CN113980919A - 调控玉米穗腐病抗性的dna序列及其突变体、分子标记和应用 - Google Patents
调控玉米穗腐病抗性的dna序列及其突变体、分子标记和应用 Download PDFInfo
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Abstract
本发明公开了调控玉米穗腐病抗性的DNA序列及其突变体、分子标记和应用。本发明首先提供了碱基序列为SEQ ID No.1、SEQ ID No.2和SEQ ID No.14所示的调控玉米穗腐病抗性的关键DNA序列及其突变体，其能够调控ZmLOX3基因在玉米果穗和籽粒中的表达，可改良玉米穗腐病抗性或培育玉米抗穗腐病新品种。本发明提供了Indel151或Indel363作为ZmLOX3基因在玉米果穗和籽粒中表达的分子标记、调控ZmLOX3基因在玉米表达量以及在改良玉米抗穗腐抗性中的用途。本发明进一步提供了检测所述DNA序列及其突变体变异情况的检测引物以及检测玉米中ZmLOX3基因表达量的检测引物。
Description
技术领域
本发明涉及调控玉米穗腐病抗性的DNA序列及其突变体，尤其涉及调控ZmLOX3基因表达的启动子区和内含子区的关键DNA序列及其突变体、ZmLOX3基因表达的分子标记和检测引物，本发明进一步涉及它们在调控玉米穗腐病抗性中的应用，属于玉米穗腐病抗性调控的DNA序列、其突变体及其应用领域。
背景技术
玉米是重要的粮食、饲料和工业原材料，也是当前中国的第一大农作物。玉米的健康安全生产对保障我国的粮食安全和农业发展有着举足轻重的作用。
玉米穗腐病(ear rot)，也叫穗粒腐病(kernel and ear rot)，是世界范围内玉米产区普遍发生的一类真菌病害。迄今已鉴定出40余种引起玉米穗腐病的病原真菌，其中常见的致病菌主要有以下几种：轮状镰孢F.verticillioides，拟轮枝镰孢F.moniliforme，禾谷镰孢F.graminearum，尖镰孢F.oxysporum，层出镰孢F.proliferatum，黄色镰孢F.culmorum，燕麦镰孢F.avenaceum，木贼镰孢F.equiseti，半裸镰孢F.semitectum，黄曲霉A.flavus，赭曲霉A.ochraceus等。一般，各地的病原菌组成差异较大，并且不同病院真菌可以单独或复合侵染引起玉米穗粒腐病的发生。
玉米穗腐病的发病率通常与材料的遗传背景关系比较紧密，也受不同环境和气候条件的影响，高温高湿的环境容易加重玉米穗粒腐病的发生。
穗腐病的发生可引起籽粒腐烂，造成产量损失高达30％-40％。除直接造成产量损失外，穗腐病的致病镰孢、曲霉等病原真菌能产生多种毒素(如镰孢菌能够产生单端孢霉烯、伏马毒素和玉米赤霉烯酮等真菌毒素，曲霉产生的黄曲霉毒素以及赭曲霉毒素等)，还影响玉米安全贮存和食用安全。真菌毒素不但能够影响真菌对宿主的侵染和破坏程度，而且对包括人类和畜禽在内的脊椎动物具有严重的毒害作用。不同类型的毒素，对寄主植物和人畜造成的伤害程度和引起的症状各异。伏马毒素是一类鞘脂类化合物，在动植物细胞中，竞争性抑制鞘脂的生物合成，从而造成鞘胺碱在细胞的积累，导致寄主组织坏死，可引起猪肺水肿、马的脑白质软化症、大鼠肝癌以及人类食道癌等；雪腐镰刀菌烯醇和脱氧雪腐镰刀菌烯醇具有结合真核生物60S核糖体亚基的能力，导致蛋白质合成的抑制和诱导凋亡的作用，产生萎蔫、褪绿或坏死的症状，引起人畜呕吐、腹泻、拒食或***中毒等症状；黄曲霉毒素是重要的致癌物质，其可抑制寄主植物和人畜体内白质的合成，给动植物组织造成严重损害。整体上，玉米穗粒腐病不仅导致玉米产量和品质的降低，也给其食品与饲料的安全带来重大隐患并直接威胁到人畜健康，因此，控制玉米穗腐病的发生和危害具有重要意义。
挖掘调控玉米穗腐病的抗性基因，是提高玉米穗腐病抗性、培育抗病品种，保证玉米安全生产的关键措施。前人研究发现，脂氧合酶基因(ZmLOXs)是宿主与病原真菌间相互作用的重要信号交流分子，源自宿主玉米中的脂氧合酶基因在调控玉米穗腐病的抗性方面发挥着重要作用。其中，玉米9-LOX基因ZmLOX3可能作为致病因子被轮枝镰孢菌(F.verticillioides)利用从而引起茎/穗腐病抗性。完全敲除ZmLOX3基因会增加玉米对镰孢菌引起的粒腐病的抗性，但对黄曲霉引起的粒腐病表现出感病；表明了ZmLOX3基因在玉米穗腐病抗性调控方面的复杂性，并且也表明直接的敲除该基因并不能直接用于玉米穗腐病的抗性改良。
鉴于ZmLOX3基因在镰孢菌和黄曲霉抗性调控方面的拮抗作用，一种保证该基因的蛋白功能，但适当的降低其表达量的策略，可能能达到抗镰孢菌同时兼抗黄曲霉的作用，从而实现综合提高玉米穗腐病抗性。玉米在长达近一万年的进化过程中，积累了丰富的自然变异，自然界中很可能存在这样一种不改变ZmLOX3蛋白功能，但能适当降低其表达丰度的自然变异。该类自然变异的挖掘对玉米穗腐病抗性改良至关重要，并将在玉米育种中具有广阔的应用前景。
发明内容
本发明的目的之一是提供一种调控玉米穗腐病的关键DNA序列；
本发明的目的之二是提供所述调控玉米穗腐病的关键DNA序列的突变体；
本发明的目的之三是提供用作调控ZmLOX3基因在玉米果穗和籽粒中表达的分子标记；
本发明的目的之四提供降低玉米发生穗腐病风险或提高穗腐病抗性的方法或者培育抗穗腐病的玉米新品种的方法；
本发明的目的之五是提供一种检测所述分子标记的变异情况的检测引物；
本发明的目的之六是提供检测玉米中ZmLOX3基因表达量的检测引物；
本发明的上述目的是通过以下技术方案来实现的：
本发明一方面提供了一种调控玉米穗腐病抗性的关键DNA序列，其多核苷酸为(a)、(b)、(c)或(d)所示：
(a)SEQ ID No.1所示的多核苷酸序列，或
(b)与SEQ ID No.1的互补序列在严谨杂交条件能够进行杂交的多核苷酸序列；
(c)与SEQ ID No.1所示的多核苷酸至少有90％或以上同源性的多核苷酸序列；或
(d)在SEQ ID No.1所示的多核苷酸序列的基础上进行一个或多个碱基的缺失、取代或***的突变体，且该突变体仍具有调控玉米穗腐病的功能或活性。
本发明发现，上述的关键DNA序列能够调控ZmLOX3基因在玉米果穗和籽粒中的表达量变化进而影响玉米对于穗腐病或穗粒腐抗性的高低。
本发明中所述的“穗腐病”或“穗粒腐病”为同一种病害。
在本发明的上下文中，术语“突变体”是含有变化的DNA序列，在所述DNA序列中，优选在基本保持DNA序列的同时缺失、添加和/或替代原始序列的一个或多个核苷酸。例如，可以从DNA序列5’或3’末端缺失一个或多个碱基对以产生“截短的”DNA序列；也可以在DNA序列内部***、缺失或替代一个或多个碱基对。可以通过例如标准DNA诱变技术或通过化学合成变体DNA序列或其部分而产生变体DNA序列。突变体多核苷酸还包括合成来源的多核苷酸，例如采用定点诱变所得到的突变体，或者是通过重组的方法(例如DNA改组)所得到的突变体，或者是通过自然选择所得到的突变体。
作为所述突变体的一种具体的实施方案，本发明提供了一种调控玉米穗腐病抗性的关键DNA序列的突变体，其多核苷酸序列如SEQ ID No.2所示；该突变体是在ZmLOX3基因启动子区Indel151处造成151bp的碱基***，其***的序列如SEQ ID No.3；在ZmLOX3基因的第二个内含子区Indel363处缺失361bp或者363bp碱基，其缺失的多核苷酸序列为SEQ IDNo.4或者SEQ ID No.5所示；该突变体能够升高ZmLOX3基因在玉米果穗和籽粒中的表达量高，表现出发生穗腐病的风险提高。
作为所述突变体的一种优选的具体实施方案，本发明提供了一种调控玉米穗腐病抗性的关键DNA序列的突变体，其多核苷酸序列如SEQ ID No.14所示；该突变体是在ZmLOX3基因启动子区Indel151处缺失151bp的碱基，其缺失的序列如SEQ ID No.3所示；在ZmLOX3基因的第二个内含子区Indel363处***363bp碱基，其***的多核苷酸序列如SEQ ID No.5所示；该突变体能够降低ZmLOX3基因在玉米果穗和籽粒中的表达量，表现出发生穗腐病的风险显著降低。
本发明因此确定在ZmLOX3基因启动子区Indel151缺失SEQ ID No.3所示的151bp的片段或在ZmLOX3基因第二个内含子区Indel363处***SEQ ID No.4或SEQ ID No.5所示的碱基片段的玉米材料普遍具有较低的ZmLOX3基因表达量，进而表现出抗轮状镰孢引起的穗腐病，发生穗腐病的风险降低；相反，在ZmLOX3基因启动子区Indel151***SEQ ID No.3所示的151bp的片段或在ZmLOX3基因的第二个内含子区Indel363处缺失SEQ ID No.4或SEQID No.5所示的碱基片段的玉米材料普遍具有较高的ZmLOX3基因表达量，表现出易感轮状镰孢引起的穗腐病，发生穗腐病的风险提高。
由此，本发明提供了一种降低玉米发生穗腐病风险或提供穗腐病抗性的方法，包括：在ZmLOX3基因启动子区Indel151缺失SEQ ID No.3所示的151bp的片段；或者，在ZmLOX3基因第二个内含子区Indel363处***SEQ ID No.4或SEQ ID No.5所示的碱基片段；或者在ZmLOX3基因启动子区Indel151缺失SEQ ID No.3所示的151bp的片段以及在ZmLOX3基因第二个内含子区Indel363处***SEQ ID No.4或SEQ ID No.5所示的碱基片段。
本发明还提供了一种培育抗穗腐病的玉米新品种的方法，包括降低ZmLOX3基因在玉米果穗和籽粒中的表达量，进而提高玉米穗腐病的抗性；因此，通过降低ZmLOX3基因在玉米果穗和籽粒中的表达量来达到提高玉米穗腐病抗性的方法都应属于本发明的保护范围；包括利用其它的组成型或组织特异的启动子和ZmLOX3构建的表达盒驱动ZmLOX3基因在玉米果穗和籽粒中表达，或利用其它的自然变异来改变ZmLOX3基因在玉米果穗和籽粒中表达，或通过RNAi来降低ZmLOX3基因在玉米果穗和籽粒中表达的方法。
作为本方面一种优选的具体实施方案，降低ZmLOX3基因在玉米果穗和籽粒中的表达量的方法可以是：在ZmLOX3基因启动子区Indel151缺失SEQ ID No.3所示的151bp的片段；或者，在ZmLOX3基因第二个内含子区Indel363处***SEQ ID No.4或SEQ ID No.5所示的碱基片段；或者在ZmLOX3基因启动子区Indel151缺失SEQ ID No.3所示的151bp的片段以及在ZmLOX3基因第二个内含子区Indel363处***SEQ ID No.4或SEQ ID No.5所示的碱基片段。
