CN111647058B - 对松墨天牛等墨天牛属昆虫具有毒力的Bt毒素 - Google Patents
对松墨天牛等墨天牛属昆虫具有毒力的Bt毒素 Download PDFInfo
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Abstract
本发明涉及生物防治领域,具体涉及一种对松墨天牛等墨天牛属昆虫具有毒力的Bt毒素。所述Bt毒素为毒素BRC1912,其氨基酸序列如SEQ ID NO.2所示,该毒素对松墨天牛等墨天牛属昆虫具有良好的杀虫活性,本发明为松墨天牛等墨天牛属的害虫防控提供了有效途径。
Description
技术领域
本发明涉及林业害虫的生物防治领域,具体涉及一种对松墨天牛等墨天牛属昆虫具有毒力的Bt毒素。
背景技术
在林业上,鞘翅目天牛科墨天牛属昆虫是松材线虫的主要传播媒介(Mota etal., 2008; Futai, 2013),也是主要的蛀干害虫,又被称为次期性害虫,生活在韧皮部或者木质部之中。在韧皮部啃食输导组织,阻止树木对养分的吸收,在木质部啃食边材,对水分的运输造成阻碍,不严重的会造成树木发育不良,严重的导致树木整株坏死。蛀干害虫由于其生活习性,除了成虫会在树外活动,其它虫态几乎都生活在树木的韧皮部或者是木质部中,极其隐蔽,也较难防治。鉴于当前的防治方法效果不佳,其中生物防治具有绿色环保,持效期长的优点,进一步寻找有效的微生物制剂对松墨天牛等墨天牛属昆虫进行生物防治,对于遏制松材线虫病的扩散蔓延具有重要的意义。
苏云金芽胞杆菌(简称Bt)是一种昆虫病原微生物,对16个目3000多种害虫有活性,被广泛地应用于农、林和卫生害虫的生物防治中(Adang et al.,2014)。Bt的杀虫活性主要依赖于其在芽胞期产生的伴孢晶体蛋白(简称ICPs),主要是Cry毒素(Deist et al.,2014)。不同类别的Cry毒素的杀虫谱表现出显著的特异性,如Cry1、Cry9、Cry15类毒素对鳞翅目昆虫有效,Cry3、Cry7、Cry8和Cry34/35类毒素对鞘翅目的幼虫有活性(Bravo, 1997)。其中,有报道表明,Cry3A类毒素对天牛科幼虫有效,但其杀虫活性不高(Chen et al.,2005)。因此,为有效防治天牛科墨天牛属昆虫,迫切需要解决Cry3A毒素对其弱毒的问题。
在Cry3Aa毒素作用过程中,当其被昆虫摄取后,原毒素会被昆虫肠道蛋白酶酶解活化出55 kDa大小的活性片段,该片段对鞘翅目昆虫表现出杀虫活性(Walter et al.,2008; Li et al., 2011)。然后,单体的活性毒素寡聚化形成的低聚物与中肠上皮细胞刷状缘膜囊泡(BBNV)上的受体结合,从而引起膜穿孔并导致细胞死亡(Pardo-López et al.,2013)。而有研究表明,天牛肠道蛋白酶主要为胰蛋白酶,然而,73 kDa Cry3Aa毒素无法被天牛的肠道蛋白酶有效酶解活化出55 kDa的活性片段,即存在酶解活化障碍。已有研究证明Cry3Aa原毒素domain I的α-helix 3和α-helix 4之间的loop区域是被蛋白酶切割释放活性片段的区域,而在该区域缺乏胰蛋白酶的识别位点(Oppert et al., 1997)。此外,还发现55 kDa的Cry3Aa活性片段很容易被胰蛋白酶过度酶解为较小短肽,从而使得毒素对天牛失去活性。这表明在Cry3Aa毒素的其他结构域中存在较多的能够被胰蛋白酶识别的位点,从而使得毒素被过度酶解。这些结果表明昆虫肠道蛋白酶对Cry3A毒素的酶解活化障碍和过度酶解是造成其对目标昆虫弱毒的主要原因。因此,为解决昆虫肠道蛋白酶对毒素的酶解活化障碍和过度酶解现象,本发明以cry3Aa2基因为对象,通过分子改造的手段对毒素中的胰蛋白酶识别位点进行突变,开发能够产生对墨天牛属昆虫具有毒力的新型Cry毒素,是一项具有重大意义和应用价值的研究。
发明内容
本发明的目的在于提供了一种对松墨天牛等墨天牛属昆虫具有毒力的Bt毒素,以SEQ ID NO.4(Cry3Aa2)为支架毒素蛋白,通过分子改造的手段对毒素中的胰蛋白酶识别位点进行突变,开发能够产生对墨天牛属昆虫具有毒力的新型Cry毒素。
为实现上述目的,采用以下技术方案:
对墨天牛属昆虫具有毒力的Bt毒素,所述Bt毒素为毒素BRC1912蛋白,其氨基酸序列如SEQ ID NO.2所示。
一种编码如所述Bt毒素的基因,所述基因其核苷酸序列如SEQ ID NO.1所示。
与在SEQ ID NO: 4所列出的毒素Cry3Aa2蛋白(其基因序列如SEQ ID NO.3所示)的初始氨基酸序列相比,为促进昆虫肠道蛋白酶将毒素酶解活化出更多的55 kDa大小的活性片段,本发明的BRC1912蛋白的结构域I中的α-helix 3和α-helix 4之间的loop区域“缬氨酸-丝氨酸-丝氨酸”(第155和第157个氨基酸之间的序列)替换为短肽序列“苯丙氨酸-甲硫氨酸-精氨酸”,增加了胰蛋白酶的识别位点精氨酸(图1)。
与在SEQ ID NO: 4所列出的毒素Cry3Aa2蛋白的初始氨基酸序列相比,为克服昆虫肠道蛋白酶对毒素的过度酶解,本发明的BRC1912蛋白中包括了五个位点的氨基酸突变,将五个胰蛋白酶的识别位点突变为不被识别的丙氨酸,分别为:结构域I中在位置65处由赖氨酸置换成丙氨酸,在位置70处由赖氨酸置换成丙氨酸,在位置231处由赖氨酸置换成丙氨酸,结构域II中在位置468处由赖氨酸置换成丙氨酸,以及结构域III中在位置596处由赖氨酸置换成丙氨酸(图1)。
结构域I的基因序列和氨基酸序列分别如SEQ ID NO.5和SEQ ID NO.6所示;
结构域II基因序列和氨基酸序列分别如SEQ ID NO.7和SEQ ID NO.8所示;
结构域III基因序列和氨基酸序列分别如SEQ ID NO.9和SEQ ID NO.10所示。
所述的毒素BRC1912的三个结构域,这三个结构域中的任意一个结构域均可与Bt毒素的其他家族毒素的结构域任意组合获得对鞘翅目具有毒力的毒素。
所述的Bt毒素的其他家族毒素,这些毒素均具有三级结构域,包括但不限于Cry3、Cry7、Cry8类毒素。
所述对墨天牛属昆虫具有毒力的Bt毒素在防治天牛科墨天牛属昆虫中的应用。
所述墨天牛属昆虫包括松墨天牛(Monochamus alternatus enda、Monochamus alternatus alternatus)、云杉花墨天牛(Monochamus saltuarius)、云杉小墨天牛(Monochamus sutor)、卡来罗纳黑天牛(Monochamuscarolinensis)、密毛白点墨天牛(Monochamus scutellatus)和松墨斑墨天牛(Monochamus mutator)和樟子松墨天牛(Monochamus galloprovincialis)。
所述的对墨天牛属昆虫具有毒力的Bt毒素在防治鞘翅目害虫中的应用。
