CN101591381A - Bt蛋白Cry4Cb1、其编码基因及应用 - Google Patents
Bt蛋白Cry4Cb1、其编码基因及应用 Download PDFInfo
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- CN101591381A CN101591381A CNA2009100815961A CN200910081596A CN101591381A CN 101591381 A CN101591381 A CN 101591381A CN A2009100815961 A CNA2009100815961 A CN A2009100815961A CN 200910081596 A CN200910081596 A CN 200910081596A CN 101591381 A CN101591381 A CN 101591381A
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Abstract
本发明提供了一种新的Bt蛋白Cry4Cb1及其编码基因,所述蛋白具有SEQ ID No.2所示的氨基酸序列,或SEQ ID No.2所示的氨基酸序列经取代、缺失和/或增加一个或多个氨基酸且具有同等活性的蛋白。本发明蛋白可以用于制备Bt杀虫剂,所述基因可以转化棉花、玉米、水稻、蔬菜等农作物,使其具备相应的抗虫活性,从而降低农药的使用量,减少环境污染,具有重要的经济价值和应用前景。
Description
技术领域
本发明涉及生物技术领域,具体涉及一种新的Bt蛋白及其编码基因和应用。
背景技术
在人类生产过程中,虫害是造成农业生产损失及影响人类健康的重要因素,据FAO统计,全世界农业生产每年因虫害造成的经济损失高达14%,病害损失达12%,草害损失达11%。损失额高达1260亿美元,相当于中国农业总产值的一半,英国的4倍多。另外,蚊媒病在预防医学中占有重要位置,其中登革热和黄热病等蚊媒病传播力强、流行面广、发病率高、危害性大。据WHO统计,全世界每年感染登革热人数多达8000万,我国的海南省在1980和1986年曾经暴发过两次登革热,发病分别达到437469例和113589例。登革热和黄热病主要由埃及伊蚊传播。
为了减少这些损失,多年来,对农作物害虫及蚊虫普遍采用化学防治手段进行防治,但由于化学农药的长期、大量使用,造成了对环境的污染,农副产品中农药残留量增加,给人类的生存和健康带来了危害。此外,化学农药在杀灭害虫的同时,也杀伤了天敌及其它有益物,破坏了生态平衡。与化学防治相比,生物防治具有安全、有效、持久的特点。并且避免了化学防治带来的一系列问题。因此,生物防治技术成了人们研究的热点。在生物杀虫剂中,苏云金芽孢杆菌是目前世界上用途最广、产量最大的一类微生物杀虫剂。
苏云金芽孢杆菌(Bacillus thuringiensis,简称Bt)是一种革兰氏阳性细菌,它的分布极为广泛,在芽孢形成的同时可形成具有杀虫活性的由蛋白质组成的伴胞晶体,又名杀虫晶体蛋白(Insectididal crystalproteins,简称ICPs),ICPs是由cry基因编码的,对敏感昆虫有强烈毒性,而对高等动物和人无毒性。近几十年来,Bt已广泛应用于控制多种鳞翅目、双翅目、鞘翅目等害虫。此外,Bt还对膜翅目、同翅目、直翅目、食毛目等多种害虫及植物病原线虫、螨类、原生动物有控害作用。目前在农田害虫、森林害虫及卫生害虫的防治中Bt已成为化学合成农药的有力替代品,Bt还是转基因抗虫工程植物重要的基因来源。
自1981年Schnepf从菌株HD-1Dipel中克隆了第一个能表达杀虫活性的基因以来(Adang M.J et al,Characterized full-length andtruncated plasmid clones of the crystal protein of Bacillus thuringiensissubsp.kurstaki HD-73 and their toxicity to Manduca sexta,Gene,1985,36(3):289~300.),人们已经分离克隆了390多种编码杀虫晶体蛋白的基因,根据编码的氨基酸序列同源性它们被分别确定为不同的群、亚群、类和亚类(Crickmore N,Zeigler D R,Feitelson J,et al.Revision of the nomenclature for the Bacillus thuringiensis pesticidalcrystal proteins.Microbiol Mol Biol Rev,1998,62:807-813;http://www.biols.susx.ac.uk/Home/Neil_Crickmore/Bt/)。一般而言,Cry1,Cry2和Cry9等毒蛋白对鳞翅目害虫有效;其中研究的最多的是Cry1和Cry9类蛋白,它们编码的杀虫晶体蛋白分子量为130-140kD,许多基因目前已被广泛应用于植物的鳞翅目害虫的防治(Kozie,M.G.,Beland,G.L.,Bowman,C.,et al.Field performance of elite transgenicmaize plants expressing an insecticidal protein derived from Bacillusthuringiensis.Bio/Technology,1993,11:194-200;Perlak,F.J.,Deaton,R.W.,Armstrong,T.A.,et al.Insect resistant cotton plants.bio/technology,1990;8:939-943;Van Frankenhuyzen,K.,Gringorten,L.,and Gauhier,D.1997.Cry9Ca1 toxin,a Bacillus thuringiensis insecticidal crystalprotein with high activity against the spruce bud worm(Choristoneurafnniferana).Appl.Environ,Microbviol.63:4132-4134;王飞,2001,苏云金芽孢杆菌特异菌株生物学特性及cry9新基因的研究,硕士论文,南开大学)。苏云金芽胞杆菌以色列亚种(B.thuringiensis subsp.israelensis,简称Bti)产生的毒素蛋白对蚊虫具有很好杀虫活性,被广泛运用于蚊虫的防治(Goldberg L J,and Margalit J,1977.A bacterialspore demonstrating rapid larvicidal activity against Anopheles sergentii,Uranotaenia unguiculata,Culex univitattus,Aedes aegypti,and Culexpipiens.Mosqito News,37:355-358;)。同时,Cyt蛋白具有溶细胞性,对某些Cry蛋白具有增效作用及延缓昆虫的抗性(Wu,D.,Johnson,J.J.,and Federici,B.A.1994.Synergism of mosquitocidal toxicity betweenCytA and CryIVD Proteins using inclusion sproduced from clonedgenes of Bacillus thuringiensis.Mol.Microbiol.13:965-972;Wirth,M.C.,Georghiou,G.P.,and Federeci,B.A.1997.CytA enables CryIVendotoxins of Bacillus thuringiensis to overcome high levels of CryIVresistance in the mosquito,Culex quinquefasciatus.Proc.Natl.Acad.Sci.94:10536-10540)
