CN110079532A - 调控植物种子大小的基因及其应用 - Google Patents
调控植物种子大小的基因及其应用 Download PDFInfo
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Abstract
本发明公开了一种调控植物种子大小的基因及其应用,属于植物分子生物学、生物化学、遗传学和植物育种领域。本发明通过EMS诱变水稻获得一个大粒突变体,并利用SIMM方法定位突变性状相关基因,获得一个调控植物种子大小的基因DLZ,该基因位于12号染色体上,基因座位号为LOC_Os12g41820(MSU登陆号)。基因DLZ的突变可导致水稻产生大粒性状表型,通过调节该基因的表达可调控植物的籽粒大小,用以获得大粒突变体植株以及改良作物的方法,对于作物的高产育种工作具有重要的理论和实践意义。
Description
技术领域
本发明涉及植物分子生物学、生物化学、遗传学和植物育种领域,特别涉及一个调控种子大小的基因,更具体地涉及一个调控种子大小的DLZ基因的核酸分子及其突变体及其在育种中的应用。
技术背景
植物种子是人类赖以生存的最重要食物来源,稻米、玉米和小麦是我国的三大主粮,粮食产量和价格影响着国计民生。人口持续增长与耕地面积不断减少的现状,对我国粮食生产提出了严峻的挑战。以水稻为例,水稻产量主要决定于三个要素——有效穗数、每穗实粒数和粒重,单位面积产量还受到水稻株型的影响,这些因素都是受多基因与环境互作控制的复杂性状,并且各个子性状之间还存在相关性,要改良产量性状还需协调好各构成要素之间的关系。根据现有的经验,高产品种可以分成四个类型,即大穗偏重型、大粒偏重型、多穗偏重型和综合兼顾型。单株有效穗数的遗传力最小,很容易受环境影响,对单株有效穗数的改良不如通过合理密植来调整单位面积的有效穗数。每穗粒数的遗传力适中,改良起来仍有不小的难度。相比之下,粒重是产量构成要素中遗传力最高的,对其进行改良相对容易(张启发,绿色超级稻的构想与实践,科学出版社,2009)。
水稻粒重由种子大小即颖壳的体积和胚乳发育状况两个因素决定。颖壳通常在水稻开花之前就已经定型,受精之后灌浆形成的米粒大小和形状受制于颖壳的容积,因此,颖壳的体积是粒重大小的先决条件。对颖壳形状与大小的描述一般称为粒形,通常用粒长、粒宽、粒厚和长宽比表示。值得一提的是,粒形不仅是重要的产量性状之一,也是主要的外观品质性状,粒形与稻米的其它品质性状如垩白率、糙米率、精米率之间也存在一定的相关性。因此,在育种中通过对种子大小与形状的选择,不仅可以提高产量潜力,还可以间接调控稻米品质。
植物种子大小是多基因控制的数量性状,在前期大量的QTL扫描和定位的研究基础上,近十年来相关基因的克隆呈现一种爆发的趋势。对水稻等重要作物种子大小调控基因的挖掘和功能研究已经成为功能基因组学研究的一大热点。这些基因的发现和功能分析促进了植物种子大小基因调控网络研究的逐步深入,研究人员开始探索关键基因聚合的最佳设计,通过评价优异基因型聚合的增产效应,为创制设计型产量突破性的新品种提供理论指导和材料支撑。
SIMM(Simultaneous Identification of Multiple Mutations)(Yan et al.,Simultaneous identification of multiple causal mutations in rice.Frontiers inPlant Science,2016)是一种基于二代测序技术的快速高效的定位突变基因的方法。与其他方法相比,SIMM可在不需要野生型基因组数据的前体下,以其他同样或相近来源的突变体为背景,同时鉴定多个突变性状相关的突变位点,具有更高的灵敏度和特异性,有助于快速定位候选功能基因,辅助水稻功能基因研究及水稻设计育种。此外,SIMM还可用于极端表型池QTL(数量性状基因座,Quantitative Trait Locus)定位,有效缩小候选区间,辅助定位主效QTL基因。此方法也可有效应用于定位其他物种EMS突变体候选功能基因。
高分辨率熔解曲线分析(High-Resolution Melting Curve Analysis,HRM)是一种基于单核苷酸熔解温度不同而形成不同形态熔解曲线的基因分析新技术,具有极高的敏感性,可以检测出单个碱基的差异,并且成本低、通量高、速度快、结果准确、不受检测位点的局限。该方法无需使用序列特异性探针,而是利用一种饱和染料对PCR反应产物进行分析。其原理是:双链DNA的热稳定性受其长度和碱基组成的影响,序列变化会导致升温过程中双链DNA解链行为的改变。因为所用的荧光染料只能嵌入并结合到双链DNA上,因此利用实时PCR技术,通过实时检测双链DNA熔解过程中荧光信号值的变化,就能以生成不同形状熔解曲线的方式将PCR产物中存在的差异直观地展示出来。同时,借助于专业性的分析软件就可以对测试群体实现基于不同形状熔解曲线的基因分型或归类。
本发明通过EMS诱变水稻籼稻品种“黄华占”,筛选得到一个由单个隐性核基因控制的大粒突变体,进而对该突变体进行表型鉴定、遗传分析和遗传背景鉴定,并利用SIMM方法、HRM技术和基因信息分析,成功定位并克隆了一个种子大小调控基因DLZ,该基因位于12号染色体上,基因座位号为LOC_Os12g41820(MSU登陆号),该基因的突变可导致水稻产生大粒性状表型,从而可应用于控制作物籽粒的大小。本发明有助于提高作物产量和改善品质,为培育具有大粒重的水稻新品种提供基因资源和技术支持,对作物农艺性状的提高和高产分子育种工作具有重要的意义和应用价值。
发明内容
本文提到的所有参考文献都通过引用并入本文。
除非有相反指明,本文所用的所有技术和科学术语都具有与本发明所属领域普通技术人员通常所理解的相同的含义。除非有相反指明,本文所使用的或提到的技术是本领域普通技术人员公知的标准技术。材料、方法和例子仅作阐述用,而非加以限制。
本发明提供了一个调控种子大小的基因DLZ(大粒占,Da Li Zhan),该基因位于12号染色体上,其在水稻中的基因位点编号是LOC_Os12g41820(MSU登陆号,参考Rice GenomeAnnotation Project,http://rice.plantbiology.msu.edu/),该基因突变后可以使含有该突变的植株的种子粒型变大。
由于不同品种间的同一基因往往存在单核苷酸多态性,即同一基因的核苷酸序列往往存在个别碱基的差异,但是水稻品种数量很多,发明人不可能进行一一列举,因此本发明仅提供了在籼稻和粳稻中具有代表性的两个品种的序列。具体地,所述水稻DLZ基因的核苷酸序列选自下列组的序列之一:
(a)如调控植物种子大小的基因LOC_Os12g41820所示的核苷酸序列;
(b)如SEQ ID NO:1、2、20或21所示的核苷酸序列;
(c)其编码氨基酸序列如SEQ ID NO:3、55、56、57、58、59、60或61所示的核苷酸序列;
(d)在严谨条件下能够与(a)-(c)中所述序列的DNA杂交的DNA序列;或(e)与(a)-(d)所述序列有至少80%(优选为至少85%)序列相似性,且具有调控植物种子大小功能的DNA序列;或
(f)与(a)-(e)之任一所述序列互补的DNA序列。
上述调控种子大小的DLZ基因可从各种植物中分离获得。本领域技术人员应该知晓,本发明所述的种子大小调控基因还包括与DLZ基因的核苷酸序列或蛋白序列高度同源,并且具有同样的种子大小调控功能的同源基因。所述同源基因包括在严谨条件下能够与本发明所公开的DLZ基因的核苷酸序列杂交的DNA序列。本文中使用的“严谨条件”是公知的,包括诸如在含400mM NaCl、40mM PIPES(pH6.4)和1mM EDTA的杂交液中杂交,所述杂交的温度优选是53℃-60℃,杂交时间优选为12-16小时,然后用含0.5×SSC和0.1%SDS的洗涤液洗涤,洗涤温度优选为62℃-68℃,洗涤时间为15-60分钟。
同源基因还包括与本发明所公开的DLZ基因所示的序列有至少80%、85%、90%、95%、98%、或99%序列相似性,且具有调控种子大小功能的DNA序列,可以从任何植物中分离获得。其中,序列相似性的百分比可以通过公知的生物信息学算法来获得,包括Myers和Miller算法、Needleman-Wunsch全局比对法、Smith-Waterman局部比对法、Pearson和Lipman相似性搜索法、Karlin和Altschul的算法,这对于本领域技术人员来说是公知的。
本发明所述的基因序列可以从任何植物中分离获得,包括但不限于芸苔属、玉米、小麦、高粱、两节荠属、白芥、蓖麻子、芝麻、棉籽、亚麻子、大豆、拟南芥属、菜豆属、花生、苜蓿、燕麦、油菜籽、大麦、燕麦、黑麦(Rye)、粟、蜀黍、小黑麦、单粒小麦、斯佩尔特小麦(Spelt)、双粒小麦、亚麻、格兰马草(Gramma grass)、摩擦禾、假蜀黍、羊茅、多年生麦草、甘蔗、红莓苔子、番木瓜、香蕉、红花、油棕、香瓜、苹果、黄瓜、石斛、剑兰、菊花、百合科、棉花、桉、向日葵、芸苔、甜菜、咖啡、观赏植物和松类等。优选地,植物包括玉米、大豆、红花、芥菜、小麦、大麦、黑麦、稻、棉花和高粱。
本发明提供了一种调控植物种子大小的方法,所述方法通过影响将本发明所提供的DLZ基因表达水平,从而影响植物种子大小。所述影响植物种子大小是指通过降低DLZ基因的表达水平,从而使所述植株的种子大小发生改变,如导致大粒的表型。具体地,取决于具体应用需求,可以通过多种方法来影响DLZ基因在植物体内的表达水平,从而达到调控种子大小的效果。更具体地,调控DLZ基因的表达水平可以使用许多本领域普通技术人员可获得的工具进行,例如通过突变、诱变、反义基因的转入、共抑制或发夹结构的引入等,都可以用于破坏DLZ基因的正常表达,从而获得种子变大的植株。
本发明还提供一种获得DLZ基因的大粒突变体材料的方法,所述方法通过突变水稻内源的DLZ基因,或突变与其高度同源的基因的核苷酸序列,使该植物调控种子大小的作用途径被改变。所述DLZ基因的核苷酸序列如SEQ ID NO:1、2、20或21所示,所述DLZ基因的氨基酸序列如SEQ ID NO:3、55、56、57、58、59、60或61所示。所述“突变”包括但不限于以下方法,如用物理或化学的方法所导致的基因突变,化学方法包括用EMS等诱变剂处理所导致的诱变,所述突变还可以是点突变,也可以是DNA缺失或***突变,也可以是通过RNAi等基因沉默手段或者通过基因定点突变的方法,所述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等基因编辑方法。
本发明还提供了一种DLZ大粒突变体材料的应用方法,其特征在于所述突变材料是由DLZ基因的核苷酸序列的突变所造成,含有突变型DLZ基因的植株具有大粒种子的表型,其中所述DLZ基因的核苷酸序列优选如SEQ ID NO:1、2、20或21所示。具体地,本发明所述突变后的核苷酸序列如SEQ ID NO:4所示,氨基酸序列如SEQ ID NO:5所示,在大粒突变体中,DLZ基因的第四个外显子上发生两个碱基的突变,具体由AAA突变为TTA,导致SEQ IDNO:3蛋白的第321位氨基酸由Lys突变为Leu,导致得到的转录本与蛋白产物均发生变化,从而使植物具有大粒种子表型。本领域技术人员应该知晓,可以将所述核苷酸序列SEQ IDNO:4构建到植物表达载体,进行植物转化,从而获得新的转基因的大粒突变体材料。所述突变体材料的应用,包括但不限于在杂交育种中的应用,更具体的是指包括但不限于培育植物品种或品系、培育种子尺寸增大的植物品种或品系和培育种子尺寸变小的植物品种或品系、鉴定作物大粒品种和小粒品种的分子标记等应用。
本发明还提供了一种表达盒在调控植物种子大小中的应用,所述表达盒含有调控植物种子大小的DLZ基因的DNA序列,所述调控植物种子大小的基因的核苷酸序列选自下列组的序列之一:
(a)如调控植物种子大小的基因LOC_Os12g41820所示的核苷酸序列;
(b)如SEQ ID NO:1、2、20或21所示的核苷酸序列;
(c)其编码氨基酸序列如SEQ ID NO:3、55、56、57、58、59、60或61所示的核苷酸序列;
(d)在严谨条件下能够与(a)-(c)中所述序列的DNA杂交的DNA序列;或(e)与(a)-(d)所述序列有至少80%(优选为至少85%)序列相似性,且具有调控植物种子大小功能的DNA序列;或
(f)与(a)-(e)之任一所述序列互补的DNA序列。
具体地,上述表达盒中的种子大小调控基因还可操作性地连有一个可驱动其表达的启动子,所述构建体中的启动子可以是天然启动子或被取代的启动子,其将驱动所连核苷酸序列在植株中的表达。表达盒中的启动子包括但不限于组成型表达启动子、诱导型启动子、组织特异表达启动子、时空特异表达启动子等。本发明所述的组成型启动子的基因表达不具有组织和时间特异性,外界因素对组成型启动子启动的外源基因表达几乎没有影响。所述组成型启动子包括但不限于CaMV35S、FMV35S、水稻肌动蛋白(Actin1)启动子、玉米泛素(Ubiquitin)启动子等。本发明所述的组织特异性启动子除包含应有的一般启动子元件外,还具有增强子以及沉默子的特性,该类启动子的优点在于可启动基因在植物特定组织部位的表达,避免外源基因的不必要表达,从而节约植物体的整体能量消耗。本发明所述的诱导型启动子是指在某些特定的物理或化学信号的刺激下,可以大幅度地提高基因的转录水平的启动子,目前已经分离的诱导型启动子包括但不限于逆境诱导表达启动子、光诱导表达启动子、热诱导表达启动子、创伤诱导表达启动子、真菌诱导表达启动子和共生细菌诱导表达启动子等。本发明所述的组织特异性启动子包括但不限于LTP2种子特异表达启动子、END2种子特异表达启动子、糊粉层特异表达启动子等。
上述表达盒中还可包括其它组分,这主要取决于载体构建的目的和用途,例如可进一步包括选择标记基因、靶向或调控序列、稳定序列或引导序列、内含子等。上述构建体中还可包括其它组分,这主要取决于载体构建的目的和用途,例如可进一步包括选择标记基因、靶向或调控序列、稳定序列或引导序列、内含子等。表达盒还将在目标异源核苷酸序列的3’端包括在植物中具有功能的转录和翻译终止子。终止子可以是本发明所提供DLZ基因的自身终止子,也可以是来自外源的终止子,如胭脂氨酸合酶或章鱼碱合酶终止区域等。
在希望将异源核苷酸序列的表达产物引向特定细胞器,例如质体、造粉体,或者引向内质网,或在细胞表面或细胞外分泌的情况下,表达盒还可包含用于编码转运肽的核苷酸序列。此类转运肽是本领域所公知的,其包括但不限于Rubisco的小亚基、植物EPSP合酶、玉米Brittle-1、叶绿体转运肽等。
在制备表达盒的过程中,可对多种DNA片段加以操作,以提供处于合适方向,或是处于正确读码框中的DNA序列。为达到此目的,可使用衔接子或接头,将DNA片段连起来,或者进一步包括其它操作,以提供方便的限制性酶切位点等。
本发明上述表达盒,还进一步的可以包含一个筛选基因,所述筛选基因可以用于将含有该表达盒的植株、植物组织细胞或载体筛选出来。所述筛选基因包括但不限于抗生素抗性基因、或是抗除草剂基因、或是荧光蛋白基因等。具体地,所述筛选基因包括但不限于:氯霉素抗性基因、潮霉素抗性基因、链霉素抗性基因、奇霉素抗性基因、磺胺类抗性基因、草甘磷抗性基因、草丁膦抗性基因、bar基因、红色荧光基因DsRED、mCherry基因、青色荧光蛋白基因、黄色荧光蛋白基因、荧光素酶基因、绿色荧光蛋白基因等。
进一步地,本发明所提供的构建体中还可包括选择标记基因,用于选择经转化的细胞或组织。所述选择标记基因包括赋予抗生素抗性或对除草剂抗性的基因。合适的选择标记基因包括但不限于:氯霉素抗性基因,潮霉素抗性基因,链霉素抗性基因,奇霉素抗性基因,磺胺类抗性基因,草甘磷抗性基因,草丁膦抗性基因。所述选择标记基因还可以是红色荧光基因、青色荧光蛋白基因、黄色荧光蛋白基因、荧光素酶基因、绿色荧光蛋白基因、花青甙p1等基因。
本发明所提供的表达盒或载体可被***质粒、粘粒、酵母人工染色体、细菌人工染色体或其他适合转化进宿主细胞中的任何载体中。优选的宿主细胞是细菌细胞,尤其是用于克隆或储存多核苷酸、或用于转化植物细胞的细菌细胞,例如大肠杆菌、根瘤土壤杆菌和毛根土壤杆菌。当宿主细胞是植物细胞时,表达盒或载体可被***被转化的植物细胞的基因组中。***可以是定位的或随机的***。优选地,***通过诸如同源重组来实现。另外,表达盒或载体可保持在染色体外。本发明的表达盒或载体可存在于植物细胞的核、叶绿体、线粒体和/或质体中。优选地,本发明的表达盒或载体被***植物细胞核的染色体DNA中。
本发明所提供的DLZ基因的核苷酸序列和启动子序列或表达盒可被***载体、质粒、酵母人工染色体、细菌人工染色体或其他适合转化进宿主细胞中的任何载体中。优选的宿主细胞是细菌细胞,尤其是用于克隆或储存多核苷酸、或用于转化植物细胞的细菌细胞,例如大肠杆菌、根瘤土壤杆菌和毛根土壤杆菌。当宿主细胞是植物细胞时,表达盒或载体可***至被转化的植物细胞的基因组中。***可以是定位的或随机的***。
本发明所述的将核苷酸序列、载体或表达盒转入植株或引入植株或对植株进行转化,均指通过常规的转基因方法,将核苷酸序列、载体或表达盒转入到受体细胞或受体植株中。植物生物技术领域技术人员已知的任何转基因方法均可被用于将重组表达载体转化进植物细胞中,以产生本发明的转基因植物。转化方法可包括直接和间接的转化方法。合适的直接方法包括聚乙二醇诱导的DNA摄入、脂质体介导的转化、使用基因枪导入、电穿孔、以及显微注射。所述转化方法也包括农杆菌介导的植物转化方法等。
本发明提供了一种控制籽粒大小的杂交植物的生产方法,其特征在于,该方法包括:
(a)构建本发明所提供的表达盒;
(b)将步骤(a)获得的表达盒导入植物细胞;
(c)再生出转基因植物;和
(d)选择出转基因植物;并且
(e)任选地,增殖步骤(d)获得的植物以获得后代。
本发明的转基因植物使用植物生物技术领域技术人员已知的转化方法制备。任何方法可被用于将重组表达载体转化进植物细胞中,以产生本发明的转基因植物。转化方法可包括直接和间接的转化方法。合适的直接方法包括聚乙二醇诱导的DNA摄入、脂质体介导的转化、使用基因枪导入、电穿孔、以及显微注射等。在本发明的具体实施方式中,本发明使用了基于土壤杆菌的转化技术(可参见Horsch RB等(1985)Science 225:1229;White FF,Vectors for Gene Transfer in Higher Plants,Transgenic Plants,第1卷,Engineering and Utilization,Academic Press,1993,pp.15-38;Jenes B等.Techniquesfor Gene Transfer,Transgenic Plants,第1卷,Engineering and Utilization,Academic Press,1993,pp.128-143,等)。土壤杆菌菌株(例如根瘤土壤杆菌或毛根土壤杆菌)包含质粒(Ti或Ri质粒)和T-DNA元件,所述质粒和元件在用土壤杆菌转染后被转移至植物,而T-DNA被整合进植物细胞的基因组中。T-DNA可位于Ri-质粒或Ti-质粒上,或独立地包含在所谓的双元载体中。土壤杆菌介导的转化方法描述于例如中。土壤杆菌介导的转化最适合双子叶植物,但是也适合单子叶植物。土壤杆菌对植物的转化描述于例如中。转化可导致瞬时或稳定的转化和表达。尽管本发明的核苷酸序列可被***落入这些广泛种类中的任何植物和植物细胞中,但是其尤其适用于作物植物细胞。
与现有技术相比,本发明具有如下有益效果:
(1)本发明提供了一个水稻种子大小调控基因,该基因的突变可使作物(如水稻)的籽粒变大,从而增加作物产量,为水稻的高产育种提供了新的基因资源。
(2)本发明提供的DLZ基因可以作为控制作物籽粒大小,提高产量和品质的一个基因,应用于作物品种的改良,有助于选育出优质性状的水稻新品种。同时,DLZ基因还可用于分子标记技术,为水稻大粒高产育种等实际生产应用服务。
(3)本发明提供的DLZ基因在玉米、高粱等众多植物中具有同源基因,DLZ基因不仅可用于水稻,也可用于其它植物的新品种培育。
附图说明
图1是野生型黄华占(HHZ)和大粒突变体(dlz)的植株形态,bar=20cm。
图2是黄华占(HHZ)和大粒突变体(dlz)的种子(A)和米粒(B)形态,bar=1cm。
图3是大粒突变体(dlz)与黄华占(HHZ)杂交F2代分离群体的千粒重。
图4是SIMM方法定位大粒突变体(dlz)的突变位点,其中红色三角标示突变位点所在位置,黑色字母表示野生型的碱基,红色字母表示突变体的碱基。
图5是大粒突变体(dlz)与黄华占(HHZ)杂交F2代分离群体中三个候选基因间的重组单株与部分非重组单株的粒宽与千粒重,其中W表示野生型,H表示杂合突变型,M表示纯合突变型。
图6是DLZ基因在野生型黄华占不同组织器官中的表达量。DAP1表示受精后1天。
图7是利用CRISPR技术定点敲除粳稻中花11(ZH11)和籼稻黄华占(HHZ)的DLZ基因产生的序列变异(A)和相应的转基因植株种子大小(B),其中红色字母表示***的碱基,红色“-”表示缺失的碱基;Z系列编号表示粳稻中花11背景的转基因植株,H系列编号表示黄华占背景的转基因植株。
图8是野生型黄华占(W)、杂合突变体(H)与dlz纯合突变体(M)的各种重要农艺性状表型及单株产量比较,其中a、b、c不同字母表示数据经t测验具有显著差异,相同字母表示差异不显著。
图9是利用RNAi技术将粳稻中花11的DLZ进行基因沉默产生的转基因植株的DLZ基因表达水平及其粒宽表型,其中灰色柱子表示转基因阴性植株。
图10是利用CRISPR技术编辑粳稻中花11的DLZ基因启动子产生的转基因植株的DLZ基因表达水平、启动子片段缺失情况以及粒宽表型,其中灰色柱子表示表达量变化与粒宽表型变化不明显的单株。
图11是不同植物的同源DLZ蛋白的同源性比较,其中Brachypodium di表示二穗短柄草,Hordeum vulgare表示大麦,Oryza sativa表示水稻,Oryza brachyant表示野生稻,Panicum hallii表示黍,Setaria italica表示谷子,Zea mays表示玉米,Sorghum bicolor表示高粱。
具体实施方式
下面对本发明的实施例作详细说明,本实施例在以本发明技术方案为前提下进行实施,给出了详细的实施方式和具体的操作过程,但本发明的保护范围不限于下述的实施例。
实施例1、水稻大粒突变体(dlz)的筛选
采用含0.7%质量浓度的EMS水溶液浸泡籼稻黄华占种子(M0),诱变处理12小时,将M0代种子植株结实后混收,获得突变体库(M1)。来自M1代种子的植株在种子成熟期用于筛选,通过表型观察,获得植株发育正常、种子明显变大的植株(图1,图2A)。突变体种子变大主要是由于粒宽的增加,并且颖壳体积增大后籽粒充实度不受影响,因此米粒的宽度也显著增大(图2B),故千粒重增加。
