐性,未保藏,描述于WO2009/100188中);事件DAS68416(大豆,除草剂耐性,保藏为ATCC PTA-10442,描述于WO2011/066384或WO2011/066360中);事件DP-098140-6(玉米,除草剂耐性,保藏为ATCC PTA-8296,描述于US-A 2009-137395或WO 08/112019中);事件DP-305423-1(大豆,品质性状,未保藏,描述于US-A 2008-312082或WO2008/054747中);事件DP-32138-1(玉米,杂交系统,保藏为ATCC PTA-9158,描述于US-A 2009-0210970或WO2009/103049中);事件DP-356043-5(大豆,除草剂耐性,保藏为ATCC PTA-8287,描述于US-A 2010-0184079或WO2008/002872中);事件EE-1(茄子brinjal,昆虫控制,未保藏,描述于WO 07/091277中);事件FI117(玉米,除草剂耐性,保藏为ATCC 209031,描述于US-A2006-059581或WO 98/044140中);事件FG72(大豆,除草剂耐性,保藏为PTA-11041,描述于WO2011/063413中);事件GA21(玉米,除草剂耐性,保藏为ATCC 209033,描述于US-A2005-086719或WO98/044140中);事件GG25(玉米,除草剂耐性,保藏为ATCC 209032,描述于US-A2005-188434或WO 98/044140中);事件GHB119(棉花,昆虫控制-除草剂耐性,保藏为ATCCPTA-8398,描述于WO2008/151780中);事件GHB614(棉花,除草剂耐性,保藏为ATCC PTA-6878,描述于US-A 2010-050282或WO2007/017186中);事件GJ11(玉米,除草剂耐性,保藏为ATCC 209030,描述于US-A 2005-188434或WO98/044140中);事件GM RZ13(甜菜,病毒抗性,保藏为NCIMB-41601,描述于WO2010/076212中);事件H7-1(甜菜,除草剂耐性,保藏为NCIMB41158或NCIMB 41159,描述于US-A 2004-172669或WO 2004/074492中);事件JOPLIN1(小麦,疾病耐性,未保藏,描述于US-A 2008-064032中);事件LL27(大豆,除草剂耐性,保藏为NCIMB41658,描述于WO2006/108674或US-A 2008-320616中);事件LL55(大豆,除草剂耐性,保藏为NCIMB 41660,描述于WO 2006/108675或US-A 2008-196127中);事件LLcotton25(棉花,除草剂耐性,保藏为ATCC PTA-3343,描述于WO2003/013224或US-A 2003-097687中);事件LLRICE06(稻,除草剂耐性,保藏为ATCC 203353,描述于US 6,468,747或WO2000/026345中);事件LLRice62(稻,除草剂耐性,保藏为ATCC 203352,描述于WO2000/026345中);事件LLRICE601(稻,除草剂耐性,保藏为ATCC PTA-2600,描述于US-A 2008-2289060或WO2000/026356中);事件LY038(玉米,品质性状,保藏为ATCC PTA-5623,描述于US-A 2007-028322或WO2005/061720中);事件MIR162(玉米,昆虫控制,保藏为PTA-8166,描述于US-A 2009-300784或WO2007/142840中);事件MIR604(玉米,昆虫控制,未保藏,描述于US-A 2008-167456或WO2005/103301中);事件MON15985(棉花,昆虫控制,保藏为ATCC PTA-2516,描述于US-A 2004-250317或WO2002/100163中);事件MON810(玉米,昆虫控制,未保藏,描述于US-A 2002-102582中);事件MON863(玉米,昆虫控制,保藏为ATCC PTA-2605,描述于WO2004/011601或US-A 2006-095986中);事件MON87427(玉米,授粉控制,保藏为ATCC PTA-7899,描述于WO2011/062904中);事件MON87460(玉米,胁迫耐性,保藏为ATCC PTA-8910,描述于WO2009/111263或US-A 2011-0138504中);事件MON87701(大豆,昆虫控制,保藏为ATCCPTA-8194,描述于US-A 2009-130071或WO2009/064652中);事件MON87705(大豆,品质性状-除草剂耐性,保藏为ATCC PTA-9241,描述于US-A 2010-0080887或WO2010/037016中);事件MON87708(大豆,除草剂耐性,保藏为ATCC PTA-9670,描述于WO2011/034704中);事件MON87712(大豆,产率,保藏为PTA-10296,描述于WO2012/051199中);事件MON87754(大豆,品质性状,保藏为ATCC PTA-9385,描述于WO2010/024976中);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中);事件MON88017(玉米,昆虫控制-除草剂耐性,保藏为ATCC PTA-5582,描述于US-A 2008-028482或WO2005/059103中);事件MON88913(棉花,除草剂耐性,保藏为ATCC PTA-4854,描述于WO2004/072235或US-A 2006-059590中);事件MON88302(油菜,除草剂耐性,保藏为PTA-10955,描述于WO2011/153186中);事件MON88701(棉花,除草剂耐性,保藏为PTA-11754,描述于WO2012/134808中),事件MON89034(玉米,昆虫控制,保藏为ATCC PTA-7455,描述于WO07/140256或US-A 2008-260932中);事件MON89788(大豆,除草剂耐性,保藏为ATCC PTA-6708,描述于US-A 2006-282915或WO2006/130436中);事件MS 11(油菜,授粉控制-除草剂耐性,保藏为ATCC PTA-850或PTA-2485,描述于WO2001/031042中);事件MS8(油菜,授粉控制-除草剂耐性,保藏为ATCC PTA-730,描述于WO2001/041558或US-A 2003-188347中);事件NK603(玉米,除草剂耐性,保藏为ATCC PTA-2478,描述于US-A 2007-292854中);事件PE-7(稻,昆虫控制,未保藏,描述于WO2008/114282中);事件RF3(油菜,授粉控制-除草剂耐性,保藏为ATCC PTA-730,描述于WO2001/041558或US-A 2003-188347中);事件RT73(油菜,除草剂耐性,未保藏,描述于WO2002/036831或US-A 2008-070260中);事件SYHT0H2/SYN-000H2-5(大豆,除草剂耐性,保藏为PTA-11226,描述于WO2012/082548中),事件T227-1(甜菜,除草剂耐性,未保藏,描述于WO2002/44407或US-A 2009-265817中);事件T25(玉米,除草剂耐性,未保藏,描述于US-A 2001-029014或WO2001/051654中);事件T304-40(棉花,昆虫控制-除草剂耐性,保藏为ATCC PTA-8171,描述于US-A 2010-077501或WO2008/122406中);事件T342-142(棉花,昆虫控制,未保藏,描述于WO2006/128568中);事件TC1507(玉米,昆虫控制-除草剂耐性,未保藏,描述于US-A 2005-039226或WO2004/099447中);事件VIP1034(玉米,昆虫控制-除草剂耐性,保藏为ATCC PTA-3925,描述于WO2003/052073中);事件32316(玉米,昆虫控制-除草剂耐性,保藏为PTA-11507,描述于WO2011/084632中);事件4114(玉米,昆虫控制-除草剂耐性,保藏为PTA-11506,描述于WO2011/084621中);事件EE-GM3/FG72(大豆,除草剂耐性,ATCC登录号PTA-11041),任选地与以下事件堆叠:事件EE-GM1/LL27或事件EE-GM2/LL55(WO2011/063413A2),事件DAS-68416-4(大豆,除草剂耐性,ATCC登录号PTA-10442,WO2011/066360A1),事件DAS-68416-4(大豆,除草剂耐性,ATCC登录号PTA-10442,WO2011/066384A1),事件DP-040416-8(玉米,昆虫控制,ATCC登录号PTA-11508,WO2011/075593A1),事件DP-043A47-3(玉米,昆虫控制,ATCC登录号PTA-11509,WO2011/075595A1),事件DP-004114-3(玉米,昆虫控制,ATCC登录号PTA-11506,WO2011/084621A1),事件DP-032316-8(玉米,昆虫控制,ATCC登录号PTA-11507,WO2011/084632A1),事件MON-88302-9(油菜,除草剂耐性,ATCC登录号PTA-10955,WO2011/153186A1),事件DAS-21606-3(大豆,除草剂耐性,ATCC登录号PTA-1 1028,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,堆叠的除草剂耐性,ATCC登录号,PTA-11336,WO2012/075426A1),事件DAS-14536-7(大豆,堆叠的除草剂耐性,ATCC登录号:PTA-11335,WO2012/075429A1),事件SYN-000H2-5(大豆,除草剂耐性,ATCC登录号:PTA-11226,WO2012/082548A2),事件DP-061061-7(油菜,除草剂耐性,无可获得的保藏号,WO2012071039A1),事件DP-073496-4(油菜,除草剂耐性,无可获得的保藏号,US2012131692),事件8264.44.06.1(大豆,堆叠的除草剂耐性,登录号PTA-11336,WO2012075426A2),事件8291.45.36.2(大豆,堆叠的除草剂耐性,登录号PTA-11335,WO2012075429A2),事件SYHT0H2(大豆,ATCC登录号PTA-11226,WO2012/082548A2),事件MON88701(棉花,ATCC登录号PTA-11754,WO2012/134808A1),事件KK179-2(苜蓿,ATCC登录号PTA-11833,WO2013/003558A1),事件pDAB8264.42.32.1(大豆,堆叠的除草剂耐性,ATCC登录号PTA-11993,WO2013/010094A1),事件MZDT09Y(玉米,ATCC登录号PTA-13025,WO2013/012775A1)。
