CN109912701B - 一种提高番茄抗虫性的方法 - Google Patents
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Abstract
本发明公开了一种提高番茄抗虫性的方法。本发明提供了蛋白或其相关生物材料在调控植物抗虫性中的应用;所述相关生物材料为能够表达所述蛋白的核酸分子,或含有所述核酸分子的表达盒、重组载体、重组菌或转基因细胞系;所述蛋白为SlMTB1蛋白、SlMTB2蛋白和SlMTB3蛋白。实验证明,SlMTB1,SlMTB2,SlMTB3基因沉默的转基因番茄在受到棉铃虫侵害时,会比野生型番茄表达更多的PI‑II,使其对棉铃虫的抗性显著提高。因此,SlMTB1,SlMTB2,SlMTB3蛋白及其部分编码基因可用于农牧业和生态环境治理所需的抗虫植物品种的培育与鉴定,在植物抗虫育种领域中具有重要作用。
Description
技术领域
本发明属于生物技术领域,涉及一种提高番茄抗虫性的方法。
背景技术
番茄是一种非常重要的经济作物,既是水果,又是蔬菜,其营养价值十分丰富,深受广大消费者的喜爱。番茄是我国栽培最为普遍的果菜之一。目前,中国是世界上番茄栽培面积最大、生产总量最多的国家。随着农业技术的不断提高,番茄种植面积日益扩大,病虫害也越来越严重。病虫害严重影响农业生产,影响作物的产量和品质,制约农业经济的稳定发展。长期以来人们普遍采用化学杀虫剂来控制害虫,但随着时间的推移尤其是化学农药的大量、不合理的使用,带来了如农药残留、害虫产生抗药性和再猖獗、杀灭天敌等自然生态平衡被破坏、环境污染、人类食品安全受到威胁的严重问题。越来越多的研究结果表明,植物在长期的进化过程中形成了一套极为复杂而又精细的防御机制来感知和对抗各类病虫害。减少农药施用、利用植物自身的抗性控制病虫害,对生产无公害蔬菜、水果尤其重要,是我国农业最迫切的需求之一。
利用植物抗虫基因培育的抗虫转基因作物,可以避免害虫危害,还具有成本低、保护全、特异性强等优点,成为当前农业生物工程研究的一个热点。抗虫转基因作物21年的商业化,累计种植面积已超过8亿公顷,它为种植者和生态环境的改善带来了丰厚收益,革命性地改变了农业生产的格局。
番茄是研究植物与病虫相互作用关系的经典模式系统。早在二十世纪九十年代初,科学家从番茄中发现了世界上第一个植物多肽激素--系统素。系统素以及被称为“抗虫激素”的茉莉酸在植物对咀嚼式昆虫及腐生型病原菌的抗性反应中起重要调控作用,并且通过遗传操作茉莉酸途径中的重要元件可以显著改变植物对病虫的抗性。也就是说,提高植物中茉莉酸信号途径可以提高植物对咀嚼式昆虫的抗性,提高茉莉酸信号转导途径可以通过增强茉莉酸途径的正向调控元件和抑制茉莉酸途径的负向调控元件来实现。
发明内容
本发明的一个目的是提供SlMTB1蛋白、SlMTB2蛋白和SlMTB3蛋白以及相关生物材料的新用途。
本发明所提供的新用途具体为蛋白或其相关生物材料在调控植物抗虫性中的应用;所述相关生物材料为能够表达所述蛋白的核酸分子,或含有所述核酸分子的表达盒、重组载体、重组菌或转基因细胞系;所述蛋白为SlMTB1蛋白、SlMTB2蛋白和SlMTB3蛋白。
所述SlMTB1蛋白具体为如下(A1)-(A4)中任一所示蛋白质:
(A1)氨基酸序列为SEQ ID No.1的蛋白质;
(A2)将SEQ ID No.1所示的氨基酸序列经过一个或几个氨基酸残基的取代和/或缺失和/或添加且具有相同功能的蛋白质;
(A3)与(A1)-(A2)中任一所限定的氨基酸序列具有99%以上、95%以上、90%以上、85%以上或者80%以上同源性且具有相同功能的蛋白质;
(A4)在(A1)-(A3)中任一所限定的蛋白质的N端和/或C端连接标签后得到的融合蛋白;
所述SlMTB2蛋白具体为如下(B1)-(B4)中任一所示蛋白质:
(B1)氨基酸序列为SEQ ID No.2的蛋白质;
(B2)将SEQ ID No.2所示的氨基酸序列经过一个或几个氨基酸残基的取代和/或缺失和/或添加且具有相同功能的蛋白质;
(B3)与(B1)-(B2)中任一所限定的氨基酸序列具有99%以上、95%以上、90%以上、85%以上或者80%以上同源性且具有相同功能的蛋白质;
(B4)在(B1)-(B3)中任一所限定的蛋白质的N端和/或C端连接标签后得到的融合蛋白;
所述SlMTB3蛋白具体为如下(C1)-(C4)中任一所示蛋白质:
(C1)氨基酸序列为SEQ ID No.3的蛋白质;
(C2)将SEQ ID No.3所示的氨基酸序列经过一个或几个氨基酸残基的取代和/或缺失和/或添加且具有相同功能的蛋白质;
(C3)与(C1)-(C2)中任一所限定的氨基酸序列具有99%以上、95%以上、90%以上、85%以上或者80%以上同源性且具有相同功能的蛋白质;
(C4)在(C1)-(C3)中任一所限定的蛋白质的N端和/或C端连接标签后得到的融合蛋白。
在所述应用中,所述蛋白在所述植物中的表达量和/或活性越低,所述植物的抗虫性越强。
本发明还保护能够使受体植物中蛋白的表达量和/或活性降低的物质在提高植物抗虫性中的应用;所述蛋白为SlMTB1蛋白、SlMTB2蛋白和SlMTB3蛋白;所述SlMTB1蛋白为前文(A1)-(A4)中任一所示蛋白质;所述SlMTB2蛋白为前文(B1)-(B4)中任一所示蛋白质;所述SlMTB3蛋白为前文(C1)-(C4)中任一所示蛋白质。
进一步地,所述“能够使受体植物中蛋白的表达量和/或活性降低的物质”可为如下任一:
a)用于沉默所述SlMTB1蛋白的编码基因表达的RNA片段;
b)用于沉默所述SlMTB2蛋白的编码基因表达的RNA片段;
c)用于沉默所述SlMTB3蛋白的编码基因表达的RNA片段;
d)用于同时沉默所述SlMTB1蛋白、所述SlMTB2蛋白和所述SlMTB3蛋白的编码基因表达的RNA片段;
e)编码a)所述RNA片段的DNA片段;
f)编码b)所述RNA片段的DNA片段;
g)编码c)所述RNA片段的DNA片段;
h)编码d)所述RNA片段的DNA片段;
i)含有e)-h)中任一所述DNA片段的重组载体、表达盒、转基因细胞系或重组菌。