实施例2、水稻大粒突变体(dlz)的遗传分析
将dlz突变体与野生型黄华占杂交,获得F1代杂交种,再取F1植株的种子种植F2分离群体,种子完熟后收种、完全干燥,并进行千粒重的测量。野生型黄华占的平均千粒重为21克左右,而F2群体的考种结果表明,大粒的单株在群体中占少数(图3)。由于千粒重是数量性状,其大小在群体中呈连续分布,以23克为分界点,千粒重大于23克的定为大粒性状,小于或等于23克的定为正常性状,则F2-1和F2-2两个群体的正常性状与大粒性状的分离比符合3:1(χ2=1.44<χ2 (0.05,df=1)=3.84),表明突变体的大粒性状是由一个隐性核基因突变产生的。
实施例3、水稻大粒突变体DLZ基因的克隆
突变体的基因克隆采取SIMM方法,即利用突变体与原野生亲本杂交构建F2代群体,通过重测序进行基因定位的方法。具体地,将dlz突变体与野生型黄华占杂交,从F2群体中选取30个极端表型(千粒重25克以上)的突变体植株,分别提取叶片的基因组DNA并等量混合,按照Illumina Hiseq2000测序平台测序建库标准流程进行建库,并通过PE101重测序。测序数据应用SIMM方法分析定位突变位点,最终定位到水稻第12号染色体末端的一个450kb区间,其中包括四个单碱基突变,分别位于LOC_Os12g41220、LOC_Os12g41820和LOC_Os12g41910基因的编码区,且LOC_Os12g41820基因有两个相邻碱基均发生了突变(图4)。
另取三个F1单株的种子种植F2群体,每个群体种植约600至800个单株,共计种植2100株。利用HRM法鉴定上述每个单株的三个突变位点的基因型,并从中筛选出80个重组单株,考查了重组单株与非重组单株的粒宽和千粒重。结果发现,在非重组单株中,杂合型的千粒重介于野生型和突变型之间,表明dlz基因具有半显性效应:当LOC_Os12g41820和LOC_Os12g41910两个位点都是杂合型时,LOC_Os12g41220位点的野生型与突变型之间的千粒重并无差异;当LOC_Os12g41220和LOC_Os12g41820两个位点都是杂合型时,LOC_Os12g41910位点的野生型与突变型之间的千粒重也无差异;只有LOC_Os12g41820位点的野生型与杂合型、突变型之间的千粒重差异显著,并且该差异趋势与非重组单株相同(图5)。实验结果表明LOC_Os12g41820即为DLZ基因。
在野生型黄华占(籼稻)中,DLZ基因的基因组DNA全长6146bp,序列如SEQ ID NO:1所示。该基因有9个外显子,分别在SEQ ID NO:1的第831至1151位、第1502至1541位、第2298至2427位、第2569至3056位、第3148至3228位、第3364至3462位、第3643至3756位、第3840至3911位、第4498至4566位核苷酸。该基因编码区(CDS)全长1635bp,序列如SEQ ID NO:2所示,其CDS编码1个含有544个氨基酸的蛋白质,其氨基酸序列如SEQ ID NO:3所示。在本发明所提供的水稻大粒突变体(dlz)中,DLZ基因的第4个外显子上发生两个碱基的突变,具体为SEQ ID NO:2的第961、962、963位碱基从AAA突变为TTA(见SEQ ID NO:4),从而导致SEQ IDNO:3的第321位氨基酸由赖氨酸(Lys)突变为亮氨酸(Leu)(见SEQ ID NO:5)。将粳稻品种日本晴(典型粳稻基因组供体)与籼稻品种黄华占相比发现,DLZ基因的核苷酸序列(见SEQ IDNO:20)差异主要存在于非编码区,编码区仅在第6个外显子上存在一个SNP位点,具体为在SEQ ID NO:2的第1170位碱基位置上,黄华占是G,日本晴是A,该SNP位点位于SEQ ID NO:3的第390位Lys的密码子第三位(见SEQ ID NO:21),但并不改变所编码的氨基酸(见SEQ IDNO:3),说明DLZ基因在粳稻和籼稻之间差异较小。
实施例4、DLZ基因在水稻各组织器官中的表达模式分析
根据DLZ基因的cDNA序列设计引物,上游引物为820qF:5’-AGTCCAGGCGTATACAGTGC-3’(SEQ ID NO:6),下游引物为820qR:5’-TCAGAGCAATCCTGACACCA-3’(SEQ ID NO:7)。同时以水稻Ubiquitin基因为内参设计引物,上游引物为UBqF:5’-CAACCAGCTGAGGCCCAAGAA-3’(SEQ ID NO:8),下游引物为UBqR:5’-CCAGGGAGATAACAACGGAAGC-3’(SEQ ID NO:9)。分别提取野生型黄华占的根、茎、叶、分蘖芽、外稃、内稃、雌蕊、花药、不同长度的幼穗、受精后1至7天的种子等组织的总RNA并合成cDNA模板,采取实时荧光定量PCR的方法分析DLZ基因的表达水平。结果如图6所示,该基因在所检测的各个组织器官中都有较高表达,表达量差异不大。
实施例5、定点敲除籼稻和粳稻的DLZ基因
在DLZ基因的第1个外显子上选择第1个CRISPR定点诱变的靶位点序列Target 1:5’-CCTTCCTGGTCGACCGGCATTGG-3’(SEQ ID NO:10),在第3个外显子上选择第2个CRISPR定点诱变的靶位点序列Target 2:5’-GCTGTTCGTGTTGGATCGCTTGG-3’(SEQ ID NO:11)。合成带有粘性末端的接头引物U3-Target1-linkerF:5’-ggcACCTTCCTGGTCGACCGGCAT-3’(SEQ IDNO:12),U3-Target1-linkerR:5’-aaacATGCCGGTCGACCAGGAAGG-3’(SEQ ID NO:13),U6a-Target2-linkerF:5’-gccGCTGTTCGTGTTGGATCGCT-3’(SEQ ID NO:14),U6a-Target2-linkerR:5’-aaacAGCGATCCAACACGAACAG-3’(SEQ ID NO:15)。将两对接头引物分别用ddH2O溶解成10μM工作液,F引物和R引物各取10μL加入到80μL ddH2O中混合稀释到1μM,于90℃处理30s,移至室温冷却完成退火。再用T4DNA ligase分别连入经BsaI酶切线性化的pYLsgRNA-U3或pYLsgRNA-U6a质粒,得到微量的U3::Target1-gRNA和U6a::Target2-gRNA表达盒。将上述表达盒作为模板,经过两轮PCR扩增并加上BsaI酶切位点,连入经BsaI酶切线性化的pYLCRISPR/Cas9-MH(B)终载体,最终得到Pubi::Cas9-U3::Target1-gRNA-U6a::Target2-gRNA表达盒。PCR模板与引物组合如表1所示,通用引物序列如表2所示,阳性克隆的鉴定分别用SP-L和U3-Target1-linkerR、U3-Target1-linkerF和U6a-Target2-linkerR、U6a-Target2-linkerF和SP-R等三对引物进行菌液PCR检测,SP-L引物用于U3-Target1的测序,SP-R用于U6a-Target2的测序。
将Pubi::Cas9-U3::Target1-gRNA-U6a::Target2-gRNA表达盒通过农杆菌介导的水稻遗传转化方法分别转入粳稻品种中花11和黄华占种子诱导产生的愈伤组织中,经过潮霉素筛选和抗性愈伤组织的分化再生得到转基因植株。利用CTAB法抽提上述植株叶片的基因组DNA,并用引物对U3-Target1-linkerF和U6a-Target2-linkerR做转基因植株T-DNA***阳性检测分析,阳性植株可扩增出600bp左右的条带。将阳性植株的基因组DNA再用引物对820-Target1-F:5’-CTGACATGGGCGCACATG-3’(SEQ ID NO:16)和820-Target2-R:5’-CCTCGTATCCTTGCAGCAACTT-3’(SEQ ID NO:17)扩增出包含两个靶位点的约1.84kb的DNA区段,再利用引物对820-T1-SEQ:5’-GGTGATGCACACGAAGAAGC-3’(SEQ ID NO:18)和820-T2-SEQ:5’-TCCCTAGTTGCATCCGTTTG-3’(SEQ ID NO:19)分别对Target1和Target2进行测序。测序结果如图7A所示,在两个靶位点上都发生了不同的碱基***或缺失,进而改变了DLZ基因的读码框,并且两条染色体都发生了突变,因此表明产生的阳性植株是DLZ基因功能丧失的突变体,其粒宽也比野生型中花11或黄华占大大增加(图7B),证明定点敲除粳稻和籼稻的DLZ基因均导致种子大小的显著增加。
实施例6、dlz突变基因在杂合状态下具有显著增产效应
如图8所示,大粒突变体(dlz)的粒宽变大和千粒重增加,株高与野生型黄华占相比有所下降,但茎秆更粗壮,穗数减少,穗长变短,每穗颖花数也显著减少,综合以上特点,纯合突变体产量下降。大粒突变基因在杂合状态下株高、穗数、穗长和每穗颖花数都与野生型黄华占相似,没有显著差异,而杂合株的粒宽与千粒重均高于野生型黄华占而低于纯合突变体,茎秆也比野生型材料粗壮,单株产量比野生型黄华占显著增加,表明大粒突变基因在杂合状态下具有增加水稻产量的潜力,可应用于杂交稻生产。
实施例7、抑制DLZ基因的表达水平可增加水稻粒宽
以DLZ基因为基础,分别设计引物
RNAi-SpeI-1F:
5’-CACGTGGACCACTAGTATGTGGATGTATGGCTATTTCTGGA-3’(SEQ ID NO:22),
RNAi-SpeI-1R:
5’-GTCCGTACCAACTAGTTCGTATCCTTGCAGCAACTTATTCA-3’(SEQ ID NO:23),
RNAi-BamHI-2F:
5’-TGAATTCGCTGGATCCTCGTATCCTTGCAGCAACTTATTCA-3’(SEQ ID NO:24),
RNAi-BamHI-2R:
5’-GTCGACTGGAGGATCCATGTGGATGTATGGCTATTTCTGGA-3’(SEQ ID NO:25)。
以DLZ基因的cDNA为模板,分别扩增出214bp的片段,通过In-Fusion方法正反向分别连接于表达载体Ubi-intron的酶切位点SpeI和BamHI上,构建成RNAi载体。将该载体通过农杆菌介导的水稻遗传转化法转入粳稻中花11中,对所得的T0代转基因植株进行表达量分析。结果如图9所示,转基因阳性植株的内源DLZ基因表达水平受到了显著的抑制,并且粒宽也显著增大,表明抑制DLZ基因的表达水平可增加水稻粒宽,且粒宽的大小与DLZ基因的表达水平呈正相关。
实施例8、通过启动子编辑实现对DLZ基因表达水平的精细调控
在DLZ基因翻译起始位点ATG上游2087bp的启动子区(SEQ ID NO:26)设置8个CRISPR靶位点,分别为TP1:5’-TTTGACAGCTTCCTGATCTT-3’(SEQ ID NO:27),TP2:5’-CAAGTAAGATGCCAAGAATG-3’(SEQ ID NO:28),TP3:5’-TTGTCAACGGGAGAACAAC-3’(SEQ IDNO:29),TP4:5’-TAGGATATTTGAGCTACGG-3’(SEQ ID NO:30),TP5:5’-TAGAAAGAAGTCTGGAGCA-3’(SEQ ID NO:31),TP6:5’-AACGCCAGCTTGAGGGCAG-3’(SEQ ID NO:32),TP7:5’-TTCTCGTCGTTTCTTGCGTG-3’(SEQ ID NO:33),TP8:5’-GTGTGTGGGTTGACCGAAT-3’(SEQ ID NO:34)。分别合成带有粘性末端的接头引物,如表3所示,按照实施例5的方法构建Pubi::Cas9-U3::TP3-gRNA-U3::TP6-gRNA-U6a::TP1-gRNA-U6a::TP2-gRNA-U6b::TP4-gRNA-U6b::TP5-gRNA-U6c::TP7-gRNA-U6c::TP8-gRNA表达盒,PCR模板与引物组合如表1所示,通用引物序列如表2所示,最后通过农杆菌介导的水稻遗传转化法转入粳稻中花11中。用引物820Pro-2087bp-F:5’-GGAAAGGAAGAAAAGGCTAATATGCTCATC-3’(SEQ ID NO:51)和820Pro-2087bp-R:5’-ATGTCAGGATGTGCTTCTGGGACAC-3’(SEQ ID NO:52)扩增T0代转基因植株的DLZ基因2087bp启动子区,电泳结果如图10所示,很多转基因植株的扩增产物明显小于2087bp,表明这些植株中的DLZ基因启动子区发生了不同程度的片段缺失。将扩增产物连入pEASY-Blunt载体,用M13F和M13R通用引物测序,部分植株的启动子测序结果如SEQ ID NO:53(8-5号单株),SEQ ID NO:54(8-9号单株)所示。其中8-5号单株在TP2和TP5之间发生了750bp左右的缺失;8-9号单株在TP2和TP8之间发生了大片段的缺失,并倒置***了TP5和TP6之间350bp左右的片段。这些结果表明启动子编辑可以产生丰富的序列变异。
进一步检测转基因植株的DLZ基因表达水平变化,结果如图10所示,少数植株的DLZ基因表达水平被抑制到与RNAi基因沉默的效果相当(如8-22号单株),大部分植株的DLZ基因表达水平下降了一半左右。考察转基因植株的粒宽表型,发现DLZ基因表达水平受抑制程度越高,粒宽越大,表达水平下降一半左右的植株,其粒宽的大小介于野生型与基因敲除植株之间,即与dlz杂合突变体的表型相当,表明通过启动子编辑技术可以实现对DLZ基因表达水平的精细调控,实现增产的效果。
实施例9、不同作物中的同源基因分析
将DLZ基因编码的蛋白质序列输入NCBI数据库中进行BLASTP搜索,获得了二穗短柄草(Brachypodium distachyon)(SEQ ID NO:55)、大麦(Hordeum vulgare)(SEQ ID NO:56)、野生稻(Oryza brachyantha)(SEQ ID NO:57)、黍(Panicum hallii)(SEQ ID NO:58)、谷子(Setaria italica)(SEQ ID NO:59)、玉米(Zea mays)(SEQ ID NO:60)、高粱(Sorghumbicolor)(SEQ ID NO:61)等作物基因组中预测的同源蛋白。将水稻DLZ蛋白与这些同源蛋白的氨基酸序列输入ClustalW2网站(https://www.ebi.ac.uk/Tools/msa/clustalw2/)进行序列比对,结果显示来自不同植物的同源蛋白都具有非常相似的保守序列,彼此之间同源性很高(图11),表明DLZ蛋白在不同植物的种子发育中功能保守,起着非常重要的作用。
利用CRISPR/Cas9技术分别对上述7个作物的DLZ基因进行突变,对获得的转基因阳性植株进行种子颗粒大小的性状观察,结果显示转基因阳性植株是DLZ基因功能丧失的突变体,DLZ基因无法正常表达,其种子粒宽也较相应作物的野生型大大增加,并呈现粒大穗多性状,证明定点突变不同物种DLZ的同源基因均导致种子大小的显著增加。
表1 CRISPR载体构建中扩增gRNA表达盒的模板引物组合
表2 CRISPR载体构建中扩增gRNA表达盒的通用引物序列
表3启动子编辑CRISPR载体构建中扩增gRNA表达盒的靶点特异引物序列
序列表
<110> 深圳市作物分子设计育种研究院
未名兴旺***作物设计前沿实验室(北京)有限公司
<120> 调控植物种子大小的基因及其应用
<150> 2018100674731
<151> 2018-01-24
<160> 61
<170> SIPOSequenceListing 1.0
<210> 1
<211> 6146
<212> DNA
<213> Oryza sativa
<400> 1
cactgcccaa ttgcccatgc tccagacttc tttctactcc tacattccac atatctccat 60
ggacagtaac tcctcccaag ctaccacttc aaccctaatc ccctctctct ctcttccgca 120
gaggtagagt gagagagatg gtcagatagc tagattgata tccctctctc tctctcacag 180
acatctcttt ttgcaagatc tcttcttgtt catcatcttc ttcttttttt ctcccccttt 240
tgcttcacca atccatcttt tgtcacgaga tgtgaccgag ctgaagctag tagtagtgga 300
gcagcgaaag caagtacgcc aagaaaaaaa aaaggaagaa gaaagaagaa agaaagaaag 360
aaaaaaacgc cagcttgagg gcagagggca aaagcggcga cgaggagcag tggccaaagc 420
tcagattctt cccgtgggct atttttacca cccgcatccc ctctctttga gccccttggc 480
cgattcattc accgacgcaa agatccaacc cctcttcagg tgtcggcaga tgccgccttt 540
gtgaggtttc cagtgggggg atttctcgtc gtttcttgcg tgcggttgcg ttcttgatcc 600
agtgagcgca cggatatatc cgccctggtt tagtagagag agagagagag agagagagag 660
agagagagag agagagagag agagggggtt cttgattgag ttccaagtgt tggattgggt 720
tcttggagct gttggattgg gtttttttgg gagagagatg ggggtttgga ggtgtgtggg 780
ttgaccgaat tggatcaaga ttattgcggg aggggggggg gggggttgca atggcggatt 840
tggggctgtg gaagcaaggg tggaggtggg tggtgtccca gaagcacatc ctgacatggg 900
cgcacatggc ggcgagcggc ggcaccgaga ggctggcctt cctggtcgac cggcattggc 960
ccgccgtgtc ccgggcctgc gtgagctccg gccgcctcgc gctcgccgcg ctgcggcaat 1020
ggcgcggctg cgcggcgcgc gggatcctgg agatggctag cctgggccct gcgtccgtgt 1080
tcgtcatcct ctggagcttc ttcgtgtgca tcacctcgcc ggcgtgcgcc ctctacgcgc 1140
tcctgggcat ggtacggcat gcaagtcttg cttgctttgc gctttcgcct tgatgatgta 1200
gtggattatg gataacgatt tgtgcgcgtt ctaaatcttg tcatgtgctc gtctttcttt 1260
ttttcttctt tttatcaagg gtggattgca tgttaggtta cctttctttt cgaaaagtat 1320
agttaaagtg gtaattggtg gtacaaaagt agtatgtcat tacactttca tgagattgat 1380
cagtttgatg tgtttctaga ttcatttatg ctttagttat tgcaagttta tactacactt 1440
cagtaattca cacgtgctgt ttctagatgt tattttggaa ccgttcacag tattttaagc 1500
atcatttgca ttagaaagtt ttatctagtt tgtcttgcta gaggaaggag cacatggaaa 1560
ctaacacttg catatttagg gataagcact actggttcta ttcctatttt gtgtatgtta 1620
gctaatgtgt ttcttgctga gtggttcagt ttcaggttca ttagcagatt atcttattgg 1680
ttgatttatg tcaaaatact taaggtcaat tcgtagtttg cacagtgtac ttcaataaca 1740
tgaatgcaac tcgtttcttt gttcagcctg atatttatga aaaatcttat aatgtgatac 1800
tgtgtttaat atgtatgaac ctgtctagag aattactagc tagtgaaatt ctacttgttt 1860
catttcacac aaagtcaact atgggtagac tggttcatga ccatttattt aggctctggt 1920
acacctgtaa ctactgctgt agttgactat atgacttact tatgctggtt ttctactgtg 1980
gtatgagttt ctccctttgg gataccacct gtgttcagtg gaatgtcagg tagtatctga 2040
ccttttcagc tagattgcac tggataaatt atactgaaat aagcaatagg aatgaattcc 2100
aggactatgt ctcctcttgt tctctccttg caattcctct tattgcaatg cagactgaac 2160
cacttgtttt tactgtcatc tggcatactt gttcagttag taacttctac ttgcgagtaa 2220
ggatgcaaga tttcccatgt aagatggata ctataaatat cattttgtct aattgcttaa 2280
tacctttctt ttttcaggga gctgctgggg cagtcattca ttacatgggc tatacgcctg 2340
gtcttttcat tgtaggatta tttggaatat tgattatgtg gatgtatggc tatttctgga 2400
ttacaggaat gcttctgatt gctggaggtt tgttttatct taatatttaa gtctgttcat 2460
aatgataatt ttgtgttttt gtttgtcaaa tccataaatt tttcttcctc cctagttgca 2520