进一步的，含有所述SEQ ID No.1所述的部分或全部DNA序列、SEQ ID No.2或SEQID No.14所示的部分或全部的DNA序列的突变体的表达盒、含有该表达盒的重组植物表达载体、转基因细胞系和宿主菌均属于本发明的保护范围之内。
所述的重组植物表达载体是将所述表达盒与质粒或表达载体所构建得到的重组植物表达载体并且能够将所述表达盒转入植物宿主细胞、组织或器官中。
本发明的DNA序列或其突变体可以用于制备转基因植物。譬如，通过农杆菌介导、基因枪法等方法将含有所述DNA序列或其突变体的重组植物表达载体导入植物细胞、组织或器官中，再将该转化的植物细胞、组织或器官培育成植株，获得转基因植物。用于构建所述植物表达载体的出发载体可为任意一种用于农杆菌转化植物的双元载体或可用于植物微弹轰击的载体等。
为了实施本发明，制备和使用植物表达载体和宿主细胞的常规组合物和方法为本领域技术人员所熟知，具体方法可以参考例如Sambrook等。
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。所述的标记基因包括：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。
总之，可将本发明提供的DNA序列、该序列的突变体、分子标记Indel151或Indel363以及检测该分子标记变异情况的特异性检测引物以及检测ZmLOX3基因表达量的特异性检测引物等应用于培育抗病的玉米新品种，尤其是应用在改良玉米穗腐病抗性等方面。
作为参考，本发明提供了一种调控玉米的穗腐病抗性的方法，包括：用SEQ IDNo.1所示的DNA序列，SEQ ID No.2所示的DNA序列或SEQ ID No.14所示的DNA序列的突变体调控ZmLOX3基因在玉米中进行表达。
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。在特定的实施方案中，可利用多种瞬时转化法将本发明的表达盒提供给植物。利用常规方法可使已转化的细胞再生稳定转化植株(McCormick et al.Plant CellReports.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。
本发明进一步确定Indel151或者Indel363能够作为调控ZmLOX3基因在玉米果穗和籽粒中表达的分子标记。本发明还发现，Indel151和Indel363几乎完全连锁在一起，任意的其中一个变异都可以单独作为检测ZmLOX3基因在玉米果穗和籽粒中表达的分子标记。
本发明进一步提供了用于检测Indel151变异情况的检测引物；作为一种优选的具体实施方案，所述的检测引物的核苷酸序列分别为SEQ ID No.6和SEQ ID No.7所示；采用所述特异性检测引物可对玉米品种中的Indel151的变异情况进行检测，从而用于玉米的分子辅助育种。
本发明进一步提供了用于检测Indel363变异情况的检测引物；作为一种优选的实施方案，所述的检测引物的核苷酸序列分别为SEQ ID No.8和SEQ ID No.9所示；采用所述特异性检测引物可对玉米品种中的Indel363的变异情况进行检测，从而用于玉米的分子辅助育种。
此外，其他的根据SEQ ID No.1，SEQ ID No.2或SEQ ID No.14所设计的引物，或通过其他方法设计的可用于检测Indel151或Indel363变异情况的引物也应属于本发明的保护范围。
本发明还进一步提供了用于检测ZmLOX3基因表达量的特异性扩增引物，该特异性检测引物序列分别为SEQ ID No.10和SEQ ID No.11所示；采用该特异性检测引物可对玉米品种的ZmLOX3基因表达量进行检测，进而为玉米育种中针对ZmLOX3基因的研究提供参考。
本发明中所述ZmLOX3基因，其编码区的核苷酸序列为SEQ ID No.12所示，其编码蛋白如SEQ ID No.13所示。
本发明所提供的调控玉米穗腐病的DNA序列或其突变体能够调控ZmLOX3基因在玉米果穗或籽粒中的表达，其对改良玉米穗腐病具有重要意义，还可将其进一步应用于玉米新品种的选育。本发明还进一步提供了用作调控ZmLOX3基因表达的分子标记Indel151或Indel363、检测所述分子标记变异情况的特异性检测引物以及检测玉米中ZmLOX3基因表达量的特异性检测引物，这些均能直接应用于定向改良玉米的穗腐病抗性，对于玉米抗病新品种选育也具有重要的应用潜力。
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“同源性”指多核苷酸序列之间在百分比核苷酸位置同一性(即序列相似性或同一性)方面的相似性或百分同一性的水平。此处所用的术语同源性也指不同多核苷酸分子之间相似的功能特性的概念，例如具有相似功能的启动子可能具有同源的顺式元件。当多核苷酸分子在特定条件下特异性地杂交以形成双链体分子时，它们是同源的。在这些条件下(称为严谨杂交条件)一个多核苷酸分子可以用作鉴定共有同源性的另一个多核苷酸分子的探针或引物。
本发明中所述“严谨杂交条件”意指在所属领域中已知的低离子强度和高温的条件。通常，在严谨条件下，探针与其靶序列杂交的可检测程度比与其它序列杂交的可检测程度更高(例如超过本底至少2倍。严谨杂交条件是序列依赖性的，在不同的环境条件下将会不同，较长的序列在较高温度下特异性杂交。通过控制杂交的严谨性或洗涤条件可鉴定与探针100％互补的靶序列。对于核酸杂交的详尽指导可参考有关文献(Tijssen,Techniquesin Biochemistry and Molecular Biology-Hybridization with Nucleic Probes,"Overview of principles of hybridization and the strategy of nucleic acidassays.1993)。更具体的，所述严谨条件通常被选择为低于特异序列在规定离子强度pH下的热熔点(Tm)约5-10℃。Tm为在平衡状态下50％与目标互补的探针杂交到目标序列时所处的温度(在指定离子强度、pH和核酸浓度下)(因为目标序列过量存在，所以在Tm下在平衡状态下50％的探针被占据)。严谨条件可为以下条件：其中在pH 7.0到8.3下盐浓度低于约1.0M钠离子浓度，通常为约0.01到1.0M钠离子浓度(或其它盐)，并且温度对于短探针(包括(但不限于)10到50个核苷酸)而言为至少约30℃，而对于长探针(包括(但不限于)大于50个核苷酸)而言为至少约60℃。严谨条件也可通过加入诸如甲酰胺的去稳定剂来实现。对于选择性或特异性杂交而言，正信号可为至少两倍的背景杂交，视情况为10倍背景杂交。例示性严谨杂交条件可如下：50％甲酰胺，5×SSC和1％SDS，在42℃下培养；或5×SSC，1％SDS，在65℃下培养，在0.2×SSC中洗涤和在65℃下于0.1％SDS中洗涤。所述洗涤可进行5、15、30、60、120分钟或更长时间。
本发明中所述的“多个”通常意味着2-8个，优选为2-4个；所述的“替换”是指分别用不同的氨基酸残基取代一个或多个氨基酸残基；所述的“缺失”是指氨基酸残基数量的减少，也即是分别缺少其中的一个或多个氨基酸残基；所述的“***”是指氨基酸残基序列的改变，相对天然分子而言，所述改变导致添加一个或多个氨基酸残基。
术语“编码序列”：转录成RNA的核酸序列。
术语“启动子”指多核苷酸分子，所述多核苷酸分子在其天然状态位于可读框(或蛋白质编码区)的翻译起始密码子上游或5’，并参与RNA聚合酶II及其它蛋白质(反式作用转录因子)的识别和结合以启动转录。
术语“植物启动子”是在植物细胞中有功能的天然或非天然启动子。组成型植物启动子在植物发育始终的大部分或所有组织中发挥功能。可以将任何植物启动子用作5’调节元件以用于调节与其可操作地连接的一种或多种特定基因的表达。当与可转录的多核苷酸分子可操作地连接时，启动子一般引起该可转录的多核苷酸分子的转录，其转录方式与该启动子通常连接的可转录的多核苷酸分子的转录方式类似。植物启动子可以包括通过操作已知启动子以产生人工、嵌合或杂合启动子而产生的启动子。这类启动子也可以通过例如向具有其自身部分或全部调节元件的活性启动子加入异源调节元件而组合了来自一个或多个启动子的顺式元件。
术语“顺式元件”指赋予基因表达全面控制的一方面的顺式作用转录调节元件。顺式元件可以起到结合调节转录的转录因子，反式作用蛋白质因子的作用。一些顺式元件结合超过一种转录因子，而且转录因子可以与超过一种顺式元件以不同的亲和力相互作用。
术语“可操作地连接”指第一个多核苷酸分子(例如启动子)与第二个可转录的多核苷酸分子(例如目的基因)连接，其中多核苷酸分子如此排列，从而第一个多核苷酸分子影响第二个多核苷酸分子的功能。优选地，两个多核苷酸分子是单个连续多核苷酸分子的部分，且更优选是临近的。例如，如果启动子在细胞内调节或介导目的基因的转录，则该启动子与目的基因可操作地连接。
术语“可转录的多核苷酸分子”指能够被转录为RNA分子的任何多核苷酸分子。已知以使可转录的多核苷酸分子被转录为功能mRNA分子的方式将构建体引入细胞的方法，所述功能mRNA分子得到翻译并从而表达为蛋白质产物。为了抑制特定的目的RNA分子的翻译，也可以构建能够表达反义RNA分子的构建体。
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。
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“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***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。
附图说明