更进一步地,a)所述RNA片段为SEQ ID No.7或SEQ ID No.4的第1401-1600位核苷酸编码的RNA;b)所述RNA片段为SEQ ID No.8或SEQ ID No.5的第601-800位核苷酸编码的RNA;c)所述RNA片段为SEQ ID No.9或SEQ ID No.6的第81-280位核苷酸编码的RNA;d)所述RNA片段为SEQ ID No.10编码的RNA;e)所述DNA片段的核苷酸序列为SEQ ID No.7;f)所述DNA片段的核苷酸序列为SEQ ID No.8;g)所述DNA片段的核苷酸序列为SEQ ID No.9;h)所述DNA片段的核苷酸序列为SEQ ID No.10。
本发明的另一个目的是提供一种培育抗虫性提高的植物的方法(或称为“提高番茄抗虫性的方法”)。
本发明所提供的培育抗虫性提高的植物的方法(或称为“提高番茄抗虫性的方法”),具体可包括如下步骤:使受体植物中蛋白的表达量和/或活性降低,获得抗虫植物;所述蛋白为SlMTB1蛋白、SlMTB2蛋白和SlMTB3蛋白;所述SlMTB1蛋白为前文(A1)-(A4)中任一所示蛋白质;所述SlMTB2蛋白为前文(B1)-(B4)中任一所示蛋白质;所述SlMTB3蛋白为前文(C1)-(C4)中任一所示蛋白质。
在所述方法中,所述“使受体植物中蛋白的表达量和/或活性降低”可通过对所述受体植物中所述蛋白的编码基因进行敲除或抑制表达来实现。
进一步地,可通过任何能够实现“对所述受体植物中所述蛋白的编码基因进行敲除或抑制表达”这一目的的技术手段实现,如通过RNAi手段对所述编码基因进行抑制表达。
更进一步地,对所述受体植物中的所述蛋白的编码基因进行抑制表达可通过向所述受体植物中导入前文任一所述的“能够使受体植物中蛋白的表达量和/或活性降低的物质”来实现。
更加具体的,在本发明的一个实施例中,对所述受体植物中的所述蛋白的编码基因进行抑制表达是通过向所述受体植物中导入RNAi干扰载体来实现的;所述RNAi干扰载体为在pHELLSGATE2载体(Invitrogen)的两个attP1和attP2位点分别发生BP反应,分别插入SEQ ID No.10以及SEQ ID No.10的反向互补序列后所得的重组载体。
在上述方法中,将所述RNAi干扰载体导入所述受体植物,具体可为:通过使用Ti质粒、Ri质粒、植物病毒载体、直接DNA转化、显微注射、电导、农杆菌介导等常规生物学方法转化植物细胞或组织,并将转化的植物组织培育成植株。
在所述方法中,所述抗虫性提高具体可体现为如下1)和/或2):
1)所述抗虫植物上的虫的体积和/或重量小于所述受体植物;
2)所述抗虫植物中PI-II基因的表达量高于所述受体植物。
进一步地,所述PI-II基因的序列为序列表中SEQ ID No.11。
在本发明的一个实施例中,所述PI-II基因的表达量为mRNA水平上的表达。
在上述各应用和方法中,所述虫为昆虫。进一步,所述昆虫为咀嚼式昆虫。更进一步,所述咀嚼式昆虫为棉铃虫。
在上述各应用和方法中,所述植物为双子叶植物或单子叶植物。进一步,所述双子叶植物为茄科植物。更进一步,所述茄科植物为番茄。
在上述各应用和方法中,所述“能够表达所述蛋白的核酸分子”为所述SlMTB1蛋白的编码基因、所述SlMTB2蛋白的编码基因和所述SlMTB3蛋白的编码基因。
进一步地,所述SlMTB1蛋白的编码基因可为如下(D1)-(D3)中任一:
(D1)SEQ ID No.4所示的DNA分子;
(D2)在严格条件下与(D1)限定的DNA分子杂交且编码所述SlMTB1蛋白的DNA分子;
(D3)与(D1)-(D2)中任一限定的DNA序列具有99%以上、95%以上、90%以上、85%以上或者80%以上同源性且编码所述SlMTB1蛋白的DNA分子;
进一步地,所述SlMTB2蛋白的编码基因可为如下(E1)-(E3)中任一:
(E1)SEQ ID No.5所示的DNA分子;
(E2)在严格条件下与(E1)限定的DNA分子杂交且编码所述SlMTB2蛋白的DNA分子;
(E3)与(E1)-(E2)中任一限定的DNA序列具有99%以上、95%以上、90%以上、85%以上或者80%以上同源性且编码所述SlMTB2蛋白的DNA分子;
进一步地,所述SlMTB3蛋白的编码基因可为如下(F1)-(F3)中任一:
(F1)SEQ ID No.6所示的DNA分子;
(F2)在严格条件下与(F1)限定的DNA分子杂交且编码所述SlMTB3蛋白的DNA分子;
(F3)与(F1)-(F2)中任一限定的DNA序列具有99%以上、95%以上、90%以上、85%以上或者80%以上同源性且编码所述SlMTB3蛋白的DNA分子。
上述严格条件可为用6×SSC,0.5%SDS的溶液,在65℃下杂交,然后用2×SSC,0.1%SDS和1×SSC,0.1%SDS各洗膜一次。
前文所述的“能够使受体植物中蛋白的表达量和/或活性降低的物质”也属于本发明的保护范围。
本发明是通过抑制番茄中茉莉酸途径的负向调控元件来实现番茄对咀嚼式昆虫的抗性。实验证明,SlMTB1,SlMTB2,SlMTB3基因沉默的转基因番茄在受到棉铃虫侵害时,会比野生型番茄表达更多的PI-II,使其对棉铃虫的抗性显著提高。因此,SlMTB1,SlMTB2,SlMTB3基因与番茄对棉铃虫抗性相关;SlMTB1,SlMTB2,SlMTB3蛋白及其部分编码基因可用于农牧业和生态环境治理所需的抗虫植物品种的培育与鉴定,在植物抗虫育种领域中具有重要作用。
附图说明
图1为SlMTB1,SlMTB2,SlMTB3基因沉默的转基因番茄在饲喂棉铃虫7天后的地上部分抗虫表型结果。
图2为SlMTB1,SlMTB2,SlMTB3基因沉默的转基因番茄在饲喂棉铃虫7天后棉铃虫的体积与重量统计结果。A为棉铃虫的体积;B为棉铃虫的重量统计结果。
图3为SlMTB1,SlMTB2,SlMTB3基因沉默的转基因番茄在饲喂棉铃虫2天后地上部分的PI-II mRNA相对表达量检测结果。
具体实施方式
下述实施例中所使用的实验方法如无特殊说明,均为常规方法。
下述实施例中所用的材料、试剂等,如无特殊说明,均可从商业途径得到。
下述实施例中SlMTB1蛋白氨基酸序列为SEQ ID No.1,其编码基因SlMTB1的核苷酸序列为SEQ ID No.4;SlMTB2蛋白氨基酸序列为SEQ ID No.2,其编码基因SlMTB2的核苷酸序列为SEQ ID No.5;SlMTB3蛋白氨基酸序列为SEQ ID No.3,其编码基因SlMTB3的核苷酸序列为SEQ ID No.6。
下述实施例中番茄采用番茄常规品种M82(以下也称为野生型番茄),为美国番茄遗传资源中心(TGRC,http://tgrc.ucdavis.edu/)产品。
下述实施例中pHELLSGATE2载体为Invitrogen公司产品。
