CN101932710B - 增强对稻瘟病菌的抗性的基因以及该基因的应用 - Google Patents
增强对稻瘟病菌的抗性的基因以及该基因的应用 Download PDFInfo
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Abstract
本发明涉及能够增强对稻温病菌(Magnaporthe oryzae)的抗性的Pi5-1蛋白和Pi5-2蛋白、编码这些蛋白的基因、包括这些基因的重组体载体、用重组体载体转化的植物、植物的种子、通过在植物中表达所述基因以增强对植物病原菌的抗性的方法、抗所述蛋白的抗体、以及包括能够增强对植物病原菌的抗性的基因的组合物。
Description
技术领域
本发明涉及增强对稻瘟病菌(Magnaporthe oryzae)的抗性的基因以及该基因的应用。更具体地说,本发明涉及能够增强对稻瘟病菌的抗性的Pi5-1蛋白和Pi52蛋白、编码这些蛋白的基因、包括些基因的重组体载体、用该重组体载体转化的植物、植物的种子、通过在植物中表达所述基因以增强对植物病原菌的抗性的方法、抗所述蛋白的抗体、以及包括能够增强对植物病原菌的抗性的基因的组合物。
背景技术
天然免疫应答是植物和动物存活的关键。该应答由用病原体识别受体(PRRs;也叫模式识别受体或抗病性蛋白)对与病原体相关分子模式(PAMPs)(也指微生物相关分子模式)或无毒力蛋白(Avr)的检测所介导。在动物体中,细胞质内的PRRs家族介导凋亡和防护病原体入侵关键的炎症反应,所述细胞质内的PRRs家族包括结合核苷酸的寡聚结构域(NOD)。植物体也含有一系列的细胞内PRR蛋白,称为NB-LRR(结合核苷酸的-富含亮氨酸的重复单位)R蛋白,它们与动物的NOD蛋白结构相似。这些植物NB-LRR蛋白的特征是:由N-端卷曲螺旋(CC)或Toll/白介素-1受体(TIR)结构域、中心NB结构域以及C-端LRR结构域(Hammond-Kosack和Jones(1997)Annu.Rev.Plant Physiol.Plant Mol.Biol.48:575-607)组成的三结构域结构,以及典型的识别病原体来源的Avr蛋白(也叫效应器)(Van der Biezen和Jones(1998)Trends Biochem.Sci.23:454-456)。
稻瘟病是水稻最具破坏性的疾病之一,在种植水稻的世界各地均有发生。到目前为止已鉴定了超过70个的对不同地理位置的稻瘟病病原体的稻瘟病菌(rice blast pathogen Magnaporthe oryzae)的分离株具有抗性的稻瘟病抗性基因(Ballini等(2008)Mol.Plant Microbe Interact.21:859-868)。例如,Pib对大部分的稻瘟病菌的日本分离株具有很强的抗性(Wang等(1999)PlantJ.19:55-64)。相反,Pi37对稻瘟病菌的日本分离株仅具有部分抗性,但对稻瘟病菌的中国分离株具有完全的抗性。因此,需要分离多个抗性基因,以充分认识抗稻瘟病的分子基础。通过标记辅助的育种或通过转基因方法,这些基因的特性将促进农业上的有用的水稻品种的发展。
迄今为止,共9个稻瘟病抗性基因已被克隆和表征:Pib、Pita、Pi9、Pi2和Piz-t、Pi-d2、Pi36、Pi37、以及Pikm。除了Pi-d2(非-RD受体样激酶)之外,这些基因都编码NB-LRR型蛋白。这些克隆的稻瘟病抗性基因的不同的特征已经被观察到。Pib蛋白包括重复的NB区域。Pita缺乏典型的LRR,但包括:由不同长度的不完全重复所组成的富亮氨酸结构域(LRD)。在Pita的LRD发现单个的氨酸差异,以区分易感等位基因(susceptible alleles)的抗性。等位基因Pi2和Piz-t在三个连续的LRRs内显示8个氨基酸差异,这些残基决定了抗性的特异性。Pi9基因与Pi2和Piz-t基因高度相似,位于染色体6上的相同区域。Pikm-介导的抗性需要两个相邻NB-LRR基因Pikm1-TS和Pikm2-TS。在这些克隆的R基因中,只有Pita被发现与相应的稻瘟病曲霉菌无毒力蛋白AvrPita相互作用。因而,由NB-LRR型蛋白介导的防御信号在水稻中的表征较弱。
据报道,Pi5对从韩国和菲律宾收集的许多稻瘟病曲霉菌的分离株具有抗性(Wang等(1994)Genetics 136:1421-1434)。为了获得对Pi5-介导的稻瘟病抗性的分子基础进一步理解,我们使用了基于图谱的方法来分离Pi5基因组区域。我们先前在RIL260水稻品种中把Pi5插入到染色体9短臂的170-kb的区间内(Jeon等(2003)Molecular Genetics and Genomics 269:280-289)。
根据韩国专利申请No.10-0764563,描述了用于诱导植物疾病抗性的基因、包括该基因的载体和由该载体获得的转化株。此外,根据韩国专利申请No.10-0701302,描述了分离自野生水稻的植物疾病抗性ogpr 1基因、该基因的氨基酸序列和使用该基因得到的转化株。然而,上述的基因与本发明的基因是不同的。
发明内容
本发明是考虑到上述需求而进行的。具体来说,为了进一步缩小新的图谱种群(mapping polulation)中的Pi5抗性基因座,通过对有抗性的基因组区域的序列的分析,确定了两种类型的CC-NB-LRR基因,作为Pi5的候选基因。此外,生产了表达一个或两个上述候选基因的转基因水稻株,并表征了它们抗稻瘟病曲霉菌的表型。
为了解决上述问题,本发明提供了对稻瘟病曲霉菌具有增强抗性的Pi5-1和Pi5-2蛋白。
此外,本发明还提供了编码所述蛋白的基因。
此外,本发明还提供了包括所述基因的重组体载体。
此外,本发明还提供了用所述重组体载体转化的植物和该植物的种子。
此外,本发明还提供了通过在植物中表达所述基因以增强对植物病原体抗性的方法。
此外,本发明还提供抗所述蛋白的抗体。
此外,本发明还提供了一种组合物,该组合物含有用于增强对植物病原体的抗性的基因。
根据本发明,基于Pi5-1基因和Pi5-2基因间的协同作用,增强了对植物病原体(特别是稻瘟病曲霉菌)的抗性。
附图说明
图1显示了RIL260/IR50种群的Pi5基因座的染色体定位。(上)在RIL260/CO39和RIL260/M202的标记C1454和S04G03间,显示出170-kb的Pi5抗性基因组区域。(下)在RIL260/IR50种群中确定的Pi5区域中8个罕见的重组体的示意图。相关的分子标记间显示断裂点。空白条表示假定的RIL260基因组,黑条表示IR50基因组,阴影条表示在两个基因组间的区域是杂合的。粗箭头表示通过图谱种群分析定界的携带有Pi5基因座的130-kb的最小区间。确定了每个系F3子代对稻瘟病曲霉菌PO6-6的抗性,R,抗性;S,易感性;R/S,分离系(segregating line)。
图2显示了RIL260和Nipponbare品种中Pi5基因座的基因组序列比较。显示了两个基因组的由RiceGAAS确定的预测的ORFs。NB-LRR基因,Pi5-1等位基因,以及Pi5-2和Pi5-3基因用黑色箭头表示。在RIL260中缺失的Pi5-1Nipponbare等位基因的N-端区域用绿色显示。推测的转座子(putative transposons)和假定的基因(hypothetical genes)分别用蓝箭头和灰箭头表示。有数字的空白箭头预测编码下列蛋白:1、推测的真核的翻译起始因子;2、推测的GTP-结合蛋白;3、推测的四氢叶酸酯合成酶;4、推测的醛糖1-表异构酶;5、推测的组蛋白H5;6、推测的冷休克-DEAD-盒蛋白A;7和10、锚蛋白样蛋白;8和9、含重复HGWP的蛋白。红虚线表示RIL260和Nipponbare ORFs的高相似度(>90%)。细线表示很少或没有同源性的染色体区域。箭头表示转录的方向。RIL260DNA序列的间隙(gap)用虚线框表示。
图3显示了对转基因水稻株的分析。(A)与稻瘟病曲霉菌PO6-6接种后2天,Pi5-1,Pi5-2和Pi5-1/Pi5-2F1转基因水稻株的PT-PCR分析。水稻Actin1基因被用于作为这些反应的内部对照。(B)与稻瘟病曲霉菌PO6-6接种后7天,Pi5-1,Pi5-2,和Pi5-1/Pi5-2F1转基因株的疾病症状。(C)来源于对稻瘟病曲霉菌PO6-6感染有反应的Pi5-1-63/Pi5-2-74 F1子代的转基因株的F2子代的基因组DNA PCR分析和疾病反应。抗性品种(携带Pi5的RIL260)和易感品种(缺失Pi5基因的Dongjin(DJ))用作对照。
图4显示了Pi5-1和Pi5-2的基因组结构,以及他们的基因产物。外显子以浅灰色框表示,内含子以粗线表示。5′-及3′-非翻译区(UTR)以深黑框表示。ATG和TGA分别表示翻译起始密码子和翻译终止密码子,数字显示氨基酸位置。
图5和6分别显示了Pi5-1蛋白和Pi5-2蛋白的氨基酸序列。这两种蛋白都含有卷曲螺旋(CC)、核苷酸结合部位(NB)、富含亮氨酸的重复单位(LRR)、以及C-端区域(CT)。用加下划线的斜体字表示的Pi5-1的31-67位氨基酸和Pi5-2的26-87位氨基酸包括CC基序。以NB蛋白为特征的保守的内部基序(即P-环、激酶-2、RNBS-B、GLPL、RNBS-D和MHDV结构域)用下划线和粗体表示。在许多NB-LRR蛋白的LRR中发现的保守的xLDL基序也是加下划线的。
图7显示了对用稻瘟病曲霉菌PO6-6接种的RIL260品种的Pi5基因和PBZ1基因的RT-PCR分析。在病原体接种后0、4、12、24、48和72小时,从RIL260叶组织制备的cDNAs用于试验。水稻Actin1基因被用作内部对照。
具体实施方式
为了实现上述发明目的,本发明提供了能增强对稻瘟病曲霉菌抗性的Pi5-1和Pi5-2蛋白。在这方面,每种Pi5-1和Pi5-2蛋白不能独立地增强对稻瘟病曲霉菌的抗性。相反,对稻瘟病曲霉菌抗性的增强只有通过Pi5-1和Pi5-2蛋白间的协同作用而获得。
本发明的Pi5-1和Pi5-2蛋白的范围包括:具有从水稻分离的SEQ IDNO:1或SEQ ID NO:2表示的氨基酸序列的蛋白和上述蛋白的功能等价物。术语“功能等价物”是指:氨基酸残基的替换或缺失的结果,所述功能等价物与SEQ ID NO:1或SEQ ID NO:2代表的氨基酸序列具有至少70%、优选至少80%、更优选至少90%、仍然更优选95%的同源性,因此,所述功能等价物表示与由SEQ ID NO:1或SEQ ID NO:2表达的蛋白具有基本上相同的生物学活性的蛋白。术语“基本上相同的生物学活性”是指植物对稻瘟病曲霉菌增强的抗性。
本发明还提供了编码上述Pi5-1和Pi5-2蛋白的基因。本发明中,Pi5-1和Pi5-2基因中的每一种均可以包括蜷曲螺旋(CC)、核苷酸结合结构域(NB)和富亮氨酸的重复单位(LRR)结构域(见图5和图6)。本发明的基因包括基因组DNA和编码Pi5-1和Pi5-2蛋白的cDNA。更具体地说,Pi5-1的cDNA序列包括5′和3′非翻译区域(分别包括70bp和220bp),并且Pi5-1的cDNA序列编码1,025个氨基酸。Pi5-2的cDNA序列包括5′和3′非翻译区域(分别包括73bp和164bp),并且Pi5-2的cDNA序列编码1,063个氨基酸。
优选地,本发明的Pi5-1和Pi5-2中的每一种的基因组DNAs均可以能包括如SEQ ID NO:3或SEQ ID NO:4所示的核苷酸序列。此外,本发明的Pi5-1和Pi5-2中的每一种的cDNAs均可以包括如SEQ ID NO:5或SEQ IDNO:6所示的核苷酸序列。上述核苷酸序列的变体也在本发明范围内。具体地说,所述基因可以包括与如SEQ ID NO:3至SEQ ID NO:6所示的核苷酸序列具有至少70%,优选至少80%,更优选至少90%,仍然更优选95%的同源的核苷酸序列。对于某一多核苷酸,术语“序列同源性”由对最佳排列的具有待比对区域的两个核苷酸序列的比对来确定的。在这方面,待比对的区域中的部分核苷酸序列可以包括相对于与最佳排列的两个序列有关的参比序列(无任何插入或缺失)而言的插入或缺失(即,间隙)。
此外,本发明提供了一个重组体载体,其中,该重组体载体包括本发明的Pi5-1和Pi5-2。
术语“重组体”是指能够复制异源核苷酸或表达该异源核苷酸的细胞、肽、异源肽或由所述异源核苷酸编码的蛋白质。重组体细胞能够表达自然状态下的细胞中未发现的正义或反义形式的基因或基因片段。此外,当通过人工方法将所述基因修饰并再次引入到所述细胞中时,所述重组体细胞能够表达在自然状态中发现的基因。
此处所用术语“载体”是指被递送至细胞的DNA片段和核苷酸分子。载体能复制DNA,并在宿主细胞中独立地复制。术语“递送系统”和“载体”通常可以互换地使用。术语“表达载体”是指包括期望的编码序列和对于在特定的宿主有机体中表达可操作连接的编码序列所必需的其他适当的核苷酸序列的重组体DNA分子。
优选地,本发明的重组体载体是一个重组的植物表达载体。
所述植物表达载体的优选例子是Ti-质粒载体,当所述载体位于合适的宿主(诸如根癌土壤杆菌)中时,所述Ti-质粒载体能转移它本身的一部分(即所谓的T区域)到植物细胞中。目前,其他类型的Ti-质粒载体(见EP0 116 718 B1)用于将杂和基因转移到原生质体中,该原生质体通过将所述杂合DNA适当地插入到植物基因组中而产生新的植物。特别地,Ti质粒载体的优选形式是在EP 0 120 516 B1和USP No.4,940,838中描述的所谓二元载体。其他可以用于将本发明的DNA转移到植物宿主中的载体,可以选自双链植物病毒(如,CaMV)、单链植物病毒、以及来源于复染色体病毒等的病毒载体(例如,不完整的植物病毒载体)。所述载体的使用是有益的,特别是当植物宿主不适宜被转化的时候。
所述表达载体可以包括至少一个选择性标记。所述选择性标记是具有允许基于通常的化学方法进行选择的特性的核苷酸序列。能被用来从非转化细胞中区分出转化的细胞的任何种类基因均可以作为选择性标记。例子包括:对除草剂(如草甘膦和磷酸麦黄酮)有抗性的基因;以及对抗生素(如卡那霉素、G418、博来霉素、潮霉素和氯霉素)有抗性的基因,但不局限于此。
对于根据本发明的一个具体实施方式的所述植物表达载体,启动子可以是CaMV 35S、肌动蛋白、泛素、pEMU、MAS或组蛋白启动子中的任意一种,但不局限于此。术语“启动子”是指:为了起始DNA的转录,RNA聚合酶所结合的DNA分子,并且启动子对应于结构基因的上游DNA区域。术语“植物启动子”表示能在植物细胞中启动转录的启动子。术语“结构启动子”表示在大多数环境情况和发育状态或细胞分化状态下有活性的启动子。由于转化株可以在不同阶段通过不同的机制筛选出,因此,结构启动子对本发明就是优选的。因而,本发明中并不限制筛选结构启动子的可能性。
对于所述终止子,任何常规的终止子均可用于本发明。所述终止子的例子包括:胭脂碱合酶(NOS)、水稻α-淀粉酶RAmy1A终止子、菜豆碱终止子和用于根癌土壤杆菌的optopine基因的终止子等,但是不局限此。关于终止子的必要性,已知该区域能增加植物细胞转录的可靠性和有效性。因而,由本发明的上下文看来,优选使用终止子。
此外,本发明提供了一种植物,其中,该植物由根据本发明的重组体载体转化。
根据本发明,所述植物是单子叶植物,包括水稻、大麦、玉蜀黍、小麦、黑麦、燕麦、草坪草(turfgrass)、干草、粟、甘蔗、黑麦草、果树草(turfgrass)等等。最优选的是水稻。
此外,本发明提供了所述植物的种子。优选地,所述种子是水稻种子。
此外,本发明提供了增强对植物病原体的抗性的方法,其中,该方法包括:用包括Pi5-1和Pi5-2基因的本发明的重组体载体转化植物,并在该植物中表达Pi5-1和Pi5-2基因的步骤。
优选地,该病原体为稻瘟病曲霉菌。更优选地,所述植物是单子叶植物,包括水稻、大麦、玉蜀黍、小麦、黑麦、燕麦、草坪草、干草、粟、甘蔗、黑麦草、果树草等等。最优选水稻。