图1A为通过测序方法发现ZmLOX3的启动子和第二个内含子区分别存在一个***缺失自然变异(Indel)；Indel151，启动子区的151bp的Indel；Indel363，第二个内含子区的363bp的Indel。图片的右侧是所测序的12个自交系中授粉当天的胚珠(Ovule_0DAP)和授粉后5天的籽粒(Kernel_5DAP)中ZmLOX3基因的表达情况。整体上，有Indel151***或Indel363缺失的自交系中ZmLOX3基因的表达量较高；B为GWAS分析表明Indel151和Indel363与ZmLOX3基因的表达高低显著关联，曼哈顿图的下边为所有标记的连锁不平衡模块(LD-Block)图；C为自交系中Indel151的不同基因型与ZmLOX3基因表达量间的关系情况，当存在Indel151的***时，ZmLOX3基因的表达量较高；D为Indel151的不同基因型在不同年代的玉米品种间的等位基因频率变化情况，其中降低ZmLOX3基因表达所对应的基因型在育种过程中受到选择；E为Indel151处的两种基因型在不同杂种优势群材料间的等位基因频率分布情况。
图2为12个自交系的Indel151基因型鉴定结果。
图3为12个自交系的Indel363基因型鉴定结果。
图4为10个自交系的玉米籽粒接菌结果；其中有Indel151***、在Indel363处缺失的5个自交系与其余不同基因型自交系相比明显更易感病。
图5为在杂交种中不同Indel151基因型对应ZmLOX3基因的表达量情况(左图)及Indel151的不同基因型在不同年代杂交种间的等位基因频率变化情况(右图)。
具体实施方式
以下结合具体实施例来进一步描述本发明，本发明的优点和特点将会随着描述而更为清楚。但这些实施例仅是范例性的，并不对本发明的范围构成任何限制。本领域技术人员应该理解的是，在不偏离本发明的精神和范围下可以对本发明的细节和形式进行修改或替换，但这些修改和替换均落入本发明的保护范围内。
以下实施例中所用的自交系可从“中国作物种质信息网”得到相关信息和申请获取对应的种子。
实施例1 ZmLOX3基因启动子区和第二个内含子区的***缺失(Indel)变异调控ZmLOX3基因的表达量变化的试验
1、ZmLOX3基因启动子区和第二个内含子区变异位点的发掘
利用平均>55×的PacBio SMRT reads数据，>94×的二代Illumina 150bp双末端测序数据，及>139×的BioNano单分子数据，对世界范围内广泛使用的12个玉米骨干自交系亲本进行全基因组组装。这12个自交系为：PH207、A632、OH43、Zheng58、Ye478、Dan340、Huangzaosi、Chang7-2、Jing92、Jing724、Xu178、S37；加上已发表的B73和Mo17基因组，这14个高质量基因组几乎涵盖了玉米育种上所有杂种优势群的核心代表自交系。
将14个核心自交系中ZmLOX3基因上下游10kb的基因组序列调取出来，进行DNA序列的Blast比对，发现14个自交系中，在ZmLOX3基因的启动子和第二个内含子中各存在一个***缺失变异(Indel，图1中A)。其中启动子区的Indel长度为151bp，其序列为SEQ IDNo.3；第二个外显子区的Indel长度为363bp，其碱基序列为SEQ ID No.5所示，或者该外显子区的Indel长度为361bp，其碱基序列为SEQ ID No.4所示。
2、Indel151和Indel363与ZmLOX3基因的表达量变异显著关联
中国历史上经历了6次玉米品种(杂交种)的更新换代，本发明人前期收集组配了116份不同年代品种更新过程中广泛利用的杂交种及其对应的137份亲本自交系(部分杂交种有共享的亲本)。2019年夏天，对这253份材料(116杂交种和137份自交系)吐丝期的雌穗进行了取样(每个材料取样3个重复)，然后进行RNA-Seq分析，得到20000余个基因在253份材料果穗中的表达量数据。其中，本发明中单独把ZmLOX3基因的表达量数据抽出来用于后续分析。
利用137份玉米自交系的深度(>10×)二代重测序的数据结合已公布的玉米B73V3基因组，挖掘出了25,320,664单核苷酸多态性分子标记(SNPs)和4,319,510个***缺失多态性分子标记(Indel)。此外，利用开发的PCR引物LOX3-2(序列为SEQ ID No.6和SEQ IDNo.7)及LOX3-3序列为SEQ ID No.8和SEQ ID No.9)鉴定了137份玉米自交系中Indel151(图2)和Indel363(图3)的基因型。
利用这些挖掘到的分子标记估算出137份玉米自交系的群体结构和亲缘关系，然后将检测到的ZmLOX3基因的表达量数据作为表型进行全基因组关联分析(GWAS)。其中在ZmLOX3基因附近鉴定到一个显著的关联信号，最显著的信号落在Indel151标记上(p-value＝1.40E-9)，Indel363为第二显著的关联标记(图1B)，这表明Indel151和Indel363与ZmLOX3基因的表达量变异显著关联。
本试验进一步根据Indel151处的基因型将137份材料分成两类，一类为Indel151处缺失151bp序列的基因型材料，共计95份；另一类为Indel151处存在151bp序列的基因型材料，共计33份；比较两类材料间的ZmLOX3基因表达量，发现缺失Indel151的材料中ZmLOX3基因的表达量更低(图1C)，表明Indel151可以正调控ZmLOX3基因的表达。连锁不平衡分析还表明，Indel151和Indel363位于同一个LD-block(连锁不平衡模块)内(图1B)；并且14个自交系中的序列分析也表明，这两个变异总是以相斥相的形式连锁在一起出现，即当一个材料中Indel151处为缺失151bp序列的基因型时，往往在Indel363处为***363bp或361bp序列的基因型(图1A)。表明有可能通过两者之一的基因型推测出另一个Indel处的基因型；不过，Indel151与ZmLOX3基因的表达量关联性更强，因此Indel151可作为更优的检测ZmLOX3基因表达量的分子标记。
实施例2 Indel151和Indel363通过调控ZmLOX3基因表达量影响玉米穗腐病抗性
适当降低籽粒中ZmLOX3基因的表达量，但不改变其蛋白功能，有可能起到抗镰孢菌同时兼抗黄曲霉的作用。
本发明人前期研究结果显示，缺失Indel151或***Indel363的材料，往往具有较低的ZmLOX3基因表达量。为了进一步验证Indel151和Indel363与ZmLOX3基因表达量间的关系，将前文中的12份代表性自交系(PH207，A632，OH43，Zheng58，Ye478，Dan340，Huangzaosi，Chang7-2，Jing92，Jing724，Xu178，S37)，分别种在田间，对其吐丝期的胚珠和授粉5天后的籽粒进行取样和液氮速冻。每5个单株的样品混合成一个样品，每种自交系取3个生物学重复，之后用TRIzol法提取RNA，利用特异引物(SEQ ID No.10和SEQ ID No.11)检测ZmLOX3基因的表达量。发现与前文GWAS分析结果一样，缺失Indel151或***Indel363的材料(PH207，A632，Zheng58，Ye478，Jing724，Xu178，S37)，不论是在未授粉的胚珠，还是授粉5天后的籽粒中，普遍具有较低的ZmLOX3基因表达量(图1中A)，再次验证了Indel151和Indel363与ZmLOX3基因表达量间的关系。
进一步的对12个代表性自交系中的10个材料的玉米籽粒进行轮状镰孢F.verticillioides接菌实验(图4)。结果表明，ZmLOX3基因表达低的5个自交系(B73、Ye478、Mo17、PH207、A632，缺失Indel151，***Indel363)普遍比ZmLOX3基因表达高的5个自交系(Jing92、Dan340、Huangzaosi、OH43、Chang7-2，***Indel151，缺失Indel363)更抗轮状镰孢引起的穗腐病。这些结果表明Indel151和Indel363能通过调控ZmLOX3基因表达量进而影响玉米穗腐病抗性。
实施例3 Indel151处的优异等位基因在现代玉米育种过程中受到了强的人工选择
本发明人前期收集了来自于中国和美国不同的育种时期的350份玉米自交系材料，包括美国早期(Public-US)和近现代(Ex-PVP)育种材料163份，中国早期(CN1960&70s)、中期(CN1980&90s)和当前(CN2000&10s)主栽玉米育种材料187份。对Indel151的基因型(Indel363与Indel151紧密连锁，可以被Indel151的基因型所代表)在这些材料中的频率分布进行分析发现，随着育种时期的推进，缺失Indel151(对应较低的ZmLOX3基因表达量和较高的穗腐病抗性)类的基因型频率在中国和美国玉米育种过程中都发生了显著的提升(图1D)，表明缺失Indel151类的基因型在现代玉米育种过程中受到了强的人工选择。
另外，350份玉米自交系材料还可以被划分到8个杂种优势类群中，分别为SS、PA、X、NSS、TSPT、IDT、PB、Mixed。前人报道显示，TSPT种质普遍比较易感穗腐病，其中，前文中的Huangzaosi、Chang7-2、Jing92都是典型的TSPT种质，本发明的接菌实验也证明其确实比较易感穗腐病(图4)，印证了前人的报道。本发明还发现，在TSPT种质中，***Indel151类的基因型频率较高，而Indel151的***对应较高的ZmLOX3基因表达量，进而易感穗腐病，揭示了TSPT种质易感穗腐病的原因。同时，也预示了Indel151与Indel363在TSPT种质穗腐病抗性改良方面的重要潜力。
进一步的，本发明还分析了116个不同年代的杂交种中ZmLOX3基因的表达情况及Indel151不同基因型的分布情况。分析发现在杂交种中，缺失Indel151的材料中ZmLOX3的表达量显著低于杂合和***Indel151的材料(图5中的左图)。ZmLOX3低表达量对应的是抗穗腐病，所以缺失Indel151的基因型为优良基因型，***Indel151的基因型为不利基因型。等位基因频率分析发现，Indel151的纯合优良等位基因型在杂交种的育种过程中逐渐上升，而纯合不利基因型在2000年之后的杂交种中几乎消失；并且随着育种年代的推进，杂合基因型的比例也越来越低，进一步证明Indel151在育种过程中受到了强的选择。这些杂交种全部是玉米生产中直接栽培利用的品种，在杂交种中受到选择，进一步证实了该自然变异(Indel151及其紧密连锁的Indel363)的重要性。
SEQUENCE LISTING
<110> 中国农业科学院生物技术研究所
<120> 调控玉米穗腐病抗性的DNA序列及其突变体、分子标记和应用
<130> BJ-2002-210706A
<160> 14
<170> PatentIn version 3.5
<210> 1
<211> 6655
<212> DNA
<213> Zea mays L.