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农害基因以获得或增强植物细胞中的表达。通常,表达这种蛋白的构建体含有驱动基因转录的启动子,以及允许转录终止和聚腺苷酸化的3'非翻译区。这种构建体的组织在本领域中是公知的。在一些情况下,可能有用的是,工程化基因使得所得肽在植物细胞内分泌或以其他方式靶向。例如,可以工程化基因以含有信号肽以促进肽向内质网的转移。也可以优选工程化植物表达盒以含有内含子,从而为了表达需要进行内含子的mRNA加工。
通常将“植物表达盒”插入“植物转化载体”中。该植物转化载体可以由实现植物转化所需的一种或多种DNA载体组成。例如,本领域的常见做法是利用由多于一个的连续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,由此将功能分开到分开的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(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“感兴趣基因”(经工程化能够在植物细胞中表达的基因,其中期望从所述的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。顺式作用序列以允许有效转移到植物细胞中并在其中表达的方式排列。例如,可选择的标记基因和杀农害基因位于左边界和右边界之间。第二质粒载体常含有介导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,转移DNA,如本领域所理解的(Hellens and Mullineaux(2000)Trends 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转化植物所必需的,所述其他方法如微弹射、微注射、电穿孔、聚乙二醇等。
通常,植物转化方法包括将异源DNA转移到靶植物细胞(例如未成熟或成熟胚、悬浮培养物、未分化的愈伤组织、原生质体等),然后应用最大阈值水平的适当选择(取决于可选择的标记基因)以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,置于补充有最大阈值水平的选择剂的再生培养基上,转化的细胞将分化出芽。然后将芽转移到选择性生根培养基中以回收生根的芽或小植株。然后转基因小植株生长成成熟植物并产生可育种子(例如Hiei等人(1994)ThePlant Journal 6:271-282;Ishida等人(1996)Nature Biotechnology 14:745-750)。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 and Park(1994)Critical Reviews in Plant Science 13:219-239和Bommineni和Jauhar(1997)Maydica 42:107-120。由于转化的物质含有许多细胞;转化的和未转化的细胞都存在于受试靶标愈伤组织或组织或细胞组的任何部分中。杀死未转化细胞并允许转化细胞增殖的能力将导致转化的植物培养物。通常,去除未转化细胞的能力,是快速回收转化植物细胞和成功产生转基因植物的限制因素。
转化方案以及将核苷酸序列导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。可以通过几种方法之一进行转基因植物的产生,包括但不限于微注射、电穿孔、直接基因转移、通过农杆菌将异源DNA导入植物细胞(农杆菌介导的转化)、用粘附于颗粒的异源外来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)、Lecl转化、以及各种其他非粒子直接介导的DNA转移方法。
用于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 and Maliga(1993)Proc.Natl.Acad.Sci.USA 90:913-917;Svab and Maliga(1993)AffiO J 12:601-606。该方法依赖于粒子枪递送含有可选择标记物的DNA并通过同源重组将DNA靶向质体基因组。另外,可通过核编码和质体定向的RNA聚合酶的组织偏好表达,反式激活沉默的质体携带的转基因,来完成质体转化。McBride等人(1994)Proc.Natl.Acad.Sci.USA 91:7301-7305报道了这种系统。
在将异源外来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,并通过定期转移至新鲜培养基分离和增殖从该选择处理中存活的推定转化的细胞。通过连续传代和以适当选择进行攻击,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转基因植物的基因组中整合的异源感兴趣基因的存在。
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,并用相同的转化株或不同株授粉,并鉴定具有期望的表型特征的组成型表达的所得杂合体。可以生长两代或更多代,以确保期望的表型特征的表达被稳定维持和遗传,然后收获种子以确保已经实现期望的表型特征的表达。以这种方式,本发明提供了转化的种子(也称为“转基因种子”),其具有稳定地并入其基因组中的本发明核苷酸构建体,例如,本发明的表达盒。
植物转化的评估
在将异源外来DNA导入植物细胞后,通过各种方法确认异源基因在植物基因组中的转化或整合,如与整合基因相关的核酸、蛋白和代谢物分析。
PCR分析是一种快速的方法,用于在移植到土壤之前的早期阶段筛选转化的细胞、组织或枝中并入基因的存在(Sambrook and Russell(2001)Molecular Cloning:ALaboratory Manual.Cold Spring Harbor;Laboratory Press,Cold Spring Harbor,NY)。使用对感兴趣基因或农杆菌载体背景等特异的寡核苷酸引物进行PCR。
可以通过基因组DNA的Southern印迹分析确认植物转化(Sambrook and 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。然后根据标准技术,用例如放射性标记的32P靶DNA片段探测膜或“印迹”,以证实导入的基因在植物基因组中的整合(Sambrook and Russell,2001,同上引文)。
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 and Russell,2001,同上)。然后通过本领域已知的方法(Sambrook and Russell,2001,同上)、通过将滤膜与源自杀农害基因的放射性探针杂交来测试由杀农害基因编码的RNA的表达。
可以对转基因植物进行蛋白印迹、生物化学测定法等,以通过标准程序(Sambrookand Russell,2001,同上),使用与杀农害蛋白上存在的一个或多个表位结合的抗体来确认由杀农害基因编码的蛋白的存在。
植物中的杀农害活性
在本发明的另一个方面,可以产生表达具有杀农害活性的杀农害蛋白的转基因植物。上述通过实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,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。可以通过本领域描述的常规方法分离表达杀农害蛋白的植物,例如通过转化愈伤组织、选择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可选择标记物,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物,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可选择标记物。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。本领域熟知用于检测植物、植物器官(例如,叶、茎、根等)、种子、植物细胞、繁殖体、其胚胎或后代中转基因的存在的方法。在一个实施方案中,通过测试杀农害活性来检测转基因的存在。
可以测试表达杀农害蛋白的可育植物的杀农害活性,并且选择表现最佳活性的植物用于进一步的育种。本领域中可获得用于测定农害活性的方法。通常,将蛋白混合并用于摄食测定法。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(corn,maize)、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、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。
蔬菜包括但不限于番茄、生菜、青豆、利马豆、豌豆、和黄瓜属(Cucumis)成员,如黄瓜、甜瓜和香瓜。观赏植物包括但不限于杜鹃花、绣球花、芙蓉、玫瑰、郁金香、水仙花、矮牵牛花、康乃馨、一品红和菊花。优选地,本发明的植物是农作物(例如,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、油菜等)。
在杀农害控制中的用途
本领域已知在农害控制或在工程化其他生物体作为杀农害剂中,使用包含本发明核苷酸序列或其变体的株系的一般方法。参见,例如美国专利号5,039,523和EP0480762A2。
含有本发明核苷酸序列或其变体的芽孢杆菌菌株,或经遗传改变以含有本发明的杀农害基因和蛋白的微生物,可用于保护农作物和产品免受农害侵害。在本发明的一个方面,用将细胞应用于一种或多种靶农害环境时,用可以延长细胞中产生的毒素的活性的试剂,处理产生毒素(杀农害剂)的生物体的整个(即未裂解的)细胞。
或者,通过将杀农害基因导入细胞宿主中来产生杀农害剂。杀农害基因的表达直接或间接地导致杀农害剂的细胞内产生和维持。在本发明的一个方面,在将细胞应用于一种或多种靶农害环境时,在延长细胞中产生的毒素的活性的条件下处理这些细胞。所得产物保留毒素的毒性。然后可以根据常规技术配制这些天然包封的杀农害剂,以应用于容纳靶农害的环境,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,以允许将所得物质应用为杀农害剂。
本发明的活性成分通常以组合物的形式应用,并且可以与其他化合物同时或相继应用于待处理的作物区或植物。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、杀农害皂、休眠油、聚合物、和/或时间释放或可生物降解的载体制剂(其允许在单次应用制剂后长期给药靶区域)。如果需要,其还可以是选择性除草剂、化学杀昆虫剂、杀病毒剂、杀微生物剂、杀变形虫剂(amoebicides)、农药、杀真菌剂、杀细菌剂、杀线虫剂、杀软体动物剂或这些制剂中几种的混合物,可以进一步组合通常用于制剂领域的农业上可接受的载体、表面活性剂或促进应用的助剂。合适的载体和助剂可以是固体或液体,并且对应于制剂技术中通常使用的物质,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”或制成农害“诱捕器”,以允许杀农害制剂的靶农害摄食或摄取。
施用本发明的活性成分、或施用含有至少一种由本发明的细菌菌株产生的杀农害蛋白的本发明农业化学组合物的方法,包括叶施用、种子包衣和土壤施用。施用次数和施用率取决于相应农害的侵染强度。
