CN114790458B - 一种水稻稻瘟病非小种专化抗性基因qbr1、分子标记及应用 - Google Patents
一种水稻稻瘟病非小种专化抗性基因qbr1、分子标记及应用 Download PDFInfo
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- C12N15/8271—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield for stress resistance, e.g. heavy metal resistance
- C12N15/8279—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield for stress resistance, e.g. heavy metal resistance for biotic stress resistance, pathogen resistance, disease resistance
- C12N15/8282—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield for stress resistance, e.g. heavy metal resistance for biotic stress resistance, pathogen resistance, disease resistance for fungal resistance
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- Breeding Of Plants And Reproduction By Means Of Culturing (AREA)
Abstract
本发明属于生物技术领域，具体涉及一种水稻稻瘟病非小种专化抗性基因QBR1、分子标记及应用。本发明获得一种具有水稻稻瘟病非小种专化抗性的新基因QBR1。本发明还公开了QBR1基因的cDNA全长和氨基酸序列；本发明的QBR1基因可以用于培育抗病能力增强的植物新品种，或制备抗病能力增强的转基因植物。本发明还公开了一种QBR1基因的分子标记。本发明的QBR1基因和分子标记有利于对农作物品种抗病原物侵染能力的改良，对扩大作物种植范围，提高作物产量具有重要意义。
Description
技术领域
本发明属于生物技术领域；更具体地，本发明涉及一种水稻稻瘟病非小种专化抗性基因QBR1、分子标记及应用。
背景技术
水稻是我国乃至全世界最重要的粮食作物之一，水稻稻瘟病是由子囊菌稻瘟菌(Magnaporthe grisea)引起的广泛发生在世界各稻区的最重要的病害之一，每年造成全球10％-30％的水稻减产。我国的稻瘟病危害也相当严重，每年都因稻瘟病危害造成粮食生产重大损失，这与人口增加和耕地减少而带来的粮食增产压力形成尖锐矛盾。目前，防治稻瘟病的方法主要为化学防治和应用抗病品种。相对于化学防治方法，利用抗病品种更为经济有效，而且可以避免因大量使用农药而造成的对环境的危害和对自然界生物多样性的不良影响。然而，由于稻瘟病真菌生理小种演化迅速，部分抗病新品种在推广数年内抗性就会明显衰退，抗病无法持久化。因此改良水稻稻瘟病抗性需要发掘和利用新的稻瘟病菌生理小种抗谱不同的种质或基因资源。
由于稻瘟病危害的严重性，稻瘟病抗性基因遗传研究一直是分子生物学研究的热点。到目前为止，已经有27个抗稻瘟病基因被先后克隆和鉴定。它们分别为Pia，Pib，Pita，Pi1,Pi2，Pi9，Piz-t，Pid2，Pid3，Pid3-A4，Pi5，pi21，Pi25，Pi35，Pi36，Pi37，Pi50，Pi63，Pii，Pit，Pish，Pik，Pi-CO39，Pik-KA，Pikh/Pi-54，Pikm，Pikp、Pb1。这些抗性基因分布在除水稻基因组第3染色体以外的其余11条染色体上。在水稻NBS-LRR基因家族中，基因呈串连重复分布于同一遗传座位是一种非常普遍的现象。在已克隆的上述27个抗病基因中，除Pb1，Pi-ta，Pid2和Pi36位于单基因座位上，其余23个基因分布于10个多基因座位上。主要包括1号染色体的Pi37座位、6号染色体的Pi2/9座位、9号染色体的Pi5座位、11号染色体的Pik座位。其中，位于靠近水稻6号染色体着丝粒区域的Pi2/9遗传座位，已经克隆了Pi2，Pi9，Piz-t和Pi50等抗性基因，目前还发现Piz、Pigm、Pi26、Pi40、Pi2-1、Pi2-2等基因也都位于该位点，它们分别来源于不同的抗病品种，并且多数具有广谱抗性。
根据是否依赖于稻瘟菌无毒基因，水稻抗稻瘟病基因可以分为小种专化性(racespecific)和非小种专化性(non-race specific)抗病基因。小种专化性抗稻瘟病基因虽然表现为完全抗性，但也容易因为稻瘟病菌生理小种的变化而丧失抗性。相对小种专化性抗病基因而言，非小种专化性抗病基因一般属于数量性状，效应较小而表现部分抗性，它不能阻止稻瘟病菌的侵染，但具有抗扩展和减轻病害的作用，因其降低了对病原菌的选择压而呈现广谱持久抗病的特点。因此，非小种专化性抗病基因在水稻抗稻瘟病育种中具有极高的利用价值。目前，已克隆的非小种专化性抗稻瘟病基因有pi21，Pi35，Pb1，均由日本科学家克隆。pi21是唯一被克隆的隐性抗稻瘟病基因，位于第4染色体上2个分子标记G271～G317之间。pi21是一个非小种专化性抗病基因，它激发的是一种慢速抗病反应，而这种低速诱导的抗病反应可能是一种新的持久抗病反应机制。Pb1基因来源于部分抗性籼稻品种Modan，编码一个包含1296个氨基酸的典型CCNBS-LRR类抗病蛋白。Pb1在成株期抗性最强，在不同生育期其抗性的强弱与基因表达水平的高低有关。Pi35是最近被克隆的一个稻瘟病小种非专化性抗病基因，位于1号染色体的Pi37座位。在该座位上的3个基因Pi37，Pish和Pi35序列高度一致，彼此间只有几个至十几个氨基酸残基的差异。Pi35基因的克隆提供了一个很重要的提示，就是在小种专化性抗病基因位点，通过序列变异，可以进化出非小种专化性抗病基因。携带Pi35基因的粳稻品种北海188和藤系138，在日本30多年来一直保持稳定的高水平叶瘟抗性，进一步证明了非小种专化性抗病基因对稻瘟病的持久稳定抗性。然而，非小种专化性抗病基因克隆数目有限，需要克隆更多的基因资源以提供给抗病分子育种应用；同时，对非小种专化性抗病基因的抗性机制基础理论研究还非常不够，需要进一步深入。
发明内容
为了解决现有技术中稻瘟病非小种专化性抗病基因数量不够的技术问题，本发明提供以下技术方案：
一种水稻稻瘟病非小种专化抗性基因QBR1，其核苷酸序列全长如SEQ ID NO:1所示。
一种水稻稻瘟病非小种专化抗性基因QBR1的cDNA，所述cDNA的核苷酸序列如SEQID NO:2所示。
非小种专化抗性基因QBR1或非小种专化抗性基因QBR1的cDNA编码的蛋白质，所述蛋白质的氨基酸序列如SEQ ID NO:3所示。
非小种专化抗性基因QBR1、非小种专化抗性基因QBR1的cDNA在提高水稻稻瘟病抗性中的应用。QBR1蛋白在提高水稻稻瘟病抗性中的应用。
优选地，所述稻瘟病抗性为子囊菌稻瘟菌抗性。
一种培育抗稻瘟病水稻品种的方法，所述方法为：将非小种专化抗性基因pi21、Pi35中至少一个基因与非小种专化抗性基因QBR1或非小种专化抗性基因QBR1的cDNA在水稻中聚合。
一种水稻稻瘟病非小种专化抗性基因QBR1的分子标记，所述分子标记包括下列七组引物组中至少一种:
第一组引物组序列包含SEQ ID NO:4和SEQ ID NO:5
SEQ ID NO:4(16640-F):5’-TGGCGACAATAAGCTTGGGA-3’