自本世纪初发现苏云金芽胞杆菌至今已有100多年的历史,在农作物和园艺植物害虫、森林害虫以及卫生害虫的防治方面得到广泛的应用,也起到良好的效果。但是,由于大规模和反复使用苏云金芽胞杆菌,许多昆虫种群已相继在不同程度上对杀虫晶体蛋白产生了抗性。以Bt杀虫晶体蛋白为基础的杀虫剂的使用已有50多年的历史,最初一直没有检测到昆虫对Bt的抗性,但是,上世纪80年中期开始,抗性问题不断在实验室及田间试验中得到证实(M cGaughey,W.H.1985.Insect resistance to the biological insecticide Bacillus thuringiensis.Science.229:193-195),原因主要是持续使用单品种及亚致剂量的Bt以及Bt转基因抗虫植物的应用造成昆虫种群长期受到杀虫剂的选择压力。1985年,McGaughey报道仓库谷物害虫印度谷螟(Plodia interpunctella)在Dipel(Bt subsp.kurstaik HD-1的商品制剂)的选择压力下,繁殖15代后,抗性增加97倍;在高剂量选择压力下,抗性可增加250倍。1990年,在夏威夷首次证实大田中的小菜蛾对Bt杀虫剂产生了明显的抗性(Tabashnik,B.E.,Finson,N.,Groeters,F.R.,et al.1994.Reversal of resistance to Bacillus thuringiensisin Plutella xylostella.Proc.Natl.Acad.Sci.USA.91:4120-4124),上世纪90年代以来,在我国应用Bt杀虫剂时间较长的深圳、广州、上海等地,发现Bt杀虫剂对小菜蛾防治效果明显下降,意味着抗性已经形成(冯夏.1996.广东小菜蛾对苏云金杆菌的抗性研究.昆虫学报,39(3):238-244;Hofte,H.,Van Rie,J.,Jansens,S.,Van Houtven,A.,Vanderbruggen,H.,and Vaeck,M.,1988.Monoclonal antibody analysis and insecticidal spectrum ofthree types of lepidopteran-specific insecticidal crystal proteins ofBacillus thuringiensis.Appl.Environ.Microbiol.54:2010-2017)。目前发现在实验室及田间至少有十几种昆虫对Bt及其杀虫晶体蛋白产生了抗性,用选择压力数学模型预测到,在Bt转基因抗虫植物选择压力的条件下,昆虫将会产生抗性(Schnepf,E.,Crickmore,N.,Van Pie,J.,et al. 1998.Bacillus thuringiensis and its pesticidal Crystalproteins.Microbiol.Mol.Biol.Rev.65(3):775-806)。另外,有研究证明Bti在大田的使用中尚未发现抗性问题(Regis L,et al.,2000.The useof bacterial larvicides in mosquito and black fly control programsinBrazil.Mem.Instituto Oswaldo Cruz,95:207-210.),但是蚊虫对其抗性问题不断在实验室中得到证实,这种情况也可能会在大田中出现(Georghiou G P,and Wirth M C,1997.Influence of exposure to singleversus multiple toxins of Bacillus thuringiensis subsp.israelensis ondevelopment of resistance in the mosquito Culex quinquefasciatus(Diptera:Culicidae).Applied and Environmental Microbiology,63:1095-1101.)。
为避免抗性昆虫所造成的损失,寻找新的高毒力基因资源是解决这个问题的有效途径,这对我国的生物防治有着十分重要的意义。
发明内容
本发明的第一个目的在于针对上述不足提供一种新的BT毒力蛋白资源。
本发明的第二个目的在于提供编码所述蛋白的基因。
本发明的目的还在于提供上述蛋白及基因的应用。
本发明从四川省成都平原土壤中分离得到的苏云金芽孢杆菌(Bacillus thuringiensis)新菌株HS18-1。通过对HS18-1的毒力测试表明,HS18-1对鞘翅目害虫、鳞翅目害虫、双翅目害虫等等,均具有极高的毒力。
根据Cry4类基因保守序列设计1对特异引物,扩增其基因组DNA,结果表明该菌株存在Cry4类基因,进一步设计其全长基因引物,克隆得到Cry4Ba基因,其核苷酸序列如序列表SEQ ID No.1所示,序列SEQ ID No.1的全长为3474bp,分析表明,GC含量为35.90%,编码1157个氨基酸组成的蛋白。经测定,其氨基酸序列如SEQ IDNo.2所示。在softberry网站采用bacterial sigma7.0 promoter程序对全序列进行预测表明,在基因编码区上游含有RNA聚合酶活化位点的序列,将该基因命名为cry4Cb1。本发明进一步分析了Cry4Cb1蛋白的氨基酸组成(见表1)。
表1 Cry4Cb1蛋白的氨基酸组成
氨基酸 | 百分比% | 氨基酸 | 百分比% |
Ala(A): | 5.96 | Met(M): | 2.07 |
Cys(C): | 1.12 | Asn(N): | 7.78 |
Asp(D): | 5.53 | Pro(P): | 3.98 |
Glu(E): | 5.01 | Gln(Q): | 4.67 |
Phe(F): | 3.28 | Arg(R): | 3.37 |
Gly(G): | 5.96 | Ser(S): | 6.91 |
His(H): | 2.68 | Thr(T): | 7.95 |
Ile(I): | 6.40 | Val(V): | 5.70 |
Lys(K): | 5.70 | Trp(W): | 1.30 |
Leu(L): | 8.82 | Tyr(Y): | 5.79 |
应当理解,本领域技术人员可根据本发明公开的氨基酸序列,在不影响其活性的前提下,取代、缺失和/或增加一个或几个氨基酸,得到所述蛋白的突变序列。例如在非活性区段,将第20位的Ser替换为Met。因此,本发明Bt蛋白还包括SEQ ID No.2所示氨基酸序列经取代、替换和/或增加一个或几个氨基酸,具有Cry4Cb1蛋白同等活性的由Cry4Cb1衍生得到的蛋白质。本发明基因包括编码所述蛋白的核酸序列。
此外,应理解,考虑到密码子的简并性以及不同物种密码子的偏爱性,本领域技术人员可以根据需要使用适合特定物种表达的密码子。
本发明的基因和蛋白质可以从菌株HS18-1中克隆或分离得到,或者通过DNA或肽合成的方法得到。
可将本发明基因与表达载体可操作地连接,得到能够表达本发明蛋白的重组表达载体,进而可以通过诸如农杆菌介导法、基因枪法、花粉管通道法等转基因方法,将所述表达载体导入宿主,得到转Cry4Cb1基因的转化体,例如农作物或者果树等植物,使其具备抗虫活性。
此外,还可以通过发酵本发明菌株HS18-1,得到含有Cry4Cb1蛋白的发酵液,将其制备成杀虫剂,用于农作物害虫的防治。本领域技术人员还可以将上述基因转化细菌或真菌,通过大规模发酵生产本发明Bt蛋白。
本领域技术人员还可以根据本发明公开的基因,将其转化棉花、玉米、水稻、蔬菜等农作物,使其具备相应的抗虫活性。从而降低农药的使用量,减少环境污染,具有重要的经济价值和应用前景。
附图说明
图1显示的是cry4Cb1全长基因克隆,其中M,marker;1,cry4Cb1基因。