实施例1、抑制SlMTB1,SlMTB2,SlMTB3基因表达的重组载体的获得
提取番茄(Solanum lycopersicon)品种M82的总RNA,反转录得到的cDNA为模板,先用引物1和引物2进行PCR扩增SlMTB1,用引物3和引物4进行PCR扩增SlMTB2,用引物5和引物6进行PCR扩增SlMTB3;再以前三步扩增产物为模板用引物1和引物6进行PCR扩增SlMTB1/2/3。
引物1:
5′-GGGGACAAGTTTGTACAAAAAAGCAGGCTCCTCTTAGGAGATGCCAT-3′(上游引物,下划线部分碱基为attB1位点);
引物2:5′-CCCTGATTTCAGCGGAGCTCACCCTTACAA-3′;
引物3:5′-CCGCTGAAATCAGGGGTTGTGGAGCTT-3′;
引物4:5′-GCAGAAACTAAACTATCCACCTTCGGAGAA-3′;
引物5:5′-TAGTTTAGTTTCTGCTGAATGTTCTTT-3′;
引物6:
5′-GGGGACCACTTTGTACAAGAAAGCTGGGTCTTCCCTAGCTTCCCT-3′(下游引物,下划线部分碱基为attB2位点)。
得到658bpPCR产物,利用Omega DNA纯化试剂盒纯化后待用。
经过测序,该PCR产物的核苷酸序列含有序列表中序列4自5’末端第1401-1600位核苷酸(即SEQ ID No.7)followed by序列表中序列5自5’末端第601-800位核苷酸(即SEQID No.8)followed by序列表中序列6自5’末端第81-280位核苷酸(即SEQ ID No.9),为SlMTB1,SlMTB2,SlMTB3基因部分片段(即SEQ ID No.10)。
将上述PCR产物与pHELLSGATE2载体,进行一步BP反应,将BP反应产物转化大肠杆菌DH5α,挑选阳性克隆。提取阳性克隆的质粒进行测序,该质粒为在pHELLSGATE2载体的两个attP1和attP2位点分别发生BP反应,分别插入SEQ ID No.10,以及SEQ ID No.10的反向互补序列,表明构建的重组载体正确,记作pHELLSGATE2-SlMTB1/2/3–RNAi,为沉默SlMTB1,SlMTB2,SlMTB3基因表达的重组载体。
实施例2、SlMTB1,SlMTB2,SlMTB3基因沉默转基因番茄的获得及功能鉴定
一、SlMTB1,SlMTB2,SlMTB3基因沉默转基因番茄的获得
1、SlMTB1,SlMTB2,SlMTB3基因沉默重组菌的获得
将由实施例1得到的沉默SlMTB1,SlMTB2,SlMTB3基因表达的重组载体pHELLSGATE2-SlMTB1/2/3-RNAi转化农杆菌AGLO(记载Li C,Liu G,Xu C,Lee GI,Bauer P,Ling HQ,Ganal MW,Howe GA.The tomato suppressor of prosystemin-mediatedresponses2 gene encodes a fatty acid desaturase required for the biosynthesisof jasmonic acid and the production of a systemic wound signal for defensegene expression.Plant Cell.2003.15(7):1646-1661.,公众可从中国科学院遗传与发育生物学研究所获得),挑取单克隆,于含有壮观霉素、利福平、链霉素抗生素的LB液体培养基中28℃过夜振荡培养,得到转化子。
将转化子进行菌液PCR鉴定,采用引物1和引物6(见上文),得到658bp的为阳性重组菌,将阳性重组菌命名为AGL0/pHELLSGATE2-SlMTB1/2/3-RNAi并保存于-70℃备用。
2、SlMTB1,SlMTB2,SlMTB3基因沉默转基因番茄的获得
1)、番茄转化相关培养基的配制
液体MS培养基:将4.4g MS盐、30g蔗糖和水混合,用水定容至1L,用1mol/L KOH调pH至5.8~6.0之间,高压灭菌。
种子生长培养基(1/2MS培养基):将2.2g MS盐、30g蔗糖和水混合,用水定容至1L,用1mol/L KOH调pH至5.8~6.0之间,加入0.8%的琼脂,高压灭菌。
预(共)培养培养基(D1):将4.4g MS、1.0mg玉米素(Zeatin)和30g蔗糖溶于水中,用水定容至1L,用1mol/L KOH调pH至5.8~6.0之间,加0.8%的琼脂,高压灭菌。
筛选分化培养基(2Z):将4.4g MS盐、2.0mg玉米素、50mg卡那霉素、100mg肌醇、0.5mg叶酸和20g蔗糖溶于水中,用水定容至1L,用1mol/L KOH调pH至5.8~6.0之间,加0.8%的琼脂,高压灭菌。
生根培养基:将4.4g MS盐、50mg卡那霉素、0.5mg叶酸、0.5mg吲哚丁酸和30g蔗糖溶于水中,用水定容至1L,用1mol/L KOH调pH至5.8~6.0之间,加0.8%的琼脂,高压灭菌。
2)、SlMTB1,SlMTB2,SlMTB3基因沉默转基因番茄的制备
(1)转化外植体的准备
取野生型番茄(品种M82),挑选饱满、粒大种子,用75%乙醇浸泡2min,接着用30%NaClO浸泡10min,用无菌水冲洗5遍,播种于种子生长培养基上,于25℃、16h光照/8h黑暗条件下光照培养。萌发8天后,在无菌条件下用锋利的剪刀将子叶切成小方块(动作应快),将子叶小方块接种于预培养培养基中,于25℃、16h光照/8h黑暗条件下培养,2d后,可用于番茄转化。
(2)侵染液的准备
将保存备用的AGL0/pHELLSGATE2-SlMTB1/2/3-RNAi接种于含相应抗生素的LB液体培养基中,28℃、200rpm过夜培养。次日以1:100的比例转接至新的LB液体培养基中,28℃、200rpm培养至OD600=0.7。菌液以5000rpm离心10min,弃上清液,收集菌体。用液体MS培养基重新悬浮菌体,稀释至OD600=0.4,备用。
(3)外植体的转化、筛选与生根
将步骤(1)得到的子叶块分别浸入(2)制备好的侵染液10min,然后接种于D1培养基上(培养基上放滤纸)共培养2天,转入到筛选分化培养基(2Z)中筛选培养,每2周继代一次,培养8周后产生抗性芽。当不定芽伸长至3cm时,用解剖刀将抗性芽切下,并转接至生根培养基上,进行生根培养,将生根的T0代转基因植株移入土中常规管理,收获T1代pHELLSGATE2-SlMTB1/2/3-RNAi的转基因番茄种子。