SEQ ID NO:5(16640-R):5’-GGCTACTACTCCCTCCGTCC-3’；
第二组引物组序列包含SEQ ID NO:6和SEQ ID NO:7
SEQ ID NO:6(17000-F):5’-ACGGGTAGAGCTGCCAAATC-3’
SEQ ID NO:7(17000-R):5’-TAGCGAACTCAACCACCGAC-3’；
第三组引物组序列包含SEQ ID NO:8和SEQ ID NO:9
SEQ ID NO:8(17570-F):5’-TCATCGTCATCACCGGCTTC-3’
SEQ ID NO:9(17570-R):5’-TTGAGGGAGGGCTTTCAACG-3’；
第四组引物组序列包含SEQ ID NO:10和SEQ ID NO:11
SEQ ID NO:10(Nbs1-1F):5’-GACAGACAAACGTTGGGCAC-3’
SEQ ID NO:11(Nbs1-1R):5’-AAGATCAGCGTGTGACCGTT-3’；
第五组引物组序列包含SEQ ID NO:12和SEQ ID NO:13
SEQ ID NO:12(Nbs7-2F)：5’-TGGAAAACAGAGTCCTCGGC-3’
SEQ ID NO:13(Nbs7-2R)：5’-GGCAAAGATTTGCGTCCTCC-3’；
第六组引物组序列包含SEQ ID NO:14和SEQ ID NO:15
SEQ ID NO:14(1066-F):5’-GAAGGTGAAGGAGATGATGG-3’
SEQ ID NO:15(1066-R):5’-GGTTCAAAACTACGCGGAAG-3’；
第七组引物组序列包含SEQ ID NO:16和SEQ ID NO:17
SEQ ID NO:16(19010-F):5’-TGTACAGGCCAGCCATTCAG-3’
SEQ ID NO:17(19010-R):5’-CAATCGTCGTGTTCATCGCC-3’。
一种水稻稻瘟病非小种专化抗性基因QBR1分子标记在种质资源选育抗稻瘟病水稻中的应用。
本发明获得一种具有水稻稻瘟病非小种专化抗性的新基因QBR1。本发明还公开了QBR1基因的cDNA全长和氨基酸序列；本发明的QBR1基因可以用于培育抗病能力增强的植物新品种，或制备抗病能力增强的转基因植物。本发明有利于对农作物品种抗病原物侵染能力的改良，对扩大作物种植范围，提高作物产量具有重要意义。
附图说明
图1为QBR1基因内标记与抗性表型共连锁关系。
图2为BC4F4重组子基因分型与表型结果。
图3为QBR1位点结构图。
图4为部分生理小种对QBR1的接种结果。
图5为QBR1基因编辑敲除突变体丧失抗性。
图6为3个小种非专化抗性基因导入系与聚合系抗性表现。
具体实施方式
为使本发明的目的、技术方案和优点更加清楚，下面将结合本发明实施例，对本发明实施例中的技术方案进行清楚、完整地描述，显然，所描述的实施例是本发明一部分实施例，而不是全部的实施例。基于本发明中的实施例，本领域普通技术人员在没有作出创造性劳动前提下所获得的所有其他实施例，都属于本发明保护的范围。
实施例1QBR1是一个不同于Pi2/9位点已克隆基因的新抗病基因
对400份种质资源进行Pi2/9位点等位基因挖掘，发现交流自国际水稻研究所的种质资源CO39-A35中，其Pi2/9位点存在1个功能等位基因((命名为QBR1，其核苷酸序列如SEQID NO:1所示，其cDNA的核苷酸序列如SEQ ID NO:2所示，其氨基酸序列如SEQ ID NO:3所示))。通过下列7组引物序列至少一种以上作为分子标记辅助；
第一组引物组序列包含SEQ ID NO:4和SEQ ID NO:5
SEQ ID NO:4(16640-F):5’-TGGCGACAATAAGCTTGGGA-3’
SEQ ID NO:5(16640-R):5’-GGCTACTACTCCCTCCGTCC-3’；
第二组引物组序列包含SEQ ID NO:6和SEQ ID NO:7
SEQ ID NO:6(17000-F):5’-ACGGGTAGAGCTGCCAAATC-3’
SEQ ID NO:7(17000-R):5’-TAGCGAACTCAACCACCGAC-3’；
第三组引物组序列包含SEQ ID NO:8和SEQ ID NO:9
SEQ ID NO:8(17570-F):5’-TCATCGTCATCACCGGCTTC-3’
SEQ ID NO:9(17570-R):5’-TTGAGGGAGGGCTTTCAACG-3’；
第四组引物组序列包含SEQ ID NO:10和SEQ ID NO:11
SEQ ID NO:10(Nbs1-1F):5’-GACAGACAAACGTTGGGCAC-3’
SEQ ID NO:11(Nbs1-1R):5’-AAGATCAGCGTGTGACCGTT-3’；
第五组引物组序列包含SEQ ID NO:12和SEQ ID NO:13
SEQ ID NO:12(Nbs7-2F)：5’-TGGAAAACAGAGTCCTCGGC-3’
SEQ ID NO:13(Nbs7-2R)：5’-GGCAAAGATTTGCGTCCTCC-3’；
第六组引物组序列包含SEQ ID NO:14和SEQ ID NO:15
SEQ ID NO:14(1066-F):5’-GAAGGTGAAGGAGATGATGG-3’
SEQ ID NO:15(1066-R):5’-GGTTCAAAACTACGCGGAAG-3’；
第七组引物组序列包含SEQ ID NO:16和SEQ ID NO:17
SEQ ID NO:16(19010-F):5’-TGTACAGGCCAGCCATTCAG-3’
SEQ ID NO:17(19010-R):5’-CAATCGTCGTGTTCATCGCC-3’；
将上述该Pi2/9的功能等位基因QBR1导入到易感稻瘟病品种CO39中构建近等基因系。对BC3F3代株系接种，所有28个测试稻瘟菌菌株都表现出部分抗性或部分抗性与感病的分离(表1)，连锁分析进一步证实了抗性与QBR1基因标记共连锁关系(图1)，上述结果表明QBR1是一个非小种专化性抗稻瘟病基因，这明显区别于该基因所在的Pi2/9位点目前已克隆和定位的小种专化性等位基因。
表1 QBR1基因BC3F3代株系对28个测试菌株都有部分抗性的表现
注：“R”、“PR”分别表示“完全抗性”和“部分抗性”
实施例2QBR1基因的精细定位
2017年通过遗传连锁分析获得BC4F2-51,BC4F2-186,BC4F2-303和BC4F2-568四个重组子植株，在海南加代，对其进行接种后表型验证，结果表明4个重组子都表现抗病。因此选取杂合区段最长的BC4F2-186后代进行进一步重组子筛选。我们首先进行了精细定位标记开发，得到分别位于物理距离9.7Mb、9.86Mb、10.0Mb、10.2Mb、10.3Mb、10.45Mb、10.66Mb、11.0Mb位置的多态性标记。利用这些标记对BC4F2-186子代共483个单株基因组DNA进行分析，筛选出11-1,15-6,17-5,30-6,38-2,39-4,44-4,57-4,61-4,71-1共10个重组子。其中，57-4单株整体表型完全与亲本不一致，推断为杂株。对另外9个重组子单株进行基因型分析，并收获子代种子进行稻瘟病小种接种(表1)。基因型与表型结果对照分析表明，QBR1基因位于10.21Mb-10.66Mb之间450kb的物理区间内。选取该区段处于杂合状态的30-6和38-2两个重组子大量收种，利用9838株的BC4F4群体筛选重组子代进行精细定位，将候选基因定位在Pi2/9位点的两个dCAPS标记Nbs1F/R+NdeI和Nbs7-2F/R+SacI之间(图2)。
表1 BC4F3重组子表型与基因型