图2显示的是重组质粒pET-4Cb的酶切鉴定图谱,其中1重组质粒pET-4Cb;2,用Nde I+EcoR I双酶切pET-30a;3,Nde I+EcoR I双酶切pET-4Cb;4,插入的DNA;M1、M2为Marker。
图3显示的是在E.coli BL21(DE3)中表达Cry4Cb1的SDS-PAGE检测,其中M为蛋白marker;1.阴性对照(E.coiiBL21(DE3)(pET-30a));2.裂解上清;3.Cry4Cb1包涵体。
具体实施方式
以下实施例进一步说明本发明的内容,但不应理解为对本发明的限制。在不背离本发明精神和实质的情况下,对本发明方法、步骤或条件所作的修改或替换,均属于本发明的范围。
若未特别指明,实施例中所用的技术手段为本领域技术人员所熟知的常规手段。
实施例1 Cry4Cb1基因的克隆
本发明从四川省成都平原土壤中分离得到的苏云金芽孢杆菌(Bacillus thuringiensis)新菌株,该菌株已于2008年10月21日在中国微生物菌种保藏管理委员会普通微生物中心(地址:北京市朝阳区大屯路甲3号,中国科学院微生物研究所,邮编100101)保藏,分类命名为苏云金芽孢杆菌(Bacillus thuringiensis),保藏号为CGMCC No.2718。
本例通过如下方法克隆得到Cry4Cb1基因的全长序列。
采用基因组DNA纯化试剂盒(购自赛百盛公司)提取菌株HS18-1的总DNA。设计引物序列如下:
P1:5’ATGTCTAATCGTTATCAACGGTACCC 3’
P2:5’TCACTCGTTCATACAAATCAACTCGA 3’
PCR反应体系:
10×buffer 2.5μl
MgCl2(25mM) 1.5μl
Taq酶 0.2μl
dNTPs(2.5mM) 2μl
引物 2μl
模板 5μl
最终反应体积 25μl
热循环反应:94℃预变性5min;94℃变性1min,52℃退火,72℃延伸2min,30个循环;72℃延伸5min;4℃停止反应。扩增反应产物在1%琼脂糖凝胶上电泳,置凝胶成像系统中观察PCR扩增结果。结果如图1所示,通过扩增得到了约为3.5kb的序列,将该序列进行测序,其核苷酸序列如SEQ ID No.1所示,与目的序列一致。
实施例2 Cry4Cb1基因的表达及杀虫活性测定
根据cry4Cb1基因开放阅读框两端序列,设计并合成一对特异性引物cry4F:5′-GCGCATATG(NdeI)ATGTCTAATCGTTATCA ACGGTACCC-3′;cry4R:5′-CGGAATTC(EcoR I)TCACTCGTTCATACAAATCAACTCGA-3′,分别在5’端引物Nde I和EcoR I酶切位点。以BtMC28质粒为模板进行扩增,扩增的产物采用Nde I和EcoR I进行双酶切,酶切产物与同样进行双酶切后的载体pET-30a(+)连接,转化E.coli DH5α感受态细胞,提取其质粒酶切电泳验证了插入片断大小符合预期目的后(图2),再转入受体菌E.coli.BL21(DE3)。将重组质粒命名为pET-4Cb,含重组质粒的重组子命名为E.coli.BL21(4Cb)。SDS-PAGE分析表明cry4Cb1基因的表达产物在菌体超声破碎后的沉淀中(图3),分子量约为130kDa左右,与预测的蛋白分子量相符。cry4Cb1基因表达产物分别对甜菜夜蛾,棉铃虫及伊蚊的生测结果表明:表达产物对这三种虫都具有较好的杀虫活性。对棉铃虫杀虫活性最高,LC50为10.06μg/mL;对伊蚊的LC50为18.41μg/mL;对甜菜夜蛾杀虫活性最低,LC50为30.29μg/mL。蛋白对鳞翅目杀虫活性的的测定方法参见(Song FP,Zhang J,Gu AX,et al.,2003.Identification of cry1I-typegenes from Bacillus thuringiensis strains and characterization of a novelcry1I-type gene.Appl.Environ.Microbiol 69:5207-5211),蛋白对双翅目杀虫活性的的测定方法参见(Ibarra JE,del Rincón MC,Sergio Ordúz,et al.,2003. Diversity of Bacillus thuringienisis Strains from LatinAmerica with Insecticidal Activity against Different Mosquito Species.Appl Environ Microbiol 69:5269-5274)。
序列表
<110>四川农业大学
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atg aat tca tat caa aat aaa aat gaa tat gaa ata ttg gat gct tca 48
Met Asn Ser Tyr Gln Asn Lys Asn Glu Tyr Glu Ile Leu Asp Ala Ser
1 5 10 15
caa aac aac tct aat atg tct aat cgt tat caa cgg tac cca cta gca 96
Gln Asn Asn Ser Asn Met Ser Asn Arg Tyr Gln Arg Tyr Pro Leu Ala
20 25 30
cat aat cca caa act tct ata caa act acg aat tat aag gat tgg ctg 144
His Asn Pro Gln Thr Ser Ile Gln Thr Thr Asn Tyr Lys Asp Trp Leu
35 40 45
aaa atg tgt caa aat cct cat caa aat ccc tta gac atg gaa ggg tat 192
Lys Met Cys Gln Asn Pro His Gln Asn Pro Leu Asp Met Glu Gly Tyr
50 55 60
gat agt aat tca gtc gtt gtg gta agt aca ggt ttg att gtt gtt ggt 240
Asp Ser Asn Ser Val Val Val Val Ser Thr Gly Leu Ile Val Val Gly
65 70 75 80
act tta att agt att ttg agt gcg gga ttg gga tct ata cct ata att 288
Thr Leu Ile Ser Ile Leu Ser Ala Gly Leu Gly Ser Ile Pro Ile Ile
85 90 95
tat ggt act tta ttg cct gtt cta tgg aac gat cca aac aat ccg caa 336
Tyr Gly Thr Leu Leu Pro Val Leu Trp Asn Asp Pro Asn Asn Pro Gln
100 105 110
aaa act tgg cat gaa ttt atg agt cat ggt gaa aca ctt ttg aac caa 384
Lys Thr Trp His Glu Phe Met Ser His Gly Glu Thr Leu Leu Asn Gln
115 120 125
aca ata tca aca gtt gag agg aat aga gca gca gcc tat ttg gag gga 432
Thr Ile Ser Thr Val Glu Arg Asn Arg Ala Ala Ala Tyr Leu Glu Gly
130 135 140
tac aca aca gca gta aaa aat gtg aag aag cac tta aat gtg tgg ctc 480