组合物可以配制成粉末、撒粉、丸剂、颗粒、喷雾、乳剂、胶体、溶液等,并且可以通过常规方法制备,例如干燥、冻干、均质、萃取、过滤、离心、沉降、或浓缩包含多肽的细胞培养物等。在所有此类含有至少一种这样的杀农害多肽的组合物中,多肽可以以约1%至约99%重量的浓度存在。
可以通过本发明的方法在给定区域内杀死或减少鳞翅目、半翅目、双翅目或鞘翅目农害的数目,或者可以预防性地将其应用于环境区域以防止易感农害的侵染。优选地,农害摄取杀农害有效量的多肽或与杀农害有效量的多肽接触。“杀农害有效量”意指能够使至少一个农害死亡,或显著减少农害生长、摄食或正常生理发育的杀农害剂量。该量将取决于以下因素,例如待控制的特定靶农害、特定环境、位置、植物、作物或待处理的农业场所、环境条件、以及杀农害有效多肽组合物的应用方法、速率、浓度、稳定性和量。制剂还可以根据气候条件、环境因素、和/或应用频率和/或农害侵染的严重性而变化。
可以通过将细菌细胞、晶体和/或孢子悬浮液、或分离的蛋白组分与期望的农业上可接受的载体一起配制,来制备期望的杀农害组合物。组合物可在施用前以适当的方式配制,如冻干、冷冻干燥、干燥、或在水性载体、培养基或合适的稀释剂如盐水或其他缓冲液中配制。配制的组合物可以是撒粉剂或颗粒物质、或油(植物或矿物质)悬浮液、或水或油/水乳剂、或可湿性粉末、或与任何其他适用于农业应用的载体物质组合。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”包括通常用于杀农害制剂技术的所有助剂、惰性组分、分散剂、表面活性剂、增粘剂、粘合剂等;这些是杀农害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助剂混合,并通过各种方法制备,例如通过使用常规制剂技术将杀农害组合物与合适的助剂均匀混合、共混和/或研磨。合适的制剂和应用方法描述于美国专利号No.6,468,523中,其通过引用并入本文。
“农害”包括但不限于昆虫、真菌、细菌、线虫、螨虫,蜱等。昆虫农害包括选自鞘翅目(Coleoptera)、双翅目(Diptera)、膜翅目(Hymenoptera)、鳞翅目(Lepidoptera)、食毛目(Mallophaga)、同翅目(Homoptera)、半翅目(Hemiptera)、直翅目(Orthroptera)、缨翅目(Thysanoptera)、革翅目(Dermaptera)、等翅目(Isoptera)、虱目(Anoplura)、蚤目(Siphonaptera)、毛翅目(Trichoptera)等的昆虫,特别是鞘翅目、鳞翅目和双翅目。
鞘翅目包括肉食亚目(Adephaga)和多食亚目(Polyphaga)。肉食亚目包括超家族步甲总科(Caraboidea)和豉甲总科(Gyrinoidea),而多食亚目包括超家族水龟甲总科(Hydrophiloidea)、隐翅虫总科(Staphylinoidea)、花萤总科(Cantharoidea)、郭公甲总科(Cleroidea)、叩甲总科(Elateroidea)、花甲总科(Dascilloidea)、泥甲总科(Dryopoidea)、丸甲总科(Byrrhoidea)、扁甲总科(Cucujoidea)、芫菁总科(Meloidea)、Mordelloidea、拟步甲总科(Tenebrionoidea)、长蠹总科(Bostrichoidea)、金龟总科(Scarabaeoidea)、天牛总科(Cerambycoidea)、叶甲总科(Chrysomeloidea)和象甲总科(Curculionoidea)。超家族步甲总科包括家族虎甲科(Cicindelidae)、步甲科(Carabidae)和龙虱科(Dytiscidae)。超家族豉甲总科包括家族豉甲科(Gyrinidae)。超家族水龟甲总科包括家族水龟甲科(Hydrophilidae)。超家族隐翅虫总科包括家族埋葬甲科(Silphidae)和隐翅虫科(Staphylinidae)。超家族花萤总科包括家族花萤科(Cantharidae)和萤科(Lampyridae)。超家族郭公甲总科包括家族郭公甲科(Cleridae)和皮蠹科(Dermestidae)。超家族叩甲总科包括家族叩甲科(Elateridae)和吉丁虫科(Buprestidae)。超家族扁甲总科包括家族瓢虫科(Coccinellidae)。超家族芫菁总科包括家族芫菁科(Meloidae)。超家族拟步甲总科包括家族拟步甲虫科(Tenebrionidae)。超家族金龟总科包括家族黑蜣科(Passalidae)和金龟科(Scarabaeidae)。超家族天牛总科包括家族天牛科(Cerambycidae)。超家族叶甲总科包括家族叶甲科(Chrysomelidae)。超家族象甲总科包括家族象甲科(Curculionidae)和小蠹科(Scolytidae)。
双翅目包括长角亚目(Nematocera)、短角亚目(Brachycera)和环裂亚目(Cyclorrhapha)。长角亚目包括家族大蚊科(Tipulidae)、蛾蚋科(Psychodidae)、蚊科(Culicidae)、蠓科(Ceratopogonidae)、摇蚊科(Chironomidae)、蚋科(Simuliidae)、毛蚋科(Bibionidae)和瘿蚊科(Cecidomyiidae)。短角亚目包括家族水虻科(Stratiomyidae)、虻科(Tabanidae)、剑虻科(Therevidae)、食虫虻科(Asilidae)、拟食虫虻科(Mydidae)、蜂虻科(Bombyliidae)和长足虻科(Dolichopodidae)。环裂亚目包括无缝组(Aschiza)和有缝组。无缝组包括家族蚤蝇科(Phoridae)、食蚜蝇科(Syrphidae)和眼蝇科(Conopidae)。有缝组包括无瓣类(Acalyptratae)和有瓣类(Calyptratae)。无瓣类包括家族斑蝇科(Otitidae)、实蝇科(Tephritidae)、潜蝇科(Agromyzidae)和果蝇科(Drosophilidae)。有瓣类包括家族虱蝇科(Hippoboscidae)、狂蝇科(Oestridae)、寄蝇科(Tachinidae)、花蝇科(Anthomyiidae)、蝇科(Muscidae)、丽蝇科(Calliphoridae)和麻蝇科(Sarcophagidae)。
鳞翅目包括家族凤蝶科(Papilionidae)、粉蝶科(Pieridae)、灰蝶科(Lycaenidae)、蛱蝶科(Nymphalidae)、斑蝶科(Danaidae)、眼蝶科(Satyridae)、弄蝶科(Hesperiidae)、天蛾科(Sphingidae)、天蚕蛾科(Saturniidae)、尺蛾科(Geometridae)、灯蛾科(Arctiidae)、夜蛾科(Noctuidae)、毒蛾科(Lymantriidae)、透翅蛾科(Sesiidae)和谷蛾科(Tineidae)。
线虫包括线虫寄生虫,如根结、胞囊和病变线虫(lesion nematode),包括异皮线虫属(Heterodera spp.)、根结线虫属(Meloidogyne spp.)和球异皮线虫属(Globoderaspp.);特别地,胞囊线虫的成员包括但不限于Heterodera glycines(大豆胞囊线虫);Heterodera schachtii(甜菜胞囊线虫);Heterodera avenae(禾谷胞囊线虫);和马铃薯金线虫(Globodera rostochiensis)和马铃薯白线虫(Globodera pallida)(马铃薯胞囊线虫)。病变线虫包括短体线虫属(Pratylenchus spp.)。
半翅目农害(包括命名为半翅亚目(Hemiptera)、同翅亚目(Homoptera)或异翅亚目(Heteroptera)的物种)包括但不限于,盲椿属(Lygus spp.),如西方黑斑盲蝽(Lygushesperus)、黑斑盲蝽(Lygus ineolaris)和绿盲蝽(Lygus elisus);蚜虫类,如绿桃蚜(Myzus persicae)、棉蚜(Aphis gossypii)、樱桃蚜虫或黑樱桃蚜虫(Myzus cerasi)、大豆蚜虫(Aphis glycines Matsumura);褐飞虱(Nilaparvata lugens)和稻绿叶蝉(Nephotettix spp.);和臭椿(stink bug),如拟绿蝽(Acrosternum hilare,green stinkbug)、茶翅臭椿(Halyomorpha halys)、南方绿臭椿(Nezara viridula)、稻臭椿(Oebaluspugnax)、森林臭虫(Pentatoma rufipes)、欧洲臭椿(Rhaphigaster nebulosa)和盾椿象Troilus luridus。
针对主要作物的本发明昆虫农害包括:
玉米:玉米螟(Ostrinia nubilalis),欧洲玉米螟(European corn borer);小地老虎(Agrotis ipsilon),黑切根虫(black cutworm);谷实夜蛾(Helicoverpa zea),棉铃虫(cotton bollworm);草地贪夜蛾(Spodoptera frugiperda),秋粘虫(fall armyworm);Diatraea grandiosella,西南玉米螟(southwestern corn borer);南美玉米苗斑螟(Elasmopalpus lignosellus),小玉米茎蛀虫(lesser cornstalk borer);小蔗螟(Diatraea saccharalis),甘蔗螟虫(sugarcane borer);玉米根萤叶甲(D.virgiferavirgifera),西方玉米根虫(western corn rootworm);Diabrotica longicornisbarberi,北方玉米根虫(northern corn rootworm);黄瓜十一星叶甲食根亚种(Diabrotica undecimpunctata howardi),南方玉米根虫(southern corn rootworm);梳爪叩头虫属(Melanotus spp.),金针虫(wireworms);Cyclocephala borealis,北方假面金龟子(northern masked chafer)(白色蛴螬white grub);Cyclocephala immaculata,南方假面金龟子(southern masked chafer)(白色蛴螬white grub);日本弧丽金龟(Popilliajaponica),日本甲虫(Japanese beetle);玉米铜色跳甲(Chaetocnema pulicaria),玉米跳甲(corn leaf beetle);玉米长喙象(Sphenophorus maidis,maize billbug);玉米蚜(Rhopalosiphum maidis),玉米叶蚜虫;Anur aphis maidiradicis,玉米根蚜(corn rootaphid);美洲谷杆长蝽(Blissus leucopterus leucopterus,chinch bug);红足黑蝗(Melanoplus femurrubrum,redleged grasshopper);迁飞黑蝗(Melanoplussanguinipes,migratory grasshopper);灰地种蝇(Hylemya platura,seedcorn maggot);玉米斑潜蝇(Agromyza parvicornis,corn blotch leafminer);黄呆蓟马(Anaphothripsobscrurus,grass thrips);窃蚁(Solenopsis ilesta,thief ant);二点叶螨(Tetranychus urticae,twospotted spider mite);