在此基础上，构建了QBR1单基因系的BAC文库，对BAC克隆筛选、测序，获得覆盖Pi2/9位点的完整序列。序列分析表明，QBR1的Pi2/9位点含有9个NBS-LRR基因(图3A)，其中，只有NBS6-QBR1的CDS区是完整的。结合上述精细定位情况，推测NBS6-QBR1为QBR1的功能基因。Pi2/9基因作为抗病基因的富集区，已克隆和定位10个以上复等位功能基因，已发现的基因虽然具有广谱抗病的特点，但都是小种专化性抗病基因。本研究在该位点发现小种非专化性抗病基因QBR1尚属首次，并且显著不同于之前克隆的Pi2、Pi9、Pi50等基因都是NBS4的直系同源物，此次也是首次发现NBS6为功能基因。
实施例3QBR1对来源于国内和国外的106个菌株均表现部分抗性，基因编辑敲除QBR1丧失对稻瘟病菌生理小种的部分抗性
为了进一步明确QBR1的非小种专化抗性特性，我们利用来自国外的28个稻瘟病菌生理小种和78个国内四川、湖北、安徽、湖南、广东、江西等地分离的生理小种，对CO39-A35(QBR1)、QBR1导入CO39构建近等基因系重组子中的AA(含基因)和CC(不含基因)纯合株系进行接种鉴定。结果表明，QBR1供体亲本和AA型导入系对所有生理小种均表现部分抗性，而CO39和CC型导入系对所有生理小种均表现感病(图4)。进一步的，以QBR1导入CO39的AA型纯系作为受体进行基因编辑，构建了NBS6的敲除突变系20TJ102和20TJ103。随机选取敲除突变系20TJ102和20TJ103各5个株系(编号为102-1、102-2、102-3、102-4、102-5和103-1、103-3、103-4、103-9、103-15)进行稻瘟菌接种，发现20TJ102、20TJ103等敲除系对稻瘟菌的抗性显著的下降(图5)，更加明确NBS6-QBR1即为目的基因。
实施例4创制了QBR1、Pi35、pi21共3个小种非专化抗性基因的聚合系，并明确其高抗效果
将3个非小种转化抗性基因Pi35、Pi21和QBR1分别导入感病水稻对照品种CO39中，构建单基因系。然后再将这3个单基因系两两聚合以及3基因聚合形成聚合系。我们利用10个稻瘟病小种17-1-1、17-5-2、17-6-1、17-6-2、17-8-2、17-8-7、17-8-8、18-4-2、V86010、Ca89对这些单基因系和聚合系进行接种。结果表明，3个单基因系(CO39+QBR1、CO39+Pi35、CO39+pi21)对所有小种都表现部分抗性，其中QBR1的抗性效果最好；2个两两基因聚合系(CO39+Pi35+QBR1、CO39+Pi35+pi21)的抗性比单基因系抗性级别明显提升；3基因聚合系(CO39+Pi35+QBR1+pi21)的抗性则提升到高抗甚至免疫级别水平(图6)。
序列表
<110> 湖南杂交水稻研究中心
<120> 一种水稻稻瘟病非小种专化抗性基因QBR1、分子标记及应用
<141> 2021-08-31
<160> 17
<170> SIPOSequenceListing 1.0
<210> 1
<211> 6205
<212> DNA
<213> Oryza sativa
<400> 1
atggcggaga cggtgctgag catggcgagg tcgctggtgg gagcgccatc agcaaggccg 60
cctccgccgc tgccgacgag accagcctcc tgctcggcgt cgagaaagac atctggtacg 120
taatatgtac tgtggctctc gtttattctg tactagctta ctgatcagca tttattgtat 180
cgacatcctt cttagcctcg attttgttgg ccatcaatcc tgatcggaac aaatcactaa 240
gtcaagcaaa tcaattgatg agtgcacaaa cttttttttt tgcatgacca atcttgagtt 300
cttgagggcg aaacagacgc acccgaccgc agtcaccctc tccctcctcc cacagtccca 360
ctcctcccct ccgccttgcc gctacccgag cgaccgccgg aaagccaagt ggctgcaagg 420
acggtggcgg tggggcatgc tctctctctc tctcggcgct gcaaaggagg gggcaccgac 480
gcatctagga tggtaaaggc gtctccacgc tagtcagatc cggtgagtgg ctggcagaaa 540
acgaggtgga atagcggagg gccgagccag tgcttggtgt ggctagcaac atgggaggct 600
agcgatgcgg tagcaagcag tggtggtaac tacaaatcta tagctccctt gccagatctg 660
acgaccccgt aaccggatct ggcgatggcc caacggtgtc ggaggctacg catccgagaa 720
tgttttggta aggcgacggc acagtcatgc gacgaccagt aatccatggg ggtgaggagg 780
caacaaggtg caaccaagaa agcaccgtga catgccgagg aggatgcggc ggctcgccat 840
gagtggatct ggaagcgaca gatgtagcta gatttggtgg cggggtcaca gcagcaacat 900
gtgccgccga tgcgacgtgg cagtggctgc ttgagatgcc agatggggcc catgtgatgg 960
aggccagagg agcagccagt gggccgacgg ggaggatgca aggcgtggtg ggctgcagcg 1020
acggctcggt gcgagcaagt tcgacgtctg tgcatgctgg gacatcctag cgggcggcgt 1080
gatggtagga ggtcggcaga caaaatagcg tgccacggaa ggtgggccat ggctggtcgg 1140
cggtggggca tcggcgcagc gaacccacag gctagcggag gctgttgggg tggtggaacg 1200
cagggtggtg atgacaggga agggtgaaaa tctagctcgg tgtttcgtca ggccggcaac 1260
gatgacacct tcaagcgccg tcttccccct ttggggcgtt gtcgagctat gaccctctcc 1320
ctcctcacag gactctccag atgaaaacct agtccattta gatgggtcat ggcggcaacc 1380
ttggcatcat gattttcttg gaggcattgt ccaggaggtc ttgtttcctc accttgcaat 1440
gccccgatca tttttgtctt tgggattctt tcagtcgttg tcatcggatt accgtgaggg 1500
acaagtggat tgtcttgtct ctctcgccct ctcactcctc aacccttcta cgtgataatg 1560
attgggagcc cgttgatgac cttctgtttg gatctagtgc tcggtgggcc tttgcaacct 1620
agtgtaggac tagcggtgat cggtcacgca tagaggcggt cggttcggtg ctagtgcttc 1680
tcttggggta tgtaaggagt cgctagaatg tggtggtgtg ctttttattt ttcctttccc 1740