tccgtttgat tcttttgacc taaaggagga tcctctctgg taatgcaggc tgtatgtgct 2580
ctttgaaaca tgcacgattt gtgatacctg tgttggctat gtatgctgtt tattgtgtgg 2640
ctgttcgtgt tggatcgctt ggtgtcttct tgacattgaa tctttctttc ctgacaaatg 2700
atcttctgaa taagttgctg caaggatacg agggaagcac agaagaaaga cagtttgaag 2760
agccaaaaca ttctgatcct gtcatggatg agttctatcg cagttgtgaa tttccctctg 2820
ctcctgatag tgaacctgag actgtttctt ctgcaaagcc cttttgctca acacccgtcc 2880
aggatgtgtt gcatgtacag aaagaggcat ctcctagcaa agtagtgaaa tcggattctg 2940
tttcattgga tgagatgaag aggatcatgg atggtttgac ccattatgaa gttttgggta 3000
ttcctcggaa tagaagtatt gatcaaaaga ttctgaaaaa ggagtaccac agaatggtaa 3060
taaaccacgg ccttctatac aagggaaaat gagaaattca tgttacaatt acttcatttt 3120
catggtacgt atgctttatt tgtctaggtc ctgcttgtac atcctgataa aaatatggga 3180
aatccactgg cctgtgaatc attcaaaaag cttcagtcag cttatgaggt aaactacaat 3240
ggaagtttat gtcttttctc ttccttgatt atattacagt taaatctggt tgaatatctg 3300
ctcttgatac caaccatggc ttctatacct ggataaaggg taatcattgt agttatgctg 3360
caggtactct cagatttcac aaagaaaaac acttacgacg accaactgag gaaagaagaa 3420
tcacgtaaaa tgactcagag atcacgtgtt gtctctcaac aggtgggttc tagttttcac 3480
aaatttagaa tccacatggt tggattattt ctttaacata tcttatcaat tatccaagca 3540
tacgaatgca gtttattcat gctctcatgt ccttgaccta ctgacctact tgctgttttc 3600
ctttatgggg cccatttgta atttgataaa ctcatcttgc agactggggt agagtttctc 3660
tccgaagagt ccaggcgtat acagtgcaca aagtgtggta attttcatct gtggatatgt 3720
accaagaaaa gcaaagcaaa agcaagatgg tgtcaggttt ggaggccaga attttttttt 3780
caggtacttt taatcgagag tgttcttaca gctaattttg tgggaaccat gtactgtagg 3840
attgctctga ttttcatcca gctaaggatg gagatggatg ggtggaaaat aaattttcgt 3900
catccttcaa ggtaatgttt tataagcaca tcatatgaag agttcacttt attttactta 3960
atgcttgcct tctacagtac tcatagacag agatctagtg tcaatacaat tttaactact 4020
agaaaatgga aattgagtac atattgattt cgaacaaatg gagaatgagg ttttatgaat 4080
ggaagcacaa tgttctgaat gttttgatac aaaattaccg ggcgctgttt cccactgtca 4140
agcttcagtt cctagtactt gttattgcct gaagttagtc atgtgtgttc cgagaccaac 4200
tttggacttg agcaagctca gttttagctg tgtcaagctg atgatctttt atcttctaat 4260
tgtattccac ctaaagaaag catctcattc caagtgttag gtacagtcat tttgttcatt 4320
ccataagcaa cttattctga ctataaggtg agattcagaa attactcagc ttaaaaatgt 4380
gcacacattt tgtagtttcc aactataatg tgtaaattct tcacttctct ttattgaact 4440
ataatgtgta aattcttcag tactctttgt tgaatcaaag tgcattgttc acttcaggaa 4500
atacctcgag cttttgtttg tgcggagagt aaggtatttg atgtgtctga atgggctact 4560
tgccaggtga gtgtctgacg atgttttata tgtttgattt aagttgacat gtatgtgcat 4620
ttgcagcagt gatttttgga tgtctcaatt gatttgatgt catctccata tgcatatttt 4680
tatactcggt tctctgctgt tttcgatgtc ttaactgact atagatatgc ctttggtcaa 4740
ttgactttgt tcagttttgt atttgatgca tattcaaacg tccagattga ctgttttact 4800
ttaaaaattg tttcagttgg taaatgaaaa tttgcttact tcattggaga taggataatt 4860
catgcatgcc atagcccata gccttatttt tctgtgtcaa gtttgtcatg gctataataa 4920
acacaacata ttaatcgcac ccgcatgtca tcccgactga tatctcaatt attgacatac 4980
ctatctaaga gaagagccaa caatgatgaa agtaaaggac taatttggct gtgcaaaatt 5040
ggaccaaaag tttattttac atttcatact tgcttcattc aacataaaca tcaaaatctg 5100
gtacgccaat tttctggcga tacatacagg gtcttgtgat aggttcattt gcatacatta 5160
aaaatgggag cctttctaac tctgttttct ttgcttgatt gtctagggca tggagtgcaa 5220
acctaacact cacggcccat cttttatggt aaacatggtt ggcgcagata ggatgtctca 5280
gagatcctac agttctcgct atccctttag tttgaatgct gagatgatcc ctgaagatga 5340
atttgagcta tggcttcaac aagcattggc atcaggtgtc ttctctgaca gcccgaaacg 5400
caggaaaagc tggagcccct tcaaactacc tcaaaaaggg ataaaaagtt ggcggcgatc 5460
ctcataaggg catagcatta aacagcatgg atctcacctg agtacaacac tgaaaaaggc 5520
tatactcttt gtgaatgtaa atagactgac caacaatttg cctggatgag caactaattt 5580
tgtccaaaaa gagacactga aacaaggggg gtaaaaggaa caaacgctta agacatgact 5640
gcaatgaatc tgactgttga aattagtgtt ctctgcaatg agatcccgcg agttttatcc 5700
gaaaaggtca gatactggga tggcgtgtca ttcatcagtt catcctaaag ctcggaaggg 5760
tatctctgta gcatgttaac ttcagtagtt ttaggggatc ggcatctgag agaatttcaa 5820
aacttcatac ctggttgcca gcataagttc tgcaggtgtt gaaaagttgt tgatcagagt 5880
agcaatttaa ggtctgatgt ttctggggaa cagtaggaga gaaaaaaatg acaaaaaaaa 5940
gagagagttg gttgtaaata catgaaaagt tttcatcaga aattagtatt gtaacattgt 6000
acactgtgat tacatcctgt gcaatactcc cataattcag atctgtgttg taatacacta 6060
catacatcct acaattttct ggtgataata gagatctaat tctcacctat tatcgttatt 6120
tatggttagt cagttactgc tctgta 6146
<210> 2
<211> 1635
<212> DNA
<213> Oryza sativa
<400> 2
atggcggatt tggggctgtg gaagcaaggg tggaggtggg tggtgtccca gaagcacatc 60
ctgacatggg cgcacatggc ggcgagcggc ggcaccgaga ggctggcctt cctggtcgac 120
cggcattggc ccgccgtgtc ccgggcctgc gtgagctccg gccgcctcgc gctcgccgcg 180
ctgcggcaat ggcgcggctg cgcggcgcgc gggatcctgg agatggctag cctgggccct 240
gcgtccgtgt tcgtcatcct ctggagcttc ttcgtgtgca tcacctcgcc ggcgtgcgcc 300
ctctacgcgc tcctgggcat gggagctgct ggggcagtca ttcattacat gggctatacg 360
cctggtcttt tcattgtagg attatttgga atattgatta tgtggatgta tggctatttc 420
tggattacag gaatgcttct gattgctgga ggctgtatgt gctctttgaa acatgcacga 480
tttgtgatac ctgtgttggc tatgtatgct gtttattgtg tggctgttcg tgttggatcg 540
cttggtgtct tcttgacatt gaatctttct ttcctgacaa atgatcttct gaataagttg 600
ctgcaaggat acgagggaag cacagaagaa agacagtttg aagagccaaa acattctgat 660
cctgtcatgg atgagttcta tcgcagttgt gaatttccct ctgctcctga tagtgaacct 720
gagactgttt cttctgcaaa gcccttttgc tcaacacccg tccaggatgt gttgcatgta 780
cagaaagagg catctcctag caaagtagtg aaatcggatt ctgtttcatt ggatgagatg 840
aagaggatca tggatggttt gacccattat gaagttttgg gtattcctcg gaatagaagt 900
attgatcaaa agattctgaa aaaggagtac cacagaatgg tcctgcttgt acatcctgat 960
aaaaatatgg gaaatccact ggcctgtgaa tcattcaaaa agcttcagtc agcttatgag 1020
gtactctcag atttcacaaa gaaaaacact tacgacgacc aactgaggaa agaagaatca 1080
cgtaaaatga ctcagagatc acgtgttgtc tctcaacaga ctggggtaga gtttctctcc 1140
gaagagtcca ggcgtataca gtgcacaaag tgtggtaatt ttcatctgtg gatatgtacc 1200
aagaaaagca aagcaaaagc aagatggtgt caggattgct ctgattttca tccagctaag 1260
gatggagatg gatgggtgga aaataaattt tcgtcatcct tcaaggaaat acctcgagct 1320
tttgtttgtg cggagagtaa ggtatttgat gtgtctgaat gggctacttg ccagggcatg 1380
gagtgcaaac ctaacactca cggcccatct tttatggtaa acatggttgg cgcagatagg 1440
atgtctcaga gatcctacag ttctcgctat ccctttagtt tgaatgctga gatgatccct 1500
gaagatgaat ttgagctatg gcttcaacaa gcattggcat caggtgtctt ctctgacagc 1560
ccgaaacgca ggaaaagctg gagccccttc aaactacctc aaaaagggat aaaaagttgg 1620
cggcgatcct cataa 1635
<210> 3
<211> 544
<212> PRT
<213> Oryza sativa
<400> 3
Met Ala Asp Leu Gly Leu Trp Lys Gln Gly Trp Arg Trp Val Val Ser
1 5 10 15
Gln Lys His Ile Leu Thr Trp Ala His Met Ala Ala Ser Gly Gly Thr
20 25 30
Glu Arg Leu Ala Phe Leu Val Asp Arg His Trp Pro Ala Val Ser Arg
35 40 45
Ala Cys Val Ser Ser Gly Arg Leu Ala Leu Ala Ala Leu Arg Gln Trp
50 55 60
Arg Gly Cys Ala Ala Arg Gly Ile Leu Glu Met Ala Ser Leu Gly Pro
65 70 75 80
Ala Ser Val Phe Val Ile Leu Trp Ser Phe Phe Val Cys Ile Thr Ser
85 90 95
Pro Ala Cys Ala Leu Tyr Ala Leu Leu Gly Met Gly Ala Ala Gly Ala
100 105 110
Val Ile His Tyr Met Gly Tyr Thr Pro Gly Leu Phe Ile Val Gly Leu
115 120 125
Phe Gly Ile Leu Ile Met Trp Met Tyr Gly Tyr Phe Trp Ile Thr Gly
130 135 140
Met Leu Leu Ile Ala Gly Gly Cys Met Cys Ser Leu Lys His Ala Arg
145 150 155 160
Phe Val Ile Pro Val Leu Ala Met Tyr Ala Val Tyr Cys Val Ala Val
165 170 175
Arg Val Gly Ser Leu Gly Val Phe Leu Thr Leu Asn Leu Ser Phe Leu
180 185 190
Thr Asn Asp Leu Leu Asn Lys Leu Leu Gln Gly Tyr Glu Gly Ser Thr
195 200 205
Glu Glu Arg Gln Phe Glu Glu Pro Lys His Ser Asp Pro Val Met Asp
210 215 220
Glu Phe Tyr Arg Ser Cys Glu Phe Pro Ser Ala Pro Asp Ser Glu Pro
225 230 235 240
Glu Thr Val Ser Ser Ala Lys Pro Phe Cys Ser Thr Pro Val Gln Asp
245 250 255
Val Leu His Val Gln Lys Glu Ala Ser Pro Ser Lys Val Val Lys Ser
260 265 270
Asp Ser Val Ser Leu Asp Glu Met Lys Arg Ile Met Asp Gly Leu Thr
275 280 285
His Tyr Glu Val Leu Gly Ile Pro Arg Asn Arg Ser Ile Asp Gln Lys
290 295 300
Ile Leu Lys Lys Glu Tyr His Arg Met Val Leu Leu Val His Pro Asp
305 310 315 320
Lys Asn Met Gly Asn Pro Leu Ala Cys Glu Ser Phe Lys Lys Leu Gln
325 330 335
Ser Ala Tyr Glu Val Leu Ser Asp Phe Thr Lys Lys Asn Thr Tyr Asp
340 345 350
Asp Gln Leu Arg Lys Glu Glu Ser Arg Lys Met Thr Gln Arg Ser Arg
355 360 365
Val Val Ser Gln Gln Thr Gly Val Glu Phe Leu Ser Glu Glu Ser Arg
370 375 380
Arg Ile Gln Cys Thr Lys Cys Gly Asn Phe His Leu Trp Ile Cys Thr
385 390 395 400
Lys Lys Ser Lys Ala Lys Ala Arg Trp Cys Gln Asp Cys Ser Asp Phe
405 410 415
His Pro Ala Lys Asp Gly Asp Gly Trp Val Glu Asn Lys Phe Ser Ser
420 425 430
Ser Phe Lys Glu Ile Pro Arg Ala Phe Val Cys Ala Glu Ser Lys Val
435 440 445
Phe Asp Val Ser Glu Trp Ala Thr Cys Gln Gly Met Glu Cys Lys Pro
450 455 460
Asn Thr His Gly Pro Ser Phe Met Val Asn Met Val Gly Ala Asp Arg
465 470 475 480
Met Ser Gln Arg Ser Tyr Ser Ser Arg Tyr Pro Phe Ser Leu Asn Ala
485 490 495
Glu Met Ile Pro Glu Asp Glu Phe Glu Leu Trp Leu Gln Gln Ala Leu
500 505 510
Ala Ser Gly Val Phe Ser Asp Ser Pro Lys Arg Arg Lys Ser Trp Ser
515 520 525
Pro Phe Lys Leu Pro Gln Lys Gly Ile Lys Ser Trp Arg Arg Ser Ser
530 535 540
<210> 4
<211> 6146
<212> DNA
<213> Oryza sativa
<400> 4
cactgcccaa ttgcccatgc tccagacttc tttctactcc tacattccac atatctccat 60
ggacagtaac tcctcccaag ctaccacttc aaccctaatc ccctctctct ctcttccgca 120
gaggtagagt gagagagatg gtcagatagc tagattgata tccctctctc tctctcacag 180
acatctcttt ttgcaagatc tcttcttgtt catcatcttc ttcttttttt ctcccccttt 240
tgcttcacca atccatcttt tgtcacgaga tgtgaccgag ctgaagctag tagtagtgga 300
gcagcgaaag caagtacgcc aagaaaaaaa aaaggaagaa gaaagaagaa agaaagaaag 360
aaaaaaacgc cagcttgagg gcagagggca aaagcggcga cgaggagcag tggccaaagc 420
tcagattctt cccgtgggct atttttacca cccgcatccc ctctctttga gccccttggc 480
cgattcattc accgacgcaa agatccaacc cctcttcagg tgtcggcaga tgccgccttt 540
gtgaggtttc cagtgggggg atttctcgtc gtttcttgcg tgcggttgcg ttcttgatcc 600
agtgagcgca cggatatatc cgccctggtt tagtagagag agagagagag agagagagag 660