<400> 1
ggcgaactcc gctccgcccg accccagggc tcggactcgg gctaagaccc ggaagacggc 60
gaactccgct ccgcccgacc ccagggctcg gactcgggct aagacccgga agacggcgaa 120
ctctgctccg cccgacccca gggctcggac tcgggctaag acccggaaga cggcgaatct 180
ccgcctcgcc cgaccccagg gctcagactc cgccctggcc tcggccaaac gatctccgcc 240
tcgcccgacc ccagggctcg gactccgccc tagcctcggc caaacgatct ccgcctcgcc 300
cgacccgggg gctcgggctc ggcctcggca acggaaggca gactcgacct cgacttcgga 360
ggagccccca cgtcgccctg cctagggcac aggtccgcca cgtcaacagg aagcgccatc 420
accaacctac cccgagccga cttgggacac gaaggacaag accggcgtcc catctggcca 480
gctccgccgg atgggcaatg atggcgcccc ccgagctctg tgacgacggc ggctcttagc 540
tctcttacgg cagcagagcg acgtcagcaa ggactcgacc gctccaacag ctgtccctcc 600
gtcaggctcc gtcgctcctc cgacagccac gacatcacgc cagcaaggtg ccaagacctc 660
tccggctgcc acattggcat gtacccaggg cgttagctct ctctctctcc gctagacacg 720
tagcactctg ctacccccgt tgtacacctg gatcctctcc ttacgactat aaaaggaagg 780
accagggcct tcttagagaa ggttggccgc gcgggaccga ggacgggaca ggcgctctct 840
tggggccgct cgcttccctc acccgcgtgg acgcttgtaa cccccctact gcaagcgcac 900
ctgacctggg cgcgggacga acacgaaggc cgcgggactt ccacctctct cacgctcgac 960
tctggccacc tcgcctctcc ccccttcgcg ctcgcccacg cgctcgaccc atctgggctg 1020
gggcacgcag cacactcact cgtcggctta gggacccccc tgtctcgaaa cgccgacaat 1080
aagcttatct cgaattcatg tggtggagga ttggaaatga tttttatgta ttagtagaat 1140
ttgtttctac tctgtaaatt acatgaccct cttcgtctca ctcctctata gtaaaaatat 1200
agcacataaa tatctccgac atcttgctaa taatagtata caaatatatt tttcatcaaa 1260
ccgaattaac ttaattgata tatgtctaaa ttactgttat tagaatggaa ttcaattcca 1320
atgaaccaaa cggggcgtaa gtgatttctt agtgaggtgg cacctatgga aatcacttag 1380
tagtttctgg gttgtaattg ctcttatggc atgaagaaaa ttttgcattc gagctcgaac 1440
gctacaaggt cgttttagaa gctacatcag taatgcttcg agtatgacaa acaacatcac 1500
aagtaacatc tacctatatg caccgaatct ccaccaccac cacaatatcg atgtcagaag 1560
gctagggaag gacgaccaat aatatagaag acacataggc cacactcaat tcaatcaaca 1620
tcacgacaat ctacttgatg tcctatgtag ccatgatctc tacgagcaat caatatcaat 1680
gcactagaat cccaactacg tagagtatta ttggacgaac gagtaaaaca ccaaagagca 1740
gccactggac gactttcttg catctccaca aggataaatt ctctcagata gtcagatgcc 1800
taatgaaatt caagtccatc aatattgaca tgtgcgaagg caagaaggat ccaaattagt 1860
tgattcgaca ctacttggtt accatcaatc tcaagggagg aaacaaagat atcaaagcat 1920
catgctttca aatgtactca aagctactct agtgacttgg cttgaatagc tcccaagagg 1980
ctacatcgat aacaggctag cactctccaa ggaattaatg gacaacttcc aaagtgcatg 2040
gcctcgttcg gacaaccggt acaatcttca aaatggaagt agaagattgg tgaaagcata 2100
tgcaactagt acaagctctt gatcgaaatt tgatgctaca ggcctaacta cttcgacgat 2160
gatcatgggg aaaagttcta gtatcagaat ttcataaaaa caaaccaaaa acaattcatg 2220
agttctaaca catgatcaac aactgaatga atgtcgaaga cacagtacga gataagtttg 2280
acaattgaaa ccgtagatat aacaataaag ataacattca tagagacaac aagggcagct 2340
tatcggataa gaagtgaggg aaagataaca ctatgataag cactcgcatc aaagaagggt 2400
ggcaagaaca aatcattcaa ttagcaactc tttgagcaac acccatacca cctacatcat 2460
aaatcataag tgctctacca ctaaatgcat gcaactcaag aactttggca tcatcttagc 2520
tcggaagcaa gacaaaacca aacagagttg ataaaaaata ttaggaaaaa tagaataatg 2580
gagacttcta gagggccaca aacatagtta acctcatctt caaaggagca tcttccttaa 2640
tacctaagag gaagactaaa cttacccttt tgtgagatga tggttgttga accatcacct 2700
acaacatata tgtgatggtt cgagatccca attcaattta taagggagga ttagtttctc 2760
aaacatagga tgctatggtc tagtcctggc cccgatggtg gcaggacttc aacttagtaa 2820
ggaaggtgaa ttaccaatat gttcaaaagg atggggtaaa aggaaggtga attatgcaat 2880
ctatttttcg cactttttca ttaatcaaaa cctatatgga taaccaatcg ttcatatgtg 2940
caactaaggt tttgactaag tgttgctatc tctaccgtaa aaggagtttt gctaccccaa 3000
acctatcaac tagtctatga ctaagctaag aagataaatc acacaaccac aagtaacaat 3060
ataaatgtgg aattttaaat atggtagaga tacaaactct cgttgatgtg tcagtatttt 3120
tactggagta tcaagaaacg cgcaagcttc ttactaatcc ttcttagagc ctcacgcaag 3180
gctaagctcc cgctcaggta accctgtata gatacaaacc ctaccaatga ctatagtaac 3240
ttttatgaag atatagggaa acacacaagt tttaccctag ttctcgttgg agcctcttat 3300
aaacatgtcc acaaggggat gaagctaaga gtcgagtaag ggtttgtttg atttctttag 3360
tcttaatgac taaaactaag caaagagctt taattattgt gcaaattgat tacattatcc 3420
ctaattaatg ctatcttcga ctactgttta cccacacgca gagctactgt tcgtgcgcat 3480
tcacaatgct gcatgcgtgc acgttgcttt ggggaggagc ctacttcatg cgcgttggtt 3540
agggagggca tatgaagtaa aaatgtcact ttatggttag tttgacaccc tcattttttt 3600
caagggattg tattttcaca aaagaaatta atttattttt cttgaaaaat aggaatccct 3660
tagaaaaaaa tagagttgtc aaactaaccc ttattcattt tagtcactct tttggtaatt 3720
agaggactat aatttagttg gaggatttta gtcacactgt gttgattctt tagtgactaa 3780
aaatgactaa aatttaatca attaaatgta gtcacccaaa ccaaacagag tcgcgcgtcc 3840
gacgtgcctc ctcaaccggc gcgtgggagt aggtctggaa aaaagctcaa gctcgatgag 3900
ttggcttggg ctcacggtaa cttgggtcgg ctcgacgctc aaaacgagtc caagtcctat 3960
ttttgtggct cgtgataagt gtgagctagc tcggctcggc tcacgaaccg acaacaattt 4020
actacataaa attctgatta gcatataatg ttagtaccaa atatacaatt agtatatgtt 4080
atttgatatt tatatacaaa attaatttat ttctatgtta tattagtact ataattattt 4140
attaatttaa gagatgtaat ttgattattt ttgatatttt atgttataat ttgtaacgta 4200
agctggtgct tgtggttaga ctctctacta gtcgagtttt cacgctcgtc aaaggaccga 4260
gccgagacga ggcaagctca acacattacc gagccgaccg gctcgtttcc gccacgtggt 4320
agagcggcag cacccttccc agatctcgtg tcccgcgcgc cgtgcgcaca gccgcgcaca 4380
cacacggact gccgcgtccg ggcgcgatcc gttctcgcga acacaccggt cgtagccgtc 4440
cgtaatttgt actagtatgg cacgcggaac acgagtcggc gccagcaccg gcacgtcacc 4500
atcggggccg ctggccccat acgggcagcc aattaataaa tccttgctct gcgacagacg 4560
cgcaggggca cgtcacgcag ccgtagtcga tcggtgcacg gtgctcgcgc ccccgttgca 4620
gcgcgcagct ctcccgcgct ataaatgccc cggctcggcc tcgctcccac agccacagcc 4680
tcacacagac accaacgcca ctgcactgca aaagcaagag cagctagcta gtaaagatgc 4740
tgagcgggat catcgacggg ctgacggggg cgaacaagca tgcgcggctc aagggcacgg 4800
tggtgctcat gcgcaagaac gtgctggacc tcaacgactt cggcgccacc gtcgttgaca 4860
gcatcagcga gttcctcggc aagggggtca cctgccagct catcagctcc accctcgtcg 4920
acgccagtga gtaccgcgcc gcgccgccgg cacctctccg atctgcgctt ccccatgtcg 4980
atcgatctcg atctctctag gctctagagc tatagctctc tcggccccca ctttttacct 5040
ttgcaagcat tttccctgca tgcgaaacaa gcgatagttt actatttggg cggccatgct 5100
gctgctgctt cggctacctt gcctccgtca tctttgacgg gacatggaaa gaaagaaaga 5160
atagagagag agacagagag agagagagag caaacaccga gaaaaagaca gcaaagctag 5220
ttgtagcctg ggcgcagaac acagcaccag atgctggcta gctcgtgaca aagtaaaaaa 5280
agagaggaac gaacacagta gtaccaagag atcagggacg agacctttct actttgaact 5340
ggattatata ttggattctt tatcagtaac ttactgctgc tagtataccc ctaccctagt 5400
ctcggggcga cgtgcctgcg tgcatgcccg acgcgtacga aacacatcga cggactcaca 5460
tgggccaccg cgcgcgcgcg tgcctgcttt aactttcgct gtgcagacaa cggcaaccgc 5520
gggcgggtcg gggcggaggc gaacctggag cagtggctga cgagcctgcc gtcgctgacg 5580
accggcgagt ccaagttcgg cgtcacgttc gactgggagg tggagaagct gggagtgccg 5640
ggggccgtcg tcgtcaagaa caaccacgcc gccgagttct tcctcaagac aatcaccctc 5700
gacgacgtgc ccggccgcgg cgccgtcacc ttcgtcgcca actcctgggt ctaccccgcg 5760
ggcaagtacc gctacaaccg cgtcttcttc tccaacgatg tgagtccttt ctcgatagat 5820
cattatgttt gtttgttatt ttgtttcttg gtatcatgta ggcatgtagc tagccgccat 5880
gtcgtcagtt ggagtgcagt aggtaggaaa aaggacgaca tgggatggga gtggttaaga 5940
aaatccatgc aagtgggact agtgtgtaac tggtagtata gctgaagaat ctagtggtag 6000
aatgatcttg tacgtgaata atgtttctga cgctgagcgc tgaggctatc cgcaaccgtt 6060
aaccctaaat ttttccctct atatcatttt ttcctctatt ttcctcccta ttttttcatc 6120
tcccgcagcg gttcccccta aatactcccc ctatatctca ctaccactat aaaatattat 6180
tttctatacc aattatcaat tttttatcta ctaacaatta ctcgtggacc cacagcacag 6240
tgtttagggt gatgaacagt gacacgctag atctgaaggg agagagaagg ggaccgacac 6300
gtagggagcc tgtagagggc accgctgcgg ccgtagggtg ctccctacgc gccgcataca 6360
aggggagggg ggagaggcag cggtaaccgc tgcgcacagc ctgagggcga ggcatgtgag 6420
ttcaccacgt gagtagcagc aaaaggaaac aacccttctt cacccggcta tcatctaacg 6480
tatcgccccg ggagaatcaa taactctaac gagatgacga aaagtcaaaa ataaagtcgt 6540
gtgatggcca tgaaagtcag tcaagcaaat cagctgctaa cacgtgtccc ttatctacag 6600
gtgtaagaga gtagagtctt gtcaatcaac ctgggttgtt ttctatctgc gtttt 6655
<210> 2
<211> 6434
<212> DNA
<213> Zea mays L.