tgattataac cttctagggc tgtaattttg tttttttttc ttgctatatt aatatgaaac 1800
ttcacactgc cttgtgcagc tcgtttaaaa aaaaacttga gttcttgaca ccataagtat 1860
gttagtactg gagtcgacat tagtttatct aaaacatctc ttcatatata tgaaggccac 1920
taatgatttt tctttcctag ggaaccaaag ttccgcggta tgttaatcca caaagcccct 1980
tgtaagatat cctatggcaa atgaactaaa ggcatgtaca atgataataa ttaataggag 2040
aatcttaaca tttctaatta gttctaatta ggaatattaa ctgatatgga agagagagag 2100
aaagaggaga gatagaacat tgttgttacg gttaacaatg gctcagcaac tacttgcctc 2160
tttaaaatgg aaacttggtt gcgagggtga aaaaaaggaa agaatattag taaagaatat 2220
attttttgtt tagtagttaa aatatttgtt gtctcaattg tttaagggca cactctaaat 2280
aattttgtgt tatctaagag tccataccat gcagaggacc cgcgttttcc acgtggaaaa 2340
ccagagcagt aacaaaccta gcgcctccac cccttagatg ggctgtggca gcgctttcgg 2400
cattgcgttt tctttggaag tatcatttag aaaatcctat tattgcctag tctgccttaa 2460
ggaagttcag gcgacaccct tggatggcga tgttcgagag cccatggatg tttagttgtt 2520
tagacatggt gttggacggt cgaatggtgg gcctgttgta ggtatggtgg catctggcaa 2580
ccagtcatgc ttagcaatag ccggatcggt gctagttctt ttcttggtgg tttgtgcacg 2640
cgttgtcgat gcatggtggt gttttttttt tctgtttttc ctgattataa gctcataggg 2700
cttactcttg tatttatttt tctgctatat taatatgaaa acttggtatg gtgatcaagg 2760
gcatgtttgg ttcaaattca ttcctagtct taccaatttt tttttttggc aatggcaata 2820
atcggttatt gaaagaaaat ggcataaatg tgctaggact aagctttgtt tactaccaaa 2880
gttgttcatt ggcagtcaac taagccaaat agttcggtaa tgccattttc ttgtctaggg 2940
accaaaccaa atatatgggt aatattttgg cgttactatt ttctttgctt ggttgagctt 3000
ggtacaaacc aaacagaccc taaatagaca gtgtcatgaa tcacgcctgc taaattccat 3060
tgaactgaac tagaatatac ttcaaagttt tcttgttttc acttggtaca atttcctggt 3120
acaaacttac atttaaattt aaaaacaaaa tcaaaacatt gttacagcac atcatggaaa 3180
aattagtcat ttcattactt ttgtaatata tacatcagtt ggggttatac tcgagatatc 3240
ctaaattgtt ttgagatgaa taattgctag gtatatcaaa gatgagctaa aaacgatgca 3300
agcattcctt agagctgctg aagttatgaa gaagaaagac gaactattaa aggtttgggc 3360
agagcaaata cgtgatctgt catatgacat tgaagattcc cttgatgaat ttaaggtcca 3420
tattgaaagc caaaccctat ttcgtcagtt ggtgaaactt agagagcgcc accggatcgc 3480
tatccgtatc cacaacctca aatcaagagt tgaagaagtg agtagcagga acacacgcta 3540
caatttagtc gagcctattt cctccggcac agaggatgac atggattcct atgcagaaga 3600
cattcgcaat caatcagctc gaaatgtgga tgaagctgag cttgttgggt tttctgactc 3660
caagaaaagg ctgcttgaaa tgatcgatac caatgctaat gatggtccgg ccaaggtaat 3720
ctgtgttgtt gggatgggtg gtttaggcaa gacagctctt tcgaggaaga tctttgaaag 3780
cgaagaagac attaggaaga acttcccttg caatgcttgg attacagtgt cacaatcatt 3840
tcacaggatt gagctactta aagatatgat acgccaactt cttggcccca gttctctgga 3900
tcaactcttg caagaattgc aagggaaggt ggtggtgcaa gtacatcatc tttctgagta 3960
cctgatagaa gagctcaagg agaagaggta ctttgttgtt ctagatgatc tatggatttt 4020
acatgattgg aattggataa atgaaattgc atttcctaag aacaataaga agggcagtcg 4080
aatagtaata accactcgga atgttgatct agcggagaag tgtgccacag cgtcactggt 4140
gtaccacctt gatttcttgc agatgaacga tgccataaca ttgctactga gaaaaacaaa 4200
taaaaatcat gaagacatgg aatcaaataa aaatatgcaa aagatggttg aacgaattgt 4260
aaataaatgt ggtcgtctac cattagcaat acttacaata ggagctgtgc ttgcaactaa 4320
acaggtgtca gaatgggaga aattctatga acaccttcct tcagaactag aaataaaccc 4380
aagcctggaa gctttgagga gaatggtgac cctaggttac aaccacctac catcccatct 4440
gaaaccatgt tttttgtatc taagtatctt tcctgaggat tttgaaatca aaaggaatcg 4500
tctagtaggt agatggatag cagaagggtt tgttagacca aaggttggga tgacgactaa 4560
ggatgtcgga gaaagttact ttaatgagct aatcaaccga agtatgattc aacgatcaag 4620