Tyr Thr Thr Ala Val Lys Asn Val Lys Lys His Leu Asn Val Trp Leu
145 150 155 160
aaa act cca aat caa gct aat gca cga aca gta gca gat tta tac aag 528
Lys Thr Pro Asn Gln Ala Asn Ala Arg Thr Val Ala Asp Leu Tyr Lys
165 170 175
gac act gat ttt tta ttt ttt aca act ttg ccc cac ctt aaa ctt cgt 576
Asp Thr Asp Phe Leu Phe Phe Thr Thr Leu Pro His Leu Lys Leu Arg
180 185 190
ggc tat gag aca tta ctt ctg agt tct tat aca caa gct gca aat atg 624
Gly Tyr Glu Thr Leu Leu Leu Ser Ser Tyr Thr Gln Ala Ala Asn Met
195 200 205
cat tta ata tta tta aag caa gct tca aaa tac gct gat caa tgg aat 672
His Leu Ile Leu Leu Lys Gln Ala Ser Lys Tyr Ala Asp Gln Trp Asn
210 215 220
gct caa ctt tct gtc tat gtt cag aaa aca gca aac gat tat tat act 720
Ala Gln Leu Ser Val Tyr Val Gln Lys Thr Ala Asn Asp Tyr Tyr Thr
225 230 235 240
gat tta gta aaa ctg ata gga gaa tat aca gat tat tgt att gca act 768
Asp Leu Val Lys Leu Ile Gly Glu Tyr Thr Asp Tyr Cys Ile Ala Thr
245 250 255
tac aga tta ggc tta act aca att aaa tct aga gct act tca tgg aac 816
Tyr Arg Leu Gly Leu Thr Thr Ile Lys Ser Arg Ala Thr Ser Trp Asn
260 265 270
ata tac aat atg tat cgt aga gag atg act att ttg gtg tta gat ctc 864
Ile Tyr Asn Met Tyr Arg Arg Glu Met Thr Ile Leu Val Leu Asp Leu
275 280 285
gta gct ctt ttc cct gca cat gat att aaa aaa tat cct agt ggg act 912
Val Ala Leu Phe Pro Ala His Asp Ile Lys Lys Tyr Pro Ser Gly Thr
290 295 300
aaa gta gag ctt act aga gaa att tat act gat gca ctt ggt gct gta 960
Lys Val Glu Leu Thr Arg Glu Ile Tyr Thr Asp Ala Leu Gly Ala Val
305 310 315 320
gcg ctt cca caa aac att gat gct ata gag caa ttg gcg acc cgt gcg 1008
Ala Leu Pro Gln Asn Ile Asp Ala Ile Glu Gln Leu Ala Thr Arg Ala
325 330 335
cct aat tta ttt agt tgg tta aag ggc ttt aaa ttt att acg act cag 1056
Pro Asn Leu Phe Ser Trp Leu Lys Gly Phe Lys Phe Ile Thr Thr Gln
340 345 350
tca aca aat agg tat tat tta tca ggt att gcg aat caa tat agc ttt 1104
Ser Thr Asn Arg Tyr Tyr Leu Ser Gly Ile Ala Asn Gln Tyr Ser Phe
355 360 365
acc aat tct aat gga gag ata tgg gga cct att tct ggg aat cct act 1152
Thr Asn Ser Asn Gly Glu Ile Trp Gly Pro Ile Ser Gly Asn Pro Thr
370 375 380
ggc gta tcg tct gat tta acc ata gat aat aat ttt tct att tac aaa 1200
Gly Val Ser Ser Asp Leu Thr Ile Asp Asn Asn Phe Ser Ile Tyr Lys
385 390 395 400
ctt tca ata tta cgt ggt tat caa ctc tca cca gat ttt tca ttt cat 1248
Leu Ser Ile Leu Arg Gly Tyr Gln Leu Ser Pro Asp Phe Ser Phe His
405 410 415
aat cca gtt cac caa att gat ttt tct aca acg aat aac caa cag gga 1296
Asn Pro Val His Gln Ile Asp Phe Ser Thr Thr Asn Asn Gln Gln Gly
420 425 430
cga gtt cag tca tat aaa tca ggc gga cct act cct gtt aat ccg gag 1344
Arg Val Gln Ser Tyr Lys Ser Gly Gly Pro Thr Pro Val Asn Pro Glu
435 440 445
acg aca gct att cat tta ccg ata gat tca aaa tgt aca caa aac tgt 1392
Thr Thr Ala Ile His Leu Pro Ile Asp Ser Lys Cys Thr Gln Asn Cys
450 455 460
aat cct aca ttt aat aat tac agt cat ata tta tct tac gca aaa act 1440
Asn Pro Thr Phe Asn Asn Tyr Ser His Ile Leu Ser Tyr Ala Lys Thr
465 470 475 480
ttc aca tca aat tta acc att ggt acc aca tca aat atc cac ttc gtt 1488
Phe Thr Ser Asn Leu Thr Ile Gly Thr Thr Ser Asn Ile His Phe Val
485 490 495
tgg ttg gac gca caa agt gtg gat cgt gaa aat aca att gat tta aat 1536
Trp Leu Asp Ala Gln Ser Val Asp Arg Glu Asn Thr Ile Asp Leu Asn
500 505 510
aat att aca cag att cca gct gta aag gcc agt caa gtt tat cca gaa 1584
Asn Ile Thr Gln Ile Pro Ala Val Lys Ala Ser Gln Val Tyr Pro Glu
515 520 525
aac tct gta att aaa ggt cct ggt cat aca ggt gga aat ctg gtt aga 1632
