CN101173288B - 人工合成的多核苷酸及获得转基因植物的方法 - Google Patents
人工合成的多核苷酸及获得转基因植物的方法 Download PDFInfo
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Abstract
本发明公开了一种人工合成的核苷酸序列,为SEQ ID NO:1所示。该核苷酸序列编码的杀虫蛋白质的氨基酸序列为SEQ ID NO:2所示。本发明还公开了包含上述核苷酸序列的质粒。本发明又公开了上述核苷酸序列的用途,即一种获得转基因植物的方法:稳定地导入了上述核苷酸序列。利用本发明的方法能用于检测转基因植物中是否含有上述核苷酸序列。
Description
技术领域
本发明涉及一种基于人工合成的核苷酸序列及其用途,即通过植物转化而获得抗虫转基因植物的方法。
背景技术
杀虫毒素是一种具有重要运用价值的蛋白质,它可以用来改造杀虫微生物农药,也可以用于转基因抗虫作物。目前转基因抗虫技术已经大量商业化运用(James C.2004 ISAAA Briefs No.32.Ithaca,NY:ISAAA.43pp),为害虫的治理提供了一种低成本的新型技术。目前植物转基因技术也已经是一种成熟技术。
目前杀虫蛋白质有多种。比较常用的是Bacillus thuringiensis(Bt)晶体蛋白质,例如Cry1Ab,Cry1C,Cry2,Cry3等(Crickmore,N.,Zeigler,D.R.,Feitelson,J.,Schnepf,E.,Van Rie,J.,Lereclus,D.,Baum,J.&Dean,D.H.1998 Microbiol.Mol.Biol.Rev.62,807-813)。现在已经商业化推广的转基因抗虫植物都是基于Bt晶体杀虫蛋白质。但是目前已发现多种昆虫可以产生对晶体杀虫蛋白质的抗性。抗性的发展影响了这些晶体毒素的抗虫性能和可利用性(Ferre,J.&Van Rie,J.(2002)Annu.Rev.Entomol.47,501-533)。没有交叉抗性的不同杀虫蛋白质共同使用或轮流使用可以有效减缓害虫抗性的发生(Zhao J-Z,Cao J,Li YX,Collins HL,RoushRT,Earle ED&Shelton AM.2003 Nature Biotechnol.21,1493-1497)。因此,获得基于不同杀虫毒素的抗虫植物,对提高杀虫能力和防止抗性发展具有重要应用价值。
发明内容
本发明要解决的技术问题是提供一种人工合成的核苷酸序列,其可用于降低害虫对农作物的危害。
为了解决上述技术问题,本发明提供一种人工合成的核苷酸序列,如SEQID NO:1所示。
本发明还提供根据上述人工合成的核苷酸序列所编码的杀虫蛋白质,其氨基酸序列为SEQ ID NO:2。
本发明还提供了另一种人工合成的核苷酸序列,其编码的蛋白质包含一个与SEQ ID NO:3有80%或者80%以上的氨基酸序列相同性的区域。
本发明还提供了一种质粒,其包含上述核苷酸序列。
本发明还提供了一种获得转基因植物的方法:稳定地导入了上述核苷酸序列。
作为本发明的获得转基因植物的方法的改进:在农作物中表达的杀虫蛋白质中包含一个与SEQ ID NO:3有80%或者80%以上相同性的区域。
作为本发明的获得转基因植物的方法的进一步改进:植物为水稻、玉米、大豆、棉花、高粱、大麦或小麦。
本发明还提供了一种检测转基因植物中是否含有上述核苷酸序列的方法,包括如下方法中的至少一种:1)、根据上述核苷酸序列设计引物,利用PCR的方法检测;2)、从植物中提取DNA进行Southern杂交;3)、从植物中提取蛋白质进行western分析。
本发明的人工合成的核苷酸序列(SEQ ID NO:1)及其编码的杀虫蛋白质(SEQ ID NO:2)Z1-2,其与以前描述过的核苷酸序列及它们所编码的蛋白质是明显不同的(例如U.S.Pat.No.5,877,012,6,107,279,6,137,033,WO98/18932,WO 99/33991,WO 99/5782,和WO 98/00546)。特别是本发明提供的核苷酸序列所编码的蛋白质在其宿主Bacillus thuringiensis(Bt)中并不表达,也没有分泌;因此不是一种以前定义的Bt营养期杀虫蛋白质(Estruch et al.1996,Proc Natl Acad Sci USA 93,5389-5394)。但是本发明的蛋白质Z1-2和营养期杀虫蛋白质VIP3在氨基酸序列上有同源性。例如,它与Vip3A有79.4%的氨基酸序列相同性(U.S.Pat.No.5849870/5,990,383);与Vip3B有77.5%的相同性(U.S.Pat No.7,244,820);与ISP3a、ISP3b和ISP3c分别有77.6%、78.3%和78.4%的氨基酸序列相同性(U.S.Pat.No.7,265,268);与Vip3C有76.3%的相同性(U.S.Pat申请20050210545)。而Vip3A与Vip3B有82.2%的相同性。Vip3A与Vip3C有81.3%的相同性,Vip3B与Vip3C有84.1%的相同性。