agtgggcata gcaggaaaaa ttaagacttg tcgaattcat gatatcatcc gtgatatcac 4680
agtttcaatc tcgagacagg aaaattttgt attattacca atgggagatg gctctgattt 4740
agttcaggaa aacactcgcc acatagcatt ccatgggagt atgtcctgca aaacaggatt 4800
ggattggagc attattcgat cattagctat ttttggtgac agacccaaga gtctagcaca 4860
tgcagtttgt ccagatcaat tgaggatgtt acgggtcttg gatcttgaag atgtgacatt 4920
cttaatcact caaaaagatt tcgaccgtat tgcattgttg tgccacttga aatacttgag 4980
tattggatat tcgtcatcca tatattcact tcccagatcc attggtaaac tacagggcct 5040
acagactttg aacatgtcaa gcacatacat tgcagcacta ccaagtgaga tcagtaaact 5100
ccaatgtctg catactcttc gttgtagaag agagcttgat tttgacaaat ttagtctaaa 5160
ccacccaatg aagtgcataa ctaacacaat atgcctgcct aaagtattca cacctttagt 5220
tagtcgcgat gatcgtgcaa tacaaattgc tgaattgcac atggccacca aaagttgctg 5280
gtctgaatca ttcggtgtga aggtacccaa aggaataggt aagttgcgag acttacaggt 5340
tctagagtat gtagatatca ggcggaccag tagtagagca atcaaagagc tggggcagtt 5400
aagcaagttg aggaaattag gtgtgataac aaaaggctcg acaaaggaaa aatgtaagat 5460
actttatgca gccattgaga agctctcttc cctccaatat ctctatgtga atgctgcgtt 5520
tttatcagat attgaaacac ttgagtgcct agattctatt tcatctcctc ctcccctact 5580
gaggacactc aagttgaatg gaagtcttga agagatgcct aactggattg agcagctcac 5640
tcacctgaag aagatctact tattgaggag caaactaaag gaaggtaaaa ccatgctgat 5700
acttggggca ttgcccaacc tcatgggcct tgatctttat tggaaagctt accttgggga 5760
gaagctagta ttcaaaacag gagcattccc aaatcttaga acactttcga tttacgattt 5820
ggatcagcta agagagatta gatttgagga cggcagctcg ccccagttgg aaaagataga 5880
aataggcgag tgcaggttgg aatcagggat tattggtatt atccaccttc caaggctcaa 5940
ggagatttca cttgaatacg aaagtaaagt ggctggactt gctcagctgg agggagaagt 6000
gcgcacacac ccaaatcacc ccgtgctgcg aatgagggag gaccgaagtg atcacgacct 6060
tgcttgtgac gccgaaggat cccctgttga agtggaagca acggatcccc tcccagagca 6120
ggagggagag agctcgcagg tgatcacgtt gacgtcgaac aacagcgaag agataagcac 6180
agctcaagct ggttttatca agtga 6205
<210> 2
<211> 3054
<212> DNA
<213> 人工序列(Artificial sequence)
<400> 2
atggcggaga cggtgctgag catggcgagg tcgctggtgg gagcgccatc agcaaggccg 60
cctccgccgc tgccgacgag accagcctcc tgctcggcgt cgagaaagac atctggtacg 120
tatatcaaag atgagctaaa aacgatgcaa gcattcctta gagctgctga agttatgaag 180
aagaaagacg aactattaaa ggtttgggca gagcaaatac gtgatctgtc atatgacatt 240
gaagattccc ttgatgaatt taaggtccat attgaaagcc aaaccctatt tcgtcagttg 300
gtgaaactta gagagcgcca ccggatcgct atccgtatcc acaacctcaa atcaagagtt 360
gaagaagtga gtagcaggaa cacacgctac aatttagtcg agcctatttc ctccggcaca 420
gaggatgaca tggattccta tgcagaagac attcgcaatc aatcagctcg aaatgtggat 480
gaagctgagc ttgttgggtt ttctgactcc aagaaaaggc tgcttgaaat gatcgatacc 540
aatgctaatg atggtccggc caaggtaatc tgtgttgttg ggatgggtgg tttaggcaag 600
acagctcttt cgaggaagat ctttgaaagc gaagaagaca ttaggaagaa cttcccttgc 660
aatgcttgga ttacagtgtc acaatcattt cacaggattg agctacttaa agatatgata 720
cgccaacttc ttggccccag ttctctggat caactcttgc aagaattgca agggaaggtg 780
gtggtgcaag tacatcatct ttctgagtac ctgatagaag agctcaagga gaagaggtac 840
tttgttgttc tagatgatct atggatttta catgattgga attggataaa tgaaattgca 900
tttcctaaga acaataagaa gggcagtcga atagtaataa ccactcggaa tgttgatcta 960
gcggagaagt gtgccacagc gtcactggtg taccaccttg atttcttgca gatgaacgat 1020
gccataacat tgctactgag aaaaacaaat aaaaatcatg aagacatgga atcaaataaa 1080
aatatgcaaa agatggttga acgaattgta aataaatgtg gtcgtctacc attagcaata 1140
cttacaatag gagctgtgct tgcaactaaa caggtgtcag aatgggagaa attctatgaa 1200
