CN110079535B - 玉米ZmPIF3s突变型蛋白、其编码基因及其在育种上的应用 - Google Patents
玉米ZmPIF3s突变型蛋白、其编码基因及其在育种上的应用 Download PDFInfo
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Abstract
本发明公开了玉米ZmPIF3s突变型蛋白、其编码基因及其在育种上的应用。本发明利用基因编辑对玉米ZmPIF3s蛋白进行突变,获得具有使株高变矮以及中胚轴长度缩短功能的ZmPIF3s突变蛋白。本发明还提供了ZmPIF3s基因编辑载体的构建方法。本发明进一步公开了利用基因编辑技术创制短中胚轴和矮杆玉米的育种方法。本发明利用CRISPR/Cas9基因编辑技术对ZmPIF3s基因进行编辑,通过后代的筛选在T2代就可获得剔除T‑DNA、具有短中胚轴和矮杆特性稳定遗传的新材料。本发明所提供的突变蛋白以及基因编辑技术能够应用于植物的分子育种,具有快速、精准、高效等优点,能有效提高育种效率并加快育种进程。
Description
技术领域
本发明涉及基于基因编辑技术获得的ZmPIF3s突变型蛋白及其编码基因,本发明进一步涉及它们在植物育种上的应用,属于玉米ZmPIF3s突变型蛋白及其应用领域。
背景技术
玉米的株高往往受到许多微效数量性状位点(QTL)的控制,现在的玉米育种依然需要通过不断累积这些微效QTL来降低株高,选择效率低下。在水稻和小麦中,半矮杆基因sd1和rht1的利用直接导致了第一次绿色革命,作物的产量得到了很大的提升,然而在玉米中类似这种绿色革命基因的、既能有效降低株高又没有明显穗部负向效应的主效半矮杆基因依然没有找到。
光敏色素(Phytochrome,Phy)是植物体内重要的光受体,能够感知红光和远红光。光激活后的光敏色素进入细胞核,通过与光敏色素相互作用因子(Phytochrome-interacting factors,PIFs)互作,调控靶基因的表达。PIFs属于basic helix-loop-helix(b HLH)转录因子家族,通过介导光信号,调控植物的生长发育过程。拟南芥中共含有7个PIF(PIF1、PIF3、PIF4、PIF5、PIF6、PIF7、PIF8)成员。拟南芥PIFs在黑暗条件下抑制植物的光形态建成,功能缺失型单突变体和多突变体不论在黑暗还是光照条件下都表现出明显的光形态建成表型。pif3突变体在持续红光下,胚轴生长比野生型要短。目前已报道的玉米中含有3个PIF3同源基因,分别是ZmPIF3.1,ZmPIF.2和ZmPIF3.3。结构上,PIF3蛋白包含三个特征结构域,即同时含有phy A结合域和phy B结合域,以及和DNA结合的b HLH结构域。
基因编辑技术是对基因组进行定向编辑的一项新技术。其原理是通过序列特异性核酸酶对目标DNA片段进行剪切,在修复过程中使靶位点发生插入、缺失和替换等突变,人工使基因组发生定点改变而获得预期目标性状。其中,CRISPR/Cas9(Clustered RegularlyInterspaced Short Palindromic Repeat and Cas9)作为新一代基因组编辑技术,具有操作简便、切割效率高、靶位点多等优点,在模式植物和作物中得到广泛的应用。玉米是世界范围上种植范围最广,产量最高的粮食作物,同时也是饲料和工业加工的重要原料,因此通过CRISPR/Cas9基因编辑技术对玉米基因组的定向准确修饰,可以快速转化现有的研究成果,实现重要农艺性状的改良,这对于玉米功能基因的基础研究与应用研究均具有重要的意义。
采用基因编辑技术对玉米ZmPIF3s蛋白进行突变获得能够使株高矮化或使中胚轴长度缩短的突变蛋白,这对于解析其遗传调控网络以及对于玉米进行分子改良育种均具有重要的理论和实践意义。
发明内容
本发明的目的之一是提供玉米ZmPIF3s突变型蛋白及其编码基因;
本发明的目的之二是提供ZmPIF3s基因编辑载体;
本发明的目的之三是将ZmPIF3s突变型蛋白及其编码基因以及所构建的ZmPIF3s基因编辑载体应用于玉米分子改良育种。
本发明的上述目的是通过以下技术方案来实现的:
本发明首先提供了玉米ZmPIF3s突变型蛋白,由ZmPIF3.1突变型蛋白、ZmPIF3.2突变型蛋白以及ZmPIF3.3突变型蛋白组成;其中,所述ZmPIF3.1突变型蛋白的氨基酸序列为SEQ ID No.34或SEQ ID No.36所示,其编码基因的核苷酸序列为SEQ ID No.33或SEQ IDNo.35所示;所述ZmPIF3.2突变型蛋白的氨基酸序列为SEQ ID No.38或SEQ ID No.40所示,其编码基因的核苷酸序列为SEQ ID No.37或SEQ ID No.39所示;所述ZmPIF3.3突变型蛋白的氨基酸序列为SEQ ID No.42或SEQ ID No.44所示,其编码基因的核苷酸序列为SEQ IDNo.41或SEQ ID No.43所示。
本发明还提供了含有所述编码基因的重组植物表达载体以及含有该重组植物表达载体的宿主细胞;将所述编码基因可操作的与表达调控元件相连接得到可以在植物中表达该编码基因的重组植物表达载体。
所述重组植物表达载体还可含有用于选择转化细胞的选择性标记基因;所述选择性标记基因用于选择经转化的细胞或组织。标记基因包括:编码抗生素抗性的基因以及赋予除草化合物抗性的基因等。此外,所述的标记基因还包括表型标记,例如β-半乳糖苷酶和荧光蛋白等。
本发明中所述的转化方案以及将所述多核苷酸或多肽引入植物的方案可视用于转化的植物(单子叶植物或双子叶植物)或植物细胞的类型而变化。将所述多核苷酸或多肽引入植物细胞的合适方法包括:显微注射、电穿孔、农杆菌介导的转化、直接基因转移以及高速弹道轰击等。利用常规方法可使已转化的细胞再生稳定转化植株(McCormick etal.Plant Cell Reports.1986.5:81-84)。
本发明可用于转化任何植物种类,包括但不限于:单子叶植物或双子叶植物,优选是玉米。
本发明的另一个目的是提供一种ZmPIF3s基因编辑载体,其构建方法包括:
(Ⅰ)、获取玉米U6-1启动子片段;
(Ⅱ)、制备6个sgRNA表达盒:
(Ⅲ)、将步骤(Ⅱ)获得的6个sgRNA表达盒依次连接CPB-Ubi-hspcas9载体,即得。
其中,步骤(Ⅰ)中所述的制备sgRNA表达盒包括以下步骤:
(一)、采用重叠PCR的方法将带有接头的靶点序列和sgR骨架序列融合在一起获得6个PCR产物;其中,PCR的上游引物分别分别为SEQ ID No.19-SEQ ID No.24所示;下游引物为SEQ ID No.27所示,模板序列为SEQ ID No.28所示;该重叠PCR反应程序如下:(1)94℃2min;(2)98℃,10s;46℃,30s;68℃10s;共35个循环;(3)68℃5min。
(二)、再次用重叠PCR的方法将步骤(一)获得的6个融合PCR片段和U6-1启动子片段分别连接获得的6个PCR产物得到sgRNA连接产物;其中,该重叠PCR中所用到的上游、下游引物序列为SEQ ID No.29和SEQ ID No.30所示;所用到的模板是U6-1启动子片段以及步骤(一)扩增得到的6个PCR产物;该重叠PCR反应程序如下:(1)94℃2min;(2)98℃,10s;52-55℃,30s;68℃40s;共35个循环;(3)68℃5min。
本发明选取背景材料为xiang249,通过所构建的CRISPR/Cas9基因编辑载体同时对玉米中ZmPIF3.1,ZmPIF3.2和ZmPIF3.3三个目标基因进行打靶进行定点编辑,经过筛选从中获得了相对于野生型材料具有短中胚轴和矮杆的剔除外源标记基因的Zmpif3s-#1突变体材料以及Zmpif3s-#2突变体材料:
在Zmpif3s-#2突变体材料中,ZmPIF3.1基因从37bp处开始切割删除,共删除842bp,删除后导致移码突变,提前出现终止子;ZmPIF3.2基因从137bp处开始切割删除,共删除814bp,删除后导致移码突变,提前出现终止子;ZmPIF3.3基因从631bp处开始切割删除,而且删除后又导致后面的序列发生倒位,最终导致提前出现终止子。
在Zmpif3s-#2突变体材料中,ZmPIF3.1基因从38bp处开始切割删除,共删除630bp,而且删除后又有CCTG 4bp的插入,删除后导致移码突变,提前出现终止子;ZmPIF3.2基因从第2bp处开始切割删除,删除基因组序列共51bp,删除后导致移码突变,提前出现终止子;ZmPIF3.3基因从630bp处开始切割删除,共删除960bp,删除导致移码突变,最后往3’端延伸,出现新的终止子。
所筛选得到的Zmpif3s-#1突变体材料和Zmpif3s-#2突变体材料因基因ZmPIF3s功能异常导致中胚轴缩短和株高降低,这表明ZmPIF3s蛋白的缺失对玉米的株型(中胚轴和株高)调控起着至关重要的作用。本发明通过CRISPR/Cas9基因编辑技术对ZmPIF3s基因进行敲除突变,最终获得突变后的ZmPIF3s蛋白,属于功能缺失型突变,只要是通过CRISPR/Cas9技术导致的ZmPIF3s基因的突变,尽管突变后的序列可能多样化,但是可以合理的预测这些多样化的突变序列都会导致株高降低或中胚轴长度缩短表型的出现,相应的,采用CRISPR/Cas9技术对ZmPIF3s基因进行突变所获得的突变蛋白以及编码基因均应属于本发明的保护范畴之内
因此,本发明进一步将获得的ZmPIF3s突变蛋白或所构建的ZmPIF3s基因编辑载体应用于降低植物株高或缩短中胚轴长度;具体的,可以将ZmPIF3s突变蛋白或ZmPIF3s基因编辑载体应用于培育植物新品种,其中所述的植物新品种可以是矮杆或短中胚轴植物新品种。