agagagagag agagagagag agagggggtt cttgattgag ttccaagtgt tggattgggt 720
tcttggagct gttggattgg gtttttttgg gagagagatg ggggtttgga ggtgtgtggg 780
ttgaccgaat tggatcaaga ttattgcggg aggggggggg gggggttgca atggcggatt 840
tggggctgtg gaagcaaggg tggaggtggg tggtgtccca gaagcacatc ctgacatggg 900
cgcacatggc ggcgagcggc ggcaccgaga ggctggcctt cctggtcgac cggcattggc 960
ccgccgtgtc ccgggcctgc gtgagctccg gccgcctcgc gctcgccgcg ctgcggcaat 1020
ggcgcggctg cgcggcgcgc gggatcctgg agatggctag cctgggccct gcgtccgtgt 1080
tcgtcatcct ctggagcttc ttcgtgtgca tcacctcgcc ggcgtgcgcc ctctacgcgc 1140
tcctgggcat ggtacggcat gcaagtcttg cttgctttgc gctttcgcct tgatgatgta 1200
gtggattatg gataacgatt tgtgcgcgtt ctaaatcttg tcatgtgctc gtctttcttt 1260
ttttcttctt tttatcaagg gtggattgca tgttaggtta cctttctttt cgaaaagtat 1320
agttaaagtg gtaattggtg gtacaaaagt agtatgtcat tacactttca tgagattgat 1380
cagtttgatg tgtttctaga ttcatttatg ctttagttat tgcaagttta tactacactt 1440
cagtaattca cacgtgctgt ttctagatgt tattttggaa ccgttcacag tattttaagc 1500
atcatttgca ttagaaagtt ttatctagtt tgtcttgcta gaggaaggag cacatggaaa 1560
ctaacacttg catatttagg gataagcact actggttcta ttcctatttt gtgtatgtta 1620
gctaatgtgt ttcttgctga gtggttcagt ttcaggttca ttagcagatt atcttattgg 1680
ttgatttatg tcaaaatact taaggtcaat tcgtagtttg cacagtgtac ttcaataaca 1740
tgaatgcaac tcgtttcttt gttcagcctg atatttatga aaaatcttat aatgtgatac 1800
tgtgtttaat atgtatgaac ctgtctagag aattactagc tagtgaaatt ctacttgttt 1860
catttcacac aaagtcaact atgggtagac tggttcatga ccatttattt aggctctggt 1920
acacctgtaa ctactgctgt agttgactat atgacttact tatgctggtt ttctactgtg 1980
gtatgagttt ctccctttgg gataccacct gtgttcagtg gaatgtcagg tagtatctga 2040
ccttttcagc tagattgcac tggataaatt atactgaaat aagcaatagg aatgaattcc 2100
aggactatgt ctcctcttgt tctctccttg caattcctct tattgcaatg cagactgaac 2160
cacttgtttt tactgtcatc tggcatactt gttcagttag taacttctac ttgcgagtaa 2220
ggatgcaaga tttcccatgt aagatggata ctataaatat cattttgtct aattgcttaa 2280
tacctttctt ttttcaggga gctgctgggg cagtcattca ttacatgggc tatacgcctg 2340
gtcttttcat tgtaggatta tttggaatat tgattatgtg gatgtatggc tatttctgga 2400
ttacaggaat gcttctgatt gctggaggtt tgttttatct taatatttaa gtctgttcat 2460
aatgataatt ttgtgttttt gtttgtcaaa tccataaatt tttcttcctc cctagttgca 2520
tccgtttgat tcttttgacc taaaggagga tcctctctgg taatgcaggc tgtatgtgct 2580
ctttgaaaca tgcacgattt gtgatacctg tgttggctat gtatgctgtt tattgtgtgg 2640
ctgttcgtgt tggatcgctt ggtgtcttct tgacattgaa tctttctttc ctgacaaatg 2700
atcttctgaa taagttgctg caaggatacg agggaagcac agaagaaaga cagtttgaag 2760
agccaaaaca ttctgatcct gtcatggatg agttctatcg cagttgtgaa tttccctctg 2820
ctcctgatag tgaacctgag actgtttctt ctgcaaagcc cttttgctca acacccgtcc 2880
aggatgtgtt gcatgtacag aaagaggcat ctcctagcaa agtagtgaaa tcggattctg 2940
tttcattgga tgagatgaag aggatcatgg atggtttgac ccattatgaa gttttgggta 3000
ttcctcggaa tagaagtatt gatcaaaaga ttctgaaaaa ggagtaccac agaatggtaa 3060
taaaccacgg ccttctatac aagggaaaat gagaaattca tgttacaatt acttcatttt 3120
catggtacgt atgctttatt tgtctaggtc ctgcttgtac atcctgattt aaatatggga 3180
aatccactgg cctgtgaatc attcaaaaag cttcagtcag cttatgaggt aaactacaat 3240
ggaagtttat gtcttttctc ttccttgatt atattacagt taaatctggt tgaatatctg 3300
ctcttgatac caaccatggc ttctatacct ggataaaggg taatcattgt agttatgctg 3360
caggtactct cagatttcac aaagaaaaac acttacgacg accaactgag gaaagaagaa 3420
tcacgtaaaa tgactcagag atcacgtgtt gtctctcaac aggtgggttc tagttttcac 3480
aaatttagaa tccacatggt tggattattt ctttaacata tcttatcaat tatccaagca 3540
tacgaatgca gtttattcat gctctcatgt ccttgaccta ctgacctact tgctgttttc 3600
ctttatgggg cccatttgta atttgataaa ctcatcttgc agactggggt agagtttctc 3660
tccgaagagt ccaggcgtat acagtgcaca aagtgtggta attttcatct gtggatatgt 3720
accaagaaaa gcaaagcaaa agcaagatgg tgtcaggttt ggaggccaga attttttttt 3780
caggtacttt taatcgagag tgttcttaca gctaattttg tgggaaccat gtactgtagg 3840
attgctctga ttttcatcca gctaaggatg gagatggatg ggtggaaaat aaattttcgt 3900
catccttcaa ggtaatgttt tataagcaca tcatatgaag agttcacttt attttactta 3960
atgcttgcct tctacagtac tcatagacag agatctagtg tcaatacaat tttaactact 4020
agaaaatgga aattgagtac atattgattt cgaacaaatg gagaatgagg ttttatgaat 4080
ggaagcacaa tgttctgaat gttttgatac aaaattaccg ggcgctgttt cccactgtca 4140
agcttcagtt cctagtactt gttattgcct gaagttagtc atgtgtgttc cgagaccaac 4200
tttggacttg agcaagctca gttttagctg tgtcaagctg atgatctttt atcttctaat 4260
tgtattccac ctaaagaaag catctcattc caagtgttag gtacagtcat tttgttcatt 4320
ccataagcaa cttattctga ctataaggtg agattcagaa attactcagc ttaaaaatgt 4380
gcacacattt tgtagtttcc aactataatg tgtaaattct tcacttctct ttattgaact 4440
ataatgtgta aattcttcag tactctttgt tgaatcaaag tgcattgttc acttcaggaa 4500
atacctcgag cttttgtttg tgcggagagt aaggtatttg atgtgtctga atgggctact 4560
tgccaggtga gtgtctgacg atgttttata tgtttgattt aagttgacat gtatgtgcat 4620
ttgcagcagt gatttttgga tgtctcaatt gatttgatgt catctccata tgcatatttt 4680
tatactcggt tctctgctgt tttcgatgtc ttaactgact atagatatgc ctttggtcaa 4740
ttgactttgt tcagttttgt atttgatgca tattcaaacg tccagattga ctgttttact 4800
ttaaaaattg tttcagttgg taaatgaaaa tttgcttact tcattggaga taggataatt 4860
catgcatgcc atagcccata gccttatttt tctgtgtcaa gtttgtcatg gctataataa 4920
acacaacata ttaatcgcac ccgcatgtca tcccgactga tatctcaatt attgacatac 4980
ctatctaaga gaagagccaa caatgatgaa agtaaaggac taatttggct gtgcaaaatt 5040
ggaccaaaag tttattttac atttcatact tgcttcattc aacataaaca tcaaaatctg 5100
gtacgccaat tttctggcga tacatacagg gtcttgtgat aggttcattt gcatacatta 5160
aaaatgggag cctttctaac tctgttttct ttgcttgatt gtctagggca tggagtgcaa 5220
acctaacact cacggcccat cttttatggt aaacatggtt ggcgcagata ggatgtctca 5280
gagatcctac agttctcgct atccctttag tttgaatgct gagatgatcc ctgaagatga 5340
atttgagcta tggcttcaac aagcattggc atcaggtgtc ttctctgaca gcccgaaacg 5400
caggaaaagc tggagcccct tcaaactacc tcaaaaaggg ataaaaagtt ggcggcgatc 5460
ctcataaggg catagcatta aacagcatgg atctcacctg agtacaacac tgaaaaaggc 5520
tatactcttt gtgaatgtaa atagactgac caacaatttg cctggatgag caactaattt 5580
tgtccaaaaa gagacactga aacaaggggg gtaaaaggaa caaacgctta agacatgact 5640
gcaatgaatc tgactgttga aattagtgtt ctctgcaatg agatcccgcg agttttatcc 5700
gaaaaggtca gatactggga tggcgtgtca ttcatcagtt catcctaaag ctcggaaggg 5760
tatctctgta gcatgttaac ttcagtagtt ttaggggatc ggcatctgag agaatttcaa 5820
aacttcatac ctggttgcca gcataagttc tgcaggtgtt gaaaagttgt tgatcagagt 5880
agcaatttaa ggtctgatgt ttctggggaa cagtaggaga gaaaaaaatg acaaaaaaaa 5940
gagagagttg gttgtaaata catgaaaagt tttcatcaga aattagtatt gtaacattgt 6000
acactgtgat tacatcctgt gcaatactcc cataattcag atctgtgttg taatacacta 6060
catacatcct acaattttct ggtgataata gagatctaat tctcacctat tatcgttatt 6120
tatggttagt cagttactgc tctgta 6146
<210> 5
<211> 544
<212> PRT
<213> Oryza sativa
<400> 5
Met Ala Asp Leu Gly Leu Trp Lys Gln Gly Trp Arg Trp Val Val Ser
1 5 10 15
Gln Lys His Ile Leu Thr Trp Ala His Met Ala Ala Ser Gly Gly Thr
20 25 30
Glu Arg Leu Ala Phe Leu Val Asp Arg His Trp Pro Ala Val Ser Arg
35 40 45
Ala Cys Val Ser Ser Gly Arg Leu Ala Leu Ala Ala Leu Arg Gln Trp
50 55 60
Arg Gly Cys Ala Ala Arg Gly Ile Leu Glu Met Ala Ser Leu Gly Pro
65 70 75 80
Ala Ser Val Phe Val Ile Leu Trp Ser Phe Phe Val Cys Ile Thr Ser
85 90 95
Pro Ala Cys Ala Leu Tyr Ala Leu Leu Gly Met Gly Ala Ala Gly Ala
100 105 110
Val Ile His Tyr Met Gly Tyr Thr Pro Gly Leu Phe Ile Val Gly Leu
115 120 125
Phe Gly Ile Leu Ile Met Trp Met Tyr Gly Tyr Phe Trp Ile Thr Gly
130 135 140
Met Leu Leu Ile Ala Gly Gly Cys Met Cys Ser Leu Lys His Ala Arg
145 150 155 160
Phe Val Ile Pro Val Leu Ala Met Tyr Ala Val Tyr Cys Val Ala Val
165 170 175
Arg Val Gly Ser Leu Gly Val Phe Leu Thr Leu Asn Leu Ser Phe Leu
180 185 190
Thr Asn Asp Leu Leu Asn Lys Leu Leu Gln Gly Tyr Glu Gly Ser Thr
195 200 205
Glu Glu Arg Gln Phe Glu Glu Pro Lys His Ser Asp Pro Val Met Asp
210 215 220
Glu Phe Tyr Arg Ser Cys Glu Phe Pro Ser Ala Pro Asp Ser Glu Pro
225 230 235 240
Glu Thr Val Ser Ser Ala Lys Pro Phe Cys Ser Thr Pro Val Gln Asp
245 250 255
Val Leu His Val Gln Lys Glu Ala Ser Pro Ser Lys Val Val Lys Ser
260 265 270
Asp Ser Val Ser Leu Asp Glu Met Lys Arg Ile Met Asp Gly Leu Thr
275 280 285
His Tyr Glu Val Leu Gly Ile Pro Arg Asn Arg Ser Ile Asp Gln Lys
290 295 300
Ile Leu Lys Lys Glu Tyr His Arg Met Val Leu Leu Val His Pro Asp
305 310 315 320
Leu Asn Met Gly Asn Pro Leu Ala Cys Glu Ser Phe Lys Lys Leu Gln
325 330 335
Ser Ala Tyr Glu Val Leu Ser Asp Phe Thr Lys Lys Asn Thr Tyr Asp
340 345 350
Asp Gln Leu Arg Lys Glu Glu Ser Arg Lys Met Thr Gln Arg Ser Arg
355 360 365
Val Val Ser Gln Gln Thr Gly Val Glu Phe Leu Ser Glu Glu Ser Arg
370 375 380
Arg Ile Gln Cys Thr Lys Cys Gly Asn Phe His Leu Trp Ile Cys Thr
385 390 395 400
Lys Lys Ser Lys Ala Lys Ala Arg Trp Cys Gln Asp Cys Ser Asp Phe
405 410 415
His Pro Ala Lys Asp Gly Asp Gly Trp Val Glu Asn Lys Phe Ser Ser
420 425 430
Ser Phe Lys Glu Ile Pro Arg Ala Phe Val Cys Ala Glu Ser Lys Val
435 440 445
Phe Asp Val Ser Glu Trp Ala Thr Cys Gln Gly Met Glu Cys Lys Pro
450 455 460
Asn Thr His Gly Pro Ser Phe Met Val Asn Met Val Gly Ala Asp Arg
465 470 475 480
Met Ser Gln Arg Ser Tyr Ser Ser Arg Tyr Pro Phe Ser Leu Asn Ala
485 490 495
Glu Met Ile Pro Glu Asp Glu Phe Glu Leu Trp Leu Gln Gln Ala Leu
500 505 510
Ala Ser Gly Val Phe Ser Asp Ser Pro Lys Arg Arg Lys Ser Trp Ser
515 520 525
Pro Phe Lys Leu Pro Gln Lys Gly Ile Lys Ser Trp Arg Arg Ser Ser
530 535 540
<210> 6
<211> 20
<212> DNA
<213> 人工合成(Artificial synthesized)
<400> 6
agtccaggcg tatacagtgc 20
<210> 7
<211> 20
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 7
tcagagcaat cctgacacca 20
<210> 8
<211> 21
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 8
caaccagctg aggcccaaga a 21
<210> 9
<211> 22
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 9
ccagggagat aacaacggaa gc 22
<210> 10
<211> 23
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 10
ccttcctggt cgaccggcat tgg 23
<210> 11
<211> 23
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 11