Asn Ser Val Ile Lys Gly Pro Gly His Thr Gly Gly Asn Leu Val Arg
530 535 540
att gat agt agt ggt tat atg tca att gtt tgt aaa ttc cca cta caa 1680
Ile Asp Ser Ser Gly Tyr Met Ser Ile Val Cys Lys Phe Pro Leu Gln
545 550 555 560
gta aag gga tat cgt gtt cgt att aga tat gca gca aat aat aga gct 1728
Val Lys Gly Tyr Arg Val Arg Ile Arg Tyr Ala Ala Asn Asn Arg Ala
565 570 575
gaa ctt tat ata tcg tca gct gga aat agt cca agt aaa aat gtt gat 1776
Glu Leu Tyr Ile Ser Ser Ala Gly Asn Ser Pro Ser Lys Asn Val Asp
580 585 590
cta gac cct aca ttt tca ggt act aac tat gaa agc tta aat tat aca 1824
Leu Asp Pro Thr Phe Ser Gly Thr Asn Tyr Glu Ser Leu Asn Tyr Thr
595 600 605
aat ttt aaa gat aaa gaa act gag ttt ata ata aca gaa gga cag ctc 1872
Asn Phe Lys Asp Lys Glu Thr Glu Phe Ile Ile Thr Glu Gly Gln Leu
610 615 620
gtt aaa caa tca ata ata ttc tca acc aat gga aat gtt ctc ctg gat 1920
Val Lys Gln Ser Ile Ile Phe Ser Thr Asn Gly Asn Val Leu Leu Asp
625 630 635 640
aag att gaa ttt att cca ctg gga acg aca acc tat gag tat gaa gag 1968
Lys Ile Glu Phe Ile Pro Leu Gly Thr Thr Thr Tyr Glu Tyr Glu Glu
645 650 655
aag cag aat cta gaa aaa gcg cga aaa gcg ttg aac gct ttg ttt acg 2016
Lys Gln Asn Leu Glu Lys Ala Arg Lys Ala Leu Asn Ala Leu Phe Thr
660 665 670
gat ggc acg aat ggc tat cta caa atg gat acc att gat tat gat atc 2064
Asp Gly Thr Asn Gly Tyr Leu Gln Met Asp Thr Ile Asp Tyr Asp Ile
675 680 685
aat caa act gca aac tta ata gaa tgt gta tca gat gaa ttg tat gca 2112
Asn Gln Thr Ala Asn Leu Ile Glu Cys Val Ser Asp Glu Leu Tyr Ala
690 695 700
aaa gaa aag ata gtt tta tta gat gaa gtc aaa tat gcg aag cgg ctt 2160
Lys Glu Lys Ile Val Leu Leu Asp Glu Val Lys Tyr Ala Lys Arg Leu
705 710 715 720
agc ata tca cgt aac cta ctt tcg aaa gat tat tta gaa ttt tca gat 2208
Ser Ile Ser Arg Asn Leu Leu Ser Lys Asp Tyr Leu Glu Phe Ser Asp
725 730 735
gta ttt gaa gaa aac gga tgg acg aca agt gat aat att tca atc cag 2256
Val Phe Glu Glu Asn Gly Trp Thr Thr Ser Asp Asn Ile Ser Ile Gln
740 745 750
gcg gat aat cct att ttt aag ggg aat tat tta aaa atg ttt ggg gca 2304
Ala Asp Asn Pro Ile Phe Lys Gly Asn Tyr Leu Lys Met Phe Gly Ala
755 760 765
aga gat att gat gga acc cta ttt cca act tat ctc tat caa aaa ata 2352
Arg Asp Ile Asp Gly Thr Leu Phe Pro Thr Tyr Leu Tyr Gln Lys Ile
770 775 780
gag gag tcc aag tta aaa ccc tat aca cgt tat cga gta aga ggg ttt 2400
Glu Glu Ser Lys Leu Lys Pro Tyr Thr Arg Tyr Arg Val Arg Gly Phe
785 790 795 800
gtg gga agt agt aaa gat cta aaa tta gtg gta aca cgc tat gag aaa 2448
Val Gly Ser Ser Lys Asp Leu Lys Leu Val Val Thr Arg Tyr Glu Lys
805 810 815
gaa att gat gcc att atg aat gtt cca aat gat ttg gca cat atg cag 2496
Glu Ile Asp Ala Ile Met Asn Val Pro Asn Asp Leu Ala His Met Gln
820 825 830
ctt aac cct tca tgt gga gat tat cgc tgt gaa tca tcg tcc cag ttt 2544
Leu Asn Pro Ser Cys Gly Asp Tyr Arg Cys Glu Ser Ser Ser Gln Phe
835 840 845
ttg gtg aac caa gtg cat cct aca tca aca gct gga tat gct ctt gat 2592
Leu Val Asn Gln Val His Pro Thr Ser Thr Ala Gly Tyr Ala Leu Asp
850 855 860
atg tat gca tgc ccg tta agt tca gat aaa aac cat gtt atg tgt cac 2640
Met Tyr Ala Cys Pro Leu Ser Ser Asp Lys Asn His Val Met Cys His
865 870 875 880
gat cgt cat cca ttt gat ttt cat att gac acc gga gaa gta ggt aca 2688
Asp Arg His Pro Phe Asp Phe His Ile Asp Thr Gly Glu Val Gly Thr
885 890 895
aat aca aac gta ggt att gat gtt tta ttt aaa att tct aat cca gat 2736
Asn Thr Asn Val Gly Ile Asp Val Leu Phe Lys Ile Ser Asn Pro Asp