本发明提供的蛋白质Z1-2和中国专利申请号200610049780.4所描述的杀虫蛋白质也有明显不同。Z1-2与200610049780.4中所描述的核苷酸序列所编码的蛋白质有6个氨基酸不同。重要的是两者的杀虫能力也有区别。依照本发明提供的核苷酸序列所获得的转基因水稻在抗稻纵卷叶螟上比利用200610049780.4提供的核苷酸序列具有更高的杀虫能力。利用本发明提供的核苷酸序列获得的转基因水稻有50%的转化事件对稻纵卷叶螟有100%的杀虫活性,而利用200610049780.4的只有7%。此外,本发明更是进一步提供了一种人工合成的核苷酸序列和以这种核苷酸序列转化的抗虫转基因植物。这些转基因植物对至少4种鳞翅目昆虫,二化螟(Chilo suppressalis)、稻纵卷叶螟(Cnaphalocrocis medinalis)、甜菜夜蛾(Spodoptera exigua)和玉米螟(Ostrinia nubilalis)有明显的杀虫能力。利用本发明提供的核苷酸序列获得的转基因水稻有50%的转化事件对稻纵卷叶螟有100%的杀虫活性,而利用200610049780.4的只有7%。
通常一个杀虫基因在E.coli表达的蛋白质的杀虫活性可以预测其转基因植物的抗虫能力。本发明发现SEQ ID NO:1编码的杀虫蛋白质在E.coli中表达的蛋白质对二化螟(C.suppressalis)和稻纵卷叶螟(C.medinalis)没有杀虫能力,对甜菜夜蛾(S.exigua)仅仅有低的杀虫能力(LC50>20,000ng/cm2)。
针对这个特性,本发明提供了一种利用Z1-2控制害虫的方法。本发明是通过这样的技术方案来实现的:1)、提供一种SEQ ID NO:1所示的核苷酸序列;2)、将上述核苷酸序列和一个启动子以及一个终止子功能性地连接而形成一个能够在植物中表达的人工基因;在单子叶植物中启动子可以选择玉米ubiquitin-1启动子、水稻actin启动子等,在双子叶植物中启动子可以是CaMV 35S(Hull and Howell,1987)启动子、Arapdopsis的Ubiquitin-3启动子等;3)这个人工基因被克隆到转化载体,转化植物细胞并且进一步获得转基因植物。
本发明进一步提供了表达Z1-2的转基因水稻。它们对二化螟和稻纵卷叶螟的杀虫能力相当高,其中50%以上的转化事件同时对二化螟和稻纵卷叶螟达100%的杀虫率。因此,可以得出如下结论:根据本发明提供的核苷酸序列所编码的蛋白质在转基因植物上的抗虫能力不能通过E.coli中表达的重组蛋白质的杀虫能力而进行判断和推测。
本发明还进一步提供了一种获得杀虫农作物的方法,在农作物中表达的杀虫蛋白质中包含一个与SEQ ID NO:3有80%或者以上相同性的区域。
本发明还提供了检测上述转基因植物中编码杀虫蛋白质的核苷酸序列和表达的杀虫蛋白质的方法。
本发明还提供了Z1-2的抗体。这种抗体可以用来检测转基因植物中Z1-2的表达。利用抗体检测抗原的方法是已知的技术。
本发明中的杀虫蛋白质Z1-2是指任何包含保持具有杀虫活性的SEQ IDNO:2氨基酸序列最小片段的蛋白质,特别是包含SEQ ID NO:2氨基酸序列中第676到795的120个氨基酸序列片段。Z1-2包括含SEQ ID NO:2氨基酸序列片段的杂交分子,它也包括SEQ ID NO:2的变异体分子。一般认为,功能蛋白质的部分插入、删除变异和点突变对其功能可以没有明显的影响。一种蛋白质的氨基酸序列与SEQ ID NO:2有85%或者以上相同性,可以仍然具有杀虫活性。进一步,一个杀虫蛋白质可以与其他蛋白质分子构建形成有杀虫活性的杂交分子。Z1-2也可以与其他分子,包括VIP3和Bt晶体杀虫蛋白质,构建有杀虫活性的杂交分子;特别是它与其他Bt晶体杀虫蛋白质构建的融合分子。
具体实施方式
本发明的以下实施例所使用分子生物学的方法均为已知的技术。在Ausubel编写的John Wiley and Sons公司出版的Current Protocols in MolecularBiology,和J.Sambrook等编写Cold Spring Harbor Laboratory Press(2001)出版的Molecular Cloning:A Labortory Manual,3rd ED.等文献均有详细的说明。
实施例1、Bt宿主中杀虫蛋白质的检测和杀虫活性的测定:
Z2-1是从苏云菌杆菌的分离菌株HJC-1(Bacillus thuringiensis)克隆获得的。HJC-1在Luria Broth(LB)培养液中培养,280C,250转/分钟震荡。在培养开始后12、24、36、48、60、72、84、96小时取样,分别进行对玉米螟和甜菜夜蛾的杀虫活性进行生物测定。同时,对这些样品利用Z2-1抗体通过Western分析方法进行检测。结果表明离株HJC-1在各个培养时间都没有对玉米螟和甜菜夜蛾产生明显的杀虫或者抑制生长的作用。Western分析进一步表明Z1-2在各个培养时间都没有明显的表达。
