CN102559646A - 小麦抗除草剂蛋白及其在植物育种中的应用 - Google Patents
小麦抗除草剂蛋白及其在植物育种中的应用 Download PDFInfo
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Abstract
本发明提供了使植物具有除草剂抗性的ALS蛋白质，其衍生自小麦品种宁春4号，其带有Ala179和/或Ser627突变。另外，本发明还提供了相应的核酸、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
技术领域
本发明属于植物蛋白质领域，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中的应用。
背景技术
农田中杂草是影响农作物产量的最重要因素之一，而人工除草所需要消耗的人力资源和成本很高，而且不便于农业集约化生产，严重制约了农作物种植向高产、优质和低成本方向的发展进程。因此，除草剂就应运而生，其使用为解决农田草害、促进栽培方式的革新及增产中起了很大的作用。
常见的除草剂，包括有磺酰脲类、咪唑啉酮类、嘧啶并三唑类、水杨酸嘧啶类等除草剂，目前已经大面积商业推广使用。然而，由于这些除草剂通常也能杀死农作物本身，因此对一般不具有除草剂抗性（耐受性）的农作物而言，除草剂的使用在时间和空间上受到了极大限制，如需要在农作物播种前一段时间使用除草剂。
为此，人们研究开发具有除草剂抗性（耐受性）的植物（农作物），这样可以在种植期间使用除草剂，杀死杂草，但不影响植物本身的生长，由此拓宽了除草剂的使用范围。然而，具有除草剂抗性的植物及其起决定性的物质（如蛋白、基因等）目前还没有理论来定向设计出，只能依赖于科研人员艰苦实践而获得，尤其是依赖长期筛选，并且凭借一些运气才能获得。
本发明人正是长期而艰苦的研究实践，偶然地在小麦宁春4号突变植株中发现了一系列（突变）蛋白，其能够在除草剂存在的条件下基本保持原有蛋白的正常植物生理功能，从而使得植物具有除草剂抗性（耐受性）。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，如小麦等。
发明内容
本发明要解决的技术问题在于提供新的使植物具有除草剂抗性的蛋白质。另外，本发明还提供了编码这些蛋白质的核酸以及基因工程中间体（如，表达盒、载体和细胞等），获得具有除草剂抗性的植物的方法和应用，并且提供了判断植物是否采用本发明方法获得的鉴定方法。
具体而言，在第一方面，本发明提供了使植物具有除草剂抗性的ALS蛋白质，其带有Ala179和/或Ser627突变，即带有Ala179突变、或者Ser627突变、或者Ala179和Ser627双突变。在本文中，ALS就是乙酰乳酸合成酶（acetolactate synthase）的简称，其酶分类编号为EC4,1.3.18。乙酰乳酸合成酶是支链氨基酸合成中的一个关键酶，如在缬氨酸和亮氨酸的合成中催化丙酮酸生成乙酰乳酸和二氧化碳，在异亮氨酸的合成中催化丙酮酸与丁酮酸生成2-乙醛基-2-羟基丁酸（James C. Global Status of Commercialized Biotech/GM Crops: 2006. ISAAA Briefs. Ithaca, NY）。
本发明第一方面的蛋白质是从本发明人突变的水稻植株中发现的。目前，已经发现ALS突变蛋白在不同植物（包括玉米、小麦、水稻、油菜、和向日葵等）的同源蛋白的第122、155、197、205、574、653、和/或654等位点会产生突变，从而使得相应植物对咪唑啉酮类、磺酰脲类、三唑嘧啶类和嘧啶水杨酸类等除草剂产生抗性（Tan S, Evzns R R, Dahmer M L, et al. Imidazolinone-tolerant crops: History， current status and Future. Pest Management Science. 2005, 61: 246-257；Tan S, Evzns R R, Dahmer M L, et al. Imidazolinone-tolerant crops: History, current status and Future [J]. Pest Management Science. 2005, 61: 246-257）。然而，没有报道表明，在来自水稻的ALS的第179和/或627位突变，会使得相应植物（尤其是水稻）具有除草剂抗性。
优选本发明第一方面的蛋白质带有Ala179Val和/或Ser627Asn突变，即带有Ala179Val突变、或者Ser627Asn突变、或者Ala179Val和Ser627Asn双突变，即从原有氨基酸序列的第179位的Ala突变为Val和/或第627位的Ser突变为Asn。除了本发明具体实施方式所述的突变植株筛选和扩增方法之外，在蛋白质序列已知的情况下，本领域技术人员有能力通过改变已知蛋白质的编码基因序列并将其导入表达载体，就可以制备出取代、添加或缺失了氨基酸残基的蛋白质，这些方法记载于《分子克隆实验指南》（北京：科学出版社，2002年）等文献中。在本发明的具体实施方式中，本发明第一方面的蛋白质的氨基酸序列如SEQ ID NO：2、4或6所示。这些蛋白质经证实是对除草剂不敏感的乙酰乳酸合成酶。