gctgttcgtg ttggatcgct tgg 23
<210> 12
<211> 24
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 12
ggcaccttcc tggtcgaccg gcat 24
<210> 13
<211> 24
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 13
aaacatgccg gtcgaccagg aagg 24
<210> 14
<211> 23
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 14
gccgctgttc gtgttggatc gct 23
<210> 15
<211> 23
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 15
aaacagcgat ccaacacgaa cag 23
<210> 16
<211> 18
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 16
ctgacatggg cgcacatg 18
<210> 17
<211> 22
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 17
cctcgtatcc ttgcagcaac tt 22
<210> 18
<211> 20
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 18
ggtgatgcac acgaagaagc 20
<210> 19
<211> 20
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 19
tccctagttg catccgtttg 20
<210> 20
<211> 6118
<212> DNA
<213> Oryza sativa
<400> 20
cactgcccaa ttgcccatgc tccagacttc tttctactcc tacattccac atatctccat 60
ggacagtaac tcctcccaag ctaccacttc aaccctaatc ccctctctct ctcttccgca 120
gaggtagagt gagagagatg gtcagatagc tagattgata tccctctctc tctctcacac 180
acatctcttt ttgcaagatc tcttcttgtt catcatcttc ttcttttttt ctcccccttt 240
tgcttcacca atccatcttt tgtcacgaga tgtggccgag ctgaagctag tagtagtgga 300
gcagcgaaag caagtacgcc aagaaaaaaa aaaggaagaa gaaagaagaa agaaagaaag 360
aaaaaaacgc cagcttgagg gcagagggca aaagcggcga cgaggagcag tggccaaagc 420
tcagattctt cccgtgggct atttttacca cccgcatccc ctctctttga gccccttggc 480
cgattcattc accgacgcaa agatccaacc cctcttcagg tgtcggcaga tgccgccttt 540
gtgaggtttc cagtgggggg atttctcgtc gtttcttgcg tgcggttgcg ttcttgatcc 600
agtgagcgca cggatatatc cgccctggtt tagtagagag agagagagag agagagagag 660
agagagagag aggttcttga ttgagttcca agtgttggat tgggttcttg gagctgttgg 720
attgggtttt tttgggagag agatgggggt ttggaggtgt gtgggttgac cgaattggat 780
caagattatt gcgggagggg gggggggggg ttgcaatggc ggatttgggg ctgtggaagc 840
aagggtggag gtgggtggtg tcccagaagc acatcctgac atgggcgcac atggcggcga 900
gcggcggcac cgagaggctg gccttcctgg tcgaccggca ttggcccgcc gtgtcccggg 960
cctgcgtgag ctccggccgc ctcgcgctcg ccgcgctgcg gcaatggcgc ggctgcgcgg 1020
cgcgcgggat cctggagatg gctagcctgg gccctgcgtc cgtgttcgtc atcctctgga 1080
gcttcttcgt gtgcatcacc tcgccggcgt gcgccctcta cgcgctcctg ggcatggtac 1140
ggcatgcaag tcttgcttgc tttgcgcttt cgccttgatg atgtagtgga ttatggataa 1200
cgatttgtgc gcgttctaaa tcttgtcatg tgctcgtctt tctttttttc ttctttttat 1260
caagggtgga ttgcatgtta ggttaccttt cttttcgaaa agtatagtta aagtggtaat 1320
tggtggtaca aaagtagtat gtcattacac tttcatgaga ttgatcagtt tgatgtgttt 1380
ctagattcat ttatgcttta gttattgcaa gtttatacta cacttcagta attcacacgt 1440
gctgtttcta gatgttattt tggaaccgtt cacagtattt taagcagaga aagttttatc 1500
tagtttgtct tgctagagga aggagcacat ggaaactaac acttgcatat ttagggataa 1560
gcactactgg ttctattcct attttgtgta tgttagctaa tgtgtttctt gctgagtggt 1620
tcagtttcag gttcattagc agattatctt attggttgat ttatgtcaaa atacttaagg 1680
tcaattcgta gtttgcacag tgtacttcaa taacatgaat gcaactcgtt tctttgttca 1740
gcctgatatt tatgaaaaat cttataatgt gatactgtgt ttaatatgta tgaacctgtc 1800
tagagaatta ctagctagtg aaattctact tgtttcattt cacacaaagt caactatggg 1860
tagactggtt catgaccatt tatttaggct ctggtacacc tgtaactact gctgtagttg 1920
actatatgac ttacttctgc tggttttcta ctgtggtatg agtttctccc tttgggatac 1980
cacctgtgtt cagtggaatg tcaggtagta tctgaccttt tcagctagat tgcactggat 2040
aaattatact gaaataagca ataggaatga attccaggac tatgtctcct cttgttctct 2100
ccttgcaatt cctcttattg caatgcagac tgaaccactt gtttttacta tcatctggca 2160
tacttgttca gttagtaact tctacttgcg agtaaggatg caagatttcc catgtaagat 2220
ggatactata aatatcattt tgtctaattg cttaatacct ttcttttttc agggagctgc 2280
tggggcagtc attcattaca tgggctatac gcctggtctt ttcattgtag gattatttgg 2340
aatattgatt atgtggatgt atggctattt ctggattaca ggaatgcttc tgattgctgg 2400
aggtttgttt tatcttaata tttaagtctg ttcataatga taattttgtg tttttgtttg 2460
tcaaatccat aaatttttct tcctccctag ttgcatccgt ttgattcttt tgacctaaag 2520
gaggatcctc tctggtaatg caggctgtat gtgctctttg aaacatgcac gatttgtgat 2580
acctgtgttg gctatgtatg ctgtttattg tgtggctgtt cgtgttggat cgcttggtgt 2640
cttcttgaca ttgaatcttt ctttcctgac aaatgatctt ctgaataagt tgctgcaagg 2700
atacgaggga agcacagaag aaagacagtt tgaagagcca aaacattctg atcctgtcat 2760
ggatgagttc tatcgcagtt gtgaatttcc ctctgctcct gatagtgaac ctgagactgt 2820
ttcttctgca aagccctttt gctcaacacc cgtccaggat gtgttgcatg tacagaaaga 2880
ggcatctcct agcaaagtag tgaaatcgga ttctgtttca ttggatgaga tgaagaggat 2940
catggatggt ttgacccatt atgaagtttt gggtattcct cggaatagaa gtattgatca 3000
aaagattctg aaaaaggagt accacagaat ggtaataaac cacggccttc tatacaaggg 3060
aaaatgagaa attcatgtta caattacttc attttcatgg tacgtatgct ttatttgtct 3120
aggtcctgct tgtacatcct gataaaaata tgggaaatcc actggcctgt gaatcattca 3180
aaaagcttca gtcagcttat gaggtaaact acaatggaag tttatgtctt ttctcttcct 3240
tgattatatt acagttaaat ctggttgaat atctgctctt gataccaacc atggcttcta 3300
tacctggata aagggtaatc attgtagtta tgctgcaggt actctcagat ttcacaaaga 3360
aaaacactta cgacgaccaa ctgaggaaag aagaatcacg taaaatgact cagagatcac 3420
gtgttgtctc tcaacaggtg ggttctagtt ttcacaaatt tagaatccac atggttggat 3480
tatttcttta acatatctta tcaattatcc aagcatacga atgcagttta ttcatgctct 3540
catgtccttg acctactgac ctacttgctg ttttccttta tggggcccat ttgtaatttg 3600
ataaactcat cttgcagact ggggtagagt ttctctccga agagtccagg cgtatacagt 3660
gcacaaaatg tggtaatttt catctgtgga tatgtaccaa gaaaagcaaa gcaaaagcaa 3720
gatggtgtca ggtttggagg ccagaatttt tttttcaggt acttttaatc gagagtgttc 3780
ttacagctaa ttttgtggga accatgtact gtaggattgc tctgattttc atccagctaa 3840
ggatggagat ggatgggtgg aaaataaatt ttcgtcatcc ttcaaggtaa tgttttataa 3900
gcacatcata tgaagagttc actttatttt acttaatgct tgccttctac agtactcata 3960
gacagagatc tagtgtcaat agaattttaa ctactagaaa atggaaattg agtacatgtt 4020
gatttcgaac aaatggagaa tgaggtttta tgaatggaag cacaatgttc tgaatgtttt 4080
gatacaaaat taccgggcgc tgtttcccac tgtcaagctt cagttcctag tacttgttat 4140
tgcctgaagt tagtcatgtg tgttccgaga ccaactttgg acttgagcaa gctcagtttt 4200
agctgtgtca agctgatgat cttttatctt ctaattgtat tccacctaaa gaaagcatct 4260
cattccaagt gttaggtaca gtcattttgt tcattccata agcaacttat tctgactata 4320
gggtgagatt cagaaattac tcagcttaaa aatgtgcaca cattttgtag tttccaacta 4380
taatgtgtaa attcttcact tctctttatt gaactataat gtgtaaattc ttcagtactc 4440
tttgttgaat caaagtgcat tgttcacttc aggaaatacc tcgagctttt gtttgtgcgg 4500
agagtaaggt atttgatgtg tctgaatggg ctacttgcca ggtgagtgtc tgacgatgtt 4560
ttatatgttt gatttaagtt gacatgtatg tgcatttgca gcagtgattt ttggatgtct 4620
caattgattt gatgtcatct ccatatgcat atttttatac tcggttctct gctgttttcg 4680
atgtcttaac tgactataga tatgcctttg gtcaattgac tttgttcagt tttgtatttg 4740
atgcatattc aaacgtccag attgactgtt ttactttaaa aattgtttca gttggtaaat 4800
gaaaatttgc ttacttcatt ggagatagga taattcatgc atgccatagc ccatagcctt 4860
atttttctgt gtcaagtttg tcatggctat aataaacaca acatattaat tgcacctgca 4920
tgtcatcccg actgatatct caattattga catacctatc taagagaaga gccaacaatg 4980
atgaaagtaa aggactaatt tggctgtgca aaattggacc ataagtttat tttacatttc 5040
atacttgctt cattcaacat aaacatcaaa atctggtacg ccaattttct ggcgatacat 5100
acagggtctt gtgataggtt catttgcata cattaaaaat gggagccttt ctaactctgt 5160
tttctttgct tgattgtcta gggcatggag tgcaaaccta acactcacgg cccatctttt 5220
atggtaaaca tggttggcgc agataggatg tctcagagat cctacagttc tcgctatccc 5280
tttagtttga atgctgagat gatccctgaa gatgaatttg agctatggct tcaacaagca 5340
ttggcatcag gtgtcttctc tgacagcccg aaacgcagga aaagctggag ccccttcaaa 5400
ctacctcaaa aagggataaa aagttggcgg cgatcctcat aagggcatag cattgaacag 5460
catggatctc acctgagtac aacactgaaa aaggctatac tctttgtgaa tgtaaataga 5520
ctgaccaaca atttgcctgg atgagcaact aattttgtcc aaaaagagac tgaaacaagg 5580
ggggtaaaag gaacaaacgc ttaagacatg actgcaataa atctgactgt tgaaattagt 5640
gttctctgca atgagatccc gcgagtttta tccgaaaagg tcagacactg ggatggtgtg 5700
tcattcatca gttcatccta agctcggaag ggtatctctg tagcatgtta acttcagtag 5760
ttttagggga tcggcatctg agagaatttc aaaacttcat acctggttgc cagcataagt 5820
tctgcaggtg ttgaaaagtt gttgatcaga gtagcaattt aaggtctgat gtttctgggg 5880
aacagtagga gagaaaaaaa tgacaaaaaa aagagagagt aggttgtaaa tacatgaaaa 5940
gttttcatca gaaattagtg ttgtaacatt gtacactgtg attacatcct gtgcaatact 6000
cccataattc agatctgtgt tgtaatacac tacatacatc ctacaatttt ctggtgataa 6060
tagagatcta attctcacct attatcgtta tttatggtta gtcagttact gctctgta 6118
<210> 21
<211> 1635
<212> DNA
<213> Oryza sativa
<400> 21
atggcggatt tggggctgtg gaagcaaggg tggaggtggg tggtgtccca gaagcacatc 60
ctgacatggg cgcacatggc ggcgagcggc ggcaccgaga ggctggcctt cctggtcgac 120
cggcattggc ccgccgtgtc ccgggcctgc gtgagctccg gccgcctcgc gctcgccgcg 180
ctgcggcaat ggcgcggctg cgcggcgcgc gggatcctgg agatggctag cctgggccct 240
gcgtccgtgt tcgtcatcct ctggagcttc ttcgtgtgca tcacctcgcc ggcgtgcgcc 300
ctctacgcgc tcctgggcat gggagctgct ggggcagtca ttcattacat gggctatacg 360
cctggtcttt tcattgtagg attatttgga atattgatta tgtggatgta tggctatttc 420
tggattacag gaatgcttct gattgctgga ggctgtatgt gctctttgaa acatgcacga 480
tttgtgatac ctgtgttggc tatgtatgct gtttattgtg tggctgttcg tgttggatcg 540
cttggtgtct tcttgacatt gaatctttct ttcctgacaa atgatcttct gaataagttg 600
ctgcaaggat acgagggaag cacagaagaa agacagtttg aagagccaaa acattctgat 660
cctgtcatgg atgagttcta tcgcagttgt gaatttccct ctgctcctga tagtgaacct 720
gagactgttt cttctgcaaa gcccttttgc tcaacacccg tccaggatgt gttgcatgta 780
cagaaagagg catctcctag caaagtagtg aaatcggatt ctgtttcatt ggatgagatg 840
aagaggatca tggatggttt gacccattat gaagttttgg gtattcctcg gaatagaagt 900
attgatcaaa agattctgaa aaaggagtac cacagaatgg tcctgcttgt acatcctgat 960
aaaaatatgg gaaatccact ggcctgtgaa tcattcaaaa agcttcagtc agcttatgag 1020
gtactctcag atttcacaaa gaaaaacact tacgacgacc aactgaggaa agaagaatca 1080
cgtaaaatga ctcagagatc acgtgttgtc tctcaacaga ctggggtaga gtttctctcc 1140
gaagagtcca ggcgtataca gtgcacaaaa tgtggtaatt ttcatctgtg gatatgtacc 1200
aagaaaagca aagcaaaagc aagatggtgt caggattgct ctgattttca tccagctaag 1260
gatggagatg gatgggtgga aaataaattt tcgtcatcct tcaaggaaat acctcgagct 1320
