CN106467908B - 耐受除草剂的植物及其应用 - Google Patents

Abstract

本发明耐受除草剂的植物及其应用，提供了使植物具有除草剂抗性的ALS蛋白质，和相应的核酸、表达盒、载体、细胞、植物、获得具有除草剂抗性的植物的应用和方法、以及鉴定本发明的植物的方法。

Description

耐受除草剂的植物及其应用

技术领域

本发明属于植物除草剂抗性领域，具体而言，本发明涉及能赋予植物(尤其是小麦)除草剂抗性(耐受性)的蛋白及其在植物育种和杂草管理中的应用。

背景技术

在农业生产中，农田杂草是影响作物产量的最重要因素之一，杂草会与作物争夺阳光、肥料、水分和空间，降低作物的产量和品质，孳生病虫害，而且会妨碍农事操作。除草剂的应用是农业上的一项现代先进技术，为防除农田杂草起到了极大的推动作用。但是，由于作物和部分农田杂草的亲缘关系非常近，施用除草剂如何才能做到选择性地杀死杂草而不伤害农作物，已经成为除草剂应用中所面临的一个重要问题。针对这一现状，发展抗除草剂作物新品种将是最方便、经济、有效地控制田间杂草的措施。

目前发展抗除草剂作物的途径主要有两种：一种是通过转基因手段将抗除草剂基因转入作物中，最成功的案例是美国孟山都公司开发的一系列转基因抗草甘膦除草剂作物品种农达(Roundup)系列，包括抗农达大豆、玉米、棉花、油菜、向日葵和甜菜。但是利用转基因基因技术培育的抗除草剂的作物品种需要经过较长时间的安全性评价，而且，从品种选育成功到生产应用的周期较长。第二种是利用遗传学非转基因的手段筛选作物抗除草剂突变体。该方法代表了目前国际抗除草剂作物培育的新方向。首先通过遗传诱变的方法构建突变体库，从中筛选到抗除草剂突变体，这样得到的突变体可以直接作为抗除草剂新品种加以利用，或通过回交转育的方法杂交到其他栽培品种。其优点是非转基因作物，不需要经过安全评估直接应用，产业化所需的批准时间短。因此我们选择了后者作为获得抗除草剂作物的方法。

乙酰羟基合成酶(AHAS，EC2.2.1.6)或者叫乙酰乳酸合成酶(ALS)是支链氨基酸(缬氨酸、亮氨酸和异亮氨酸)生物合成途径的第一个酶，也是几种重要除草剂的靶位点，主要包括：咪唑啉酮类除草剂、磺酰脲类除草剂、三唑并嘧啶类除草剂、嘧啶基氧基苯甲酸酯类除草剂和磺酰基氨基羧基三唑啉酮类除草剂。这五类除草剂与其靶位点乙酰羟基合成酶结合后，能够抑制蛋白活性，从而阻止易感植物的生长发育。在模式植物拟南芥中的研究表明，AHAS蛋白某些氨基酸突变后能够使拟南芥获得除草剂抗性，突变后的蛋白不再与除草剂结合，同时又保持原有的生物学活性。

正是基于以上原理，本发明人通过长期而艰苦的研究实践，在冬小麦矮抗58和济麦22中构建了突变体库，并从中筛选到一系列抗除草剂突变体，并鉴定出赋予其除草剂抗性的蛋白。本发明人还开发了这些蛋白及其编码基因在转基因或者非转基因植物育种中的应用，可用于培育具有除草剂抗性的植物，尤其是农作物，如抗除草剂小麦等。

发明内容

本发明提供了一种来源于小麦的IMI核酸序列，所述IMI核酸序列具有咪唑啉酮类除草剂抗性和磺酰脲类除草剂抗性，所述IMI基因的核苷酸序列或氨基酸序列在植物中表达时能赋予植物增加的对咪唑啉酮类除草剂和/或磺酰脲类除草剂的耐受性。在本发明中，所述IMI核苷酸序列的编号及其对应的核苷酸和氨基酸序列分别是：抗性基因IMI1-286，其核苷酸序列如SEQ ID NO:1所示，其编码的氨基酸序列如SEQ ID NO:2所示；抗性基因IMI1-512，其核苷酸序列如SEQ ID NO:3所示，其编码的氨基酸序列如SEQ ID NO:4所示；抗性基因IMI1-536，其核苷酸序列如SEQ ID NO:5所示，其编码的氨基酸序列如SEQ ID NO:6所示；抗性基因IMI1-1880，其核苷酸序列如SEQ ID NO:7所示，其编码的氨基酸序列如SEQ IDNO:8所示；抗性基因IMI2-292，其核苷酸序列如SEQ ID NO:9所示，其编码的氨基酸序列如SEQ ID NO:10所示；抗性基因IMI2-518，其核苷酸序列如SEQ ID NO:11所示，其编码的氨基酸序列如SEQ ID NO:12所示；抗性基因IMI2-542，其核苷酸序列如SEQ ID NO:13所示，其编码的氨基酸序列如SEQ ID NO:14所示；抗性基因IMI2-1886，其核苷酸序列如SEQ ID NO:15所示，其编码的氨基酸序列如SEQ ID NO:16所示；抗性基因IMI3-295，其核苷酸序列如SEQID NO:17所示，其编码的氨基酸序列如SEQ ID NO:18所示；抗性基因IMI3-521，其核苷酸序列如SEQ ID NO:19所示，其编码的氨基酸序列如SEQ ID NO:20所示；抗性基因IMI3-545，其核苷酸序列如SEQ ID NO:21所示，其编码的氨基酸序列如SEQ ID NO:22所示；抗性基因IMI3-1889，其核苷酸序列如SEQ ID NO:23所示，其编码的氨基酸序列如SEQ ID NO:24所示。

本发明提供了一种用于增强植物、植物组织、植物细胞或其他宿主细胞对于至少一种除草剂的耐受性或抗性的方法，所述除草剂干扰ALS酶的活性，更具体的所述除草剂包括但不限于磺酰脲类、咪唑啉酮类、嘧啶并三唑类、水杨酸嘧啶类等除草剂(PeterBabczinski,Thomas Zelinski.Mode of action herbicidal ALS-inhibitors onacetolactate synthase from green plant cell cultures，yeast，and Escherichiacoli.Pesticide science,1991,31:305-323；郑培忠,沈健英.乙酰乳酸合成酶抑制剂的种类及其耐药性研究进展.杂草科学,2009,2:4-8)。因此，在本文中，除草剂抗性指的是对包括磺酰脲类、咪唑啉酮类、嘧啶并三唑类、水杨酸嘧啶类在内的除草剂具有耐受性。优选在本发明中，除草剂是咪唑啉酮类除草剂，如在本发明的具体实施方式中，除草剂是咪草烟(其化学名称为(RS)5-乙基-2-(4-异丙基-4-甲基-5-氧代-3-咪唑啉-2-基)烟酸，CAS号为81335-77-5)。即优选在本发明中，除草剂抗性是咪唑啉酮类除草剂抗性，更优选是咪草烟抗性。

本发明所述的增强植物、植物组织、植物细胞或其他宿主细胞对至少一种除草剂的耐受性或抗性的方法，可以通过转化的方法实施，或是杂交、回交、自交或无性繁殖等方式实施，也可以通过化学诱变或基因定点突变的方式获得，使改变后的植株具有本发明所提供的核苷酸序列SEQ ID NO:1、SEQ ID NO:3、SEQ ID NO:5、SEQ ID NO:7、SEQ ID NO:9、SEQ ID NO:11、SEQ ID NO:13、SEQ ID NO:15、SEQ ID NO:17、SEQ ID NO:19、SEQ ID NO:21和/或SEQ ID NO:23。更具体地，所述突变后的植株可以具有一种或一种以上的上述突变后的核苷酸序列。

转化的方法是指通过农杆菌介导的方法，或基因枪法，将本发明所提供的IMI核苷酸序列SEQ ID NO:1、SEQ ID NO:3、SEQ ID NO:5、SEQ ID NO:7、SEQ ID NO:9、SEQ ID NO:11、SEQ ID NO:13、SEQ ID NO:15、SEQ ID NO:17、SEQ ID NO:19、SEQ ID NO:21和/或SEQID NO:23转入到植物、植物组织、植物细胞或其他宿主细胞中，以增强其对除草剂的耐受性或抗性。化学诱变的方法包括用EMS等诱变剂处理所导致的诱变；基因定点突变的方法包括但不限于ZFN定点突变方法、TALEN定点突变方法、和/或CRISPR/Cas9等定点突变方法。

本发明还提供了一种提高小麦杂交种子纯度的方法，包括：通过转化的方法、化学诱变、回交转育或定点突变的方法，使得小麦具有本发明所提供的核苷酸序列SEQ ID NO:1、SEQ ID NO:3、SEQ ID NO:5、SEQ ID NO:7、SEQ ID NO:9、SEQ ID NO:11、SEQ ID NO:13、SEQ ID NO:15、SEQ ID NO:17、SEQ ID NO:19、SEQ ID NO:21和/或SEQ ID NO:23，并且同时具有咪唑啉酮类除草剂和/或磺酰脲类除草剂的抗性或耐受性，将该小麦材料当做父本材料，或是将该材料的除草剂抗性回交转育到其他父本材料中，再将父本材料与母本材料进行杂交，产生的杂交种子均具有除草剂抗性，而母本材料自己结实的种子由于不具备除草剂抗性，将所有结出的种子在浸种阶段用咪唑啉酮类除草剂和/或磺酰脲类除草剂溶液进行浸泡，则可以杀死假的杂交种子，或是在秧苗期，对小苗进行除草剂的喷施，杀死非杂交产生的种子秧苗，从而提高杂交种子生产的制种纯度，降低杂交种子的种植风险。

本发明还提供了一种表达盒(cassette)，所述表达盒包含本发明所提供的核酸序列SEQ ID NO:1、SEQ ID NO:3、SEQ ID NO:5、SEQ ID NO:7、SEQ ID NO:9、SEQ ID NO:11、SEQ ID NO:13、SEQ ID NO:15、SEQ ID NO:17、SEQ ID NO:19、SEQ ID NO:21和/或SEQ IDNO:23。所述表达盒中的核酸序列由一个启动子驱动表达，所述启动子包括但不限于CaMV35S启动子或者稻肌动蛋白I基因、玉米醇脱氢酶基因、玉米矮缩I基因或烟草花叶病毒(TMV)Ω元件的启动子。进一步的，本发明所提供的表达盒还可以包括，在细菌中复制所需的复制起点(例如，来自pBR322或P15A ori的ORI区)，土壤杆菌T-DNA转移所需要的元件(例如，T-DNA的左边界和/或右边界)，从而方便从细菌往植物中的转化过程。另外，本发明所述的表达盒还可以包含增强子、内含子、多克隆位点、操纵基因、阻遏物结合位点、转录因子结合位点等，例如来自TMV、玉米褪绿斑点病毒(MCMV)或苜蓿花叶病毒(AMV)的前导序列等作为增强子，又如可以包括来自Adh 1、bronze 1、肌动蛋白1或肌动蛋白2的内含子。

本发明还提供了一种载体，所述载体包含本发明上述所提供的含有SEQ ID NO:1、SEQ ID NO:3、SEQ ID NO:5、SEQ ID NO:7、SEQ ID NO:9、SEQ ID NO:11、SEQ ID NO:13、SEQID NO:15、SEQ ID NO:17、SEQ ID NO:19、SEQ ID NO:21和/或SEQ ID NO:23的表达盒。在本文中，载体是指本领域中常用的细菌质粒、粘粒、噬菌粒、酵母质粒、植物细胞病毒、动物病毒及其它各种病毒载体。根据所应用的目的不同，载体可以分为克隆载体、表达载体和转化载体，指的是所使用的目的分别针对克隆并验证基因、表达相应基因和将相应基因转化。本发明中可用的载体包括但不限于：在细菌中表达用的载体(原核表达载体)、在酵母中表达用的载体(如毕赤酵母载体)、在昆虫细胞中表达的杆状病毒载体、在哺乳动物细胞中表达用的载体(痘苗病毒载体、逆转录病毒载体、腺病毒载体、腺伴病毒载体等)、在植物中表达用的植物病毒载体以及在哺乳动物乳腺中表达用的各种载体。本发明所述的载体是转化表达载体，尤其是植物转化载体，特别是适合农杆菌转化植物的载体。这些载体很多已经商品化了。

本发明还提供了一种细胞，所述细胞包含本发明所提供的包含SEQ ID NO:1、SEQID NO:3、SEQ ID NO:5、SEQ ID NO:7、SEQ ID NO:9、SEQ ID NO:11、SEQ ID NO:13、SEQ IDNO:15、SEQ ID NO:17、SEQ ID NO:19、SEQ ID NO:21和/或SEQ ID NO:23序列的表达盒或载体。细胞可以是原核细胞，也可以是真核细胞，如，细菌细胞、酵母细胞、植物细胞、昆虫细胞、哺乳动物细胞等。在本文中，植物细胞指的是来自植物的细胞(可以包括细胞群，但优选是单个细胞)，但是这些细胞不能仅仅借助光合作用而以水、二氧化碳和无机盐等无机物合成碳水化合物、蛋白质来发育成存活的单个植株。植物细胞可以是必须使用有机物(如某些植物激素)才能发育的植物细胞或细胞群，也可以是完全无发育成植株能力的植物细胞或细胞群，通俗地说，即死亡的植物细胞或细胞群，如干燥粉碎过的市售小麦面等。

本发明中所述的植物可以是双子叶植物，也可以是单子叶植物。示例性的植物包括小麦、水稻、玉米、黑麦、燕麦、大麦、大豆、马铃薯、油菜、甜菜、甘蔗、高粱、西红柿、南瓜、辣椒、生菜、向日葵、咖啡、茶、棉花、苜蓿、烟草、或拟南芥等。在本发明的具体实施方式中，优选本发明的植物是小麦，更优选是宁春4号。其中，宁春4号是我国在1980年代初期就在宁夏培育出的非转基因小麦品种，已经可以在我国推广使用了30年，是我国解放以来累计种植面积最大的小麦品种，其品种审定编号为宁审种8101。本发明优选的植物，可以继承了宁春4号丰产性突出、适应性广泛、品质优良等特性，并具有除草剂抗性，尤其具有咪唑啉酮类除草剂抗性，特别是咪草烟抗性。

本发明还提供了本发明上述方法或核苷酸序列、氨基酸序列在获得具有除草剂抗性的植物过程中的应用。在获得具有除草剂抗性的植物过程中，可以应用到本发明前述各方面所述的方法中。本发明所提供的还包括，本发明所提供的蛋白质SEQ ID NO:2、SEQ IDNO:4、SEQ ID NO:6、SEQ ID NO:8、SEQ ID NO:10、SEQ ID NO:12、SEQ ID NO:14、SEQ IDNO:16、SEQ ID NO:18、SEQ ID NO:20、SEQ ID NO:22和/或SEQ ID NO:24在获得具有除草剂抗性的植物过程中的应用，本发明所提供的核酸SEQ ID NO:1、SEQ ID NO:3、SEQ ID NO:5、SEQ ID NO:7、SEQ ID NO:9、SEQ ID NO:11、SEQ ID NO:13、SEQ ID NO:15、SEQ ID NO:17、SEQ ID NO:19、SEQ ID NO:21和/或SEQ ID NO:23在获得具有除草剂抗性的植物过程中的应用，本发明所提供的表达盒在获得具有除草剂抗性的植物过程中的应用，本发明所提供的载体在获得具有除草剂抗性的植物过程中的应用，本发明所提供的细胞在获得具有除草剂抗性的植物过程中的应用，本发明所提供的植物在获得具有除草剂抗性的植物过程中的应用。优选在本发明中，除草剂是咪唑啉酮类除草剂和磺酰脲类除草剂，优选是咪草烟。

在本文中，获得包括制备、培育、生产或以其他方式产生得到，包括通过转基因育种方式获得，如将本发明所提供的核酸SEQ ID NO:1、SEQ ID NO:3、SEQ ID NO:5、SEQ IDNO:7、SEQ ID NO:9、SEQ ID NO:11、SEQ ID NO:13、SEQ ID NO:15、SEQ ID NO:17、SEQ IDNO:19、SEQ ID NO:21和/或SEQ ID NO:23转化入植物后使之表达本发明的蛋白质；也包括通过非转基因育种方式获得，如通过杂交、回交、自交或无性繁殖本发明所获得的植物并分选出仍旧包含本发明所提供的核酸并表达本发明所提供的蛋白质的植物。因此，本发明还提供了一种应用，包括：在获得具有除草剂抗性的转基因植物过程中的应用和在获得具有除草剂抗性的非转基因植物过程中的应用。