实施例2、合成核苷酸序列的克隆和表达:
通过PCR从Bt离株HJC-1中获得了编码Z1-2的核苷酸序列。根据玉米的密码子所有频率,经过人工设计而得到核苷酸序列SEQ ID No:1;上海生工(上海,中国)合成了SEQ ID No:1。进一步利用通用的标准方法将含上述核苷酸序列克隆到表达载体pET28a的BamHI和SacI位点之间,得到表达载体pET28a-VEsyn。此表达载体然后被转入大肠杆菌BL21star。单个菌落接种到100毫升LB细菌培养液,在37℃下震动培养至OD600=0.6,然后加IPTG(lsopropyl-β-D-thiogalactoside)到0.5mM,并继续在同样的条件下培养4小时。培养液经过5000g离心10分钟沉淀大肠杆菌细胞,然后弃上清收集沉淀。沉淀中加30毫升20mM Tris-HCl缓冲液,超声粉碎。这样获得的重组蛋白质用来制备抗体和杀虫活性的测定。
实施例3、抗体的产生:
通过实施例2而表达的Z1-2蛋白质,经过SDS-PAGE分离后从胶上切出。使用100μg的Z1-2蛋白质加完全福氏佐剂,皮下多点注射进行初次免疫,以后每间隔2周后进行第2次免疫、第3次免疫。在Western分析检测效价结果为5000倍时,收集、离心分离血清,放置-80度冰箱保存。
实施例4、E.coli表达的重组Z1-2蛋白质的杀虫活性:
将实施例2所获得的蛋白涂在昆虫人工饲料的表面,放置2小时后,接上新生一龄幼虫进行杀虫活性测定。阴性对照的准备方法与实施例2相同,但质粒为不含任何插入DNA的pET 28a载体本身。在25℃条件下,7天后记录杀虫数目。杀虫活性结果如下表1所示:
表1
二化螟 | 稻纵卷叶螟 | 甜菜夜蛾 | |
本发明 | 0% | 0% | 80% |
阴性对照 | 0% | 0% | 0% |
Bt杀虫剂 | 100% | 100% | 100% |
实施例5、农杆菌转化T-DNA载体的构建:
农杆菌转化T-DNA载体是基于pCambia 1300载体而构建的。编码杀虫蛋白质Z1-1(SEQ ID No:4)和Z1-2的合成核苷酸序列分别被与玉米ubiquitin-1启动子和Nos终止子功能性地连接而形成一个能够在植物中表达的人工基因。它的5’端被设计上BamHI位点,3’端被设计上KpnI位点。这个人工基因被进一步克隆到pCambia 1300多克隆位点的BamHI和KpnI之间,获得T-DNA载体pCAM1300-Z1-1和pCAM1300-Z1-2。由于玉米ubiquitin-1启动子在禾本科植物中均有启动子活性,pCAM1300-Z1-1和pCAM1300-Z1-2可以作为多种禾本科植物的抗虫转化T-DNA载体。
实施例6转基因水稻的获得:
转基因植物的获得方法是采用现有技术(卢雄斌龚祖埙1998生命科学10:125-131;刘凡等,2003分子植物育种1:108-115)。选取成熟饱满种子去壳,诱导产生愈伤组织作为转化材料。取含目的基因的农杆菌(pCAM1300-Z1-1和pCAM1300-Z1-2)划板,挑单菌落接种准备转化用农杆菌。将待转化的愈伤组织放入适当浓度的农杆菌液中(含乙酰丁香酮),让农杆菌结合到愈伤组织表面,然后把愈伤组织转移到共培养基中,共培养2~3天。用无菌水冲洗转化后的愈伤,转移到含适当抗生素的筛选培养基上,筛选培养(50ng/mL潮霉素)两个月(中间继代一次)。把筛选后,生长活力良好的愈伤转移到预分化培养基上培养20大左右,然后将预分化好的愈伤组织移到分化培养基,14小时光照分化发芽。2-3周后,把抗性再生植株转移到生根培养基上壮苗生根,最后将再生植株洗去琼脂移植于温室,作为鉴定材料。
实施例7、转基因玉米的获得:
取授粉后8-10天的Hi-2玉米穗。收集所有的未成熟胚(大小为1.0-1.5mm)。将含有T-DNA载体pCAM1300-Z1-2的农杆菌与未成熟胚共培育共培养2-3天(22℃)。转移未成熟胚到愈伤诱导培养基上(含200mg/L的Timentin杀农杆菌),28℃暗培养10-14天。将所有的愈伤转到带有50ng/mL潮霉素的筛选培养基上,28℃暗培养2-3周。
转移所有的组织到新鲜潮霉素的筛选培养基上,28℃暗培养2-3周。然后,转移所有筛选后成活的胚性组织到再生培养基上,28℃暗培养10-14天,每皿一个株系。转移胚性组织到新鲜的再生培养基上,26℃光照培养10-14天。转移所有发育完全的植株到生根培养基上,26℃光照培养直到根发育完全。
实施例8、转基因棉花的获得:
利用CaMV 35S启动子或者Arapdopsis的Ubiquitin-3启动子控制的Z1-2基因可以通过Agribacterium介导的方法或者花粉管方法导入棉花。棉花的Agribacterium介导方法或者花粉管导入方法是有知的技术。