tttgtttgtg cggagagtaa ggtatttgat gtgtctgaat gggctacttg ccagggcatg 1380
gagtgcaaac ctaacactca cggcccatct tttatggtaa acatggttgg cgcagatagg 1440
atgtctcaga gatcctacag ttctcgctat ccctttagtt tgaatgctga gatgatccct 1500
gaagatgaat ttgagctatg gcttcaacaa gcattggcat caggtgtctt ctctgacagc 1560
ccgaaacgca ggaaaagctg gagccccttc aaactacctc aaaaagggat aaaaagttgg 1620
cggcgatcct cataa 1635
<210> 22
<211> 41
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 22
cacgtggacc actagtatgt ggatgtatgg ctatttctgg a 41
<210> 23
<211> 40
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 23
gtccgtacca actagttcgt atccttgcag caacttattc 40
<210> 24
<211> 41
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 24
tgaattcgct ggatcctcgt atccttgcag caacttattc a 41
<210> 25
<211> 41
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 25
gtcgactgga ggatccatgt ggatgtatgg ctatttctgg a 41
<210> 26
<211> 2087
<212> DNA
<213> Oryza sativa
<400> 26
ggaaaggaag aaaaggctaa tatgctcatc ttttttcata gattatactc catatcagta 60
gtatattgtg tcataaagaa taaaagagat atcagactgc cccccctctc tctctcccct 120
ttctcttgca cacaatatca tgatcacact atatttttag tcataggaag agatatttga 180
gaattttgac agcttcctga tcttaggttt ctttattgaa ctgatcttct tttatcagtg 240
ggataaaatg ttgccctata gctatattta caaggacaaa ccaaatgttt tagtatatac 300
cagaatcaaa catgcaacaa ttaattaagt attatagaac taaaaccact ttgttaaaag 360
caaggtctaa attatctgga gaaagtaaga agcaacatgt gatatattat aatattgtct 420
agtttttgta ctaaggtgtg tgttgcaatt gatgcaagtg gggtgtagca taatccatac 480
aagtaagatg ccaagaatgg ggaggagaga ctgtgattat ggcaggaaca tgctcttaat 540
cagtatacag aagtactact actaactact tgcaattact ccaatctctc tcttttctca 600
ttaactgcaa tgcataatcc gtactatccc gtgcaagtaa ctcaaaactt aaggcctcgg 660
ttagggctac taaatgaact atctgcaaat cccgttgttc tcccgttgac aatcatatac 720
ttagcatatt actcattgct tgtttgttag cttatcaagc acatcaaaaa aataaaattt 780
ttaaacttag ttttaagtta tcttgaatca tcgtttattg tcaatattat ctttttgaac 840
cgtcaataaa aaatataaaa aattatctat gaactttcct ttttctgctt cattcttttt 900
tatggcttat cagccatagt tcaaacgatc caccgtagct caaatatcct actactaatt 960
atttttcagc taaaaaagtt agcttccatt ttccaacctt acaatcaagc taacacagtc 1020
actgtcatat aaatagtata ctcaccctaa tcaagctaaa tcttttattt tcctaatgac 1080
tgaactccga aataatatta aattagaaat ctaatgatct agaagatgaa aaccacctct 1140
tttctaatca agctctcttt tgtaaccacc caccaccaca gccatcaaca ccaccaacag 1200
tccaacactg cccaattgcc catgctccag acttctttct actcctacat tccacatatc 1260
tccatggaca gtaactcctc ccaagctacc acttcaaccc taatcccctc tctctctctt 1320
ccgcagaggt agagtgagag agatggtcag atagctagat tgatatccct ctctctctct 1380
cacacacatc tctttttgca agatctcttc ttgttcatca tcttcttctt tttttctccc 1440
ccttttgctt caccaatcca tcttttgtca cgagatgtgg ccgagctgaa gctagtagta 1500
gtggagcagc gaaagcaagt acgccaagaa aaaaaaaagg aagaagaaag aagaaagaaa 1560
gaaagaaaaa aacgccagct tgagggcaga gggcaaaagc ggcgacgagg agcagtggcc 1620
aaagctcaga ttcttcccgt gggctatttt taccacccgc atcccctctc tttgagcccc 1680
ttggccgatt cattcaccga cgcaaagatc caacccctct tcaggtgtcg gcagatgccg 1740
cctttgtgag gtttccagtg gggggatttc tcgtcgtttc ttgcgtgcgg ttgcgttctt 1800
gatccagtga gcgcacggat atatccgccc tggtttagta gagagagaga gagagagaga 1860
gagagagaga gagagaggtt cttgattgag ttccaagtgt tggattgggt tcttggagct 1920
gttggattgg gtttttttgg gagagagatg ggggtttgga ggtgtgtggg ttgaccgaat 1980
tggatcaaga ttattgcggg aggggggggg gggggttgca atggcggatt tggggctgtg 2040
gaagcaaggg tggaggtggg tggtgtccca gaagcacatc ctgacat 2087
<210> 27
<211> 20
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 27
tttgacagct tcctgatctt 20
<210> 28
<211> 20
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 28
caagtaagat gccaagaatg 20
<210> 29
<211> 19
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 29
ttgtcaacgg gagaacaac 19
<210> 30
<211> 19
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 30
taggatattt gagctacgg 19
<210> 31
<211> 19
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 31
tagaaagaag tctggagca 19
<210> 32
<211> 19
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 32
aacgccagct tgagggcag 19
<210> 33
<211> 20
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 33
ttctcgtcgt ttcttgcgtg 20
<210> 34
<211> 19
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 34
gtgtgtgggt tgaccgaat 19
<210> 35
<211> 23
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 35
ggcattgtca acgggagaac aac 23
<210> 36
<211> 23
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 36
aaacgttgtt ctcccgttga caa 23
<210> 37
<211> 23
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 37
ggcaaacgcc agcttgaggg cag 23
<210> 38
<211> 23
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 38
aaacctgccc tcaagctggc gtt 23
<210> 39
<211> 24
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 39
gccgtttgac agcttcctga tctt 24
<210> 40
<211> 24
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 40
aaacaagatc aggaagctgt caaa 24
<210> 41
<211> 24
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 41
gccgcaagta agatgccaag aatg 24
<210> 42
<211> 23
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 42
aaaccattct tggcatctta ctt 23
<210> 43
<211> 23
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 43
gttgtaggat atttgagcta cgg 23
<210> 44
<211> 23
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 44
aaacccgtag ctcaaatatc cta 23
<210> 45
<211> 23
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 45
gttgtagaaa gaagtctgga gca 23
<210> 46
<211> 23
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 46
aaactgctcc agacttcttt cta 23
<210> 47
<211> 24
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 47
tcagttctcg tcgtttcttg cgtg 24
<210> 48
<211> 24
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 48
aaaccacgca agaaacgacg agaa 24
<210> 49
<211> 23
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 49
tcaggtgtgt gggttgaccg aat 23
<210> 50
<211> 23
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 50
aaacattcgg tcaacccaca cac 23
<210> 51
<211> 30
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 51
ggaaaggaag aaaaggctaa tatgctcatc 30
<210> 52
<211> 25
<212> DNA
<213> 人工合成(Aritificial synthesized)
<400> 52
atgtcaggat gtgcttctgg gacac 25
<210> 53
<211> 1344
<212> DNA
<213> Oryza sativa
<400> 53
ggaaaggaag aaaaggctaa tatgctcatc ttttttcata gattatactc catatcagta 60
gtatattgtg tcataaagaa taaaagagat atcagactgc cccccctctc tctctcccct 120
ttctcttgca cacaatatca tgatcacact atatttttag tcataggaag agatatttga 180
gaattttgac agcttcctga tggtttcttt attgaactga tcttctttta tcagtgggat 240
aaaatgttgc cctatagcta tatttacaag gacaaaccaa atgttttagt atataccaga 300
atcaaacatg caacaattaa ttaagtatta tagaactaaa accactttgt taaaagcaag 360
gtctaaatta tctggagaaa gtaagaagca acatgtgata tattataata ttgtctagtt 420
tttgtactaa ggtgtgtgtt gcaattgatg caagtggggt gtagcataat ccatacaagt 480
aagatgccaa gatccagact tctttctact cctacattcc acatatctcc atggacagta 540
actcctccca agctaccact tcaaccctaa tcccctctct ctctcttccg cagaggtaga 600
gtgagagaga tggtcagata gctagattga tatccctctc tctctctcac acacatctct 660
ttttgcaaga tctcttcttg ttcatcatct tcttcttttt ttctccccct tttgcttcac 720
caatccatct tttgtcacga gatgtggccg agctgaagct agtagtagtg gagcagcgaa 780
agcaagtacg ccaagaaaaa aaaaggaaga agaaagaaga aagaaagaaa gaaaaaaacg 840
ccagcttgag ggacagaggg caaaagcggc gacgaggagc agtggccaaa gctcagatcc 900
ttcccgtggg ctatttttac cacccgcatc ccctctcttt gagccccttg gccgattcat 960
tcaccgacgc aaagatccaa cccctcttca ggtgtcggca gatgccgcct ttgtgaggtt 1020
tccagtgggg ggatttctcg tcgtttcttg cgtgcggttg cgttcttgat ccagtgagcg 1080
cacggatata tccgccctgg tttagtagag agagagagag agagagagag agagaggttc 1140
ttgattgagt tccaagtgtt ggattgggtt cttggagctg ttggattggg tttttttggg 1200
agagagatgg gggtttggag gtgtgtgggt tgaccaattg gatcaagatt attgcgggag 1260
ggggggggga ggttgcaatg gcggatttgg ggctgtggaa gcaagggtgg aggtgggtgg 1320
tgtcccagaa gcacatcctg acat 1344
<210> 54
<211> 981
<212> DNA
<213> Oryza sativa
<400> 54
ggaaaggaag aaaaggctaa tatgctcatc ttttttcata gattatactc catatcagta 60
gtatattgtg tcataaagaa taaaagagat atcagactgc cccccctctc tctctcccct 120
ttctcttgca cacaatatca tgatcacact atatttttag tcataggaag agatatttga 180
gaattttgac agcttcctga tattgtgtga tcatgatatc aggtttcttt attgaactga 240
tcttctttta tcagtgggat aaaatgttgc cctatagcta tatttacaag gacaaaccaa 300
atgttttagt atataccaga atcaaacatg caacaattaa ttaagtatta tagaactaaa 360
accactttgt taaaagcaag gtctaaatta tctggagaaa gtaagaagca acatgtgata 420
tattataata ttgtctagtt tttgtactaa ggtgtgtgtt gcaattgatg caagtggggt 480
gtagcataat ccatacaagt aagatgccaa gaccctcaag ctggcgtttt tttctttctt 540
tctttcttct ttcttcttcc tttttttttc ttggcgtact tgctttcgct gctccactac 600
tactagcttc agctcggcca catctcgtga caaaagatgg attggtgaag caaaaggggg 660
agaaaaaaag aagaagatga tgaacaagaa gagatcttgc aaaaagagat gtgtgtgaga 720
gagagagagg gatatcaatc tagctatctg accatctctc tcactctacc tctgcggaag 780
agagagagag gggattaggg ttgaagtggt agcttgggag gagttactgt ccatggagat 840
atgtggaatg taggagtaga aagaagtctg gagaattgga tcaagattat tgcgggaggg 900
gggggggggt tgcaatggcg gatttggggc tgtggaagca agggtggagg tgggtggtgt 960
cccagaagca catcctgaca t 981
<210> 55
<211> 528
<212> PRT
<213> Brachypodium_distachyon
<400> 55
Met Asp Ile Met Thr Trp Ala His Met Ala Ala Gly Cys Gly Arg Glu
1 5 10 15
Arg Val Ala Ser Leu Val Asp Arg His Trp Pro Ala Val Ser Arg Ala
20 25 30
Cys Val Cys Ser Ser Cys Phe Val Leu Ala Ala Leu Arg Gln Trp Gln
35 40 45
Gly Cys Thr Ala Arg Gly Phe Leu Gly Leu Ala Ser Leu Gly Pro Ala
50 55 60
Ala Val Phe Val Ile Leu Trp Ser Phe Phe Val Cys Met Thr Ser Pro
65 70 75 80
Val Cys Ala Leu Tyr Ala Leu Leu Ile Leu Gly Ala Thr Gly Ala Val
85 90 95
Ile His Tyr Met Gly Tyr Thr Pro Gly Leu Leu Ile Val Gly Leu Phe
100 105 110
Gly Ile Leu Ile Met Trp Met Tyr Gly Tyr Phe Trp Ile Thr Gly Met
115 120 125
Leu Leu Val Ala Gly Gly Ser Met Cys Ser Leu Lys His Ala Arg Phe
130 135 140
Val Ile Pro Val Leu Ala Val Tyr Ala Val Tyr Cys Val Ala Val Arg
145 150 155 160
Val Gly Trp Leu Gly Val Phe Leu Thr Leu Asn Leu Ser Phe Leu Thr