本发明提供了一种使植物株高变矮或缩短中胚轴长度的方法,包括:
(1)将所筛选到的编码ZmPIF3s突变蛋白的编码基因可操作的与调控元件连接后构建得到重组植物表达载体;(2)将所构建的重组植物表达载体转化到植物中筛选得到株高变矮或中胚轴长度缩短的植物新品种;或者,将构建的ZmPIF3s基因编辑载体转化到植物中筛选得到株高变矮或中胚轴长度缩短的植物新品种。
所述重组植物表达载体还可含有用于选择转化细胞的选择性标记基因。选择性标记基因用于选择经转化的细胞或组织。标记基因包括:编码抗生素抗性的基因以及赋予除草化合物抗性的基因等。此外,所述的标记基因还包括表型标记,例如β-半乳糖苷酶和荧光蛋白等。
本发明中所述的转化方案以及将所述多核苷酸或多肽引入植物的方案可视用于转化的植物(单子叶植物或双子叶植物)或植物细胞的类型而变化。将所述多核苷酸或多肽引入植物细胞的合适方法包括:显微注射、电穿孔、农杆菌介导的转化、直接基因转移以及高速弹道轰击等。利用常规方法可使已转化的细胞再生稳定转化植株(McCormick etal.Plant Cell Reports.1986.5:81-84)。
本发明可用于转化任何植物种类,包括但不限于:单子叶植物或双子叶植物,优选是玉米。
本发明首次利用基因编辑技术对玉米ZmPIF3s基因进行定点突变创制新的矮杆玉米新等位基因并剔除T-DNA外源序列,获得表型稳定遗传的株系;本发明所提供的编辑分子育种方法具有精准、效率高等常规育种不具备的优势,具有广泛的应用前景。
本发明所涉及到的术语定义
除非另外定义,否则本文所用的所有技术及科学术语都具有与本发明所属领域的普通技术人员通常所了解相同的含义。虽然在本发明的实践或测试中可使用与本文所述者类似或等效的任何方法、装置和材料,但现在描述优选方法、装置和材料。
术语“多核苷酸”或“核苷酸”意指单股或双股形式的脱氧核糖核苷酸、脱氧核糖核苷、核糖核苷或核糖核苷酸及其聚合物。除非特定限制,否则所述术语涵盖含有天然核苷酸的已知类似物的核酸,所述类似物具有类似于参考核酸的结合特性并以类似于天然产生的核苷酸的方式进行代谢。除非另外特定限制,否则所述术语也意指寡核苷酸类似物,其包括PNA(肽核酸)、在反义技术中所用的DNA类似物(硫代磷酸酯、磷酰胺酸酯等等)。除非另外指定,否则特定核酸序列也隐含地涵盖其保守修饰的变异体(包括(但不限于)简并密码子取代)和互补序列以及明确指定的序列。特定而言,可通过产生其中一个或一个以上所选(或所有)密码子的第3位经混合碱基和/或脱氧肌苷残基取代的序列来实现简并密码子取代(Batzer等人,Nucleic Acid Res.19:5081(1991);Ohtsuka等人,J.Biol.Chem.260:2605-2608(1985);和Cassol等人,(1992);Rossolini等人,Mol Cell.Probes 8:91-98(1994))。
术语“多肽”、“肽”和“蛋白”在本文中互换使用意指氨基酸残基的聚合物。即,针对多肽的描述同样适用于描述肽和描述蛋白,且反之亦然。所述术语适用于天然产生氨基酸聚合物以及其中一个或一个以上氨基酸残基为非天然编码氨基酸的氨基酸聚合物。如本文中所使用,所述术语涵盖任何长度的氨基酸链,其包括全长蛋白(即抗原),其中氨基酸残基经由共价肽键连接。
术语“重组宿主细胞株”或“宿主细胞”意指包含本发明多核苷酸的细胞,而不管使用何种方法进行插入以产生重组宿主细胞,例如直接摄取、转导、f配对或所属领域中已知的其它方法。外源性多核苷酸可保持为例如质粒的非整合载体或者可整合入宿主基因组中。宿主细胞可为原核细胞或真核细胞,宿主细胞还可为单子叶或双子叶植物细胞。
术语“可操作的连接”指两个或更多个元件之间功能性的连接,可操作的连接的元件可为邻接或非邻接的。
术语“转化”:将异源性DNA序列引入到宿主细胞或有机体的方法。
术语“表达”:内源性基因或转基因在植物细胞中的转录和/或翻译。
术语“编码序列”:转录成RNA的核酸序列。
术语“重组植物表达载体”:一种或多种用于实现植物转化的DNA载体;本领域中这些载体常被称为二元载体。二元载体连同具有辅助质粒的载体是大多常用于土壤杆菌介导转化的。二元载体通常包括:T-DNA转移所需要的顺式作用序列、经工程化处理以便能够在植物细胞中表达的选择标记物,待转录的异源性DNA序列等。
附图说明
图1 ZmPIF3s基因编辑前后测序比对结果。
图2大田中野生型和Zmpif3s突变体植株的株高分析结果。
图3野生型和Zmpif3s突变体植株的暗形态建成的表型。
具体实施方式
下面将结合实施例对本发明的实施方案进行详细描述,但是本领域技术人员将会理解,下列实施例仅用于说明本发明,而不应视为限定本发明的范围。实施例中未注明具体条件者,按照常规条件或制造商建议的条件进行。所用试剂或仪器未注明生产厂商者,均为可以通过市购获得的常规产品。
实施例1玉米短中胚轴和矮杆突变体Zmpif3s的构建
1、玉米ZmPIF3s基因的靶位点设计
首先从玉米Gramene数据库中获取ZmPIF3.1(GRMZM2G115960-T03)(其核苷酸序列为SEQ ID No.1所示,氨基酸序列为SEQ ID No.2所示),ZmPIF3.2(GRMZM2G387528-T02)(其核苷酸序列为SEQ ID No.3所示,其氨基酸序列为SEQ ID No.4所示)和ZmPIF3.3(GRMZM2G062541-T01)(其核苷酸序列为SEQ ID No.5所示,其氨基酸序列为SEQ ID No.6所示)的基因组序列,然后用Snap Gene Viewer 3.2软件对每个ZmPIF3s基因设计两个靶位点。靶点序列分别是:
ZmPIF3.1-Guide 1:GTCGAGCTGCTATGGGAGAA(SEQ ID No.7)
ZmPIF3.1-Guide 2:GGGGAAAATGGAGACCAAGA(SEQ ID No.8)
ZmPIF3.2-Guide 1:GCAGCAGCAGAGCTCAACAA(SEQ ID No.9)
ZmPIF3.2-Guide 2:GAAAG AAGTAGAGAC CAGGA(SEQ ID No.10)
ZmPIF3.3-Guide 1:GCTGTTCTCCGAGCAGAGGA(SEQ ID No.11)
ZmPIF3.3-Guide 2:GCAAGTGCAGATGATGTCGA(SEQ ID No.12);
然后利用CTAB的方法提取玉米野生型自交系Xiang249的基因组DNA。
ZmPIF3.1-F1:5'GTCCATTTCCATCCAGGTCC3'(SEQ ID No.13),
ZmPIF3.1-R1:5'CCACCGGACTCATCATCAAG3'(SEQ ID No.14);
ZmPIF3.2-F1:5'GTGAGCGGTTCCTTCTCATT3'(SEQ ID No.15),
ZmPIF3.2-R1:5'CTCCTCTGCAGGTTTGTTGT3'(SEQ ID No.16);
ZmPIF3.3-F1:5'AGGGGAGGGCGTCTTGAACT3'(SEQ ID No.17),
ZmPIF3.3-R1:5'GGAACGGGAGCATGTCGAAC3'(SEQ ID No.18);
分别用上述引物(SEQ ID No.13-SEQ ID No.18)对所提的基因组DNA进行常规PCR扩增,将扩增产物送测序公司测序。测序结果与玉米Gramene数据库中的ZmPIF3.1,ZmPIF3.2和ZmPIF3.3的参考序列进行Blast比对分析,发现Xiang249的ZmPIF3.1、ZmPIF3.2和ZmPIF3.3的所设计的靶点序列与参考基因组B73相同,以确认转基因受体材料Xiang249的靶点序列。
最后合成带有接头的靶点序列引物,用于后续的载体构建。引物序列如下:
ZmPIF3.1-1F:
GAGCCGCAAGCACCGAATTGTCGAGCTGCTATGGGAGAAGTTTTAGAGCTAGAAATAGCAAGTT(SEQID No.19)
ZmPIF3.1-2F:
GAGCCGCAAGCACCGAATTGGGGAAAATGGAGACCAAGAGTTTTAGAGCTAGAAATAGCAAGTT(SEQID No.20)
ZmPIF3.2-1F:
GAGCCGCAAGCACCGAATTGCAGCAGCAGAGCTCAACAAGTTTTAGAGCTAGAAATAGCAAGTT(SEQID No.21)
ZmPIF3.2-2F:
GAGCCGCAAGCACCGAATTGAAAGAAGTAGAGACCAGGAGTTTTAGAGCTAGAAATAGCAAGTT(SEQID No.22)
ZmPIF3.3-1F:
GAGCCGCAAGCACCGAATTGCTGTTCTCCGAGCAGAGGAGTTTTAGAGCTAGAAATAGCAAGTT(SEQID No.23)
ZmPIF3.3-2F:
GAGCCGCAAGCACCGAATTGCAAGTGCAGATGATGTCGAGTTTTAGAGCTAGAAATAGCAAGTT(SEQID No.24)
其中,黑色字体部分为连接U6-1启动子片段的接头引物序列,斜体字部分为设计的靶点序列,下划线部分为连接sgR骨架片段的接头引物序列。
2、CRISPR/Cas9基因编辑载体构建
基因编辑载体构建参考Li C等(Li C,et al.,Plant Biotechnol J,2017,15:1566-1576)的报道方法,按以下步骤进行:
(1)玉米U6-1启动子片段的制备
引物序列为
MU61-1F:5'TGCTTTTTTTAAGCTGCTGTTTTTGTTAGCCCCATCG3'(SEQ ID No.25),
MU61-1R:
5'AATTCGGTGCTTGCGGCTC3'(SEQ ID No.26);
模板为B73基因组DNA,
PCR扩增体系为表1
表1 PCR扩增体系
成分 | 体积 |
2×PCR Buffer for KOD Fx | 25μL |
2mM dNTPs | 10μL |
MU61-1F | 1.5μL |
MU61-1R | 1.5μL |
KOD Fx | 1μL |
模板 | 1μL |
Add ddH<sub>2</sub>O | Up to 50μL |
PCR反应程序如下:
获得的PCR产物即为玉米U6-1启动子片段。
(2)sgRNA表达盒的制备
首先用重叠PCR的方法将带有接头的靶点序列和sgR骨架序列融合在一起获得的6段PCR产物,分别命名为ZmPIF3.1-1片段,ZmPIF 3.1-2片段,ZmPIF 3.2-1片段,ZmPIF 3.2-2片段,ZmPIF 3.3-1片段,ZmPIF 3.3-2片段。PCR扩增体系为表2。
表2 PCR扩增体系
其中上游引物分别是ZmPIF3.1-1F,ZmPIF3.1-2F,ZmPIF3.2-1F,ZmPIF3.2-2F,ZmPIF3.3-1F,ZmPIF3.3-2F,下游引物为MUsgR-R:GGCCAGTGCCAAGCTTAAAAAAAGCACCGACTCG(SEQ ID No.27);
模板序列为sgR骨架序列,人工合成sgR骨架片段序列如下:
GTTTTAGAGCTAGAAATAGCAAGTTAAAATAAGGCTAGTCCGTTATCAACTTGAAAAAGTGGCACCGAGTCGGTGCTTTTTTT(SEQ ID No.28)
PCR反应程序如下:
然后再次用重叠PCR的方法将上一步获得的融合PCR片段和U6-1启动子片段分别连接上,获得的PCR产物即为sgRNA连接产物,分别命名为U61-ZmPIF3.1-1片段,U61-ZmPIF3.1-2片段,U61-ZmPIF 3.2-1片段,U61-ZmPIF 3.2-2片段,U61-ZmPIF 3.3-1片段,U61-ZmPIF3.3-2片段。
引物序列为:
MU61-2F:TGCACTGCACAAGCTGCTGTTTTTGTTAGCCCCATCG(SEQ ID No.29)
MU61-1F:TGCTTTTTTTAAGCTGCTGTTTTTGTTAGCCCCATCG(SEQ ID No.30)
PCR扩增体系见表3。
表3 PCR扩增体系
其中,片段1为U6-1启动子片段,片段2分别为ZmPIF3.1-1片段,ZmPIF 3.1-2片段,ZmPIF 3.2-1片段,ZmPIF 3.2-2片段,ZmPIF 3.3-1片段,ZmPIF 3.3-2片段。在连接U6-1启动子片段和第一个靶位点PCR片段(如ZmPIF3.1-1片段)要用MU61-2F及MusgR-R引物,连接第二个或更多个靶位点PCR片段(如ZmPIF3.1-2片段及其他片段)用MU61-1F及MusgR-R引物。
PCR反应程序如下:
(3)sgRNA表达盒连接CPB-Ubi-hspcas9载体
按以下反应体系和过程,将每个sgRNA表达盒依次连接CPB-Ubi-hspcas9载体,获得连接产物。由于每个ZmPIF3基因都设计了两个靶位点,而且每个靶点都是用的玉米U6-1启动子,故总共需要连接6次,才能将ZmPIF3s的所有靶点连接到CPB-Ubi-hspcas9载体中。反应体系及过程见表4。
表4反应体系及过程
成分 | 体积 |
sgRNA连接产物 | 1μL |
CPB-pubi-hspcas9载体片段 | 1μL |
重组酶 | 0.5μL |
重组酶为Clontech公司的In-Fusion酶,反应条件为50℃,30min。
其中CPB-pubi-hspcas9载体片段是通过CPB-pubi-hspcas9质粒经HindIII单酶切后获得。sgRNA连接产物为U61-ZmPIF3.1-1片段,U61-ZmPIF 3.1-2片段,U61-ZmPIF 3.2-1片段,U61-ZmPIF 3.2-2片段,U61-ZmPIF 3.3-1片段,U61-ZmPIF 3.3-2片段,但需要注意的是,每次只能连接1个片段,每次连接完一个片段后需经测序核实后再用HindIII单酶切连接后的质粒载体,用于后续的连接构建。
(4)转化大肠杆菌DH5α及验证
将连接产物用热激法42℃转化大肠杆菌DH5α,菌液涂布于含有50mg/L卡那霉素的平板上,37℃培养约12h。挑取平板上长出的单菌落,摇菌扩繁。以菌液为模板进行PCR验证。PCR扩增体系见表5。
表5 PCR扩增体系
成分 | 体积 |
10×Taq Buffer | 2.5μL |
dNTPs(2.5mM each) | 2μL |
上游引物 | 0.5μL |
下游引物 | 0.5μL |
rTaq(2.5U/μL) | 0.5μL |
菌液 | 1μL |
Add ddH<sub>2</sub>O | Up to 25μL |
其中根据载体设计的引物如下:
上游引物为Ubi-4F:5’CTTAGACATGCAATGCTCATTATCTC3’(SEQ ID No.31),
下游引物为PstI-R:5’CTGGCGAAAGGGGGATGT3’(SEQ ID No.32),
用于检测阳性克隆,PCR反应程序如下:
提取检测含目的条带的菌液的质粒送公司测序。测序结果正确的质粒经HindIII单酶切后获得第一次的连接产物载体片段,然后再重复步骤(3)和(4),依次将其他的sgRNA连接产物连接到CPB-pubi-hspcas9载体中。
3、纯合的去除外源标记基因的Zmpif3s突变体的获得
将上述阳性质粒转入农杆菌EHA105。采用常规农杆菌介导法转化玉米自交系Xiang249获得T0代转基因植株。将T0代植株以单株收种,收获的种子播于大田中,在玉米长至两叶一心时,用Basta试剂(1/1000,V/V)涂抹叶尖,筛选去除载体的(Basta涂抹显示阴性)T1代植株。然后取样提取T1代植株的基因组DNA为模板,鉴定编辑位点的删除片段大小。PCR扩增体系如下见表6。
表6 PCR扩增体系
成分 | 体积 |
2×Msater Mix | 7.5μL |
上游引物 | 0.3μL |
下游引物 | 0.3μL |
模板DNA | 1μL |
Add ddH<sub>2</sub>O | Up to 15μL |
其中每个ZmPIF3基因检测引物分别用ZmPIF3.1-F1/ZmPIF3.1-R1,ZmPIF3.2-F1/ZmPIF3.2-R1,ZmPIF3.3-F1/ZmPIF3.3-R1。模板DNA分别为野生型植株Xiang249和突变体植株的基因组DNA。PCR反应程序如下:
由于每个ZmPIF3基因都设计了两个靶点,故理论上如果Cas9同时在两个靶点准确切割,则每个Zmpif3纯合突变体经PCR检测后应该获得单一的小带片段。野生型植株则PCR扩增后应该获得没有切割的单一的大带片段(表7):
表7 PCR扩增结果
大带bp | 小带bp | 检测基因 |
1500 | 800 | ZmPIF3.1 |
1420 | 800 | ZmPIF3.2 |
1177 | 320 | ZmPIF3.3 |
经初步PCR筛选鉴定后,选取单一的小带片段连接到克隆载体中进行测序鉴定,测序比对结果如图1所示。
在Zmpif3s#1突变体材料中,ZmPIF3.1和ZmPIF3.2基因经Cas9在所设计的两靶位点之间切割后导致删除突变,其中ZmPIF3.1删除842bp,ZmPIF3.2删除814bp,而且删除序列导致删除之后的基因序列移码突变。ZmPIF3.3基因从630bp处开始删除,删除后又有部分序列的倒位,而且也出现移码突变,提前出现终止子。
在Zmpif3s#2突变体材料中,ZmPIF3.1基因经Cas9在所设计的两靶位点之间切割后导致630bp的删除突变和4bp的插入突变,导致插入后的碱基序列移码突变;ZmPIF3.2和ZmPIF3.3基因经Cas9在所设计的两靶位点之间切割后导致删除突变,其中ZmPIF3.2删除51bp,ZmPIF3.3删除960bp。
上述在Zmpif3s突变体材料中发生的删除、插入和倒位突变,都使ZmPIF3s蛋白被成功敲除。
试验例1 Zmpif3s突变体的表型观察试验
1、将野生型与取实施例1中鉴定到的阴性的纯合Zmpif3s突变体(#1,#2)分别种植于海南省三亚市乐东黎族自治县尖峰镇翁毛村万钟公司产学研示范园和河北省廊坊市广阳区万庄镇伊指挥营村国际高新技术产业园基地,野生型与突变体种植在同一小区,每小区15株苗,三次重复。对大田中野生型和突变体植株的株高进行分析发现,Zmpif3s突变体的株高显著低于野生型(P<0.01)(图2)。这一结果说明,基因编辑获得的ZmPIF3s突变蛋白具有降低株高的功能。
2、同样,为了观察突变体植株的暗形态建成的表型,将野生型植株和Zmpif3s突变体(#1,#2)种植于培养箱中,统一种植深度为2cm,然后暗培养10d后取黄化苗照相观察表型。结果发现Zmpif3s突变体黄化苗的中胚轴极端,统计分析结果表明Zmpif3s突变体的中胚轴长度极显著短于野生型(P<0.001)(图3)。这一结果说明,基因编辑获得的ZmPIF3s突变蛋白具有缩短中胚轴长度的功能。
SEQUENCE LISTING
<110> 华南农业大学、中国农业科学院生物技术研究所
<120> 玉米ZmPIF3s突变型蛋白、其编码基因及其在植物育种上的应用
<130> BJ-2002-190319A
<160> 44
<170> PatentIn version 3.5
<210> 1
<211> 1704
<212> DNA
<213> Zea mays L.