目前，以抑制ALS为靶位的除草剂包括磺酰脲类、咪唑啉酮类、嘧啶并三唑类、水杨酸嘧啶类等除草剂（Peter Babczinski, Thomas Zelinski. Mode of action herbicidal ALS-inhibitors on acetolactate synthase from green plant cell cultures，yeast，and Escherichia coli. 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。
在第二方面，本发明提供了核酸，其编码本发明第一方面的蛋白质。在本文中，核酸可以是DNA，也可以是RNA，优选是DNA。在知晓所编码蛋白序列或核酸序列的前提下，通过常规的密码子对应关系和宿主表达频率，运用PCR方法、DNA重组法或人工合成的方法，本领域技术人员有能力获得并优化编码本发明第一方面的蛋白质的核酸。一旦获得该核酸，就可以将其克隆入载体，再转化或转染入相应的细胞，然后通过常规的宿主细胞进行增殖，从中分离得到大量的核酸。优选本发明第二方面的核酸的核苷酸序列如SEQ ID NO：1、3或5所示。
在第三方面，本发明提供了表达盒（cassette），其包含本发明第二方面的核酸。本发明第三方面的表达盒优选在5’-3’的转录方向上包含启动子区、本发明第二方面的核酸和转录和翻译的终止区。本发明第三方面的表达盒优选适合在植物中表达本发明第二方面的核酸。示例性的启动子包括来自CaMV 35S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；示例性的转录和翻译的终止区包括转录终止元件或多聚腺苷酸化信号等。本发明第三方面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，本发明第三方面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。
在第四方面，本发明提供了载体，其包含本发明第二方面的核酸。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。本发明第四方面的载体是转化载体，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。
在第五方面，本发明提供了细胞，其包含本发明第二方面的核酸。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。
本发明的第二方面的核酸可被插入转化进细胞中的任何载体中。优选的细胞是细菌细胞，尤其是用于克隆或储存核酸、或是用于转化植物细胞的细菌细胞，例如农杆菌、大肠杆菌，包括根瘤土壤杆菌和毛根土壤杆菌等。另外优选的细胞是植物细胞，优选是小麦细胞，更优选是宁春4号细胞，所述植物细胞中包含的本发明第二方面的核酸可以是通过转基因技术导入植物细胞的，包括导入到植物细胞的核、叶绿体、线粒体和/或质体中，也可以是通过突变技术（如本发明具体实施方式所述的技术）使得本发明第二方面的核酸存在于植物细胞中的。
在第六方面，本发明提供了植物，其包含本发明第二方面的核酸，和/或其表达本发明第一方面的蛋白质。由于引入了本发明第二方面的核酸和本发明第一方面的蛋白质，本发明第六方面的植物具有除草剂抗性。在本文中，植物指的是借助光合作用，以水、二氧化碳和无机盐等无机物就能合成碳水化合物、蛋白质来维系生存的单个植株、植株群或其繁殖材料，包括植物、植物品种、植株、植物事件、植物后代、植物种子或其他植物的可繁殖部分。其中，植物后代本身就是植物，包括通过转基因技术产生的植物后代、与其他植物品种杂交产生的植物后代、以及回交或自交产生的植物后代。
本发明第六方面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，优选本发明第六方面的植物是小麦，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。本发明第六方面优选的植物，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。
在第七方面，本发明提供了本发明前述各方面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。本发明第七方面所提供的应用包括，本发明第一方面的蛋白质在获得具有除草剂抗性的植物过程中的应用，本发明第二方面的核酸在获得具有除草剂抗性的植物过程中的应用，本发明第三方面的表达盒在获得具有除草剂抗性的植物过程中的应用，本发明第四方面的载体在获得具有除草剂抗性的植物过程中的应用，本发明第五方面的细胞在获得具有除草剂抗性的植物过程中的应用，本发明第六方面的植物在获得具有除草剂抗性的植物过程中的应用。优选在本发明的第七方面中，除草剂是咪唑啉酮类除草剂，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，如将本发明第二方面的核酸转化入植物后使之表达本发明第一方面的蛋白质；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第六方面的植物并分选出仍旧包含本发明第二方面的核酸并表达本发明第一方面的蛋白质的植物。因此，本发明的第七方面提供的应用包括，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。