植物转化是指能够将DNA转入到植物的任何方法。这样的转化方法不必须具有再生和/或组织培养的时期。目前植物物种的转化不只对于双子叶植物,对单子叶植物也非常普遍。原则上,任何转化办法均可以用于将本发明的杂合DNA引入到适当的祖细胞。所述转化方法可以选自以下方法:用于原生质体的钙/聚乙烯甘醇方法(Krens,F.A.等人,1982,Nature 296,72-74;Negrutiu I.等人,June 1987,Plant Mol.Biol.8,363-373);用于原生质体的电穿孔方法(Shillito R.D.等人,1985 Bio/Technol.3,1099-1102);用于植物成分的纤维注射方法(Crossway A.等人,1986,Mol.Gen.Genet.202,179-185);用于多种植物成分的粒子轰击方法(DNA或RNA包被的)(Klein T.M.等人,1987,Nature 327,70);或通过植物入侵或完整成熟花粉或小孢子转化(EP 0 301 316)而由根癌土壤杆菌介导的基因转移中的(非完整)病毒感染方法。本发明中优选的方法包括土壤杆菌属介导的DNA转移。尤其,在EP A 120 516和USP No.4,940,838中描述的双载体技术优选用于本发明。
用于根据本发明的植物转化的术语“植物细胞”可以是任何植物细胞。植物细胞可以是培养细胞、培养组织、培养器官或整体植物;优选地是培养细胞、培养组织或培养器官;更优选的是培养细胞的任何形式。优选地,该植物是水稻。
术语“植物组织”包括具有用于培养的各种形态的分化或未分化的植物组织(包括根、茎、叶、花粉、种子),例如单细胞、原生质体、蓓蕾和愈合组织,但不局限于此。植物组织可以是整株的,或为器官培养、组织培养或细胞培养的状态。
此外,本发明还提供了抗本发明的Pi5-1和Pi5-2蛋白的抗体。根据本发明,术语“抗体”包括单克隆抗体、多特异性抗体、人抗体、人源化抗体、骆驼源抗体(camelised antibody)、嵌合体抗体、单链Fvs(scFv)、单链抗体、单结构域抗体、Fab片段、F(ab)片段、结合有二硫化物的Fvs(sdFv)、抗个体遗传型(抗-Id)抗体或能够结合上述任何表位的片段。尤其是免疫球蛋白分子和免疫球蛋白分子的免疫活性片段(例如包括抗原结合区域的分子),也包括在本发明的抗体内。免疫球蛋白分子可以是IgG、IgE、IgM、IgD、IgA和IgY中的任意一类,或IgG1、IgG2、IgG3、IgG4、IgA1和IgA2或它们的亚类中的任何一类。
本发明的抗体可以根据常规的方法制备,该方法包括:通过下述典型的操作将本发明的基因克隆在表达型载体中,以获得蛋白并从这种蛋白中制备抗体。因此,也包括能够从所述蛋白中生成的部分肽。关于本发明的部分肽,它含有至少7个氨基酸,优选至少9个氨基酸,更优选至少12个氨基酸。本发明的抗体的类型是没有明确的限制的。单克隆抗体、多克隆抗体和具有抗原结合特性的部分抗体都包括在本发明的抗体中。各种免疫球蛋白抗体也包括在内。而且,如人源化抗体的特殊抗体等也包括在本发明的抗体内。
此外,本发明还提供了含有能够增强对植物病原体抗性的Pi5-1和Pi5-2基因的组合物。由于本发明的Pi5-1和Pi5-2蛋白基于它们之间的协同作用能够增强对植物病原体的抗性,因此,含有Pi5-1和Pi5-2基因的组合物可以被用来增强对植物病原体的抗性。优选地,该植物病原体是稻瘟病曲霉菌。
参照下述实施例对本发明进行更详细的说明。然而,这只是具体地举例说明本发明,本发明绝不局限于这些例子。
实施例
植物材料
携带Pi5等位基因的RIL260水稻品种和稻瘟病易感品种(IR50)被用作本研究中的亲本品系。RIL260和IR50品种杂交生成用于基因连锁分析的图谱种群。RIL260/IR50F1个体的自花授粉种子(F2)被收集,以获得足够大的图谱种群。一种山茶水稻品种(Dongjin)被用作稻瘟病曲霉菌接种和水稻转化实验的易感对照。RIL260和携带Pi5的单基因水稻品系IRBL5-M被用来作为在稻瘟病曲霉菌实验有抗性的对照品种。另外的8个单基因水稻品系,IRBLi-F5、IRBL9-W、IRBLb-B、IRBLta-K1、IRBLz-Fu、IRBLks-F5、IRBLkm-Ts和IRBLsh-S以及这些单基因品系易感背景品种Lijiangxintuanheigu(LTH)也被用于确定稻瘟病曲霉菌分离株的毒力模式的接种实验。水稻幼苗生在白天室温30℃和夜里20℃的14/10小时周期的光/暗循环的温室。
病原体接种和疾病评估
与Pi5抗性基因座不相容的一种菲律宾分离株稻瘟病曲霉菌PO6-6已被通常地用于检测这个基因座。为了在Pi5转基因水稻株中分析稻瘟病的抗性,使用了另外5个不同的韩国稻瘟病曲霉菌分离株KJ105a、KJ107、KJ401、KI215和R01-1。所有接种和疾病评估在Kyung Hee大学温室设施内,使用从Liu等(2002,Mol.Genet.Genomics 267:472-480)轻微改良的方法进行。每种确定的重组体品系和转基因植物的F3子代的3周龄植物被用于接种实验。稻瘟病曲霉菌在燕麦片琼脂培养基上于24℃黑暗中生长2周。分生孢子在收集前4天通过用消毒的环刮盘表面而被诱导。接种的植物被放置于密封的容器内在黑暗中24℃保持湿度24小时,然后转到在14/10小时(光/暗)光周期下24℃和80%湿度的生长室。接种后7天,完成疾病评估。
来自RIL260/IR50图谱种群子代的基因型分析
C1454和JJ817的酶切扩增多态序列(CAPS)标记和一个序列特征扩增区域(SCAR)标记JJ803(符合先前报道的占主导地位的标记JJ803)被用于RIL260/IR50分离子代的分析(表1)。按需使用占主导地位的标记JJ113-T3和S04G03。
表1.本研究中使用的PCR引物
通过简单的小量方法从水稻株幼叶中分离基因组DNA(Chen和Ronald(1999)Plant Mol.Biol.Rep.17:53-57)。用50ng基因组DNA做为模板,在最终的30μl体系(每种引物100pM、每种dNTP 200μM、10mM pH 9.0的Tris-盐酸、2mM MgCl2、50mM KCl、0.1%聚乙二醇辛基苯基醚X-100和0.5UTaq聚合酶)进行PCR分析。针对CAPS标记,C1454和JJ817PCR产物随后分别被MluI和AseI消化,然后在琼脂糖凝胶上分级(size-fractionated)。
DNA测序和基因预测
跨越Pi5基因座的RIL260双BAC(BIBAC)克隆被选作DNA序列分析。由小量制备纯化的质粒被Sau3AI部分消化,并进行琼脂糖凝胶电泳分离。用商业试剂盒(凝胶提取试剂盒,Qiagen)分离该0.5-3.0kb的基因组DNA片段,亚克隆到pBluescriptII SK(-)(Clontech)的BamHI部位,然后用电穿孔转化到E.coli DH10B。为了对每种带有25-kb平均大小的插入的BIBAC克隆进行DNA测序,筛选约60个克隆并用T3和T7引物在一个或两个方向上进行测序。
用BLAST(基本的局部对比检索工具)执行对NCBI资料库(http://www.ncbi.nlm.nih.gov/)的相似性检索。为了预测蛋白质编码基因区域,使用水稻基因组自动注释系统(水稻GAAS)(http://RiceGAAS.dna.affrc.go.jp/)。
用于遗传互补实验的载体构建
通过来自于BIBAC克隆的亚克隆重建Pi5-1和Pi5-2的基因组DNA区域(Jeon等人.(2003)Mol.Genet.Genomics 269:280-289)。为了构建携带整个Pi5-1编码区的克隆,将包括0.5-kb预测启动子的JJ80载体的6.6-kbBamHI-SacI片段亚克隆到双载体pC1300intC(基因库登记号.AF294978)中。合成的质粒JJ104用BamHI和BstEII进行消化,并融合到JJ106的7.3-kbHindIII-BstEII插入片段,构建具有5.2-kb启动子区域的JJ105。0.5-kbSacI-XhoI片段用引物5′-GTCCAAAGAGAAATGCGACAACAC-3′(SEQ IDNO:21)和5′-CGCTCGAGGTGGCATTTCATCCAATAGGCAAC-3′(SEQ IDNO:22)通过PCR进行扩增。将合成的产物插入到JJ105延伸终止子区域,产生携带11,516-bp Pi5-1基因组区域的JJ204构建体。
Pi5-2基因由以下4个片段的多重配位构建:4.2-kbJJ113的EcoRI-BglIIDNA片段;用引物5′-GGATGATGTGATCTGCAGAGAAAC-3′(SEQ ID NO:23)和5′-CAGCCTCACTGAAATTGCGAAGCA-3′(SEQ ID NO:24)扩增的200-bp BglII-ClaI PCR产物;4.2-kb JJ120的ClaI-XbaI DNA片段;以及EcoRI-XbaI消化的pC1300intC载体片段。在合成的构建体JJ117中,通过克隆JJ120的3.7-kb NsiI-EcoRI片段,启动子区域被延伸。最后,通过插入0.9kb-延伸的终止序列到JJ142质粒的Eco065I部位,构成JJ212中Pi5-2的13,250-bp的整个基因组序列。在JJ204和JJ212的克隆的基因组序列通过DNA测序被确定。
转基因水稻株的产生
通过电穿孔将Pi5-1和Pi5-2的基因组克隆转化到根癌土壤杆菌EHA105或LBA4404,并根据已经确定的程序由土壤杆菌属转入到易感的水稻品种(Jeon等人.(2000)Plant J.22:561-570)。转基因植物(T0)是自花授粉的,T1种子被收集。因基于转基因的分离模式的T1品系的自花授粉,继而从T2子代中选取纯合子Pi5-1(Pi5-1-63)和Pi5-2(Pi5-2-74)转基因品系。携带Pi5-1和Pi5-2的F1株从Pi5-1-63和Pi5-2-74品系间的交叉产生,并自花授粉产生F2株。
Pi5-1和Pi5-2cDNAs的分离
从与稻瘟病曲霉菌PO6-6接种后24和48小时收集的水稻叶,并用Trizol试剂制备两种总RNA制品。用PolyATtract mRNA隔离系统(Promega)从每批总RNA中获得纯化的mRNA,并以1∶1比例混合以合成cDNA。经由凝胶过滤按大小进行分级分离,以筛选超过0.5kb的cDNA,用单ZAP XR载体构建(Stratagene)cDNA文库。然后,经由集落印记杂交(colony blothydridization)分别用Pi5-1和Pi5-2编码区域的相应的探针(570-bp JJ204的HindIII-KpnI片段和589-bp JJ212的EcoRV-SpeI片段部分),筛选文库。通过DNA测序分析对分离出的cDNA克隆进行分析。
RT-PCR分析
为了检测响应病原体治疗在转录本积累上的变化,收集不同时期的来自于10RIL260、IRBL5-M和与稻瘟病曲霉菌PO6-6接种的转基因水稻株中的每一种的叶子,以进行RT-PCR分析。总RNA用Trizol试剂和一种寡-dT的逆转录引物以及第一链cDNA合成试剂盒(Roche)制备。在PCR反应中使用第一链cDNA和基因特异性引物。水稻Actin1基因和发病机制相关的可诱导的烯丙苯噻唑(PBZ1)基因的引物用于内部对照(表1)。PCR条件如下:94℃5min,随后28-35个以下循环:94℃,1min;56℃,1min;72℃,1min,以及最后在72℃5min的延伸。对每个引物设定三个独立的扩增。
实施例1:携带Pi5的130-kb的染色体区域的基因特征
先前,Pi5抗病基因被划分到在水稻染色体9的两个侧翼标记S04G03和C1454之间的170-kb的区间。这个发现是我们先前分析产生于携带Pi5的RIL260和易感品种CO39之间的杂交,以及RIL260和另一个易感品种M202之间的杂交的两个种群的结果(Jeon等人.(2003)Mol.Genet.Genomics269:280-289)。为了进一步描绘Pi5基因,在本研究中,我们产生了来源于RIL260和另一个易感品种IR50之间杂交的第三个图谱种群。通过PCR筛选,我们发现在检测过的易感品种中,只有IR50包括占主导地位的标记JJ817,这也在抗性品种RIL260被发现(未显示数据)。相反,我们不能在包括CO39和M202的其他易感品种中扩增JJ817的PCR产物。我们选定IR50作为基于RIL260和IR50基因组区域之间的相似性的一个标测亲本,我们推测其能促进在这个区间的重组。
为了确定在170-kbPi5基因座内的罕见重组,运用CAPS标记JJ817和C1454以及SCAR标记JJ803的预先筛选策略,在我们目前的RIL260/IR50F2种群分析中使用。在被分析的2,014个F2个体中,我们确定了JJ817和JJ803之间的8个重组体,但没有位于JJ803和C1454之间的重组体(图1)。用占主导地位的标记JJ113-T3和S04G03,我们随后确定了我们在它们的子代(F3)株分离的这8个重组体的断裂点,这使我们能从杂合子基因型区分出纯合子。总的来说,发现所有8个品系中含有JJ113-T3和JJ817之间的重组体。
在每例的F3子代中确定了源自这8个确定品系的稻瘟病曲霉菌PO6-6感染的疾病表型。这些实验进一步将Pi5基因定位在标记JJ817和C1454间一个130-kb的区间(图1)。我们先前的和目前的结果显示标记JJ803和JJ113-T3都与Pi5介导的抗性共分离(图1)。我们不能进一步在Pi5基因座上细致标测R基因。
实施例2:包括Pi5基因座的130-kb染色体区域的基因序列分析
为了确定候选在Pi5基因座的R基因,涵盖130-kbPi5区域的7个BIBAC克隆——JJ80、JJ98、JJ106、JJ110、JJ113、JJ120和JJ123被筛选和测序。用这些序列对照公共数据库的BLAST搜索和并且用水稻GAAS程序基因注释分析在RIL260品种的Pi5基因座预测整体18个可读框(ORFs):7个假定轭蛋白、2个NB-LRR蛋白、2个推测的转座子蛋白、1个推测的真核生物的翻译起始因子、1个推测的GTP-结合蛋白、1个推测的四氢叶酸酯合成酶、1个推测的醛糖1-表异构酶、1个推测的组蛋白H5、1个推测的冷休克-DEAD-盒-蛋白A和1个锚定蛋白(图2)。从这基因组序列分析,2个显示与NB-LRR抗病基因同源性的Pi5候选基因在RIL260和指定的Pi5-1和Pi5-2被确定。
130-kb RIL260 Pi5区间的从JJ803到JJ817的接近90kb的区域,与由已测序的品系Nipponbare表示的山茶基因组的相应区域进行比较(国际水稻基因组测序计划2005;图2)。作为结果的序列分析显示,Nipponbare Pi5区间包括2个NB-LRR基因、Os09g15840(一个Pi5-1等位基因)和在RIL260、Os09g15850、被指示的Pi5-3中没有确定的基因。相反,Nipponbare缺乏相应的Pi5-2。值得注意的是,RIL260和Nipponbare的Pi5-1等位基因的5个上游序列非常不同,显示在这些等位基因调节序列内的极端序列趋势。另外,我们没有在RIL260和Nipponbare的90-kbPi5区间的任何其他部分发现显著的序列相似性(图2)。这些结果提示Pi5抗病基因座在这些抗病和易感水稻品种有显著的差异。
由于该基因座的大间隙,我们没有比较Pi5抗病基因座与公开序列的籼稻品种93-11的基因座。在一个接种实验,我们发现Nipponbare和93-11都对稻瘟病曲霉菌PO6-6易感(未显示数据),说明它们都没有携带Pi5抗病基因。
实施例3:表达Pi5候选基因的转基因水稻株的特征
要确定Pi5-1和Pi5-2两个候选基因哪个为Pi5对M.曲霉菌介导的抗病性负责,我们分别用携带Pi5-1和Pi5-2的基因组克隆JJ204和JJ212,在天然启动子的对照下,转化用土壤杆菌介导转化的易感粳稻品种Dongjin。生成的转基因品系的RT-PCR分析显示:独立转化的品系的15个Pi5-1中的13个以及13个Pi5-2中的12个,表达它们的在稻瘟病曲霉菌PO6-6接种的转基因(图3A)。主要的携带Pi5-1或Pi5-2的转基因品系(T0)被与稻瘟病曲霉菌PO6-6接种。令人惊讶的是,无论如何,13个Pi5-1或12个Pi5-2转基因株中没有显示对稻瘟病曲霉菌分离株PO6-6显示出抗性(图3B)。为了确定这些结果,我们接种来自于这些T0品系的T1子代,并发现所有子代对对稻瘟病曲霉菌分离株的易感性与野生型对照Dongjin品系相同。这显示无论Pi5-1或Pi5-2都不能单独赋予对稻瘟病曲霉菌PO6-6的抗性。