caccttcctt cagaactaga aataaaccca agcctggaag ctttgaggag aatggtgacc 1260
ctaggttaca accacctacc atcccatctg aaaccatgtt ttttgtatct aagtatcttt 1320
cctgaggatt ttgaaatcaa aaggaatcgt ctagtaggta gatggatagc agaagggttt 1380
gttagaccaa aggttgggat gacgactaag gatgtcggag aaagttactt taatgagcta 1440
atcaaccgaa gtatgattca acgatcaaga gtgggcatag caggaaaaat taagacttgt 1500
cgaattcatg atatcatccg tgatatcaca gtttcaatct cgagacagga aaattttgta 1560
ttattaccaa tgggagatgg ctctgattta gttcaggaaa acactcgcca catagcattc 1620
catgggagta tgtcctgcaa aacaggattg gattggagca ttattcgatc attagctatt 1680
tttggtgaca gacccaagag tctagcacat gcagtttgtc cagatcaatt gaggatgtta 1740
cgggtcttgg atcttgaaga tgtgacattc ttaatcactc aaaaagattt cgaccgtatt 1800
gcattgttgt gccacttgaa atacttgagt attggatatt cgtcatccat atattcactt 1860
cccagatcca ttggtaaact acagggccta cagactttga acatgtcaag cacatacatt 1920
gcagcactac caagtgagat cagtaaactc caatgtctgc atactcttcg ttgtagaaga 1980
gagcttgatt ttgacaaatt tagtctaaac cacccaatga agtgcataac taacacaata 2040
tgcctgccta aagtattcac acctttagtt agtcgcgatg atcgtgcaat acaaattgct 2100
gaattgcaca tggccaccaa aagttgctgg tctgaatcat tcggtgtgaa ggtacccaaa 2160
ggaataggta agttgcgaga cttacaggtt ctagagtatg tagatatcag gcggaccagt 2220
agtagagcaa tcaaagagct ggggcagtta agcaagttga ggaaattagg tgtgataaca 2280
aaaggctcga caaaggaaaa atgtaagata ctttatgcag ccattgagaa gctctcttcc 2340
ctccaatatc tctatgtgaa tgctgcgttt ttatcagata ttgaaacact tgagtgccta 2400
gattctattt catctcctcc tcccctactg aggacactca agttgaatgg aagtcttgaa 2460
gagatgccta actggattga gcagctcact cacctgaaga agatctactt attgaggagc 2520
aaactaaagg aaggtaaaac catgctgata cttggggcat tgcccaacct catgggcctt 2580
gatctttatt ggaaagctta ccttggggag aagctagtat tcaaaacagg agcattccca 2640
aatcttagaa cactttcgat ttacgatttg gatcagctaa gagagattag atttgaggac 2700
ggcagctcgc cccagttgga aaagatagaa ataggcgagt gcaggttgga atcagggatt 2760
attggtatta tccaccttcc aaggctcaag gagatttcac ttgaatacga aagtaaagtg 2820
gctggacttg ctcagctgga gggagaagtg cgcacacacc caaatcaccc cgtgctgcga 2880
atgagggagg accgaagtga tcacgacctt gcttgtgacg ccgaaggatc ccctgttgaa 2940
gtggaagcaa cggatcccct cccagagcag gagggagaga gctcgcaggt gatcacgttg 3000
acgtcgaaca acagcgaaga gataagcaca gctcaagctg gttttatcaa gtga 3054
<210> 3
<211> 1017
<212> PRT
<213> 人工序列(Artificial sequence)
<400> 3
Met Ala Glu Thr Val Leu Ser Met Ala Arg Ser Leu Val Gly Ala Pro
1 5 10 15
Ser Ala Arg Pro Pro Pro Pro Leu Pro Thr Arg Pro Ala Ser Cys Ser
20 25 30
Ala Ser Arg Lys Thr Ser Gly Thr Tyr Ile Lys Asp Glu Leu Lys Thr
35 40 45
Met Gln Ala Phe Leu Arg Ala Ala Glu Val Met Lys Lys Lys Asp Glu
50 55 60
Leu Leu Lys Val Trp Ala Glu Gln Ile Arg Asp Leu Ser Tyr Asp Ile
65 70 75 80
Glu Asp Ser Leu Asp Glu Phe Lys Val His Ile Glu Ser Gln Thr Leu
85 90 95
Phe Arg Gln Leu Val Lys Leu Arg Glu Arg His Arg Ile Ala Ile Arg
100 105 110
Ile His Asn Leu Lys Ser Arg Val Glu Glu Val Ser Ser Arg Asn Thr
115 120 125
Arg Tyr Asn Leu Val Glu Pro Ile Ser Ser Gly Thr Glu Asp Asp Met
130 135 140
Asp Ser Tyr Ala Glu Asp Ile Arg Asn Gln Ser Ala Arg Asn Val Asp
145 150 155 160
Glu Ala Glu Leu Val Gly Phe Ser Asp Ser Lys Lys Arg Leu Leu Glu
165 170 175
Met Ile Asp Thr Asn Ala Asn Asp Gly Pro Ala Lys Val Ile Cys Val
180 185 190
Val Gly Met Gly Gly Leu Gly Lys Thr Ala Leu Ser Arg Lys Ile Phe
195 200 205
Glu Ser Glu Glu Asp Ile Arg Lys Asn Phe Pro Cys Asn Ala Trp Ile
210 215 220