900 905 910
gga tac gct aca gta ggg aat cta gaa gtc att gaa gaa gga cca cta 2784
Gly Tyr Ala Thr Val Gly Asn Leu Glu Val Ile Glu Glu Gly Pro Leu
915 920 925
aca ggc gac gca ttg gca cat gtg aaa cat aag gaa aag aaa tgg aag 2832
Thr Gly Asp Ala Leu Ala His Val Lys His Lys Glu Lys Lys Trp Lys
930 935 940
caa cac atg gag aaa aaa cgt tgg aaa aca caa caa gcc tac gat cct 2880
Gln His Met Glu Lys Lys Arg Trp Lys Thr Gln Gln Ala Tyr Asp Pro
945 950 955 960
gca aaa cag gct gta gat gca tta ttt aca aat gaa caa gag tta cac 2928
Ala Lys Gln Ala Val Asp Ala Leu Phe Thr Asn Glu Gln Glu Leu His
965 970 975
tat cat att act tta gat cat att caa aac gct gat cga ctg ata cag 2976
Tyr His Ile Thr Leu Asp His Ile Gln Asn Ala Asp Arg Leu Ile Gln
980 985 990
gcg att ccc tat gta tac cat gct tgg tta ccg agt gct cca ggt atg 3024
Ala Ile Pro Tyr Val Tyr His Ala Trp Leu Pro Ser Ala Pro Gly Met
995 1000 1005
aac tat gat gga tat caa ggg tta aac gca cgt atc atg caa gca 3069
Asn Tyr Asp Gly Tyr Gln Gly Leu Asn Ala Arg Ile Met Gln Ala
1010 1015 1020
cgc tat tta tat gat gca cgg aat atc ata aca aat ggt gac ttt 3114
Arg Tyr Leu Tyr Asp Ala Arg Asn Ile Ile Thr Asn Gly Asp Phe
1025 1030 1035
aca cag ggg tta acg gga tgg cac gca gca ggg aag gca acg gta 3159
Thr Gln Gly Leu Thr Gly Trp His Ala Ala Gly Lys Ala Thr Val
1040 1045 1050
caa cag atg aat ggc gct tct gta tta gtt cta tca aat tgg agt 3204
Gln Gln Met Asn Gly Ala Ser Val Leu Val Leu Ser Asn Trp Ser
1055 1060 1065
gcg ggg gta tct caa aac ttg cat gtc caa gac cat cat gga tat 3249
Ala Gly Val Ser Gln Asn Leu His Val Gln Asp His His Gly Tyr
1070 1075 1080
gtg cta cgt gtg att gcc aaa aaa gaa gga cct gga aaa ggg tat 3294
Val Leu Arg Val Ile Ala Lys Lys Glu Gly Pro Gly Lys Gly Tyr
1085 1090 1095
gta acg atg atg gat tgt aat gga aat cag gaa aca ctg aag ttc 3339
Val Thr Met Met Asp Cys Asn Gly Asn Gln Glu Thr Leu Lys Phe
1100 1105 1110
act tct tgt gaa gaa gga tat atg aca aaa aca gta gag gta ttc 3384
Thr Ser Cys Glu Glu Gly Tyr Met Thr Lys Thr Val Glu Val Phe
1115 1120 1125
cca gaa agt gat cgt gta cga ata gag atg gga gaa acc gaa ggt 3429
Pro Glu Ser Asp Arg Val Arg Ile Glu Met Gly Glu Thr Glu Gly
1130 1135 1140
acg ttt tat ata gat agc atc gag ttg att tgt atg aac gag tga 3474
Thr Phe Tyr Ile Asp Ser Ile Glu Leu Ile Cys Met Asn Glu
1145 1150 1155
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Met Asn Ser Tyr Gln Asn Lys Asn Glu Tyr Glu Ile Leu Asp Ala Ser
1 5 10 15
Gln Asn Asn Ser Asn Met Ser Asn Arg Tyr Gln Arg Tyr Pro Leu Ala
20 25 30
His Asn Pro Gln Thr Ser Ile Gln Thr Thr Asn Tyr Lys Asp Trp Leu
35 40 45
Lys Met Cys Gln Asn Pro His Gln Asn Pro Leu Asp Met Glu Gly Tyr
50 55 60
Asp Ser Asn Ser Val Val Val Val Ser Thr Gly Leu Ile Val Val Gly
65 70 75 80
Thr Leu Ile Ser Ile Leu Ser Ala Gly Leu Gly Ser Ile Pro Ile Ile
85 90 95
Tyr Gly Thr Leu Leu Pro Val Leu Trp Asn Asp Pro Asn Asn Pro Gln
100 105 110
Lys Thr Trp His Glu Phe Met Ser His Gly Glu Thr Leu Leu Asn Gln
115 120 125
Thr Ile Ser Thr Val Glu Arg Asn Arg Ala Ala Ala Tyr Leu Glu Gly
130 135 140
Tyr Thr Thr Ala Val Lys Asn Val Lys Lys His Leu Asn Val Trp Leu
145 150 155 160
Lys Thr Pro Asn Gln Ala Asn Ala Arg Thr Val Ala Asp Leu Tyr Lys
165 170 175
Asp Thr Asp Phe Leu Phe Phe Thr Thr Leu Pro His Leu Lys Leu Arg
180 185 190
Gly Tyr Glu Thr Leu Leu Leu Ser Ser Tyr Thr Gln Ala Ala Asn Met
195 200 205
His Leu Ile Leu Leu Lys Gln Ala Ser Lys Tyr Ala Asp Gln Trp Asn
210 215 220