在第八方面，本发明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）使植物包含本发明第二方面的核酸；和/或，
（2）使植物表达本发明第一方面的蛋白质。
可以采用前述或者本领域技术人员能获知的转基因和非转基因方法，使植物包含本发明第二方面的核酸，或者使植物表达本发明第一方面的蛋白质。实施本发明第八方面的方法，能够获得本发明第六方面的植物。优选在本发明的第八方面中，可以采用的步骤包括转基因、杂交、回交、自交或无性繁殖步骤。这些步骤本身对于转基因或非转基因育种领域的技术人员来说，都是可以获知并实施的。
在第九方面，本发明提供了鉴定本发明第六方面的植物或者本发明第八方面的方法获得的植物的方法，其包括如下步骤：
（1）测定所述植物是否包含本发明第二方面的核酸；和/或，
（2）测定所述植物是否表达本发明第一方面的蛋白质。
通过该方法，可以判断植物是否属于本发明的植物，即是否属于本发明第六方面的植物，或者是否属于本发明第八方面的方法获得的植物。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Trp548突变或其相应核酸突变的方法都可以，优选能够检测Trp548Cys或Trp548Met突变或其相应核酸突变。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显示了抗咪唑啉酮类除草剂的小麦突变植株。
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），《Plant Propagation by Tissue Culture》(Edwin F. George,Michael A. Hall,Geert-Jan De Klerk, 2008, Springer)。
实施例1 从宁春4号突变植株中提取抗咪唑啉酮类除草剂的蛋白突变体
取市售的小麦品种——宁春4号种子400kg，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。将收割的种子在田间种植，待长至三叶期时，用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）以640g/公顷的用药量田间喷施。调查秧苗枯死情况，发现其中有绿色秧苗持续生长（图 1），保留并繁殖这些绿色秧苗，其为抗咪唑啉酮类除草剂的宁春4号突变植株3株。
分别取各宁春4号突变植株的叶片，连同野生型的宁春4号植株叶片，分别提取它们的基因组DNA，委托北京三博远志生物技术有限公司进行测序，发现：突变植株1相对于野生型宁春4号，在乙酰乳酸合成酶（ALS）基因的第536位点发生突变，由C变为T，导致相应编码蛋白的第179位由丙氨酸变为缬氨酸，即该突变植株的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；突变植株2相对于野生型宁春4号，在ALS基因的第1880位点发生突变，由G变为A，导致相应编码蛋白的第627位由丝氨酸变为天冬酰胺，即该突变植株的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；突变植株3相对于野生型宁春4号，在ALS基因的第536和1880位点发生突变，在第536位由C变为T，且在第1880位由G变为A，导致相应编码蛋白的第179位由丙氨酸变为缬氨酸，且第627位由丝氨酸变为天冬酰胺，即该突变植株的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 ID NO：6所示。
实施例2 蛋白突变体的抗咪唑啉酮类除草剂的离体活性测定
参照Fan ZJ等的方法（Specific activity determination of acetolactate synthase from maize（Zea mays L.）. 参见Han WN等编. The Proceedings of the 18 th Asia-Pacific Weed Science Society Conference . 515 - 522 . May 28 -June 2，2001 . Beijing：Standards Press of China.），提取上述宁春4号的野生型和各突变植株的ALS酶，并测定相应酶活性被咪唑啉酮类除草剂抑制的比率。过程简而言之，分别取各植株苗5g切碎后，加入10mL pH 7的50 mmol/L K2HPO4-KH2PO4缓冲液（其中含1mmol/L丙酮酸钠，0.5mmol/L MgCl2，0.5mmol/L TPP，10μmol/L FAD），用石英砂研磨粉碎后用8 层纱布过滤，滤液于4℃、20000rpm离心30分钟。