高粱:Chilo partellus,高粱螟;草地贪夜蛾(Spodoptera frugiperda),秋粘虫;Spodoptera cosmioides;南部灰翅夜蛾(Spodoptera eridania);谷实夜蛾(Helicoverpazea),棉铃虫(cotton bollworm);南美玉米苗斑螟(Elasmopalpus lignosellus),小玉米茎蛀虫;粒肤脏切叶蛾(Feltia subterranea,granulate cutworm);长毛食叶然金龟(Phyllophaga crinita Burmeister,white grub);伪金针虫属物种(Eleodes spp.)、单叶叩甲属(Conoderus spp.)和Aeolus spp.,铁线虫;黑角负泥虫(Oulema melanopus),谷物叶甲虫;玉米铜色跳甲(Chaetocnema pulicaria),玉米跳甲;玉米长喙象(Sphenophorusmaidis,maize billbug);玉米蚜(Rhopalosiphum maidis),玉米叶蚜虫;Sipha flava,黄甘蔗蚜虫;美洲谷杆长蝽(Blissus leucopterus leucopterus,chinch bug);高粱瘿蚊(Contarinia sorghicola,sorghum midge);朱砂叶螨(Tetranychus cinnabarinus,carmine spider mite);二点叶螨(Tetranychus urticae,twospotted spider mite);
小麦:Pseudaletia unipunctata,粘虫;草地贪夜蛾(Spodoptera frugiperda),秋粘虫;南美玉米苗斑螟(Elasmopalpus lignosellus),小玉米茎蛀虫;Agrotisorthogonia,西方切根虫(west cutworm);南美玉米苗斑螟(Elasmopalpus lignosellus),小玉米茎蛀虫;黑角负泥虫(Oulema melanopus),谷物叶甲;Hypera punctata,苜蓿叶象鼻虫;黄瓜十一星叶甲食根亚种(Diabrotica undecimpunctata howardi),南方玉米根虫(southern corn rootworm);俄国小麦蚜虫;Schizaphis graminum,麦二叉蚜;Macrosiphum avenae,麦长管蚜;红足黑蝗(Melanoplus femurrubrum,redlegedgrasshopper);特异黑蝗(Melanoplus differentialis,differential grasshopper);迁飞黑蝗(Melanoplus sanguinipes,migratory grasshopper);黑森瘿蚊(Mayetioladestructor,Hessian fly);麦红吸浆虫(Sitodiplosis mosellana,wheat midge);美洲麦杆蝇(Meromyza americana,wheat stem maggot);麦地种蝇(Hylemya coarctata,wheatbulb fly);烟褐蓟马(Frankliniella fusca,tobacco thrips);麦茎蜂(Cephus cinctus,wheat stem sawfly);郁金香瘤瘿螨(Aceria tulipae,wheat curl mite);
向日葵:向日葵芽蛾(Suleima helianthana,sunflower bud moth);向日葵螟(Homoeosoma electellum,sunflower moth);向日葵叶甲(zygogramma exclamationis,sunflower beetle);Bothyrus gibbosus,胡萝卜甲虫;向日葵籽瘿蚊(Neolasiopteramurtfeldtiana,sunflower seed midge);
棉花:烟芽夜蛾(Heliothis virescens,cotton budworm);棉铃虫(Helicoverpazea,cotton bollworm);甜菜夜蛾(Spodoptera exigua,beet armyworm);红铃麦蛾(Pectinophora gossypiella,pink bollworm);墨西哥棉铃象(Anthonomus grandis,bollweevil);棉蚜(Aphis gossypii,cotton aphid);棉伪斑腿盲蝽(Pseudatomoscelisseriatus,cotton fleahopper);纹翅粉虱(Trialeurodes abutilonea,bandedwingedwhitefly);美国牧草盲蝽(Lygus lineolaris,tarnished plant bug);红足黑蝗(Melanoplus femurrubrum,redleged grasshopper);特异黑蝗(Melanoplusdifferentialis,differential grasshopper);烟蓟马(Thrips tabaci,onion thrips);烟褐蓟马(Frankliniella fusca,tobacco thrips);朱砂叶螨(Tetranychuscinnabarinus,carmine spider mite);二点叶螨(Tetranychus urticae,twospottedspider mite);
稻:小蔗螟(Diatraea saccharalis,sugarcane borer);草地贪夜蛾(Spodopterafrugiperda),秋粘虫;Spodoptera cosmioides;南部灰翅夜蛾(Spodoptera eridania);棉铃虫(Helicoverpa zea,cotton bollworm);葡萄肖叶甲(Colaspis brunnea,grapecolaspis);稻水象甲(Lissorhoptrus oryzophilus,rice water weevil);米象(Sitophilus oryzae,rice weevil);二条黑尾叶蝉(Nephotettix nigropictus,稻叶蝉rice leafhopper);美洲谷杆长蝽(Blissus leucopterus leucopterus,chinch bug);拟绿蝽(Acrosternum hilare,green stink bug);二化螟(Chilu suppressalis,亚洲稻螟asiatic rice borer);
大豆:大豆尺蠖(Pseudoplusia includens,soybean looper);大豆夜蛾(Anticarsia gemmatalis,velvetbean caterpillar);苜蓿绿夜蛾(Plathypena scabra,green cloverworm);玉米螟(Ostrinia nubilalis),欧洲玉米螟;小地老虎(Agrotisipsilon,黑切根虫(black cutworm));甜菜夜蛾(Spodoptera exigua,beet armyworm);Spodoptera cosmioides;南部灰翅夜蛾(Spodoptera eridania);烟芽夜蛾(Heliothisvirescens,cotton budworm);棉铃虫(Helicoverpa zea,cotton bollworm);墨西哥豆甲(Epilachna varivestis,Mexican bean beetle);绿桃蚜(Myzus persicae,green peachaphid);蚕豆小绿叶蝉(Empoasca fabae,马铃薯叶蝉potato leafhopper);拟绿蝽(Acrosternum hilare,green stink bug);红足黑蝗(Melanoplus femurrubrum,redlegedgrasshopper);特异黑蝗(Melanoplus differentialis,differential grasshopper);灰地种蝇(Hylemya platura,seedcorn maggot);大豆蓟马(Sericothrips variabilis,soybean thrips);烟蓟马(Thrips tabaci,onion thrips);草莓红蜘蛛(Tetranychwturkestani,strawberry spider mite);二点叶螨(Tetranychus urticae,twospottedspider mite);
大麦:玉米螟(Ostrinia nubilalis),欧洲玉米螟;小地老虎(Agrotis ipsilon,黑切根虫(black cutworm));Schizaphis graminum,麦二叉蚜;美洲谷杆长蝽(Blissusleucopterus leucopterus,chinch bug);拟绿蝽(Acrosternum hilare,green stinkbug);褐椿象(Euschistus servus,brown stink bug);新热带褐椿(Euschistus heros,neotropical brown stink bug);灰地种蝇(Hylemya platura,seedcorn maggot);黑森瘿蚊(Mayetiola destructor,Hessian fly);麦岩螨(Petrobia latens,褐色小麦螨(brownwheat mite));
油菜:甘蓝蚜(Brevicoryne brassicae,cabbage aphid);十字花科跳甲(Phyllotreta cruciferae,Flea beetle);蓓带夜蛾(Mamestra configurata,披肩粘虫(bertha armyworm));小菜蛾(Plutella xylostella,diamondback moth);地种蝇属物种(Delia ssp.,Root maggots)。
增加植物产率的方法
本发明提供增加植物产率的方法。该方法包括提供表达编码本文公开的杀农害多肽序列的多核苷酸的植物或植物细胞,和在侵染农害(或易受农害侵染)的田间生长植物或其种子,所述多肽对所述农害具有杀农害活性。在一些实施方案中,所述多肽对鳞翅目、鞘翅目、双翅目、半翅目、或线虫农害具有杀农害活性,所述田间侵染鳞翅目、半翅目、鞘翅目、双翅目、或线虫农害。如本文所定义,植物的“产率”是指植物产生的生物质的质量和/或数量。“生物质”意指任何测量的植物产品。生物质产量的增加是测量的植物产品的产率中任何的改善。增加植物产率具有一些商业应用。例如,增加植物叶生物质可以增加用于人或动物消费的叶类蔬菜的产率。另外,增加叶生物质可用于增加植物源的药物或工业产品的产量。产率的增加可以包括任何统计学上显著的增加,包括但不限于:与不表达杀农害序列的植物相比,产率至少增加1%、至少增加3%、至少增加5%、至少增加10%、至少增加20%、至少30%、至少50%、至少70%、至少100%或更高。在具体的方法中,植物产率由于表达本文公开的杀农害蛋白的植物改善的农害抗性而增加。杀农害蛋白的表达导致农害侵染或摄食的能力降低。