实施例9、转基因水稻的抗虫能力测定:
分别取转Z1-1和Z1-2基因的水稻4叶期的叶片,每个叶片分别接5头二化螟或者稻纵卷叶螟初孵幼虫。在养虫室观测5天。结果如下表2所示:
表2
基因 | Z1-1 | Z1-2(本发明) |
总转化事件数目 | 15 | 20 |
100%杀二化螟的转化事件数目(%) | 10(67% ) | 16(80%) |
100%杀稻纵卷叶螟的转化事件数目(%) | 1(7%) | 10(50%) |
实施例10、转基因水稻的检测:
PCR检测:PCR引物分别是:5’TG GAC AGC AGC GAC GTG TAC CTGGTG和5’CCT GAT GCT GAA GTT GTA GAA GTA G。水稻的总基因组DNA为模板。结果表明转Z1-2基因水稻有460bp DNA片段放大,而非转基因水稻对照则没有。
Western分析:利用Z1-2抗体,通用的Western分析方法对水稻叶片中的总蛋白质中的Z1-2进行检测。取水稻叶片,在液氮中粉碎,再与SDS样品溶液混合、加热制备SDS-PAGE样品。结果表明转Z1-2基因水稻有大约88kD的Z1-2蛋白质表达,而非转基因水稻对照没有。
最后,还需要注意的是,以上列举的仅是本发明的若干个具体实施例。显然,本发明不限于以上实施例,还可以有许多变形。本领域的普通技术人员能从本发明公开的内容直接导出或联想到的所有变形,均应认为是本发明的保护范围。
序列表
SEQ ID N0:1
1 ATGAACATGA ACAACACCAA GCTCAACGCC CGCGCCCTCC CGTCCTTCAT CGACTACTTC
61 AACGGCATCT ACGGCTTCGC CACCGGCATC AAGGACATCA TGAACATGAT CTTCAAGACC
121 GACACCGGCG GCGACCTCAC CCTCGACGAG ATCCTCAAGA ACCAGCAGCT CCTCAACGAG
181 ATCTCCGGCA AGCTCGACGG CGTGAACGGC TCCCTCAACG ACCTCATCGC CCAGGGCAAC
241 CTCAACACCG AGCTATCCAA GGAGATCCTC AAGATCGCCA ACGAGCAGAA CCAGGTGCTC
301 AACGACGTGA ACAACAAGCT CGACGCCATC AACACCATGC TCCACATCTA CCTCCCGAAG
361 ATCACCTCCA TGCTCTCCGA CGTGATGAAG CAGAACTACG CCCTCTCCCT CCAGATCGAG
421 TACCTCTCCA AGCAGCTCCA GGAGATCTCC GACAAGCTCG ACATCATCAA CGTGAACGTG
481 CTCATCAACT CCACCCTCAC CGAGATCACC CCGGCCTACC AGCGCATCAA GTACGTGAAC
541 GAGAAGTTCG AGGAGCTGAC CTTCGCCACC GAGACCAACC TCAAGGTGAA GAAGGACGGC
601 TCCCCGGCCG ACATCCTCGA CGAGCTAACC GAGCTTACCG AGCTGGCCAA GTCCGTGACC
661 AAGAACGACG TGGACGGCTT CGAGTTCTAC CTCAACACCT TCCACGACGT GATGGTGGGC
721 AACAACCTCT TCGGCCGCTC CGCCCTCAAG ACCGCCTCCG AACTCATCAC CAAGGAGAAC
781 GTGAAGACCT CCGGCTCCGA GGTGGGCAAC GTGTACAACT TCCTCATCGT GCTCACCGCC
841 CTCCAGGCCA AGGCCTTCCT CACCCTCACC ACCTGCCGCA AGCTCCTCGG CCTCGCCGAC
901 ATCGACTACA CCTCCATCAT GAACGAGCAC CTCAACAAGG AGAAGGAGGA GTTCCGCGTG
961 AACATCCTCC CGACCCTCTC CAACACCTTC TCCAACCCGA ACTACATCAA GACCAAGGGC
1021 TCCGACGAGG ACGCCGAGGT GATCATCCAG GCCGAGCCGG GCCACGCCCT CGTGGGCTTC
1081 GAGATGATCA ACGACCCGTC CCCGGCCCTC AAGGTGTACC AGGCCAAGCT CACCACCAAC
1141 TACCAGGTGG ACAAGCAGAG CCTGAGCGAG ACCGTGTACG GCGACATGGA CAAGATCCTG
1201 TGCCCGGACA AGAGCCAGCA GATGTACTAC CTGCACAACA TCACCTTCCC GAACGAGTAC