<400> 1
atgtccgaca gcaacgactt cgtcgagctg ctatgggaga acggccaggc ggtggtgcac 60
gggaggagga agcagcatcc ccaggccgcc ttcccgccct tcacctgcgg cgcggccagc 120
agcagcagag ctcaagaaaa ccagccgggt actaccgacc cggtctcgct gttcaagacg 180
ggaggcctct tcgccgactt ctcgtccggc ctagacgccg cccgtggcaa cggtgacctt 240
gacgacaccg tgccctggat ccactgcccc atcgtcgagg aagacagcgc cgcacccgca 300
cctgccctcg cggagggcta cagcccagat ttcttctcgg agctccatgc agcggcgtta 360
gcagcggcgg cggcagagac gaacctcagc cctctgccac caccggtcca gcacaacaga 420
agcaccccgg ttgccaccac cagcagagga ccagaaccct cgaaggaagc ccagcgcata 480
ccagtaccag gtccaggcag caggcccgag ccacaatctg agttcgcggc caccaggaaa 540
cctcggccgg agagcggtgg agagggcttg atgaacttct ccctcttctc taggcctgca 600
gccttggtcc gggcaagcct acagaggcca ccgccaccgc agacaggcac tgacaaggtg 660
tccaatgtca ccacgagcac ccgcgtggag tcgacggttc tacagtcggc tagtgggcca 720
agaatcgccc ctgtgttcac ggaccagagg acggcgtggt cacaatccaa ggaggtgcgg 780
ttttcatgcg caccagcgct ggcggctggt aacctgcacc aggatatgcc tctgggaaga 840
cctggcaaca acatgactcc ccaggggaaa atggagacca agaaggcttg tgaggttgca 900
gtcgctactc catcgctctg ctccggcaat ggcgagtcct ggcgcgaaca aaagaggaag 960
agtcaggccg aatgttctgc aagtcaagac gatgatcttg atgatgagtc cggtgggatg 1020
cgaggatcag gcggcagagg tacgaagcgc agccgcaccg cggaggtgca caatttgtcc 1080
gaaaggagga gaagggacag gatcaatgag aaaatgcgcg ccctgcaaga gctcatcccc 1140
aactgcaaca agattgacaa agcctccatg ctggatgaag cgatcgagta cctcaaaacc 1200
ctccagcttc aagttcagat gatggccatg ggaagtggac tgtgcattcc tcccatgctg 1260
ctgccgcgag ccatgcagct gcaaattcct tcaattgctc atttccatca tctcggcatg 1320
ggattgggtt atgggatggg tgtcctcgat atgaacagca cagcggcagt tccatttcag 1380
cccatccctg gtgctcacta tacttgtccg atgatcccag gcacgccacc acaaggtctt 1440
ggaattggaa tgcctggcac aaacacgatg ccagtgtttg gactccctgg gcaagcaatc 1500
catccctcag cgtctagcgt acagccattc ccatcgttgg cgggtcttcc tgttaggcca 1560
aacctggcac ctcaagtccc agccgtcatg gctaacatgg tgcaggagca acagcaaggc 1620
gtagcgactc agcagcagca gagtctgaat aatgaagctc aacagaaagc aaataccgga 1680
gatccacaaa tactacaaaa atag 1704
<210> 2
<211> 567
<212> PRT
<213> Zea mays L.
<400> 2
Met Ser Asp Ser Asn Asp Phe Val Glu Leu Leu Trp Glu Asn Gly Gln
1 5 10 15
Ala Val Val His Gly Arg Arg Lys Gln His Pro Gln Ala Ala Phe Pro
20 25 30
Pro Phe Thr Cys Gly Ala Ala Ser Ser Ser Arg Ala Gln Glu Asn Gln
35 40 45
Pro Gly Thr Thr Asp Pro Val Ser Leu Phe Lys Thr Gly Gly Leu Phe
50 55 60
Ala Asp Phe Ser Ser Gly Leu Asp Ala Ala Arg Gly Asn Gly Asp Leu
65 70 75 80
Asp Asp Thr Val Pro Trp Ile His Cys Pro Ile Val Glu Glu Asp Ser
85 90 95
Ala Ala Pro Ala Pro Ala Leu Ala Glu Gly Tyr Ser Pro Asp Phe Phe
100 105 110
Ser Glu Leu His Ala Ala Ala Leu Ala Ala Ala Ala Ala Glu Thr Asn
115 120 125
Leu Ser Pro Leu Pro Pro Pro Val Gln His Asn Arg Ser Thr Pro Val
130 135 140
Ala Thr Thr Ser Arg Gly Pro Glu Pro Ser Lys Glu Ala Gln Arg Ile
145 150 155 160
Pro Val Pro Gly Pro Gly Ser Arg Pro Glu Pro Gln Ser Glu Phe Ala
165 170 175
Ala Thr Arg Lys Pro Arg Pro Glu Ser Gly Gly Glu Gly Leu Met Asn
180 185 190
Phe Ser Leu Phe Ser Arg Pro Ala Ala Leu Val Arg Ala Ser Leu Gln
195 200 205
Arg Pro Pro Pro Pro Gln Thr Gly Thr Asp Lys Val Ser Asn Val Thr
210 215 220
Thr Ser Thr Arg Val Glu Ser Thr Val Leu Gln Ser Ala Ser Gly Pro
225 230 235 240
Arg Ile Ala Pro Val Phe Thr Asp Gln Arg Thr Ala Trp Ser Gln Ser
245 250 255
Lys Glu Val Arg Phe Ser Cys Ala Pro Ala Leu Ala Ala Gly Asn Leu
260 265 270
His Gln Asp Met Pro Leu Gly Arg Pro Gly Asn Asn Met Thr Pro Gln
275 280 285
Gly Lys Met Glu Thr Lys Lys Ala Cys Glu Val Ala Val Ala Thr Pro
290 295 300
Ser Leu Cys Ser Gly Asn Gly Glu Ser Trp Arg Glu Gln Lys Arg Lys
305 310 315 320
Ser Gln Ala Glu Cys Ser Ala Ser Gln Asp Asp Asp Leu Asp Asp Glu
325 330 335
Ser Gly Gly Met Arg Gly Ser Gly Gly Arg Gly Thr Lys Arg Ser Arg
340 345 350
Thr Ala Glu Val His Asn Leu Ser Glu Arg Arg Arg Arg Asp Arg Ile
355 360 365
Asn Glu Lys Met Arg Ala Leu Gln Glu Leu Ile Pro Asn Cys Asn Lys
370 375 380
Ile Asp Lys Ala Ser Met Leu Asp Glu Ala Ile Glu Tyr Leu Lys Thr
385 390 395 400
Leu Gln Leu Gln Val Gln Met Met Ala Met Gly Ser Gly Leu Cys Ile
405 410 415
Pro Pro Met Leu Leu Pro Arg Ala Met Gln Leu Gln Ile Pro Ser Ile
420 425 430
Ala His Phe His His Leu Gly Met Gly Leu Gly Tyr Gly Met Gly Val
435 440 445
Leu Asp Met Asn Ser Thr Ala Ala Val Pro Phe Gln Pro Ile Pro Gly
450 455 460
Ala His Tyr Thr Cys Pro Met Ile Pro Gly Thr Pro Pro Gln Gly Leu
465 470 475 480
Gly Ile Gly Met Pro Gly Thr Asn Thr Met Pro Val Phe Gly Leu Pro
485 490 495
Gly Gln Ala Ile His Pro Ser Ala Ser Ser Val Gln Pro Phe Pro Ser
500 505 510
Leu Ala Gly Leu Pro Val Arg Pro Asn Leu Ala Pro Gln Val Pro Ala
515 520 525
Val Met Ala Asn Met Val Gln Glu Gln Gln Gln Gly Val Ala Thr Gln
530 535 540
Gln Gln Gln Ser Leu Asn Asn Glu Ala Gln Gln Lys Ala Asn Thr Gly
545 550 555 560
Asp Pro Gln Ile Leu Gln Lys
565
<210> 3
<211> 1923
<212> DNA
<213> Zea mays L.