Ala Gln Leu Ser Val Tyr Val Gln Lys Thr Ala Asn Asp Tyr Tyr Thr
225 230 235 240
Asp Leu Val Lys Leu Ile Gly Glu Tyr Thr Asp Tyr Cys Ile Ala Thr
245 250 255
Tyr Arg Leu Gly Leu Thr Thr Ile Lys Ser Arg Ala Thr Ser Trp Asn
260 265 270
Ile Tyr Asn Met Tyr Arg Arg Glu Met Thr Ile Leu Val Leu Asp Leu
275 280 285
Val Ala Leu Phe Pro Ala His Asp Ile Lys Lys Tyr Pro Ser Gly Thr
290 295 300
Lys Val Glu Leu Thr Arg Glu Ile Tyr Thr Asp Ala Leu Gly Ala Val
305 310 315 320
Ala Leu Pro Gln Asn Ile Asp Ala Ile Glu Gln Leu Ala Thr Arg Ala
325 330 335
Pro Asn Leu Phe Ser Trp Leu Lys Gly Phe Lys Phe Ile Thr Thr Gln
340 345 350
Ser Thr Asn Arg Tyr Tyr Leu Ser Gly Ile Ala Asn Gln Tyr Ser Phe
355 360 365
Thr Asn Ser Asn Gly Glu Ile Trp Gly Pro Ile Ser Gly Asn Pro Thr
370 375 380
Gly Val Ser Ser Asp Leu Thr Ile Asp Asn Asn Phe Ser Ile Tyr Lys
385 390 395 400
Leu Ser Ile Leu Arg Gly Tyr Gln Leu Ser Pro Asp Phe Ser Phe His
405 410 415
Asn Pro Val His Gln Ile Asp Phe Ser Thr Thr Asn Asn Gln Gln Gly
420 425 430
Arg Val Gln Ser Tyr Lys Ser Gly Gly Pro Thr Pro Val Asn Pro Glu
435 440 445
Thr Thr Ala Ile His Leu Pro Ile Asp Ser Lys Cys Thr Gln Asn Cys
450 455 460
Asn Pro Thr Phe Asn Asn Tyr Ser His Ile Leu Ser Tyr Ala Lys Thr
465 470 475 480
Phe Thr Ser Asn Leu Thr Ile Gly Thr Thr Ser Asn Ile His Phe Val
485 490 495
Trp Leu Asp Ala Gln Ser Val Asp Arg Glu Asn Thr Ile Asp Leu Asn
500 505 510
Asn Ile Thr Gln Ile Pro Ala Val Lys Ala Ser Gln Val Tyr Pro Glu
515 520 525
Asn Ser Val Ile Lys Gly Pro Gly His Thr Gly Gly Asn Leu Val Arg
530 535 540
Ile Asp Ser Ser Gly Tyr Met Ser Ile Val Cys Lys Phe Pro Leu Gln
545 550 555 560
Val Lys Gly Tyr Arg Val Arg Ile Arg Tyr Ala Ala Asn Asn Arg Ala
565 570 575
Glu Leu Tyr Ile Ser Ser Ala Gly Asn Ser Pro Ser Lys Asn Val Asp
580 585 590
Leu Asp Pro Thr Phe Ser Gly Thr Asn Tyr Glu Ser Leu Asn Tyr Thr
595 600 605
Asn Phe Lys Asp Lys Glu Thr Glu Phe Ile Ile Thr Glu Gly Gln Leu
610 615 620
Val Lys Gln Ser Ile Ile Phe Ser Thr Asn Gly Asn Val Leu Leu Asp
625 630 635 640
Lys Ile Glu Phe Ile Pro Leu Gly Thr Thr Thr Tyr Glu Tyr Glu Glu
645 650 655
Lys Gln Asn Leu Glu Lys Ala Arg Lys Ala Leu Asn Ala Leu Phe Thr
660 665 670
Asp Gly Thr Asn Gly Tyr Leu Gln Met Asp Thr Ile Asp Tyr Asp Ile
675 680 685
Asn Gln Thr Ala Asn Leu Ile Glu Cys Val Ser Asp Glu Leu Tyr Ala
690 695 700
Lys Glu Lys Ile Val Leu Leu Asp Glu Val Lys Tyr Ala Lys Arg Leu
705 710 715 720
Ser Ile Ser Arg Asn Leu Leu Ser Lys Asp Tyr Leu Glu Phe Ser Asp
725 730 735
Val Phe Glu Glu Asn Gly Trp Thr Thr Ser Asp Asn Ile Ser Ile Gln
740 745 750
Ala Asp Asn Pro Ile Phe Lys Gly Asn Tyr Leu Lys Met Phe Gly Ala
755 760 765
Arg Asp Ile Asp Gly Thr Leu Phe Pro Thr Tyr Leu Tyr Gln Lys Ile
770 775 780
Glu Glu Ser Lys Leu Lys Pro Tyr Thr Arg Tyr Arg Val Arg Gly Phe
785 790 795 800
Val Gly Ser Ser Lys Asp Leu Lys Leu Val Val Thr Arg Tyr Glu Lys
805 810 815
Glu Ile Asp Ala Ile Met Asn Val Pro Asn Asp Leu Ala His Met Gln
820 825 830
Leu Asn Pro Ser Cys Gly Asp Tyr Arg Cys Glu Ser Ser Ser Gln Phe
835 840 845
Leu Val Asn Gln Val His Pro Thr Ser Thr Ala Gly Tyr Ala Leu Asp
850 855 860
Met Tyr Ala Cys Pro Leu Ser Ser Asp Lys Asn His Val Met Cys His
86 5870 875 880
Asp Arg His Pro Phe Asp Phe His Ile Asp Thr Gly Glu Val Gly Thr
885 890 895
Asn Thr Asn Val Gly Ile Asp Val Leu Phe Lys Ile Ser Asn Pro Asp
900 905 910
Gly Tyr Ala Thr Val Gly Asn Leu Glu Val Ile Glu Glu Gly Pro Leu
915 920 925
Thr Gly Asp Ala Leu Ala His Val Lys His Lys Glu Lys Lys Trp Lys
930 935 940