上述共培养、筛选培养、生根培养的条件均为:温度为25℃、16h光照/8h黑暗。
3)、SlMTB1,SlMTB2,SlMTB3基因沉默转基因番茄的鉴定
将生长18天的两叶一心期的番茄叶片用止血钳作机械损伤,在每个小叶垂直主叶脉的方向平行夹两下,1小时后剪取植株地上部分,迅速用液氮速冻,-80℃保存。提取T1代pHELLSGATE2-SlMTB1/2/3-RNAi转基因番茄叶片的RNA,反转录得到cDNA,以野生型番茄M82为对照,进行RT-PCR。
用于RT-PCR检测SlMTB1表达水平的引物(扩增得到位于SlMTB1编码区3’末端的178bp片段)为:
上游引物SlMTB1-QF:5′-AATCATCCAAATATTCAAAGAG-3′;
下游引物SlMTB1-QR:5′-CGGTGAAAGTTGCCTTAA-3′。
用于RT-PCR检测SlMTB2表达水平的引物(扩增得到位于SlMTB2编码区3’末端的82bp片段)为:
上游引物SlMTB2-QF:5′-AGTTGGAATCTGTTTCGGAC-3′;
下游引物SlMTB2-QR:5′-GGACTAGAAGGCTTATCGAA-3′。
用于RT-PCR检测SlMTB3表达水平的引物(扩增得到位于SlMTB3编码区3’末端的135bp片段)为:
上游引物SlMTB3-QF:5′-GATCAGGGACGCGGAATA-3′;
下游引物SlMTB3-QR:5′-TTTGCTACAGGATGGGTT-3′。
以actin2为内参,用于扩增内参基因的上游引物为:
上游引物SlACT2-QF:5′-TTGCTGACCGTATGAGCAAG-3′;
下游引物SlACT2-QR:5′-GGACAATGGATGGACCAGAC-3′。
结果T1代pHELLSGATE2-SlMTB1/2/3-RNAi的转基因番茄中SlMTB1,SlMTB2,SlMTB3基因的相对表达量为0.01985,0.00346,0.00616;
野生型番茄中SlMTB1,SlMTB2,SlMTB3基因的相对表达量为0.05756,0.02145,0.03843;
可以看出,与野生型番茄比,T1代pHELLSGATE2-SlMTB1/2/3-RNAi的转基因番茄中的SlMTB1,SlMTB2,SlMTB3基因的表达受到抑制,为阳性转基因植物。
4)、纯合转基因植物的获得
将上述鉴定为阳性的T1代pHELLSGATE2-SlMTB1/2/3-RNAi的转基因番茄植株收种得到T2代种子,消毒后播在800μL/L卡那霉素的1/2MS培养基上筛选(野生型M82在含卡那霉素的培养基上生长,其侧根会受到严重抑制,没有侧根生长;转基因材料由于具有卡那霉素抗性侧根生长基本不受抑制),若某一个T1代pHELLSGATE2-SlMTB1/2/3-RNAi转基因植株收获的种子都表现为对卡那霉素抗性,则该T1代转基因植株为纯合体,其收获的种子也都是纯合体,可以用于后续的昆虫饲喂实验。
经鉴定,T2代pHELLSGATE2-SlMTB1/2/3-RNAi转基因番茄6#T1代收获的种子都表现为对卡那霉素抗性,即T2代pHELLSGATE2-SlMTB1/2/3-RNAi转基因番茄6#这个株系是纯合体。
采用同样的方法将空载体pHELLSGATE2转入野生型番茄中,得到T0代转空载体番茄,收种、播种,直到得到T2代转空载体番茄。采用同样的方法检测其SlMTB1,SlMTB2,SlMTB3基因相对表达量,结果表明,T2代转空载体番茄叶片中SlMTB1,SlMTB2,SlMTB3表达量与野生型番茄相比,无显著差异。
二、pHELLSGATE2-SlMTB1/2/3-RNAi的转基因番茄对昆虫抗性影响
棉铃虫(Helicoverpaarmigera)记载在文献“Wu K.,Y.Lu,H.Feng,Y.Jiang,andJ.Zhao.2008.Suppression of cotton bollworm in multiple crops in China inareas with Bttoxin-containing cotton.Science.321(5896):1676-1678.”中公开过,公众可从中国科学院遗传与发育生物学研究所获得。
在昆虫饲喂前,将野生型番茄、T2代转空载体番茄和T2代pHELLSGATE2-SlMTB1/2/3-RNAi的转基因番茄在常规条件下培养:光照16h,黑暗8h,光照强度70microeinsteins m- 2s-2,温度25℃。待植物长至4周时,取长势一致的2龄期棉铃虫小心放到植物上,任其自由取食。上述每个株系取20株进行试验,每棵植株放3头棉铃虫。待到饲喂第2天,采集地上部分叶片提取RNA用于后续分析。
另一批野生型和转基因植株饲喂到第7天,对植物进行拍照记录,对饲喂相应植株的棉铃虫拍照记录并做重量统计。
实验结果表明,与野生型番茄M82相比,纯合的T2代pHELLSGATE2-SlMTB1/2/3-RNAi转基因番茄在受到棉铃虫咀食后会表达更多的PI-II,SlMTB1,SlMTB2,SlMTB3基因沉默的转基因番茄pHELLSGATE2-SlMTB1/2/3-RNAi对棉铃虫的抗性显著提高,说明SlMTB1,SlMTB2,SlMTB3基因与番茄对棉铃虫抗性相关。具体表现为:
(1)棉铃虫饲喂到第7天,对植物进行拍照记录。
结果如图1所示,饲喂7天后T2代pHELLSGATE2-SlMTB1/2/3-RNAi转基因番茄(图1中转基因植株)的生长点和叶片基本没受到破坏,基本保持完整;而作为对照的野生型番茄M82(图1中野生型)的生长点出现很多咀食的痕迹,叶片出现不少较大的空洞;野生型番茄M82与T2代转空载体番茄表现相同。
(2)棉铃虫饲喂到第7天,对饲喂相应植株的棉铃虫拍照记录并做重量统计。
结果如图2所示,饲喂7天后,咀食纯合的T2代pHELLSGATE2-SlMTB1/2/3-RNAi转基因番茄(图2中转基因植株)的棉铃虫体积(图2中A)和重量(图2中B)都显著低于咀食野生型番茄M82(图2中野生型)的棉铃虫;而咀食野生型番茄M82与T2代转空载体番茄的棉铃虫体积和重量都没有显著差异。
(3)PI-II基因(SEQ ID No.11)作为番茄茉莉酸信号转导途径中的一个重要的标记基因,长期以来一直被用来衡量对昆虫的抗性。