本发明还提供了获得具有除草剂抗性的植物的方法，其包括如下步骤：

(1)使植物包含本发明所提供的核酸SEQ ID NO:1、SEQ ID NO:3、SEQ ID NO:5、SEQ ID NO:7、SEQ ID NO:9、SEQ ID NO:11、SEQ ID NO:13、SEQ ID NO:15、SEQ ID NO:17、SEQ ID NO:19、SEQ ID NO:21和/或SEQ ID NO:23；和/或，

(2)使植物表达本发明所提供的蛋白质SEQ ID NO:2、SEQ ID NO:4、SEQ ID NO:6、SEQ ID NO:8、SEQ ID NO:10、SEQ ID NO:12、SEQ ID NO:14、SEQ ID NO:16、SEQ ID NO:18、SEQ ID NO:20、SEQ ID NO:22和/或SEQ ID NO:24。

此外，本发明还提供了一种鉴定本植物中是否含有本发明核苷酸序列的方法，其包括如下步骤：

(1)测定所述植物是否包含本发明所提供的核酸SEQ ID NO:1、SEQ ID NO:3、SEQID NO:5、SEQ ID NO:7、SEQ ID NO:9、SEQ ID NO:11、SEQ ID NO:13、SEQ ID NO:15、SEQ IDNO:17、SEQ ID NO:19、SEQ ID NO:21和/或SEQ ID NO:23；和/或，

(2)测定所述植物是否表达本发明所提供的蛋白质SEQ ID NO:2、SEQ ID NO:4、SEQ ID NO:6、SEQ ID NO:8、SEQ ID NO:10、SEQ ID NO:12、SEQ ID NO:14、SEQ ID NO:16、SEQ ID NO:18、SEQ ID NO:20、SEQ ID NO:22和/或SEQ ID NO:24。

测定的步骤可以通过常规的核酸检测和/或蛋白质检测方法来进行，只需要能够检测出蛋白质或其编码核酸带有相应核酸突变的方法都可以，示例性的方法包括蛋白质测序、核酸测序、聚合酶链式反应(PCR)检测、探针杂交检测等。

本发明取得的有益效果在于获得可替代现有技术的使植物具有除草剂抗性的蛋白质、核酸、表达盒、载体、细胞、植物、获得具有除草剂抗性的植物的应用和方法、以及鉴定本发明的植物的方法，并且可以通过转基因或非转基因方法获得具有除草剂抗性的植物品种，尤其是能继承宁春4号优良特性的小麦品种。

本发明引用了公开文献，这些文献是为了更清楚地描述本发明，它们的全文内容均纳入本文进行参考，就好像它们的全文已经在本文中重复叙述过一样。为了便于理解，以下将通过具体的附图和实施例对本发明进行详细地描述。需要特别指出的是，这些描述仅仅是示例性的描述，并不构成对本发明范围的限制。依据本说明书的论述，本发明的许多变化、改变对所属领域技术人员来说都是显而易见的了。

附图说明

图1显示了抗除草剂小麦阳性植株的筛选,其中红色箭头指示筛选到的阳性植株，黑色箭头指示非抗性植株。

图2显示了分子鉴定获得的小麦中3个IMI蛋白中突变的氨基酸位点。

图3显示了三个抗除草剂小麦株系(抗性位点分别是IMI1-S627N、IMI2-S629N、IMI3-S630N)后代M5代在三叶期喷施500毫克/升的咪草烟后一个月的生长状况。

图4显示了9个抗性株系(IMI1-A96T、IMI2-A98T、IMI1-A179V、IMI2-A181、IMI1-P171L、IMI2-P173L、IMI3-A99T、IMI3-P174L和IMI3-A182V)M4代植株在三叶期喷施500毫克/升的咪草烟后一个月的生长状况，其中，1.抗性植株IMI1-A96T；2.抗性植株IMI2-A98T；3.野生型对照；4.野生型对照；5.抗性植株IMI1-A179V；6.抗性植株IMI2-A181V；7.抗性植株IMI1-P171L；8.抗性植株IMI2-P173L；9.抗性植株IMI3-A99T；10.抗性植株IMI3-P174L；11.抗性植株IMI3-A182V。

具体实施方式

下述实施例中所用方法如无特别说明，均为常用分子生物学、组织培养技术和农学手册所记载的方法。例如，具体步骤可参见：《Molecular Cloning:A Laboratory Manual(3rd edition)》(Sambrook，J.，Russell,David W.，2001，Cold Spring Harbor)，《PlantPropagation by Tissue Culture》(Edwin F.George,Michael A.Hall,Geert-Jan DeKlerk,2008,Springer)。

实施例1.小麦突变体库的构建及抗除草剂突变体的筛选

取市售的小麦品种——春小麦宁春四号种子400kg、冬小麦矮抗58和济麦22种子各20kg，用水浸没种子4小时，将种子捞出，控干水分。然后，用0.5％(w/w)甲磺酸乙酯(EMS)浸泡种子12小时，保证液面浸没种子。12小时后弃去EMS溶液，换自来水浸泡种子，弃自来水，浸泡共重复5次，每次浸泡5分钟，然后用自来水冲洗种子2小时，冲洗过程中翻动种子，从而将EMS洗净，稍微风干后播种，待植株成熟后收割，其中春小麦宁春四号收获M2代(突变体库2代)种子3500公斤，矮抗58和济麦22两个品种各收获120公斤。

将M2代种子用浓度为500毫克/升的咪唑啉酮类除草剂咪草烟((RS)5-乙基-2-(4-异丙基-4-甲基-5-氧代-3-咪唑啉-2-基)烟酸，CAS号81335-77-5)在23度环境下浸泡18小时，然后将除草剂倒掉，将种子均匀地铺撒在沙床上萌发。3周后，发现除草剂浸泡处理的种子中有绿色秧苗持续生长(图1)，其中矮抗58品种获得119株疑似阳性植株，济麦22获得117株疑似阳性植株。

实施例2.抗除草剂阳性植株的分子水平鉴定

为了证明本发明获得的除草剂抗性突变体与植株体内的乙酰羟基合成酶突变的关系，本实施例对获得的抗除草剂阳性植株进行了分子水平的鉴定。

小麦基因组中的三个乙酰羟基合成酶基因IMI1、IMI2和IMI3分别位于D基因组、B基因组和A基因组中。取筛选获得的疑似抗除草剂阳性植株叶片，连同野生型植株叶片，分别单株提取基因组DNA，并以基因组DNA为模板将小麦基因组中编码的3个乙酰羟基合成酶IMI1、IMI2和IMI3通过聚合酶链式反应扩增其全长基因组DNA序列，扩增引物分别是：

TaImi1-F:TAAAGAAGCTAGCCCAATCTGAAC(SEQ ID NO:25)和TaImi1-R:AGAACATAGACTACAAAACTACAGG(SEQ ID NO:26)用于扩增IMI1基因；

TaImi2-F:CACAACCGCGAGTGGAAATACC(SEQ ID NO:27)和TaImi2-R:GAGAACAGAGACTACAAAACTACACG(SEQ ID NO:28)用于扩增IMI2基因；

TaImi3-F:GTTTGAGCATCAGCCTTTCTGCAC(SEQ ID NO:29)和

TaImi3-R:TCCATCGGTCATTATGTCATAACCG(SEQ ID NO:30)用于扩增IMI3基因。

PCR结束后，通过电泳胶回收，将获得的回收液委托北京三博远志生物技术有限公司进行测序，并对测序结果进行分析。分析结果显示，小麦由于是异源六倍体(AABBDD)，其三个拷贝的乙酰羟基合成酶基因的序列同源性非常高，其中IMI1与IMI2之间的核苷酸和蛋白质一致性分别为97％和99％，IMI1与IMI3之间的核苷酸和蛋白质一致性分别为98％和98％，IMI2与IMI3之间的核苷酸和蛋白质一致性分别为97％和98％。

分析结果发现，在鉴定的236株阳性植株中，有229株发现了3个乙酰羟基合成酶中某一位点发生了突变，在3个基因上各有4个突变位点(见图2)，分别是IMI1第286位由G到A的突变，导致相应编码氨基酸由第96位丙氨酸变为苏氨酸，该突变植株的ALS基因的核苷酸序列如SEQ ID NO：1所示，其编码的ALS蛋白的氨基酸序列如SEQ ID NO：2所示；IMI1第512位由C到T的突变，导致相应编码氨基酸由第171位脯氨酸变为亮氨酸，该突变植株的ALS基因的核苷酸序列如SEQ ID NO：3所示，其编码的ALS蛋白的氨基酸序列如SEQ ID NO：4所示；IMI1第536位由C到T的突变，导致相应编码氨基酸由第179位丙氨酸变为缬氨酸，该突变植株的ALS基因的核苷酸序列如SEQ ID NO：5所示，其编码的ALS蛋白的氨基酸序列如SEQ IDNO：6所示；IMI1第1880位由G到A的突变，导致相应编码氨基酸由第627位丝氨酸变为天冬酰胺，该突变植株的ALS基因的核苷酸序列如SEQ ID NO：7所示，其编码的ALS蛋白的氨基酸序列如SEQ ID NO：8所示。IMI2第292位由G到A的突变，导致相应编码氨基酸由第98位丙氨酸变为苏氨酸，该突变植株的ALS基因的核苷酸序列如SEQ ID NO：9所示，其编码的ALS蛋白的氨基酸序列如SEQ ID NO：10所示；IMI2第518位由C到T的突变，导致相应编码氨基酸由第173位脯氨酸变为亮氨酸，该突变植株的ALS基因的核苷酸序列如SEQ ID NO：11所示，其编码的ALS蛋白的氨基酸序列如SEQ ID NO：12所示；IMI2第542位由C到T的突变，导致相应编码氨基酸由第181位丙氨酸变为缬氨酸，该突变植株的ALS基因的核苷酸序列如SEQ ID NO：13所示，其编码的ALS蛋白的氨基酸序列如SEQ ID NO：14所示；IMI2第1886位由G到A的突变，导致相应编码氨基酸由第629位丝氨酸变为天冬酰胺，该突变植株的ALS基因的核苷酸序列如SEQ ID NO：15所示，其编码的ALS蛋白的氨基酸序列如SEQ ID NO：16所示。IMI3第295位由G到A的突变，导致相应编码氨基酸由第99位丙氨酸变为苏氨酸，该突变植株的ALS基因的核苷酸序列如SEQ ID NO：17所示，其编码的ALS蛋白的氨基酸序列如SEQ ID NO：18所示；IMI3第521位由C到T的突变，导致相应编码氨基酸由第174位脯氨酸变为亮氨酸，该突变植株的ALS基因的核苷酸序列如SEQ ID NO：19所示，其编码的ALS蛋白的氨基酸序列如SEQ ID NO：20所示；IMI3第545位由C到T的突变，导致相应编码氨基酸由第182位丙氨酸变为缬氨酸，该突变植株的ALS基因的核苷酸序列如SEQ ID NO：21所示，其编码的ALS蛋白的氨基酸序列如SEQ ID NO：22所示；IMI3第1889位由G到A的突变，导致相应编码氨基酸由第630位丝氨酸变为天冬酰胺，该突变植株的ALS基因的核苷酸序列如SEQ ID NO：23所示，其编码的ALS蛋白的氨基酸序列如SEQ ID NO：24所示。

实施例3.抗除草剂阳性植株后代田间咪唑啉酮类除草剂的耐受表型

将鉴定为纯合突变体的株系后代单株收种，在田间种约一平方米的小区，同时旁边种植野生型作为对照。待小麦长至三叶期时，喷施以浓度为500毫克/升的咪草烟，浓度是商用推荐剂量的2倍。喷施一个月后调查秧苗枯死情况，我们发现野生型对照在喷施咪草烟之后几乎停止生长，新长出来的叶片完全枯黄，进而逐渐枯死，而我们筛选到的不同位点的抗除草剂阳性苗则全部呈现对咪草烟完全耐受的表型，其正常的生长发育几乎没有受到影响(见图3和图4)，最终正常拔节抽穗。这表明我们筛选到的抗除草剂突变体确实能够稳定遗传地获得对咪唑啉酮类除草剂的耐受。

实施例4.抗性材料的两两杂交

为了检测是否具有双重或双重以上突变位点的抗性植株会比具有单位点抗性植株的除草剂耐受能力更强，我们将实施例1和2中获得的具有不同突变位点的抗性植株进行了两两杂交。

选择IMI1-G1880A突变位点分子鉴定为纯合的株系G38(序列对应SEQ ID NO：7)、IMI2-G1886A突变位点分子鉴定为纯合的株系E88(序列对应SEQ ID NO：15)以及IMI3-G1889A突变位点分子鉴定为纯合的株系E14(序列对应SEQ ID NO：23)，将这三个株系在温室种植，开花期将G38与E88、G38与E14、E88与E14进行杂交。具体方法是：先对母本穗提前进行人工去雄并套袋，父本植株开花的时候，将父本颖壳剪掉，光下放置约10分钟，待花药从颖壳伸长出来后，将父本花粉抖到母本穗上，重复两次，待种子成熟后收种，即为F1代杂交种。收获的F1代杂交种繁殖一代，收获F2代群体，在G38与E88杂交F2代群体中可以鉴定到同时带有纯合的IMI1-G1880A和IMI2-G1886A突变的植株，在G38与E14杂交F2代群体中可以鉴定到同时带有纯合的IMI1-G1880A和IMI3-G1889A突变的植株，在E888与E14杂交F2代群体中可以鉴定到同时带有纯合的IMI2-G1886A和IMI3-G1889A突变的植株。用除草剂对这些带有双位点突变的纯合植株进行喷施实验，结果表明，同时突变两个位点的植株相对于只有一个突变位点的植株，对除草剂的耐受性得到了显著的提高。

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acctccggcc cgggggccac caacctcgtc tccgcgctcg ccgacgccct cctcgactcc 480

atccccatgg tcgccatcac gggccaggtc ctccgccgca tgatcggcac ggacgcgttc 540

caggagacgc ccatagtgga ggtcacgcgc tccatcacca agcacaacta cctggtcctt 600

gacgtggagg atatcccccg cgtcatccag gaagccttct tccttgcatc ctctggccgc 660

ccggggccgg tgctagttga tatccccaag gacatccagc agcagatggc tgtgcccgtc 720

tgggacactc caatgagttt gccagggtac atcgcccgcc tgcccaagcc accatctact 780

gaatcgcttg agcaggtcct gcgtctggtt ggcgagtcac ggcgcccaat tctgtatgtt 840

ggtggtggct gcgctgcgtc tggcgaggag ttgcgccgct ttgttgagct tactgggatt 900

ccagttacaa ctactctgat gggccttggc aacttcccca gcgacgaccc actgtctctg 960

cgcatgcttg ggatgcatgg cactgtgtat gcaaattatg cagtagataa ggctgacctg 1020

ttgctcgcat ttggtgtgcg gtttgatgat cgtgtgactg ggaaaatcga ggcttttgca 1080

agcaggtcca agattgtgca cattgacatt gacccagctg agattggcaa gaacaagcag 1140

ccacatgtct ccatttgtgc agatgttaag cttgctttac aggggttgaa tgatctatta 1200

aatgggagca aagcacaaca gggtctggat tttggtccat ggcacaagga gttggatcag 1260

cagaagaggg agtttcctct aggattcaag acttttggcg aggccatccc gccgcaatat 1320

gctatccagg tactggatga gctgacaaaa ggggaggcga tcattgccac tggtgttggg 1380

cagcaccaga tgtgggcggc tcagtattac acttacaagc ggccacggca gtggctgtct 1440

tcgtctggtt tgggggcaat gggatttggg ttaccagctg cagctggcgc tgctgtggcc 1500

aacccaggtg ttacagttgt tgacattgat ggtgatggta gtttcctcat gaacattcag 1560

gagttggcgt tgatccgcat tgagaacctc ccagtgaagg tgatgatatt gaacaaccag 1620

catctgggaa tggtggtgca gtgggaggat aggttttaca aggccaatcg ggcgcacaca 1680

taccttggca acccagaaaa tgagagtgag atatatccag attttgtgac gattgctaaa 1740

ggattcaacg ttccagcagt tcgagtgacg aagaagagcg aagtcactgc agcaatcaag 1800

aagatgcttg agaccccagg gccatacttg ttggatatca tagtcccgca tcaggagcac 1860

gtgctgccta tgatcccaag cggtggtgct ttcaaggaca tgatcatgga gggtgatggc 1920

aggacctcgt actga 1935

<210> 4

<211> 644

<212> PRT

<213> 小麦(Triticum aestivum)