取上清液，加入硫酸铵使之达到50%饱和度，然后0℃放置2小时后，于4℃、20000rpm离心30分钟。弃上清液，将沉淀溶解于5mL pH 7的50 mmol/L K2HPO4-KH2PO4缓冲液（其中含20mmol/L丙酮酸钠，0.5 mmol/L MgCl2），分别得各植株的ALS酶液。将0.1mL 0.5%咪草烟水剂（对照用水替代）、0.5mL 酶反应液（其中含24mmol/L丙酮酸钠，0.6mmol/L MgCl2，1mmol/L TPP，20μmol/L FAD，50 mmol/L K2HPO4-KH2PO4，pH 7）和0.4mL ALS酶液混合，于35℃避光静置反应1小时，然后加入0.1mL 3mol/L硫酸终止反应，于60℃放置15分钟脱羧。然后加入显色剂（0.5ml 0.5%（w/w）肌酸和0.5 ml 5%（w/w）α-萘酚的混合溶液（溶于2.5mol/L的NaOH 溶液））于60℃放置15分钟显色，测定525 nm 处的吸光度。将加入对照的野生型宁春4号的ALS酶活性分别记为100%，计算咪唑啉酮类除草剂对宁春4号的野生型和各突变ALS酶的活性的影响。
结果如下表所示，宁春4号的各突变ALS酶都基本保持了野生型ALS酶的活性，而且在存在咪唑啉酮类除草剂的情况下，野生型ALS酶的活性都有十分显著的下降，而各突变ALS酶却都保持了原有酶活性，表明突变ALS酶能够带来抗咪唑啉酮类除草剂的性能。
咪唑啉酮类除草剂对宁春4号的野生型和突变ALS酶活性的影响表
| 除草剂浓度 | 0 | 0.5% |
| 野生型ALS的相对酶活性 | 100% | 18% |
| 突变植株1的ALS的相对酶活性 | 93% | 90% |
| 突变植株2的ALS的相对酶活性 | 108% | 101% |
| 突变植株3的ALS的相对酶活性 | 86% | 88% |
实施例3转蛋白突变体基因的转基因植物的获得
委托北京未名凯拓农业生物技术有限公司用常规的农杆菌介导法将上述宁春4号的突变ALS酶的编码基因分别转入野生型拟南芥植株。过程简而言之，利用正向引物（5’ -AACTTCCAACCCTCTCTCTGCCTC-3’）和反向引物（5’ - TCAGTACGAGGTCCTGCCATC -3’），分别从上述宁春4号的各突变植株的基因组DNA中PCR扩增出突变ALS基因，测序正确后，将核苷酸序列如SEQ ID NO：1、3和5所示的ALS基因克隆入pCAMBIA1303质粒（购自Cambia公司）中，挑选阳性克隆转化农杆菌AGL0，培养菌体并转化拟南芥，PCR鉴定转化阳性的拟南芥收种后得到T1代的转基因种子，种植时用咪唑啉酮类除草剂咪草烟喷雾处理，发现生长状态良好，而未转基因的拟南芥苗全部枯死。
<110> 未名兴旺系统作物设计前沿实验室(北京)有限公司
<120> 小麦抗除草剂蛋白及其在植物育种中的应用
<160> 6
<170> PatentIn version 3.3
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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
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ggattcaacg ttccagcagt tcgagtgacg aagaagagcg aagtcactgc agcaatcaag 1800
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ggattcaacg ttccagcagt tcgagtgacg aagaagagcg aagtcactgc agcaatcaag 1800
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gtgctgccta tgatcccaaa cggtggtgct ttcaaggaca tgatcatgga gggtgatggc 1920
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Ala Ala Met Pro Lys Pro Ala Arg His Pro Leu Pro Arg His Gln Pro
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Ser Pro Ala Ala Thr Ser Ala Ala Pro Pro Ala Thr Ala Leu Arg Pro
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Ala Arg Ala Ser Gly Arg Val Gly Val Cys Val Ala Thr Ser Gly Pro