Thr Val Ser Gln Ser Phe His Arg Ile Glu Leu Leu Lys Asp Met Ile
225 230 235 240
Arg Gln Leu Leu Gly Pro Ser Ser Leu Asp Gln Leu Leu Gln Glu Leu
245 250 255
Gln Gly Lys Val Val Val Gln Val His His Leu Ser Glu Tyr Leu Ile
260 265 270
Glu Glu Leu Lys Glu Lys Arg Tyr Phe Val Val Leu Asp Asp Leu Trp
275 280 285
Ile Leu His Asp Trp Asn Trp Ile Asn Glu Ile Ala Phe Pro Lys Asn
290 295 300
Asn Lys Lys Gly Ser Arg Ile Val Ile Thr Thr Arg Asn Val Asp Leu
305 310 315 320
Ala Glu Lys Cys Ala Thr Ala Ser Leu Val Tyr His Leu Asp Phe Leu
325 330 335
Gln Met Asn Asp Ala Ile Thr Leu Leu Leu Arg Lys Thr Asn Lys Asn
340 345 350
His Glu Asp Met Glu Ser Asn Lys Asn Met Gln Lys Met Val Glu Arg
355 360 365
Ile Val Asn Lys Cys Gly Arg Leu Pro Leu Ala Ile Leu Thr Ile Gly
370 375 380
Ala Val Leu Ala Thr Lys Gln Val Ser Glu Trp Glu Lys Phe Tyr Glu
385 390 395 400
His Leu Pro Ser Glu Leu Glu Ile Asn Pro Ser Leu Glu Ala Leu Arg
405 410 415
Arg Met Val Thr Leu Gly Tyr Asn His Leu Pro Ser His Leu Lys Pro
420 425 430
Cys Phe Leu Tyr Leu Ser Ile Phe Pro Glu Asp Phe Glu Ile Lys Arg
435 440 445
Asn Arg Leu Val Gly Arg Trp Ile Ala Glu Gly Phe Val Arg Pro Lys
450 455 460
Val Gly Met Thr Thr Lys Asp Val Gly Glu Ser Tyr Phe Asn Glu Leu
465 470 475 480
Ile Asn Arg Ser Met Ile Gln Arg Ser Arg Val Gly Ile Ala Gly Lys
485 490 495
Ile Lys Thr Cys Arg Ile His Asp Ile Ile Arg Asp Ile Thr Val Ser
500 505 510
Ile Ser Arg Gln Glu Asn Phe Val Leu Leu Pro Met Gly Asp Gly Ser
515 520 525
Asp Leu Val Gln Glu Asn Thr Arg His Ile Ala Phe His Gly Ser Met
530 535 540
Ser Cys Lys Thr Gly Leu Asp Trp Ser Ile Ile Arg Ser Leu Ala Ile
545 550 555 560
Phe Gly Asp Arg Pro Lys Ser Leu Ala His Ala Val Cys Pro Asp Gln
565 570 575
Leu Arg Met Leu Arg Val Leu Asp Leu Glu Asp Val Thr Phe Leu Ile
580 585 590
Thr Gln Lys Asp Phe Asp Arg Ile Ala Leu Leu Cys His Leu Lys Tyr
595 600 605
Leu Ser Ile Gly Tyr Ser Ser Ser Ile Tyr Ser Leu Pro Arg Ser Ile
610 615 620
Gly Lys Leu Gln Gly Leu Gln Thr Leu Asn Met Ser Ser Thr Tyr Ile
625 630 635 640
Ala Ala Leu Pro Ser Glu Ile Ser Lys Leu Gln Cys Leu His Thr Leu
645 650 655
Arg Cys Arg Arg Glu Leu Asp Phe Asp Lys Phe Ser Leu Asn His Pro
660 665 670
Met Lys Cys Ile Thr Asn Thr Ile Cys Leu Pro Lys Val Phe Thr Pro
675 680 685
Leu Val Ser Arg Asp Asp Arg Ala Ile Gln Ile Ala Glu Leu His Met
690 695 700
Ala Thr Lys Ser Cys Trp Ser Glu Ser Phe Gly Val Lys Val Pro Lys
705 710 715 720
Gly Ile Gly Lys Leu Arg Asp Leu Gln Val Leu Glu Tyr Val Asp Ile
725 730 735
Arg Arg Thr Ser Ser Arg Ala Ile Lys Glu Leu Gly Gln Leu Ser Lys
740 745 750
Leu Arg Lys Leu Gly Val Ile Thr Lys Gly Ser Thr Lys Glu Lys Cys
755 760 765
Lys Ile Leu Tyr Ala Ala Ile Glu Lys Leu Ser Ser Leu Gln Tyr Leu
770 775 780
Tyr Val Asn Ala Ala Phe Leu Ser Asp Ile Glu Thr Leu Glu Cys Leu
785 790 795 800
Asp Ser Ile Ser Ser Pro Pro Pro Leu Leu Arg Thr Leu Lys Leu Asn
805 810 815
Gly Ser Leu Glu Glu Met Pro Asn Trp Ile Glu Gln Leu Thr His Leu
820 825 830
Lys Lys Ile Tyr Leu Leu Arg Ser Lys Leu Lys Glu Gly Lys Thr Met
835 840 845
Leu Ile Leu Gly Ala Leu Pro Asn Leu Met Gly Leu Asp Leu Tyr Trp
850 855 860
Lys Ala Tyr Leu Gly Glu Lys Leu Val Phe Lys Thr Gly Ala Phe Pro
865 870 875 880
Asn Leu Arg Thr Leu Ser Ile Tyr Asp Leu Asp Gln Leu Arg Glu Ile
885 890 895
Arg Phe Glu Asp Gly Ser Ser Pro Gln Leu Glu Lys Ile Glu Ile Gly
900 905 910
Glu Cys Arg Leu Glu Ser Gly Ile Ile Gly Ile Ile His Leu Pro Arg
915 920 925