也可以用一种或多种化学组合物处理植物,包括一种或多种除草剂、杀昆虫剂或杀真菌剂。示例性的化学组合物包括:
水果/蔬菜类除草剂:莠去津(Atrazine)、除草定(Bromacil)、敌草隆(Diuron)、草甘膦(Glyphosate)、利谷隆(Linuron)、赛克津(Metribuzin)、西玛津(Simazine)、氟乐灵(Trifluralin)、吡氟禾草灵(Fluazifop)、草铵膦(Glufosinate)、氯吡嘧高恩(Halosulfuron Gowan)、百草枯(Paraquat)、拿草特(Propyzamide)、稀禾定(Sethoxydim)、氟丙嘧草酯(Butafenacil)、氯吡嘧磺隆(Halosulfuron)、茚嗪氟草胺(Indaziflam);
水果/蔬菜杀昆虫剂:涕灭威(Aldicarb)、苏云金芽孢杆菌(Bacillusthuriengiensis)、西维因(Carbaryl)、卡巴呋喃(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)、高氰戊菊酯(Esfenvalerate)、λ-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-三氟氯氰菊酯(Gamma-cyhalothrin)、螺甲螨酯(Spiromesifen)、多杀菌素(Spinosad)、氯虫酰胺(Rynaxypyr)、溴氰虫酰胺(Cyazypyr)、杀农害隆(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)、啶虫胺腈(Sulfoxaflor)、丁氟螨酯(Cyflumetofen),、Cyanopyrafen、噻虫胺(Clothianidin)、噻虫嗪(Thiamethoxam)、Spinotoram、硫双威(Thiodicarb)、氟啶虫酰胺(Flonicamid)、灭虫威(Methiocarb)、甲氨基阿维菌素-苯甲酸盐(Emamectin-benzoate)、茚虫威(Indoxacarb)、克线磷(Fenamiphos)、吡丙醚(Pyriproxifen)、六苯丁锡氧(Fenbutatin-oxid);
水果/蔬菜杀真菌剂:唑嘧菌胺(Ametoctradin)、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)、环唑醇(Cyproconazole)、环丙嘧啶(Cyprodinil)、恶醚唑(Difenoconazole)、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)、乙磷铝(Fosetyl)、异菌脲(Iprodione)、异丙菌(Iprovalicarb)、异皮姆(Isopyrazam)、醚菌酯(Kresoxim-methyl)、代森锰锌(Mancozeb)、双炔酰菌(Mandipropamid)、甲霜灵/精甲霜灵(Metaiaxyl/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)、丙森锌(Propineb)、丙氧喹啉(Proquinazid)、丙硫菌唑(Prothioconazole)、唑菌胺酯(Pyraclostrobin)、嘧霉胺(Pyrimethanil)、喹喔啉(Quinoxyfen)、螺恶嗪(Spiroxamine)、硫(Sulphur)、戊唑醇(Tebuconazole)、甲基托布津(Thiophanate-methyl)、三氟嘧啶(Trifloxystrobin);
谷类除草剂:
2.4-D酰嘧磺隆(Amidosulfuron)、溴苯腈(Bromoxynil)、氟酮唑草-E(Carfentrazone-E)、绿麦隆(Chlorotoluron)、氯磺隆(Chlorsulfuron)、炔草酯-P(Clodinafop-P)、二氯吡啶酸(Clopyralid)、麦草畏(Dicamba)、禾草灵-M(Diclofop-M)、吡氟草胺(Diflufenican)、精恶唑禾草灵(Fenoxaprop)、氟磺草胺(Florasulam)、氟卡巴腙-NA(Flucarbazone-NA)、氟噻草胺(Flufenacet)、Flupyrosulfuron-M、氯氟吡氧乙酸(Fluroxypyr)、呋草酮(Flurtamone)、草甘膦(Glyphosate)、碘甲磺隆(Iodosulfuron)、碘苯腈(Ioxynil)、异丙隆(Isoproturon)、MCPA、甲磺胺磺隆(Mesosulfuron)、甲磺隆(Metsulfuron)、二甲戊乐灵(Pendimethalin)、唑啉草酯(Pinoxaden)、丙氧基卡巴(Propoxycarbazone)、苄草丹(Prosulfocarb)、甲氧磺草胺(Pyroxsulam)、磺酰磺隆(Sulfosulfuron)、噻吩磺隆(Thifensulfuron)、苯草酮(Tralkoxydim)、醚苯磺隆(Triasulfuron)、苯磺隆(Tribenuron)、氟乐灵(Trifluralin)、三氟甲磺隆(Tritosulfuron);
谷类杀真菌剂:嘧菌酯(Azoxystrobin)、联苯吡菌胺(Bixafen)、啶酰菌胺(Boscalid)、多菌灵(Carbendazim)、百菌清(Chlorothalonil)、环氟菌胺(Cyflufenamid)、环丙唑醇(Cyproconazole)、嘧菌环胺(Cyprodinil)、醚菌胺(Dimoxystrobin)、氟环唑(Epoxiconazole)、苯锈啶(Fenpropidin)、丁苯吗啉(Fenpropimorph)、氟吡菌酰胺(Fluopyram)、氟嘧菌酯(Fluoxastrobin)、氟喹唑(Fluquinconazole)、氟唑菌酰胺(Fluxapyroxad)、吡唑萘菌胺(Isopyrazam)、醚菌酯(Kresoxim-methyl)、叶菌唑(Metconazole)、苯菌酮(Metrafenone)、吡噻菌胺(Penthiopyrad)、啶氧菌酯(Picoxystrobin)、咪鲜胺(Prochloraz)、丙环唑(Propiconazole)、丙氧喹啉(Proquinazid)、丙硫菌唑(Prothioconazole)、唑菌胺酯(Pyraclostrobin)、喹喔啉(Quinoxyfen)、螺螨胺(Spiroxamine)、戊唑醇(Tebuconazole)、甲基托布津(Thiophanate-methyl)、肟菌酯(Trifloxystrobin);
谷类杀昆虫剂:乐果(Dimethoate)、λ-三氟氯氰菊酯(Lambda-cyhalthrin)、溴氰菊酯(Deltamethrin)、α-氯氰菊酯(alpha-Cypermethrin)、β-氟氯氰菊酯(β-cyfluthrin)、联苯菊酯(Bifenthrin)、吡虫啉(Imidacloprid)、噻虫胺(Clothianidin)、噻虫嗪(Thiamethoxam)、噻虫啉(Thiacloprid)、啶虫脒(Acetamiprid)、呋虫胺(Dinetofuran)、毒死蜱(Clorphyriphos)、抗蚜威(Pirimicarb)、灭虫威(Methiocarb)、啶虫胺腈(Sulfoxaflor);
玉米除草剂:阿特拉津(Atrazine)、甲草胺(Alachlor)、溴苯腈(Bromoxynil)、乙草胺(Acetochlor)、麦草畏(Dicamba)、二氯吡啶酸(Clopyralid)、(S-)二甲酚草胺((S-)Dimethenamid)、草铵膦(Glufosinate)、草甘膦(Glyphosate)、异恶唑草酮(Isoxaflutole)、(S-)甲草胺((S-)Metolachlor)、硝磺草酮(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)、氟磺草酮(Tembotrione)、嘧磺草胺(Saflufenacil)、噻酮磺隆(Thiencarbazone)、氟噻草胺(Flufenacet)、皮罗威(Pyroxasulfon);
玉米杀昆虫剂:卡巴呋喃(Carbofuran)、毒死蜱(Chlorpyrifos)、联苯菊酯(Bifenthrin)、氟虫腈(Fipronil)、吡虫啉(Imidacloprid)、λ-三氟氯氰菊酯(Lambda-Cyhalothrin)、七氟菊酯(Tefluthrin)、特丁磷(Terbufos)、噻虫嗪(Thiamethoxam)、噻虫胺(Clothianidin)、螺甲螨酯(Spiromesifen)、氟虫双酰胺(Flubendiamide)、杀铃脲(TrifJumuron)、氯虫酰胺(Rynaxypyr)、溴氰菊酯(Deltamethrin)、硫双威(Thiodicarb)、β-氟氯氰菊酯(β-Cyfluthrin)、氯氰菊酯(Cypermethrin)、联苯菊酯(Bifenthrin)、虱螨脲(Lufenuron)、嘧丙磷(Tebupirimphos)、乙虫腈(Ethiprole)、溴氰虫酰胺(Cyazypyr)、噻虫啉(Thiacloprid)、啶虫脒(Acetamiprid)、呋虫胺(Dinetofuran)、阿维菌素(Avermectin);
玉米杀真菌剂:嘧菌酯(Azoxystrobin)、联苯吡菌胺(Bixafen)、啶酰菌胺(Boscalid)、环丙唑醇(Cyproconazole)、醚菌胺(Dimoxystrobin)、氟环唑(Epoxiconazole)、Fenitropan、氟吡菌酰胺(Fluopyram)、氟嘧菌酯(Fluoxastrobin)、氟唑菌酰胺(Fluxapyroxad)、吡唑萘菌胺(Isopyrazam)、叶菌唑(Metconazole)、吡噻菌胺(Penthiopyrad)、啶氧菌酯(Picoxystrobin)、丙环唑(Propiconazole)、丙硫菌唑(Prothioconazole)、唑菌胺酯(Pyraclostrobin)、戊唑醇(Tebuconazole)、肟菌酯(Trifloxystrobin);
稻除草剂:丁草胺(Butachlor)、敌稗(Propanil)、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)、恶嗪草酮(Oxaziclomefone)、吡嘧磺隆(Pyrazosulfuron)、稗草畏(Pyributicarb)、快杀稗(Quinclorac)、禾草丹(Thiobencarb)、茚草酮(Indanofan)、氟噻草胺(Flufenacet)、四唑酰草胺(Fentrazamide)、氯吡嘧磺隆(Halosulfuron)、恶嗪草酮(Oxaziclomefone)、双环磺草酮(Benzobicyclon)、环酯草醚(Pyriftalid)、五氟磺草胺(Penoxsulam)、双草醚(Bispyribac)、丙炔恶草酮(Oxadiargyl)、乙氧嘧磺隆(Ethoxysulfuron)、丙草胺(Pretilachlor)、甲基磺草酮(Mesotrione)、特呋三酮(Tefuryltrione)、恶草灵(Oxadiazone)、精恶唑禾草灵(Fenoxaprop)、吡丙醚(Pyrimisulfan);