165 170 175
Asn Asp Leu Leu Asn Lys Leu Leu Gln Gly Tyr Glu Gly Ser Thr Glu
180 185 190
Glu Met Glu Phe Glu Glu Met Lys Asp Pro His Pro Gly Met Asp Glu
195 200 205
Phe Tyr Pro Ser Tyr Glu Tyr Pro Pro Ala Pro Asp Ser Glu Pro Glu
210 215 220
Thr Val Ser Ser Ala Lys Pro Phe Cys Ala Ser Pro Thr Gln Asp Val
225 230 235 240
Leu His Val Gln Lys Glu Ala Ser Pro Ser Lys Ile Val Lys Ser Asp
245 250 255
Ser Thr Ala Leu Asp Glu Met Lys Arg Ile Met Asp Gly Ser Thr Tyr
260 265 270
Tyr Glu Ile Phe Gly Ile Pro Arg Asn Arg Ser Ala Asp Leu Lys Ile
275 280 285
Leu Lys Gly Glu Tyr Arg Arg Met Ala Met Leu Val His Pro Asp Lys
290 295 300
Asn Met Gly Asn Ser Leu Ala Cys Glu Ser Phe Lys Lys Leu Gln Ser
305 310 315 320
Ala Tyr Glu Val Leu Ser Asp Leu Thr Lys Lys Asn Ser Tyr Asp Glu
325 330 335
Gln Leu Arg Lys Glu Glu Ser Arg Gln Met Thr Gln Arg Ser Arg Val
340 345 350
Val Ser Gln Gln Ser Gly Val Glu Phe Leu Ser Glu Glu Ser Arg Arg
355 360 365
Ile Gln Cys Thr Lys Cys Gly Asn Phe His Leu Trp Ile Cys Thr Lys
370 375 380
Arg Ser Lys Ala Lys Ala Arg Trp Cys Gln Asp Cys Ser Gln His His
385 390 395 400
Val Ala Lys Asp Gly Asp Gly Trp Val Glu Asn Gly Tyr Ser Thr Ser
405 410 415
Leu Lys Ile Glu Ile Pro Arg Ala Phe Val Cys Ala Glu Ser Lys Ile
420 425 430
Phe Asp Val Ser Glu Trp Ala Thr Cys Gln Gly Met Glu Cys Lys Pro
435 440 445
Asn Thr His Gly Pro Thr Phe Met Val Asn Met Val Gly Ala Asp Arg
450 455 460
Met Pro Gln Arg Ser Tyr Ser Ser Arg Tyr Pro Phe Ser Leu Asp Ala
465 470 475 480
Glu Met Ile Pro Glu Asp Glu Phe Asp Leu Trp Leu Gln Gln Ala Leu
485 490 495
Ala Thr Gly Val Phe Ser Asp Ser Pro Lys Arg Arg Lys Ser Trp Ser
500 505 510
Pro Phe Lys Leu Thr Gln Lys Gly Val Arg Ser Trp Arg Arg Ser Ser
515 520 525
<210> 56
<211> 545
<212> PRT
<213> Hordeum_vulgare
<400> 56
Met Ala Gly Leu Gly Leu Trp Asn Gln Gly Trp Thr Trp Val Leu Ser
1 5 10 15
Gln Lys His Val Val Ala Trp Ala His Ala Ala Ala Gly Cys Gly Arg
20 25 30
Asp Arg Leu Ala Phe Leu Val Asp Arg His Trp Pro Ala Val Ser Arg
35 40 45
Ala Cys Ala Thr Ser Ser Arg Leu Val Leu Glu Ala Leu Arg Gln Trp
50 55 60
Arg Gly Cys Thr Ala Arg Gly Leu Leu Ala Leu Ala Ser Leu Gly Pro
65 70 75 80
Ala Ala Val Phe Val Ile Leu Trp Ser Cys Phe Val Cys Met Thr Ser
85 90 95
Ser Ala Cys Ala Leu Tyr Ala Leu Leu Ala Leu Gly Ala Val Gly Ala
100 105 110
Val Ile His Tyr Met Gly Tyr Thr Pro Gly Leu Leu Ile Val Gly Leu
115 120 125
Phe Gly Ile Met Ile Met Trp Met Tyr Gly Tyr Phe Trp Ile Thr Gly
130 135 140
Met Leu Leu Val Ala Gly Gly Cys Met Cys Ser Leu Lys His Ala Arg
145 150 155 160
Phe Val Ile Pro Val Leu Ala Met Tyr Ala Val Tyr Cys Val Ala Val
165 170 175
Arg Val Gly Trp Leu Gly Val Phe Phe Met Leu Asn Leu Ser Phe Leu
180 185 190
Thr Asn Asp Leu Leu Asn Lys Leu Leu Gln Gly Tyr Glu Gly Ser Thr
195 200 205
Glu Glu Arg Pro Phe Glu Glu Met Lys Asp Ser Asp Pro Ala Thr Asp
210 215 220
Ala Phe Phe Arg Gly Cys Glu Tyr Pro Pro Ala Pro Glu Ser Glu Pro
225 230 235 240
Glu Thr Val Ser Ser Ala Lys Pro Phe Cys Ala Ala Pro Thr Gln Asp
245 250 255
Val Leu His Val Gln Lys Glu Pro Ser Pro Thr Lys Ile Val Lys Ser
260 265 270
Asn Ser Thr Ser Leu Asp Glu Met Lys Arg Ile Met Asp Gly Ser Thr
275 280 285
Tyr Tyr Glu Val Leu Gly Ile Pro Arg Ser Lys Ser Ile Asn Gln Ile
290 295 300
Glu Leu Lys Lys Glu Tyr Arg Lys Leu Ala Val Leu Val His Pro Asp
305 310 315 320
Lys Asn Met Gly Asn Pro Leu Ala Cys Glu Ser Phe Lys Lys Leu Gln
325 330 335
Ser Ala Phe Glu Val Leu Ser Asp Leu Thr Lys Lys Asn Gly Tyr Asp
340 345 350
Glu Gln Leu Arg Lys Glu Glu Ser Arg Gln Met Thr Gln Arg Ser Arg
355 360 365
Val Val Ser Gln Pro Ser Gly Val Glu Phe Leu Ser Glu Glu Ser Arg
370 375 380
Arg Ile Gln Cys Thr Lys Cys Gly Asn Phe His Leu Trp Ile Cys Thr
385 390 395 400
Lys Arg Ser Lys Ala Lys Ala Arg Trp Cys Gln Glu Cys Ser Gln Tyr
405 410 415
His Val Ala Lys Asp Gly Asp Gly Trp Val Glu Asn Arg Tyr Ser Thr
420 425 430
Ser Leu Lys Ile Glu Ile Pro Arg Ala Phe Val Cys Ala Glu Ser Lys
435 440 445
Ile Phe Asp Val Ser Glu Trp Ala Thr Cys Gln Gly Met Glu Cys Lys
450 455 460
Pro Asn Thr His Gly Pro Thr Phe Met Val Asn Met Val Gly Ala Asp
465 470 475 480
Arg Met Pro Gln Arg Ser His Ser Ser Arg Tyr Pro Phe Ser Leu Asp
485 490 495
Ala Glu Met Ile Pro Glu Asp Glu Phe Glu Leu Trp Leu Gln Gln Ala
500 505 510
Leu Ala Thr Gly Val Phe Ser Asp Ser Pro Lys Arg Arg Lys Ser Trp
515 520 525
Ser Pro Phe Lys Leu Pro Gln Lys Gly Ile Arg Ser Trp Arg Arg Ser
530 535 540
Ser
545
<210> 57
<211> 545
<212> PRT
<213> Oryza_brachyantha
<400> 57
Met Ala Asp Leu Gly Leu Trp Lys Gln Gly Trp Arg Trp Val Leu Ser
1 5 10 15
Gln Lys His Ile Leu Thr Trp Ala His Met Ala Ala Ser Gly Gly Thr
20 25 30
Glu Arg Leu Ala Phe Leu Val Asp Arg His Trp Pro Ala Val Ser Arg
35 40 45
Thr Cys Val Ser Ser Gly Arg Leu Ala Leu Ala Ala Leu Arg Gln Trp
50 55 60
Arg Gly Cys Ala Ala Arg Gly Ile Leu Glu Met Ala Ser Leu Gly Pro
65 70 75 80
Ala Ser Val Phe Val Ile Leu Trp Ser Cys Phe Val Cys Met Thr Ser
85 90 95
Pro Ala Cys Ala Leu Tyr Ala Leu Leu Ser Leu Gly Ala Ala Gly Ala
100 105 110
Val Ile His Tyr Met Gly Tyr Thr Pro Gly Leu Phe Ile Val Gly Leu
115 120 125
Phe Gly Ile Leu Ile Met Trp Met Tyr Gly Tyr Phe Trp Ile Thr Gly
130 135 140
Met Leu Leu Ile Ser Gly Gly Cys Met Cys Ser Leu Lys His Ala Arg
145 150 155 160
Phe Val Ile Pro Val Leu Ala Met Tyr Ala Val Tyr Cys Val Ala Val
165 170 175
Arg Val Gly Leu Leu Gly Val Phe Leu Thr Leu Asn Leu Ser Phe Leu
180 185 190
Thr Asn Asp Leu Met Asn Lys Leu Leu Gln Gly Tyr Glu Gly Ser Thr
195 200 205
Glu Glu Arg Gln Phe Glu Glu Thr Lys His Ser Asp Pro Val Met Asp
210 215 220
Glu Phe Tyr Arg Ser Cys Glu Tyr Pro Thr Ala Pro Asp Ser Glu Pro
225 230 235 240
Glu Thr Val Ser Ser Ala Lys Pro Phe Cys Ser Thr Pro Val Gln Asp
245 250 255
Val Leu His Val Gln Lys Glu Ala Ser Pro Ser Lys Val Val Lys Ser
260 265 270
Asp Ser Val Ser Leu Asp Glu Met Lys Arg Ile Met Asp Gly Leu Thr
275 280 285
His Tyr Glu Val Leu Gly Ile Pro Arg Asn Arg Ser Ile Asp Gln Lys
290 295 300
Thr Leu Lys Lys Glu Tyr His Arg Met Val Leu Leu Val His Pro Asp
305 310 315 320
Lys Asn Met Gly Asn Pro Leu Ala Cys Glu Ser Phe Lys Lys Leu Gln
325 330 335
Ser Ala Tyr Glu Val Leu Ser Asp Phe Thr Lys Lys Asn Thr Tyr Asp
340 345 350
Asp Gln Leu Arg Lys Glu Glu Ser Arg Lys Met Thr Gln Arg Ser Arg
355 360 365
Val Val Ser Gln Gln Thr Gly Val Glu Phe Leu Ser Glu Glu Ser Arg
370 375 380
Arg Ile Gln Cys Thr Lys Cys Gly Asn Phe His Leu Trp Ile Cys Thr
385 390 395 400
Lys Lys Ser Lys Ala Lys Ala Arg Trp Cys Gln Asp Cys Ser Asp Phe
405 410 415
His Pro Ala Lys Asp Gly Asp Gly Trp Val Glu Asn Lys Phe Ser Ala
420 425 430
Ser Phe Lys Met Glu Ile Pro Arg Ala Phe Val Cys Ala Glu Ser Lys
435 440 445
Ile Phe Asp Val Ser Glu Trp Ala Thr Cys Gln Gly Met Glu Cys Lys
450 455 460
Pro Asn Thr His Gly Pro Ser Phe Met Val Asn Met Val Gly Ala Asp
465 470 475 480
Arg Met Ser Gln Arg Ser Tyr Ser Ser Arg Tyr Pro Phe Ser Leu Asn
485 490 495
Ala Glu Met Val Pro Glu Asp Glu Phe Glu Leu Trp Leu Gln Gln Ala
500 505 510
Leu Ala Ser Gly Val Phe Ala Asp Ser Pro Lys Arg Arg Lys Ser Trp
515 520 525
Ser Pro Phe Lys Leu Pro Gln Lys Gly Ile Lys Ser Trp Arg Arg Ser
530 535 540
Ser
545
<210> 58
<211> 545
<212> PRT
<213> Panicum_hallii
<400> 58
Met Ala Asp Leu Gly Leu Trp Lys Gln Ala Trp Arg Trp Val Leu Ser
1 5 10 15
Gln Lys His Ile Leu Ala Trp Ala His Thr Ala Ala Cys Gly Ser Arg
20 25 30
Glu Arg Leu Ala Phe Leu Val Asp Arg His Trp Pro Ala Val Ser Arg
35 40 45
Ala Cys Ala Thr Ser Ser Arg Leu Ala Leu Ala Ala Leu Arg Gln Trp
50 55 60
Arg Gly Cys Met Ala Arg Gly Val Leu Ala Val Ala Ser Leu Gly Pro
65 70 75 80
Ala Ala Val Phe Val Ile Leu Trp Ser Phe Phe Val Cys Met Thr Ser
85 90 95
Pro Ala Trp Ala Leu Phe Ala Leu Leu Ser Leu Gly Ala Ala Gly Ala
100 105 110
Val Val His Tyr Met Gly Tyr Thr Pro Gly Leu Phe Ile Val Gly Leu
115 120 125
Phe Gly Ile Leu Ile Met Trp Met Tyr Gly Tyr Phe Trp Ile Thr Gly
130 135 140
Met Leu Leu Val Ala Gly Gly Cys Met Cys Ser Leu Lys His Ala Arg
145 150 155 160
Tyr Val Ile Pro Ile Leu Thr Thr Tyr Ala Ile Tyr Cys Val Ala Val
165 170 175
Arg Val Gly Trp Leu Gly Val Phe Leu Thr Leu Asn Leu Ser Phe Leu
180 185 190
Thr Asn Asp Leu Leu Asn Lys Leu Leu Gln Gly Tyr Glu Gly Cys Thr
195 200 205
Glu Glu Glu Gln Phe Glu Asp Met Lys Asp Ser Asp Pro Val Met Asp
210 215 220
Glu Phe Tyr Arg Ser Cys Glu Phe Pro Pro Thr Pro Asp Ser Glu Pro
225 230 235 240
Glu Thr Val Ser Ser Ala Lys Pro Tyr Cys Ser Ala Pro Thr Gln Asp
245 250 255
Val Leu His Val Gln Lys Glu Glu Pro Pro Ser Lys Val Val Lys Ser
260 265 270
Asp Ser Ser Ser Leu Asp Glu Ile Lys Arg Ile Met Asp Gly Ser Asn
275 280 285
Tyr Tyr Glu Val Leu Gly Ile Pro Arg Asn Arg Ser Ile Asp Gln Lys
290 295 300
Ser Leu Lys Lys Glu Tyr His Arg Met Val Leu Leu Val His Pro Asp
305 310 315 320
Lys Asn Met Gly Asn Pro Leu Ala Cys Glu Ser Phe Lys Lys Leu Gln
325 330 335
Ser Ala Tyr Glu Val Leu Ser Asp Phe Thr Lys Lys Asn Ser Tyr Asp
340 345 350
Glu Gln Leu Arg Lys Glu Glu Ser Gln Asn Met Thr Pro Arg Ser Arg
355 360 365
Val Val Ser Gln Gln Ser Gly Val Glu Phe Leu Ser Glu Glu Ser Arg
370 375 380
Arg Ile Gln Cys Thr Lys Cys Gly Asn Phe His Ile Trp Ile Cys Thr
385 390 395 400
Lys Arg Ser Lys Thr Lys Ala Arg Phe Cys Gln Gly Cys Asp Gln Phe
405 410 415
His Gln Ala Lys Asp Gly Asp Gly Trp Val Glu Thr Arg Phe Ser Thr
420 425 430
Ser Val Lys Met Glu Ile Pro Arg Ala Phe Val Cys Ala Glu Ser Lys
435 440 445
Ile Phe Asp Val Ser Glu Trp Ala Thr Cys Gln Gly Met Glu Cys Lys
450 455 460
Pro Asn Thr His Gly Pro Thr Phe Met Val Asn Met Val Gly Ala Asp
465 470 475 480
Arg Met Pro Gln Arg Ser Tyr Ser Ser Arg Tyr Pro Phe Ser Leu Asp
485 490 495
Ala Glu Met Ile Pro Glu Asp Glu Phe Glu Leu Trp Leu Gln Gln Ala
500 505 510
Leu Ala Ser Gly Val Phe Ala Asp Ser Pro Lys Arg Arg Lys Ser Trp
515 520 525
Ser Pro Phe Lys Leu Pro Gln Lys Gly Ile Lys Ser Trp Arg Arg Ser
530 535 540
Ser
545
<210> 59
<211> 544
<212> PRT
<213> Setaria_italica
<400> 59