<400> 3
atgtccgaca gcagcgactt cgccgagctg ctatgggaga acggccaggc ggtggtgcac 60
gcgcggacga ggaaacagcc gccccagcag cccgccttcc cgcccttcac cttcacctgc 120
agcagcagag ctcaacaaag gcagccggtg ccggcgctgc tcaagacggg aggaggtctc 180
tttgccgact tctcctcggc caccgccggc ctagacgccc cccgtcacaa cggcgacctc 240
gacgacaccg tgccgtggat ccactacccc atccccatcg tcgacgaagc cagtcccgcc 300
gcgcctgccc tggcagatag cttcatccca gatttcttct cggagctgca tgcggcggcg 360
gcgacgacga gcctcggccc tctgccacca ccaccaccac caccaccagc agtccagcac 420
accggcgacg acaacgacaa ccggagcacc ccggttccca atcccaccac cggcagagga 480
ccagagccct caaaggaaac ccaccgcgca ccagtaccag gtccagccgg caggcctgag 540
ccacaaccac aagctgagct cgcgcccgcc aggaagcctc ggcaggagag cggcggcggg 600
gaggctttga tgaacttctc catcttctct aggccggcag ccatggcgcg tgccagcctg 660
cggcagaggc cgccacagac aggcaccgac aaggcgtcca atgccaccac gagcacccgc 720
gtggagtcga cggtgctaca gtcggccagt gccagtggcc cgagaactgc aactgccccc 780
gccctgttcg tggaccagag gacggcgtgg ccgtcgcagc agcccaagga cgtgcggttt 840
ccttgcgcag cagcagcgcc accgccgacg ccgccgccga cggctgctaa cctgcagcaa 900
gaaagaccta gcaacaacaa catgactcct ccgcagaaag aagtagagac caggaaggct 960
tcagaggctg cgggcgccac tgccacctca tcggtctgct ccggcaatgg tgctgggaca 1020
ggaaaggacg acgagtcctg gcgccaacag aagaggaaga gcctgcaggc cgagtgttcc 1080
gcgagtcaag atgatgatcc tgacgatgag tccggtggga tgagaagatc gtgcagcaga 1140
ggtgcgaagc gcagccgcac cgcggaggtg cacaacctgt ccgaaaggag gagaagggac 1200
aggatcaatg aaaaaatgcg tgccctgcaa gagctcatcc ccaactgcaa caagattgac 1260
aaggcctcca tgctggacga agcgatcgag tacctcaaaa ccctccagct tcaagtgcag 1320
atgatgtcca tgggaagtgg gctgtgcatt cctcccatgc tgctgccacc agccatgcag 1380
cacctgcaga ttcctccggc tgctgctcac ttccctcatc tcggcatggg attgggttac 1440
gggatgggtg tcctcgatat gaacagcgcg gcggcggcgg cggcggcggt tccatttccg 1500
cacatgcccg gtgctcactt cccttgcccg atgatcccag gcgcgccact accacggggt 1560
ctcggaatcg gaatgcctgc cagaaacacg atgccggtgt tcggagtccc tcctgggcaa 1620
ccaatccagc caccattccc atctttggcg ggccttcctg tcaggccaaa cctggcacct 1680
cctcacgcct cctcagccgc catggctaac atggtgcagg agcaacagca gcagcagcag 1740
catcagagcc tgaataatgg agcatcaaca gcagcagcaa cagcatcaga tcctcagact 1800
atcgtgcagg ctgagagaca gcattttagt gtgccctctt cagcacaaac cgagagcgat 1860
cagtttttgg atggtggtgg caacagggct atttgtacag gaagaaatga ggctgaaaca 1920
tga 1923
<210> 4
<211> 640
<212> PRT
<213> Zea mays L.
<400> 4
Met Ser Asp Ser Ser Asp Phe Ala Glu Leu Leu Trp Glu Asn Gly Gln
1 5 10 15
Ala Val Val His Ala Arg Thr Arg Lys Gln Pro Pro Gln Gln Pro Ala
20 25 30
Phe Pro Pro Phe Thr Phe Thr Cys Ser Ser Arg Ala Gln Gln Arg Gln
35 40 45
Pro Val Pro Ala Leu Leu Lys Thr Gly Gly Gly Leu Phe Ala Asp Phe
50 55 60
Ser Ser Ala Thr Ala Gly Leu Asp Ala Pro Arg His Asn Gly Asp Leu
65 70 75 80
Asp Asp Thr Val Pro Trp Ile His Tyr Pro Ile Pro Ile Val Asp Glu
85 90 95
Ala Ser Pro Ala Ala Pro Ala Leu Ala Asp Ser Phe Ile Pro Asp Phe
100 105 110
Phe Ser Glu Leu His Ala Ala Ala Ala Thr Thr Ser Leu Gly Pro Leu
115 120 125
Pro Pro Pro Pro Pro Pro Pro Pro Ala Val Gln His Thr Gly Asp Asp
130 135 140
Asn Asp Asn Arg Ser Thr Pro Val Pro Asn Pro Thr Thr Gly Arg Gly
145 150 155 160
Pro Glu Pro Ser Lys Glu Thr His Arg Ala Pro Val Pro Gly Pro Ala
165 170 175
Gly Arg Pro Glu Pro Gln Pro Gln Ala Glu Leu Ala Pro Ala Arg Lys
180 185 190
Pro Arg Gln Glu Ser Gly Gly Gly Glu Ala Leu Met Asn Phe Ser Ile
195 200 205
Phe Ser Arg Pro Ala Ala Met Ala Arg Ala Ser Leu Arg Gln Arg Pro
210 215 220
Pro Gln Thr Gly Thr Asp Lys Ala Ser Asn Ala Thr Thr Ser Thr Arg
225 230 235 240
Val Glu Ser Thr Val Leu Gln Ser Ala Ser Ala Ser Gly Pro Arg Thr
245 250 255
Ala Thr Ala Pro Ala Leu Phe Val Asp Gln Arg Thr Ala Trp Pro Ser
260 265 270
Gln Gln Pro Lys Asp Val Arg Phe Pro Cys Ala Ala Ala Ala Pro Pro
275 280 285
Pro Thr Pro Pro Pro Thr Ala Ala Asn Leu Gln Gln Glu Arg Pro Ser
290 295 300
Asn Asn Asn Met Thr Pro Pro Gln Lys Glu Val Glu Thr Arg Lys Ala
305 310 315 320
Ser Glu Ala Ala Gly Ala Thr Ala Thr Ser Ser Val Cys Ser Gly Asn
325 330 335
Gly Ala Gly Thr Gly Lys Asp Asp Glu Ser Trp Arg Gln Gln Lys Arg
340 345 350
Lys Ser Leu Gln Ala Glu Cys Ser Ala Ser Gln Asp Asp Asp Pro Asp
355 360 365
Asp Glu Ser Gly Gly Met Arg Arg Ser Cys Ser Arg Gly Ala Lys Arg
370 375 380
Ser Arg Thr Ala Glu Val His Asn Leu Ser Glu Arg Arg Arg Arg Asp
385 390 395 400
Arg Ile Asn Glu Lys Met Arg Ala Leu Gln Glu Leu Ile Pro Asn Cys
405 410 415
Asn Lys Ile Asp Lys Ala Ser Met Leu Asp Glu Ala Ile Glu Tyr Leu
420 425 430
Lys Thr Leu Gln Leu Gln Val Gln Met Met Ser Met Gly Ser Gly Leu
435 440 445
Cys Ile Pro Pro Met Leu Leu Pro Pro Ala Met Gln His Leu Gln Ile
450 455 460
Pro Pro Ala Ala Ala His Phe Pro His Leu Gly Met Gly Leu Gly Tyr
465 470 475 480
Gly Met Gly Val Leu Asp Met Asn Ser Ala Ala Ala Ala Ala Ala Ala
485 490 495
Val Pro Phe Pro His Met Pro Gly Ala His Phe Pro Cys Pro Met Ile
500 505 510
Pro Gly Ala Pro Leu Pro Arg Gly Leu Gly Ile Gly Met Pro Ala Arg
515 520 525
Asn Thr Met Pro Val Phe Gly Val Pro Pro Gly Gln Pro Ile Gln Pro
530 535 540
Pro Phe Pro Ser Leu Ala Gly Leu Pro Val Arg Pro Asn Leu Ala Pro
545 550 555 560
Pro His Ala Ser Ser Ala Ala Met Ala Asn Met Val Gln Glu Gln Gln
565 570 575
Gln Gln Gln Gln His Gln Ser Leu Asn Asn Gly Ala Ser Thr Ala Ala
580 585 590
Ala Thr Ala Ser Asp Pro Gln Thr Ile Val Gln Ala Glu Arg Gln His
595 600 605
Phe Ser Val Pro Ser Ser Ala Gln Thr Glu Ser Asp Gln Phe Leu Asp
610 615 620
Gly Gly Gly Asn Arg Ala Ile Cys Thr Gly Arg Asn Glu Ala Glu Thr
625 630 635 640
<210> 5
<211> 1518
<212> DNA
<213> Zea mays L.