<400> 2
ggcgaactcc gctccgcccg accccagggc tcggactcgg gctaagaccc ggaagacggc 60
gaactccgct ccgcccgacc ccagggctcg gactcgggct aagacccgga agacggcgaa 120
ctctgctccg cccgacccca gggctcggac tcgggctaag acccggaaga cggcgaatct 180
ccgcctcgcc cgaccccagg gctcagactc cgccctggcc tcggccaaac gatctccgcc 240
tcgcccgacc ccagggctcg gactccgccc tagcctcggc caaacgatct ccgcctcgcc 300
cgacccgggg gctcgggctc ggcctcggca acggaaggca gactcgacct cgacttcgga 360
ggagccccca cgtcgccctg cctagggcac aggtccgcca cgtcaacagg aagcgccatc 420
accaacctac cccgagccga cttgggacac gaaggacaag accggcgtcc catctggcca 480
gctccgccgg atgggcaatg atggcgcccc ccgagctctg tgacgacggc ggctcttagc 540
tctcttacgg cagcagagcg acgtcagcaa ggactcgacc gctccaacag ctgtccctcc 600
gccaggctcc gtcgctcctc cgacagccac gacatcacgc cagcaaggtg ccaagacctc 660
tccggctgcc acattggcat gtacccaggg cgttagctct ctctctctcc gctagacacg 720
tagcactctg ctaccccccg ttgtacacct ggatcctctc cttacgacta taaaaggaag 780
gaccagggcc ttcttagaga aggttggccg cgcgggaccg aggacgggac aggcgctctc 840
ttggggccgc tcgcttccct cacccgcgtg gacgcttgta acccccctac tgcaagcgca 900
cctgacctgg gcgcgggacg aacacgaagg ccgcgggact tccacctctc tcacgctcga 960
ctccggccac ctcgcctctc cccccttcgc gctcgcccac gcgctcgacc catctgggct 1020
ggggcacgca gcacactcac tcgtcggctt agggaccccc ctgtctcgaa acgccgacaa 1080
taagcttatc tcgaattcat ggggtggagg attggaaatg atttttatgt attagtagaa 1140
tttgtttcta ctctgtaaat tacatgaccc tcttcgtctc actcctctat agtaaaaata 1200
tagcacataa atatctccga catcttgcta ataatagtat acaaatatat ttttcatcaa 1260
accgaattaa cttaattgat atatgtctaa attactgtta ttagaatgga attcaattcc 1320
aatgaaccaa acggggcgta agtgatttct tagtgaggtg gcacctatgg aaatcactta 1380
gtagtttctg ggttgtaatt gctcttatgg catgaagaaa attttgcatt cgagctcgaa 1440
cgctacaagg tcgttttaga agctacatca gtaatgcttc gagtatgaca aacaacatca 1500
caagtaacat ctacctatat gcaccgaatc tccaccacca ccacaatatc gatgtcagaa 1560
ggctagggaa ggacgaccaa taatatagaa gacacatagg ccacactcaa ttcaatcaac 1620
atcacgacaa tctacttgat gtcctatgta gccatgatct ctacgagcaa tcaatatcaa 1680
tgcactagaa tcccaactac gtagagtatt attggacgaa cgagtaaaac accaaagagc 1740
agccactgga cgactttctt gcatctccac aaggataaat tctctcagat agtcagatgc 1800
ctaatgaaat tcaagtccat caatattgac atgtgcgaag gcaagaagga tccaaattag 1860
ttgattcgac actacttggt taccatcaat ctcaagggag gaaacaaaga tatcaaagca 1920
tcatgctttc aaatgtactc aaagctactc tagtgacttg gcttgaatag ctcccaagag 1980
gctacatcga taacaggcta gcactctcca aggaattaat ggacaacttc caaagtgcat 2040
ggcctcgttc ggacaaccgg tacaatcttc aaaatggaag tagaagattg gtgaaagcat 2100
atgcaactag tacaagctct tgatcgaaat ttgatgctac aggcctaact acttcgacga 2160
tgatcatggg gaaaagttct agtatcagaa tttcataaaa acaaaccaaa aacaattcat 2220
gagttctaac acatgatcaa caactgaatg aatgtcgaag acacagtacg agataagttt 2280
gacaattgaa accgtagata taacaataaa gataacattc atagagacaa caagggcagc 2340
ttatcggata agaagtgagg gaaagataac actatgataa gcactcgcat caaagaaggg 2400
tggcaagaac aaatcattca attagcaact ctttgagcaa cacccatacc acctacatca 2460
taaatcataa gtgctctacc actaaatgca tgcaactcaa gaactttggc atcatcttag 2520
ctcggaagca agacaaaacc aaacagagtt gataaaaaat attaggaaaa atagaataat 2580
ggagacttct agagggccac aaacatagtt aacctcatct tcaaaggagc atcttcctta 2640
atacctaaga ggaagactaa acttaccctt ttgtgagatg atggttgttg aaccatcacc 2700
tacaacatat atgtgatggt tcgagatccc aattcaattt ataagggagg attagtttct 2760
caaacatagg atgctatggt ctagtcctgg ccccgatggt ggcaggactt caacttagta 2820
aggaaggtga attaccaata tgttcaaaag gatggggtaa aaggaaggtg aattatgcaa 2880
tctatttttc gcactttttc attaatcaaa acctatatgg ataaccaatc gttcatatgt 2940
gcaactaagg ttttgactaa gtgttgctat ctctaccgta aaaggagttt tgctacccca 3000
aacctatcaa ctagtctatg actaagctaa gaagataaat cacacaacca caagtaacaa 3060
tataaatgtg gaattttaaa tatggtagag atacaaactc tcgttgatgt gtcagtattt 3120
ttactggagt atcaagaaac gcgcaagctt cttactaatc cttcttagag cctcacgcaa 3180
ggctaagctc ccgctcaggt aaccctgtat agatacaaac cctaccaatg actatagtaa 3240
cttttatgaa gatataggga aacacacaag ttttacccta gttctcgttg gagcctctta 3300
taaacatgtc cacaagggga tgaagctaag agtcgagtaa gggtttgttt gatttcttta 3360
gtcttaatga ctaaaactaa gcaaagagct ttaattattg tgcaaattga ttacattatc 3420
cctaattaat gctatcttcg actactgttt acccacacgc agagctactg ttcgtgcgca 3480
ttcacaatgc tgcatgcgtg cacgttgctt tggggaggag cctacttcat gcgcgttggt 3540
tagggagggc atatgaagta aaaatgtcac tttatggtta gtttgacacc ctcatttttt 3600
tcaagggatt gtattttcac aaaagaaatt aatttatttt tcttgaaaaa taggaatccc 3660
ttagaaaaaa atagagttgt caaactaacc cttattcatt ttagtcactc ttttggtaat 3720
tagaggacta taatttagtt ggaggatttt agtcacactg tgttgattct ttagtgacta 3780
aaaatgacta aaatttaatc aattaaatgt agtcacccaa accaaacaga gtcgcgcgtc 3840
cgacgtgcct cctcaaccgg cgcgtgggag taggtctgga aaaaagctcg agctcgatga 3900
gttggcttgg gctcacggta acttgggtcg gctcgacgct caaaacgagt ccaagtccta 3960
tttttgtggc tcgtgataag tgtgagctag ctcggctcgg ctcacgaacc gacaacaatt 4020
tactacataa aattctgatt agcatataat gttagtacca aatatacaat tagtatatgt 4080
tatttgatat ttatatacaa aattaattta tttctatgtt atattagtac tataattatt 4140
tattaattta agagatgtaa tttgattatt tttgatattt tatgttataa tttgtaacgt 4200
aagctggtgc ttgtggttag actctctact agtcgagttt tcacgctcgt caaaggaccg 4260
agccgagacg aggcaagctc aacacattac cgagccgacc ggctcgtttc cgccacgtgg 4320
tagagcggca gcacccttcc cagatctcgt gtcccgcgcg ccgtgcgcac agccgcgcac 4380
acacacggac tgccgcgtcc gggcgcgatc cgttctcgcg aacacaccgg tcgtagccgt 4440
ccgtaatttg tactagtatg gcacgcggaa cacgagtcgg cgccagcacc ggcacgtcac 4500
catcggggcc gctggcccca tacgggcagc caattaataa aagagccgaa gcgaccgacg 4560
gcgcaggtac acgcaggcgc ctcctgcacc attcaccatt cacgtctctg tggcccaata 4620
ataaaaccgc attaattacg ctcgcgcaga cacggtagca cactggccgc cacagtgcca 4680
cccacccacc aatccttgct ctgcgacaga cgcgcagggg cacgtcacgc agccgtagtc 4740
gatcggtgca cggtgctcgc gcccccgttg cagcgcgcag ctctcccgcg ctataaatgc 4800
cccggctcgg cctcgctccc acagccacag cctcacacag acaccaacgc cactgcactg 4860
caaaagcaag agcagctagc tagtaaagat gctgagcggg atcatcgacg ggctgacggg 4920
ggcgaacaag catgcgcggc tcaagggcac ggtggtgctc atgcgcaaga acgtgctgga 4980
cctcaacgac ttcggcgcca ccgtcgttga cagcatcagc gagttcctcg gcaagggggt 5040
cacctgccag ctcatcagct ccaccctcgt cgacgccagt gagtaccgcg ccgcgccgcc 5100
ggcacctctc cgatctgcgc ttccccatgt cgatcgatct cgatctctct aggctctaga 5160
gctatagctc tctcggcccc cactttttac ctttgcaagc attttccctg catgcgaaac 5220
aagcgatagt ttactatttg ggcggccatg ctgctgctgc ttcggctacc ttgcctccgt 5280
catctttgac gggacatgga aagaaagaaa gaatagagag agagacagag agagagagag 5340
agcaaacacc gagaaaaaga cagcaaagct agttgtagcc tgggcgcaga acacagcacc 5400
agatgctggc tagctcgtga caaagtaaaa aaagagagga acgaacacag tagtaccaag 5460
agatcaggga cgagaccttt ctactttgaa ctggattata tattggattc tttatcagta 5520
acttactgct gctagtatac ccctacccta gtctcggggc gacgtgcctg cgtgcatgcc 5580
cgacgcgtac gaaacacatc gacggactca catgggccac cgcgcgcgcg cgtgcctgct 5640
ttaactttcg ctgtgcagac aacggcaacc gcgggcgggt cggggcggag gcgaacctgg 5700
agcagtggct gacgagcctg ccgtcgctga cgaccggcga gtccaagttc ggcgtcacgt 5760
tcgactggga ggtggagaag ctgggagtgc cgggggccgt cgtcgtcaag aacaaccacg 5820
ccgccgagtt cttcctcaag acaatcaccc tcgacgacgt gcccggccgc ggcgccgtca 5880
ccttcgtcgc caactcctgg gtctaccccg cgggcaagta ccgctacaac cgcgtcttct 5940
tctccaacga tgtgagtcct ttctcgatag atcattatgt ttgtttgttt attggtatca 6000
tgtagctagc cgccatgtcg tcagttggag tgcagtaggt aggaaaaagg acgacatggg 6060
atgggagtgg ttaagaaaat ccatgcaagt gggactagtg tgtaactggt agtatagctg 6120
aagaatctag tggtagaatg atcttgtacg tgaataatgt ttctgacgct gagcgctgag 6180
ggcgaggcat gtgagttcac cacgtgagta gcagcaaaag gaaacgaccc ttcttcaccc 6240
ggctatcatc taacgtatcg ccccgggaga atcaataact ttattaacga gatgacgaaa 6300
agtcaaaaaa aaaagtcgtg tgatggccat gatagtcagt caagcaaatc agcgtccaac 6360
acgtgtccct tatctacagg tgtaagagag tagagtcttg tcaatcaacc tgggttgttt 6420
tctatctgcg tttt 6434
<210> 3
<211> 151
<212> DNA
<213> Zea mays L.