实施例4:表达Pi5-1和Pi5-2的转基因水稻株的特征
因为最近的报道(Sinapidou等人.(2004)Plant J.38:898-909)已经证实对于病原体感染的抗性2个R基因是必需的,我们决定检测表达抗稻瘟病的2个候选基因株。我们因而通过一个高度易感纯合子Pi5-1品系#63(Pi5-1-63)与高度易感纯合子Pi5-2品系#74(Pi5-2-74)杂交生成携带Pi5-1和Pi5-2的转基因株。基因表达分析显示来源于在稻瘟病曲霉菌PO6-6接种上Pi5-1和Pi5-2基因均表达的杂交的F1株。引人注目的是,23个Pi5-1-63/Pi5-2-74F1株检测均显示完全抗稻瘟病曲霉菌PO6-6。携带Pi5-1或Pi5-2转基因品系像先前确定的那样易感(图3B)。
为了确定这个发现,我们把来自于Pi5-1-63/Pi5-2-74 F1品系的F2子代株与瘟病曲霉菌分离株PO6-6接种。72个被检测的F2子代中37个携带2个转基因,并赋予抗稻瘟病曲霉菌PO6-6。相反,缺乏Pi5-1和Pi5-2的F2子代是易感的(图3C)。RT-PCR分析证实Pi5-1-63/Pi5-2-74品系在稻瘟病曲霉菌PO6-6接种之前和之后在相似于RIL260的水平表达它们的转基因。为了检测如果Pi5-1和Pi5-2对抗其他稻瘟病曲霉菌分离株是必需的,我们接种与Pi5不相容的带有4个另外的分离株的转基因株。这些分离株显示在携带不同单R基因水稻品系的不同的毒力模式,确认这些是真正不同的稻瘟病分离株。我们发现共表达Pi5-1和Pi5-2的转基因株抗所有被检测的稻瘟病曲霉菌分离株。抗病供者RIL260和携带Pi5的单基因品系IRBL5-M也抗这4个分离株。相反,Dongjin和只携带Pi5-1或Pi5-2的植株对检测的稻瘟病曲霉菌分离株易感(表2)。这些结果证实2个NB-LRR基因Pi5-1和Pi5-2对Pi5-介导的抗稻瘟病曲霉菌分离株是必需的。
表2.转基因株对稻瘟病曲霉菌分离株的疾病反应
a转基因植株
bR,抗性;S,易感性。
Pi5单基因品系IRBL5-M对稻瘟病曲霉菌KI215易感。基因组序列分析显示携带Pi5的IRBL5-M基因组区域与RIL260的相同(未显示数据)。另外,RT-PCR分析进一步证实IRBL5-M在稻瘟病曲霉菌PO6-6接种之前和之后,表达Pi5-1和Pi5-2的水平与RIL260相似。基于这些结果,我们假定Pi5-1和Pi5-2均表达的转基因株将也对稻瘟病曲霉菌KI215易感。实际上,我们的接种结果显示Pi5-1和Pi5-2均表达的转基因株对稻瘟病曲霉菌KI215易感。相反,RIL260被发现抗稻瘟病曲霉菌KI215,这显示可能包含另一个对这个分离株赋予抗性的R基因(表2)。
实施例5:由Pi5-1和Pi5-2编码的蛋白的特性和系统进化分析
为了分离研究中相对于2个Pi5基因的cDNA克隆,用从稻瘟病曲霉菌PO6-6接种24和48小时后收集的水稻叶分离的mRNA的Uni-ZAP XR载体,建立RIL260的cDNA文库。该文库用集落杂交法筛选,用Pi5-1和Pi5-2的基因特异区域作为探针。我们分别确定Pi5-1和Pi5-2的7个和5个cDNA克隆。序列分析进一步显示包含整个开放阅读框(ORF)的Pi5-1 cDNA的3个克隆,而其他缺失包绕ATG翻译起始密码子的N-末端。在全部ORF克隆中,最长的克隆(#1-7)被全部测序。这些实验显示T Pi5-1编码1,025个氨基酸的蛋白,ORF分别侧临70bp和220bp的5′-和3′-非翻译区(基因库登记号.EU869185;图4和5)。克隆的序列分析显示了包括整个ORF的5个克隆中的3个。在它们中间,最长的克隆(#2-4)被进一步通过测序特征化。该分析显示Pi5-2编码1,063个氨基酸的ORF,并且这个ORF分别侧临73bp和164bp的5′-和3′-非翻译区(基因库登记号.EU869186;图4和6)。
其推导的氨基酸序列比较显示,Pi5-1和Pi5-2编码N-末端CC,中心定位的NB和LRR,并且也编码C-末端区域(图5和6)。用Pfam和SMART数据库的保守的区域扫描预测Pi5-1的109-576位和Pi5-2的109-567位残基包括NB区域,这是被植物R基因产物共享的信号基序。以NB-包含R基因产物为特征的保守的内部结构域也在Pi5-1和Pi5-2中确定,包括P-环、激酶-2、RNBS-B、GLPL、RNBS-D和MHDV结构域。另外用Paircoil2程序分析(http://groups.csail.mit.edu/cb/paircoil2/)用阈值0.1预测一个在Pi5-1的31-67位和Pi5-2的26-87位氨基酸之间潜在的CC结构域,显示这些蛋白属于NB-LRR抗蛋白CC亚群。
Pi5-1和Pi5-2的LRR区域分别由24.3%和22.6%的亮氨酸残基组成,并包含一系列不同长度的不完整的重复单位(10-12)(图5和6)。值得注意的是,Pi5-1和Pi5-2蛋白的一些重复单位与在其他细胞浆R蛋白发现的共有序列LxxLxxLxxLxLxxC/N/Sx(x)LxxLPxx匹配。Pi5-1的第一和第三个重复区域以及Pi5-2的第三和第六个重复区域包括xLDL基序,该基序在许多NB-LRR蛋白的第三个LRR是保守的(图5和6)。仍然值得注意的是,Pi5-1和Pi5-2蛋白包含一个与其他NB-LRR蛋白不同的独特的C末端,不与任何已知蛋白基序匹配。
cDNA和这些R基因的基因组序列之间的序列比较显示,Pi5-1和Pi5-2分别携带5个和6个外显子(图4)。Pi5基因比较其他对M.曲霉菌赋予抗性的克隆的水稻R基因,在它们的编码区域内有更大数量的内含子。而且,Pi5-1和Pi5-2基因在它们的RNBS-D和MHDV结构域均包含内含子。
实施例6:Pi5-1和Pi5-2基因的表达分析
为了检测是否这两个确定的R基因的表达在病原体治疗时有变化,我们在稻瘟病曲霉菌PO6-6感染的RIL260、IRBL5-M和Pi5-1-63/Pi5-2-74转基因植物进行这两个基因的RT-PCR分析(图7)。为了这个目的,使用从在稻瘟病曲霉菌PO6-6接种后不同时间点收集的3周龄植株的叶子分离总RNA。该结果显示Pi5-1在病原体攻击12小时后表达增加,而Pi5-2基因在感染前后的RIL260均持续低水平表达(图7)。IRBL5-M和Pi5-1-63/Pi5-2-74品系也展示与Pi5基因相似的表达模式。这些发现显示Pi5-1和Pi5-2均在病原体感染期间表达,提示被编码的蛋白也共表达。PBZ1转录本——病原体诱导基因——在M.曲霉菌处理的叶子累积到高水平(图7)。
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740 745 750
Leu Pro Glu Ser Leu Lys Asn Leu Thr Asn Leu Glu Tyr Ile Asn Leu
755 760 765
Ser Asn Ile Gly Glu Ser Ile Asp Phe Asn Gln Ile Gln Gln Leu Arg
770 775 780
His Ile Leu Lys Lys Thr Phe Phe Ser Gly Asp Ile Gly Gly Ser Glu
785 790 795 800
Leu Gln Thr Cys Glu His Ala Ala Asp Ser Ala Asp Ser Lys Lys Glu
805 810 815
Ile Thr Met Asp Phe Ser Ala Asn Leu His Gly Asn Ile Thr Leu Pro
820 825 830
Pro Lys Cys Ser Thr Glu Glu Lys Ser Gly Glu Asn Ser Glu Arg Phe
835 840 845
Leu Ser Ala Ala Val Arg Glu Asp Ser Ser Ser Thr Asp Val Ser Thr
850 855 860
Tyr Val Lys Pro Val Val Ser Ser Leu Ile Gly Val Leu Arg Arg Pro
865 870 875 880
Thr Arg Leu Asp Val Pro Ala Gly Ala Met Ala Ser Gln Val Gly Leu
885 890 895
Ala Gln Met Pro Ser Ser Asn Asn Gly Lys Ala Gly Pro His Pro Thr
900 905 910
Met Ala Ala Ala Gln Thr Pro Glu Ile Asp Gln Pro Val His Lys Arg
915 920 925
Val Arg Trp Asp Asp Ile Ile Asp Tyr Ser Arg Pro Pro Asn Ser Lys
930 935 940
Pro Ala Arg Ser Ala Ser Leu Val Gln Ser Thr Asp Leu Ser Thr Pro
945 950 955 960
Lys Lys Ser Tyr Lys Lys Ile His Ser Met Pro Val Val Tyr Ser Ser
965 970 975
Ile Pro Lys Gly Ser Ser Gly Gly Thr Tyr Leu Met Pro Ala Lys Ala
980 985 990
Ile Ala Ser Ser Tyr Arg Arg Tyr Ser Pro Gln Arg Trp Glu Gln His
995 1000 1005
Ile Gly Tyr Gln Gly Thr Asp Glu Asp Glu Leu Met Val Val Pro Pro
1010 1015 1020
Phe Gly Glu Trp Asp Gln Ser Pro Thr Leu Arg Lys Ser Asp Phe Arg
1025 1030 1035 1040
Tyr Glu Lys Val Phe Ala Lys Leu Thr Glu Glu Lys Met Ser Gly Gln
1045 1050 1055
Arg Gln Lys Pro Gln Gln Val
1060
<210>3
<211>11516
<212>DNA
<213>水稻
<400>3
aagcttgaaa gtgtgtgaag aaatgaagaa agtgaagtct acttatagga caggtataac 60
ggttatgaga agtggaaatg tccgaactgc cccgcaaccg ccatcaggga gtgatctgga 120
gcgtccacgc caaaccccag gatcaacggt gatcaccagg gttcggctga acctgggctg 180
ggccagatgg cttccaaacc tagtctgggt ctcctcctcg acctgcctta tccaattctt 240
gcaattttgg gatgtttctt tagtagttta aaagcccatt cttgcattga tagatttctc 300
cttcactttt agtctggatt tcctcccaac ttctgggttt atcacctgca taaaaatata 360
catcaacaat tgtggaatat gtgagtttta acacctatca ctatgttgga tattttatta 420
tctggacttt atgcagacgt tgacggtata gatttggcat ttaacagtcg ccaacattat 480
acaatcgggt aaatagtatt gatgtattcc atgaatctgt gtatagaagg tatgccatat 540
ccttgacact gacaccatat gtgtttccgt ttttgagaaa acatttccat ttccaaaaat 600
ttctgaccaa cactctctat ttccgaaaac tcatctgaaa tccgaaaact tttcaaaccg 660
ttttcatccc agtaatgaat ggcaggctct acgtgcagac ccacaaatgc tccacaaaga 720
tctcaaaaaa tttatcgtat aaatttcata cgtagactaa atttcacaaa aattcacttg 780
tgtacccatt gaggtatgta ttttgttttc aaaaaaaaac acagtttacc ttgtggtgct 840
cgggacatcg gccggagctc atgatcataa tggagtcagt cagggtggct actggagctt 900
atggcggagg aggtcagggt ggtgggacgt ggggcagagt ttagtccggt cggcaaccag 960
agctccgaga ggagatctgg gcagtagtga gagaagggtc aggaggcgcc agagtggcca 1020
cgacggaggc tgggaaagcg atgccgaaca gcaaagcatg cgttgaagta ggcggggtgg 1080
cagcagcctc ggggcggagc gggcatggaa caagggtaga gctggagtcg gtggccggct 1140
actgagacca ttcacagtgc gtcaccaagt tttgatattc ttcacatgcc acgtagatca 1200
gagcggtatt agccacatgt tacatagtgc aaaactctac aagaccacat atgggttgct 1260
tgtagtgggt cccactaata ccacttttac acttctcttt ttacttttcc cctctcctct 1320
ctcccctctc ttcctttccc caccggccac cgcagagagc cgggcggaga cggaggttgc 1380
gggggtcggg cggagcggcg gctcaagcgg aggtcgtgga ggcgagcagg agacggcgcc 1440
gggccgagtg gcgacggcgg tggggcctag caccgtgcct cccctcctcc tcctcctcta 1500
gtggtggcga atcgaggaag ccgccgccgc cgcgccccac ctcttcatct gatgcgggcg 1560
ggaagctgcc gccaccgccg tcgggcagat ccagatctcc ggcctccgtg tcgggcggct 1620
acgcggcggc gggcgtcggc gtcggtcagc gccctcctca cctggccagc ccaccgacta 1680
gccctaccgt cgtcatcatc cctggtcagc accctcctcc cctggcattc ggctgcctca 1740
cccctctcct ctctctcctc tagcggtggt gtgggtccca cctgtgggtc cctctgagac 1800
gaagaacgcg tagtcaactt tggctacgcg cggagctgag gaacgcacac gattggataa 1860
agtggcaaag gaatatttgc ttttttggtt gaggctaaat gctacgtaga tctactgtga 1920
atggcctgat ggcgcagcgg ctctcttgcg ggacagagtg ggacgtgcac gacgagaaat 1980