所述鞘翅目害虫包括赤拟谷盗(Tribolium castaneum)、红棕象甲(Rhynchophorus ferrugineus)、椰心叶甲(Brontispa longissima)、星天牛(Anoplophora chinensis)、光肩星天牛(Anoplophora glabripennis)和云斑天牛(Batocera horsfieldi)。
本发明的BRC1912晶体蛋白用于墨天牛属昆虫及其他鞘翅目昆虫的生物测定。在具体实施例中,优选地用于防治松墨天牛(Monochamus alternatusalternatus)、云杉花墨天牛(Monochamus saltuarius)、云杉小墨天牛(Monochamus sutor)、光肩星天牛(Anoplophora glabripennis)、云斑天牛(Batocera horsfieldi)和红棕象甲(Rhynchophorus ferrugineus)。
本发明的优点在于:
本发明的BRC1912毒素与SEQ ID NO: 4所列出的毒素Cry3Aa2蛋白在体外酶解活化效率、杀虫活性和目标昆虫种类谱相比,BRC1912毒素能够克服被昆虫肠道蛋白酶过度酶解的现象,即酶解活化出更多的55 KDa大小的活性片段,从而提高了对目标昆虫的杀虫活性,且表现出更大的目标昆虫种类谱。
附图说明:
图1为BRC1912与Cry3Aa2蛋白氨基酸序列的比对。
图2为通用载体pGEX-6P-1的质粒图谱。
图3为pHT304-G穿梭载体图谱。
图4为BRC1912晶体的显微镜形态特征图。
图5为BRC1912晶体蛋白的SDS-PAGE的结果图。其中泳道1为蛋白Marker,泳道2为BRC1912晶体蛋白。
具体实施方式
以下叙述本发明的实施例。应说明的是,本发明的实施例对于本发明只有说明作用,并不对本发明的范围构成任何限制。任何熟悉本领域的技术人员,在不脱离本发明技术方案的情况下,对本发明做出改进,变动或修饰均等同于等效实施案例,均属于本发明所附权力要求保护范围内。
实施例1 BRC1912基因的克隆及其工程菌的构建
对cry3Aa2基因上五位点进行氨基酸的突变获得BRC1912基因序列,分别为:在位置65处由赖氨酸(K)置换成丙氨酸(A),在位置70处由赖氨酸(K)置换成丙氨酸(A),在位置231处由赖氨酸(K)置换成丙氨酸(A),在位置468处由赖氨酸(K)置换成丙氨酸(A),以及在位置596处由赖氨酸(K)置换成丙氨酸(A)。且domain I中的α-helix 3和α-helix 4之间的loop区域短肽序列“缬氨酸-丝氨酸-丝氨酸”(“VSS”)(第155和第157个氨基酸之间的序列)替换为短肽序列“苯丙氨酸-甲硫氨酸-精氨酸”(“FMR”)(图1)。生物合成的BRC1912基因直接克隆在通用载体pGEX-6P-1的BamHI和XohI位点上(图2),将该质粒转入大肠杆菌中进行表达。
另一方面,以克隆在pGEX-KG-6P载体上的BRC1912基因为模板进行PCR扩增,PCR扩增引物为:
Cry3Aa2-C-F: 5’-CCCTCGAGATGAATCCGAACAATCGAAGT-3’
Cry3Aa2-C-R: 5’-CGGGATCCAAGCTTTCAGTCAGTCATGGA-3’
PCR扩增纯化产物通过XhoI和BamHI酶切位点连入pHT304-G载体中。并将该质粒电转入苏云金芽胞杆菌无晶体突变株HD73Cry-,获得能够表达对松墨天牛幼虫具有毒力的BRC1912晶体蛋白的工程菌Cry3Aa-C-BT,该菌株已于2020年1月13日保藏于中国微生物菌种保藏管理委员会普通微生物中心,分类号为苏云金芽孢杆菌 (Bacillus thuringiensis),保藏编号为CGMCC NO.19338,保藏地址为北京市朝阳区北辰西路1号院3号。
实施例2本发明所述的重组的pHT304-G穿梭载体的构建
以苏云金杆菌/大肠杆菌的pHT304穿梭载体为基础,设计合成cry3Aa2基因的启动子,包括启动cyt1A基因的BtI和BtII启动序列,STAB-SD序列以及多克隆位点序列(XhoI,SphI,XbaI,BamHI,KpnI和SacI)。该合成片段通过HindIII和EcoRI酶切位点连入pHT304质粒中(图3)。
实施例3 BRC1912毒素在苏云金芽胞杆菌中的表达及纯化
将获得的Cry3Aa-C-BT工程菌(CGMCC NO.19338)接入PM培养基(酵母粉2 g/L,胰蛋白胨10 g/L,KH2PO4 1 g/L,FeSO4•7H2O 0.02 g/L,MgSO4•7H2O 0.3 g/L,MnSO4•7H2O 0.02g/L,ZnSO4•7H2O 0.02 g/L,pH 7.0)中30℃培养至晶体释放期(60 h),经过受体菌体,晶体蛋白的溶解后,利用盐酸将pH调至4.7获得BRC1912晶体蛋白(图4、5)。
实施例4 BRC1912毒素在大肠杆菌中的表达及纯化
将毒素BRC1912转入大肠杆菌的工程菌在LB培养基中37℃培养至对数期并加入IPTG至终浓度为0.8 nM,恒温培养诱导表达蛋白,依照Glutathione Sepharose 4B GST产品说明书进行蛋白纯化。
实施例5 BRC1912毒素对松墨天牛(Monochamus alternatusalternatus)的生物测定
天牛人工饲料的配方:麸皮60 g,虾粉10 g,山梨酸2 g,苯钾酸钠4 g,酵母粉25g,琼脂30 g,松树韧皮部100 g,松树木质部50 g,蔗糖40 g,水300 mL。
生物测定方法:松墨天牛幼虫分别培养在含有0.1g人工饲料的500 µL离心管中。每一支离心管对应一只三龄初期幼虫,分别加入150 µL不同浓度梯度的Cry3Aa2原毒素和BRC1912毒素(1000、500、250、125、62.5和6.25 µg/mL)。各浓度梯度三组重复,每组含有10只幼虫作为重复(各浓度梯度测试30只)。以单蒸水作为空白对照。所有离心管放置在25℃,75%湿度的环境下培养,并对其进行光周期16:8处理(光照培养16小时,黑暗培养8小时)。观察各组幼虫的生长状况并在两周后计算各组幼虫死亡率,运用概率单位分析计算出致死中浓度(LC50)。BRC1912毒素对松墨天牛三龄初期幼虫两周的LC50值为12.3 μg/mL,与Cry3Aa2原始毒素相比,杀虫活性提高了9.5倍(详见表1)。
表1 Cry3Aa2和BRC1912毒素对松墨天牛幼虫的杀虫活性
实施例6 BRC1912毒素对云杉花墨天牛(Monochamus saltuarius)的生物测定
实验设计:将纯化的Cry3Aa2原毒素和BRC1912毒素分别稀释成1500、1000、500、250、125、62.5和6.25 µg/mL6个梯度,以单蒸水作为空白对照。其他方法同实施例5。BRC1912毒素对云杉花墨天牛二龄幼虫两周的LC50值为48.4 μg/mL,与Cry3Aa2原始毒素相比,杀虫活性提高了7.9倍(详见表2)。