稻杀昆虫剂:二嗪农(Diazinon)、仲丁威(Fenobucarb)、丙硫克百威(Benfuracarb)、噻嗪酮(Buprofezin)、呋虫胺(Dinotefuran)、氟虫腈(Fipronil)、吡虫啉(Imidacloprid)、异丙威(Isoprocarb)、噻虫啉(Thiacloprid)、环虫酰肼(Chromafenozide)、噻虫胺(Clothianidin)、乙虫腈(Ethiprole)、氟虫双酰胺(Flubendiamide)、氯虫酰胺(Rynaxypyr)、溴氰菊酯(Deltamethiin)、啶虫脒(Acetamiprid)、噻虫嗪(Thiamethoxam)、溴氰虫酰胺(Cyazypyr)、多杀菌素(Spinosad)、Spinotoram、甲氨基阿维菌素-苯甲酸盐(Emamectin-Benzoate)、氯氰菊酯(Cypermethrin)、毒死蜱(Chlorpyriphos)、醚菊酯(Etofenprox)、克百威(Carbofuran)、丙硫克百威(Benfuracarb)、啶虫胺腈(Sulfoxaflor);
稻杀真菌剂:嘧菌酯(Azoxystrobin)、多菌灵(Carbendazim)、环丙酰菌胺(Carpropamid)、双氯氰菌胺(Diclocymet)、苯醚甲环(Difenoconazole)、克瘟散(Edifenphos)、嘧菌腙(Ferimzone)、庆大霉素(Gentamycin)、异菌脲(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(Mancozeb)、苯氧菌胺(Metominostrobin)、肟醚菌胺(Orysastrobin)、戊菌隆(Pencycuron)、噻菌灵(Probenazole)、丙环唑(Propiconazole)、丙森锌(Propineb)、咯喹酮(Pyroquilon)、戊唑醇(Tebuconazole)、甲基托布津(Thiophanate-methyl)、噻酰菌胺(Tiadinil)、三环唑(Tricyclazole)、肟菌酯(Trifloxystrobin)、有效霉素(Validamycin);
棉花除草剂:敌草隆(Diuron)、伏草隆(Fluometuron)、MSMA、乙氧氟草醚(Oxyfluorfen)、扑草净(Prometryn)、氟乐灵(Trifluralin)、氟酮唑草(Carfentrazone)、烯草酮(Clethodim)、吡氟禾草灵(Fluazifop-butyl)、草甘膦(Glyphosate)、达草灭(Norflurazon)、二甲戊灵(Pendimethalin)、嘧草硫醚(Pyrithiobac-sodium)、三氟啶磺隆(Trifloxysulfuron)、吡喃草酮(Tepraloxydim)、草铵膦(Glufosinate)、丙炔氟草酮(Flumioxazin)、噻苯隆(Thidiazuron);
棉花杀农害剂:乙酰甲胺磷(Acephate)、涕灭威(Aldicarb)、毒死蜱(Chlorpyrifos)、氯氰菊酯(Cypermethrin)、溴氰菊酯(Deltamethiin)、阿维菌素(Abamectin)、啶虫脒(Acetamiprid)、甲氨基阿维菌素-苯甲酸盐(Emamectin Benzoate)、吡虫啉(Imidacloprid)、茚虫威(Indoxacarb)、λ-三氟氯氰菊酯(Lambda-Cyhalothrin)、多杀菌素(Spinosad)、硫双威(Thiodicarb)、γ-三氟氯氰菊酯(Gamma-Cyhalothiin)、螺螨虫(Spiromesifen)、吡啶胺(Pyridalyl)、氟虫胺(Flonicamid)、氟虫双酰胺(Flubendiamide)、杀玲脲(Triflumuron)、氯虫酰胺(Rynaxypyr)、β-氟氯氰菊酯(Beta-Cyfluthrin)、螺虫乙酯(Spirotetramat)、噻虫胺(Clothianidin)、噻虫嗪(Thiamethoxam)、噻虫啉(Thiacloprid)、呋虫胺(Dinetofuran)、氟虫双酰胺(Flubendiamide)、溴氰虫酰胺(Cyazypyr)、多杀菌素(Spinosad)、Spinotoram、γ-三氟氯氰菊酯(gamma Cyhalothrin)、4-[[(6-氯吡啶-3-基)甲基](22-二氟乙基)氨基]呋喃-2(5H)-酮、硫双威(Thiodicarb)、阿维菌素(Avermectin)、氟啶虫酰胺(Flonicamid)、啶虫丙醚(Pyridalyl)、螺甲螨酯(Spiromesifen)、氟啶虫胺腈(Sulfoxaflor);
棉花杀真菌剂:嘧菌酯(Azoxystrobin)、联苯吡菌胺(Bixafen)、啶酰菌胺(Boscalid)、多菌灵(Carbendazim)、百菌清(Chlorothalonil)、铜(Copper)、环丙唑醇(Cyproconazole)、苯醚甲环唑(Difenoconazole)、醚菌胺(Dimoxystrobin)、氟环唑(Epoxiconazole)、咪唑菌酮(Fenamidone)、氟啶胺(Fluazinam)、氟吡菌酰胺(Fluopyram)、氟嘧菌酯(Fluoxastrobin)、氟唑菌酰胺(Fluxapyroxad)、异菌脲(Iprodione)、吡唑萘菌胺(Isopyrazam)、异噻菌胺(Isotianil)、代森锰锌(Mancozeb)、代森锰(Maneb)、苯氧菌胺(Metominostrobin)、吡噻菌胺(Penthiopyrad)、啶氧菌酯(Picoxystrobin)、丙森锌(Propineb)、丙硫菌唑(Prothioconazole)、唑菌胺酯(Pyraclostrobin)、五氯硝基苯(Quintozene)、戊唑醇(Tebuconazole)、氟醚唑(Tetraconazole)、甲基托布津(Thiophanate-methyl)、肟菌酯(Trifloxystrobin);
大豆除草剂:甲草胺(Alachlor)、灭草松(Bentazone)、氟乐灵(Trifluralin)、氯嘧磺隆(Chlorimuron-Ethyl)、甲基氯酯磺草胺酸(Cloransulam-Methyl)、精恶唑禾草灵(Fenoxaprop)、氟磺胺草醚(Fomesafen)、吡氟禾草灵(Fluazifop)、草甘膦(Glyphosate)、咪草啶酸(Imazamox)、灭草喹(Imazaquin)、咪草烟(Imazethapyr)、(S-)异丙甲草胺((S-)Metolachlor)、嗪草酮(Metribuzin)、二甲戊乐灵(Pendimethalin)、吡喃草酮(Tepraloxydim)、草铵膦(Glufosinate);
大豆杀昆虫剂:λ-三氟氯氰菊酯(Lambda-cyhalothrin)、灭多威(Methomyl)、吡虫啉(Imidacloprid)、噻虫胺(Clothianidin)、噻虫嗪(Thiamethoxam)、噻虫啉(Thiacloprid)、啶虫脒(Acetamiprid)、呋虫胺(Dinetofuran)、氟虫双酰胺(Flubendiamide)、氯虫酰胺(Rynaxypyr)、溴氰虫酰胺(Cyazypyr)、多杀菌素(Spinosad)、Spinotoram、甲氨基阿维菌素-苯甲酸盐(Emamectin-Benzoate)、氟虫腈(Fipronil)、乙虫腈(Ethiprole)、溴氰菊酯(Deltamethrin)、β-氟氯氰菊酯(β-Cyfluthrin)、γ和λ-三氟氯氰菊酯(gamma and lambda Cyhalothrin),4-[[(6-氯吡啶-3-基)甲基](22-二氟乙基)氨基]呋喃-2(5H)-酮、螺虫乙酯(Spirotetramat)、Spinodiclofen、杀铃脲(Triflumuron)、氟啶虫酰胺(Flonicamid)、硫双威(Thiodicarb)、β-氟氯氰菊酯(beta-Cyfluthrin);
大豆杀真菌剂:嘧菌酯(Azoxystrobin)、联苯吡菌胺(Bixafen)、啶酰菌胺(Boscalid)、多菌灵(Carbendazim)、百菌清(Chlorothalonil)、铜(Copper)、环唑醇(Cyproconazole)、醚唑(Difenoconazole)、醚菌胺(Dimoxystrobin)、氟环唑(Epoxiconazole)、氟啶胺(Fluazinam)、氟吡菌酰胺(Fluopyram)、氟嘧菌酯(Fluoxastrobin)、粉唑醇(Flutriafol)、氟唑菌酰胺(Fluxapyroxad)、吡唑萘菌胺(Isopyrazam)、扑海因(Iprodione)、异噻菌胺(Isotianil)、代森锰锌(Mancozeb)、代森锰(Maneb)、叶菌唑(Metconazole)、苯氧菌胺(Metominostrobin)、腈菌唑(Myclobutanil)、吡噻菌胺(Penthiopyrad)、啶氧菌酯(Picoxystrobin)、丙环唑(Propiconazole)、丙森锌(Propineb)、丙硫菌唑(Prothioconazole)、唑菌胺酯(Pyraclostrobin)、戊唑醇(Tebuconazole)、氟醚唑(Tetraconazole)、甲基托布津(Thiophanate-methyl)、肟菌酯(Trifloxystrobin);
甜菜除草剂:杀草敏(Chloridazon)、甜菜安(Desmedipham)、甜菜呋(Ethofumesate)、甜菜宁(Phenmedipham)、野麦畏(Triallate)、二氯吡啶酸(Clopyralid)、吡氟禾草灵(Fluazifop)、环草定(Lenacil)、苯嗪草酮(Metamitron)、喹草酸(Quinmerac)、噻草酮(Cycloxydim)、氟胺磺隆(Triflusulfuron)、吡喃草酮(Tepraloxydim)、喹禾灵(Quizalofop);
甜菜杀昆虫剂:吡虫啉(Imidacloprid)、噻虫胺(Clothianidin)、噻虫嗪(Thiamethoxam)、噻虫啉(Thiacloprid)、乙酰咪唑(Acetamiprid)、呋虫胺(Dinetofuran)、溴氰菊酯(Deltamethrin)、β-氟氯氰菊酯(β-Cyfluthrin)、γ/λ-三氟氯氰菊酯(gamma/lambda Cyhalothrin)、4-[[(6-氯吡啶-3-基)甲基](22-二氟乙基)氨基]呋喃-2(5H)-酮、七氟菊酯(Tefluthrin)、氯虫酰胺(Rynaxypyr)、Cyaxypyr、氟虫腈(Fipronil)、卡巴呋喃(Carbofuran);
卡诺拉油菜(Canola)除草剂:二氯吡啶酸(Clopyralid)、禾草灵(Diclofop)、吡氟禾草灵(Fluazifop)、草铵膦(Glufosinate)、草甘膦(Glyphosate)、吡草胺(Metazachlor)、氟乐灵胺苯磺隆(Trifluralin Ethametsulfuron)、喹草酸(Quinmerac)、精喹禾灵(Quizalofop)、烯草酮(Clethodim)、吡喃草(Tepraloxydim);