1261 GTGATCACCG AGATCATCTT CACCAAGAAG AAGAACAGCC TGAGGTACGA GGTGATCGCC
1321 AACTACTACG AGTTCAGCAG CGGCGACATC GACCTGAACA AGAAGCTGGT GAAGAGCAGC
1381 GAGGCCGAGT ACAGCACCCT GAGCGTGAGC AACGACGCCA TCTACATGCC GCTGGGCGTG
1441 ATCAGCGAGA CCTTCCTGAC CCCGATCAAG GGCTTCGGCC TGACCGTGGA CGAGAGCAGC
1501 AGGCTGGTGA CCCTGACCTG CAAGAGCTAC CTGAGGGAGA TCCTGCTGGC CACCGACCTG
1561 AGCAACAAGG CCACCAAGCT GATCGTGCCG CCGAACGGCT TCATCAGCAA CCTGGTGGAG
1621 AACGGCGACA TCGAGGCCGA CAACATCGAG CCGTGGAAGG GCAACAACAA GAACGCCTAC
1681 GTGGACCACA CCGGCGGCGT GAACGGCACC AAGGCCCTGT ACACCCAGGA CGACGGCGAG
1741 TTCAGCCAGT TCATCGGCGA CAAGCTGAAG AGCAAGACCG AGTACATCAT CCAGTACACC
1801 GTGAAGGGCA ACACCAGCAT CTACCTGAAG GACAAGAAGA ACGAGAACGT GATCTACGAG
1861 GACAAGAACA ACAACCTGGA GGCCTTCCAG ACCATCACCA AGAGGTTCAC CACCGAGCTG
1921 GACAGCAGCG ACGTGTACCT GGTGTTCAAG TGCAAGAACG GCTACAAGGC CTGGGGCGAC
1981 AACTTCCTGA TCACCGAGAT CAGGCCGAAG GAGGTGGTGA GCCCGGAGCT GATCAAGGTG
2041 GAGAACTGGA TCGGCATGGG CGGCAGCAAC CACGTGAACC CGGACAGCCT GCTGCTGTTC
2101 ACCGGCGGCA GGAGCATCCT GAAGCAGAAC CTGCAGCTGG ACAGCTACAG CACCTACAGG
2161 GTGAGGTTCA GCCTGATGGT GATCGGCAAG GCCAAGGTGA TCATCAGGAA CAGCAGCGAG
2221 GTGCTGTTCG AGAAGAGCTA CGTGAACGAC AGCGAGGGCG TGCTGGAGGG CGTGAGCGAG
2281 ACCTTCACCA CCAAGAGCAT CCAGGACAAC TTCTACGTGG AGCTGAGCAA CGAGGGCACC
2341 TTCGGCAGCA AGGACGTGGC CTACTTCTAC AACTTCAGCA TCAGGTAA
SEQ ID NO:2:
1 Met Asn Met Asn Asn Thr Lys Leu Asn Ala Arg Ala Leu Pro Ser Phe Ile Asp Tyr Phe
21 Asn Gly Ile Tyr Gly Phe Ala Thr Gly Ile Lys Asp Ile Met Asn Met Ile Phe Lys Thr
41 Asp Thr Gly Gly Asp Leu Thr Leu Asp Glu Ile Leu Lys Asn Gln Gln Leu Leu Asn Glu
61 Ile Ser Gly Lys Leu Asp Gly Val Asn Gly Ser Leu Asn Asp Leu Ile Ala Gln Gly Asn
81 Leu Asn Thr Glu Leu Ser Lys Glu Ile Leu Lys Ile Ala Asn Glu Gln Asn Gln Val Leu
101 Asn Asp Val Asn Asn Lys Leu Asp Ala Ile Asn Thr Met Leu His Ile TVr Leu Pro Lys
121 Ile Thr Ser Met Leu Ser Asp Val Met Lys Gln Asn Tyr Ala Leu Ser Leu Gln Ile Glu
141 Tyr Leu Ser Lys Gln Leu Gln Glu Ile Ser Asp Lys Leu Asp Ile Ile Asn Val Asn Val
161 Leu Ile Asn Ser Thr Leu Thr Glu Ile Thr Pro Ala Tyr Gln Arg Ile Lys Tyr Val Asn