表2 Cry3Aa2和BRC1912毒素对云杉花墨天牛幼虫的杀虫活性
实施例7 BRC1912毒素对云杉小墨天牛(Monochamus sutor)的生物测定
实验设计:将纯化的Cry3Aa2原毒素和BRC1912毒素稀释成1500、1000、500、250、125、62.5和6.25 µg/mL6个梯度,以单蒸水作为空白对照。其他方法同实施例5。BRC1912毒素对云杉小墨天牛二龄幼虫两周的LC50值为28.9 μg/mL,与Cry3Aa2原始毒素相比,杀虫活性提高了9.9倍(详见表3)。
表3 Cry3Aa2和BRC1912毒素对云杉小墨天牛幼虫的杀虫活性
实施例8 BRC1912毒素对光肩星天牛(Anoplophora glabripennis)的生物测定
实验设计:将纯化的Cry3Aa2原毒素和BRC1912毒素稀释成1500、1000、500、250、125和62.5 µg/mL6个梯度,以单蒸水作为空白对照。其他方法同实施例5。BRC1912毒素对云杉花墨天牛二龄幼虫两周的LC50值为142.2 μg/mL,与Cry3Aa2原始毒素相比,杀虫活性提高了5.5倍(详见表4)。
表4 Cry3Aa2和BRC1912毒素对光肩星天牛幼虫的杀虫活性
实施例9 BRC1912毒素对云斑天牛(Batocera horsfieldi)的生物测定
实验设计:将纯化的Cry3Aa2原毒素和BRC1912毒素稀释成1500、1000、500、250、125和62.5 µg/mL6个梯度,以单蒸水作为空白对照。其他方法同实施例5。BRC1912毒素对云杉花墨天牛二龄幼虫两周的LC50值为249.1 μg/mL,与Cry3Aa2原始毒素相比,杀虫活性提高了6.6倍(详见表5)。
表5 Cry3Aa2和BRC1912毒素对云斑天牛幼虫的杀虫活性
实施例10 BRC1912毒素对红棕象甲(Rhynchophorus ferrugineus)的生物测定
将纯化所得的Cry3Aa2原毒素和BRC1912毒素分别用单蒸水稀释到3.8、1.9、0.95和0.47 mg/mL。在24孔板中的人工饲料中分别加入50 μL不同浓度的毒素,每个孔放置一只二龄幼虫,各3个重复,每个重复10只幼虫。以单蒸水作为空白对照。放置人工气候箱中进行培养。培养条件:温度为28℃,相对湿度70%以上,光周期为8 h/16 h(白天/黑夜),观察各组幼虫的生长状况并在一周后计算各组幼虫死亡率,运用概率单位分析计算出致死中浓度(LC50),再通过Microsoft Excel 2016以及SPSS 18.0进行数据的总结与方差分析。
BRC1912毒素对红棕象甲二龄幼虫两周的LC50值为2.7 mg/mL,与Cry3Aa2原始毒素相比,对红棕象甲表现出了杀虫活性(详见表6)。
表6 Cry3Aa2和BRC1912毒素对红棕象甲幼虫的杀虫活性
SEQUENCE LISTING
序 列 表
<110> 福建农林大学
<120> 对松墨天牛等墨天牛属昆虫具有高毒力的Bt毒素
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<213> 苏云金杆菌(Bacillus thuringiensis)人工合成
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cagaaaatag ctgattatgc aaaaaataaa gctcttgcag agttacaggg ccttcaaaat 420
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aatccacata gccaggggcg gataagagag ctgttttctc aagcagaaag tcattttcgt 540
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caagctgcca acacacattt atttttacta aaagacgctc aaatttatgg agaagaatgg 660
ggatacgaaa aagaagatat tgctgaattt tatgcaagac aactaaaact tacgcaagaa 720
tatactgacc attgtgtcaa atggtataat gttggattag ataaattaag aggttcatct 780
tatgaatctt gggtaaactt taaccgttat cgcagagaga tgacattaac agtattagat 840
ttaattgcac tatttccatt gtatgatgtt cggctatacc caaaagaagt taaaaccgaa 900
ttaacaagag acgttttaac agatccaatt gtcggagtca acaaccttag gggctatgga 960
acaaccttct ctaatataga aaattatatt cgaaaaccac atctatttga ctatctgcat 1020
agaattcaat ttcacacgcg gttccaacca ggatattatg gaaatgactc tttcaattat 1080
tggtccggta attatgtttc aactagacca agcataggat caaatgatat aatcacatct 1140
ccattctatg gaaataaatc cagtgaacct gtacaaaatt tagaatttaa tggagaaaaa 1200
gtctatagag ccgtagcaaa tacaaatctt gcggtctggc cgtccgctgt atattcaggt 1260
gttacaaaag tggaatttag ccaatataat gatcaaacag atgaagcaag tacacaaacg 1320
tacgactcaa aaagaaatgt tggcgcggtc agctgggatt ctatcgatca attgcctcca 1380
gaaacaacag atgaacctct agaagcggga tatagccatc aactcaatta tgtaatgtgc 1440
tttttaatgc agggtagtag aggaacaatc ccagtgttaa cttggacaca taaaagtgta 1500
gactttttta acatgattga ttcgaaaaaa attacacaac ttccgttagt aaaggcatat 1560
aagttacaat ctggtgcttc cgttgtcgca ggtcctaggt ttacaggagg agatatcatt 1620
caatgcacag aaaatggaag tgcggcaact atttacgtta caccggatgt gtcgtactct 1680