卡诺拉油菜杀真菌剂:嘧菌酯(Azoxystrobin)、联苯吡菌胺(Bixafen)、啶酰菌胺(Boscalid)、多菌灵(Carbendazim)、环唑醇(Cyproconazole)、苯醚甲环唑(Difenoconazole)、醚菌胺(Dimoxystrobin)、氟环唑(Epoxiconazole)、氟啶胺(Fluazinam)、氟吡菌酰胺(Fluopyram)、氟嘧菌酯(Fluoxasu'obin)、氟硅唑(Flusilazole)、氟唑菌酰胺(Fluxapyroxad)、异菌脲(Iprodione)、吡唑萘菌胺(Isopyrazam)、甲哌鎓(Mepiquat-chloride)、叶菌唑(Metconazole)、苯氧菌胺(Metominostrobin)、多效唑(Paclobutrazole)、吡噻菌胺(Penthiopyrad.)、啶氧菌酯(Picoxystrobin)、咪鲜胺(Prochloraz)、丙硫菌唑(Prothioconazole)、吡唑醚菌酯(Pyraclostrobin)、戊唑醇(Tebuconazole)、甲基托布津(Thiophanate-methyl)、三氟嘧啶(Trifloxystrobin)、农利灵(Vinclozolin);
卡诺拉油菜杀昆虫剂:卡巴呋喃(Carbofuran)、噻虫啉(Thiacloprid)、溴氰菊酯(Deltamethrin)、吡虫啉(Imidacloprid)、噻虫胺(Clothianidin)、噻虫嗪(Thiamethoxam)、啶虫脒(Acetamiprid)、呋虫胺(Dinetofuran)、β-氟氯氰菊酯(β-Cyfluthrin)、γ和λ-三氟氯氰菊酯(gamma and lambda Cyhalothrin)、tau-氟戊酸(tau-Fluvaleriate)、乙虫腈(Ethiprole)、多杀菌素(Spinosad)、Spinotoram、氟虫双酰胺(Flubendiamide)、氯虫酰胺(Rynaxypyr)、氰虫酰胺(Cyazypyr)、4-[[(6-氯吡啶-3-基)甲基](22-二氟乙基)氨基]呋喃-2(5H)-酮。
将本发明的基因导入另一植物的方法
本文还提供了将本发明的核酸导入另一植物的方法。可以通过轮回选择、回交、谱系育种、品系选择、混合选择(mass selection)、突变育种和/或遗传标记物增强的选择,将本发明的核酸或其片段导入第二植物中。
因此,在一个实施方案中,本发明的方法包括使包含本发明的核酸的第一植物与第二植物杂交以产生F1后代植物并选择包含本发明核酸的F1后代植物。所述方法可以进一步包括使选择的后代植物与包含本发明核酸的第一植物杂交以产生回交后代植物并选择包含本发明核酸的回交后代植物。用于评估杀农害活性的方法在本文其他地方提供。所述方法可以进一步包括连续重复这些步骤一次或多次以产生选定的包含本发明核酸的第二代或更高世代的回交后代植物。
任何涉及选择期望表型植物的育种方法都可用于本发明的方法中。在一些实施方案中,F1植物可以自花授粉以产生分离的F2代。然后可以在每代(F3,F4,F5等)中选择代表期望表型(例如杀农害活性)的单株植物,直到在育种群体中该性状是纯合的或固定的。
第二植物可以是具有期望性状的植物,如除草剂耐性、昆虫耐性、干旱耐性、线虫控制、水分利用效率、氮利用效率、改善的营养价值、疾病抗性、改善的光合作用、改善的纤维质量、胁迫耐性、改进的生殖等。第二植物可以是如本文其他地方所述的原种事件(eliteevent)。
在各种实施方案中,可以从得到的杂交中收获植物部分(完整植物、植物器官(例如,叶、茎、根等)、种子、植物细胞、繁殖体、胚胎等),并且繁殖或收集用于下游使用(如食物、饲料、生物燃料、油、面粉、膳食等)。
获得植物产品的方法
本发明还涉及获得商品的方法,包括从包含本发明核酸的作物中收获和/或研磨籽粒以获得商品。农艺和商业上重要的产品和/或物质组合物,包括但不限于动物饲料、商品和植物产品以及副产品(旨在用作人消耗的食品或用于人消耗的组合物和商品中),特别是失活的(devitalized)种子/籽粒产品,包括由这种籽粒/种子产生的(半)加工产品,其中所述产品是或包含完全或加工的种子或籽粒、动物饲料、玉米或大豆粉状物(meal)、玉米或大豆粉、玉米、玉米淀粉、大豆粉状物、大豆粉、薄片、大豆蛋白浓缩物、大豆蛋白分离物、组织化(texturized)大豆蛋白浓缩物、化妆品、护发产品、大豆果仁酱、纳豆、豆豉、水解大豆蛋白、人造黄油(whipped topping)、起酥油、卵磷脂、可食用的整粒大豆(生的、烤的或作为毛豆),大豆酸奶、大豆奶酪、豆腐、腐竹、以及煮熟(cooked)、抛光(polished)、蒸(steamed)、烤(baked)或煮半熟(parboiled)的籽粒等,都旨在本发明的范围内,只要这些产品和物质组合物含有可检测量的本文所述的核苷酸和/或氨基酸序列,其中通过所述的核苷酸和/或氨基酸序列,可以对含有这种核苷酸序列的任何植物进行诊断。
提供以下实施例是为了举例说明而不是为了限制。
实验实施例
实施例1.发现新的杀农害基因
使用以下步骤从细菌菌株鉴定新的杀农害基因:
·从菌株制备总DNA。总DNA含有基因组DNA和染色体外DNA。染色体外DNA含有以下中的一些或全部的混合物:各种大小的质粒;噬菌体染色体;其他未表征的染色体外分子。
·DNA的测序。通过二代测序方法,对总DNA进行测序。
·通过同源性和/或其他计算分析,鉴定推定的毒素基因。
·当需要时,通过几种PCR或克隆策略(例如TAIL-PCR)之一对感兴趣基因进行序列整理。
表1.鉴定的新基因;
通过PCR从pAX980扩增本文公开的毒素基因,通过本领域熟知的方法将PCR产物克隆到芽孢杆菌表达载体pAX916或其他合适的载体中。将所得的含有带axmi基因的载体的芽孢杆菌菌株在常规生长培养基上培养,如CYS培养基(10g/l Bacto-酪胨;3g/l酵母提取物;6g/l KH2PO4;14g/l K2HPO4;0.5mM MgSO4;0.05mM MnCl2;0.05mM FeSO4),直到通过显微镜检查显现孢子形成。制备样品并生物测定法中测试活性。
实施例2.杀农害活性测定法
可以测试本发明的核苷酸序列产生杀农害蛋白的能力。通常以多种方式评估杀农害蛋白在农害上作为杀农害剂发挥作用的能力。本领域熟知的一种方法是进行摄食测定法。在这样的摄食测定法中,将农害暴露于含有待测化合物的样品或对照样品。通常,这通过将待测物质或这种物质的合适稀释液放置在农害将摄取的材料(例如人工饲料)上来进行。待测物质可以由液体、固体或浆料组成。可将待测物质置于表面上,然后使其干燥。或者,可以将待测物质与熔融人工饲料混合,然后分配到测定室中。测定室可以是例如杯子、盘子或微量滴定板的孔。
用于吮吸式农害(例如蚜虫)的测定法可以包括通过分隔将测试物质与昆虫分开,理想地分隔是可以被吮吸式昆虫的吸嘴部分刺穿的部分,以允许摄取测试物质。通常将测试物质与摄食刺激剂(如蔗糖)混合以促进测试化合物的摄取。
其他类型的测定法可以包括将测试物质微注射到农害的口中或者肠道,以及开发转基因植物,然后测试农害以转基因植物为食的能力。植物测试可以包括隔离通常食用的植物部分,例如附着叶子的小笼子,或者将整个植物隔离在含有昆虫的笼子中。
用于测定农害的其他方法和处理是本领域已知的,并且可以在例如Robertsonand Preisler,编辑(1992)Pesticide bioassays with arthropods,CRC,Boca Raton,FL中可见。或者,测定法通常描述于期刊Arthropod Management Tests and Journal ofEconomic Entomology或由美国昆虫学会(Entomological Society of America(ESA))成员讨论。
在一些实施方案中,将编码本文公开的杀农害蛋白毒素区的DNA区克隆到大肠杆菌表达载体pMAL-C4x中,在编码麦芽糖结合蛋白(MBP)的malE基因之后。这些框内融合导致MBP-Axmi融合蛋白在大肠杆菌中表达。
为了在大肠杆菌中表达,用各质粒转化BL21*DE3。将单菌落接种在补充有羧苄青霉素和葡萄糖的LB中,并在37℃下生长过夜。第二天,用1%的过夜培养物接种新鲜培养基,并在37℃下生长至对数期。随后,在20℃用0.3mM IPTG过夜诱导培养物。将每个细胞沉淀悬浮于20mM Tris-Cl缓冲液,pH7.4+200mM NaCl+1mM DTT+蛋白酶抑制剂中并超声处理。通过SDS-PAGE的分析可用于确认融合蛋白的表达。
然后使整个无细胞提取物在与快速蛋白液相色谱法(FPLC)连接的直链淀粉柱上运行,用于MBP-axmi融合蛋白的亲和纯化。用10mM麦芽糖溶液从树脂上洗脱结合的融合蛋白。然后用因子Xa或胰蛋白酶切割纯化的融合蛋白,以从Axmi蛋白中除去氨基末端MBP标签。可通过SDS-PAGE测定蛋白的切割和溶解。
实施例3.表达和纯化
表达Axmi669和Axmi991并测定其生物活性。通过Genscipt合成基因并将其克隆到pMalC4X载体中以分别产生质粒pGen669和pGen991。通过测序确认克隆,然后转化B121感受态细胞。将每个的单菌落接种在LB培养基中并在37℃生长至对数期,并用0.5mM IPTG在20℃诱导16小时。根据标准方案,对纯化的Axmi669和Axmi991分别进行针对所选昆虫农害的生物测定法。结果显示在表2和表3中。
表2.Axmi669的活性
农害组 | 发育障碍得分 | 死亡率百分数 |
棉铃虫(HELIZE) | 4 | 0% |
小菜蛾(PLUTMA) | 4 | 100% |
西南玉米螟(GIATGR) | 4 | 0% |
大豆夜蛾(THEMGE) | 4 | 0% |
大豆尺蠖(PSEPIN) | 4 | 100% |
表3.Axmi991的活性
农害组 | 发育障碍得分 | 死亡率百分数 |
棉铃虫(Hz) | 1 | 0% |
小菜蛾(DBM) | 4 | 100% |
南部灰翅夜蛾(SAW) | 4 | 100% |
大豆夜蛾(VBC) | 4 | 0% |
发育障碍的分数:
0-没有活性;
1-非一致发育障碍
2-轻微一致的发育障碍(对照大小的75%)
3-强的一致的发育障碍(对照大小的74-26%)
4-严重的一致的发育障碍(小于对照大小的25%)
实施例4.用于植物表达的基因的载体构建
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。此类序列是本领域熟知的,可包括用于在单子叶植物中表达的稻肌动蛋白启动子或玉米泛素启动子,用于在双子叶植物中表达的拟南芥UBQ3启动子或CaMV 35S启动子,以及nos或PinII终止子。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,设计和产生合成的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。
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,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,在本领域中已知来自白羽扇豆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)导致异源蛋白靶向内质网。如果得到的融合蛋白在C-末端还含有内质网保留序列,即包含肽N-末端-赖氨酸-天冬氨酸-谷氨酸-亮氨酸(即,“KDEL”基序,SEQ ID NO:5),则融合蛋白将靶向内质网。如果融合蛋白缺乏在C-末端的内质网靶向序列,则蛋白将靶向内质网,但最终将被隔离在质外体中。