181 Glu Lys Phe Glu Glu Leu Thr Phe Ala Thr Glu Thr Asn Leu Lys Val Lys Lys Asp Gly
201 Ser Pro Ala Asp Ile Leu Asp Glu Leu Thr Glu Leu Thr Glu Leu Ala Lys Ser Val Thr
221 Lys Asn Asp Val Asp Gly Phe Glu Phe Tyr Leu Asn Thr Phe His Asp Val Met Val Gly
241 Asn Asn Leu Phe Gly Arg Ser Ala Leu Lys Thr Ala Ser Glu Leu Ile Thr Lys Glu Asn
261 Val Lys Thr Ser Gly Ser Glu Val Gly Asn Val Tyr Asn Phe Leu Ile Val Leu Thr Ala
281 Leu Gln Ala Lys Ala Phe Leu Thr Leu Thr Thr Cys Arg Lys Leu Leu Gly Leu Ala Asp
301 Ile Asp Tyr Thr Ser Ile Met Asn Glu His Leu Asn Lys Glu Lys Glu Glu Phe Arg Val
321 Asn Ile Leu Pro Thr Leu Ser Asn Thr Phe Ser Asn Pro Asn Tyr Ile Lys Thr Lys Gly
341 Ser Asp Glu Asp Ala Glu Val Ile Ile Gln Ala Glu Pro Gly His Ala Leu Val Gly Phe
361 Glu Met Ile Asn Asp Pro Ser Pro Ala Leu Lys Val Tyr Gln Ala Lys Leu Thr Thr Asn
381 Tyr Gln Val Asp Lys Gln Ser Leu Ser Glu Thr Val Tyr Gly Asp Met Asp Lys Ile Leu
401 Cys Pro Asp Lys Ser Gln Gln Met Tyr Tyr Leu His Asn Ile Thr Phe Pro Asn Glu Tyr
421 Val Ile Thr Glu Ile Ile Phe Thr Lys Lys Lys Asn Ser Leu Arg Tyr Glu Val Ile Ala
441 Asn Tyr Tyr Glu Phe Ser Ser Gly Asp Ile Asp Leu Asn Lys Lys Leu Val Lys Ser Ser
461 Glu Ala Glu Tyr Ser Thr Leu Ser Val Ser Asn Asp Ala Ile Tyr Met Pro Leu Gly Val
481 Ile Ser Glu Thr Phe Leu Thr Pro Ile Lys Gly Phe Gly Leu Thr Val Asp Glu Ser Ser
501 Arg Leu Val Thr Leu Thr Cys Lys Ser Tyr Leu Arg Glu Ile Leu Leu Ala Thr Asp Leu
521 Ser Asn Lys Ala Thr Lys Leu Ile Val Pro Pro Asn GlV Phe Ile Ser Asn Leu Val Glu
541 Asn Gly Asp Ile Glu Ala Asp Asn Ile Glu Pro Trp Lys Gly Asn Asn Lys Asn Ala Tyr
561 Val Asp His Thr Gly Gly Val Asn Gly Thr Lys Ala Leu Tyr Thr Gln Asp Asp Gly Glu
581 Phe Ser Gln Phe Ile Gly Asp Lys Leu Lys Ser Lys Thr Glu Tyr Ile Ile Gln Tyr Thr
601 Val Lys Gly Asn Thr Ser Ile Tyr Leu Lys Asp Lys Lys Asn Glu Asn Val Ile Tyr Glu
621 Asp Lys Asn Asn Asn Leu Glu Ala Phe Gln Thr Ile Thr Lys Arg Phe Thr Thr Glu Leu
641 Asp Ser Ser Asp Val Tyr Leu Val Phe Lys Cys Lys Asn Gly Tyr Lys Ala Trp Gly Asp
661 Asn Phe Leu Ile Thr Glu Ile Arg Pro Lys Glu Val Val Ser Pro Glu Leu Ile Lys Val