<400> 4

Met Ala Ala Ala Thr Ser Pro Ala Val Ala Phe Ser Gly Ala Thr Ala

1 5 10 15

Ala Ala Met Pro Lys Pro Ala Arg His Pro Leu Pro Arg His Gln Pro

20 25 30

Val Ser Arg Arg Ala Leu Pro Ala Arg Val Val Arg Cys Cys Ala Ala

35 40 45

Ser Pro Ala Ala Thr Ser Ala Ala Pro Pro Ala Thr Ala Leu Arg Pro

50 55 60

Trp Gly Pro Ser Glu Pro Arg Lys Gly Ala Asp Ile Leu Val Glu Ala

65 70 75 80

Leu Glu Arg Cys Gly Ile Val Asp Val Phe Ala Tyr Pro Gly Gly Ala

85 90 95

Ser Met Glu Ile His Gln Ala Leu Thr Arg Ser Pro Val Ile Thr Asn

100 105 110

His Leu Phe Arg His Glu Gln Gly Glu Ala Phe Ala Ala Ser Gly Tyr

115 120 125

Ala Arg Ala Ser Gly Arg Val Gly Val Cys Val Ala Thr Ser Gly Pro

130 135 140

Gly Ala Thr Asn Leu Val Ser Ala Leu Ala Asp Ala Leu Leu Asp Ser

145 150 155 160

Ile Pro Met Val Ala Ile Thr Gly Gln Val Leu Arg Arg Met Ile Gly

165 170 175

Thr Asp Ala Phe Gln Glu Thr Pro Ile Val Glu Val Thr Arg Ser Ile

180 185 190

Thr Lys His Asn Tyr Leu Val Leu Asp Val Glu Asp Ile Pro Arg Val

195 200 205

Ile Gln Glu Ala Phe Phe Leu Ala Ser Ser Gly Arg Pro Gly Pro Val

210 215 220

Leu Val Asp Ile Pro Lys Asp Ile Gln Gln Gln Met Ala Val Pro Val

225 230 235 240

Trp Asp Thr Pro Met Ser Leu Pro Gly Tyr Ile Ala Arg Leu Pro Lys

245 250 255

Pro Pro Ser Thr Glu Ser Leu Glu Gln Val Leu Arg Leu Val Gly Glu

260 265 270

Ser Arg Arg Pro Ile Leu Tyr Val Gly Gly Gly Cys Ala Ala Ser Gly

275 280 285

Glu Glu Leu Arg Arg Phe Val Glu Leu Thr Gly Ile Pro Val Thr Thr

290 295 300

Thr Leu Met Gly Leu Gly Asn Phe Pro Ser Asp Asp Pro Leu Ser Leu

305 310 315 320

Arg Met Leu Gly Met His Gly Thr Val Tyr Ala Asn Tyr Ala Val Asp

325 330 335

Lys Ala Asp Leu Leu Leu Ala Phe Gly Val Arg Phe Asp Asp Arg Val

340 345 350

Thr Gly Lys Ile Glu Ala Phe Ala Ser Arg Ser Lys Ile Val His Ile

355 360 365

Asp Ile Asp Pro Ala Glu Ile Gly Lys Asn Lys Gln Pro His Val Ser

370 375 380

Ile Cys Ala Asp Val Lys Leu Ala Leu Gln Gly Leu Asn Asp Leu Leu

385 390 395 400

Asn Gly Ser Lys Ala Gln Gln Gly Leu Asp Phe Gly Pro Trp His Lys

405 410 415

Glu Leu Asp Gln Gln Lys Arg Glu Phe Pro Leu Gly Phe Lys Thr Phe

420 425 430

Gly Glu Ala Ile Pro Pro Gln Tyr Ala Ile Gln Val Leu Asp Glu Leu

435 440 445

Thr Lys Gly Glu Ala Ile Ile Ala Thr Gly Val Gly Gln His Gln Met

450 455 460

Trp Ala Ala Gln Tyr Tyr Thr Tyr Lys Arg Pro Arg Gln Trp Leu Ser

465 470 475 480

Ser Ser Gly Leu Gly Ala Met Gly Phe Gly Leu Pro Ala Ala Ala Gly

485 490 495

Ala Ala Val Ala Asn Pro Gly Val Thr Val Val Asp Ile Asp Gly Asp

500 505 510

Gly Ser Phe Leu Met Asn Ile Gln Glu Leu Ala Leu Ile Arg Ile Glu

515 520 525

Asn Leu Pro Val Lys Val Met Ile Leu Asn Asn Gln His Leu Gly Met

530 535 540

Val Val Gln Trp Glu Asp Arg Phe Tyr Lys Ala Asn Arg Ala His Thr

545 550 555 560

Tyr Leu Gly Asn Pro Glu Asn Glu Ser Glu Ile Tyr Pro Asp Phe Val

565 570 575

Thr Ile Ala Lys Gly Phe Asn Val Pro Ala Val Arg Val Thr Lys Lys

580 585 590

Ser Glu Val Thr Ala Ala Ile Lys Lys Met Leu Glu Thr Pro Gly Pro

595 600 605

Tyr Leu Leu Asp Ile Ile Val Pro His Gln Glu His Val Leu Pro Met

610 615 620

Ile Pro Ser Gly Gly Ala Phe Lys Asp Met Ile Met Glu Gly Asp Gly

625 630 635 640

Arg Thr Ser Tyr

<210> 5

<211> 1935

<212> DNA

<213> 小麦(Triticum aestivum)

<400> 5

atggccgccg ccacctcccc cgccgtcgca ttctcgggcg ccaccgccgc cgccatgccc 60

aaacccgccc gccatcctct cccgcgccac cagcccgtct cgcgccgcgc gctccccgcc 120

cgcgtcgtca ggtgttgcgc cgcgtccccc gccgccacct ccgccgcgcc tcccgcaacc 180

gcgctccggc cctggggccc gtccgagccc cgcaagggcg ccgacatcct cgtcgaggcg 240

ctcgagcgct gcggcatcgt cgacgtcttc gcctaccccg gcggcgcctc catggagatc 300

caccaggcgc tgacgcgctc gcccgtcatc accaaccacc tcttccgcca cgagcagggg 360

gaggcgttcg cggcgtccgg ctacgcccgc gcgtccggcc gcgtcggcgt ctgcgtcgcc 420

acctccggcc cgggggccac caacctcgtc tccgcgctcg ccgacgccct cctcgactcc 480

atccccatgg tcgccatcac gggccaggtc ccccgccgca tgatcggcac ggacgtgttc 540

caggagacgc ccatagtgga ggtcacgcgc tccatcacca agcacaacta cctggtcctt 600

gacgtggagg atatcccccg cgtcatccag gaagccttct tccttgcatc ctctggccgc 660

ccggggccgg tgctagttga tatccccaag gacatccagc agcagatggc tgtgcccgtc 720

tgggacactc caatgagttt gccagggtac atcgcccgcc tgcccaagcc accatctact 780

gaatcgcttg agcaggtcct gcgtctggtt ggcgagtcac ggcgcccaat tctgtatgtt 840

ggtggtggct gcgctgcgtc tggcgaggag ttgcgccgct ttgttgagct tactgggatt 900

ccagttacaa ctactctgat gggccttggc aacttcccca gcgacgaccc actgtctctg 960

cgcatgcttg ggatgcatgg cactgtgtat gcaaattatg cagtagataa ggctgacctg 1020

ttgctcgcat ttggtgtgcg gtttgatgat cgtgtgactg ggaaaatcga ggcttttgca 1080

agcaggtcca agattgtgca cattgacatt gacccagctg agattggcaa gaacaagcag 1140

ccacatgtct ccatttgtgc agatgttaag cttgctttac aggggttgaa tgatctatta 1200

aatgggagca aagcacaaca gggtctggat tttggtccat ggcacaagga gttggatcag 1260

cagaagaggg agtttcctct aggattcaag acttttggcg aggccatccc gccgcaatat 1320

gctatccagg tactggatga gctgacaaaa ggggaggcga tcattgccac tggtgttggg 1380

cagcaccaga tgtgggcggc tcagtattac acttacaagc ggccacggca gtggctgtct 1440

tcgtctggtt tgggggcaat gggatttggg ttaccagctg cagctggcgc tgctgtggcc 1500

aacccaggtg ttacagttgt tgacattgat ggtgatggta gtttcctcat gaacattcag 1560

gagttggcgt tgatccgcat tgagaacctc ccagtgaagg tgatgatatt gaacaaccag 1620

catctgggaa tggtggtgca gtgggaggat aggttttaca aggccaatcg ggcgcacaca 1680

taccttggca acccagaaaa tgagagtgag atatatccag attttgtgac gattgctaaa 1740

ggattcaacg ttccagcagt tcgagtgacg aagaagagcg aagtcactgc agcaatcaag 1800

aagatgcttg agaccccagg gccatacttg ttggatatca tagtcccgca tcaggagcac 1860

gtgctgccta tgatcccaag cggtggtgct ttcaaggaca tgatcatgga gggtgatggc 1920

aggacctcgt actga 1935

<210> 6

<211> 644

<212> PRT

<213> 小麦(Triticum aestivum)

<400> 6

Met Ala Ala Ala Thr Ser Pro Ala Val Ala Phe Ser Gly Ala Thr Ala

1 5 10 15

Ala Ala Met Pro Lys Pro Ala Arg His Pro Leu Pro Arg His Gln Pro

20 25 30

Val Ser Arg Arg Ala Leu Pro Ala Arg Val Val Arg Cys Cys Ala Ala

35 40 45

Ser Pro Ala Ala Thr Ser Ala Ala Pro Pro Ala Thr Ala Leu Arg Pro

50 55 60

Trp Gly Pro Ser Glu Pro Arg Lys Gly Ala Asp Ile Leu Val Glu Ala

65 70 75 80

Leu Glu Arg Cys Gly Ile Val Asp Val Phe Ala Tyr Pro Gly Gly Ala

85 90 95

Ser Met Glu Ile His Gln Ala Leu Thr Arg Ser Pro Val Ile Thr Asn

100 105 110

His Leu Phe Arg His Glu Gln Gly Glu Ala Phe Ala Ala Ser Gly Tyr

115 120 125

Ala Arg Ala Ser Gly Arg Val Gly Val Cys Val Ala Thr Ser Gly Pro

130 135 140

Gly Ala Thr Asn Leu Val Ser Ala Leu Ala Asp Ala Leu Leu Asp Ser

145 150 155 160

Ile Pro Met Val Ala Ile Thr Gly Gln Val Pro Arg Arg Met Ile Gly

165 170 175

Thr Asp Val Phe Gln Glu Thr Pro Ile Val Glu Val Thr Arg Ser Ile

180 185 190

Thr Lys His Asn Tyr Leu Val Leu Asp Val Glu Asp Ile Pro Arg Val

195 200 205

Ile Gln Glu Ala Phe Phe Leu Ala Ser Ser Gly Arg Pro Gly Pro Val

210 215 220

Leu Val Asp Ile Pro Lys Asp Ile Gln Gln Gln Met Ala Val Pro Val

225 230 235 240

Trp Asp Thr Pro Met Ser Leu Pro Gly Tyr Ile Ala Arg Leu Pro Lys

245 250 255

Pro Pro Ser Thr Glu Ser Leu Glu Gln Val Leu Arg Leu Val Gly Glu

260 265 270

Ser Arg Arg Pro Ile Leu Tyr Val Gly Gly Gly Cys Ala Ala Ser Gly

275 280 285

Glu Glu Leu Arg Arg Phe Val Glu Leu Thr Gly Ile Pro Val Thr Thr

290 295 300

Thr Leu Met Gly Leu Gly Asn Phe Pro Ser Asp Asp Pro Leu Ser Leu

305 310 315 320

Arg Met Leu Gly Met His Gly Thr Val Tyr Ala Asn Tyr Ala Val Asp

325 330 335

Lys Ala Asp Leu Leu Leu Ala Phe Gly Val Arg Phe Asp Asp Arg Val

340 345 350

Thr Gly Lys Ile Glu Ala Phe Ala Ser Arg Ser Lys Ile Val His Ile

355 360 365

Asp Ile Asp Pro Ala Glu Ile Gly Lys Asn Lys Gln Pro His Val Ser

370 375 380

Ile Cys Ala Asp Val Lys Leu Ala Leu Gln Gly Leu Asn Asp Leu Leu

385 390 395 400

Asn Gly Ser Lys Ala Gln Gln Gly Leu Asp Phe Gly Pro Trp His Lys

405 410 415

Glu Leu Asp Gln Gln Lys Arg Glu Phe Pro Leu Gly Phe Lys Thr Phe

420 425 430

Gly Glu Ala Ile Pro Pro Gln Tyr Ala Ile Gln Val Leu Asp Glu Leu

435 440 445

Thr Lys Gly Glu Ala Ile Ile Ala Thr Gly Val Gly Gln His Gln Met

450 455 460

Trp Ala Ala Gln Tyr Tyr Thr Tyr Lys Arg Pro Arg Gln Trp Leu Ser

465 470 475 480

Ser Ser Gly Leu Gly Ala Met Gly Phe Gly Leu Pro Ala Ala Ala Gly

485 490 495

Ala Ala Val Ala Asn Pro Gly Val Thr Val Val Asp Ile Asp Gly Asp

500 505 510

Gly Ser Phe Leu Met Asn Ile Gln Glu Leu Ala Leu Ile Arg Ile Glu

515 520 525

Asn Leu Pro Val Lys Val Met Ile Leu Asn Asn Gln His Leu Gly Met

530 535 540

Val Val Gln Trp Glu Asp Arg Phe Tyr Lys Ala Asn Arg Ala His Thr

545 550 555 560

Tyr Leu Gly Asn Pro Glu Asn Glu Ser Glu Ile Tyr Pro Asp Phe Val

565 570 575

Thr Ile Ala Lys Gly Phe Asn Val Pro Ala Val Arg Val Thr Lys Lys

580 585 590

Ser Glu Val Thr Ala Ala Ile Lys Lys Met Leu Glu Thr Pro Gly Pro

595 600 605

Tyr Leu Leu Asp Ile Ile Val Pro His Gln Glu His Val Leu Pro Met

610 615 620

Ile Pro Ser Gly Gly Ala Phe Lys Asp Met Ile Met Glu Gly Asp Gly

625 630 635 640

Arg Thr Ser Tyr

<210> 7

<211> 1935

<212> DNA

<213> 小麦(Triticum aestivum)

<400> 7

atggccgccg ccacctcccc cgccgtcgca ttctcgggcg ccaccgccgc cgccatgccc 60

aaacccgccc gccatcctct cccgcgccac cagcccgtct cgcgccgcgc gctccccgcc 120

cgcgtcgtca ggtgttgcgc cgcgtccccc gccgccacct ccgccgcgcc tcccgcaacc 180

gcgctccggc cctggggccc gtccgagccc cgcaagggcg ccgacatcct cgtcgaggcg 240

ctcgagcgct gcggcatcgt cgacgtcttc gcctaccccg gcggcgcctc catggagatc 300

caccaggcgc tgacgcgctc gcccgtcatc accaaccacc tcttccgcca cgagcagggg 360

gaggcgttcg cggcgtccgg ctacgcccgc gcgtccggcc gcgtcggcgt ctgcgtcgcc 420

acctccggcc cgggggccac caacctcgtc tccgcgctcg ccgacgccct cctcgactcc 480

atccccatgg tcgccatcac gggccaggtc ccccgccgca tgatcggcac ggacgcgttc 540

caggagacgc ccatagtgga ggtcacgcgc tccatcacca agcacaacta cctggtcctt 600

gacgtggagg atatcccccg cgtcatccag gaagccttct tccttgcatc ctctggccgc 660

ccggggccgg tgctagttga tatccccaag gacatccagc agcagatggc tgtgcccgtc 720

tgggacactc caatgagttt gccagggtac atcgcccgcc tgcccaagcc accatctact 780

gaatcgcttg agcaggtcct gcgtctggtt ggcgagtcac ggcgcccaat tctgtatgtt 840

ggtggtggct gcgctgcgtc tggcgaggag ttgcgccgct ttgttgagct tactgggatt 900

ccagttacaa ctactctgat gggccttggc aacttcccca gcgacgaccc actgtctctg 960

cgcatgcttg ggatgcatgg cactgtgtat gcaaattatg cagtagataa ggctgacctg 1020

ttgctcgcat ttggtgtgcg gtttgatgat cgtgtgactg ggaaaatcga ggcttttgca 1080

agcaggtcca agattgtgca cattgacatt gacccagctg agattggcaa gaacaagcag 1140

ccacatgtct ccatttgtgc agatgttaag cttgctttac aggggttgaa tgatctatta 1200

aatgggagca aagcacaaca gggtctggat tttggtccat ggcacaagga gttggatcag 1260

cagaagaggg agtttcctct aggattcaag acttttggcg aggccatccc gccgcaatat 1320

gctatccagg tactggatga gctgacaaaa ggggaggcga tcattgccac tggtgttggg 1380

cagcaccaga tgtgggcggc tcagtattac acttacaagc ggccacggca gtggctgtct 1440

tcgtctggtt tgggggcaat gggatttggg ttaccagctg cagctggcgc tgctgtggcc 1500

aacccaggtg ttacagttgt tgacattgat ggtgatggta gtttcctcat gaacattcag 1560

gagttggcgt tgatccgcat tgagaacctc ccagtgaagg tgatgatatt gaacaaccag 1620

catctgggaa tggtggtgca gtgggaggat aggttttaca aggccaatcg ggcgcacaca 1680

taccttggca acccagaaaa tgagagtgag atatatccag attttgtgac gattgctaaa 1740

ggattcaacg ttccagcagt tcgagtgacg aagaagagcg aagtcactgc agcaatcaag 1800

aagatgcttg agaccccagg gccatacttg ttggatatca tagtcccgca tcaggagcac 1860

gtgctgccta tgatcccaaa cggtggtgct ttcaaggaca tgatcatgga gggtgatggc 1920

aggacctcgt actga 1935

<210> 8

<211> 644

<212> PRT

<213> 小麦(Triticum aestivum)