caaaaatatc gagctagaat tcattatgct tctacatctc agataacatt tacactcagt 1740
ttagacgggg caccatttaa tcaatactat ttcgataaaa cgataaatgc aggagacaca 1800
ttaacgtata attcatttaa tttagcaagt ttcagcacac cattcgaatt atcagggaat 1860
aacttacaaa taggcgtcac aggattaagt gctggagata aagtttatat agacaaaatt 1920
gaatttattc cagtgaatta atgaggctcc atgactgact gaaagctt 1968
<210> 2
<211> 646
<212> PRT
<213> 苏云金杆菌(Bacillus thuringiensis)人工合成
<400> 2
Gly Ser Met Asn Pro Asn Asn Arg Ser Glu His Asp Thr Ile Lys Thr
1 5 10 15
Thr Glu Asn Asn Glu Val Pro Thr Asn His Val Gln Tyr Pro Leu Ala
20 25 30
Glu Thr Pro Asn Pro Thr Leu Glu Asp Leu Asn Tyr Lys Glu Phe Leu
35 40 45
Arg Met Thr Ala Asp Asn Asn Thr Glu Ala Leu Asp Ser Ser Thr Thr
50 55 60
Ala Asp Val Ile Gln Ala Gly Ile Ser Val Val Gly Asp Leu Leu Gly
65 70 75 80
Val Val Gly Phe Pro Phe Gly Gly Ala Leu Val Ser Phe Tyr Thr Asn
85 90 95
Phe Leu Asn Thr Ile Trp Pro Ser Glu Asp Pro Trp Lys Ala Phe Met
100 105 110
Glu Gln Val Glu Ala Leu Met Asp Gln Lys Ile Ala Asp Tyr Ala Lys
115 120 125
Asn Lys Ala Leu Ala Glu Leu Gln Gly Leu Gln Asn Asn Val Glu Asp
130 135 140
Tyr Val Ser Ala Leu Ser Ser Trp Gln Lys Asn Pro Phe Met Arg Arg
145 150 155 160
Asn Pro His Ser Gln Gly Arg Ile Arg Glu Leu Phe Ser Gln Ala Glu
165 170 175
Ser His Phe Arg Asn Ser Met Pro Ser Phe Ala Ile Ser Gly Tyr Glu
180 185 190
Val Leu Phe Leu Thr Thr Tyr Ala Gln Ala Ala Asn Thr His Leu Phe
195 200 205
Leu Leu Lys Asp Ala Gln Ile Tyr Gly Glu Glu Trp Gly Tyr Glu Lys
210 215 220
Glu Asp Ile Ala Glu Phe Tyr Ala Arg Gln Leu Lys Leu Thr Gln Glu
225 230 235 240
Tyr Thr Asp His Cys Val Lys Trp Tyr Asn Val Gly Leu Asp Lys Leu
245 250 255
Arg Gly Ser Ser Tyr Glu Ser Trp Val Asn Phe Asn Arg Tyr Arg Arg
260 265 270
Glu Met Thr Leu Thr Val Leu Asp Leu Ile Ala Leu Phe Pro Leu Tyr
275 280 285
Asp Val Arg Leu Tyr Pro Lys Glu Val Lys Thr Glu Leu Thr Arg Asp
290 295 300
Val Leu Thr Asp Pro Ile Val Gly Val Asn Asn Leu Arg Gly Tyr Gly
305 310 315 320
Thr Thr Phe Ser Asn Ile Glu Asn Tyr Ile Arg Lys Pro His Leu Phe
325 330 335
Asp Tyr Leu His Arg Ile Gln Phe His Thr Arg Phe Gln Pro Gly Tyr
340 345 350
Tyr Gly Asn Asp Ser Phe Asn Tyr Trp Ser Gly Asn Tyr Val Ser Thr
355 360 365
Arg Pro Ser Ile Gly Ser Asn Asp Ile Ile Thr Ser Pro Phe Tyr Gly
370 375 380
Asn Lys Ser Ser Glu Pro Val Gln Asn Leu Glu Phe Asn Gly Glu Lys
385 390 395 400
Val Tyr Arg Ala Val Ala Asn Thr Asn Leu Ala Val Trp Pro Ser Ala
405 410 415
Val Tyr Ser Gly Val Thr Lys Val Glu Phe Ser Gln Tyr Asn Asp Gln
420 425 430
Thr Asp Glu Ala Ser Thr Gln Thr Tyr Asp Ser Lys Arg Asn Val Gly
435 440 445
Ala Val Ser Trp Asp Ser Ile Asp Gln Leu Pro Pro Glu Thr Thr Asp
450 455 460
Glu Pro Leu Glu Ala Gly Tyr Ser His Gln Leu Asn Tyr Val Met Cys
465 470 475 480
Phe Leu Met Gln Gly Ser Arg Gly Thr Ile Pro Val Leu Thr Trp Thr
485 490 495
His Lys Ser Val Asp Phe Phe Asn Met Ile Asp Ser Lys Lys Ile Thr
500 505 510
Gln Leu Pro Leu Val Lys Ala Tyr Lys Leu Gln Ser Gly Ala Ser Val
515 520 525
Val Ala Gly Pro Arg Phe Thr Gly Gly Asp Ile Ile Gln Cys Thr Glu