Met Ala Asp Leu Gly Leu Trp Lys Gln Ala Trp Arg Trp Val Leu Ser
1 5 10 15
Gln Lys His Ile Leu Ala Trp Ala His Thr Ala Ala Cys Gly Ser Arg
20 25 30
Glu Arg Leu Ala Phe Leu Val Asp Arg His Trp Pro Ala Val Ser Arg
35 40 45
Ala Cys Ala Thr Ser Ser Arg Leu Ala Leu Ala Ala Leu Leu Gln Trp
50 55 60
Arg Gly Cys Met Ala Arg Gly Val Leu Ala Val Ala Ser Leu Gly Pro
65 70 75 80
Ala Ala Val Phe Val Ile Leu Trp Ser Phe Phe Val Cys Met Thr Ser
85 90 95
Pro Ala Trp Ala Leu Phe Ala Leu Leu Leu Leu Gly Ala Ala Gly Ala
100 105 110
Val Val His Tyr Met Gly Tyr Thr Pro Gly Leu Phe Ile Val Gly Leu
115 120 125
Phe Gly Ile Leu Ile Met Trp Met Tyr Gly Tyr Phe Trp Ile Thr Gly
130 135 140
Met Leu Leu Val Ala Gly Gly Cys Met Cys Ser Leu Lys His Ala Arg
145 150 155 160
Tyr Val Ile Pro Ile Leu Thr Thr Tyr Ala Ile Tyr Cys Val Ala Ile
165 170 175
Arg Val Gly Trp Leu Gly Val Phe Leu Thr Leu Asn Leu Ser Phe Leu
180 185 190
Ala Asn Asp Leu Leu Asn Lys Leu Leu Gln Gly Tyr Glu Glu Ser Thr
195 200 205
Glu Glu Lys Phe Glu Asp Met Lys Asp Ser Asp Pro Val Met Asp Glu
210 215 220
Phe Tyr Arg Ser Cys Glu Phe Pro Pro Ala Pro Asp Ser Glu Pro Glu
225 230 235 240
Thr Val Ser Ser Ala Lys Pro Tyr Cys Ser Ser Pro Thr Gln Asp Val
245 250 255
Leu His Val Gln Lys Glu Glu Pro Pro Ser Lys Val Val Lys Ser Asp
260 265 270
Ser Ser Ser Leu Asp Glu Ile Lys Arg Ile Met Asp Gly Ser Asn His
275 280 285
Tyr Glu Val Leu Gly Ile Pro Arg Asn Arg Ser Ile Asp Gln Lys Ser
290 295 300
Leu Lys Lys Glu Tyr His Arg Met Val Leu Leu Val His Pro Asp Lys
305 310 315 320
Asn Met Gly Asn Pro Leu Ala Cys Glu Ser Phe Lys Lys Leu Gln Ser
325 330 335
Ala Tyr Glu Val Leu Ser Asp Phe Thr Lys Arg Asn Ser Tyr Asp Glu
340 345 350
Gln Leu Arg Lys Glu Glu Ser Gln Lys Met Thr Pro Arg Ser Arg Val
355 360 365
Val Ser Gln Gln Gly Gly Val Glu Phe Leu Ser Glu Glu Ser Arg Arg
370 375 380
Ile Gln Cys Thr Lys Cys Gly Asn Phe His Ile Trp Ile Cys Thr Lys
385 390 395 400
Arg Ser Lys Thr Lys Ala Arg Phe Cys Gln Gly Cys Asp Gln Tyr His
405 410 415
Gln Ala Lys Asp Gly Asp Gly Trp Val Glu Thr Arg Phe Ser Thr Ser
420 425 430
Tyr Lys Met Glu Ile Pro Arg Ala Phe Val Cys Ala Glu Ser Lys Ile
435 440 445
Phe Asp Val Ser Glu Trp Ala Thr Cys Gln Gly Met Glu Cys Lys Pro
450 455 460
Asn Thr His Gly Pro Thr Phe Met Val Asn Met Val Gly Ala Asp Arg
465 470 475 480
Met Pro Gln Arg Ser Tyr Ser Ser Arg Tyr Pro Phe Ser Leu Asp Ala
485 490 495
Glu Met Ile Pro Glu Asp Glu Phe Glu Leu Trp Leu Gln Gln Ala Leu
500 505 510
Ala Ser Gly Val Phe Ala Asp Ser Pro Lys Arg Arg Lys Ser Trp Ser
515 520 525
Pro Phe Lys Leu Pro Gln Lys Gly Ile Lys Ser Trp Arg Arg Ser Ser
530 535 540
<210> 60
<211> 545
<212> PRT
<213> Zea_mays
<400> 60
Met Glu Asp Leu Gly Leu Trp Asn Gln Ala Trp Met Trp Val Leu Ser
1 5 10 15
Gln Lys His Ile Leu Ala Trp Ala His Thr Ala Ala Cys Gly Ser Arg
20 25 30
Glu Arg Leu Ala Phe Leu Val Asp Arg His Trp Pro Ala Val Ser Arg
35 40 45
Gly Cys Ala Thr Ser Ser Arg Leu Thr Leu Ala Ala Leu Arg Gln Trp
50 55 60
Arg Gly Cys Met Ala Arg Gly Val Leu Ala Val Ala Ser Leu Gly Pro
65 70 75 80
Ala Ala Val Phe Val Ile Leu Trp Ser Phe Phe Val Cys Met Thr Ser
85 90 95
Pro Ala Cys Ala Leu Tyr Ala Leu Leu Ser Leu Gly Ala Ala Ala Ala
100 105 110
Val Val His Tyr Met Gly Tyr Thr Pro Gly Leu Leu Ile Val Gly Leu
115 120 125
Phe Gly Ile Leu Ile Met Trp Met Tyr Gly Tyr Phe Trp Ile Thr Gly
130 135 140
Met Leu Leu Val Ala Gly Gly Cys Met Cys Ser Leu Lys His Ala Arg
145 150 155 160
Tyr Val Thr Pro Val Leu Thr Ser Tyr Ala Ile Tyr Cys Val Ala Val
165 170 175
Arg Val Gly Trp Leu Gly Val Phe Leu Thr Phe Asn Leu Ser Phe Leu
180 185 190
Thr Asn Asp Leu Leu Asn Lys Leu Ala Gln Gly Tyr Glu Gly Ser Thr
195 200 205
Glu Glu Ser Gln Phe Glu Asp Met Lys Asp Ser Asp Pro Val Met Asp
210 215 220
Glu Phe Tyr Arg Ser Cys Glu Phe Pro Ser Val Pro Asp Ser Glu Pro
225 230 235 240
Glu Thr Val Ser Ser Ala Lys Pro Tyr Cys Ser Ala Pro Ile Gln Asp
245 250 255
Val Leu His Val Gln Lys Glu Glu Pro Pro Ser Lys Ile Val Lys Ser
260 265 270
Asp Ser Ser Ser Ser Asp Glu Ile Lys Arg Ile Met Asp Gly Ser Asn
275 280 285
His Tyr Glu Val Leu Gly Val Pro Arg Asn Arg Ser Ile Asp Gln Lys
290 295 300
Ala Leu Lys Lys Glu Tyr His Arg Met Val Leu Leu Val His Pro Asp
305 310 315 320
Lys Asn Met Gly Asn Pro Leu Ala Cys Glu Ser Phe Lys Lys Leu Gln
325 330 335
Ser Ala Tyr Glu Val Leu Ser Asp Phe Thr Lys Lys Asn Ser Tyr Asp
340 345 350
Gln Gln Leu Arg Lys Glu Glu Ser Gln Lys Met Thr Pro Arg Ser Arg
355 360 365
Ala Val Ser Gln Gln Ser Gly Val Glu Phe Leu Ser Glu Glu Ser Arg
370 375 380
Arg Ile Gln Cys Thr Lys Cys Gly Asn Phe His Ile Trp Ile Cys Thr
385 390 395 400
Lys Arg Ser Lys Thr Lys Ala Arg Phe Cys Gln Gly Cys Asp Gln Phe
405 410 415
His Gln Ala Lys Asp Gly Asp Gly Trp Val Glu Thr Arg Phe Ser Ser
420 425 430
Ser Ile Lys Met Glu Ile Pro Arg Ala Phe Val Cys Ala Glu Ser Lys
435 440 445
Ile Phe Asp Val Ser Glu Trp Ala Thr Cys Gln Gly Met Glu Cys Lys
450 455 460
Pro Asn Thr His Gly Pro Thr Phe Met Val Asn Met Val Gly Ala Asp
465 470 475 480
Arg Met Pro Gln Arg Ser Tyr Ser Ser Arg Tyr Pro Phe Ser Leu Asp
485 490 495
Ala Glu Met Ile Pro Asp Asp Glu Phe Glu Met Trp Leu Gln Gln Ala
500 505 510
Leu Ala Ser Gly Val Phe Ala Asp Ser Pro Lys Arg Arg Lys Ser Trp
515 520 525
Ser Pro Phe Lys Leu Pro Gln Lys Gly Ile Lys Ser Trp Arg Arg Ser
530 535 540
Ser
545
<210> 61
<211> 545
<212> PRT
<213> Sorghum_bicolor
<400> 61
Met Ala Asp Leu Gly Leu Trp Lys Gln Ala Trp Met Trp Val Leu Ser
1 5 10 15
Gln Lys His Ile Leu Ala Trp Ala His Thr Ala Ala Cys Gly Ser Arg
20 25 30
Glu Arg Leu Ala Phe Leu Val Asp Arg His Trp Pro Ala Val Ser Arg
35 40 45
Ala Cys Ala Thr Ser Ser Arg Leu Ala Leu Ala Ala Leu Arg Gln Trp
50 55 60
Arg Gly Cys Thr Ala Arg Gly Val Leu Ala Val Ala Ser Leu Gly Pro
65 70 75 80
Ala Ala Val Phe Val Ile Leu Trp Ser Phe Phe Val Cys Met Thr Ser
85 90 95
Pro Ala Cys Ala Leu Tyr Ala Leu Leu Ser Leu Gly Ala Ala Ala Ala
100 105 110
Val Val His Tyr Met Gly Tyr Thr Pro Gly Leu Phe Ile Val Gly Leu
115 120 125
Phe Gly Ile Leu Ile Met Trp Met Tyr Gly Tyr Phe Trp Ile Thr Gly
130 135 140
Met Leu Leu Val Ala Gly Gly Cys Met Cys Ser Leu Lys His Ala Arg
145 150 155 160
Tyr Val Ile Pro Val Leu Thr Ser Tyr Ala Ile Tyr Ser Val Ala Val
165 170 175
Arg Val Gly Trp Leu Gly Val Phe Leu Thr Leu Asn Leu Ser Phe Leu
180 185 190
Thr Asn Asp Leu Leu Asn Lys Leu Ala Gln Gly Tyr Glu Gly Ser Thr
195 200 205
Glu Glu Ser Gln Phe Glu Asp Ile Lys Gly Ser Asp Pro Val Met Asp
210 215 220
Glu Phe Tyr Arg Ser Cys Glu Phe Pro Pro Val Pro Asp Ser Glu Pro
225 230 235 240
Glu Thr Val Ser Ser Ala Lys Pro Tyr Cys Thr Ala Pro Val Gln Asp
245 250 255
Val Leu His Val Gln Lys Glu Glu Pro Pro Ser Lys Val Val Lys Ser
260 265 270
Asp Ser Ser Ser Leu Asp Glu Ile Lys Arg Ile Met Asp Gly Ser Asn
275 280 285
His Tyr Glu Val Leu Gly Val Pro Arg Asn Arg Ser Ile Asp Gln Lys
290 295 300
Thr Leu Lys Lys Glu Tyr His Arg Met Val Leu Leu Val His Pro Asp
305 310 315 320
Lys Asn Met Gly Asn Pro Leu Ala Cys Glu Ser Phe Lys Lys Leu Gln
325 330 335
Ser Ala Tyr Glu Val Leu Ser Asp Phe Thr Lys Lys Asn Ser Tyr Asp
340 345 350
Glu Gln Leu Arg Lys Glu Glu Ser Leu Lys Met Thr Pro Arg Ser Arg
355 360 365
Val Val Ser Gln Gln Ser Gly Val Glu Phe Leu Ser Glu Glu Ser Arg
370 375 380
Arg Ile Gln Cys Thr Lys Cys Gly Asn Phe His Ile Trp Ile Cys Thr
385 390 395 400
Lys Arg Ser Lys Thr Arg Ala Arg Phe Cys Gln Gly Cys Asp Gln Phe
405 410 415
His Gln Ala Lys Asp Gly Asp Gly Trp Val Glu Thr Arg Phe Ser Ser
420 425 430
Ser Ile Lys Met Glu Ile Pro Arg Ala Phe Val Cys Ala Glu Ser Lys
435 440 445
Ile Phe Asp Val Ser Glu Trp Ala Thr Cys Gln Gly Met Glu Cys Lys
450 455 460
Pro Asn Thr His Gly Pro Thr Phe Met Val Asn Met Val Gly Thr Asp
465 470 475 480
Arg Met Pro Gln Arg Ser Tyr Ser Ser Arg Tyr Pro Phe Ser Leu Asp
485 490 495
Ala Glu Met Ile Pro Glu Asp Glu Phe Glu Leu Trp Leu Gln Gln Ala
500 505 510
Leu Ala Ser Gly Val Phe Ala Asp Ser Pro Lys Arg Arg Lys Ser Trp
515 520 525
Ser Pro Phe Lys Leu Pro Gln Lys Gly Ile Lys Ser Trp Arg Arg Ser
530 535 540
Ser
545
Claims (11)
1.一种调控植物种子大小的方法,通过影响调控植物种子大小的基因的表达调控植物的种子大小,其特征在于所述调控植物种子大小的基因的核苷酸序列选自下列组的序列之一:
(a)如调控植物种子大小的基因LOC_Os12g41820所示的核苷酸序列;
(b)如SEQ ID NO:1、2、20或21所示的核苷酸序列;
(c)其编码氨基酸序列如SEQ ID NO:3、55、56、57、58、59、60或61所示的核苷酸序列;
(d)在严谨条件下能够与(a)-(c)中所述序列的DNA杂交的DNA序列;或
(e)与(a)-(d)所述序列有至少80%(优选为至少85%)序列相似性,且具有调控植物种子大小功能的DNA序列;或
(f)与(a)-(e)之任一所述序列互补的DNA序列。
2.权利要求1所述的方法,其特征在于,通过突变调控植物种子大小的基因的核苷酸序列以使该基因表达的蛋白质失活,从而获得具有大粒性状表型的植物材料,其中所述调控植物种子大小的基因的核苷酸序列选自下列组的序列之一:
(a)如调控植物种子大小的基因LOC_Os12g41820所示的核苷酸序列;
(b)如SEQ ID NO:1、2、20或21所示的核苷酸序列;
(c)其编码氨基酸序列如SEQ ID NO:3、55、56、57、58、59、60或61所示的核苷酸序列;
(d)在严谨条件下能够与(a)-(c)中所述序列的DNA杂交的DNA序列;或
(e)与(a)-(d)所述序列有至少80%(优选为至少85%)序列相似性,且具有调控植物种子大小功能的DNA序列;或
(f)与(a)-(e)之任一所述序列互补的DNA序列。
3.权利要求2所述的方法,其中所述的突变包括在调控植物种子大小的基因的核苷酸序列上进行取代、缺失或添加一个或多个核苷酸。
4.权利要求3所述的方法,其中所述的突变通过物理诱变、化学诱变或RNAi、TALEN、CRISPR/Cas9等定点突变技术获得。
5.权利要求1-4之任一所述的方法在调控植物种子大小中的应用。
6.一种表达盒、表达载体和工程菌在调控植物种子大小中的应用,其特征在于所述表达盒、表达载体和工程菌包含调控植物种子大小的基因,所述调控植物种子大小的基因的核苷酸序列选自下列组的序列之一:
(a)如调控植物种子大小的基因LOC_Os12g41820所示的核苷酸序列;
(b)如SEQ ID NO:1、2、20或21所示的核苷酸序列;
(c)其编码氨基酸序列如SEQ ID NO:3、55、56、57、58、59、60或61所示的核苷酸序列;
(d)在严谨条件下能够与(a)-(c)中所述序列的DNA杂交的DNA序列;或
(e)与(a)-(d)所述序列有至少80%(优选为至少85%)序列相似性,且具有调控植物种子大小功能的DNA序列;或
(f)与(a)-(e)之任一所述序列互补的DNA序列。
7.一种突变体材料的获得方法,其特征在于所述突变体材料是由调控植物种子大小的基因的突变所造成,含有该突变后的核苷酸序列的植株具有大粒种子的表型,其中所述调控植物种子大小的基因的核苷酸序列选自下列组的序列之一:
(a)如调控植物种子大小的基因LOC_Os12g41820所示的核苷酸序列;
(b)如SEQ ID NO:1、2、20或21所示的核苷酸序列;
(c)其编码氨基酸序列如SEQ ID NO:3、55、56、57、58、59、60或61所示的核苷酸序列;
(d)在严谨条件下能够与(a)-(c)中所述序列的DNA杂交的DNA序列;或
(e)与(a)-(d)所述序列有至少80%(优选为至少85%)序列相似性,且具有调控植物种子大小功能的DNA序列;或
(f)与(a)-(e)之任一所述序列互补的DNA序列。
8.权利要求7所述的方法,其中所述的突变是点突变,或是DNA缺失或***突变,或是通过RNAi、定点突变等基因沉默手段产生。
9.权利要求8所述的方法,其中所述的突变后的核苷酸序列如SEQ ID NO:4所示,其编码的氨基酸序列如SEQ ID NO:5所示。
10.权利要求7-9之任一所述的方法、及其获得的突变体材料在育种中的应用。
11.权利要求10所述的应用,其中所述的育种是指利用调控植物种子大小的基因控制作物籽粒的大小或粒重,或作为鉴定作物大粒品种和小粒品种的分子标记。
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