ttcctcttgc atgaataacc cacggatgta ggcctcctta actaggagct acgaggtggg 2040
atttgggagg ctttttctat ttcaaggctt aaagcctatc caatatgatc tctaatctta 2100
aaatttagtg tttatttttg aaaaaaagtt ggctccaata attttcctaa ttagcttcca 2160
aaaattggaa tcttactccc tatccttctc gctctccact agtggagaag cgtgctagtt 2220
cttaacacac gcatgggtgg agggatcaca tggatattat ggaaatatcc tcatggactg 2280
gtgattaaca tagtcctact aggagagaat tgagtttatt ctatatcgtt atcgacgttt 2340
ggtatcacga ctacgttatt tggatggtat ggggatcatc ggtactagag tatatgcgag 2400
attgaggtaa aagagatgga gacagatatt tttatactgg ttcgcccctt atctaacagg 2460
taatagccct acatcctgtt ggctaaagcc ggtattgctc ttattcatct gaatcgcaca 2520
agtataatat ttaggataac ctgtctagct gtcatcgact tggcggcatg gataaccaac 2580
tcgtagtcga tgacggggta gtatttctct tcgaatatga actcgtcgag atcagagatg 2640
gctctagatc tctcttgccg gtctcaggag gcaccagatg gggtatgcct aggctaatct 2700
ctaatgtcga tatttagcgg cgtattggct tgtgtgtatg ttatgtgttg tggctcgtca 2760
tctctcctcc tagggggctt gtatttatac ccatagatgc ccccttgtct aagtagaact 2820
agggagataa atatggatac gatccgagta gtccttgtcg tttccatata gaactctttt 2880
tgtcctttct tatccgaaac tccttttata tacgaggtat gattccatat aagacatggt 2940
atatggtggg ccctgccgag cttagtcagg aatgtggtat ccacaaccct gacaatcatg 3000
cagggttcat tgtctccgag ttctacgcgg agactaagac aactcaaggt ataaaaggcg 3060
cactctctaa ggggttcaaa agagcaaatc atagcacgaa cacacccacc atagtttacg 3120
aagccagagg ctgtgaagcc tactcgccag gagatcttgt cgactcatct cgacaaggat 3180
ctcgccggta acgctggatt catctcttct ctttgtactc cgtggtttcc atatcaatct 3240
catataaact ggattatggt tattatctta cgaggggtct aaaccagtat aatctttgtc 3300
tctctgattg ttttaatatc gtatcatgta gatcctcata ccaacttacc ctaatacact 3360
atttatccga tctacaggta tcccctatcg acaataggta tataagatat tataaaaaag 3420
atggagacat atgtgagctg atgatttagg ctgcattcgt tgcagcttct ttccaaccca 3480
tctccctcgt tttccgtgcg catgcttttc aaactgctaa acggtgcgtt ttttacaaaa 3540
agtttatata cgaaagttgc ttaaaaaaat catattgatc cattttttaa aaagatagca 3600
aaaaattaag taatcacacg ctaatggact actctgtttt ccgtgccgga ggatagcttt 3660
cccaacccag ggaaacgaac ccaaccttag gtgtgttcat tgctaggtgt tcccaacccc 3720
tctcccttat attccgtgcg catgcttttc aaactgttaa acggtgtgtc tctttttaaa 3780
aaatttctat acgaaagttg ctaaaaaaat catatcaata catttttgaa aaaaaaagct 3840
aatacttaat taatcacgta cgtactaata gaccgcttcg ttttctatgc gcagaagatt 3900
tgttcccaac ccccacaaca aacacagcct taacatgctt gcatttaaaa agctataaaa 3960
tttaataaat tataaaatta tagataatat aacatgctta cttgatgtga cactttacct 4020
gttaagtttt aggagtcggt gtttagattc aaactttttt tttcaaactt ccaacttttc 4080
catcagatca aatgtttggg cacatgcatt cagcaataaa tgtggacaaa aaaaaaccag 4140
ttgtacagtt tgcatgtaaa tcacgagatg aatcttttga gcctaattac gtcatgattt 4200
gacaatgtgg tgctaaagta aatatttact aatgacagat taattaggtt taatagattt 4260
gtctcgcagt ttacaggcga aatatataat tttttttgtt attagtttat atttaatact 4320
taaaatgtat gtccatatac ttaaaaaaat tttgtaccac gaactaaaca cagccaagtg 4380
gactctaact ctctctctct ctatatatat atatatatat atatagtaat gtgttcgtat 4440
gtcctggata gaaactcatt tcctccgcat agaaaacgga gcggtctatt aatacgtgat 4500
taattaagta ttagctattt ttttcgaaaa taaattaatt taatttttta aataacattt 4560
atatagaaac tttttaaaaa acacgttgat taaccatttg aaaagcgtgc gtgcacgcgg 4620
cgtgaaaaat gaggcagaga tgttgtgaaa aggagtgccg aacacagtca aagcctcagg 4680
tggtgtttgg atccagggac ttaacttaaa ctttagtccc tatatttaga cactaattta 4740
tagtattaaa tatagactac ttataaaact aattacataa atgaaagcta attcacgaga 4800
caattttttt aagcctaatt aatccataat taaagaatgt ttactgtagc atcacatagg 4860
ctaatcatgg attaattaga cttaatagat tcatctcgtg aattagtcca agattataga 4920
tgggctttat taatagtcta cgtttaatat ttataattaa ttttcaatcc aatatgatag 4980
gacttaaaat ttagtcccat ctacagggtc agaggattcg gtcggtctca gggcagtcct 5040
ctccgtataa cgcagcgccc gatatttttt atgggcataa atagtctgat tgctactgta 5100
gcattttgtt caactaactc cccaatgggc aatggctagt cgtcgagtgt cagtagtcaa 5160
gagtagactc cgcttcgctt cgccgtacct tctctcttct cgttctccta tttcacaaat 5220
cacaaccgga ttgctttctt ccttcctccg gtctggtccc caacctccgc ggcagccatg 5280
gttggcgccg agatgcttgt ggccgcggcg gtgagccagg tcgcccggaa gatcaacgac 5340
atcgtggggg tcgcgcaggg cgaggtgaag ctgtgctgca atttcagcga cgatttggag 5400
ggcatcaagg atacccttgt gtacctggaa accttgctga aaaatgcgga gaataactcc 5460
ttcggaagcg acagggccaa cctgcgccac tggcttggcc agatcaagtc cctggcttac 5520
gatatcgaag atatcgttga tgggtactac tcttccaagg agcagttcga tgggggcagc 5580
tatgcacaga aggtaacaga atctcattcc tttttcttca tcggtaaaat ttcttcaatt 5640
tcaactcaat tttagaatgc cccgcaaaaa aaaaatcaat tttagaatgg atctacatta 5700
atgagatgta gaggtgtatt actatgggca ggggaagcac cgtgtgtgca tccttagtta 5760
ctgcataggc aataagttat cctttccatt gtccaaaaaa attttgaagc aaggaggaaa 5820
agcttgaagc ttagtctttt agttttttct ttttgattat tttgttattt atctcagtta 5880
tgagattagg agtgtatatg cactcgtgta gcttttgtct gtgtgtggat atagaggctg 5940
gatttctatc cattatctta aataaattgt ccaagaaatt tatcagggga aatgcagtta 6000
gatgcaccat tagaattgct tcattgcctg tggaagagtg gaacagctct gagaattgtg 6060
attatgtttt atatatttaa acaatattga ttactagtac tagatttact ctcttttttt 6120
ttccttatga aattccttga tactggtagt agtgagagat aaacctaata attacatgcc 6180
acatacctgt agattgtact atacttcaac acccttttgc aaatagtgag agataaacct 6240
aatacttact tacatgccac atcgtcagag cactcaattc ttttttgttt tggtaagcaa 6300
tcagcttttg cctttacata gacaaaaatt gatcgacgaa tattttgaaa aaaaaaacct 6360
ttgtatttat atggaaaaga aaatgcaagg tactttacca aacaattgtg catctatgat 6420
ctatctgcta tgtaggggtt gataccggcc ttggttcctt gctgcaccaa ctgttattta 6480
cctatctcca tttttgttct tccagcaaca atagtatcaa aatactaaat gtctttcctg 6540
actaaggcta ccttcataaa tttacagggg tcattattct gctcgctatc caatccaatg 6600
cttctgaaag gtagcatggt ttataagatg aaatccaaga gagagatgct acagcaaagc 6660
caacagttgc ccaatcagta tcatttcctt tcatatatca attcagctgt gcattatttt 6720
gaggagaagc aaacaacatc atacagaaat actgacattg caattgtcgg gagggatgct 6780
gatttggatc atctcatgga tcttttaatg caaaacagcg ctgaagagct ttgtattata 6840
cccatagttg ggcctgtagg ttttggaaag acaagccttg cacagttagt tttcaatgat 6900
acaagaacag aggtattcag ctttaggata tgggttcatg tttccatggg taatatcaac 6960
cttgaaaaaa ttgggagaga tatagtttca caaactacag aaaaaattga gggaaatatg 7020
cagctgcagt caatcaagaa tgctgttcag cgtgtgctaa ataaatatag ttgcttgatc 7080
ataatagaca gcctttgggg aaaggatgaa gaagtgaatg aattgaagca gatgttgctt 7140
acaggtagac acacagaaag caagatcata gtgaccactc atagcaataa agtagctaag 7200
ctgatttcca ccgttccact gtacaagttg gcagctttat ctgaggatga ttgtttaaaa 7260
atattctctc aaagggcaat gacaggtccg ggtgacccgt tgttcaggga atatggagaa 7320
gaaatcgtta gaaggtgtga aggcacaccc ttggtagcca attttctcgg ttctgtggtg 7380
aatgctcaac gacaaaggcg tgagatttgg caagctgcaa aggatgaaga aatgtggaag 7440
atagaggaag attatcccca agacaaaatt tcaccactat ttccatcatt caagataata 7500
tattataata tgccccatga gctaagatta tgctttgtat attgttcaat cttccctaaa 7560
ggaactgtta tagaaaagaa gaaacttatt cagcaatgga ttgcacttga catgattgag 7620
tccaaacatg gaaccttgcc acttgatgta actgcggaga aatatattga tgaacttaaa 7680
gcaatctatt tccttcaagt tttagagcgg tctcaggtaa gttcatgggt tgctttttac 7740
cttctgtaca tatcctatgt aactagaatg tggttaaata tctccattaa gcatagtagc 7800