530 535 540
Asn Gly Ser Ala Ala Thr Ile Tyr Val Thr Pro Asp Val Ser Tyr Ser
545 550 555 560
Gln Lys Tyr Arg Ala Arg Ile His Tyr Ala Ser Thr Ser Gln Ile Thr
565 570 575
Phe Thr Leu Ser Leu Asp Gly Ala Pro Phe Asn Gln Tyr Tyr Phe Asp
580 585 590
Lys Thr Ile Asn Ala Gly Asp Thr Leu Thr Tyr Asn Ser Phe Asn Leu
595 600 605
Ala Ser Phe Ser Thr Pro Phe Glu Leu Ser Gly Asn Asn Leu Gln Ile
610 615 620
Gly Val Thr Gly Leu Ser Ala Gly Asp Lys Val Tyr Ile Asp Lys Ile
625 630 635 640
Glu Phe Ile Pro Val Asn
645
<210> 3
<211> 1952
<212> DNA
<213> 苏云金杆菌(Bacillus thuringiensis)
<400> 3
tccatggatg aatccgaaca atcgaagtga acatgataca ataaaaacta ctgaaaataa 60
tgaggtgcca actaaccatg ttcaatatcc tttagcggaa actccaaatc caacactaga 120
agatttaaat tataaagagt ttttaagaat gactgcagat aataatacgg aagcactaga 180
tagctctaca acaaaagatg tcattcaaaa aggcatttcc gtagtaggtg atctcctagg 240
cgtagtaggt ttcccgtttg gtggagcgct tgtttcgttt tatacaaact ttttaaatac 300
tatttggcca agtgaagacc cgtggaaggc ttttatggaa caagtagaag cattgatgga 360
tcagaaaata gctgattatg caaaaaataa agctcttgca gagttacagg gccttcaaaa 420
taatgtcgaa gattatgtga gtgcattgag ttcatggcaa aaaaatcctg tgagttcacg 480
aaatccacat agccaggggc ggataagaga gctgttttct caagcagaaa gtcattttcg 540
taattcaatg ccttcgtttg caatttctgg atacgaggtt ctatttctaa caacatatgc 600
acaagctgcc aacacacatt tatttttact aaaagacgct caaatttatg gagaagaatg 660
gggatacgaa aaagaagata ttgctgaatt ttataaaaga caactaaaac ttacgcaaga 720
atatactgac cattgtgtca aatggtataa tgttggatta gataaattaa gaggttcatc 780
ttatgaatct tgggtaaact ttaaccgtta tcgcagagag atgacattaa cagtattaga 840
tttaattgca ctatttccat tgtatgatgt tcggctatac ccaaaagaag ttaaaaccga 900
attaacaaga gacgttttaa cagatccaat tgtcggagtc aacaacctta ggggctatgg 960
aacaaccttc tctaatatag aaaattatat tcgaaaacca catctatttg actatctgca 1020
tagaattcaa tttcacacgc ggttccaacc aggatattat ggaaatgact ctttcaatta 1080
ttggtccggt aattatgttt caactagacc aagcatagga tcaaatgata taatcacatc 1140
tccattctat ggaaataaat ccagtgaacc tgtacaaaat ttagaattta atggagaaaa 1200
agtctataga gccgtagcaa atacaaatct tgcggtctgg ccgtccgctg tatattcagg 1260
tgttacaaaa gtggaattta gccaatataa tgatcaaaca gatgaagcaa gtacacaaac 1320
gtacgactca aaaagaaatg ttggcgcggt cagctgggat tctatcgatc aattgcctcc 1380
agaaacaaca gatgaacctc tagaaaaggg atatagccat caactcaatt atgtaatgtg 1440
ctttttaatg cagggtagta gaggaacaat cccagtgtta acttggacac ataaaagtgt 1500
agactttttt aacatgattg attcgaaaaa aattacacaa cttccgttag taaaggcata 1560
taagttacaa tctggtgctt ccgttgtcgc aggtcctagg tttacaggag gagatatcat 1620
tcaatgcaca gaaaatggaa gtgcggcaac tatttacgtt acaccggatg tgtcgtactc 1680
tcaaaaatat cgagctagaa ttcattatgc ttctacatct cagataacat ttacactcag 1740
tttagacggg gcaccattta atcaatacta tttcgataaa acgataaata aaggagacac 1800
attaacgtat aattcattta atttagcaag tttcagcaca ccattcgaat tatcagggaa 1860
taacttacaa ataggcgtca caggattaag tgctggagat aaagtttata tagacaaaat 1920
tgaatttatt ccagtgaatt aatgaggatc ca 1952
<210> 4
<211> 644
<212> PRT
<213> 苏云金杆菌(Bacillus thuringiensis)
<400> 4