检测饲喂第2天后的纯合的T2代pHELLSGATE2-SlMTB1/2/3-RNAi转基因番茄和野生型番茄M82,以及T2代转空载体番茄中PI-II的mRNA相对表达量,方法如下:
A、番茄叶片RNA提取
总RNA的提取用Trizol法提取:
1)将各株系上述不同时间取的番茄叶片在液氮中磨成粉末后,取100mg叶片粉末转入加有1mL Trizol提取液的RNase-free离心管中,震荡混匀。
2)研磨液室温(25℃)放置5分钟,加入0.2mL氯仿,盖紧离心管,剧烈摇荡离心管20秒,室温放置3分钟。4℃12000g离心15分钟。
3)取上层水相约600μL于一新离心管,加0.5mL异丙醇,室温放置10分钟,4℃,12000g离心10分钟。
4)弃去上清液,加入1mL的75%乙醇,涡旋混匀,4℃下12000g离心1分钟。
5)重复步骤4),小心弃去上清液,然后室温或真空干燥10分钟,溶解于DEPC处理过的双蒸水中,得到RNA。
B、单链cDNA的合成
1)在200μL的RNase-free离心管中加入下列试剂:
模板RNA(总RNA) 2μg
引物Oligo(dT)18(0.5μg/μL) 1μL
RNase-free ddH2O 补至13μL
2)轻轻混匀后离心3~5s,70℃反应5分钟,迅速置于冰上冷却5分钟,然后加入:
3)终体积为25μL。轻轻混匀后稍离心,42℃反应60分钟,94℃反应5分钟后,得到cDNA,置冰上进行后续实验或-20℃冷冻保存,得到cDNA。
C、荧光定量RT-PCR分析
1)反应体系
2)反应程序
95℃,30s;45个循环((95℃,10s;60℃,20s;72℃,20s)
3)数据分析
Ct值为PCR管中荧光信号达到设定的域值时所经历的循环数。
ΔCt=Ct(Gene)-Ct(Actin),以2-ΔCt的值衡量基因的转录水平。
涉及到的定量PCR分析表达的基因及所用引物如下所示:
内参基因为Actin2,用于扩增内参基因的RT-PCR引物为:
SlACT2-QF:5′-TTGCTGACCGTATGAGCAAG-3′;
SlACT2-QR:5′-GGACAATGGATGGACCAGAC-3′。
检测PI-II的RT-PCR引物为:
SlPI-II-QF:5′-AATTATCCATCATGGCTGTTCAC-3′;
SlPI-II-QR:5′-CCTTTTTGGATCAGATTCTCCTT-3′。
结果如图3所示(图3中,将作为对照的野生型番茄M82未接虫时PI-II的mRNA相对表达量设定为1),饲喂棉铃虫前,T2代pHELLSGATE2-SlMTB1/2/3-RNAi转基因番茄(图3中转基因植株)与对照植物野生型番茄M82(图3中野生型)相比,PI-II的mRNA相对表达量没有显著差异;而当饲喂棉铃虫2天后,与对照植物野生型番茄M82相比,SlMTB1,SlMTB2,SlMTB3基因沉默的转基因番茄在受到昆虫嚼食后表达更多的PI-II,表明SlMTB1,SlMTB2,SlMTB3基因沉默的转基因番茄对昆虫的抗性显著提高,与图1中植物地上部分叶片被咀食程度和图2中棉铃虫的大小与重量的结果一致;而野生型番茄M82与T2代转空载体番茄相比,在未接虫和接虫2天后PI-II的mRNA相对表达量都没有显著差异。上述实验均重复3次,结果一致。
上述结果证明SlMTB1,SlMTB2,SlMTB3基因与番茄对棉铃虫抗性相关,说明SlMTB1,SlMTB2,SlMTB3蛋白及其编码基因可用于农牧业和生态环境治理所需的抗虫植物品种的培育与鉴定,在植物抗虫育种领域中具有重要作用。
最后说明的是,以上实施例仅用以说明本发明的技术方案而非限制,尽管通过参照本发明的优选实施例已经对本发明进行了描述,但本领域的普通技术人员应当理解,可以在形式上和细节上对其作出各种各样的改变,而不偏离所附权利要求书所限定的本发明的精神和范围。
<110> 中国科学院遗传与发育生物学研究所
<120> 一种提高番茄抗虫性的方法
<130> GNCLN172157
<160> 11
<170> PatentIn version 3.5
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atggttactg ggaatatgtt gtggagtggt gaggataagg ctatggtggc gtctgtttta 60
ggaaaggaag cttttgaata tttgatgtct ggctctgttt cagcagaatg ttctttaatg 120
gcaataggga atgatcagaa tttgcagaat aagctttcag atctcgtgga acgcccaaac 180
gccgctaatt ttagctggaa ttatgccatc ttttggcaaa tttcgcggtc taagtcaggg 240
gaattggtgc tagggtgggg ggatgggtgt tgcagagagc ctaaggaagc agaggagcgt 300
gaagttaaaa agattctcaa tctacgcctc gatgatgagg gtcaacaaag gatgaggaag 360
agggtacttc agaagctgca tatgttattt ggtggaacag atgaagataa ctatgctttt 420
ggattagata gagttactga tactgaaatg ttcttccttg cctcgatgta tttttcgttc 480
cctcgaggag agggaggtcc cgggaagtgt tttggatcag gtaagtattt gtggttatca 540
gatgcattga catctaatct agattattgt gctagatctt tcttagctaa gtctgctggt 600
atgcaaacta ttgctttgat tccaactgat gtaggagttg tggaattggg gtcagtgaga 660
tcgataccgg agagtttaga gctattacag aatataaaat cttgcttctc gtcgttttta 720
tcacttgtta gggataagca agcagcaggt atagcagctg tacctgagaa aaacgaggga 780
aacaatcccc gcctttccaa ctctggcgct gttactgaac gaacagatgg aaatcctaag 840