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Arg Met Leu Gly Met His Gly Thr Val Tyr Ala Asn Tyr Ala Val Asp
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Thr Gly Lys Ile Glu Ala Phe Ala Ser Arg Ser Lys Ile Val His Ile
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Asn Gly Ser Lys Ala Gln Gln Gly Leu Asp Phe Gly Pro Trp His Lys
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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> 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 tgatcccaaa 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 Asn Gly Gly Ala Phe Lys Asp Met Ile Met Glu Gly Asp Gly
625 630 635 640
Arg Thr Ser Tyr
Claims (10)
1.使植物具有除草剂抗性的ALS蛋白质，其带有Ala179和/或Ser627突变，优选带有Ala179Val和/或Ser627Asn突变。
2.权利要求1所述的蛋白质，其中除草剂是咪唑啉酮类除草剂，优选是咪草烟。
3.权利要求1或2所述的蛋白质，其氨基酸序列如SEQ ID NO：2、4或6所示。
4.核酸，其编码权利要求1~3之任一所述的蛋白质，优选其核苷酸序列如SEQ ID NO：1、3或5所示。
5.表达盒、载体或细胞，其包含权利要求4所述的核酸。
6.植物，其包含权利要求4所述的核酸，或者其表达权利要求1~3之任一所述的蛋白质，优选所述植物是小麦。
7.权利要求1~3之任一所述的蛋白质、权利要求4所述的核酸、权利要求5所述的表达盒、载体或细胞、或权利要求6所述的植物在获得具有除草剂抗性的植物中的应用，优选其中除草剂是咪唑啉酮类除草剂，优选是咪草烟。
8.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4所述的核酸；或，
（2）使植物表达权利要求1~3之任一所述的蛋白质。
9.权利要求8所述的方法，其包括转基因、杂交、回交、自交或无性繁殖步骤。
10.鉴定权利要求6所述的植物或者权利要求8或9所述的方法获得的植物的方法，其包括如下步骤：
（1）测定所述植物是否包含权利要求4所述的核酸；或，
（2）测定所述植物是否表达权利要求1~3之任一所述的蛋白质。
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| CN106467908A (zh) * | 2015-08-18 | 2017-03-01 | 未名兴旺系统作物设计前沿实验室（北京）有限公司 | 耐受除草剂的植物及其应用 |
| CN106636027A (zh) * | 2017-02-08 | 2017-05-10 | 上海市农业科学院 | 具有抗除草剂活性的小麦蛋白质、其编码基因及应用 |
| CN106754809A (zh) * | 2017-02-08 | 2017-05-31 | 上海市农业科学院 | 具有抗咪唑啉酮类除草剂活性的小麦蛋白质、其编码基因及应用 |
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| CN108004224A (zh) * | 2017-08-03 | 2018-05-08 | 江苏省农业科学院 | 使植物具有除草剂抗性的水稻als突变型蛋白及其应用 |
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| CN112813045A (zh) * | 2021-02-09 | 2021-05-18 | 江苏省农业科学院 | 棉花GhALS突变型蛋白、基因及其分子标记和应用 |
| CN113005105A (zh) * | 2019-12-21 | 2021-06-22 | 科稷达隆生物技术有限公司 | 一组抗除草剂蛋白及其在植物育种中的应用 |
| CN113717957A (zh) * | 2021-11-04 | 2021-11-30 | 科稷达隆(北京)生物技术有限公司 | 一种als突变蛋白及其应用 |
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| CN107354139A (zh) * | 2017-09-18 | 2017-11-17 | 江苏省农业科学院 | 小麦als突变型蛋白、核酸及其应用 |
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