Met Asn Pro Asn Asn Arg Ser Glu His Asp Thr Ile Lys Thr Thr Glu
1 5 10 15
Asn Asn Glu Val Pro Thr Asn His Val Gln Tyr Pro Leu Ala Glu Thr
20 25 30
Pro Asn Pro Thr Leu Glu Asp Leu Asn Tyr Lys Glu Phe Leu Arg Met
35 40 45
Thr Ala Asp Asn Asn Thr Glu Ala Leu Asp Ser Ser Thr Thr Lys Asp
50 55 60
Val Ile Gln Lys Gly Ile Ser Val Val Gly Asp Leu Leu Gly Val Val
65 70 75 80
Gly Phe Pro Phe Gly Gly Ala Leu Val Ser Phe Tyr Thr Asn Phe Leu
85 90 95
Asn Thr Ile Trp Pro Ser Glu Asp Pro Trp Lys Ala Phe Met Glu Gln
100 105 110
Val Glu Ala Leu Met Asp Gln Lys Ile Ala Asp Tyr Ala Lys Asn Lys
115 120 125
Ala Leu Ala Glu Leu Gln Gly Leu Gln Asn Asn Val Glu Asp Tyr Val
130 135 140
Ser Ala Leu Ser Ser Trp Gln Lys Asn Pro Val Ser Ser Arg Asn Pro
145 150 155 160
His Ser Gln Gly Arg Ile Arg Glu Leu Phe Ser Gln Ala Glu Ser His
165 170 175
Phe Arg Asn Ser Met Pro Ser Phe Ala Ile Ser Gly Tyr Glu Val Leu
180 185 190
Phe Leu Thr Thr Tyr Ala Gln Ala Ala Asn Thr His Leu Phe Leu Leu
195 200 205
Lys Asp Ala Gln Ile Tyr Gly Glu Glu Trp Gly Tyr Glu Lys Glu Asp
210 215 220
Ile Ala Glu Phe Tyr Lys Arg Gln Leu Lys Leu Thr Gln Glu Tyr Thr
225 230 235 240
Asp His Cys Val Lys Trp Tyr Asn Val Gly Leu Asp Lys Leu Arg Gly
245 250 255
Ser Ser Tyr Glu Ser Trp Val Asn Phe Asn Arg Tyr Arg Arg Glu Met
260 265 270
Thr Leu Thr Val Leu Asp Leu Ile Ala Leu Phe Pro Leu Tyr Asp Val
275 280 285
Arg Leu Tyr Pro Lys Glu Val Lys Thr Glu Leu Thr Arg Asp Val Leu
290 295 300
Thr Asp Pro Ile Val Gly Val Asn Asn Leu Arg Gly Tyr Gly Thr Thr
305 310 315 320
Phe Ser Asn Ile Glu Asn Tyr Ile Arg Lys Pro His Leu Phe Asp Tyr
325 330 335
Leu His Arg Ile Gln Phe His Thr Arg Phe Gln Pro Gly Tyr Tyr Gly
340 345 350
Asn Asp Ser Phe Asn Tyr Trp Ser Gly Asn Tyr Val Ser Thr Arg Pro
355 360 365
Ser Ile Gly Ser Asn Asp Ile Ile Thr Ser Pro Phe Tyr Gly Asn Lys
370 375 380
Ser Ser Glu Pro Val Gln Asn Leu Glu Phe Asn Gly Glu Lys Val Tyr
385 390 395 400
Arg Ala Val Ala Asn Thr Asn Leu Ala Val Trp Pro Ser Ala Val Tyr
405 410 415
Ser Gly Val Thr Lys Val Glu Phe Ser Gln Tyr Asn Asp Gln Thr Asp
420 425 430
Glu Ala Ser Thr Gln Thr Tyr Asp Ser Lys Arg Asn Val Gly Ala Val
435 440 445
Ser Trp Asp Ser Ile Asp Gln Leu Pro Pro Glu Thr Thr Asp Glu Pro
450 455 460
Leu Glu Lys Gly Tyr Ser His Gln Leu Asn Tyr Val Met Cys Phe Leu
465 470 475 480
Met Gln Gly Ser Arg Gly Thr Ile Pro Val Leu Thr Trp Thr His Lys
485 490 495
Ser Val Asp Phe Phe Asn Met Ile Asp Ser Lys Lys Ile Thr Gln Leu
500 505 510
Pro Leu Val Lys Ala Tyr Lys Leu Gln Ser Gly Ala Ser Val Val Ala
515 520 525
Gly Pro Arg Phe Thr Gly Gly Asp Ile Ile Gln Cys Thr Glu Asn Gly
530 535 540
Ser Ala Ala Thr Ile Tyr Val Thr Pro Asp Val Ser Tyr Ser Gln Lys
545 550 555 560
Tyr Arg Ala Arg Ile His Tyr Ala Ser Thr Ser Gln Ile Thr Phe Thr
565 570 575
Leu Ser Leu Asp Gly Ala Pro Phe Asn Gln Tyr Tyr Phe Asp Lys Thr
580 585 590
Ile Asn Lys Gly Asp Thr Leu Thr Tyr Asn Ser Phe Asn Leu Ala Ser
595 600 605
Phe Ser Thr Pro Phe Glu Leu Ser Gly Asn Asn Leu Gln Ile Gly Val