Leu Lys Glu Ile Ser Leu Glu Tyr Glu Ser Lys Val Ala Gly Leu Ala
930 935 940
Gln Leu Glu Gly Glu Val Arg Thr His Pro Asn His Pro Val Leu Arg
945 950 955 960
Met Arg Glu Asp Arg Ser Asp His Asp Leu Ala Cys Asp Ala Glu Gly
965 970 975
Ser Pro Val Glu Val Glu Ala Thr Asp Pro Leu Pro Glu Gln Glu Gly
980 985 990
Glu Ser Ser Gln Val Ile Thr Leu Thr Ser Asn Asn Ser Glu Glu Ile
995 1000 1005
Ser Thr Ala Gln Ala Gly Phe Ile Lys
1010 1015
<210> 4
<211> 20
<212> DNA
<213> 人工序列(Artificial sequence)
<400> 4
tggcgacaat aagcttggga 20
<210> 5
<211> 20
<212> DNA
<213> 人工序列(Artificial sequence)
<400> 5
ggctactact ccctccgtcc 20
<210> 6
<211> 20
<212> DNA
<213> 人工序列(Artificial sequence)
<400> 6
acgggtagag ctgccaaatc 20
<210> 7
<211> 20
<212> DNA
<213> 人工序列(Artificial sequence)
<400> 7
tagcgaactc aaccaccgac 20
<210> 8
<211> 20
<212> DNA
<213> 人工序列(Artificial sequence)
<400> 8
tcatcgtcat caccggcttc 20
<210> 9
<211> 20
<212> DNA
<213> 人工序列(Artificial sequence)
<400> 9
ttgagggagg gctttcaacg 20
<210> 10
<211> 20
<212> DNA
<213> 人工序列(Artificial sequence)
<400> 10
gacagacaaa cgttgggcac 20
<210> 11
<211> 20
<212> DNA
<213> 人工序列(Artificial sequence)
<400> 11
aagatcagcg tgtgaccgtt 20
<210> 12
<211> 20
<212> DNA
<213> 人工序列(Artificial sequence)
<400> 12
tggaaaacag agtcctcggc 20
<210> 13
<211> 20
<212> DNA
<213> 人工序列(Artificial sequence)
<400> 13
ggcaaagatt tgcgtcctcc 20
<210> 14
<211> 20
<212> DNA
<213> 人工序列(Artificial sequence)
<400> 14
gaaggtgaag gagatgatgg 20
<210> 15
<211> 20
<212> DNA
<213> 人工序列(Artificial sequence)
<400> 15
ggttcaaaac tacgcggaag 20
<210> 16
<211> 20
<212> DNA
<213> 人工序列(Artificial sequence)
<400> 16
tgtacaggcc agccattcag 20
<210> 17
<211> 20
<212> DNA
<213> 人工序列(Artificial sequence)
<400> 17
caatcgtcgt gttcatcgcc 20
Claims (6)
1.一种水稻稻瘟病非小种专化抗性基因QBR1，其特征在于，其核苷酸序列全长如SEQID NO:1所示。
2.一种水稻稻瘟病非小种专化抗性基因QBR1的cDNA，其特征在于，所述cDNA的核苷酸序列如SEQ ID NO:2所示。
3.由权利要求1所述的水稻稻瘟病非小种专化抗性基因QBR1或权利要求2所述非小种专化抗性基因QBR1的cDNA编码的蛋白质，其特征在于，所述蛋白质的氨基酸序列如SEQ IDNO:3所示。
4.由权利要求1所述的水稻稻瘟病非小种专化抗性基因QBR1、权利要求2所述的非小种专化抗性基因QBR1的cDNA或权利要求3所述的蛋白质在提高水稻稻瘟病抗性中的应用。
5.根据权利要求4所述的应用，其特征在于，所述稻瘟病抗性为子囊菌稻瘟菌抗性。
6.一种培育抗稻瘟病水稻品种的方法，其特征在于，所述方法为：将非小种专化抗性基因pi21、Pi35中至少一个基因与权利要求1所述的非小种专化抗性基因QBR1或权利要求2所述的非小种专化抗性基因QBR1的cDNA在水稻中聚合。
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| CN202111512039.8A CN114790458B (zh) | 2021-12-07 | 2021-12-07 | 一种水稻稻瘟病非小种专化抗性基因qbr1、分子标记及应用 |
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| CN101824418A (zh) * | 2009-12-10 | 2010-09-08 | 中国水稻研究所 | 水稻抗稻瘟病基因Pi 25编码区及其应用 |
| CN102051368A (zh) * | 2010-02-02 | 2011-05-11 | 华南农业大学 | 稻瘟病抗性基因Pik及其应用 |
| CN103642803A (zh) * | 2013-12-11 | 2014-03-19 | 中国农业科学院作物科学研究所 | 稻瘟病抗性基因Pi63的功能特异性分子标记及其方法与应用 |
| CN109182590A (zh) * | 2018-11-02 | 2019-01-11 | 江苏省农业科学院 | 检测广谱抗稻瘟病基因PigmR的功能标记及其应用 |
| CN113061171A (zh) * | 2021-04-13 | 2021-07-02 | 四川农业大学 | 抗稻瘟病蛋白和基因、分离的核酸及其应用 |
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| CN101824418A (zh) * | 2009-12-10 | 2010-09-08 | 中国水稻研究所 | 水稻抗稻瘟病基因Pi 25编码区及其应用 |
| CN102051368A (zh) * | 2010-02-02 | 2011-05-11 | 华南农业大学 | 稻瘟病抗性基因Pik及其应用 |
| CN103642803A (zh) * | 2013-12-11 | 2014-03-19 | 中国农业科学院作物科学研究所 | 稻瘟病抗性基因Pi63的功能特异性分子标记及其方法与应用 |
| CN109182590A (zh) * | 2018-11-02 | 2019-01-11 | 江苏省农业科学院 | 检测广谱抗稻瘟病基因PigmR的功能标记及其应用 |
| CN113061171A (zh) * | 2021-04-13 | 2021-07-02 | 四川农业大学 | 抗稻瘟病蛋白和基因、分离的核酸及其应用 |
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