<400> 5
atgcccgttt cgatttccat ctgtaggact gggccgggcg aggagctggc cgagctgctg 60
tgggaccgtg gcccggcgct gcggagggcg ccgccgccgt tccagccctt cacctgcagc 120
gccgccggca gcagcaggtc gcaggagctg aagcggcatg cctcggacac caccaaggcg 180
tcagcgttcg tgaccgccgt ctccgtcccg ctgggcacgc acgacgccgg ctccgggctc 240
gggctcgccg gcctccccgt ccacgacgac gacgacgccg tgccgtggtt gcattgcccc 300
gtcgccgacg acggcgacgg cgacacggcg ccgctgccgc cggagttctg cgccggcctc 360
ctgtccgagt actcggaggt ggccgcgccg gcgccggcct tccacgccgc ggccacgccg 420
ccggccgagg ccgcggccaa caagctggcc ccgccgagcg ccgccggcgg aggggagggc 480
gtcttgaact tcaccttctt ctcgcggccc ctccagcgac cgcaggcggc ggcggcgccc 540
gccgccgcgg ccgcgagcaa ccccgtcgag tccacggtcg tccaggcggc agcgaaccgg 600
ctgcggagca cgccgctgtt ctccgagcag aggatggcgt ggctgcagcc acccaaggcg 660
ccgcgcacca cagcggcagc ggcggcgcca cctcctcctc cgctggcgcc tctgctccca 720
gatagccgcc atggggagac ggtcggcacg gtagctcaac ctcagccccg gtcgcaacca 780
gaagcaagac ctccggatgc ggcggcggtg acgacctctt cggtctgctc cggcaacggt 840
ggtcggagcc agctcaagag gagccgccac ctggccgcgg actgctcggt cagtccggac 900
gaggacctgg acgacgagcc cggcgcgacg aggaggtcgg cggcgcggag cgccaagcgc 960
tgccgcaccg ccgaggtgca caacctgtcg gagaggagga gacgggaccg gatcaacgag 1020
aagatgcgcg ccctgcagga gctcattccc aactgcaaca aggtcgacaa gtcgtcgatg 1080
ctggaggagg cgatcgagta cctgaagacg ctgcagctgc aagtgcagat gatgtcgatg 1140
gggaccgggc tgtgcatgcc accggcggcg atgctgctgc cagcgatgca gcagcagctc 1200
ctgcaccacc accccatggc gcacttcccc catctcggca tgggcctggg cttcggcatg 1260
ggcgcggcgg cggggttcga catgctcccg ttcccgtgcg tcgcggccgg cgcccacttc 1320
ccgtgcccgc cgggggccat gttcggcgtg ccggggcagg cgatgccctc gctgccggcg 1380
gcgttcgctc acatgtacgg cgctggcagt ggcgccgggc cggctgggca gacggaagct 1440
gctgatgcgg ctgctcctgc acggccagga gaggcagagc agggtgatca gcaggtgcag 1500
cacgcgaagc agacgtga 1518
<210> 6
<211> 505
<212> PRT
<213> Zea mays L.
<400> 6
Met Pro Val Ser Ile Ser Ile Cys Arg Thr Gly Pro Gly Glu Glu Leu
1 5 10 15
Ala Glu Leu Leu Trp Asp Arg Gly Pro Ala Leu Arg Arg Ala Pro Pro
20 25 30
Pro Phe Gln Pro Phe Thr Cys Ser Ala Ala Gly Ser Ser Arg Ser Gln
35 40 45
Glu Leu Lys Arg His Ala Ser Asp Thr Thr Lys Ala Ser Ala Phe Val
50 55 60
Thr Ala Val Ser Val Pro Leu Gly Thr His Asp Ala Gly Ser Gly Leu
65 70 75 80
Gly Leu Ala Gly Leu Pro Val His Asp Asp Asp Asp Ala Val Pro Trp
85 90 95
Leu His Cys Pro Val Ala Asp Asp Gly Asp Gly Asp Thr Ala Pro Leu
100 105 110
Pro Pro Glu Phe Cys Ala Gly Leu Leu Ser Glu Tyr Ser Glu Val Ala
115 120 125
Ala Pro Ala Pro Ala Phe His Ala Ala Ala Thr Pro Pro Ala Glu Ala
130 135 140
Ala Ala Asn Lys Leu Ala Pro Pro Ser Ala Ala Gly Gly Gly Glu Gly
145 150 155 160
Val Leu Asn Phe Thr Phe Phe Ser Arg Pro Leu Gln Arg Pro Gln Ala
165 170 175
Ala Ala Ala Pro Ala Ala Ala Ala Ala Ser Asn Pro Val Glu Ser Thr
180 185 190
Val Val Gln Ala Ala Ala Asn Arg Leu Arg Ser Thr Pro Leu Phe Ser
195 200 205
Glu Gln Arg Met Ala Trp Leu Gln Pro Pro Lys Ala Pro Arg Thr Thr
210 215 220
Ala Ala Ala Ala Ala Pro Pro Pro Pro Pro Leu Ala Pro Leu Leu Pro
225 230 235 240
Asp Ser Arg His Gly Glu Thr Val Gly Thr Val Ala Gln Pro Gln Pro
245 250 255
Arg Ser Gln Pro Glu Ala Arg Pro Pro Asp Ala Ala Ala Val Thr Thr
260 265 270
Ser Ser Val Cys Ser Gly Asn Gly Gly Arg Ser Gln Leu Lys Arg Ser
275 280 285
Arg His Leu Ala Ala Asp Cys Ser Val Ser Pro Asp Glu Asp Leu Asp
290 295 300
Asp Glu Pro Gly Ala Thr Arg Arg Ser Ala Ala Arg Ser Ala Lys Arg
305 310 315 320
Cys Arg Thr Ala Glu Val His Asn Leu Ser Glu Arg Arg Arg Arg Asp
325 330 335
Arg Ile Asn Glu Lys Met Arg Ala Leu Gln Glu Leu Ile Pro Asn Cys
340 345 350
Asn Lys Val Asp Lys Ser Ser Met Leu Glu Glu Ala Ile Glu Tyr Leu
355 360 365
Lys Thr Leu Gln Leu Gln Val Gln Met Met Ser Met Gly Thr Gly Leu
370 375 380
Cys Met Pro Pro Ala Ala Met Leu Leu Pro Ala Met Gln Gln Gln Leu
385 390 395 400
Leu His His His Pro Met Ala His Phe Pro His Leu Gly Met Gly Leu
405 410 415
Gly Phe Gly Met Gly Ala Ala Ala Gly Phe Asp Met Leu Pro Phe Pro
420 425 430
Cys Val Ala Ala Gly Ala His Phe Pro Cys Pro Pro Gly Ala Met Phe
435 440 445
Gly Val Pro Gly Gln Ala Met Pro Ser Leu Pro Ala Ala Phe Ala His
450 455 460
Met Tyr Gly Ala Gly Ser Gly Ala Gly Pro Ala Gly Gln Thr Glu Ala
465 470 475 480
Ala Asp Ala Ala Ala Pro Ala Arg Pro Gly Glu Ala Glu Gln Gly Asp
485 490 495
Gln Gln Val Gln His Ala Lys Gln Thr
500 505
<210> 7
<211> 20
<212> DNA
<213> Artifical sequence
<400> 7
gtcgagctgc tatgggagaa 20
<210> 8
<211> 20
<212> DNA
<213> Artifical sequence
<400> 8
ggggaaaatg gagaccaaga 20
<210> 9
<211> 20
<212> DNA
<213> Artifical sequence
<400> 9
gcagcagcag agctcaacaa 20
<210> 10
<211> 20
<212> DNA
<213> Artifical sequence
<400> 10
gaaagaagta gagaccagga 20
<210> 11
<211> 20
<212> DNA
<213> Artifical sequence
<400> 11
gctgttctcc gagcagagga 20
<210> 12
<211> 20
<212> DNA
<213> Artifical sequence
<400> 12
gcaagtgcag atgatgtcga 20
<210> 13
<211> 20
<212> DNA
<213> Artifical sequence
<400> 13
gtccatttcc atccaggtcc 20
<210> 14
<211> 20
<212> DNA
<213> Artifical sequence
<400> 14
ccaccggact catcatcaag 20
<210> 15
<211> 20
<212> DNA
<213> Artifical sequence
<400> 15
gtgagcggtt ccttctcatt 20
<210> 16
<211> 20
<212> DNA
<213> Artifical sequence
<400> 16
ctcctctgca ggtttgttgt 20
<210> 17
<211> 20
<212> DNA
<213> Artifical sequence
<400> 17
aggggagggc gtcttgaact 20
<210> 18
<211> 20
<212> DNA
<213> Artifical sequence
<400> 18
ggaacgggag catgtcgaac 20
<210> 19
<211> 64
<212> DNA
<213> Artifical sequence
<400> 19
gagccgcaag caccgaattg tcgagctgct atgggagaag ttttagagct agaaatagca 60
agtt 64
<210> 20
<211> 64
<212> DNA
<213> Artifical sequence
<400> 20
gagccgcaag caccgaattg gggaaaatgg agaccaagag ttttagagct agaaatagca 60