610 615 620
Thr Gly Leu Ser Ala Gly Asp Lys Val Tyr Ile Asp Lys Ile Glu Phe
625 630 635 640
Ile Pro Val Asn
<210> 5
<211> 681
<212> DNA
<213> 人工序列
<400> 5
acaacagcag atgtcattca agcaggcatt tccgtagtag gtgatctcct aggcgtagta 60
ggtttcccgt ttggtggagc gcttgtttcg ttttatacaa actttttaaa tactatttgg 120
ccaagtgaag acccgtggaa ggcttttatg gaacaagtag aagcattgat ggatcagaaa 180
atagctgatt atgcaaaaaa taaagctctt gcagagttac agggccttca aaataatgtc 240
gaagattatg tgagtgcatt gagttcatgg caaaaaaatc ctttcatgcg tcgaaatcca 300
catagccagg ggcggataag agagctgttt tctcaagcag aaagtcattt tcgtaattca 360
atgccttcgt ttgcaatttc tggatacgag gttctatttc taacaacata tgcacaagct 420
gccaacacac atttattttt actaaaagac gctcaaattt atggagaaga atggggatac 480
gaaaaagaag atattgctga attttatgca agacaactaa aacttacgca agaatatact 540
gaccattgtg tcaaatggta taatgttgga ttagataaat taagaggttc atcttatgaa 600
tcttgggtaa actttaaccg ttatcgcaga gagatgacat taacagtatt agatttaatt 660
gcactatttc cattgtatga t 681
<210> 6
<211> 227
<212> PRT
<213> 人工序列
<400> 6
Thr Thr Ala Asp Val Ile Gln Ala Gly Ile Ser Val Val Gly Asp Leu
1 5 10 15
Leu Gly Val Val Gly Phe Pro Phe Gly Gly Ala Leu Val Ser Phe Tyr
20 25 30
Thr Asn Phe Leu Asn Thr Ile Trp Pro Ser Glu Asp Pro Trp Lys Ala
35 40 45
Phe Met Glu Gln Val Glu Ala Leu Met Asp Gln Lys Ile Ala Asp Tyr
50 55 60
Ala Lys Asn Lys Ala Leu Ala Glu Leu Gln Gly Leu Gln Asn Asn Val
65 70 75 80
Glu Asp Tyr Val Ser Ala Leu Ser Ser Trp Gln Lys Asn Pro Phe Met
85 90 95
Arg Arg Asn Pro His Ser Gln Gly Arg Ile Arg Glu Leu Phe Ser Gln
100 105 110
Ala Glu Ser His Phe Arg Asn Ser Met Pro Ser Phe Ala Ile Ser Gly
115 120 125
Tyr Glu Val Leu Phe Leu Thr Thr Tyr Ala Gln Ala Ala Asn Thr His
130 135 140
Leu Phe Leu Leu Lys Asp Ala Gln Ile Tyr Gly Glu Glu Trp Gly Tyr
145 150 155 160
Glu Lys Glu Asp Ile Ala Glu Phe Tyr Ala Arg Gln Leu Lys Leu Thr
165 170 175
Gln Glu Tyr Thr Asp His Cys Val Lys Trp Tyr Asn Val Gly Leu Asp
180 185 190
Lys Leu Arg Gly Ser Ser Tyr Glu Ser Trp Val Asn Phe Asn Arg Tyr
195 200 205
Arg Arg Glu Met Thr Leu Thr Val Leu Asp Leu Ile Ala Leu Phe Pro
210 215 220
Leu Tyr Asp
225
<210> 7
<211> 615
<212> DNA
<213> 人工序列
<400> 7
gttaaaaccg aattaacaag agacgtttta acagatccaa ttgtcggagt caacaacctt 60
aggggctatg gaacaacctt ctctaatata gaaaattata ttcgaaaacc acatctattt 120
gactatctgc atagaattca atttcacacg cggttccaac caggatatta tggaaatgac 180
tctttcaatt attggtccgg taattatgtt tcaactagac caagcatagg atcaaatgat 240
ataatcacat ctccattcta tggaaataaa tccagtgaac ctgtacaaaa tttagaattt 300
aatggagaaa aagtctatag agccgtagca aatacaaatc ttgcggtctg gccgtccgct 360
gtatattcag gtgttacaaa agtggaattt agccaatata atgatcaaac agatgaagca 420
agtacacaaa cgtacgactc aaaaagaaat gttggcgcgg tcagctggga ttctatcgat 480
caattgcctc cagaaacaac agatgaacct ctagaagcgg gatatagcca tcaactcaat 540
tatgtaatgt gctttttaat gcagggtagt agaggaacaa tcccagtgtt aacttggaca 600
cataaaagtg tagac 615
<210> 8
<211> 205
<212> PRT
<213> 人工序列
<400> 8
Val Lys Thr Glu Leu Thr Arg Asp Val Leu Thr Asp Pro Ile Val Gly