<400> 8

Met Ala Ala Ala Thr Ser Pro Ala Val Ala Phe Ser Gly Ala Thr Ala

1 5 10 15

Ala Ala Met Pro Lys Pro Ala Arg His Pro Leu Pro Arg His Gln Pro

20 25 30

Val Ser Arg Arg Ala Leu Pro Ala Arg Val Val Arg Cys Cys Ala Ala

35 40 45

Ser Pro Ala Ala Thr Ser Ala Ala Pro Pro Ala Thr Ala Leu Arg Pro

50 55 60

Trp Gly Pro Ser Glu Pro Arg Lys Gly Ala Asp Ile Leu Val Glu Ala

65 70 75 80

Leu Glu Arg Cys Gly Ile Val Asp Val Phe Ala Tyr Pro Gly Gly Ala

85 90 95

Ser Met Glu Ile His Gln Ala Leu Thr Arg Ser Pro Val Ile Thr Asn

100 105 110

His Leu Phe Arg His Glu Gln Gly Glu Ala Phe Ala Ala Ser Gly Tyr

115 120 125

Ala Arg Ala Ser Gly Arg Val Gly Val Cys Val Ala Thr Ser Gly Pro

130 135 140

Gly Ala Thr Asn Leu Val Ser Ala Leu Ala Asp Ala Leu Leu Asp Ser

145 150 155 160

Ile Pro Met Val Ala Ile Thr Gly Gln Val Pro Arg Arg Met Ile Gly

165 170 175

Thr Asp Ala Phe Gln Glu Thr Pro Ile Val Glu Val Thr Arg Ser Ile

180 185 190

Thr Lys His Asn Tyr Leu Val Leu Asp Val Glu Asp Ile Pro Arg Val

195 200 205

Ile Gln Glu Ala Phe Phe Leu Ala Ser Ser Gly Arg Pro Gly Pro Val

210 215 220

Leu Val Asp Ile Pro Lys Asp Ile Gln Gln Gln Met Ala Val Pro Val

225 230 235 240

Trp Asp Thr Pro Met Ser Leu Pro Gly Tyr Ile Ala Arg Leu Pro Lys

245 250 255

Pro Pro Ser Thr Glu Ser Leu Glu Gln Val Leu Arg Leu Val Gly Glu

260 265 270

Ser Arg Arg Pro Ile Leu Tyr Val Gly Gly Gly Cys Ala Ala Ser Gly

275 280 285

Glu Glu Leu Arg Arg Phe Val Glu Leu Thr Gly Ile Pro Val Thr Thr

290 295 300

Thr Leu Met Gly Leu Gly Asn Phe Pro Ser Asp Asp Pro Leu Ser Leu

305 310 315 320

Arg Met Leu Gly Met His Gly Thr Val Tyr Ala Asn Tyr Ala Val Asp

325 330 335

Lys Ala Asp Leu Leu Leu Ala Phe Gly Val Arg Phe Asp Asp Arg Val

340 345 350

Thr Gly Lys Ile Glu Ala Phe Ala Ser Arg Ser Lys Ile Val His Ile

355 360 365

Asp Ile Asp Pro Ala Glu Ile Gly Lys Asn Lys Gln Pro His Val Ser

370 375 380

Ile Cys Ala Asp Val Lys Leu Ala Leu Gln Gly Leu Asn Asp Leu Leu

385 390 395 400

Asn Gly Ser Lys Ala Gln Gln Gly Leu Asp Phe Gly Pro Trp His Lys

405 410 415

Glu Leu Asp Gln Gln Lys Arg Glu Phe Pro Leu Gly Phe Lys Thr Phe

420 425 430

Gly Glu Ala Ile Pro Pro Gln Tyr Ala Ile Gln Val Leu Asp Glu Leu

435 440 445

Thr Lys Gly Glu Ala Ile Ile Ala Thr Gly Val Gly Gln His Gln Met

450 455 460

Trp Ala Ala Gln Tyr Tyr Thr Tyr Lys Arg Pro Arg Gln Trp Leu Ser

465 470 475 480

Ser Ser Gly Leu Gly Ala Met Gly Phe Gly Leu Pro Ala Ala Ala Gly

485 490 495

Ala Ala Val Ala Asn Pro Gly Val Thr Val Val Asp Ile Asp Gly Asp

500 505 510

Gly Ser Phe Leu Met Asn Ile Gln Glu Leu Ala Leu Ile Arg Ile Glu

515 520 525

Asn Leu Pro Val Lys Val Met Ile Leu Asn Asn Gln His Leu Gly Met

530 535 540

Val Val Gln Trp Glu Asp Arg Phe Tyr Lys Ala Asn Arg Ala His Thr

545 550 555 560

Tyr Leu Gly Asn Pro Glu Asn Glu Ser Glu Ile Tyr Pro Asp Phe Val

565 570 575

Thr Ile Ala Lys Gly Phe Asn Val Pro Ala Val Arg Val Thr Lys Lys

580 585 590

Ser Glu Val Thr Ala Ala Ile Lys Lys Met Leu Glu Thr Pro Gly Pro

595 600 605

Tyr Leu Leu Asp Ile Ile Val Pro His Gln Glu His Val Leu Pro Met

610 615 620

Ile Pro Asn Gly Gly Ala Phe Lys Asp Met Ile Met Glu Gly Asp Gly

625 630 635 640

Arg Thr Ser Tyr

<210> 9

<211> 1941

<212> DNA

<213> 小麦(Triticum aestivum)

<400> 9

atggccgcag ccacctcccc cgccgtcgca ttctcgggcg ccgccgccgc cgccgccgcc 60

atacccaaac ccgcccgcca gcctctcccg cgccaccagc ccgcctcgcg ccgcgcgctc 120

cccgcccgca tcgtcaggtg ctgcgccgcg tcccccgccg ccacctccgt cgcgcctccc 180

gccaccgcgc tccggccgtg gggcccctcc gagccccgca agggcgccga catcctcgtc 240

gaggcgctgg agcgctgcgg catcgtcgac gtcttcgcct accctggcgg cacgtccatg 300

gagatccacc aggcgctgac gcgctcgcca gtcatcacca accacctctt ccgccacgag 360

cagggggagg cgttcgcggc gtccgggtac gcccgcgcgt ccggccgcgt cggcgtctgc 420

gtcgccacct ccggcccggg ggccaccaac ctcgtctccg cgctcgccga cgctctcctc 480

gactccatcc ccatggtcgc catcacgggc caggtccccc gccgcatgat cggcacggat 540

gcgttccagg agacgcccat cgtggaggtc acgcgctcca tcaccaagca caactacctg 600

gtccttgacg tggaggatat cccccgcgtc atccaggaag ccttcttcct cgcatcctct 660

ggccgcccgg ggccggtgct ggttgatatc cccaaggaca tccagcagca gatggctgtg 720

cctgtctggg acacgccgat gagtttgcca gggtacatcg cccgcctgcc caagccacca 780

tctactgaat cgcttgagca ggtcctgcgt ctggttggcg agtcacggcg cccaattctg 840

tatgttggtg gtggctgcgc tgcatctggt gaggagttgc gccgctttgt tgagctcact 900

gggattccag ttacaactac tcttatgggc cttggcaact tccccagtga cgacccactg 960

tctctgcgca tgctggggat gcatggcact gtgtatgcaa attatgcagt agataaggct 1020

gacctgttgc ttgcatttgg tgtgcggttt gatgatcgtg tgaccgggaa aatcgaggct 1080

tttgcaagca ggtccaagat tgtgcacatt gacattgacc cagctgagat tggcaagaac 1140

aagcagccac atgtctccat ttgtgcagat gttaagcttg ctttacaggg gttgaatgct 1200

ctattaaatg ggagcaaagc acaacagggt ctggattttg gtccatggca caaggagttg 1260

gatcagcaga agagggagtt tcctctagga ttcaagactt ttggtgaggc catcccgccg 1320

caatatgcta tccaggtact ggatgagctg acaaaagggg aggcgatcat tgccaccggt 1380

gttgggcagc atcagatgtg ggcggctcag tattacactt acaagcggcc acggcagtgg 1440

ctgtcttcat ccggtttggg tgcaatggga tttgggttgc cagctgcagc tggcgctgct 1500

gtggccaacc caggtgttac agttgttgac attgatgggg atggtagttt cctcatgaac 1560

attcaggagt tggcgttgat ccgtattgag aacctcccag tgaaggtgat gatattgaac 1620

aaccagcatc tgggaatggt ggtgcagtgg gaggataggt tttacaaggc caaccgggcg 1680

cacacatacc ttggcaaccc agaaaatgag agtgagatat atccagattt tgtgacgatt 1740

gctaaaggat tcaacgttcc ggcagttcgt gtgacgaaga agagcgaagt cactgcagca 1800

atcaagaaga tgcttgagac cccagggcca tacttgttgg atatcattgt cccgcatcag 1860

gagcacgtgc tgcctatgat cccaagcggt ggtgctttta aggacatgat catggagggt 1920

gatggcagga cctcgtactg a 1941

<210> 10

<211> 646

<212> PRT

<213> 小麦(Triticum aestivum)

<400> 10

Met Ala Ala Ala Thr Ser Pro Ala Val Ala Phe Ser Gly Ala Ala Ala

1 5 10 15

Ala Ala Ala Ala Ile Pro Lys Pro Ala Arg Gln Pro Leu Pro Arg His

20 25 30

Gln Pro Ala Ser Arg Arg Ala Leu Pro Ala Arg Ile Val Arg Cys Cys

35 40 45

Ala Ala Ser Pro Ala Ala Thr Ser Val Ala Pro Pro Ala Thr Ala Leu

50 55 60

Arg Pro Trp Gly Pro Ser Glu Pro Arg Lys Gly Ala Asp Ile Leu Val

65 70 75 80

Glu Ala Leu Glu Arg Cys Gly Ile Val Asp Val Phe Ala Tyr Pro Gly

85 90 95

Gly Thr Ser Met Glu Ile His Gln Ala Leu Thr Arg Ser Pro Val Ile

100 105 110

Thr Asn His Leu Phe Arg His Glu Gln Gly Glu Ala Phe Ala Ala Ser

115 120 125

Gly Tyr Ala Arg Ala Ser Gly Arg Val Gly Val Cys Val Ala Thr Ser

130 135 140

Gly Pro Gly Ala Thr Asn Leu Val Ser Ala Leu Ala Asp Ala Leu Leu

145 150 155 160

Asp Ser Ile Pro Met Val Ala Ile Thr Gly Gln Val Pro Arg Arg Met

165 170 175

Ile Gly Thr Asp Ala Phe Gln Glu Thr Pro Ile Val Glu Val Thr Arg

180 185 190

Ser Ile Thr Lys His Asn Tyr Leu Val Leu Asp Val Glu Asp Ile Pro

195 200 205

Arg Val Ile Gln Glu Ala Phe Phe Leu Ala Ser Ser Gly Arg Pro Gly

210 215 220

Pro Val Leu Val Asp Ile Pro Lys Asp Ile Gln Gln Gln Met Ala Val

225 230 235 240

Pro Val Trp Asp Thr Pro Met Ser Leu Pro Gly Tyr Ile Ala Arg Leu

245 250 255

Pro Lys Pro Pro Ser Thr Glu Ser Leu Glu Gln Val Leu Arg Leu Val

260 265 270

Gly Glu Ser Arg Arg Pro Ile Leu Tyr Val Gly Gly Gly Cys Ala Ala

275 280 285

Ser Gly Glu Glu Leu Arg Arg Phe Val Glu Leu Thr Gly Ile Pro Val

290 295 300

Thr Thr Thr Leu Met Gly Leu Gly Asn Phe Pro Ser Asp Asp Pro Leu

305 310 315 320

Ser Leu Arg Met Leu Gly Met His Gly Thr Val Tyr Ala Asn Tyr Ala

325 330 335

Val Asp Lys Ala Asp Leu Leu Leu Ala Phe Gly Val Arg Phe Asp Asp

340 345 350

Arg Val Thr Gly Lys Ile Glu Ala Phe Ala Ser Arg Ser Lys Ile Val

355 360 365

His Ile Asp Ile Asp Pro Ala Glu Ile Gly Lys Asn Lys Gln Pro His

370 375 380

Val Ser Ile Cys Ala Asp Val Lys Leu Ala Leu Gln Gly Leu Asn Ala

385 390 395 400

Leu Leu Asn Gly Ser Lys Ala Gln Gln Gly Leu Asp Phe Gly Pro Trp

405 410 415

His Lys Glu Leu Asp Gln Gln Lys Arg Glu Phe Pro Leu Gly Phe Lys

420 425 430

Thr Phe Gly Glu Ala Ile Pro Pro Gln Tyr Ala Ile Gln Val Leu Asp

435 440 445

Glu Leu Thr Lys Gly Glu Ala Ile Ile Ala Thr Gly Val Gly Gln His

450 455 460

Gln Met Trp Ala Ala Gln Tyr Tyr Thr Tyr Lys Arg Pro Arg Gln Trp

465 470 475 480

Leu Ser Ser Ser Gly Leu Gly Ala Met Gly Phe Gly Leu Pro Ala Ala

485 490 495

Ala Gly Ala Ala Val Ala Asn Pro Gly Val Thr Val Val Asp Ile Asp

500 505 510

Gly Asp Gly Ser Phe Leu Met Asn Ile Gln Glu Leu Ala Leu Ile Arg

515 520 525

Ile Glu Asn Leu Pro Val Lys Val Met Ile Leu Asn Asn Gln His Leu

530 535 540

Gly Met Val Val Gln Trp Glu Asp Arg Phe Tyr Lys Ala Asn Arg Ala

545 550 555 560

His Thr Tyr Leu Gly Asn Pro Glu Asn Glu Ser Glu Ile Tyr Pro Asp

565 570 575

Phe Val Thr Ile Ala Lys Gly Phe Asn Val Pro Ala Val Arg Val Thr

580 585 590

Lys Lys Ser Glu Val Thr Ala Ala Ile Lys Lys Met Leu Glu Thr Pro

595 600 605

Gly Pro Tyr Leu Leu Asp Ile Ile Val Pro His Gln Glu His Val Leu

610 615 620

Pro Met Ile Pro Ser Gly Gly Ala Phe Lys Asp Met Ile Met Glu Gly

625 630 635 640

Asp Gly Arg Thr Ser Tyr

645

<210> 11

<211> 1941

<212> DNA

<213> 小麦(Triticum aestivum)