<400> 3
aaagagccga agcgaccgac ggcgcaggta cacgcaggcg cctcctgcac cattcaccat 60
tcacgtctct gtggcccaat aataaaaccg cattaattac gctcgcgcag acacggtagc 120
acactggccg ccacagtgcc acccacccac c 151
<210> 4
<211> 361
<212> DNA
<213> Zea mays L.
<400> 4
ctgaggctat ccgcaaccgt taaccctaaa tttttccctc tatatcattt tttcctctat 60
tttcctccct attttttcat ctcccgcagc ggttccccct aaatactccc cctatatctc 120
actaccacta taaaatatta ttttctatac caattatcaa ttttttatct actaacaatt 180
actcgtggac ccacagcaca gtgtttaggg tgatgaacag tgacacgcta gatctgaagg 240
gagagagaag gggaccgaca cgtagggagc ctgtagaggg caccgctgcg gccgtagggt 300
gctccctacg cgccgcatac aaggggaggg gggagaggca gcggtaaccg ctgcgcacag 360
c 361
<210> 5
<211> 363
<212> DNA
<213> Zea mays L.
<400> 5
ctgaggctat ccgcaaccgt taaccctaaa tttttccctc tatatcattt tttcccctat 60
tttcctccct attttttcat ctcccgcagc ggttccccct aaatactccc cctatatccc 120
actaccacta taaaatatta ttttctatac caattatcaa ttttttatct actaacaatt 180
actcgtggac ccacagcaca gtgtttaggg tgatgaacag tgacacgcta gatctgaagg 240
gagagagaag gggaccgaca cgtagggagc ctgtagaggg caccgctgcg gccgtagggt 300
gctccctacg cgccgcatac aaggggaggg gggggagagg cagcggtaac cgctgcgcac 360
agc 363
<210> 6
<211> 20
<212> DNA
<213> Artifical sequence
<400> 6
gctcacgaac cgacaacaat 20
<210> 7
<211> 21
<212> DNA
<213> Artifical sequence
<400> 7
tcccgctcag catctttact a 21
<210> 8
<211> 20
<212> DNA
<213> Artifical sequence
<400> 8
tacgaaacac atcgacggac 20
<210> 9
<211> 20
<212> DNA
<213> Artifical sequence
<400> 9
gttattgatt ctcccggggc 20
<210> 10
<211> 22
<212> DNA
<213> Artifical sequence
<400> 10
gttcttcctc aagacaatca cc 22
<210> 11
<211> 21
<212> DNA
<213> Artifical sequence
<400> 11
gagaagaaga cgcggttgta g 21
<210> 12
<211> 2604
<212> DNA
<213> Zea mays L.
<400> 12
atgctgagcg ggatcatcga cgggctgacg ggggcgaaca agcatgcgcg gctcaagggc 60
acggtggtgc tcatgcgcaa gaacgtgctg gacctcaacg acttcggcgc caccgtcgtt 120
gacagcatca gcgagttcct cggcaagggg gtcacctgcc agctcatcag ctccaccctc 180
gtcgacgcca acaacggcaa ccgcgggcgg gtcggggcgg aggcgaacct ggagcagtgg 240
ctgacgagcc tgccgtcgct gacgaccggc gagtccaagt tcggcgtcac gttcgactgg 300
gaggtggaga agctgggagt gccgggggcc gtcgtcgtca agaacaacca cgccgccgag 360
ttcttcctca agacaatcac cctcgacgac gtgcccggcc gcggcgccgt caccttcgtc 420
gccaactcct gggtctaccc cgcgggcaag taccgctaca accgcgtctt cttctccaac 480
gatacgtacc tgccaagcca gatgccggcg gcgctgaagc cgtaccgcga cgacgagctc 540
cgcaacctcc gcggcgacga ccagcagggc ccctaccagg agcacgaccg cgtgtaccgc 600
tacgacgtct acaacgacct cggcgagccc gacggcggca acccgcgccc catcctcggc 660
ggctccgccg accacccgta cccgcgccgc tgccgcacgg gccgcaagcc caccaaaacc 720
gaccccaact cggagagccg actgtcgctg gtggagcaga tctacgtgcc gcgggacgag 780
cgcttcggcc acctcaagat gtccgacttc ctgggctact ccatcaaggc catcacgcag 840
ggcatcatcc cggcggtgcg cacgtacgtg gacaccaccc cgggcgagtt cgactccttc 900
caggacatca tcaacctgta cgagggcggg atcaagctgc ccaagatcca ggcgctcgag 960
gacatgcgca agctcttccc gctccagctc gtcaaggacc tcctccccgc cggcggggac 1020
tacctgctca agctccccat cccacagatc atccaaggca cgtcacagga caagaacgcg 1080
tggaggaccg acgaggagtt cgcgcgggag gtgctcgccg gcgtcaaccc gatggtgatc 1140
acgcgcctca cggagttccc gcccaagagc acgctggacc ccagcaagta cggcgaccac 1200
accagcacga tcacggcgga gcacatcgag aagaacctcg agggcctcac ggtgcagcag 1260
gcgctggacg gcaacaggct ctacatcctg gaccaccacg accgcttcat gccgttcctc 1320
atcgacgtca acaacctgga gggcaacttc atctacgcca ccaggacgct cttcttcctg 1380
cgcggcgacg gcaggctcgc gcccctcgcc atcgagctca gcgagccgta catcgacggg 1440
gacctcaccg tggccaagag caaggtctac acgccggcgt ccagcggcgt cgaggcctgg 1500
gtgtggcagc tcgccaaggc ctatgtcgcc gtcaacgact ctggctggca ccaactcgtc 1560
agccactggc tgaacaccca cgcggtgatg gagccgttcg tgatcgcgac gaaccggcag 1620
ctgagcgtga cgcacccggt gcacaagctc ctgagctcgc acttccgcga caccatgacc 1680
atcaacgcgc tggcgcggca gacgctcatc aacggcggcg gcatcttcga gatgaccgtc 1740
ttcccgggca agtacgcgct gggcatgtcc tccgtggtgt acaagagctg gaacttcacc 1800
gagcagggcc tccccgccga cctcgtcaag aggggcgtgg cggtggcgga cccgtccagc 1860
ccgtacaagg tgcggctgct gatcgaggac tacccgtacg cgagcgacgg gctggccatc 1920
tggcacgcca tcgagcagtg ggtgggcgag tacctggcca tctactaccc cgacgacggc 1980
gcgctgcggg gcgacgagga gctgcaggcg tggtggaagg aggtgcgcga ggtcgggcac 2040
ggcgaccaca aggacgcgcc ctggtggccc aagatgcagg ccgtgtcgga gctcgccagc 2100
gcctgcacca ccatcatctg gatcgcgtcg gcgctccacg ccgccgtcaa cttcggccag 2160
tacccgtacg cggggtacct cccgaacagg cccacggtga gccggcgccg gatgccggag 2220
cccggcagca aggagtacga ggagctggag cgcgacccgg agcgcggctt catccacacc 2280
atcacgagcc agatccagac catcatcggc atctcgctca tcgagatcct ctccaagcac 2340
tcctccgacg aggtgtacct cggccagcgc gacacccccg agtggacctc cgacgcccgg 2400
gcgctggcgg cgttcaagag gttcagcgac gcgctggtca agatcgaggg caaggtggtg 2460
ggcgagaacc gcgacccgca gctgaggaac aggaacggcc ccgccgagtt cccctacatg 2520
ctgctctatc ccaacacctc tgaccacagt ggcgccgccg cagggctcac tgccaagggc 2580
atccccaaca gcatctccat ctga 2604
<210> 13
<211> 867
<212> PRT
<213> Zea mays L.