atatttgggc atgatttgaa ttctggtact cactttaggg aaaaacttgc tgttaggaaa 900
gcggaggaga ggccatggga catgtaccaa aatggtaaca ggatgccatt tgtgaacgca 960
cggaatggtt taaatcctgc ttcttgggct caattcagta atgtgaagct gggaaagcct 1020
gtggagctct atgctcctcc aaccccagga cacaacctga tgaatggtgg gagggaagaa 1080
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tcaccggtag ggcaataa 1818
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<212> DNA
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atggcagaga aattttttct gaagggagag gataaggtta atatggaggg ggtattaggc 60
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tggacttatg caatctattg gcaagttgca aagtcgaaat ctggaaaatc agctttaata 240
tggggcgatg gacattgcag agaaacaaag atagggcaag gtgaaggtgc aaatgattcc 300
gcgcatcaga aaatgatgga cggaaacaag aaaaagatgg ttcttcaaaa gattcacact 360
tgctttggag ggtcagaaga tgataatatt gctgccaagt tggaatctgt ttcggacgtg 420
gaggtgtttt atctcacatc aatgtattat atcttcccat tcgataagcc ttctagtcct 480
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ccgctgaaat caggggttgt ggagcttggt tctgtgaagt ctattccgga agatcagaat 660
ttgattcaga tggtgaaaac atcggtggtg gtatctaatc ctccgcagcc caaagcaaat 720
acaaagatat ttggtcggga actcagtcta ggtggcgcta agtcaggtcc catcagcata 780
aatttttctc cgaaggtgga agaagagctg agttttgctt cagattcgta cgaagtacaa 840
gcagcgctag gtagttctca ggtttatggg aactcatcaa acgggtatcg aagtgatgaa 900
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gagattgatt ttcatcaaag acaagatgat gcagttgtaa gagtaggctg ccctttgaat 1260
gctcaccctg tttctagagt tttgaagaca tttcaggaac atcaagtggt ggcacaagaa 1320
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<212> DNA
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atgactatgt tgtggagtga tgaggataag actatggtgg cagcggtatt aggaacaaaa 60
gcttttgatt acttgatgtc tagtttagtt tctgctgaat gttctttaat ggcaatgggg 120
agtgatgaga atttgcagaa tatgctatca gatcttgtgg aacgtccgaa tgcaagtaat 180
tttagttgga attatgcaat tttttggcag atttcgcggt ctaagttggg ggagttggtg 240
ttaggatggg gagatggttg ttgtagggaa gctagggaag gagaggaatc tgaacttact 300
aggattctca atatacgcct cgcggatgag gctcaacaaa ggatgaggaa gagggttctt 360
cagaagttgc atatgttttt tggtggaaca gatgaagata actatgtgtc tggattggat 420
aaagtcactg atactgaaat gttctttctt gcttcaatgt acttttcgtt tcctcgaggg 480
caaggtggtc cggggaaatg ttttactgct ggtaaacatg tgtggttatc ggatgttatg 540
aggtcgtctg tagattattg ttctaggtct tttctgatga agtctgctgg tatgcaaaca 600
gttgttttga tcccaactga tataggggtt atggaattgg gatctgtacg aactatacca 660
gaaagcttgg agcttgtaca ttcaataaaa tcttgcttct cgtcgttttt ggctcaagtt 720
agggctaagc aagcggcacc tttagcagca gttgtagctg agaaaaagaa tggaaacaat 780
tctgtgtttc ccagctcttt tccttttgat cagtcaaagg aaaatccgaa gatctttggg 840
cagaatttag aatctggttc taccgagttt agggaaaaac ttgctctcag gaaaccggtg 900
gatgggccac tggaaatgta tagaaatgga aacagggctc ccatcataaa tacacaaaat 960