681 Glu Asn Trp Ile Gly Met Gly Gly Ser Asn His Val Asn Pro Asp Ser Leu Leu Leu Phe
701 Thr Gly Gly Arg Ser Ile Leu Lys Gln Asn Leu Gln Leu Asp Ser Tyr Ser Thr Tyr Arg
721 Val Arg Phe Ser Leu Met Val Ile Gly Lys Ala Lys Val Ile Ile Arg Asn Ser Ser Glu
741 Val Leu Phe Glu Lys Ser Tyr Val Asn Asp Ser Glu Gly Val Leu Glu Gly Val Ser Glu
761 Thr Phe Thr Thr Lys Ser Ile Gln Asp Asn Phe Tyr Val Glu Leu Ser Asn Glu Gly Thr
781 Phe Gly Ser Lys Asp Val Ala Tyr Phe Tyr Asn Phe SerIle Arg
SEQ ID NO:3
1 Glu Leu Ile Lys Val Glu Asn Trp Ile Gly Met Gly Gly Ser Asn His Val Asn Pro Asp
21 Ser Leu Leu Leu Phe Thr Gly Gly Arg Ser Ile Leu Lys Gln Asn Leu Gln Leu Asp Ser
41 Tyr Ser Thr Tyr Arg Val Arg Phe Ser Leu Met Val Ile Gly Lys Ala Lys Val Ile Ile
61 Arg Asn Ser Ser Glu Val Leu Phe Glu Lys Ser Tyr Val Asn Asp Ser Glu Gly Val Leu
81 Glu Gly Val Ser Glu Thr Phe Thr Thr Lys Ser Ile Gln Asp Asn Phe Tyr Val Glu Leu
101 Ser Asn Glu Gly Thr Phe Gly Ser Lys Asp Val Ala Tyr Phe Tyr Asn Phe Ser Ile Arg
SEQ ID NO:4
1 ATGAACAAGA ACAACACCAA GCTCAGCACC CGCGCCCTCC CGTCCTTCAT CGACTACTTC
61 AACGGCATCT ACGGCTTCGC CACCGGCATC AAGGACATCA TGAACATGAT CTTCAAGACC
121 GACACCGGCG GCGACCTCAC CCTCGACGAG ATCCTCAAGA ACCAGCAGCT CCTCAACGAG
181 ATCTCCGGCA AGCTCGACGG CGTGAACGGC TCCCTCAACG ACCTCATCGC CCAGGGCAAC
241 CTCAACACCG AGCTATCCAA GGAGATCCTC AAGATCGCCA ACGAGCAGAA CCAGGTGCTC
301 AACGACGTGA ACAACAAGCT CGACGCCATC AACACCATGC TCCACATCTA CCTCCCGAAG
361 ATCACCTCCA TGCTCTCCGA CGTGATGAAG CAGAACTACG CCCTCTCCCT CCAGATCGAG
421 TACCTCTCCA AGCAGCTCCA GGAGATCTCC GACAAGCTCG ACATCATCAA CGTGAACGTG
481 CTCATCAACT CCACCCTCAC CGAGATCACC CCGGCCTACC AGCGCATCAA GTACGTGAAC
541 GAGAAGTTCG AGGAGCTGAC CTTCGCCACC GAGACCAACC TCAAGGTGAA GAAGGACGGC
601 TCCCCGGCCG ACATCCTCGA CGAGCTAACC GAGCTTACCG AGCTGGCCAA GTCCGTGACC
661 AAGAACGACG TGGACGGCTT CGAGTTCTAC CTCAACACCT TCCACGACGT GATGGTGGGC
721 AACAACCTCT TCGGCCGCTC CGCCCTCAAG ACCGCCTCCG AACTCATCAC CAAGGAGAAC
781 GTGAAGACCT CCGGCTCCGA GGTGGGCAAC GTGTACAACT TCCTCATCGT GCTCACCGCC
841 CTCCAGGCCA AGGCCTTCCT CACCCTCACC ACCTGCCGCA AGCTCCTCGG CCTCGCCGAC
901 ATCGACTACA CCTCCATCAT GAACGAGCAC CTCAACAAGG AGAAGGAGGA GTTCCGCGTG