agtt 64
<210> 21
<211> 64
<212> DNA
<213> Artifical sequence
<400> 21
gagccgcaag caccgaattg cagcagcaga gctcaacaag ttttagagct agaaatagca 60
agtt 64
<210> 22
<211> 64
<212> DNA
<213> Artifical sequence
<400> 22
gagccgcaag caccgaattg aaagaagtag agaccaggag ttttagagct agaaatagca 60
agtt 64
<210> 23
<211> 64
<212> DNA
<213> Artifical sequence
<400> 23
gagccgcaag caccgaattg ctgttctccg agcagaggag ttttagagct agaaatagca 60
agtt 64
<210> 24
<211> 64
<212> DNA
<213> Artifical sequence
<400> 24
gagccgcaag caccgaattg caagtgcaga tgatgtcgag ttttagagct agaaatagca 60
agtt 64
<210> 25
<211> 37
<212> DNA
<213> Artifical sequence
<400> 25
tgcttttttt aagctgctgt ttttgttagc cccatcg 37
<210> 26
<211> 19
<212> DNA
<213> Artifical sequence
<400> 26
aattcggtgc ttgcggctc 19
<210> 27
<211> 34
<212> DNA
<213> Artifical sequence
<400> 27
ggccagtgcc aagcttaaaa aaagcaccga ctcg 34
<210> 28
<211> 83
<212> DNA
<213> Artifical sequence
<400> 28
gttttagagc tagaaatagc aagttaaaat aaggctagtc cgttatcaac ttgaaaaagt 60
ggcaccgagt cggtgctttt ttt 83
<210> 29
<211> 37
<212> DNA
<213> Artifical sequence
<400> 29
tgcactgcac aagctgctgt ttttgttagc cccatcg 37
<210> 30
<211> 37
<212> DNA
<213> Artifical sequence
<400> 30
tgcttttttt aagctgctgt ttttgttagc cccatcg 37
<210> 31
<211> 26
<212> DNA
<213> Artifical sequence
<400> 31
cttagacatg caatgctcat tatctc 26
<210> 32
<211> 18
<212> DNA
<213> Artifical sequence
<400> 32
ctggcgaaag ggggatgt 18
<210> 33
<211> 51
<212> DNA
<213> Artifical sequence
<400> 33
atgtccgaca gcaacgactt cgtcgagctg ctatgggaag aaggcttgtg a 51
<210> 34
<211> 16
<212> PRT
<213> Artifical sequence
<400> 34
Met Ser Asp Ser Asn Asp Phe Val Glu Leu Leu Trp Glu Glu Gly Leu
1 5 10 15
<210> 35
<211> 87
<212> DNA
<213> Artifical sequence
<400> 35
atgtccgaca gcaacgactt cgtcgagctg ctatgggacc tgcaccacga gcacccgcgt 60
ggagtcgacg gttctacagt cggctag 87
<210> 36
<211> 28
<212> PRT
<213> Artifical sequence
<400> 36
Met Ser Asp Ser Asn Asp Phe Val Glu Leu Leu Trp Asp Leu His His
1 5 10 15
Glu His Pro Arg Gly Val Asp Gly Ser Thr Val Gly
20 25
<210> 37
<211> 309
<212> DNA
<213> Artifical sequence
<400> 37
atgtccgaca gcagcgactt cgccgagctg ctatgggaga acggccaggc ggtggtgcac 60
gcgcggacga ggaaacagcc gccccagcag cccgccttcc cgcccttcac cttcacctgc 120
agcagcagag ctcaacaagg aaggcttcag aggctgcggg cgccactgcc acctcatcgg 180
tctgctccgg caatggtgct gggacaggaa aggacgacga gtcctggcgc caacagaaga 240
ggaagagcct gcaggccgag tgttccgcga gtcaagatga tgatcctgac gatgagtccg 300
gtgggatga 309
<210> 38
<211> 102
<212> PRT
<213> Artifical sequence
<400> 38
Met Ser Asp Ser Ser Asp Phe Ala Glu Leu Leu Trp Glu Asn Gly Gln
1 5 10 15
Ala Val Val His Ala Arg Thr Arg Lys Gln Pro Pro Gln Gln Pro Ala
20 25 30
Phe Pro Pro Phe Thr Phe Thr Cys Ser Ser Arg Ala Gln Gln Gly Arg
35 40 45
Leu Gln Arg Leu Arg Ala Pro Leu Pro Pro His Arg Ser Ala Pro Ala
50 55 60
Met Val Leu Gly Gln Glu Arg Thr Thr Ser Pro Gly Ala Asn Arg Arg
65 70 75 80
Gly Arg Ala Cys Arg Pro Ser Val Pro Arg Val Lys Met Met Ile Leu
85 90 95
Thr Met Ser Pro Val Gly
100
<210> 39
<211> 276
<212> DNA
<213> Artifical sequence
<400> 39
ataggcggtg gtgcacgcgc ggacgaggaa acagccgccc cagcagcccg ccttcccgcc 60
cttcaccttc acctgcagca gcagagctca acaaaggcag ccggtgccgg cgctgctcaa 120
gacgggagga ggtctctttg ccgacttctc ctcggccacc gccggcctag acgccccccg 180
tcacaacggc gacctcgacg acaccgtgcc gtggatccac taccccatcc ccatcgtcga 240
cgaagccagt cccgccgcgc ctgccctggc agatag 276
<210> 40
<211> 91
<212> PRT
<213> Artifical sequence
<400> 40
Ile Gly Gly Gly Ala Arg Ala Asp Glu Glu Thr Ala Ala Pro Ala Ala
1 5 10 15
Arg Leu Pro Ala Leu His Leu His Leu Gln Gln Gln Ser Ser Thr Lys
20 25 30
Ala Ala Gly Ala Gly Ala Ala Gln Asp Gly Arg Arg Ser Leu Cys Arg
35 40 45
Leu Leu Leu Gly His Arg Arg Pro Arg Arg Pro Pro Ser Gln Arg Arg
50 55 60
Pro Arg Arg His Arg Ala Val Asp Pro Leu Pro His Pro His Arg Arg
65 70 75 80
Arg Ser Gln Ser Arg Arg Ala Cys Pro Gly Arg
85 90
<210> 41
<211> 1005
<212> DNA
<213> Artifical sequence
<400> 41
atgcccgttt cgatttccat ctgtaggact gggccgggcg aggagctggc cgagctgctg 60
tgggaccgtg gcccggcgct gcggagggcg ccgccgccgt tccagccctt cacctgcagc 120
gccgccggca gcagcaggtc gcaggagctg aagcggcatg cctcggacac caccaaggcg 180
tcagcgttcg tgaccgccgt ctccgtcccg ctgggcacgc acgacgccgg ctccgggctc 240
gggctcgccg gcctccccgt ccacgacgac gacgacgccg tgccgtggtt gcattgcccc 300
gtcgccgacg acggcgacgg cgacacggcg ccgctgccgc cggagttctg cgccggcctc 360
ctgtccgagt actcggaggt ggccgcgccg gcgccggcct tccacgccgc ggccacgccg 420
ccggccgagg ccgcggccaa caagctggcc ccgccgagcg ccgccggcgg aggggagggc 480
gtcttgaact tcaccttctt ctcgcggccc ctccagcgac cgcaggcggc ggcggcgccc 540
gccgccgcgg ccgcgagcaa ccccgtcgag tccacggtcg tccaggcggc agcgaaccgg 600
ctgcggagca cgccgctgtt ctccgagcag aacatcatct gcacttgcag ctgcagcgtc 660
ttcaggtact cgatcgcctc ctccagcatc gacgacttgt cgaccttgtt gcagttggga 720
atgagctcct gcagggcgcg catcttctcg ttgatccggt cccgtctcct cctctccgac 780
aggttgtgca cctcggcggt gcggcagcgc ttggcgctcc gcgccgccga cctcctcgtc 840
gcgccgggct cgtcgtccag gtcctcgtcc ggactgaccg agcagtccgc ggccaggtgg 900
cggctcctct tgagctggct ccgaccaccg ttgccggagc agaccgaaga ggtcgtcacc 960
gccgccgcat ccggaggtct tgcttctggt tgcgaccggg gctga 1005
<210> 42
<211> 334
<212> PRT
<213> Artifical sequence
<400> 42
Met Pro Val Ser Ile Ser Ile Cys Arg Thr Gly Pro Gly Glu Glu Leu
1 5 10 15
Ala Glu Leu Leu Trp Asp Arg Gly Pro Ala Leu Arg Arg Ala Pro Pro
20 25 30
Pro Phe Gln Pro Phe Thr Cys Ser Ala Ala Gly Ser Ser Arg Ser Gln
35 40 45
Glu Leu Lys Arg His Ala Ser Asp Thr Thr Lys Ala Ser Ala Phe Val
50 55 60
Thr Ala Val Ser Val Pro Leu Gly Thr His Asp Ala Gly Ser Gly Leu
65 70 75 80