ggtgtacgcc ccgtatcatg ggcttcattt ggtaatgtaa agccagggaa ctcggtggat 1020
ctctatagtc ctcaggcacc accaaacaac ttacgagagt ttgtcaatgg tggaagagaa 1080
gagctccggt taaatagcct ccagcatcaa aagcctggtg gaatgcaaat agatttcacc 1140
aactcaagac ccgttgtttc tccagtgccc actgttgaat cggagcattc agatgttgaa 1200
gtttcatgca aggaaaaaca tgcaggacca gctgatgaaa ggaggcctcg aaagcgtgga 1260
agaaaaccag ccaatggaag ggaagagccc ctcaatcatg tagaagctga gaggcaacgg 1320
agggagaagc tgaaccagcg gttttatgca ttacgagctg ttgttccaaa tatttccaaa 1380
atggacaaag cttccctctt aggagatgcc attgctcaca tcactgacat gcagaagagg 1440
atcagggacg cggaatataa gctggaaaag cgaggaagca cctctgtgga tgcagctgat 1500
atcaacattg aagccgccag tgatgaagtc atcgtaagag ctagatgccc actgggaacc 1560
catcctgtag caaaagtcgt tgaagcgttc aaagagacac aggtcagcgt tgttgagtca 1620
aaacttgccg tgggaaatga tacagtatat catacatttg tggtcaagtc cagtggaccc 1680
gaacagctaa cgaaggaaaa gctaatggct gcatttgctg gtgaatcaaa ctctctctag 1740
<210> 7
<211> 200
<212> DNA
<213> 人工序列
<220>
<223>
<400> 7
cctcttagga gatgccattg cttacataac tgagctgcag aaaaaactaa gagatatgga 60
atctgagagg gagctgagat taggaagcac ttcaagggac gcaatcactt cagaagacag 120
ccctagttct gagattcaaa tccgaggacc cgacatcaac atagaagctg ccaatgatga 180
agtcattgta agggtgagct 200
<210> 8
<211> 200
<212> DNA
<213> 人工序列
<220>
<223>
<400> 8
ccgctgaaat caggggttgt ggagcttggt tctgtgaagt ctattccgga agatcagaat 60
ttgattcaga tggtgaaaac atcggtggtg gtatctaatc ctccgcagcc caaagcaaat 120
acaaagatat ttggtcggga actcagtcta ggtggcgcta agtcaggtcc catcagcata 180
aatttttctc cgaaggtgga 200
<210>9
<211>200
<212>DNA
<213> 人工序列
<220>
<223>
<400>9
tagtttagtt tctgctgaat gttctttaat ggcaatgggg agtgatgaga atttgcagaa 60
tatgctatca gatcttgtgg aacgtccgaa tgcaagtaat tttagttgga attatgcaat 120
tttttggcag atttcgcggt ctaagttggg ggagttggtg ttaggatggg gagatggttg 180
ttgtagggaa gctagggaag 200
<210>10
<211>600
<212>DNA
<213> 人工序列
<220>
<223>
<400>10
cctcttagga gatgccattg cttacataac tgagctgcag aaaaaactaa gagatatgga 60
atctgagagg gagctgagat taggaagcac ttcaagggac gcaatcactt cagaagacag 120
ccctagttct gagattcaaa tccgaggacc cgacatcaac atagaagctg ccaatgatga 180
agtcattgta agggtgagct ccgctgaaat caggggttgt ggagcttggt tctgtgaagt 240
ctattccgga agatcagaat ttgattcaga tggtgaaaac atcggtggtg gtatctaatc 300
ctccgcagcc caaagcaaat acaaagatat ttggtcggga actcagtcta ggtggcgcta 360
agtcaggtcc catcagcata aatttttctc cgaaggtgga tagtttagtt tctgctgaat 420
gttctttaat ggcaatgggg agtgatgaga atttgcagaa tatgctatca gatcttgtgg 480
aacgtccgaa tgcaagtaat tttagttgga attatgcaat tttttggcag atttcgcggt 540
ctaagttggg ggagttggtg ttaggatggg gagatggttg ttgtagggaa gctagggaag 600
<210> 11
<211> 660
<212> DNA
<213> 番茄(Solanum lycopersicon)
<400> 11
gaccccaaaa ttaaaagaag acaaggtact agtaatcaat tatccatcat ggctgttcac 60