961 AACATCCTCC CGACCCTCTC CAACACCTTC TCCAACCCGA ACTACATCAA GACCAAGGGC
1021 TCCGACGAGG ACGCCGAGGT GATCATCCAG GCCGAGCCGG GCCACGCCCT CGTGGGCTTC
1081 GAGATGATCA ACGACCCGTC CCCGGCCCTC AAGGTGTACC AGGCCAAGCT CAAGCCCAAC
1141 TACCAGGTGG ACAAGCAGAG CCTGAGCGAG ACCGTGTACG GCGACATGGA CAAGATCCTG
1201 TGCCCGGACA AGAGCCAGCA GATGTACTAC CTGCACAACA TCACCTTCCC GAACGAGTAC
1261 GTGATCACCG AGATCATCTT CACCAAGAAG AAGAACAGCC TGAGGTACGA GGTGATCGCC
1321 AACTACTACG AGTTCAGCAG CGGCGACATC GACCTGAACA AGAAGCTGGT GAAGAGCAGC
1381 GAGGCCGAGT ACAGCACCCT GAGCGTGAGC AACGACGCCA TCTACATGCC GCTGGGCGTG
1441 ATCAGCGAGA CCTTCCTGAC CCCGATCAAG GGCTTCGGCC TGACCGTGGA CGAGAGCAGC
1501 AGGCTGGTGA CCCTGACCTG CAAGAGCTAC CTGAGGGAGA TCCTGCTGGC CACCGACCTG
1561 AGCAACAAGG CCACCAAGCT GATCGTGCCG CCGAACGGCT TCATCAGCAA CCTGGTGGAG
1621 AACGGCGACA TCGAGGCCGA CAACATCGAG CCGTGGAAGG GCAACAACAA GAACGCCTAC
1681 GTGGACCACA CCGGCGGCGT GAACGGCACC AAGGCCCTGT ACACCCAGGA CGACGGCGAG
1741 TTCAGCCAGT TCATCGGCGA CAAGCTGAAG AGCAAGACCG AGTACATCAT CCAGTACACC
1801 GTGAAGGGCA ACACCAGCAT CTACCTGAAG GACAAGAAGA ACGAGAACGT GATCTACGAG
1861 GACAAGAACA ACAACCTGGA GGCCTTCCAG ACCATCACCA AGAGGTTCAC CACCGAGCTG
1921 GACAGCAGCG ACGTGTACCT GGTGTTCAAG TGCAAGAACG GCTACAAGGC CTGGGGCGAC
1981 AACTTCCTGA TCACCGAGAT CAGGCCGAAG GAGGTGGTGA GCCCGGAGCT GATCAAGGTG
2041 GAGAACTGGA TCGGCATGGG CGGCAGCAAC CACGTGAACC CGGACAGCCT GCTGCTGTTC
2101 ACCGGCGGCA GGAGCATCCT GAAGCAGAAC CTGCAGCTGG ACAGCTACAG CACCTACAGG
2161 GTGAGGTTCA GCCTGATGGT GATCGGCAAG GCCAAGGTGA TCATCAGGAA CAGCAGCGAG
2221 GTGCTGTTCG AGAAGAGCTA CGTGAACGAC AGCGAGGGCG TGCTGGAGGG CGTGAGCGAG
2281 ACCTTCACCA CCAAGAGCAT CCAGGACAAC TTCTACGTGG AGCTGAGCAA CGAGGGCACC
2341 TTCGGCAGCA AGGACGTGGC CTACTTCTAC AACTTCAGCA TCAAGTAA
Claims (6)
1.一种人工合成的多核苷酸,其特征是核苷酸序列如SEQ ID NO:1所示。
2.根据权利要求1所述人工合成的多核苷酸编码的杀虫蛋白质,其特征是:其氨基酸序列为SEQ ID NO:2。
3.一种质粒,其包含权利要求1所述的核苷酸序列。
4.一种获得转基因植物的方法,其特征是:稳定地导入了如权利要求1所述的核苷酸序列。
5.根据权利要求4所述的转基因植物的生产方法,其特征是:所述植物为水稻、玉米、大豆、棉花、高粱、大麦或小麦。
6.一种检测转基因植物中是否含有权利要求1所述核苷酸序列的方法,其特征是包括如下方法中的至少一种:1)、根据权利要求1或3所述的核苷酸序列设计引物,利用PCR的方法鉴测;2)、从植物中提取DNA进行Southern杂交;3)、从植物中提取蛋白质进行western分析。
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