ttataccatt gttttatttc taaattctca ataagtttct gtaagaagat tgaccatgat 7860
agaatggcca atagtgatat ctcaacaaac aagtaacact gttttcctcc acagaatgat 7920
gcagaaagat ccagtgcttc tgaggaaatg cttcgcatgc ataacttggc tcatgatctt 7980
gctagatcgg ttgctggtga agatatcctt gttattttag atgccgagaa tgagcgcaat 8040
gccagatatt gcaattaccg ttatgcacag gtgtctgctt ctagtttaga gtcaatcgat 8100
cgcaaggcat ggccttccaa ggcaaggtca ctaattttca agaatagtgg tgtggacttt 8160
gagcatgtca gtgaagttct ttcagtgaac aaatacctgc gtgttttgga tctcagtgga 8220
tgttgtgttc aagatattcc atctcctatc tttcagctga aacaattgag atacctcgac 8280
gtttcatctt tatctattac agcactccct ctgcaaatta gtagctttca taagttacaa 8340
atgttggatc tttcagaaac tgaactaaca gagttgccac cctttataag caacttaaaa 8400
ggactgaatt atttgaatct ccaaggttgc cagaaacttc aacgattgaa tagccttcat 8460
ttgttgcatg atctacatta cctaaacttg tcatgctgcc ctgaagttac tagttttcct 8520
gaatctattg aaaatctgac caaactccgt ttcttgaatc tttctggatg ctctaagctt 8580
tcaacattac ctatcagatt tttggaatca tttgctagcc tctgttcttt ggtagatctt 8640
aacttaagtg gctttgaatt ccaaatgttg cccgactttt ttggcaacat atattcactt 8700
cagtatttaa atctgtcaaa atgtttgaaa cttgaggtat taccacaatc ttttggccaa 8760
cttgcatatc tgaaaagcct aaatctttca tattgttctg atcttaaact gctggaatcc 8820
tttgaatgcc ttacctctct tcggtttttg aatctctcga actgctctag gcttgaatat 8880
ttgccatcat gctttgacaa gcttaataat ttagagtctc tgaatttatc acaatgtctt 8940
ggacttaaag cactacctga atcacttcaa aaccttaaaa atcttcagct tgatgtttct 9000
gggtgtcagg attgtatagt acaatccttt tctctaagta ccagaagttc ccagtcctgc 9060
caacggtcgg agaaagctga gcaggtcaga tcaagaaaca gtgaaatttc agagatcact 9120
tatgaggaac ctgctgagat tgaactttta aagaataatc caagtaaaga tttggcctcc 9180
atctcacacc taaatgagga tagaattgag gagcctgaag ttgtcactga ggtaaactta 9240
cattttatta aagaaataaa aacaatttgc ctagtgtttc ctttgaaatt tccttatgtc 9300
aatacttaat ttatctttga tagatttgag ttactggtga ttgagaaagt tgtatgccaa 9360
tttgaaccag tttctcacta ccaactgaaa atgatgacga acggaaactt tattgtagtt 9420
gtgctcgaat tgaagatccc ttttctaatg aggtgatcta attgggtacc agaacatgaa 9480
tcatactttt ttcagtagtt cttaacattt cgtagagaaa atacatgagt gttccttcaa 9540
ttaaaaaact ccctagaggc cagcaagtta taaatttaaa tgggattctc cttattacct 9600
agatttatgt tttcagaaaa ttttgtaata tcatactaac taattgtcca tgtccttgtt 9660
tcttgatgta gccaagtgca actagaggta tggtacaaca gattccagga aaccagctct 9720
catcgccttc atctcatctt tcttcctttg catcaagctc agcgccattt gcatcctcct 9780
cttcggacac ctcaacaagt gagcatccag tgcctaatga agaggcggca ggtatggtac 9840
ttcaaataat tttctcccga tttaatcatc tcaagatgag gttcacttct ttatttgaat 9900
ataccattat aaggaaatag atcatgcgcc ctcttgttta aaggaatttg atgttttttt 9960
atattcgctc tctttgagat atacctgtca tccagcagtg gtaaaaagga tagtatatag 10020
ctgacaaata tgttcatata atttccttgt gggtaatttt gatattctct ttctatctta 10080
atacgtgtta tgagcaccgg ctaatgaccg aatggggtat gaatggaaca ccggaggtcg 10140
aggcttttgg cagcctcttc gacgtctggc ccatgatcgg cgacgaaagc aaaagcaggg 10200
gatgagagag tagagaattg gagacgagac tatagattga atcttgcttg gttcattcat 10260
gatttcgtgg cccttaatta ggcttacgat tgaactgaat tagctaataa aaaaggataa 10320
caaagtcttc tctaaatgta agcaatcgga tcactatcgc tcgtccggca tgccgcgtca 10380
gttttcagct cgcgtgatct ccttcgacgc cgcttggacc ggttgctgca tccgcgtagt 10440
tgcgtttctt catgtagacg attccgaccc tttacattac aactacttgg aattctttga 10500
gtgttctaaa ataatttgga tagtggtaca tatgtggtta cattttctta atggtgacat 10560
atagccaagt tggactaaga ttaattttaa tttgtgtgga atgagaaaga gtcaagacag 10620
tttgatacgt agaagtttgc atattcagaa ccttatactg gtttgtcttg gtttattttc 10680
ttgacaatgc agctttgaca gttcctcggt ccaaagagaa atgcgacaac actcccatgc 10740
cggtaaaaga tggcctgata tctgaagatg atgcaccggt acatctgcat cagaagcccc 10800
tgcaggcgac agccatggca gccatatgac tgacctgtaa tcctacaaga aaccaactga 10860
agattcatat gtggactgag tgaaattatg aaagttattg gaataaattg ttgctctgta 10920
tgtgagagca agcttcagtc cgttagcctg gttcctttta gtagtgttct actattggga 10980
gatcttcatc aacattttac atgaaacgtg atgtaatgaa cctctgttaa ttgttaattg 11040
tcagagctct agttttttgt ggtataaatt ttgttgcaag ggggcaatgc actcaacttt 11100
tgaatcattt gattggtatg gaaatcattg catttgtgca gagttcaggc agagttcagt 11160
tcttgcattt gcacatcgta ttactattac ctggccttgt ttggcacatc tccagttcca 11220
gctccacctc tcctagagct ggagctcagc caaacagttt cagctccacc gaaaatggga 11280
gcggagctgg gtggagcact ctaacaaaat gaactagaga ggtggagcta ggttaagctg 11340
ttccacaact ccacttcaga tctaactcct aaagttaaat ttaaaagttg aagctctacc 11400
aaacgagaaa acggacgata tgccactgta taggttgggt tcgaatgaaa tgccactcaa 11460
tacaatgtat cggataaaat gccactcaat acgttgccta ttggatgaaa tgccac 11516
<210>4
<211>11403
<212>DNA
<213>水稻
<400>4
tctagatata ttacaggcct aaaattcgga tcacatattt aacacgtagt tgtgtagctg 60
tggtaccttc cgttcaggtc cagcagtggc tttccttgaa cgatactgta cactctagta 120
aaggacaaag aacacgtcaa agctggacac gagatcgaca actgcctgct ggcgggttct 180
tgcaagttgc atgtcggttg agttgtgatc atcgatctct cggaaggcgg agggaactat 240
gtctctctct ctctctctct ctctgtttct cgttagaact gaggtaacta tctcgaaata 300
atcatttctg aggctgcagt tgaagaagtt cagtagccac atctaatata tgctacattt 360
taatattata cgaatcttaa aaaaccttag tacgtgatgc aagaccgaat caatataaac 420
acagtgctgt gcgagcaaag aaccaggctc taccatatcg attggattgt gaaattcgag 480
acgtcgattt tcctctaatt tcaaacaaat ttgacacaaa tcgttcgcaa tacttgtgaa 540
agtcctttcc agtcagcaat tctttttttt taaaagttgt gggacccacc tgtcatactc 600
ctcccactct tcctccctct tactcttcct ctctctcttc tctctctcac tcttcccttc 660
tctctctcac aggcggcaga caggggacag agcagcgtcg gcagcagcgg caagcggcgg 720
caggggcagg tgacgcgaag tcggcagcgg caggggatga gggtgacgtg gacgaggatg 780
gtgatggagg agtggaggac gacgacgagg acgacgctcg ggcgctagca gcggcggcta 840
tggcggatag gcggcggcgc acgtcgaacg aacggcggcg gggacggacg gagcttaggg 900
tggcgggatg gcggcaggcg gtgtcggagg ctgctcgggg cgtcgagaga cagaggcggc 960
cggtggctga ggggtgggga tggcggtgag gagaggagga ctcttcctct gccaccgcgc 1020
tcgccccctc cgccgctcgc ctcgtcggcg cgctagcctg ctccaacgcc ggtagacacc 1080
aagccgccgc tcgagcccgt cccctgcgcc gtccttcctc gccgccgcgc ttgcctcgca 1140
cgcagccaga tgcagacggc aacgcgctag cgcatgggga tggcggcgcg gagcatggtg 1200
tcctccttgg ccatgacggc gacgttgacg cggaggccgt ctcggcggcc ccttcctcgg 1260
tagccacgac cgccgccgtg gcggccgaca tggcgaagct cccctccatc gtggtggctg 1320
acgcggcgaa gttcccctcc ttcctcgccg ccgtcactcc atgtcccccg cctgccacag 1380
acctacagtg cgtgttcaga gagtgaaaga agagagagag gaatagtgga ataggagtat 1440
gacaggtggg ctccacattt ttttaataaa taaaatattg gctggactgc cacgcgtacg 1500
ccacgtagac caaaaccgcc gcggattggg tcgaggggat aattcgtcct gtttgcatag 1560
ttgggtgtaa agaatgtccg gttttgtggt tcagggggta attcgaacga ccgcgatagt 1620
tgggagggta attcgtactt tttcctattt tttatctaat aaaaattaac ctataataaa 1680
ataattagtg atcaaatgtt taaagattta gaatcttaaa attcgtgctt attgagtggg 1740
acggagggag tatttttgtt gcaatataca ttaataaata tatgatttta ttaagatatt 1800
tttaggacta taagataaat gttttagtga attcaattta gtacatccat ctactagctc 1860
cacataatct ataatcaatt taataaacaa tttatacaat aattacatat aaactaccct 1920
attaatatat aatcccacct atcatacgcg cgttgtgtct tagagtccgt gctacagctg 1980
gctacaaatt tgtagcccgc tgctcctctc tctcttttat cttctcggca tatgtttata 2040