<400> 13
Met Leu Ser Gly Ile Ile Asp Gly Leu Thr Gly Ala Asn Lys His Ala
1 5 10 15
Arg Leu Lys Gly Thr Val Val Leu Met Arg Lys Asn Val Leu Asp Leu
20 25 30
Asn Asp Phe Gly Ala Thr Val Val Asp Ser Ile Ser Glu Phe Leu Gly
35 40 45
Lys Gly Val Thr Cys Gln Leu Ile Ser Ser Thr Leu Val Asp Ala Asn
50 55 60
Asn Gly Asn Arg Gly Arg Val Gly Ala Glu Ala Asn Leu Glu Gln Trp
65 70 75 80
Leu Thr Ser Leu Pro Ser Leu Thr Thr Gly Glu Ser Lys Phe Gly Val
85 90 95
Thr Phe Asp Trp Glu Val Glu Lys Leu Gly Val Pro Gly Ala Val Val
100 105 110
Val Lys Asn Asn His Ala Ala Glu Phe Phe Leu Lys Thr Ile Thr Leu
115 120 125
Asp Asp Val Pro Gly Arg Gly Ala Val Thr Phe Val Ala Asn Ser Trp
130 135 140
Val Tyr Pro Ala Gly Lys Tyr Arg Tyr Asn Arg Val Phe Phe Ser Asn
145 150 155 160
Asp Thr Tyr Leu Pro Ser Gln Met Pro Ala Ala Leu Lys Pro Tyr Arg
165 170 175
Asp Asp Glu Leu Arg Asn Leu Arg Gly Asp Asp Gln Gln Gly Pro Tyr
180 185 190
Gln Glu His Asp Arg Val Tyr Arg Tyr Asp Val Tyr Asn Asp Leu Gly
195 200 205
Glu Pro Asp Gly Gly Asn Pro Arg Pro Ile Leu Gly Gly Ser Ala Asp
210 215 220
His Pro Tyr Pro Arg Arg Cys Arg Thr Gly Arg Lys Pro Thr Lys Thr
225 230 235 240
Asp Pro Asn Ser Glu Ser Arg Leu Ser Leu Val Glu Gln Ile Tyr Val
245 250 255
Pro Arg Asp Glu Arg Phe Gly His Leu Lys Met Ser Asp Phe Leu Gly
260 265 270
Tyr Ser Ile Lys Ala Ile Thr Gln Gly Ile Ile Pro Ala Val Arg Thr
275 280 285
Tyr Val Asp Thr Thr Pro Gly Glu Phe Asp Ser Phe Gln Asp Ile Ile
290 295 300
Asn Leu Tyr Glu Gly Gly Ile Lys Leu Pro Lys Ile Gln Ala Leu Glu
305 310 315 320
Asp Met Arg Lys Leu Phe Pro Leu Gln Leu Val Lys Asp Leu Leu Pro
325 330 335
Ala Gly Gly Asp Tyr Leu Leu Lys Leu Pro Ile Pro Gln Ile Ile Gln
340 345 350
Gly Thr Ser Gln Asp Lys Asn Ala Trp Arg Thr Asp Glu Glu Phe Ala
355 360 365
Arg Glu Val Leu Ala Gly Val Asn Pro Met Val Ile Thr Arg Leu Thr
370 375 380
Glu Phe Pro Pro Lys Ser Thr Leu Asp Pro Ser Lys Tyr Gly Asp His
385 390 395 400
Thr Ser Thr Ile Thr Ala Glu His Ile Glu Lys Asn Leu Glu Gly Leu
405 410 415
Thr Val Gln Gln Ala Leu Asp Gly Asn Arg Leu Tyr Ile Leu Asp His
420 425 430
His Asp Arg Phe Met Pro Phe Leu Ile Asp Val Asn Asn Leu Glu Gly
435 440 445
Asn Phe Ile Tyr Ala Thr Arg Thr Leu Phe Phe Leu Arg Gly Asp Gly
450 455 460
Arg Leu Ala Pro Leu Ala Ile Glu Leu Ser Glu Pro Tyr Ile Asp Gly
465 470 475 480
Asp Leu Thr Val Ala Lys Ser Lys Val Tyr Thr Pro Ala Ser Ser Gly
485 490 495
Val Glu Ala Trp Val Trp Gln Leu Ala Lys Ala Tyr Val Ala Val Asn
500 505 510
Asp Ser Gly Trp His Gln Leu Val Ser His Trp Leu Asn Thr His Ala
515 520 525
Val Met Glu Pro Phe Val Ile Ala Thr Asn Arg Gln Leu Ser Val Thr
530 535 540
His Pro Val His Lys Leu Leu Ser Ser His Phe Arg Asp Thr Met Thr
545 550 555 560
Ile Asn Ala Leu Ala Arg Gln Thr Leu Ile Asn Gly Gly Gly Ile Phe
565 570 575
Glu Met Thr Val Phe Pro Gly Lys Tyr Ala Leu Gly Met Ser Ser Val
580 585 590
Val Tyr Lys Ser Trp Asn Phe Thr Glu Gln Gly Leu Pro Ala Asp Leu
595 600 605
Val Lys Arg Gly Val Ala Val Ala Asp Pro Ser Ser Pro Tyr Lys Val
610 615 620
Arg Leu Leu Ile Glu Asp Tyr Pro Tyr Ala Ser Asp Gly Leu Ala Ile
625 630 635 640
Trp His Ala Ile Glu Gln Trp Val Gly Glu Tyr Leu Ala Ile Tyr Tyr
645 650 655
Pro Asp Asp Gly Ala Leu Arg Gly Asp Glu Glu Leu Gln Ala Trp Trp
660 665 670
Lys Glu Val Arg Glu Val Gly His Gly Asp His Lys Asp Ala Pro Trp
675 680 685
Trp Pro Lys Met Gln Ala Val Ser Glu Leu Ala Ser Ala Cys Thr Thr
690 695 700
Ile Ile Trp Ile Ala Ser Ala Leu His Ala Ala Val Asn Phe Gly Gln
705 710 715 720
Tyr Pro Tyr Ala Gly Tyr Leu Pro Asn Arg Pro Thr Val Ser Arg Arg
725 730 735
Arg Met Pro Glu Pro Gly Ser Lys Glu Tyr Glu Glu Leu Glu Arg Asp
740 745 750
Pro Glu Arg Gly Phe Ile His Thr Ile Thr Ser Gln Ile Gln Thr Ile
755 760 765
Ile Gly Ile Ser Leu Ile Glu Ile Leu Ser Lys His Ser Ser Asp Glu
770 775 780
Val Tyr Leu Gly Gln Arg Asp Thr Pro Glu Trp Thr Ser Asp Ala Arg
785 790 795 800
Ala Leu Ala Ala Phe Lys Arg Phe Ser Asp Ala Leu Val Lys Ile Glu
805 810 815
Gly Lys Val Val Gly Glu Asn Arg Asp Pro Gln Leu Arg Asn Arg Asn
820 825 830
Gly Pro Ala Glu Phe Pro Tyr Met Leu Leu Tyr Pro Asn Thr Ser Asp
835 840 845
His Ser Gly Ala Ala Ala Gly Leu Thr Ala Lys Gly Ile Pro Asn Ser
850 855 860
Ile Ser Ile
865
<210> 14
<211> 6642
<212> DNA
<213> Zea mays L.
<400> 14
ggcgaactcc gctccgcccg accccagggc tcggactcgg gctaacaccc ggaagacggc 60
gaactccgct ccgcccgacc ccagggctcg gactcgggct aagacccgga agacggcgaa 120
ctctgctccg cccgacccca gggctcggac tcgggctaag acccggaaga cggcgaatct 180
ccgcctcgcc cgaccccagg gctcagactc cgccctggcc tcggccaaac gatctccgcc 240
tcgcccgacc ccagggctcg gactccgccc tagcctcggc caaacgatct ccgcctcgcc 300
cgacccgggg gctcgggctc ggcctcggca acggaaggca gactcgacct cgacttcgga 360
ggagccccca cgtcgccctg cctagggcac aggtccgcca cgtcaacagg aagcgccatc 420
accaacctac cccgagccga cttgggacac gaaggacaag accggcgtcc catctggcca 480
gctccgccgg atgggcaatg atggcgcccc ccgagctctg tgacgacggc ggctcttagc 540
tctcttacgg cagcagagcg acgtcagcaa ggactcgacc gctccaacag ctgtccctcc 600
gccaggctcc gtcgctcctc cgacagccac gacatcacgc cagcaaggtg ccaagacctc 660
tccggctgcc acattggcat gtacccaggg cgttagctct ctctctctcc gctagacacg 720
tagcactctg ctaccccccg ttgtacacct ggatcctctc cttacgacta taaaaggaag 780
gaccagggcc ttcttagaga aggttggccg cgcgggaccg aggacgggac aggcgctctc 840
ttggggccgc tcgcttccct cacccgcgtg gacgcttgta acccccctac tgcaagcgca 900
cctgacctgg gcgcgggacg aacacgaagg ccgcgggact tccacctctc tcacgctcga 960
ctccggccac ctcgcctctc cccccttcgc gctcgcccac gcgctcgacc catctgggct 1020
ggggcacgca gcacactcac tcgtcggctt agggaccccc ctgtctcgaa acgccgacaa 1080
taagcttatc tcgaattcat ggggtggagg attggaaatg atttttatgt attagtagaa 1140
tttgtttcta ctctgtaaat tacatgaccc tcttcgtctc actcctctat agtaaaaata 1200
tagcacataa atatctccga catcttgcta ataatagtat acaaatatat ttttcatcaa 1260
accgaattaa cttaattgat atatgtctaa attactgtta ttagaatgga attcaattcc 1320
aatgaaccaa acggggcgta agtgatttct tagtgaggtg gcacctatgg aaatcactta 1380
gtagtttctg ggttgtaatt gctcttatgg catgaagaaa attttgcatt cgagctcgaa 1440
cgctacaagg tcgttttaga agctacatca gtaatgcttc gagtatgaca aacaacatca 1500
caagtaacat ctacctatat gcaccgaatc tccaccacca ccacaatatc gatgtcagaa 1560
ggctagggaa ggacgaccaa taatatagaa gacacatagg ccacactcaa ttcaatcaac 1620
atcacgacaa tctacttgat gtcctatgta gccatgatct ctacgagcaa tcaatatcaa 1680
tgcactagaa tcccaactac gtagagtatt attggacgaa cgagtaaaac accaaagagc 1740
agccactgga cgactttctt gcatctccac aaggataaat tctctcagat agtcagatgc 1800
ctaatgaaat tcaagtccat caatattgac atgtgcgaag gcaagaaggg tccaaattag 1860
ttgattcgac actacttggt taccatcaat ctcaagggag gaaacaaaga tatcaaagca 1920
tcatgctttc aaatgtactc aaagctactc tagtgacttg gcttgaatag ctcccaagag 1980
gctacatcga taacaggcta gcactctcca aggaattaat ggacaacttc caaagtgcat 2040
ggcctcgttc ggacaaccgg tacaatcttc aaaatggaag tagaagattg gtgaaagcat 2100
atgcaactag tacaagctct tgatcgaaat ttgatgctac aggcctaact acttcgacga 2160
tgatcatggg gaaaagttct agtatcagaa tttcataaaa acaaaccaaa aacaattcat 2220