aaggaagtta attttgtcgc ttacctacta attgttcttg gaatgtttct atatgttgat 120
gccaaggctt gtactagaga atgtggtaat cttgggttcg ggatatgccc acgttcagaa 180
ggaagtccgc taaatcccat atgcatcaat tgttgctcag gctataaggg ttgtaattat 240
tataattctt tcggaaaatt tatttgtgaa ggagaatctg atccaaaaag gccaaatgct 300
tgcaccttta attgtgatcc aaatattgcc tattcaagat gtccccgttc acaaggaaaa 360
tcgttaattt atcccaccgg atgtaccacg tgttgcacag ggtacaaggg ttgctattat 420
tttggtaaag atggaaagtt tgtatgtgaa ggagagagtg atgaacccaa ggcaaatatg 480
taccctgtaa tgtgactcta gacttgtcca tcttctggat tgcccaaaat taagtaatta 540
atgtatgaaa taaaaggatg cacacttata taatgacatg ctaatcatta taatgtgggc 600
atcaagttgt gtgttatgtg taattactaa ttatctgaat aagagaatag atatccctaa 660
Claims (12)
1.蛋白或其相关生物材料在调控番茄抗虫性中的应用;
所述相关生物材料为能够表达所述蛋白的核酸分子,或含有所述核酸分子的表达盒、重组载体、重组菌或转基因细胞系;
所述蛋白为SlMTB1蛋白、SlMTB2蛋白和SlMTB3蛋白;
所述SlMTB1蛋白为如下(A1)或(A2)所示蛋白质:
(A1)氨基酸序列为SEQ ID No.1的蛋白质;
(A2)在(A1)所限定的蛋白质的N端和/或C端连接标签后得到的融合蛋白;
所述SlMTB2蛋白为如下(B1)或(B2)所示蛋白质:
(B1)氨基酸序列为SEQ ID No.2的蛋白质;
(B2)在(B1)所限定的蛋白质的N端和/或C端连接标签后得到的融合蛋白;
所述SlMTB3蛋白为如下(C1)或(C2)所示蛋白质:
(C1)氨基酸序列为SEQ ID No.3的蛋白质;
(C2)在(C1)所限定的蛋白质的N端和/或C端连接标签后得到的融合蛋白;
所述虫为咀嚼式昆虫。
2.根据权利要求1所述的应用,其特征在于:所述蛋白在番茄中的表达量越低,番茄的抗虫性越强。
3.根据权利要求1所述的应用,其特征在于:所述咀嚼式昆虫为棉铃虫。
4.根据权利要求1所述的应用,其特征在于:所述“能够表达所述蛋白的核酸分子”为所述SlMTB1蛋白的编码基因、所述SlMTB2蛋白的编码基因和所述SlMTB3蛋白的编码基因;
所述SlMTB1蛋白的编码基因是SEQ ID No.4所示的DNA分子;
所述SlMTB2蛋白的编码基因是SEQ ID No.5所示的DNA分子;
所述SlMTB3蛋白的编码基因是SEQ ID No.6所示的DNA分子。
5.能够使受体番茄中蛋白的表达量降低的物质在提高番茄抗虫性中的应用;
所述蛋白为SlMTB1蛋白、SlMTB2蛋白和SlMTB3蛋白;
所述SlMTB1蛋白为如下(A1)或(A2)所示蛋白质:
(A1)氨基酸序列为SEQ ID No.1的蛋白质;
(A2)在(A1)所限定的蛋白质的N端和/或C端连接标签后得到的融合蛋白;
所述SlMTB2蛋白为如下(B1)或(B2)所示蛋白质:
(B1)氨基酸序列为SEQ ID No.2的蛋白质;
(B2)在(B1)所限定的蛋白质的N端和/或C端连接标签后得到的融合蛋白;
所述SlMTB3蛋白为如下(C1)或(C2)所示蛋白质:
(C1)氨基酸序列为SEQ ID No.3的蛋白质;
(C2)在(C1)所限定的蛋白质的N端和/或C端连接标签后得到的融合蛋白;
所述“能够使受体番茄中蛋白的表达量降低的物质”为如下任一:
a)用于同时沉默所述SlMTB1蛋白、所述SlMTB2蛋白和所述SlMTB3蛋白的编码基因表达的RNA片段;
b)编码a)所述RNA片段的DNA片段;
c)含有b)中所述DNA片段的重组载体、表达盒、转基因细胞系或重组菌;
所述虫为咀嚼式昆虫。
6.根据权利要求5所述的应用,其特征在于:
a)所述RNA片段为SEQ ID No.10编码的RNA;
b)所述DNA片段的核苷酸序列为SEQ ID No.10。
7.根据权利要求5所述的应用,其特征在于:所述咀嚼式昆虫为棉铃虫。
8.一种培育抗虫性提高的番茄的方法,包括如下步骤:使受体番茄中蛋白的表达量降低,获得抗虫番茄;
所述蛋白为SlMTB1蛋白、SlMTB2蛋白和SlMTB3蛋白;
所述SlMTB1蛋白为如下(A1)或(A1)所示蛋白质:
(A1)氨基酸序列为SEQ ID No.1的蛋白质;
(A2)在(A1)所限定的蛋白质的N端和/或C端连接标签后得到的融合蛋白;
所述SlMTB2蛋白为如下(B1)或(B2)所示蛋白质:
(B1)氨基酸序列为SEQ ID No.2的蛋白质;
(B2)在(B1)所限定的蛋白质的N端和/或C端连接标签后得到的融合蛋白;
所述SlMTB3蛋白为如下(C1)或(C2)所示蛋白质:
(C1)氨基酸序列为SEQ ID No.3的蛋白质;
(C2)在(C1)所限定的蛋白质的N端和/或C端连接标签后得到的融合蛋白;
所述虫为咀嚼式昆虫。
9.根据权利要求8所述的方法,其特征在于:所述“使受体番茄中蛋白的表达量降低”是通过对所述受体番茄中所述蛋白的编码基因进行敲除或抑制表达来实现的。
10.根据权利要求9所述的方法,其特征在于:对所述受体番茄中的所述蛋白的编码基因进行抑制表达是通过向所述受体番茄中导入权利要求5或6中所述的“能够使受体番茄中蛋白的表达量降低的物质”来实现的。
11.根据权利要求8所述的方法,其特征在于:所述咀嚼式昆虫为棉铃虫。
12.根据权利要求9所述的方法,其特征在于:
所述SlMTB1蛋白的编码基因是SEQ ID No.4所示的DNA分子;
所述SlMTB2蛋白的编码基因是SEQ ID No.5所示的DNA分子;
所述SlMTB3蛋白的编码基因是SEQ ID No.6所示的DNA分子。
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