gctagattat aggttttgtt tggtttattg gactaatcca ttagtccctc cattttagtc 2100
ccttttattt tttaagaatt ggaggtactt atagtttata gttataggga ctaaattggt 2160
agctcatctc ttctctcttg gtccacgtgc ctttagtccc ttttagtctc tggatctaaa 2220
cagtatggga ctaaagtttc cagtacaggg tcatagcggt tatagcttgt tattgtaccc 2280
tctaagattt aaagtttgac tatagcttta tagctttagg cctgatttag tttcccaaat 2340
tttttaccga aaatatcaca ttgaatcttt ggacacatgc atagagtgtt aaatatagtt 2400
taaaaaaact aattgcataa ttagagagga atcgcgagac gaatcttttg agcctaatta 2460
gtccatgatt agccataagt gccatagtag cttagatata ctaatgacga attaattagg 2520
ctcaaaagat tcatctcgcg gttttcaggc aatttatgaa attagttttt tcattcatgt 2580
caaaaacccg tcgtgatatc cggtcaaaca tccaatatga caaaaaaaaa ttcttttcgc 2640
taaggctctt atgtaaaacg aacaacacgg tactccctct gtctaaaaaa aaaggcaaac 2700
cacgggtttg cgtgccaacg tttgactgtc cgtcttatat gaaatttttt tataattagt 2760
attttcatta ttgctagatg ataaaacatg atgaatattt tatgcgtgac ttatcttttt 2820
aatttttttc ataatttttt caaataaaac gaatggtcaa atgttagaca cggaaatcgg 2880
gttttttttt ttagacggag ggagtaggta ggagtaccag taataaaaac gaagaatacg 2940
cgttatagat ttctccacgg agggagaagc gaataaaatc ccccgcagca tatggcgtga 3000
cgaattgacg atatacaaat ttccacgggg accagtcttc ctcaatcctc tgccttgcat 3060
cttctccatc cagcgataca ataccagaag tccaaaaaac caccggtgcc gtatggccac 3120
agccggcgct gccgttgacc ggcttctacg ccgtctggcc tccggtgctg gccgtctgga 3180
gctgccctcg agcatagacg aggacatggc gcatgtaaag cgaaccctgg cgaggttgca 3240
agatgtgctg ctaaccgtag aagggaaata cttcaagatg ggcgcggagg tgcaggaatg 3300
gatgaggaag atcaagcaga ttgcttacgg cattcaagat ttgctggatg agtttgagga 3360
cagtagcggc accggatccc aaaggaacgg ctccaggatt tcagaggttc ggcttcggca 3420
catccattta tggaggggac aattagaatt accaacttgt cttttctcat tccaaatcag 3480
cattttctca aggaaaaact gttttcttta ataaaaaaaa aggggaaaat gggttctttt 3540
agtgaagtat ttcttatgcc agtgtccttg gatgccacaa taatcttttt gtagttctgt 3600
aaaaatatga tgattatata tttgtaaatg cccttcgagt gaccatttta gattattatt 3660
tgcagggaac gctatcgtgt tcgtcagctc catttttctg ccatcttagt agatcacaaa 3720
gaataagggt actgaaaaga aagttagatc aatcaacaaa agatacttct gtatttagtt 3780
tactgcagca cagcttatcc aatcttgaca aatccaatga gcaagaagtt ctgttacata 3840
gaactgaaat cattggaagg gatactgaca aagaaaatat aaaaaatcta ttgttacaaa 3900
atgatgtgga taaattaccc atcattccga tagttggcct tgcggggctg ggaaaaacgg 3960
ctgtagcaaa attgattttc catgaacagg gagaagggtg gaattttgat cagcgcatat 4020
gggtccattt ggacaagaaa ttggatctta acaaaattgc taacagtatt atctcacaag 4080
ttaaccaatc agtagatacc acaaagaatc aaattcagaa caacttacag tttaaaagga 4140
attgtcttca agaagttctt tgtgaccaaa gcagtttgat agtattggat gacttattta 4200
gcacagagga aaaccagatt gcagagttga aggaaatgtt gaggggtaca aagaagggaa 4260
ccaagatcat tgtgactact tccagtgaaa tatctgcaga gctaatacac acagttccac 4320
catacaagtt gggcccttta tctgaaggtg actgttcaac aatattttgt caaagagcat 4380
ttggtgatgg acatgaaaac agcagcctca ctgaaattgc gaagcaaatt gtgaaaaggt 4440
gtgaaggcat accggctgta gcttattctc ttggttcatt ggttcgtaac aagaataagg 4500
aggcctggtt atatgcaaga gacaaagaaa tatgggaatt accaacatta tttcctaatg 4560
ggtttgaatt acttgcatcg ttcagtgaaa tgtatatatg tatgccctcg gctttaaaat 4620
catgctttgc atacttatca accataccca aaggaacaat aattgatagg gagaaactta 4680
ttgaacagtg gatagcactt gacatggttg ggtcgaagca tgggacctta cctgcttatg 4740
tgcaaggaga gatgttcatc cagcaacttc tatcgatatc ttttcttcaa gtccgaaaca 4800
agccctctgt aagttgctca atatattgag ccaaaacctt ggctatgttt cgagatagtg 4860
ctaatataaa ttggcatgaa tagtaataat atatttttct gtcccttaaa atagtttttt 4920
ttttctatta ctgtcattca tgtgactcat tttgtttttt agtatgataa aatgctatat 4980
attctcaaaa gacaagaaaa actatatatt gttcataaat atttgttttt tttctgtgat 5040
atatctgcct gagtcttgag gtatatatgt cggctaagat agaaagttgg agctgaatag 5100
ctgatacagc gtgaaataag ggaggtagaa agcagaagct ggtataaatg tactattgat 5160
ttctagccat taactgtacc acgaaaagaa aattcatcat atataatgga ctcagtgatg 5220
ttgttcactg tgtatatttc ttttggatta cacatttcat gcatggtttt gtgggattat 5280
acaaaatcgc aagactgata agtaacccaa attcgaacaa ggtggttggt ttgaagcaaa 5340
tcaatgtgta acaaagtttt tcttttttca ggccaccaga atcagagaca ccaatcaatc 5400
taaggaactc cgtatccata acttggtcca tgactttgca atgtatgttg cccgtgatga 5460
tctcataatt ctggatggtg gagagaaggc cagtagcctt agaaaaaaca tccatgtctt 5520
ctatggagtt gtgaacaatg atattggaca atcagcactc cggaaaggtc tgctcagcag 5580
tgcaagggca gtacacttca agaactgtaa gtcagaaaag cttcttgtag aggcattctc 5640
agtactgaat catttgcgtg tcttggatct tagtggttgt tgtattgtag aattaccgga 5700
tttcattacc aatttgaggc atctgagata cctggatgtt tcatattcaa ggattctgtc 5760
attgtcaacc cagctaacta gtttgagtaa tctggaggta ttggatcttt cagaaacttc 5820
tcttgagttg ttaccatctt caattggctc atttgaaaaa ttaaaatact tgaatctaca 5880
aggatgtgat aaacttgtaa acttgccccc atttgtctgt gatctcaaga ggctagagaa 5940
tctcaaccta tcatactgtt atggaatcac tatgctaccc ccaaatctat ggaaacttca 6000
tgaacttcga attttggacc tctctagttg cacagatctt caagaaatgc catatttatt 6060
tggtaactta gcaagcttag aaaatctaaa catgtcgaaa tgctccaagc ttgaacaact 6120
accagaatct cttggtgatc tttgttacct acgatccttt aacctatcag gttgttctgg 6180
gcttaagatg ctgccagaat ctctgaaaaa tcttacaaat ttagagtata ttaatttgtc 6240
aaatattggg gagagtatcg atttcaatca gatacaacaa ctacggcaca ttctcaagaa 6300
aacatttttt tctggagata ttggagggag tgaactccaa acatgtgaac acgctgctga 6360
ttctgcagac agtaagaagg taatcttatc acatacatgg ttctgcaact aaacagaatg 6420
tagtgtaggt tttttctgtt tttttctttt tgatattttg attttgtgac tgatctactg 6480
tgtatacttt taccaatcaa ccatgtccac actaaaatag gtacttcaaa ctccttcaga 6540
aaactgttta tgtaaagaga cctagaccaa aagaataaca tggtttaagt ttctctgcag 6600
atcacatcat ccatgcatgc agaacatatg aaatgatctg acttgtcata gtgatatcta 6660
ggcaaaatac atcttttcaa aatatcacgt gtttcatggt tatagttgca aaagaccatc 6720
tttctaattt gcctagcttt cactaacatg aattattttg cagaagacac ttttcgtttt 6780
tatttcaatt tgcctgctat cactatcatg aattattttg catatattga tttaaagctt 6840
tttttttttt gccaacatac aactagagaa cataattaca tgtaaaagat attgaagagg 6900
cagtatttag attgtcatgt acttgtttgg tatttattga ggaataataa taaaaaatgt 6960
tggttaattt gaggtacttt tttattcgag atttgaggta ttttttaatt cattatttcc 7020
atctctactc taacttagat ttcatgaaaa ttattttcag gaaattacaa tggatttttc 7080
tgcaaatttg catggaaata ttactttgcc acccaagtgc tcaactggta catttgcttg 7140
agttgtttat ataaacaatt tttctttcga tacatttttc ataataaata tacgaaattg 7200
cattcaagtc gtgtaatttg tctattaatt ttgtgccacc aaggaataat gaatattttg 7260
aaacttccta tgttaattta gttgagtgtt tttattgtgt tctagtcctc atggagctct 7320
ttgggcatgt tccatgaatt tatagatgta tcatcatata aataatatga ctgcaatccc 7380
tttattgcat aatatagtct ggaactaagc ttaggttgtg tttttacttg aagttgggaa 7440
ctaatccctc tctatcacaa aacaaaacga ctcattagca catgattaat taagtattag 7500
atattttttt tgaaaaatag attaatataa tttattaaag caactttcgt atagaatttt 7560
tttgcaaaaa aaaaacacat catatagtaa tttgaaaaac gttttcacag aaaacgaggg 7620
agatgagtcg ggatcttcca ccaaagaact cagccttgta aattttgtta tataatattg 7680
tattctttgc aaacatcgta tcatatcagt gctctccata tcctttatat atatataaaa 7740
tgtagtgaaa tatgttgttt cataacataa atccattgcg gactatcttt gatactacct 7800
ccgtttcagg ttataagacg ttttgacttt agttaaagtc aaactactct aactttgact 7860