Gly Leu Ala Gly Leu Pro Val His Asp Asp Asp Asp Ala Val Pro Trp
85 90 95
Leu His Cys Pro Val Ala Asp Asp Gly Asp Gly Asp Thr Ala Pro Leu
100 105 110
Pro Pro Glu Phe Cys Ala Gly Leu Leu Ser Glu Tyr Ser Glu Val Ala
115 120 125
Ala Pro Ala Pro Ala Phe His Ala Ala Ala Thr Pro Pro Ala Glu Ala
130 135 140
Ala Ala Asn Lys Leu Ala Pro Pro Ser Ala Ala Gly Gly Gly Glu Gly
145 150 155 160
Val Leu Asn Phe Thr Phe Phe Ser Arg Pro Leu Gln Arg Pro Gln Ala
165 170 175
Ala Ala Ala Pro Ala Ala Ala Ala Ala Ser Asn Pro Val Glu Ser Thr
180 185 190
Val Val Gln Ala Ala Ala Asn Arg Leu Arg Ser Thr Pro Leu Phe Ser
195 200 205
Glu Gln Asn Ile Ile Cys Thr Cys Ser Cys Ser Val Phe Arg Tyr Ser
210 215 220
Ile Ala Ser Ser Ser Ile Asp Asp Leu Ser Thr Leu Leu Gln Leu Gly
225 230 235 240
Met Ser Ser Cys Arg Ala Arg Ile Phe Ser Leu Ile Arg Ser Arg Leu
245 250 255
Leu Leu Ser Asp Arg Leu Cys Thr Ser Ala Val Arg Gln Arg Leu Ala
260 265 270
Leu Arg Ala Ala Asp Leu Leu Val Ala Pro Gly Ser Ser Ser Arg Ser
275 280 285
Ser Ser Gly Leu Thr Glu Gln Ser Ala Ala Arg Trp Arg Leu Leu Leu
290 295 300
Ser Trp Leu Arg Pro Pro Leu Pro Glu Gln Thr Glu Glu Val Val Thr
305 310 315 320
Ala Ala Ala Ser Gly Gly Leu Ala Ser Gly Cys Asp Arg Gly
325 330
<210> 43
<211> 1089
<212> DNA
<213> Artifical sequence
<400> 43
atgcccgttt cgatttccat ctgtaggact gggccgggcg aggagctggc cgagctgctg 60
tgggaccgtg gcccggcgct gcggagggcg ccgccgccgt tccagccctt cacctgcagc 120
gccgccggca gcagcaggtc gcaggagctg aagcggcatg cctcggacac caccaaggcg 180
tcagcgttcg tgaccgccgt ctccgtcccg ctgggcacgc acgacgccgg ctccgggctc 240
gggctcgccg gcctccccgt ccacgacgac gacgacgccg tgccgtggtt gcattgcccc 300
gtcgccgacg acggcgacgg cgacacggcg ccgctgccgc cggagttctg cgccggcctc 360
ctgtccgagt actcggaggt ggccgcgccg gcgccggcct tccacgccgc ggccacgccg 420
ccggccgagg ccgcggccaa caagctggcc ccgccgagcg ccgccggcgg aggggagggc 480
gtcttgaact tcaccttctt ctcgcggccc ctccagcgac cgcaggcggc ggcggcgccc 540
gccgccgcgg ccgcgagcaa ccccgtcgag tccacggtcg tccaggcggc agcgaaccgg 600
ctgcggagca cgccgctgtt ctccgagtcg gcatgggcct gggcttcggc atgggcgcgg 660
cggcggggtt cgacatgctc ccgttcccgt gcgtcgcggc cggcgcccac ttcccgtgcc 720
cgccgggggc catgttcggc gtgccggggc aggcgatgcc ctcgctgccg gcggcgttcg 780
ctcacatgta cggcgctggc agtggcgccg ggccggctgg gcagacggaa gctgctgatg 840
cggctgctcc tgcacggcca ggagaggcag agcagggtga tcagcaggtg cagcacgcga 900
agcagacgtg atggcgaagg cgacgaggcg gctgcccccc gaacattccc gtttggtttt 960
atggtgatta tgagaatcta cccatcaaaa tctgctaaga gagagccaaa caaacaggat 1020
attacacctt ggattctgtg cattatctca caatcaaggt tcaagttgat tttcttggtt 1080
aatgtgtag 1089
<210> 44
<211> 362
<212> PRT
<213> Artifical sequence
<400> 44
Met Pro Val Ser Ile Ser Ile Cys Arg Thr Gly Pro Gly Glu Glu Leu
1 5 10 15
Ala Glu Leu Leu Trp Asp Arg Gly Pro Ala Leu Arg Arg Ala Pro Pro
20 25 30
Pro Phe Gln Pro Phe Thr Cys Ser Ala Ala Gly Ser Ser Arg Ser Gln
35 40 45
Glu Leu Lys Arg His Ala Ser Asp Thr Thr Lys Ala Ser Ala Phe Val
50 55 60
Thr Ala Val Ser Val Pro Leu Gly Thr His Asp Ala Gly Ser Gly Leu
65 70 75 80
Gly Leu Ala Gly Leu Pro Val His Asp Asp Asp Asp Ala Val Pro Trp
85 90 95
Leu His Cys Pro Val Ala Asp Asp Gly Asp Gly Asp Thr Ala Pro Leu
100 105 110
Pro Pro Glu Phe Cys Ala Gly Leu Leu Ser Glu Tyr Ser Glu Val Ala
115 120 125
Ala Pro Ala Pro Ala Phe His Ala Ala Ala Thr Pro Pro Ala Glu Ala
130 135 140
Ala Ala Asn Lys Leu Ala Pro Pro Ser Ala Ala Gly Gly Gly Glu Gly
145 150 155 160
Val Leu Asn Phe Thr Phe Phe Ser Arg Pro Leu Gln Arg Pro Gln Ala
165 170 175
Ala Ala Ala Pro Ala Ala Ala Ala Ala Ser Asn Pro Val Glu Ser Thr
180 185 190
Val Val Gln Ala Ala Ala Asn Arg Leu Arg Ser Thr Pro Leu Phe Ser
195 200 205
Glu Ser Ala Trp Ala Trp Ala Ser Ala Trp Ala Arg Arg Arg Gly Ser
210 215 220
Thr Cys Ser Arg Ser Arg Ala Ser Arg Pro Ala Pro Thr Ser Arg Ala
225 230 235 240
Arg Arg Gly Pro Cys Ser Ala Cys Arg Gly Arg Arg Cys Pro Arg Cys
245 250 255
Arg Arg Arg Ser Leu Thr Cys Thr Ala Leu Ala Val Ala Pro Gly Arg
260 265 270
Leu Gly Arg Arg Lys Leu Leu Met Arg Leu Leu Leu His Gly Gln Glu
275 280 285
Arg Gln Ser Arg Val Ile Ser Arg Cys Ser Thr Arg Ser Arg Arg Asp
290 295 300
Gly Glu Gly Asp Glu Ala Ala Ala Pro Arg Thr Phe Pro Phe Gly Phe
305 310 315 320
Met Val Ile Met Arg Ile Tyr Pro Ser Lys Ser Ala Lys Arg Glu Pro
325 330 335
Asn Lys Gln Asp Ile Thr Pro Trp Ile Leu Cys Ile Ile Ser Gln Ser
340 345 350
Arg Phe Lys Leu Ile Phe Leu Val Asn Val
355 360
Claims (3)
1. 玉米ZmPIF3s突变型蛋白或其编码基因在培育矮杆植物新品种或短中胚轴植物新品种中的应用;所述玉米ZmPIF3s突变型蛋白由ZmPIF3.1突变型蛋白、ZmPIF3.2突变型蛋白以及ZmPIF3.3突变型蛋白组成;其中,所述ZmPIF3.1突变型蛋白的氨基酸序列为SEQ IDNo.34或SEQ ID No.36所示;所述ZmPIF3.2突变型蛋白的氨基酸序列为SEQ ID No.38或SEQID No.40所示;所述ZmPIF3.3突变型蛋白的氨基酸序列为SEQ ID No.42或SEQ ID No.44所示;
SEQ ID No.34或SEQ ID No.36所示的ZmPIF3.1突变型蛋白的编码基因的核苷酸序列为SEQ ID No.33或SEQ ID No.35所示; SEQ ID No.38或SEQ ID No.40所示ZmPIF3.2突变型蛋白的编码基因的核苷酸序列为SEQ ID No.37或SEQ ID No.39所示; SEQ ID No.42或SEQ ID No.44所示的ZmPIF3.3突变型蛋白的编码基因的核苷酸序列为SEQ ID No.41或SEQID No.43所示。
2.根据权利要求1所述的应用,其特征在于,包括构建ZmPIF3s基因编辑载体,所述ZmPIF3s基因编辑载体的构建方法包括:
(Ⅰ)、获取玉米U6-1启动子片段;
(Ⅱ)、制备6个sgRNA表达盒;
(Ⅲ)、将步骤(Ⅱ)制备的6个sgRNA表达盒依次连接CPB-Ubi-hspcas9载体,即得;
步骤(Ⅱ)中所述的制备6个sgRNA表达盒包括以下步骤:
(一)、采用重叠PCR的方法将带有接头的靶点序列和sgR骨架序列融合在一起获得6个PCR产物;其中,PCR的上游引物分别为SEQ ID No.19- SEQ ID No.24所示;下游引物为SEQID No.27所示,模板序列为SEQ ID No.28所示;
(二)、再次用重叠PCR的方法将步骤(一)获得的6个融合PCR片段和U6-1启动子片段分别连接上获得的6个PCR产物得到sgRNA连接产物;
其中,该重叠PCR中所用到的上游、下游引物序列为SEQ ID No.29和SEQ ID No.30所示;所用到的模板是U6-1启动子片段以及步骤(一)扩增得到的6个PCR产物。
3.根据权利要求2所述的应用,其特征在于,步骤(一)中的重叠PCR反应程序如下:(1)94℃,2 min;(2)98℃,10 s;46℃,30 s;68℃,10 s;共35个循环;(3)68℃,5 min;步骤(二)中的重叠PCR反应程序如下:(1)94 ℃,2 min;(2)98℃,10 s;52-55℃,30 s;68℃,40 s;共35个循环;(3)68℃,5 min。
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