gagttctaac acatgatcaa caactgaatg aatgtcgaag acacagtacg agataagttt 2280
gacaattgaa accgtagata taacaataaa gataacattc atagagacaa caagggcagc 2340
ttatcggata agaagtgagg gaaagataac actatgataa gcactcgcat caaagaaggg 2400
tggcaagaac aaatcattca attagcaact ctttgagcaa cacccatacc acctacatca 2460
taaatcataa gtgctctacc actaaatgca tgcaactcaa gaactttggc atcatcttag 2520
ctcggaagca agacaaaacc aaacagagtt gataaaaaat attaggaaaa atagaataat 2580
ggagacttct agagggccac aaacatagtt aacctcatct tcaaaggagc atcttcctta 2640
atacctaaga ggaagactaa acttaccctt ttgtgagatg atggttgttg aaccatcacc 2700
tacaacatat atgtgatggt tcgagatccc aattcaattt ataagggagg attagtttct 2760
caaacatagg atgctatggt ctagtcctgg ccccgatggt ggcaggactt caacttagta 2820
aggaaggtga attaccaata tgttcaaaag gatggggtaa aaggaaggtg aattatgcaa 2880
tctatttttc gcactttttc attaatcaaa acctatatgg ataaccaatc gttcatatgt 2940
gcaactaagg ttttgactaa gtgttgctat ctctaccgta aaaggagttt tgctacccca 3000
aacctatcaa ctagtctatg actaagctaa gaagataaat cacacaacca caagtaacaa 3060
tataaatgtg gaattttaaa tatggtagag atacaaactc tcgttgatgt gtcagtattt 3120
ttactggagt atcaagaaac gcgcaagctt cttactaatc cttcttagag cctcacgcaa 3180
ggctaagctc ccgctcaggt aaccctgtat agatacaaac cctaccaatg actatagtaa 3240
cttttatgaa gatataggga aacacacaag ttttacccta gttctcgttg gagcctctta 3300
taaacatgtc cacaagggga tgaagctaag agtcgagtaa gggtttgttt gatttcttta 3360
gtcttaatga ctaaaactaa gcaaagagct ttaattattg tgcaaattga ttacattatc 3420
cctaattaat gctatcttcg actactgttt acccacacgc agagctactg ttcgtgcgca 3480
ttcacaatgc tgcatgcgtg cacgttgctt tggggaggag cctacttcat gcgcgttggt 3540
tagggagggc atatgaagta aaaatgtcac tttatggtta gtttgacacc ctcatttttt 3600
tcaagggatt gtattttcac aaaagaaatt aatttatttt tcttgaaaaa taggaatccc 3660
ttagaaaaaa atagagttgt caaactaacc cttattcatt ttagtcactc ttttggtaat 3720
tagaggacta taatttagtt ggaggatttt agtcacactg tgttgattct ttagtgacta 3780
aaaatgacta aaatttaatc aattaaatgt agtcacccaa accaaacaga gtcgcgcgtc 3840
cgacgtgcct cctcaaccgg cgcgtgggag taggtctgga aaaaagctcg agctcgatga 3900
gttggcttgg gctcacggta acttgggtcg gctcgacgct caaaacgagt ccaagtccta 3960
tttttgtggc tcgtgataag tgtgagctag ctcggctcgg ctcacgaacc gacaacaatt 4020
tactacataa aattctgatt agcatataat gttagtacca aatatacaat tagtatatgt 4080
tatttgatat ttatatacaa aattaattta tttctatgtt atattagtac tataattatt 4140
tattaattta agagatgtaa tttgattatt tttgatattt tatgttataa tttgtaacgt 4200
aagctggtgc ttgtggttag actctctact agtcgagttt tcacgctcgt caaaggaccg 4260
agccgagacg aggcaagctc aacacattac cgagccgacc ggctcgtttc cgccacgtgg 4320
tagagcggca gcacccttcc cagatctcgt gtcccgcgcg ccgtgcgcac agccgcgcac 4380
acacacggac tgccgcgtcc gggcgcgatc cgttctcgcg aacacaccgg tcgtagccgt 4440
ccgtaatttg tactagtatg gcacgcggaa cacgagtcgg cgccagcacc ggcacgtcac 4500
catcggggcc gctggcccca tacgggcagc caattaataa atccttgctc tgcgacagac 4560
gcgcaggggc acgtcacgca gccgtagtcg atcggtgcac ggtgctcgcg cccccgttgc 4620
agcgcgcagc tctcccgcgc tataaatgcc ccggctcggc ctcgctccca cagccacagc 4680
ctcacacaga caccaacgcc actgcactgc aaaagcaaga gcagctagct agtaaagatg 4740
ctgagcggga tcatcgacgg gctgacgggg gcgaacaagc atgcgcggct caagggcacg 4800
gtggtgctca tgcgcaagaa cgtgctggac ctcaacgact tcggcgccac cgtcgttgac 4860
agcatcagcg agttcctcgg caagggggtc acctgccagc tcatcagctc caccctcgtc 4920
gacgccagtg agtaccgcgc cgcgccgccg gcacctctcc gatctgcgct tccccatgtc 4980
gatcgatctc gatctctcta ggctctagag ctatagctct ctcggccccc actttttacc 5040
tttgcaagca ttttccctgc atgcgaaaca agcgatagtt tactatttgg gcggccatgc 5100
tgctgctgct tcggctacct tgcctccgtc atctttgacg ggacatggaa agaaagaaag 5160
aatagagaga gagacagaga gagagagaga gcaaacaccg agaaaaagac agcaaagcta 5220
gttgtagcct gggcgcagaa cacagcacca gatgctggct agctcgtgac aaagtaaaaa 5280
aagagaggaa cgaacacagt agtaccaaga gatcagggac gagacctttc tactttgaac 5340
tggattatat attggattct ttatcagtaa cttactgctg ctagtatacc cctaccctag 5400
tctcggggcg acgtgcctgc gtgcatgccc gacgcgtacg aaacacatcg acggactcac 5460
atgggccacc gcgcgcgcgc gtgcctgctt taactttcgc tgtgcagaca acggcaaccg 5520
cgggcgggtc ggggcggagg cgaacctgga gcagtggctg acgagcctgc cgtcgctgac 5580
gaccggcgag tccaagttcg gcgtcacgtt cgactgggag gtggagaagc tgggagtgcc 5640
gggggccgtc gtcgtcaaga acaaccacgc cgccgagttc ttcctcaaga ccatcaccct 5700
cgacgacgtg cccggccgcg gcgccgtcac cttcgtcgcc aactcctggg tctaccccgc 5760
gggcaagtac cgctacaacc gcgtcttctt ctccaacgat gtgagtcctt tctcgataga 5820
tcattatgtt tgtttgttta ttggtatcat gtagctagcc gccatgtcgt cagttggagt 5880
gcagtaggta ggaaaaagga cgacatggga tgggagtggt taagaaaatc catgcaagtg 5940
ggactagtgt gtaactggta gtatagctga agaatctagt ggtagaatga tcttgtacgt 6000
gaataatgtt tctgacgctg agcgctgagg ctatccgcaa ccgttaaccc taaatttttc 6060
cctctatatc attttttccc ctattttcct ccctattttt tcatctcccg cagcggttcc 6120
ccctaaatac tccccctata tcccactacc actataaaat attattttct ataccaatta 6180
tcaatttttt atctactaac aattactcgt ggacccacag cacagtgttt agggtgatga 6240
acagtgacac gctagatctg aagggagaga gaaggggacc gacacgtagg gagcctgtag 6300
agggcaccgc tgcggccgta gggtgctccc tacgcgccgc atacaagggg agggggggga 6360
gaggcagcgg taaccgctgc gcacagcctg agggcgaggc atgtgagttc accacgtgag 6420
tagcagcaaa aggaaacaac ccttcttcac ccggctatca tctaacgtat cgccccggga 6480
gaatcaataa ctttaacgag atgacgaaaa gtcaaaaata aagtcgtgtg atggccatga 6540
aagtcagtca agcaaatcag ctgctaacac gtgtccctta tctacaggtg taagagagta 6600
gagtcttgtc aatcaacctg ggttgttttc tatctgcgtt tt 6642
Claims (10)
1.一种调控玉米穗腐病抗性的关键DNA序列，其特征在于，其多核苷酸为(a)、(b)、(c)或(d)所示：
(a)SEQ ID No.1所示的多核苷酸序列；(b)与SEQ ID No.1的互补序列在严谨杂交条件能够进行杂交的多核苷酸；(c)与SEQ ID No.1所示的多核苷酸至少有90％或以上同源性的多核苷酸；或(d)在SEQ ID No.1所示的多核苷酸的基础上进行一个或多个碱基的缺失、取代或***的突变体，且该突变体仍具有调控玉米穗腐病的功能或活性。
2.权利要求1所述的关键DNA序列的突变体，其特征在于，其多核苷酸序列为SEQ IDNo.2或SEQ ID No.14所示。
3.权利要求1所述的关键DNA序列或权利要求2所述的关键DNA序列的突变体在调控ZmLOX3基因在玉米中表达量或改良玉米穗腐病抗性中的用途。
4.检测权利要求1所述的关键DNA序列的特异性引物；优选的，所述的特异性引物的核苷酸序列分别为SEQ ID No.10和SEQ ID No.11所示
5.一种降低玉米发生穗腐病风险或提高穗腐病抗性的方法，包括：在ZmLOX3基因启动子区Indel151缺失SEQ ID No.3所示的151bp的片段；或者在ZmLOX3基因第二个内含子区Indel363处***SEQ ID No.4或SEQ ID No.5所示的碱基片段；或者在ZmLOX3基因启动子区Indel151缺失SEQ ID No.3所示的151bp的片段以及在ZmLOX3基因第二个内含子区Indel363处***SEQ ID No.4或SEQ ID No.5所示的碱基片段。
6.一种培育抗穗腐病的玉米新品种的方法，其特征在于，包括降低ZmLOX3基因在玉米果穗和籽粒中的表达量，从而提高玉米穗腐病的抗性；
优选的，所述降低ZmLOX3基因在玉米果穗和籽粒中的表达量的方法包括：在ZmLOX3基因启动子区Indel151缺失SEQ ID No.3所示的151bp的片段；或者在ZmLOX3基因第二个内含子区Indel363处***SEQ ID No.4或SEQ ID No.5所示的碱基片段；或者在ZmLOX3基因启动子区Indel151缺失SEQ ID No.3所示的151bp的片段以及在ZmLOX3基因第二个内含子区Indel363处***SEQ ID No.4或SEQ ID No.5所示的碱基片段。
7.Indel151或Indel363作为ZmLOX3基因在玉米果穗和籽粒中表达的分子标记的用途；其中，所述Indel151的DNA序列为SEQ ID No.3所示，所述Indel363的DNA序列为SEQ IDNo.4或SEQ ID No.5所示。
8.Indel151或Indel363在调控ZmLOX3基因在玉米表达量、或者在调控玉米抗穗腐抗性、培育抗穗腐病玉米新品种中的用途；其中，所述Indel151的DNA序列为SEQ ID No.3所示，所述Indel363的DNA序列为SEQ ID No.4或SEQ ID No.5所示。
9.按照权利要求8所述的用途，其特征在于，所述的调控ZmLOX3基因在玉米表达量是降低ZmLOX3基因在玉米中的表达量；所述调控玉米抗穗腐抗性是提高玉米抗穗腐抗性。
10.用于检测Indel151或Indel363变异情况的检测引物；其中，所述Indel151的DNA序列为SEQ ID No.3所示，所述Indel363的DNA序列为SEQ ID No.4或SEQ ID No.5所示；优选的，所述的检测Indel151变异情况的检测引物的核苷酸序列分别为SEQ ID No.6和SEQ IDNo.7所示；所述的检测Indel363变异情况的检测引物的核苷酸序列分别为SEQ ID No.8和SEQ ID No.9所示。
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