aactgtatag aaaaaatagt aatatttaca acaccagcat agtttcatta aatctataat 7920
tgaataaatt ttcataatat attagtcttg ggttaaaaat atgactactt ttttctacaa 7980
aattagttaa acttagagtt gtttgacttt gacaaaagtc aaaacgtcta taccctgaac 8040
cggaggggag tattagagtt gatgagaaga ttaaaaataa aggggtcaca tagtaccact 8100
agtcttaatg ggggtttaca tacacacatt atgcgaacta tataccatta tatctacaca 8160
cacacacaca caaacacaaa agaagttttt atctccaact tctttttttt tttgcgggga 8220
aaggaaatat attattagaa tttactaact gtagcccata cagaaaatca ttattctgta 8280
attacgtacc gatgtgcaaa ttctaataaa tatctataca tcttcttttg caactagcag 8340
aagagaaatc tggtgaaaac tctgaacgat tcctatcagc tgcagttagg gaggattcaa 8400
gcagcactga tgtttctaca tatgtcaagc cggtggtgtc ctcacttatt ggagtgctgc 8460
gcagaccgac taggttggat gtgccagctg gcgcgatggc ttctcaggtt ggcctggccc 8520
agatgccatc tagcaacaat ggtaaggctg gaccgcatcc aacaatggct gcagctcaga 8580
ccccggagat tgatcaacct gtgcataaaa gggtccggtg ggatgatata atagactact 8640
cccgtcctcc caactcaaaa cctgccagga gtgcatctct tgtgcagtcg accgatctgt 8700
cgacaccaaa gaaaagttat aaaaaaattc actcaatgcc ggtggtctat tcgtcaattc 8760
caaaagggag ttccggagga acatacttga tgccagcaaa ggctatcgct tcttcctaca 8820
ggaggtatag cccacagaga tgggaacagc acattggtta tcaaggaacg gtgcgttcat 8880
tgtaattatt ccagaatata tatatgtgac ttgatatata tctccatttg attggaacac 8940
aggagtaata tatagctggt tttgcatttc ttacctggcc aggatgaaga tgaactgatg 9000
gtcgtgccac catttggtga gtgggatcag tcccccacat tacgaaaatc tgactttcgg 9060
tatgagaagg tcttcgctaa actcaccgaa gagaagatgt ctggtcaaag gcagaaaccg 9120
caacaggtat gaacaaactg ccaccaaaca gaacgtaaat aggcatgctc tgctgtttgc 9180
cagaattaaa tttcattgac tgctattgaa catatatata cttagtgcta atcaccaatt 9240
tgtgacatat aaaggtctca actatcaata aaaaaatcat gcaggctaac aataccaaaa 9300
aaaaaggaga aaaaatgaaa aaaaaattgg taaccaaaat tatagaggct gacattatca 9360
gttctcacca atcagtgatc ttgaaagaca caaaaccagt gccggagctt catggagatc 9420
aaagtgggtt ctggccccct ccaactccat gcaaattata ctatatgtat gtttatgttc 9480
ataaatacta ttcattagcc atctttcacg ttgattgaca taaaactcat gttaatggcc 9540
ccctcttaat agtttgtgaa gctgcgcata aaatcatctt gacaaatttt tggtaacaac 9600
tgggaagtgc taattatcaa tactgaacgt tctgaacaaa tcaaccattt atagacaatg 9660
caagagcaaa aaagaaatta gtatacccac aatatatttt gttgtaccct atctgagatt 9720
ctaggttgtc tattgtgaat ctgtttgtta ccaatcagtg aaatttagtc gtcctaaaca 9780
tcaaataaat ttataaattt ataaaaaata atttatgaac tggggaagta tatgttcaat 9840
tactctgcaa cgttatatat tgttatacta taaaatcacc atgatcaatg gcaataccct 9900
tctcttcttc cccagataac ttggtaactt taatatgttt attcttggct ttttttcgac 9960
aatactccct ccgtcccccc aaaaaaaaaa actcaattcc ttggtttccg tatctaacgt 10020
ttgaccgtcc gtcttatttg aaaaaattat gaaaaaaatt aaaaagataa gtcatgcata 10080
aaatattatt catgttttat catctaacaa taataaaaat acaatttata aaaaaatttt 10140
atataagacg gatggtcaaa tgttagacgc agaaatccaa gaattgagtc ttttttggga 10200
tggataatac aagggatttt gactttttag ttgtaatgtt tgaccactcg tcttattcaa 10260
aaaatttgtg caaatataaa aaacgaaaaa ttgtgcttaa agtattttgg ataataaagt 10320
aagccacaaa taaaataaat aatagttcta atttttttta ataagacgaa tggtcaaaca 10380
gtgcaaacaa aaagtcaaaa tccctacatt attttttttg agacaaaaat ccctacatta 10440
taggacggag ggagtagcta gatatacaga gatctatcca tatctagact ttagagtcgt 10500
gcgtacgaac ataccttttt cccgatggta atcgaattcg aatcgtagtc cagtacacta 10560
ctcctacctt tattttccgg tttcgcgggc ttcctttcca gtatcgggca tctacagggt 10620
ggtggtggtc agttactcac tcggtcagat cagggcgtct ctatttgctc cgtcatgaaa 10680
aaaaaattat atttttcttt gtgtgtgttt tctgatcagg tgacactgcg attgtcctct 10740
tgcagagaaa accacggaat gagcgatttg gcgcggtgtg gcatgccgaa ctcctgcaaa 10800
aaagatgaac taactcaacg aaggcagcga taacggcggc ggtcgctcac tctcctcaat 10860
aattgagcaa ccggcttccg ttgttttttt tttctcgtat ttttaataat tttattcttt 10920
tattgaaatg ggggttgggc acagcgcgcg atgtacttgt gcgccacaca cttgaatttg 10980
atcacggtcg atcttctatg tcggtacgag aatccacgga tgaagattaa atacaagcac 11040
gaaaaaaaag agagtattag ctaatgatta attaaatttt aattatttaa aacttgaaag 11100
atgtatttat ttgatatttt aaagcaactc tgtttttgca cgaaatatat cgtttagcag 11160
tttgaaaaac atactaacgg aaactaaggt agaatatgta tcttaatcag aaaaagaata 11220
gattttataa aaccttacat tttatggtgc tttgaaaaac cacacatatt ataactcgtg 11280
atacataact acaagtttat agtctaaatg tttcacaaga gcccaccttt atttaaattt 11340
ccattgattt cagcatatct gaaatgatat tttttataag tagtgaatag ttaaaccgat 11400
aac 11403
<210>5
<211>3078
<212>DNA
<213>水稻
<400>5
atggttggcg ccgagatgct tgtggccgcg gcggtgagcc aggtcgcccg gaagatcaac 60
gacatcgtgg gggtcgcgca gggcgaggtg aagctgtgct gcaatttcag cgacgatttg 120
gagggcatca aggataccct tgtgtacctg gaaaccttgc tgaaaaatgc ggagaataac 180
tccttcggaa gcgacagggc caacctgcgc cactggcttg gccagatcaa gtccctggct 240
tacgatatcg aagatatcgt tgatgggtac tactcttcca aggagcagtt cgatgggggc 300
agctatgcac agaaggggtc attattctgc tcgctatcca atccaatgct tctgaaaggt 360
agcatggttt ataagatgaa atccaagaga gagatgctac agcaaagcca acagttgccc 420
aatcagtatc atttcctttc atatatcaat tcagctgtgc attattttga ggagaagcaa 480
acaacatcat acagaaatac tgacattgca attgtcggga gggatgctga tttggatcat 540
ctcatggatc ttttaatgca aaacagcgct gaagagcttt gtattatacc catagttggg 600
cctgtaggtt ttggaaagac aagccttgca cagttagttt tcaatgatac aagaacagag 660
gtattcagct ttaggatatg ggttcatgtt tccatgggta atatcaacct tgaaaaaatt 720
gggagagata tagtttcaca aactacagaa aaaattgagg gaaatatgca gctgcagtca 780
atcaagaatg ctgttcagcg tgtgctaaat aaatatagtt gcttgatcat aatagacagc 840
ctttggggaa aggatgaaga agtgaatgaa ttgaagcaga tgttgcttac aggtagacac 900
acagaaagca agatcatagt gaccactcat agcaataaag tagctaagct gatttccacc 960
gttccactgt acaagttggc agctttatct gaggatgatt gtttaaaaat attctctcaa 1020
agggcaatga caggtccggg tgacccgttg ttcagggaat atggagaaga aatcgttaga 1080
aggtgtgaag gcacaccctt ggtagccaat tttctcggtt ctgtggtgaa tgctcaacga 1140
caaaggcgtg agatttggca agctgcaaag gatgaagaaa tgtggaagat agaggaagat 1200
tatccccaag acaaaatttc accactattt ccatcattca agataatata ttataatatg 1260
ccccatgagc taagattatg ctttgtatat tgttcaatct tccctaaagg aactgttata 1320
gaaaagaaga aacttattca gcaatggatt gcacttgaca tgattgagtc caaacatgga 1380
accttgccac ttgatgtaac tgcggagaaa tatattgatg aacttaaagc aatctatttc 1440
cttcaagttt tagagcggtc tcagaatgat gcagaaagat ccagtgcttc tgaggaaatg 1500
cttcgcatgc ataacttggc tcatgatctt gctagatcgg ttgctggtga agatatcctt 1560
gttattttag atgccgagaa tgagcgcaat gccagatatt gcaattaccg ttatgcacag 1620
gtgtctgctt ctagtttaga gtcaatcgat cgcaaggcat ggccttccaa ggcaaggtca 1680
ctaattttca agaatagtgg tgtggacttt gagcatgtca gtgaagttct ttcagtgaac 1740
aaatacctgc gtgttttgga tctcagtgga tgttgtgttc aagatattcc atctcctatc 1800
tttcagctga aacaattgag atacctcgac gtttcatctt tatctattac agcactccct 1860
ctgcaaatta gtagctttca taagttacaa atgttggatc tttcagaaac tgaactaaca 1920
gagttgccac cctttataag caacttaaaa ggactgaatt atttgaatct ccaaggttgc 1980
cagaaacttc aacgattgaa tagccttcat ttgttgcatg atctacatta cctaaacttg 2040
tcatgctgcc ctgaagttac tagttttcct gaatctattg aaaatctgac caaactccgt 2100
ttcttgaatc tttctggatg ctctaagctt tcaacattac ctatcagatt tttggaatca 2160
tttgctagcc tctgttcttt ggtagatctt aacttaagtg gctttgaatt ccaaatgttg 2220