Gln His Met Glu Lys Lys Arg Trp Lys Thr Gln Gln Ala Tyr Asp Pro
945 950 955 960
Ala Lys Gln Ala Val Asp Ala Leu Phe Thr Asn Glu Gln Glu Leu His
965 970 975
Tyr His Ile Thr Leu Asp His Ile Gln Asn Ala Asp Arg Leu Ile Gln
980 985 990
Ala Ile Pro Tyr Val Tyr His Ala Trp Leu Pro Ser Ala Pro Gly Met
995 1000 1005
Asn Tyr Asp Gly Tyr Gln Gly Leu Asn Ala Arg Ile Met Gln Ala
1010 1015 1020
Arg Tyr Leu Tyr Asp Ala Arg Asn Ile Ile Thr Asn Gly Asp Phe
1025 1030 1035
Thr Gln Gly Leu Thr Gly Trp His Ala Ala Gly Lys Ala Thr Val
1040 1045 1050
Gln Gln Met Asn Gly Ala Ser Val Leu Val Leu Ser Asn Trp Ser
1055 1060 1065
Ala Gly Val Ser Gln Asn Leu His Val Gln Asp His His Gly Tyr
1070 1075 1080
Val Leu Arg Val Ile Ala Lys Lys Glu Gly Pro Gly Lys Gly Tyr
1085 1090 1095
Val Thr Met Met Asp Cys Asn Gly Asn Gln Glu Thr Leu Lys Phe
1100 1105 1110
Thr Ser Cys Glu Glu Gly Tyr Met Thr Lys Thr Val Glu Val Phe
1115 1120 1125
Pro Glu Ser Asp Arg Val Arg Ile Glu Met Gly Glu Thr Glu Gly
1130 1135 1140
Thr Phe Tyr Ile Asp Ser Ile Glu Leu Ile Cys Met Asn Glu
1145 1150 1155
<210>3
<211>26
<212>DNA
<213>人工序列
<400>3
atgtctaatc gttatcaacg gtaccc 26
<210>4
<211>26
<212>DNA
<213>人工序列
<400>4
tcactcgttc atacaaatca actcga 26
<210>5
<211>35
<212>DNA
<213>人工序列
<400>5
gcgcatatga tgtctaatcg ttatcaacgg taccc 35
<210>6
<211>34
<212>DNA
<213>人工序列
<400>6
cggaattctc actcgttcat acaaatcaac tcga 34
Claims (9)
1、一种Bt蛋白Cry4Cb1,其氨基酸序列是:
1)SEQ ID No.2所示的氨基酸序列;或
2)SEQ ID No.2所示的氨基酸序列经取代、缺失和/或增加一个或多个氨基酸且具有同等活性的蛋白。
2、编码权利要求1所述蛋白的基因。
3、如权利要求2所述的基因,其核苷酸序列如SEQ ID No.1所示。
4、含有权利要求2或3所述基因的表达载体。
5、由权利要求4所述表达载体转化的宿主细胞。
6、如权利要求5所述的宿主细胞,其为植物宿主细胞。
7、含有权利要求所述蛋白的杀虫剂。
8、权利要求2或3所述基因或权利要求4所述表达载体在制备转基因植物中的应用。
9、权利要求2或3所述基因或权利要求4所述表达载体在提高植物抗虫性中的应用。
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CNA2009100815961A CN101591381A (zh) | 2009-04-13 | 2009-04-13 | Bt蛋白Cry4Cb1、其编码基因及应用 |
PCT/CN2010/000485 WO2010118633A1 (zh) | 2009-04-13 | 2010-04-13 | 杀虫晶体蛋白基因Cry4Cb1及其编码蛋白和应用 |
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CNA2009100815961A CN101591381A (zh) | 2009-04-13 | 2009-04-13 | Bt蛋白Cry4Cb1、其编码基因及应用 |
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Cited By (3)
Publication number | Priority date | Publication date | Assignee | Title |
---|---|---|---|---|
WO2010118633A1 (zh) * | 2009-04-13 | 2010-10-21 | 四川农业大学 | 杀虫晶体蛋白基因Cry4Cb1及其编码蛋白和应用 |
CN102584959A (zh) * | 2011-12-31 | 2012-07-18 | 四川农业大学 | 一种Bt蛋白Cry70Ba1、其编码基因及应用 |
CN102584960A (zh) * | 2011-12-31 | 2012-07-18 | 四川农业大学 | 一种Bt蛋白Cry70Aa1、其编码基因及应用 |
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CN101591381A (zh) * | 2009-04-13 | 2009-12-02 | 四川农业大学 | Bt蛋白Cry4Cb1、其编码基因及应用 |
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2009
- 2009-04-13 CN CNA2009100815961A patent/CN101591381A/zh active Pending
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2010
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Cited By (5)
Publication number | Priority date | Publication date | Assignee | Title |
---|---|---|---|---|
WO2010118633A1 (zh) * | 2009-04-13 | 2010-10-21 | 四川农业大学 | 杀虫晶体蛋白基因Cry4Cb1及其编码蛋白和应用 |
CN102584959A (zh) * | 2011-12-31 | 2012-07-18 | 四川农业大学 | 一种Bt蛋白Cry70Ba1、其编码基因及应用 |
CN102584960A (zh) * | 2011-12-31 | 2012-07-18 | 四川农业大学 | 一种Bt蛋白Cry70Aa1、其编码基因及应用 |
CN102584959B (zh) * | 2011-12-31 | 2013-11-27 | 四川农业大学 | 一种Bt蛋白Cry70Ba1、其编码基因及应用 |
CN102584960B (zh) * | 2011-12-31 | 2013-11-27 | 四川农业大学 | 一种Bt蛋白Cry70Aa1、其编码基因及应用 |
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