<400> 11

atggccgcag ccacctcccc cgccgtcgca ttctcgggcg ccgccgccgc cgccgccgcc 60

atacccaaac ccgcccgcca gcctctcccg cgccaccagc ccgcctcgcg ccgcgcgctc 120

cccgcccgca tcgtcaggtg ctgcgccgcg tcccccgccg ccacctccgt cgcgcctccc 180

gccaccgcgc tccggccgtg gggcccctcc gagccccgca agggcgccga catcctcgtc 240

gaggcgctgg agcgctgcgg catcgtcgac gtcttcgcct accctggcgg cgcgtccatg 300

gagatccacc aggcgctgac gcgctcgcca gtcatcacca accacctctt ccgccacgag 360

cagggggagg cgttcgcggc gtccgggtac gcccgcgcgt ccggccgcgt cggcgtctgc 420

gtcgccacct ccggcccggg ggccaccaac ctcgtctccg cgctcgccga cgctctcctc 480

gactccatcc ccatggtcgc catcacgggc caggtcctcc gccgcatgat cggcacggat 540

gcgttccagg agacgcccat cgtggaggtc acgcgctcca tcaccaagca caactacctg 600

gtccttgacg tggaggatat cccccgcgtc atccaggaag ccttcttcct cgcatcctct 660

ggccgcccgg ggccggtgct ggttgatatc cccaaggaca tccagcagca gatggctgtg 720

cctgtctggg acacgccgat gagtttgcca gggtacatcg cccgcctgcc caagccacca 780

tctactgaat cgcttgagca ggtcctgcgt ctggttggcg agtcacggcg cccaattctg 840

tatgttggtg gtggctgcgc tgcatctggt gaggagttgc gccgctttgt tgagctcact 900

gggattccag ttacaactac tcttatgggc cttggcaact tccccagtga cgacccactg 960

tctctgcgca tgctggggat gcatggcact gtgtatgcaa attatgcagt agataaggct 1020

gacctgttgc ttgcatttgg tgtgcggttt gatgatcgtg tgaccgggaa aatcgaggct 1080

tttgcaagca ggtccaagat tgtgcacatt gacattgacc cagctgagat tggcaagaac 1140

aagcagccac atgtctccat ttgtgcagat gttaagcttg ctttacaggg gttgaatgct 1200

ctattaaatg ggagcaaagc acaacagggt ctggattttg gtccatggca caaggagttg 1260

gatcagcaga agagggagtt tcctctagga ttcaagactt ttggtgaggc catcccgccg 1320

caatatgcta tccaggtact ggatgagctg acaaaagggg aggcgatcat tgccaccggt 1380

gttgggcagc atcagatgtg ggcggctcag tattacactt acaagcggcc acggcagtgg 1440

ctgtcttcat ccggtttggg tgcaatggga tttgggttgc cagctgcagc tggcgctgct 1500

gtggccaacc caggtgttac agttgttgac attgatgggg atggtagttt cctcatgaac 1560

attcaggagt tggcgttgat ccgtattgag aacctcccag tgaaggtgat gatattgaac 1620

aaccagcatc tgggaatggt ggtgcagtgg gaggataggt tttacaaggc caaccgggcg 1680

cacacatacc ttggcaaccc agaaaatgag agtgagatat atccagattt tgtgacgatt 1740

gctaaaggat tcaacgttcc ggcagttcgt gtgacgaaga agagcgaagt cactgcagca 1800

atcaagaaga tgcttgagac cccagggcca tacttgttgg atatcattgt cccgcatcag 1860

gagcacgtgc tgcctatgat cccaagcggt ggtgctttta aggacatgat catggagggt 1920

gatggcagga cctcgtactg a 1941

<210> 12

<211> 646

<212> PRT

<213> 小麦 (Triticum aestivum)

<400> 12

Met Ala Ala Ala Thr Ser Pro Ala Val Ala Phe Ser Gly Ala Ala Ala

1 5 10 15

Ala Ala Ala Ala Ile Pro Lys Pro Ala Arg Gln Pro Leu Pro Arg His

20 25 30

Gln Pro Ala Ser Arg Arg Ala Leu Pro Ala Arg Ile Val Arg Cys Cys

35 40 45

Ala Ala Ser Pro Ala Ala Thr Ser Val Ala Pro Pro Ala Thr Ala Leu

50 55 60

Arg Pro Trp Gly Pro Ser Glu Pro Arg Lys Gly Ala Asp Ile Leu Val

65 70 75 80

Glu Ala Leu Glu Arg Cys Gly Ile Val Asp Val Phe Ala Tyr Pro Gly

85 90 95

Gly Ala Ser Met Glu Ile His Gln Ala Leu Thr Arg Ser Pro Val Ile

100 105 110

Thr Asn His Leu Phe Arg His Glu Gln Gly Glu Ala Phe Ala Ala Ser

115 120 125

Gly Tyr Ala Arg Ala Ser Gly Arg Val Gly Val Cys Val Ala Thr Ser

130 135 140

Gly Pro Gly Ala Thr Asn Leu Val Ser Ala Leu Ala Asp Ala Leu Leu

145 150 155 160

Asp Ser Ile Pro Met Val Ala Ile Thr Gly Gln Val Leu Arg Arg Met

165 170 175

Ile Gly Thr Asp Ala Phe Gln Glu Thr Pro Ile Val Glu Val Thr Arg

180 185 190

Ser Ile Thr Lys His Asn Tyr Leu Val Leu Asp Val Glu Asp Ile Pro

195 200 205

Arg Val Ile Gln Glu Ala Phe Phe Leu Ala Ser Ser Gly Arg Pro Gly

210 215 220

Pro Val Leu Val Asp Ile Pro Lys Asp Ile Gln Gln Gln Met Ala Val

225 230 235 240

Pro Val Trp Asp Thr Pro Met Ser Leu Pro Gly Tyr Ile Ala Arg Leu

245 250 255

Pro Lys Pro Pro Ser Thr Glu Ser Leu Glu Gln Val Leu Arg Leu Val

260 265 270

Gly Glu Ser Arg Arg Pro Ile Leu Tyr Val Gly Gly Gly Cys Ala Ala

275 280 285

Ser Gly Glu Glu Leu Arg Arg Phe Val Glu Leu Thr Gly Ile Pro Val

290 295 300

Thr Thr Thr Leu Met Gly Leu Gly Asn Phe Pro Ser Asp Asp Pro Leu

305 310 315 320

Ser Leu Arg Met Leu Gly Met His Gly Thr Val Tyr Ala Asn Tyr Ala

325 330 335

Val Asp Lys Ala Asp Leu Leu Leu Ala Phe Gly Val Arg Phe Asp Asp

340 345 350

Arg Val Thr Gly Lys Ile Glu Ala Phe Ala Ser Arg Ser Lys Ile Val

355 360 365

His Ile Asp Ile Asp Pro Ala Glu Ile Gly Lys Asn Lys Gln Pro His

370 375 380

Val Ser Ile Cys Ala Asp Val Lys Leu Ala Leu Gln Gly Leu Asn Ala

385 390 395 400

Leu Leu Asn Gly Ser Lys Ala Gln Gln Gly Leu Asp Phe Gly Pro Trp

405 410 415

His Lys Glu Leu Asp Gln Gln Lys Arg Glu Phe Pro Leu Gly Phe Lys

420 425 430

Thr Phe Gly Glu Ala Ile Pro Pro Gln Tyr Ala Ile Gln Val Leu Asp

435 440 445

Glu Leu Thr Lys Gly Glu Ala Ile Ile Ala Thr Gly Val Gly Gln His

450 455 460

Gln Met Trp Ala Ala Gln Tyr Tyr Thr Tyr Lys Arg Pro Arg Gln Trp

465 470 475 480

Leu Ser Ser Ser Gly Leu Gly Ala Met Gly Phe Gly Leu Pro Ala Ala

485 490 495

Ala Gly Ala Ala Val Ala Asn Pro Gly Val Thr Val Val Asp Ile Asp

500 505 510

Gly Asp Gly Ser Phe Leu Met Asn Ile Gln Glu Leu Ala Leu Ile Arg

515 520 525

Ile Glu Asn Leu Pro Val Lys Val Met Ile Leu Asn Asn Gln His Leu

530 535 540

Gly Met Val Val Gln Trp Glu Asp Arg Phe Tyr Lys Ala Asn Arg Ala

545 550 555 560

His Thr Tyr Leu Gly Asn Pro Glu Asn Glu Ser Glu Ile Tyr Pro Asp

565 570 575

Phe Val Thr Ile Ala Lys Gly Phe Asn Val Pro Ala Val Arg Val Thr

580 585 590

Lys Lys Ser Glu Val Thr Ala Ala Ile Lys Lys Met Leu Glu Thr Pro

595 600 605

Gly Pro Tyr Leu Leu Asp Ile Ile Val Pro His Gln Glu His Val Leu

610 615 620

Pro Met Ile Pro Ser Gly Gly Ala Phe Lys Asp Met Ile Met Glu Gly

625 630 635 640

Asp Gly Arg Thr Ser Tyr

645

<210> 13

<211> 1941

<212> DNA

<213> 小麦 (Triticum aestivum)

<400> 13

atggccgcag ccacctcccc cgccgtcgca ttctcgggcg ccgccgccgc cgccgccgcc 60

atacccaaac ccgcccgcca gcctctcccg cgccaccagc ccgcctcgcg ccgcgcgctc 120

cccgcccgca tcgtcaggtg ctgcgccgcg tcccccgccg ccacctccgt cgcgcctccc 180

gccaccgcgc tccggccgtg gggcccctcc gagccccgca agggcgccga catcctcgtc 240

gaggcgctgg agcgctgcgg catcgtcgac gtcttcgcct accctggcgg cgcgtccatg 300

gagatccacc aggcgctgac gcgctcgcca gtcatcacca accacctctt ccgccacgag 360

cagggggagg cgttcgcggc gtccgggtac gcccgcgcgt ccggccgcgt cggcgtctgc 420

gtcgccacct ccggcccggg ggccaccaac ctcgtctccg cgctcgccga cgctctcctc 480

gactccatcc ccatggtcgc catcacgggc caggtccccc gccgcatgat cggcacggat 540

gtgttccagg agacgcccat cgtggaggtc acgcgctcca tcaccaagca caactacctg 600

gtccttgacg tggaggatat cccccgcgtc atccaggaag ccttcttcct cgcatcctct 660

ggccgcccgg ggccggtgct ggttgatatc cccaaggaca tccagcagca gatggctgtg 720

cctgtctggg acacgccgat gagtttgcca gggtacatcg cccgcctgcc caagccacca 780

tctactgaat cgcttgagca ggtcctgcgt ctggttggcg agtcacggcg cccaattctg 840

tatgttggtg gtggctgcgc tgcatctggt gaggagttgc gccgctttgt tgagctcact 900

gggattccag ttacaactac tcttatgggc cttggcaact tccccagtga cgacccactg 960

tctctgcgca tgctggggat gcatggcact gtgtatgcaa attatgcagt agataaggct 1020

gacctgttgc ttgcatttgg tgtgcggttt gatgatcgtg tgaccgggaa aatcgaggct 1080

tttgcaagca ggtccaagat tgtgcacatt gacattgacc cagctgagat tggcaagaac 1140

aagcagccac atgtctccat ttgtgcagat gttaagcttg ctttacaggg gttgaatgct 1200

ctattaaatg ggagcaaagc acaacagggt ctggattttg gtccatggca caaggagttg 1260

gatcagcaga agagggagtt tcctctagga ttcaagactt ttggtgaggc catcccgccg 1320

caatatgcta tccaggtact ggatgagctg acaaaagggg aggcgatcat tgccaccggt 1380

gttgggcagc atcagatgtg ggcggctcag tattacactt acaagcggcc acggcagtgg 1440

ctgtcttcat ccggtttggg tgcaatggga tttgggttgc cagctgcagc tggcgctgct 1500

gtggccaacc caggtgttac agttgttgac attgatgggg atggtagttt cctcatgaac 1560

attcaggagt tggcgttgat ccgtattgag aacctcccag tgaaggtgat gatattgaac 1620

aaccagcatc tgggaatggt ggtgcagtgg gaggataggt tttacaaggc caaccgggcg 1680

cacacatacc ttggcaaccc agaaaatgag agtgagatat atccagattt tgtgacgatt 1740

gctaaaggat tcaacgttcc ggcagttcgt gtgacgaaga agagcgaagt cactgcagca 1800

atcaagaaga tgcttgagac cccagggcca tacttgttgg atatcattgt cccgcatcag 1860

gagcacgtgc tgcctatgat cccaagcggt ggtgctttta aggacatgat catggagggt 1920

gatggcagga cctcgtactg a 1941

<210> 14

<211> 646

<212> PRT

<213> 小麦 (Triticum aestivum)

<400> 14

Met Ala Ala Ala Thr Ser Pro Ala Val Ala Phe Ser Gly Ala Ala Ala

1 5 10 15

Ala Ala Ala Ala Ile Pro Lys Pro Ala Arg Gln Pro Leu Pro Arg His

20 25 30

Gln Pro Ala Ser Arg Arg Ala Leu Pro Ala Arg Ile Val Arg Cys Cys

35 40 45

Ala Ala Ser Pro Ala Ala Thr Ser Val Ala Pro Pro Ala Thr Ala Leu

50 55 60

Arg Pro Trp Gly Pro Ser Glu Pro Arg Lys Gly Ala Asp Ile Leu Val

65 70 75 80

Glu Ala Leu Glu Arg Cys Gly Ile Val Asp Val Phe Ala Tyr Pro Gly

85 90 95

Gly Ala Ser Met Glu Ile His Gln Ala Leu Thr Arg Ser Pro Val Ile

100 105 110

Thr Asn His Leu Phe Arg His Glu Gln Gly Glu Ala Phe Ala Ala Ser

115 120 125

Gly Tyr Ala Arg Ala Ser Gly Arg Val Gly Val Cys Val Ala Thr Ser

130 135 140

Gly Pro Gly Ala Thr Asn Leu Val Ser Ala Leu Ala Asp Ala Leu Leu

145 150 155 160

Asp Ser Ile Pro Met Val Ala Ile Thr Gly Gln Val Pro Arg Arg Met

165 170 175

Ile Gly Thr Asp Val Phe Gln Glu Thr Pro Ile Val Glu Val Thr Arg

180 185 190

Ser Ile Thr Lys His Asn Tyr Leu Val Leu Asp Val Glu Asp Ile Pro

195 200 205

Arg Val Ile Gln Glu Ala Phe Phe Leu Ala Ser Ser Gly Arg Pro Gly

210 215 220

Pro Val Leu Val Asp Ile Pro Lys Asp Ile Gln Gln Gln Met Ala Val

225 230 235 240

Pro Val Trp Asp Thr Pro Met Ser Leu Pro Gly Tyr Ile Ala Arg Leu

245 250 255

Pro Lys Pro Pro Ser Thr Glu Ser Leu Glu Gln Val Leu Arg Leu Val

260 265 270

Gly Glu Ser Arg Arg Pro Ile Leu Tyr Val Gly Gly Gly Cys Ala Ala

275 280 285

Ser Gly Glu Glu Leu Arg Arg Phe Val Glu Leu Thr Gly Ile Pro Val

290 295 300

Thr Thr Thr Leu Met Gly Leu Gly Asn Phe Pro Ser Asp Asp Pro Leu

305 310 315 320

Ser Leu Arg Met Leu Gly Met His Gly Thr Val Tyr Ala Asn Tyr Ala

325 330 335

Val Asp Lys Ala Asp Leu Leu Leu Ala Phe Gly Val Arg Phe Asp Asp

340 345 350

Arg Val Thr Gly Lys Ile Glu Ala Phe Ala Ser Arg Ser Lys Ile Val

355 360 365

His Ile Asp Ile Asp Pro Ala Glu Ile Gly Lys Asn Lys Gln Pro His

370 375 380

Val Ser Ile Cys Ala Asp Val Lys Leu Ala Leu Gln Gly Leu Asn Ala

385 390 395 400

Leu Leu Asn Gly Ser Lys Ala Gln Gln Gly Leu Asp Phe Gly Pro Trp

405 410 415

His Lys Glu Leu Asp Gln Gln Lys Arg Glu Phe Pro Leu Gly Phe Lys

420 425 430

Thr Phe Gly Glu Ala Ile Pro Pro Gln Tyr Ala Ile Gln Val Leu Asp

435 440 445

Glu Leu Thr Lys Gly Glu Ala Ile Ile Ala Thr Gly Val Gly Gln His

450 455 460

Gln Met Trp Ala Ala Gln Tyr Tyr Thr Tyr Lys Arg Pro Arg Gln Trp

465 470 475 480

Leu Ser Ser Ser Gly Leu Gly Ala Met Gly Phe Gly Leu Pro Ala Ala

485 490 495

Ala Gly Ala Ala Val Ala Asn Pro Gly Val Thr Val Val Asp Ile Asp

500 505 510

Gly Asp Gly Ser Phe Leu Met Asn Ile Gln Glu Leu Ala Leu Ile Arg

515 520 525

Ile Glu Asn Leu Pro Val Lys Val Met Ile Leu Asn Asn Gln His Leu

530 535 540

Gly Met Val Val Gln Trp Glu Asp Arg Phe Tyr Lys Ala Asn Arg Ala

545 550 555 560

His Thr Tyr Leu Gly Asn Pro Glu Asn Glu Ser Glu Ile Tyr Pro Asp

565 570 575

Phe Val Thr Ile Ala Lys Gly Phe Asn Val Pro Ala Val Arg Val Thr

580 585 590

Lys Lys Ser Glu Val Thr Ala Ala Ile Lys Lys Met Leu Glu Thr Pro

595 600 605

Gly Pro Tyr Leu Leu Asp Ile Ile Val Pro His Gln Glu His Val Leu

610 615 620

Pro Met Ile Pro Ser Gly Gly Ala Phe Lys Asp Met Ile Met Glu Gly

625 630 635 640

Asp Gly Arg Thr Ser Tyr

645

<210> 15

<211> 1941

<212> DNA

<213> 小麦 (Triticum aestivum)