cccgactttt ttggcaacat atattcactt cagtatttaa atctgtcaaa atgtttgaaa 2280
cttgaggtat taccacaatc ttttggccaa cttgcatatc tgaaaagcct aaatctttca 2340
tattgttctg atcttaaact gctggaatcc tttgaatgcc ttacctctct tcggtttttg 2400
aatctctcga actgctctag gcttgaatat ttgccatcat gctttgacaa gcttaataat 2460
ttagagtctc tgaatttatc acaatgtctt ggacttaaag cactacctga atcacttcaa 2520
aaccttaaaa atcttcagct tgatgtttct gggtgtcagg attgtatagt acaatccttt 2580
tctctaagta ccagaagttc ccagtcctgc caacggtcgg agaaagctga gcaggtcaga 2640
tcaagaaaca gtgaaatttc agagatcact tatgaggaac ctgctgagat tgaactttta 2700
aagaataatc caagtaaaga tttggcctcc atctcacacc taaatgagga tagaattgag 2760
gagcctgaag ttgtcactga gccaagtgca actagaggta tggtacaaca gattccagga 2820
aaccagctct catcgccttc atctcatctt tcttcctttg catcaagctc agcgccattt 2880
gcatcctcct cttcggacac ctcaacaagt gagcatccag tgcctaatga agaggcggca 2940
gctttgacag ttcctcggtc caaagagaaa tgcgacaaca ctcccatgcc ggtaaaagat 3000
ggcctgatat ctgaagatga tgcaccggta catctgcatc agaagcccct gcaggcgaca 3060
gccatggcag ccatatga 3078
<210>6
<211>3192
<212>DNA
<213>水稻
<400>6
atggccacag ccggcgctgc cgttgaccgg cttctacgcc gtctggcctc cggtgctggc 60
cgtctggagc tgccctcgag catagacgag gacatggcgc atgtaaagcg aaccctggcg 120
aggttgcaag atgtgctgct aaccgtagaa gggaaatact tcaagatggg cgcggaggtg 180
caggaatgga tgaggaagat caagcagatt gcttacggca ttcaagattt gctggatgag 240
tttgaggaca gtagcggcac cggatcccaa aggaacggct ccaggatttc agagggaacg 300
ctatcgtgtt cgtcagctcc atttttctgc catcttagta gatcacaaag aataagggta 360
ctgaaaagaa agttagatca atcaacaaaa gatacttctg tatttagttt actgcagcac 420
agcttatcca atcttgacaa atccaatgag caagaagttc tgttacatag aactgaaatc 480
attggaaggg atactgacaa agaaaatata aaaaatctat tgttacaaaa tgatgtggat 540
aaattaccca tcattccgat agttggcctt gcggggctgg gaaaaacggc tgtagcaaaa 600
ttgattttcc atgaacaggg agaagggtgg aattttgatc agcgcatatg ggtccatttg 660
gacaagaaat tggatcttaa caaaattgct aacagtatta tctcacaagt taaccaatca 720
gtagatacca caaagaatca aattcagaac aacttacagt ttaaaaggaa ttgtcttcaa 780
gaagttcttt gtgaccaaag cagtttgata gtattggatg acttatttag cacagaggaa 840
aaccagattg cagagttgaa ggaaatgttg aggggtacaa agaagggaac caagatcatt 900
gtgactactt ccagtgaaat atctgcagag ctaatacaca cagttccacc atacaagttg 960
ggccctttat ctgaaggtga ctgttcaaca atattttgtc aaagagcatt tggtgatgga 1020
catgaaaaca gcagcctcac tgaaattgcg aagcaaattg tgaaaaggtg tgaaggcata 1080
ccggctgtag cttattctct tggttcattg gttcgtaaca agaataagga ggcctggtta 1140
tatgcaagag acaaagaaat atgggaatta ccaacattat ttcctaatgg gtttgaatta 1200
cttgcatcgt tcagtgaaat gtatatatgt atgccctcgg ctttaaaatc atgctttgca 1260
tacttatcaa ccatacccaa aggaacaata attgataggg agaaacttat tgaacagtgg 1320
atagcacttg acatggttgg gtcgaagcat gggaccttac ctgcttatgt gcaaggagag 1380
atgttcatcc agcaacttct atcgatatct tttcttcaag tccgaaacaa gccctctgcc 1440
accagaatca gagacaccaa tcaatctaag gaactccgta tccataactt ggtccatgac 1500
tttgcaatgt atgttgcccg tgatgatctc ataattctgg atggtggaga gaaggccagt 1560
agccttagaa aaaacatcca tgtcttctat ggagttgtga acaatgatat tggacaatca 1620
gcactccgga aaggtctgct cagcagtgca agggcagtac acttcaagaa ctgtaagtca 1680
gaaaagcttc ttgtagaggc attctcagta ctgaatcatt tgcgtgtctt ggatcttagt 1740
ggttgttgta ttgtagaatt accggatttc attaccaatt tgaggcatct gagatacctg 1800
gatgtttcat attcaaggat tctgtcattg tcaacccagc taactagttt gagtaatctg 1860
gaggtattgg atctttcaga aacttctctt gagttgttac catcttcaat tggctcattt 1920
gaaaaattaa aatacttgaa tctacaagga tgtgataaac ttgtaaactt gcccccattt 1980
gtctgtgatc tcaagaggct agagaatctc aacctatcat actgttatgg aatcactatg 2040
ctacccccaa atctatggaa acttcatgaa cttcgaattt tggacctctc tagttgcaca 2100
gatcttcaag aaatgccata tttatttggt aacttagcaa gcttagaaaa tctaaacatg 2160
tcgaaatgct ccaagcttga acaactacca gaatctcttg gtgatctttg ttacctacga 2220
tcctttaacc tatcaggttg ttctgggctt aagatgctgc cagaatctct gaaaaatctt 2280
acaaatttag agtatattaa tttgtcaaat attggggaga gtatcgattt caatcagata 2340
caacaactac ggcacattct caagaaaaca tttttttctg gagatattgg agggagtgaa 2400
ctccaaacat gtgaacacgc tgctgattct gcagacagta agaaggaaat tacaatggat 2460
ttttctgcaa atttgcatgg aaatattact ttgccaccca agtgctcaac tgaagagaaa 2520
tctggtgaaa actctgaacg attcctatca gctgcagtta gggaggattc aagcagcact 2580
gatgtttcta catatgtcaa gccggtggtg tcctcactta ttggagtgct gcgcagaccg 2640
actaggttgg atgtgccagc tggcgcgatg gcttctcagg ttggcctggc ccagatgcca 2700
tctagcaaca atggtaaggc tggaccgcat ccaacaatgg ctgcagctca gaccccggag 2760
attgatcaac ctgtgcataa aagggtccgg tgggatgata taatagacta ctcccgtcct 2820
cccaactcaa aacctgccag gagtgcatct cttgtgcagt cgaccgatct gtcgacacca 2880
aagaaaagtt ataaaaaaat tcactcaatg ccggtggtct attcgtcaat tccaaaaggg 2940
agttccggag gaacatactt gatgccagca aaggctatcg cttcttccta caggaggtat 3000
agcccacaga gatgggaaca gcacattggt tatcaaggaa cggatgaaga tgaactgatg 3060
gtcgtgccac catttggtga gtgggatcag tcccccacat tacgaaaatc tgactttcgg 3120
tatgagaagg tcttcgctaa actcaccgaa gagaagatgt ctggtcaaag gcagaaaccg 3180
caacaggtat ga 3192
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Claims (6)
1.一种增强对稻瘟病菌(Magnaporthe oryzae)的抗性的蛋白组合物,所述蛋白组合物由Pi5-1蛋白和Pi5-2蛋白组成,其中,Pi5-1蛋白的氨基酸序列如SEQ ID NO:1所示,Pi5-2蛋白的氨基酸序列如SEQ ID NO:2所示。
2.一种用于增强对稻瘟病菌(Magnaporthe oryzae)的抗性的基因组合物,所述基因组合物由编码权利要求1所述Pi5-1蛋白的基因和所述Pi5-2蛋白的基因组成。
3.根据权利要求2所述的基因组合物,其特征在于,所述Pi5-1蛋白的基因组DNA由SEQ ID NO:3的核苷酸序列组成,并且所述Pi5-1蛋白的cDNA由SEQ ID NO:5的核苷酸序列组成;所述Pi5-2蛋白的基因组DNA由SEQ ID NO:4的核苷酸序列组成,并且所述Pi5-2的cDNA由SEQ ID NO:6的核苷酸序列组成。
4.一种重组体载体,其中,该重组体载体包括权利要求2中的Pi5-1蛋白基因和Pi5-2蛋白的基因。
5.一种增加对稻瘟病菌(Magnaporthe oryzae)的抗性的方法,其中,该方法包括如下步骤:用权利要求4中的重组体载体转化植物,并继而在该植物中使Pi5-1蛋白和Pi5-2蛋白的基因表达。
6.根据权利要求5所述的方法,其特征在于,所述植物为单子叶植物。
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KR1020080069820A KR100990370B1 (ko) | 2008-07-18 | 2008-07-18 | 벼 도열병균에 대한 내성을 증진시키는 유전자 및 이의용도 |
PCT/KR2009/003898 WO2010008204A2 (en) | 2008-07-18 | 2009-07-15 | Genes for enhancing resistance to magnaporthe oryzae and uses thereof |
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AU2012242991B2 (en) | 2011-04-11 | 2017-03-02 | Targeted Growth, Inc. | Identification and the use of KRP mutants in plants |
CN102776209A (zh) * | 2012-06-20 | 2012-11-14 | 浙江大学 | 源于稻瘟病菌的真菌致病性基因MoMon1及其用途 |
KR101642694B1 (ko) * | 2014-06-03 | 2016-07-27 | 경희대학교 산학협력단 | 벼 유래 NB-LRR 면역 센서 저항성 유전자인 OsWIR1(OsWRKY67-inducible resistance gene1)을 이용한 식물의 병 저항성을 증가시키는 방법 및 그에 따른 식물체 |
CN113875579B (zh) * | 2021-10-22 | 2023-03-24 | 宁波市农业科学研究院 | 一种抗稻瘟软香粳型水稻的育种方法 |
WO2023200023A1 (ko) * | 2022-04-12 | 2023-10-19 | 세종대학교산학협력단 | 벼 도열병 저항성 증진 또는 개선용 조성물 및 벼 도열병 저항성 증진 또는 개선 방법 |
CN116987813B (zh) * | 2022-11-02 | 2024-02-13 | 江苏省农业科学院 | 稻瘟病多重抗病基因组合Pita+Pi5+Piz-t及其应用 |
CN117947094B (zh) * | 2024-03-26 | 2024-06-04 | 云南农业大学 | Pi-Pprs42基因提高水稻稻瘟病抗性的方法及应用 |
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WO2010008204A3 (en) | 2010-06-03 |
KR100990370B1 (ko) | 2010-10-29 |
US20100287664A1 (en) | 2010-11-11 |
CN101932710A (zh) | 2010-12-29 |
KR20100009107A (ko) | 2010-01-27 |
WO2010008204A2 (en) | 2010-01-21 |
US8389803B2 (en) | 2013-03-05 |
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