1 5 10 15
Val Asn Asn Leu Arg Gly Tyr Gly Thr Thr Phe Ser Asn Ile Glu Asn
20 25 30
Tyr Ile Arg Lys Pro His Leu Phe Asp Tyr Leu His Arg Ile Gln Phe
35 40 45
His Thr Arg Phe Gln Pro Gly Tyr Tyr Gly Asn Asp Ser Phe Asn Tyr
50 55 60
Trp Ser Gly Asn Tyr Val Ser Thr Arg Pro Ser Ile Gly Ser Asn Asp
65 70 75 80
Ile Ile Thr Ser Pro Phe Tyr Gly Asn Lys Ser Ser Glu Pro Val Gln
85 90 95
Asn Leu Glu Phe Asn Gly Glu Lys Val Tyr Arg Ala Val Ala Asn Thr
100 105 110
Asn Leu Ala Val Trp Pro Ser Ala Val Tyr Ser Gly Val Thr Lys Val
115 120 125
Glu Phe Ser Gln Tyr Asn Asp Gln Thr Asp Glu Ala Ser Thr Gln Thr
130 135 140
Tyr Asp Ser Lys Arg Asn Val Gly Ala Val Ser Trp Asp Ser Ile Asp
145 150 155 160
Gln Leu Pro Pro Glu Thr Thr Asp Glu Pro Leu Glu Ala Gly Tyr Ser
165 170 175
His Gln Leu Asn Tyr Val Met Cys Phe Leu Met Gln Gly Ser Arg Gly
180 185 190
Thr Ile Pro Val Leu Thr Trp Thr His Lys Ser Val Asp
195 200 205
<210> 9
<211> 429
<212> DNA
<213> 人工序列
<400> 9
aacatgattg attcgaaaaa aattacacaa cttccgttag taaaggcata taagttacaa 60
tctggtgctt ccgttgtcgc aggtcctagg tttacaggag gagatatcat tcaatgcaca 120
gaaaatggaa gtgcggcaac tatttacgtt acaccggatg tgtcgtactc tcaaaaatat 180
cgagctagaa ttcattatgc ttctacatct cagataacat ttacactcag tttagacggg 240
gcaccattta atcaatacta tttcgataaa acgataaatg caggagacac attaacgtat 300
aattcattta atttagcaag tttcagcaca ccattcgaat tatcagggaa taacttacaa 360
ataggcgtca caggattaag tgctggagat aaagtttata tagacaaaat tgaatttatt 420
ccagtgaat 429
<210> 10
<211> 143
<212> PRT
<213> 人工序列
<400> 10
Asn Met Ile Asp Ser Lys Lys Ile Thr Gln Leu Pro Leu Val Lys Ala
1 5 10 15
Tyr Lys Leu Gln Ser Gly Ala Ser Val Val Ala Gly Pro Arg Phe Thr
20 25 30
Gly Gly Asp Ile Ile Gln Cys Thr Glu Asn Gly Ser Ala Ala Thr Ile
35 40 45
Tyr Val Thr Pro Asp Val Ser Tyr Ser Gln Lys Tyr Arg Ala Arg Ile
50 55 60
His Tyr Ala Ser Thr Ser Gln Ile Thr Phe Thr Leu Ser Leu Asp Gly
65 70 75 80
Ala Pro Phe Asn Gln Tyr Tyr Phe Asp Lys Thr Ile Asn Ala Gly Asp
85 90 95
Thr Leu Thr Tyr Asn Ser Phe Asn Leu Ala Ser Phe Ser Thr Pro Phe
100 105 110
Glu Leu Ser Gly Asn Asn Leu Gln Ile Gly Val Thr Gly Leu Ser Ala
115 120 125
Gly Asp Lys Val Tyr Ile Asp Lys Ile Glu Phe Ile Pro Val Asn
130 135 140
Claims (6)
1.对墨天牛属昆虫具有毒力的Bt毒素,其特征在于,所述Bt毒素为毒素BRC1912,其氨基酸序列如SEQ ID NO.2所示,是一种由支架毒素蛋白工程化的毒素,所述支架毒素蛋白其氨基酸序列如SEQ ID NO.4所示。
2.一种编码如权利要求1所述Bt毒素的基因,其特征在于,所述基因其核苷酸序列如SEQ ID NO.1所示。
3.如权利要求1所述的对墨天牛属昆虫具有毒力的Bt毒素在防治天牛科墨天牛属昆虫中的应用。
4.根据权利要求3所述的应用,其特征在于:所述墨天牛属昆虫包括松墨天牛(Monochamus alternatus enda、Monochamus alternatus alternatus)、云杉花墨天牛(Monochamus saltuarius)、云杉小墨天牛(Monochamus sutor)、卡来罗纳黑天牛(Monochamuscarolinensis)、密毛白点墨天牛(Monochamus scutellatus)和松墨斑墨天牛(Monochamus mutator)和樟子松墨天牛(Monochamus galloprovincialis)。
5.如权利要求1所述的对墨天牛属昆虫具有毒力的Bt毒素在防治鞘翅目害虫中的应用。
6.根据权利要求5所述的应用,其特征在于,所述鞘翅目害虫包括赤拟谷盗(Tribolium castaneum)、红棕象甲(Rhynchophorus ferrugineus)、椰心叶甲(Brontispa longissima)、星天牛(Anoplophora chinensis)、光肩星天牛(Anoplophora glabripennis)和云斑天牛(Batocera horsfieldi)。
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