<400> 15

atggccgcag ccacctcccc cgccgtcgca ttctcgggcg ccgccgccgc cgccgccgcc 60

atacccaaac ccgcccgcca gcctctcccg cgccaccagc ccgcctcgcg ccgcgcgctc 120

cccgcccgca tcgtcaggtg ctgcgccgcg tcccccgccg ccacctccgt cgcgcctccc 180

gccaccgcgc tccggccgtg gggcccctcc gagccccgca agggcgccga catcctcgtc 240

gaggcgctgg agcgctgcgg catcgtcgac gtcttcgcct accctggcgg cgcgtccatg 300

gagatccacc aggcgctgac gcgctcgcca gtcatcacca accacctctt ccgccacgag 360

cagggggagg cgttcgcggc gtccgggtac gcccgcgcgt ccggccgcgt cggcgtctgc 420

gtcgccacct ccggcccggg ggccaccaac ctcgtctccg cgctcgccga cgctctcctc 480

gactccatcc ccatggtcgc catcacgggc caggtccccc gccgcatgat cggcacggat 540

gcgttccagg agacgcccat cgtggaggtc acgcgctcca tcaccaagca caactacctg 600

gtccttgacg tggaggatat cccccgcgtc atccaggaag ccttcttcct cgcatcctct 660

ggccgcccgg ggccggtgct ggttgatatc cccaaggaca tccagcagca gatggctgtg 720

cctgtctggg acacgccgat gagtttgcca gggtacatcg cccgcctgcc caagccacca 780

tctactgaat cgcttgagca ggtcctgcgt ctggttggcg agtcacggcg cccaattctg 840

tatgttggtg gtggctgcgc tgcatctggt gaggagttgc gccgctttgt tgagctcact 900

gggattccag ttacaactac tcttatgggc cttggcaact tccccagtga cgacccactg 960

tctctgcgca tgctggggat gcatggcact gtgtatgcaa attatgcagt agataaggct 1020

gacctgttgc ttgcatttgg tgtgcggttt gatgatcgtg tgaccgggaa aatcgaggct 1080

tttgcaagca ggtccaagat tgtgcacatt gacattgacc cagctgagat tggcaagaac 1140

aagcagccac atgtctccat ttgtgcagat gttaagcttg ctttacaggg gttgaatgct 1200

ctattaaatg ggagcaaagc acaacagggt ctggattttg gtccatggca caaggagttg 1260

gatcagcaga agagggagtt tcctctagga ttcaagactt ttggtgaggc catcccgccg 1320

caatatgcta tccaggtact ggatgagctg acaaaagggg aggcgatcat tgccaccggt 1380

gttgggcagc atcagatgtg ggcggctcag tattacactt acaagcggcc acggcagtgg 1440

ctgtcttcat ccggtttggg tgcaatggga tttgggttgc cagctgcagc tggcgctgct 1500

gtggccaacc caggtgttac agttgttgac attgatgggg atggtagttt cctcatgaac 1560

attcaggagt tggcgttgat ccgtattgag aacctcccag tgaaggtgat gatattgaac 1620

aaccagcatc tgggaatggt ggtgcagtgg gaggataggt tttacaaggc caaccgggcg 1680

cacacatacc ttggcaaccc agaaaatgag agtgagatat atccagattt tgtgacgatt 1740

gctaaaggat tcaacgttcc ggcagttcgt gtgacgaaga agagcgaagt cactgcagca 1800

atcaagaaga tgcttgagac cccagggcca tacttgttgg atatcattgt cccgcatcag 1860

gagcacgtgc tgcctatgat cccaaacggt ggtgctttta aggacatgat catggagggt 1920

gatggcagga cctcgtactg a 1941

<210> 16

<211> 646

<212> PRT

<213> 小麦 (Triticum aestivum)

<400> 16

Met Ala Ala Ala Thr Ser Pro Ala Val Ala Phe Ser Gly Ala Ala Ala

1 5 10 15

Ala Ala Ala Ala Ile Pro Lys Pro Ala Arg Gln Pro Leu Pro Arg His

20 25 30

Gln Pro Ala Ser Arg Arg Ala Leu Pro Ala Arg Ile Val Arg Cys Cys

35 40 45

Ala Ala Ser Pro Ala Ala Thr Ser Val Ala Pro Pro Ala Thr Ala Leu

50 55 60

Arg Pro Trp Gly Pro Ser Glu Pro Arg Lys Gly Ala Asp Ile Leu Val

65 70 75 80

Glu Ala Leu Glu Arg Cys Gly Ile Val Asp Val Phe Ala Tyr Pro Gly

85 90 95

Gly Ala Ser Met Glu Ile His Gln Ala Leu Thr Arg Ser Pro Val Ile

100 105 110

Thr Asn His Leu Phe Arg His Glu Gln Gly Glu Ala Phe Ala Ala Ser

115 120 125

Gly Tyr Ala Arg Ala Ser Gly Arg Val Gly Val Cys Val Ala Thr Ser

130 135 140

Gly Pro Gly Ala Thr Asn Leu Val Ser Ala Leu Ala Asp Ala Leu Leu

145 150 155 160

Asp Ser Ile Pro Met Val Ala Ile Thr Gly Gln Val Pro Arg Arg Met

165 170 175

Ile Gly Thr Asp Ala Phe Gln Glu Thr Pro Ile Val Glu Val Thr Arg

180 185 190

Ser Ile Thr Lys His Asn Tyr Leu Val Leu Asp Val Glu Asp Ile Pro

195 200 205

Arg Val Ile Gln Glu Ala Phe Phe Leu Ala Ser Ser Gly Arg Pro Gly

210 215 220

Pro Val Leu Val Asp Ile Pro Lys Asp Ile Gln Gln Gln Met Ala Val

225 230 235 240

Pro Val Trp Asp Thr Pro Met Ser Leu Pro Gly Tyr Ile Ala Arg Leu

245 250 255

Pro Lys Pro Pro Ser Thr Glu Ser Leu Glu Gln Val Leu Arg Leu Val

260 265 270

Gly Glu Ser Arg Arg Pro Ile Leu Tyr Val Gly Gly Gly Cys Ala Ala

275 280 285

Ser Gly Glu Glu Leu Arg Arg Phe Val Glu Leu Thr Gly Ile Pro Val

290 295 300

Thr Thr Thr Leu Met Gly Leu Gly Asn Phe Pro Ser Asp Asp Pro Leu

305 310 315 320

Ser Leu Arg Met Leu Gly Met His Gly Thr Val Tyr Ala Asn Tyr Ala

325 330 335

Val Asp Lys Ala Asp Leu Leu Leu Ala Phe Gly Val Arg Phe Asp Asp

340 345 350

Arg Val Thr Gly Lys Ile Glu Ala Phe Ala Ser Arg Ser Lys Ile Val

355 360 365

His Ile Asp Ile Asp Pro Ala Glu Ile Gly Lys Asn Lys Gln Pro His

370 375 380

Val Ser Ile Cys Ala Asp Val Lys Leu Ala Leu Gln Gly Leu Asn Ala

385 390 395 400

Leu Leu Asn Gly Ser Lys Ala Gln Gln Gly Leu Asp Phe Gly Pro Trp

405 410 415

His Lys Glu Leu Asp Gln Gln Lys Arg Glu Phe Pro Leu Gly Phe Lys

420 425 430

Thr Phe Gly Glu Ala Ile Pro Pro Gln Tyr Ala Ile Gln Val Leu Asp

435 440 445

Glu Leu Thr Lys Gly Glu Ala Ile Ile Ala Thr Gly Val Gly Gln His

450 455 460

Gln Met Trp Ala Ala Gln Tyr Tyr Thr Tyr Lys Arg Pro Arg Gln Trp

465 470 475 480

Leu Ser Ser Ser Gly Leu Gly Ala Met Gly Phe Gly Leu Pro Ala Ala

485 490 495

Ala Gly Ala Ala Val Ala Asn Pro Gly Val Thr Val Val Asp Ile Asp

500 505 510

Gly Asp Gly Ser Phe Leu Met Asn Ile Gln Glu Leu Ala Leu Ile Arg

515 520 525

Ile Glu Asn Leu Pro Val Lys Val Met Ile Leu Asn Asn Gln His Leu

530 535 540

Gly Met Val Val Gln Trp Glu Asp Arg Phe Tyr Lys Ala Asn Arg Ala

545 550 555 560

His Thr Tyr Leu Gly Asn Pro Glu Asn Glu Ser Glu Ile Tyr Pro Asp

565 570 575

Phe Val Thr Ile Ala Lys Gly Phe Asn Val Pro Ala Val Arg Val Thr

580 585 590

Lys Lys Ser Glu Val Thr Ala Ala Ile Lys Lys Met Leu Glu Thr Pro

595 600 605

Gly Pro Tyr Leu Leu Asp Ile Ile Val Pro His Gln Glu His Val Leu

610 615 620

Pro Met Ile Pro Asn Gly Gly Ala Phe Lys Asp Met Ile Met Glu Gly

625 630 635 640

Asp Gly Arg Thr Ser Tyr

645

<210> 17

<211> 1944

<212> DNA

<213> 小麦 (Triticum aestivum)

<400> 17

atggccgccg ccacctcccc cgccgtcgca ttctccggcg ccgccgccgc cgccgccgcc 60

atgcccaagc ccgcccgcca gcctctcccg cgccaccagc ccgcctcgcg ccgcgcgctc 120

cccgcccgcg tcgtcaggtg ctgcgccgcg tcccccgctg ctgccacctc cgccgcgccc 180

cccgccaccg cgctccggcc ctggggcccg tccgagcccc gcaagggcgc cgacatcctc 240

gtcgaggcgc tcgagcgctg cggcatcgtc gacgtattcg cctaccccgg cggcacgtcc 300

atggagatcc accaggcgct gacgcgctcg cccgtcatca ccaaccacct cttccgccac 360

gagcaggggg aggcgttcgc ggcgtccggc tacgcccgcg cgtccggccg cgtcggcgtc 420

tgcgtcgcca cctccggccc gggggccacc aacctcgtct ccgcgctcgc tgacgccctc 480

ctcgactcca tccccatggt cgccatcacg ggccaggtcc cccgccgcat gatcggcacg 540

gacgcgttcc aggagacgcc catagtggag gtcacgcgct ccatcaccaa gcacaactac 600

ctggtccttg acgtggagga tatcccccgc gtcatccagg aagccttctt cctcgcgtcc 660

tctggccgcc cggggccggt gctggttgat atccccaagg atatccagca gcagatggcc 720

gtgcctatct gggacacgcc gatgagtttg ccagggtaca tcgcccgcct gcccaagcca 780

ccatctactg aatcgcttga gcaggtcctg cgtctggttg gcgagtcacg gcgcccaatt 840

ctgtatgttg gtggtggctg cgctgcatcc ggcgaggagt tgcgccgctt tgttgagctc 900

actgggattc cggttacaac tactctgatg ggccttggca acttccccag cgacgaccca 960

ctgtctctgc gcatgcttgg gatgcatggc actgtgtatg caaattatgc agtcgataag 1020

gctgacctgt tgcttgcatt tggtgtgcgg tttgatgatc gcgtgactgg gaaaatcgag 1080

gcctttgcaa gcaggtccaa gattgtgcac attgacattg acccagctga gattggcaag 1140

aacaagcagc cacatgtctc catttgtgca gatgttaagc ttgctttaca ggggttgaat 1200

gctctattaa atgggagcaa agcacaacag ggtctggatt ttggtccatg gcacaaggag 1260

ttggatcagc agaagaggga gtttcctcta ggattcaaga cttttggcga ggccatcccg 1320

ccgcaatatg ctatccaggt actggatgag ctgacaaaag gggaggcgat cattgctact 1380

ggtgttgggc agcaccagat gtgggcggct cagtattaca cttacaagcg gccacggcag 1440

tggctgtctt cgtctggttt gggggcaatg ggatttgggt taccagctgc agctggcgct 1500

gctgtggcca acccaggtgt tacagttgtt gacattgatg gagatggtag tttcctcatg 1560

aacattcagg agttggcatt gatccgtatt gagaacctcc ctgtgaaggt gatgatattg 1620

aacaaccagc atctgggaat ggtggtgcaa tgggaggata ggttttacaa ggccaatcgg 1680

gcgcacacat accttggcaa cccagaaaat gagagtgaga tatatccaga ttttgtgacg 1740

attgctaaag gattcaacgt tccggcagtt cgtgtgacga agaagagcga agtcactgca 1800

gcaatcaaga agatgcttga gaccccaggg ccatacttgt tggatatcat cgtcccgcat 1860

caggagcacg tgctgcctat gatcccaagc ggtggtgctt tcaaggacat gatcatggag 1920

ggtgatggca ggacctcgta ctga 1944

<210> 18

<211> 647

<212> PRT

<213> 小麦 (Triticum aestivum)