因此,该基因编码融合蛋白,所述融合蛋白含有与本发明氨基酸序列的N末端融合的来自白羽扇豆Lupinus albus的酸性磷酸酶基因的N-末端31个氨基酸(ID GI:14276838,Miller等人(2001)同上引文),以及在C-末端的KDEL(SEQ ID NO:5)序列。因此,预测所得蛋白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可选择标记物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。这些可包括用于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。
实施例5.大豆转化
使用本领域熟知的方法实现大豆转化,如可以使用根瘤农杆菌介导的转化大豆半种子外植体、基本上使用Paz等人(2006),Plant cell Rep.25:206描述的方法进行。使用环磺酮(tembotrione)作为选择标记物鉴定转化体。观察到绿芽的出现,并记录为对除草剂异恶唑草酮(isoxaflutole)或环磺酮耐性的指标。耐受的转基因芽将显示与未用异恶唑草酮或环磺酮处理的野生型大豆芽相当的正常绿化,而用相同量的异恶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,如异恶唑草酮或环磺酮。
耐性绿色芽被转移到生根培养基或接枝。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,例如用环磺酮以100g AI/ha的比率或用补充有硫酸铵甲基酯菜籽油的甲基磺草酮(mesotrione)以300g AI/ha的比率喷洒。施用后10天,评估由应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6: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抑制剂除草剂喷洒充分生长的植物,例如环磺酮,等于100或200g AI/ha(补充硫酸铵和甲基酯菜籽油)。喷洒应用后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。
实施例7.用本文所述的杀农害蛋白基因转化玉米细胞
最好在授粉8-12天后收集玉米穗。从穗中分离胚,优选那些0.8-1.5mm大小的胚用于转化。将胚以盾片朝上铺板在适当的孵育培养基上,如DN62A5S培养基(3.98g/L N6盐;1mL/L(1000x原液的)N6维生素;800mg/L L-天冬酰胺;100mg/L Myo-肌醇;1.4g/L L-脯氨酸;100mg/L酪蛋白氨基酸;50g/L蔗糖;1mL/L(1mg/mL原液的)2,4-D)。然而,除DN62A5S之外的培养基和盐也是合适的并且是本领域已知的。将胚在25℃在黑暗中过夜孵育。然而,过夜孵育胚本身并不是必需的。
将得到的外植体转移到网格方块(每板30-40个)中,转移到渗透培养基上约30-45分钟,然后转移到照射板上(参见,例如,PCT公开号WO/0138514和美国专利号5,240,842)。
使用气溶胶束加速器,使用基本上如PCT公开号WO/0138514中所述的条件,将设计用于植物细胞的本发明基因的DNA构建体加速到植物组织中。照射后,将胚在渗透培养基上孵育约30分钟,并置于孵育培养基上在25℃在黑暗中过夜。为了避免过度损坏照射的外植体,将其孵育至少24小时,然后转移到恢复培养基中。然后将胚涂布在恢复期培养基上,在黑暗中25℃约5天,然后转移到选择培养基中。根据所用特定选择的性质和特征,将外植体在选择培养基中孵育长达八周。在选择期后,将得到的愈伤组织转移到胚成熟培养基中,直到观察到成熟体细胞胚的形成。然后将所得的成熟体细胞胚置于弱光下,并通过本领域已知的方法启动再生过程。使得到的芽在生根培养基上生根,并将所得植物转移到育苗盆中并繁殖为转基因植物。
材料
DN62A5S培养基
用1N KOH/1N KCl将溶液的pH调节至pH 5.8,以高达3g/L的浓度加入Gelrite(Sigma),并将培养基高压灭菌。冷却至50℃后,加入2ml/L的5mg/ml硝酸银原液(Phytotechnology Labs)。
实施例8.通过农杆菌介导的转化在植物细胞中转化本发明的基因
最好在授粉后8-12天收集穗。从穗中分离胚,优选那些0.8-1.5mm大小的胚用于转化。将胚按照盾片朝上铺板在适当的孵育培养基中,并在黑暗中于25℃孵育过夜。然而,过夜孵育胚本身并不是必需的。使胚与含有适当载体的农杆菌菌株接触约5-10分钟,用于Ti质粒介导的转移,然后铺板在共培养培养基上约3天(在黑暗中22℃)。共培养后,将外植体转移至恢复期培养基中5-10天(在黑暗中于25℃)。根据所用特定选择的性质和特征,将外植体在选择培养基中培养长达八周。在选择期后,将得到的愈伤组织转移到胚成熟培养基中,直到观察到成熟体细胞胚的形成。然后将所得的成熟体细胞胚置于弱光下,并如本领域已知的启动再生过程。
实施例9.稻的转化
从温室中良好控制条件下生长的供体植物收集含有处于正确发育阶段的胚的未成熟稻种子。在种子灭菌后,切下未成熟胚并在固体培养基上预诱导3天。预诱导后,将胚浸入含有期望载体的农杆菌悬浮液中几分钟。然后将胚在含有乙酰丁香酮的固体培养基上共培养,并在黑暗中孵育4天。然后将外植体转移到含有草丁膦(phosphinotricin)作为选择剂的第一选择培养基中。大约3周后,将具有愈伤组织发育的盾片切成几个较小的片并转移到相同的选择培养基中。大约每2周进行一次随后的传代培养。在每次传代培养后,将活跃生长的愈伤组织切成较小的片并在第二选择培养基上孵育。数周后,将明显对草丁膦具有抗性的愈伤组织转移到选择性再生培养基中。产生的小植株在半强度MS下培养以完全伸长。植物最终转移到土壤中并在温室中生长。
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 (26)
1.一种重组核酸分子,其包含编码具有杀农害活性的氨基酸序列的核苷酸序列,其中所述核苷酸序列选自:
a)SEQ ID NO:1或3中所示的核苷酸序列;
b)编码包含SEQ ID NO:2或4的氨基酸序列的多肽的核苷酸序列;
c)编码多肽的核苷酸序列,所述多肽包含与SEQ ID NO:2或4的氨基酸序列具有至少95%序列同一性的氨基酸序列。
2.根据权利要求1所述的重组核酸分子,其中所述核苷酸序列是设计用于在植物中表达的合成序列。
3.根据权利要求1所述的重组核酸分子,其中所述核苷酸序列与能够指导所述核苷酸序列在植物细胞中表达的启动子可操作地连接。
4.一种载体,其包含权利要求1所述的重组核酸分子。
5.根据权利要求4所述的载体,其还包含编码异源多肽的核酸分子。
6.一种宿主细胞,其含有权利要求1所述的重组核酸。
7.根据权利要求6所述的宿主细胞,其是细菌宿主细胞。
8.根据权利要求6所述的宿主细胞,其是植物细胞。
9.一种转基因植物,其包含权利要求8所述的宿主细胞。
10.根据权利要求9所述的转基因植物,其中所述植物选自玉米、高粱、小麦、卷心菜、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。
11.一种转基因种子,其包含权利要求1所述的核酸分子。
12.一种具有杀农害活性的重组多肽,其选自:
a)包含SEQ ID NO:2或4的氨基酸序列的多肽;和
b)包含与SEQ ID NO:2或4的氨基酸序列具有至少95%序列同一性的氨基酸序列的多肽。
13.根据权利要求12所述的多肽,其进一步包含异源氨基酸序列。
14.一种组合物,其包含权利要求12的多肽。
15.根据权利要求14所述的组合物,其中所述组合物选自粉末、撒粉剂、丸剂、颗粒、喷雾、乳剂、胶体和溶液。
16.根据权利要求14所述的组合物,其中所述组合物通过干燥、冻干、均质、萃取、过滤、离心、沉降或浓缩细菌细胞培养物来制备。
17.根据权利要求14所述的组合物,其包含约1%至约99%重量的所述多肽。
18.一种控制鳞翅目、半翅目、鞘翅目、线虫或双翅目农害群的方法,所述方法包括使所述群与杀农害有效量的权利要求12的多肽接触。
19.一种杀死鳞翅目、半翅目、鞘翅目、线虫或双翅目农害的方法,所述方法包括使所述农害接触,或向所述农害喂食,杀农害有效量的权利要求12的多肽。
20.一种生产具有杀农害活性的多肽的方法,所述方法包括在表达编码多肽的核酸分子的条件下培养权利要求6的宿主细胞。
21.一种植物或植物细胞,其在其基因组中稳定并入了DNA构建体,所述DNA构建体包含编码具有杀农害活性的蛋白的核苷酸序列,其中所述核苷酸序列选自:
a)SEQ ID NO:1或3中所示的核苷酸序列;
b)编码包含SEQ ID NO:2或4的氨基酸序列的多肽的核苷酸序列;和
c)编码多肽的核苷酸序列,所述多肽包含与SEQ ID NO:2或4的氨基酸序列具有至少95%序列同一性的氨基酸序列。
22.一种保护植物免受农害侵害的方法,所述方法包括在植物或其细胞中表达编码杀农害多肽的核苷酸序列,其中所述核苷酸序列选自:
a)SEQ ID NO:1或3中所示的核苷酸序列;
b)编码包含SEQ ID NO:2或4的氨基酸序列的多肽的核苷酸序列;和
c)编码多肽的核苷酸序列,所述多肽包含与SEQ ID NO:2或4的氨基酸序列具有至少95%序列同一性的氨基酸序列。
23.根据权利要求22所述的方法,其中所述植物产生杀农害多肽,其中所述多肽具有针对鳞翅目、半翅目、鞘翅目、线虫或双翅目农害的杀农害活性。
24.一种增加植物产率的方法,所述方法包括在田间生长在其基因组中稳定并入了DNA构建体的植物或其种子,所述DNA构建体包含编码具有杀农害活性的蛋白的核苷酸序列,其中所述核苷酸序列选自:
a)SEQ ID NO:1或3中所示的核苷酸序列;
b)编码包含SEQ ID NO:2或4的氨基酸序列的多肽的核苷酸序列;和
c)编码多肽的核苷酸序列,所述多肽包含与SEQ ID NO:2或4的氨基酸序列具有至少95%序列同一性的氨基酸序列;
其中所述田间被农害侵染,其中所述多肽对所述农害具有杀农害活性。
25.权利要求1的核酸用于保护植物免受农害侵害的用途,所述核酸编码的氨基酸对所述农害具有杀农害活性。
26.一种商品,其包含权利要求1的核酸分子或由其编码的蛋白,其中所述产品选自完全或加工的种子或籽粒、动物饲料、玉米或大豆粉状物、玉米或大豆粉、玉米淀粉、大豆粉状物、大豆粉、薄片、大豆蛋白浓缩物、大豆蛋白分离物、组织化大豆蛋白浓缩物、化妆品、护发产品、大豆果仁酱、纳豆、豆豉、水解大豆蛋白、人造黄油、起酥油、卵磷脂、可食用的整粒大豆、大豆酸奶、大豆奶酪、豆腐、腐竹、以及煮熟、抛光、蒸、烤或煮半熟的籽粒。
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CA3234859A1 (en) | 2018-05-31 |
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CN110267975B (zh) | 2024-04-19 |
BR112019010476A2 (pt) | 2019-09-10 |
WO2018098214A1 (en) | 2018-05-31 |
EP3544991A1 (en) | 2019-10-02 |
AU2017365169A1 (en) | 2019-05-30 |
CA3043493A1 (en) | 2018-05-31 |
AU2017365169B2 (en) | 2022-07-21 |
CN118256514A (zh) | 2024-06-28 |
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