<400> 18

Met Ala Ala Ala Thr Ser Pro Ala Val Ala Phe Ser Gly Ala Ala Ala

1 5 10 15

Ala Ala Ala Ala Met Pro Lys Pro Ala Arg Gln Pro Leu Pro Arg His

20 25 30

Gln Pro Ala Ser Arg Arg Ala Leu Pro Ala Arg Val Val Arg Cys Cys

35 40 45

Ala Ala Ser Pro Ala Ala Ala Thr Ser Ala Ala Pro Pro Ala Thr Ala

50 55 60

Leu Arg Pro Trp Gly Pro Ser Glu Pro Arg Lys Gly Ala Asp Ile Leu

65 70 75 80

Val Glu Ala Leu Glu Arg Cys Gly Ile Val Asp Val Phe Ala Tyr Pro

85 90 95

Gly Gly Thr Ser Met Glu Ile His Gln Ala Leu Thr Arg Ser Pro Val

100 105 110

Ile Thr Asn His Leu Phe Arg His Glu Gln Gly Glu Ala Phe Ala Ala

115 120 125

Ser Gly Tyr Ala Arg Ala Ser Gly Arg Val Gly Val Cys Val Ala Thr

130 135 140

Ser Gly Pro Gly Ala Thr Asn Leu Val Ser Ala Leu Ala Asp Ala Leu

145 150 155 160

Leu Asp Ser Ile Pro Met Val Ala Ile Thr Gly Gln Val Pro Arg Arg

165 170 175

Met Ile Gly Thr Asp Ala Phe Gln Glu Thr Pro Ile Val Glu Val Thr

180 185 190

Arg Ser Ile Thr Lys His Asn Tyr Leu Val Leu Asp Val Glu Asp Ile

195 200 205

Pro Arg Val Ile Gln Glu Ala Phe Phe Leu Ala Ser Ser Gly Arg Pro

210 215 220

Gly Pro Val Leu Val Asp Ile Pro Lys Asp Ile Gln Gln Gln Met Ala

225 230 235 240

Val Pro Ile Trp Asp Thr Pro Met Ser Leu Pro Gly Tyr Ile Ala Arg

245 250 255

Leu Pro Lys Pro Pro Ser Thr Glu Ser Leu Glu Gln Val Leu Arg Leu

260 265 270

Val Gly Glu Ser Arg Arg Pro Ile Leu Tyr Val Gly Gly Gly Cys Ala

275 280 285

Ala Ser Gly Glu Glu Leu Arg Arg Phe Val Glu Leu Thr Gly Ile Pro

290 295 300

Val Thr Thr Thr Leu Met Gly Leu Gly Asn Phe Pro Ser Asp Asp Pro

305 310 315 320

Leu Ser Leu Arg Met Leu Gly Met His Gly Thr Val Tyr Ala Asn Tyr

325 330 335

Ala Val Asp Lys Ala Asp Leu Leu Leu Ala Phe Gly Val Arg Phe Asp

340 345 350

Asp Arg Val Thr Gly Lys Ile Glu Ala Phe Ala Ser Arg Ser Lys Ile

355 360 365

Val His Ile Asp Ile Asp Pro Ala Glu Ile Gly Lys Asn Lys Gln Pro

370 375 380

His Val Ser Ile Cys Ala Asp Val Lys Leu Ala Leu Gln Gly Leu Asn

385 390 395 400

Ala Leu Leu Asn Gly Ser Lys Ala Gln Gln Gly Leu Asp Phe Gly Pro

405 410 415

Trp His Lys Glu Leu Asp Gln Gln Lys Arg Glu Phe Pro Leu Gly Phe

420 425 430

Lys Thr Phe Gly Glu Ala Ile Pro Pro Gln Tyr Ala Ile Gln Val Leu

435 440 445

Asp Glu Leu Thr Lys Gly Glu Ala Ile Ile Ala Thr Gly Val Gly Gln

450 455 460

His Gln Met Trp Ala Ala Gln Tyr Tyr Thr Tyr Lys Arg Pro Arg Gln

465 470 475 480

Trp Leu Ser Ser Ser Gly Leu Gly Ala Met Gly Phe Gly Leu Pro Ala

485 490 495

Ala Ala Gly Ala Ala Val Ala Asn Pro Gly Val Thr Val Val Asp Ile

500 505 510

Asp Gly Asp Gly Ser Phe Leu Met Asn Ile Gln Glu Leu Ala Leu Ile

515 520 525

Arg Ile Glu Asn Leu Pro Val Lys Val Met Ile Leu Asn Asn Gln His

530 535 540

Leu Gly Met Val Val Gln Trp Glu Asp Arg Phe Tyr Lys Ala Asn Arg

545 550 555 560

Ala His Thr Tyr Leu Gly Asn Pro Glu Asn Glu Ser Glu Ile Tyr Pro

565 570 575

Asp Phe Val Thr Ile Ala Lys Gly Phe Asn Val Pro Ala Val Arg Val

580 585 590

Thr Lys Lys Ser Glu Val Thr Ala Ala Ile Lys Lys Met Leu Glu Thr

595 600 605

Pro Gly Pro Tyr Leu Leu Asp Ile Ile Val Pro His Gln Glu His Val

610 615 620

Leu Pro Met Ile Pro Ser Gly Gly Ala Phe Lys Asp Met Ile Met Glu

625 630 635 640

Gly Asp Gly Arg Thr Ser Tyr

645

<210> 19

<211> 1944

<212> DNA

<213> 小麦 (Triticum aestivum)

<400> 19

atggccgccg ccacctcccc cgccgtcgca ttctccggcg ccgccgccgc cgccgccgcc 60

atgcccaagc ccgcccgcca gcctctcccg cgccaccagc ccgcctcgcg ccgcgcgctc 120

cccgcccgcg tcgtcaggtg ctgcgccgcg tcccccgctg ctgccacctc cgccgcgccc 180

cccgccaccg cgctccggcc ctggggcccg tccgagcccc gcaagggcgc cgacatcctc 240

gtcgaggcgc tcgagcgctg cggcatcgtc gacgtattcg cctaccccgg cggcgcgtcc 300

atggagatcc accaggcgct gacgcgctcg cccgtcatca ccaaccacct cttccgccac 360

gagcaggggg aggcgttcgc ggcgtccggc tacgcccgcg cgtccggccg cgtcggcgtc 420

tgcgtcgcca cctccggccc gggggccacc aacctcgtct ccgcgctcgc tgacgccctc 480

ctcgactcca tccccatggt cgccatcacg ggccaggtcc tccgccgcat gatcggcacg 540

gacgcgttcc aggagacgcc catagtggag gtcacgcgct ccatcaccaa gcacaactac 600

ctggtccttg acgtggagga tatcccccgc gtcatccagg aagccttctt cctcgcgtcc 660

tctggccgcc cggggccggt gctggttgat atccccaagg atatccagca gcagatggcc 720

gtgcctatct gggacacgcc gatgagtttg ccagggtaca tcgcccgcct gcccaagcca 780

ccatctactg aatcgcttga gcaggtcctg cgtctggttg gcgagtcacg gcgcccaatt 840

ctgtatgttg gtggtggctg cgctgcatcc ggcgaggagt tgcgccgctt tgttgagctc 900

actgggattc cggttacaac tactctgatg ggccttggca acttccccag cgacgaccca 960

ctgtctctgc gcatgcttgg gatgcatggc actgtgtatg caaattatgc agtcgataag 1020

gctgacctgt tgcttgcatt tggtgtgcgg tttgatgatc gcgtgactgg gaaaatcgag 1080

gcctttgcaa gcaggtccaa gattgtgcac attgacattg acccagctga gattggcaag 1140

aacaagcagc cacatgtctc catttgtgca gatgttaagc ttgctttaca ggggttgaat 1200

gctctattaa atgggagcaa agcacaacag ggtctggatt ttggtccatg gcacaaggag 1260

ttggatcagc agaagaggga gtttcctcta ggattcaaga cttttggcga ggccatcccg 1320

ccgcaatatg ctatccaggt actggatgag ctgacaaaag gggaggcgat cattgctact 1380

ggtgttgggc agcaccagat gtgggcggct cagtattaca cttacaagcg gccacggcag 1440

tggctgtctt cgtctggttt gggggcaatg ggatttgggt taccagctgc agctggcgct 1500

gctgtggcca acccaggtgt tacagttgtt gacattgatg gagatggtag tttcctcatg 1560

aacattcagg agttggcatt gatccgtatt gagaacctcc ctgtgaaggt gatgatattg 1620

aacaaccagc atctgggaat ggtggtgcaa tgggaggata ggttttacaa ggccaatcgg 1680

gcgcacacat accttggcaa cccagaaaat gagagtgaga tatatccaga ttttgtgacg 1740

attgctaaag gattcaacgt tccggcagtt cgtgtgacga agaagagcga agtcactgca 1800

gcaatcaaga agatgcttga gaccccaggg ccatacttgt tggatatcat cgtcccgcat 1860

caggagcacg tgctgcctat gatcccaagc ggtggtgctt tcaaggacat gatcatggag 1920

ggtgatggca ggacctcgta ctga 1944

<210> 20

<211> 647

<212> PRT

<213> 小麦 (Triticum aestivum)

<400> 20

Met Ala Ala Ala Thr Ser Pro Ala Val Ala Phe Ser Gly Ala Ala Ala

1 5 10 15

Ala Ala Ala Ala Met Pro Lys Pro Ala Arg Gln Pro Leu Pro Arg His

20 25 30

Gln Pro Ala Ser Arg Arg Ala Leu Pro Ala Arg Val Val Arg Cys Cys

35 40 45

Ala Ala Ser Pro Ala Ala Ala Thr Ser Ala Ala Pro Pro Ala Thr Ala

50 55 60

Leu Arg Pro Trp Gly Pro Ser Glu Pro Arg Lys Gly Ala Asp Ile Leu

65 70 75 80

Val Glu Ala Leu Glu Arg Cys Gly Ile Val Asp Val Phe Ala Tyr Pro

85 90 95

Gly Gly Ala Ser Met Glu Ile His Gln Ala Leu Thr Arg Ser Pro Val

100 105 110

Ile Thr Asn His Leu Phe Arg His Glu Gln Gly Glu Ala Phe Ala Ala

115 120 125

Ser Gly Tyr Ala Arg Ala Ser Gly Arg Val Gly Val Cys Val Ala Thr

130 135 140

Ser Gly Pro Gly Ala Thr Asn Leu Val Ser Ala Leu Ala Asp Ala Leu

145 150 155 160

Leu Asp Ser Ile Pro Met Val Ala Ile Thr Gly Gln Val Leu Arg Arg

165 170 175

Met Ile Gly Thr Asp Ala Phe Gln Glu Thr Pro Ile Val Glu Val Thr

180 185 190

Arg Ser Ile Thr Lys His Asn Tyr Leu Val Leu Asp Val Glu Asp Ile

195 200 205

Pro Arg Val Ile Gln Glu Ala Phe Phe Leu Ala Ser Ser Gly Arg Pro

210 215 220

Gly Pro Val Leu Val Asp Ile Pro Lys Asp Ile Gln Gln Gln Met Ala

225 230 235 240

Val Pro Ile Trp Asp Thr Pro Met Ser Leu Pro Gly Tyr Ile Ala Arg

245 250 255

Leu Pro Lys Pro Pro Ser Thr Glu Ser Leu Glu Gln Val Leu Arg Leu

260 265 270

Val Gly Glu Ser Arg Arg Pro Ile Leu Tyr Val Gly Gly Gly Cys Ala

275 280 285

Ala Ser Gly Glu Glu Leu Arg Arg Phe Val Glu Leu Thr Gly Ile Pro

290 295 300

Val Thr Thr Thr Leu Met Gly Leu Gly Asn Phe Pro Ser Asp Asp Pro

305 310 315 320

Leu Ser Leu Arg Met Leu Gly Met His Gly Thr Val Tyr Ala Asn Tyr

325 330 335

Ala Val Asp Lys Ala Asp Leu Leu Leu Ala Phe Gly Val Arg Phe Asp

340 345 350

Asp Arg Val Thr Gly Lys Ile Glu Ala Phe Ala Ser Arg Ser Lys Ile

355 360 365

Val His Ile Asp Ile Asp Pro Ala Glu Ile Gly Lys Asn Lys Gln Pro

370 375 380

His Val Ser Ile Cys Ala Asp Val Lys Leu Ala Leu Gln Gly Leu Asn

385 390 395 400

Ala Leu Leu Asn Gly Ser Lys Ala Gln Gln Gly Leu Asp Phe Gly Pro

405 410 415

Trp His Lys Glu Leu Asp Gln Gln Lys Arg Glu Phe Pro Leu Gly Phe

420 425 430

Lys Thr Phe Gly Glu Ala Ile Pro Pro Gln Tyr Ala Ile Gln Val Leu

435 440 445

Asp Glu Leu Thr Lys Gly Glu Ala Ile Ile Ala Thr Gly Val Gly Gln

450 455 460

His Gln Met Trp Ala Ala Gln Tyr Tyr Thr Tyr Lys Arg Pro Arg Gln

465 470 475 480

Trp Leu Ser Ser Ser Gly Leu Gly Ala Met Gly Phe Gly Leu Pro Ala

485 490 495

Ala Ala Gly Ala Ala Val Ala Asn Pro Gly Val Thr Val Val Asp Ile

500 505 510

Asp Gly Asp Gly Ser Phe Leu Met Asn Ile Gln Glu Leu Ala Leu Ile

515 520 525

Arg Ile Glu Asn Leu Pro Val Lys Val Met Ile Leu Asn Asn Gln His

530 535 540

Leu Gly Met Val Val Gln Trp Glu Asp Arg Phe Tyr Lys Ala Asn Arg

545 550 555 560

Ala His Thr Tyr Leu Gly Asn Pro Glu Asn Glu Ser Glu Ile Tyr Pro

565 570 575

Asp Phe Val Thr Ile Ala Lys Gly Phe Asn Val Pro Ala Val Arg Val

580 585 590

Thr Lys Lys Ser Glu Val Thr Ala Ala Ile Lys Lys Met Leu Glu Thr

595 600 605

Pro Gly Pro Tyr Leu Leu Asp Ile Ile Val Pro His Gln Glu His Val

610 615 620

Leu Pro Met Ile Pro Ser Gly Gly Ala Phe Lys Asp Met Ile Met Glu

625 630 635 640

Gly Asp Gly Arg Thr Ser Tyr

645

<210> 21

<211> 1944

<212> DNA

<213> 小麦 (Triticum aestivum)

<400> 21

atggccgccg ccacctcccc cgccgtcgca ttctccggcg ccgccgccgc cgccgccgcc 60

atgcccaagc ccgcccgcca gcctctcccg cgccaccagc ccgcctcgcg ccgcgcgctc 120

cccgcccgcg tcgtcaggtg ctgcgccgcg tcccccgctg ctgccacctc cgccgcgccc 180

cccgccaccg cgctccggcc ctggggcccg tccgagcccc gcaagggcgc cgacatcctc 240

gtcgaggcgc tcgagcgctg cggcatcgtc gacgtattcg cctaccccgg cggcgcgtcc 300

atggagatcc accaggcgct gacgcgctcg cccgtcatca ccaaccacct cttccgccac 360

gagcaggggg aggcgttcgc ggcgtccggc tacgcccgcg cgtccggccg cgtcggcgtc 420

tgcgtcgcca cctccggccc gggggccacc aacctcgtct ccgcgctcgc tgacgccctc 480

ctcgactcca tccccatggt cgccatcacg ggccaggtcc cccgccgcat gatcggcacg 540

gacgtgttcc aggagacgcc catagtggag gtcacgcgct ccatcaccaa gcacaactac 600

ctggtccttg acgtggagga tatcccccgc gtcatccagg aagccttctt cctcgcgtcc 660

tctggccgcc cggggccggt gctggttgat atccccaagg atatccagca gcagatggcc 720

gtgcctatct gggacacgcc gatgagtttg ccagggtaca tcgcccgcct gcccaagcca 780

ccatctactg aatcgcttga gcaggtcctg cgtctggttg gcgagtcacg gcgcccaatt 840

ctgtatgttg gtggtggctg cgctgcatcc ggcgaggagt tgcgccgctt tgttgagctc 900

actgggattc cggttacaac tactctgatg ggccttggca acttccccag cgacgaccca 960

ctgtctctgc gcatgcttgg gatgcatggc actgtgtatg caaattatgc agtcgataag 1020

gctgacctgt tgcttgcatt tggtgtgcgg tttgatgatc gcgtgactgg gaaaatcgag 1080

gcctttgcaa gcaggtccaa gattgtgcac attgacattg acccagctga gattggcaag 1140

aacaagcagc cacatgtctc catttgtgca gatgttaagc ttgctttaca ggggttgaat 1200

gctctattaa atgggagcaa agcacaacag ggtctggatt ttggtccatg gcacaaggag 1260

ttggatcagc agaagaggga gtttcctcta ggattcaaga cttttggcga ggccatcccg 1320

ccgcaatatg ctatccaggt actggatgag ctgacaaaag gggaggcgat cattgctact 1380

ggtgttgggc agcaccagat gtgggcggct cagtattaca cttacaagcg gccacggcag 1440

tggctgtctt cgtctggttt gggggcaatg ggatttgggt taccagctgc agctggcgct 1500

gctgtggcca acccaggtgt tacagttgtt gacattgatg gagatggtag tttcctcatg 1560

aacattcagg agttggcatt gatccgtatt gagaacctcc ctgtgaaggt gatgatattg 1620

aacaaccagc atctgggaat ggtggtgcaa tgggaggata ggttttacaa ggccaatcgg 1680

gcgcacacat accttggcaa cccagaaaat gagagtgaga tatatccaga ttttgtgacg 1740

attgctaaag gattcaacgt tccggcagtt cgtgtgacga agaagagcga agtcactgca 1800

gcaatcaaga agatgcttga gaccccaggg ccatacttgt tggatatcat cgtcccgcat 1860

caggagcacg tgctgcctat gatcccaagc ggtggtgctt tcaaggacat gatcatggag 1920

ggtgatggca ggacctcgta ctga 1944

<210> 22

<211> 647

<212> PRT

<213> 小麦 (Triticum aestivum)

<400> 22

Met Ala Ala Ala Thr Ser Pro Ala Val Ala Phe Ser Gly Ala Ala Ala

1 5 10 15

Ala Ala Ala Ala Met Pro Lys Pro Ala Arg Gln Pro Leu Pro Arg His

20 25 30

Gln Pro Ala Ser Arg Arg Ala Leu Pro Ala Arg Val Val Arg Cys Cys

35 40 45

Ala Ala Ser Pro Ala Ala Ala Thr Ser Ala Ala Pro Pro Ala Thr Ala

50 55 60

Leu Arg Pro Trp Gly Pro Ser Glu Pro Arg Lys Gly Ala Asp Ile Leu

65 70 75 80

Val Glu Ala Leu Glu Arg Cys Gly Ile Val Asp Val Phe Ala Tyr Pro

85 90 95

Gly Gly Ala Ser Met Glu Ile His Gln Ala Leu Thr Arg Ser Pro Val

100 105 110

Ile Thr Asn His Leu Phe Arg His Glu Gln Gly Glu Ala Phe Ala Ala

115 120 125

Ser Gly Tyr Ala Arg Ala Ser Gly Arg Val Gly Val Cys Val Ala Thr

130 135 140

Ser Gly Pro Gly Ala Thr Asn Leu Val Ser Ala Leu Ala Asp Ala Leu

145 150 155 160

Leu Asp Ser Ile Pro Met Val Ala Ile Thr Gly Gln Val Pro Arg Arg

165 170 175

Met Ile Gly Thr Asp Val Phe Gln Glu Thr Pro Ile Val Glu Val Thr

180 185 190

Arg Ser Ile Thr Lys His Asn Tyr Leu Val Leu Asp Val Glu Asp Ile

195 200 205

Pro Arg Val Ile Gln Glu Ala Phe Phe Leu Ala Ser Ser Gly Arg Pro

210 215 220

Gly Pro Val Leu Val Asp Ile Pro Lys Asp Ile Gln Gln Gln Met Ala

225 230 235 240

Val Pro Ile Trp Asp Thr Pro Met Ser Leu Pro Gly Tyr Ile Ala Arg

245 250 255

Leu Pro Lys Pro Pro Ser Thr Glu Ser Leu Glu Gln Val Leu Arg Leu

260 265 270

Val Gly Glu Ser Arg Arg Pro Ile Leu Tyr Val Gly Gly Gly Cys Ala

275 280 285

Ala Ser Gly Glu Glu Leu Arg Arg Phe Val Glu Leu Thr Gly Ile Pro

290 295 300

Val Thr Thr Thr Leu Met Gly Leu Gly Asn Phe Pro Ser Asp Asp Pro

305 310 315 320

Leu Ser Leu Arg Met Leu Gly Met His Gly Thr Val Tyr Ala Asn Tyr

325 330 335

Ala Val Asp Lys Ala Asp Leu Leu Leu Ala Phe Gly Val Arg Phe Asp

340 345 350

Asp Arg Val Thr Gly Lys Ile Glu Ala Phe Ala Ser Arg Ser Lys Ile

355 360 365

Val His Ile Asp Ile Asp Pro Ala Glu Ile Gly Lys Asn Lys Gln Pro

370 375 380

His Val Ser Ile Cys Ala Asp Val Lys Leu Ala Leu Gln Gly Leu Asn

385 390 395 400

Ala Leu Leu Asn Gly Ser Lys Ala Gln Gln Gly Leu Asp Phe Gly Pro

405 410 415

Trp His Lys Glu Leu Asp Gln Gln Lys Arg Glu Phe Pro Leu Gly Phe

420 425 430

Lys Thr Phe Gly Glu Ala Ile Pro Pro Gln Tyr Ala Ile Gln Val Leu

435 440 445

Asp Glu Leu Thr Lys Gly Glu Ala Ile Ile Ala Thr Gly Val Gly Gln

450 455 460

His Gln Met Trp Ala Ala Gln Tyr Tyr Thr Tyr Lys Arg Pro Arg Gln

465 470 475 480

Trp Leu Ser Ser Ser Gly Leu Gly Ala Met Gly Phe Gly Leu Pro Ala

485 490 495

Ala Ala Gly Ala Ala Val Ala Asn Pro Gly Val Thr Val Val Asp Ile

500 505 510

Asp Gly Asp Gly Ser Phe Leu Met Asn Ile Gln Glu Leu Ala Leu Ile

515 520 525

Arg Ile Glu Asn Leu Pro Val Lys Val Met Ile Leu Asn Asn Gln His

530 535 540

Leu Gly Met Val Val Gln Trp Glu Asp Arg Phe Tyr Lys Ala Asn Arg

545 550 555 560

Ala His Thr Tyr Leu Gly Asn Pro Glu Asn Glu Ser Glu Ile Tyr Pro

565 570 575

Asp Phe Val Thr Ile Ala Lys Gly Phe Asn Val Pro Ala Val Arg Val

580 585 590

Thr Lys Lys Ser Glu Val Thr Ala Ala Ile Lys Lys Met Leu Glu Thr

595 600 605

Pro Gly Pro Tyr Leu Leu Asp Ile Ile Val Pro His Gln Glu His Val

610 615 620

Leu Pro Met Ile Pro Ser Gly Gly Ala Phe Lys Asp Met Ile Met Glu

625 630 635 640

Gly Asp Gly Arg Thr Ser Tyr

645

<210> 23

<211> 1944

<212> DNA

<213> 小麦 (Triticum aestivum)

<400> 23

atggccgccg ccacctcccc cgccgtcgca ttctccggcg ccgccgccgc cgccgccgcc 60

atgcccaagc ccgcccgcca gcctctcccg cgccaccagc ccgcctcgcg ccgcgcgctc 120

cccgcccgcg tcgtcaggtg ctgcgccgcg tcccccgctg ctgccacctc cgccgcgccc 180

cccgccaccg cgctccggcc ctggggcccg tccgagcccc gcaagggcgc cgacatcctc 240

gtcgaggcgc tcgagcgctg cggcatcgtc gacgtattcg cctaccccgg cggcgcgtcc 300

atggagatcc accaggcgct gacgcgctcg cccgtcatca ccaaccacct cttccgccac 360

gagcaggggg aggcgttcgc ggcgtccggc tacgcccgcg cgtccggccg cgtcggcgtc 420

tgcgtcgcca cctccggccc gggggccacc aacctcgtct ccgcgctcgc tgacgccctc 480

ctcgactcca tccccatggt cgccatcacg ggccaggtcc cccgccgcat gatcggcacg 540

gacgcgttcc aggagacgcc catagtggag gtcacgcgct ccatcaccaa gcacaactac 600

ctggtccttg acgtggagga tatcccccgc gtcatccagg aagccttctt cctcgcgtcc 660

tctggccgcc cggggccggt gctggttgat atccccaagg atatccagca gcagatggcc 720

gtgcctatct gggacacgcc gatgagtttg ccagggtaca tcgcccgcct gcccaagcca 780

ccatctactg aatcgcttga gcaggtcctg cgtctggttg gcgagtcacg gcgcccaatt 840

ctgtatgttg gtggtggctg cgctgcatcc ggcgaggagt tgcgccgctt tgttgagctc 900

actgggattc cggttacaac tactctgatg ggccttggca acttccccag cgacgaccca 960

ctgtctctgc gcatgcttgg gatgcatggc actgtgtatg caaattatgc agtcgataag 1020

gctgacctgt tgcttgcatt tggtgtgcgg tttgatgatc gcgtgactgg gaaaatcgag 1080

gcctttgcaa gcaggtccaa gattgtgcac attgacattg acccagctga gattggcaag 1140

aacaagcagc cacatgtctc catttgtgca gatgttaagc ttgctttaca ggggttgaat 1200

gctctattaa atgggagcaa agcacaacag ggtctggatt ttggtccatg gcacaaggag 1260

ttggatcagc agaagaggga gtttcctcta ggattcaaga cttttggcga ggccatcccg 1320

ccgcaatatg ctatccaggt actggatgag ctgacaaaag gggaggcgat cattgctact 1380

ggtgttgggc agcaccagat gtgggcggct cagtattaca cttacaagcg gccacggcag 1440

tggctgtctt cgtctggttt gggggcaatg ggatttgggt taccagctgc agctggcgct 1500

gctgtggcca acccaggtgt tacagttgtt gacattgatg gagatggtag tttcctcatg 1560

aacattcagg agttggcatt gatccgtatt gagaacctcc ctgtgaaggt gatgatattg 1620

aacaaccagc atctgggaat ggtggtgcaa tgggaggata ggttttacaa ggccaatcgg 1680

gcgcacacat accttggcaa cccagaaaat gagagtgaga tatatccaga ttttgtgacg 1740

attgctaaag gattcaacgt tccggcagtt cgtgtgacga agaagagcga agtcactgca 1800

gcaatcaaga agatgcttga gaccccaggg ccatacttgt tggatatcat cgtcccgcat 1860

caggagcacg tgctgcctat gatcccaaac ggtggtgctt tcaaggacat gatcatggag 1920

ggtgatggca ggacctcgta ctga 1944

<210> 24

<211> 647

<212> PRT

<213> 小麦 (Triticum aestivum)

<400> 24

Met Ala Ala Ala Thr Ser Pro Ala Val Ala Phe Ser Gly Ala Ala Ala

1 5 10 15

Ala Ala Ala Ala Met Pro Lys Pro Ala Arg Gln Pro Leu Pro Arg His

20 25 30

Gln Pro Ala Ser Arg Arg Ala Leu Pro Ala Arg Val Val Arg Cys Cys

35 40 45

Ala Ala Ser Pro Ala Ala Ala Thr Ser Ala Ala Pro Pro Ala Thr Ala

50 55 60

Leu Arg Pro Trp Gly Pro Ser Glu Pro Arg Lys Gly Ala Asp Ile Leu

65 70 75 80

Val Glu Ala Leu Glu Arg Cys Gly Ile Val Asp Val Phe Ala Tyr Pro

85 90 95

Gly Gly Ala Ser Met Glu Ile His Gln Ala Leu Thr Arg Ser Pro Val

100 105 110

Ile Thr Asn His Leu Phe Arg His Glu Gln Gly Glu Ala Phe Ala Ala

115 120 125

Ser Gly Tyr Ala Arg Ala Ser Gly Arg Val Gly Val Cys Val Ala Thr

130 135 140

Ser Gly Pro Gly Ala Thr Asn Leu Val Ser Ala Leu Ala Asp Ala Leu

145 150 155 160

Leu Asp Ser Ile Pro Met Val Ala Ile Thr Gly Gln Val Pro Arg Arg

165 170 175

Met Ile Gly Thr Asp Ala Phe Gln Glu Thr Pro Ile Val Glu Val Thr

180 185 190

Arg Ser Ile Thr Lys His Asn Tyr Leu Val Leu Asp Val Glu Asp Ile

195 200 205

Pro Arg Val Ile Gln Glu Ala Phe Phe Leu Ala Ser Ser Gly Arg Pro

210 215 220

Gly Pro Val Leu Val Asp Ile Pro Lys Asp Ile Gln Gln Gln Met Ala

225 230 235 240

Val Pro Ile Trp Asp Thr Pro Met Ser Leu Pro Gly Tyr Ile Ala Arg

245 250 255

Leu Pro Lys Pro Pro Ser Thr Glu Ser Leu Glu Gln Val Leu Arg Leu

260 265 270

Val Gly Glu Ser Arg Arg Pro Ile Leu Tyr Val Gly Gly Gly Cys Ala

275 280 285

Ala Ser Gly Glu Glu Leu Arg Arg Phe Val Glu Leu Thr Gly Ile Pro

290 295 300

Val Thr Thr Thr Leu Met Gly Leu Gly Asn Phe Pro Ser Asp Asp Pro

305 310 315 320

Leu Ser Leu Arg Met Leu Gly Met His Gly Thr Val Tyr Ala Asn Tyr

325 330 335

Ala Val Asp Lys Ala Asp Leu Leu Leu Ala Phe Gly Val Arg Phe Asp

340 345 350

Asp Arg Val Thr Gly Lys Ile Glu Ala Phe Ala Ser Arg Ser Lys Ile

355 360 365

Val His Ile Asp Ile Asp Pro Ala Glu Ile Gly Lys Asn Lys Gln Pro

370 375 380

His Val Ser Ile Cys Ala Asp Val Lys Leu Ala Leu Gln Gly Leu Asn

385 390 395 400

Ala Leu Leu Asn Gly Ser Lys Ala Gln Gln Gly Leu Asp Phe Gly Pro

405 410 415

Trp His Lys Glu Leu Asp Gln Gln Lys Arg Glu Phe Pro Leu Gly Phe

420 425 430

Lys Thr Phe Gly Glu Ala Ile Pro Pro Gln Tyr Ala Ile Gln Val Leu

435 440 445

Asp Glu Leu Thr Lys Gly Glu Ala Ile Ile Ala Thr Gly Val Gly Gln

450 455 460

His Gln Met Trp Ala Ala Gln Tyr Tyr Thr Tyr Lys Arg Pro Arg Gln

465 470 475 480

Trp Leu Ser Ser Ser Gly Leu Gly Ala Met Gly Phe Gly Leu Pro Ala

485 490 495

Ala Ala Gly Ala Ala Val Ala Asn Pro Gly Val Thr Val Val Asp Ile

500 505 510

Asp Gly Asp Gly Ser Phe Leu Met Asn Ile Gln Glu Leu Ala Leu Ile

515 520 525

Arg Ile Glu Asn Leu Pro Val Lys Val Met Ile Leu Asn Asn Gln His

530 535 540

Leu Gly Met Val Val Gln Trp Glu Asp Arg Phe Tyr Lys Ala Asn Arg

545 550 555 560

Ala His Thr Tyr Leu Gly Asn Pro Glu Asn Glu Ser Glu Ile Tyr Pro

565 570 575

Asp Phe Val Thr Ile Ala Lys Gly Phe Asn Val Pro Ala Val Arg Val

580 585 590

Thr Lys Lys Ser Glu Val Thr Ala Ala Ile Lys Lys Met Leu Glu Thr

595 600 605

Pro Gly Pro Tyr Leu Leu Asp Ile Ile Val Pro His Gln Glu His Val

610 615 620

Leu Pro Met Ile Pro Asn Gly Gly Ala Phe Lys Asp Met Ile Met Glu

625 630 635 640

Gly Asp Gly Arg Thr Ser Tyr

645

<210> 25

<211> 24

<212> DNA

<213> 人工合成（Artificial Synthesize）

<400> 25

taaagaagct agcccaatct gaac 24

<210> 26

<211> 25

<212> DNA

<213> 人工合成（Artificial Synthesize）

<400> 26

agaacataga ctacaaaact acagg 25

<210> 27

<211> 22

<212> DNA

<213> 人工合成（Artificial Synthesize）

<400> 27

cacaaccgcg agtggaaata cc 22

<210> 28

<211> 26

<212> DNA

<213> 人工合成（Artificial Synthesize）

<400> 28

gagaacagag actacaaaac tacacg 26

<210> 29

<211> 24

<212> DNA

<213> 人工合成（Artificial Synthesize）

<400> 29

gtttgagcat cagcctttct gcac 24

<210> 30

<211> 25

<212> DNA

<213> 人工合成（Artificial Synthesize）

<400> 30

tccatcggtc attatgtcat aaccg 25

Claims (15)

1.ALS突变蛋白质，其氨基酸序列如SEQ ID NO:4或SEQ ID NO:12或SEQ ID NO:20所示。

2.权利要求1所述的蛋白质，其能够赋予植物除草剂抗性，其中所述除草剂是咪唑啉酮类除草剂。

3.权利要求2所述的蛋白质，其中所述除草剂是咪草烟。

4.权利要求1～3任一所述的蛋白质，其来自小麦。

5.核酸，其编码权利要求1～4之任一所述的蛋白质。

6.权利要求5所述的核酸，其核苷酸序列如SEQ ID NO:3或SEQ ID NO:11或SEQ ID NO:19所示。

7.表达盒、载体或细胞，其包含权利要求5或6所述的核酸。

8.权利要求1～4之任一所述的蛋白质、权利要求5或6所述的核酸、权利要求7所述的表达盒、载体或细胞、包含权利要求5或6所述的核酸的植物、或者表达权利要求1～4之任一所述的蛋白质的植物在获得具有除草剂抗性的植物中的应用。

9.权利要求8所述的应用，其中所述除草剂是咪唑啉酮类除草剂。

10.权利要求8所述的应用，其中所述除草剂是咪草烟。

11.获得具有除草剂抗性的植物的方法，其包括如下步骤：

(1)使植物包含权利要求5或6所述的核酸；或，

(2)使植物表达权利要求1～4之任一所述的蛋白质。

12.权利要求11所述的方法，其包括转基因、突变、杂交、回交、自交或无性繁殖步骤。

13.权利要求8～10任一项所述的应用，其中所述植物是小麦。

14.权利要求11或12所述的方法，其中所述植物为小麦。

15.鉴定植物除草剂抗性的方法，其中植物是包含权利要求5或6所述的核酸的小麦、表达权利要求1～4之任一所述的蛋白质的小麦、或者权利要求11、12或14所述的方法获得的植物，其包括如下步骤：其包括如下步骤：

(1)测定所述植物是否包含权利要求5或6所述的核酸；或，

(2)测定所述植物是否表达权利要求1～4之任一所述的蛋白质。