子预44中抗稻瘟病基因Pi-zy3(t)的定位
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摘要
稻瘟病是影响水稻高产、稳产、优质的重要病害。广谱持久抗瘟品种的培育和利用是防治稻瘟病最经济有效的措施之一,但广谱持久抗性分子机制还不清楚。子预44是一具有广谱持久稻瘟病抗性的云南地方粳稻品种,对其抗瘟基因的鉴定将有助于揭示其抗瘟分子机制。本研究通过利用子预44和感病水稻江南香糯杂交构建F2遗传群体,稻瘟病菌株LP33苗期喷雾接种亲本预44、江南香糯及F2代单株,对其抗性进行评价和主效抗瘟基因定位。研究结果显示:子预44表现高抗,江南香糯高感;F2群体稻瘟病抗感分离符合3:1的分离比,即子预44对LP33的抗性为单显性基因控制的抗性遗传,并将该基因暂定名为Pi-zy3(t)。进一步利用SSR分子标记将Pi-zy3(t)定位在水稻第六号染色体RM276到RM3827之间,为进一步的基因克隆和抗病机分子制研究奠定了基础,同时也为利用子预44进行抗病分子育种提供了辅助选择的分子标记。
R ice blast is one of the most severe diseases in rice production and causes serious yield losses and quality decreasing. Development and use of broad-spectrum durable resistant varieties is considered as the most effective and economical strategy to control this disease, but the mechanism of broad-spectrum durable blast resistance is not clear. The japonica rice cultivar Ziyu44 confers broad-spectrum durable resistance to blast.Identification and cloning of resistance genes in Ziyu44 will benefit to investigate the molecular basis of broad-spectrum durable resistance. In this study, we developed an F2 population from the cross between Ziyu44 and JNXN. The F2 progenies, Ziyu44 and JNXN, were inoculated with the blast isolate LP33 for blast disease evaluation. The results showed that Ziyu44 conferred high level resistance to LP33, and JNXN showed high susceptibility to LP33. The segregation of resistant and susceptible plants in F2 population agreed with the ratios of3:1, indicating that the blast resistance of Ziyu44 to LP33 is controlled by a single dominant gene, temporarily designated Pi-zy3(t). Further linkage analysis using susceptible F2 individuals, Pi-zy3(t) was located on rice chromosome 6 closed to the centromere between SSR markers RM276 and RM3827. These results provide essential information for the cloning of the Pi-zy3(t) gene and will facilitate marker-assisted selection in rice breeding.
R ice blast is one of the most severe diseases in rice production and causes serious yield losses and quality decreasing. Development and use of broad-spectrum durable resistant varieties is considered as the most effective and economical strategy to control this disease, but the mechanism of broad-spectrum durable blast resistance is not clear. The japonica rice cultivar Ziyu44 confers broad-spectrum durable resistance to blast.Identification and cloning of resistance genes in Ziyu44 will benefit to investigate the molecular basis of broad-spectrum durable resistance. In this study, we developed an F2 population from the cross between Ziyu44 and JNXN. The F2 progenies, Ziyu44 and JNXN, were inoculated with the blast isolate LP33 for blast disease evaluation. The results showed that Ziyu44 conferred high level resistance to LP33, and JNXN showed high susceptibility to LP33. The segregation of resistant and susceptible plants in F2 population agreed with the ratios of3:1, indicating that the blast resistance of Ziyu44 to LP33 is controlled by a single dominant gene, temporarily designated Pi-zy3(t). Further linkage analysis using susceptible F2 individuals, Pi-zy3(t) was located on rice chromosome 6 closed to the centromere between SSR markers RM276 and RM3827. These results provide essential information for the cloning of the Pi-zy3(t) gene and will facilitate marker-assisted selection in rice breeding.
引文
Ahn S.N.,Kim Y.K.,Hong H.C.,Han S.S.,Kwon S.J.,Choi H.C.,Moon H.P.,and Mc Couch S.R.,2000,Molecular mapping ofa new gene for resistance to rice blast(Pyricularia griseaSacc.),Euphytica,116(1):17-22
Chen X.W.,Li S.G.,Xu J.C.,Zhai W.X.,Ling Z.Z.,Ma B.T.,WangY.P.,Wang W.M.,Cao G.,Ma Y.Q.,Shang J.J.,Zhao X.F.,Zhou K.D,and Zhu L.H.,2004,Identification of two blast resistance genes in a rice variety,Digu,J.Phytopathol.,152(2):77-85
Chen X.W.,Shang J.J.,Chen D.X.,Lei C.L.,Zou Y.,Zhai W.X.,Liu G.Z.,Xu J.C.,Ling Z.Z.,Cao G.,Ma B.T.,Wang Y.P.,Zhao X.F.,Li S.G.,and Zhu L.H.,2006,A B-lectin receptorkinase gene conferring rice blast resistance,Plant J.,46(5):794-804
Deng Y.W.,Zhu X.D.,Shen Y.,and He Z.H.,2006,Genetic characterization and fine mapping of the blast resistance locusPigm(t)tightly linked to Pi2 and Pi9 in a broad-spectrum resistant Chinese variety,Theor.Appl.Genet.,113(4):705-713
Hayashi K.,Yoshida H.,and Ashikawa I.,2006,Development ofPCR-based allele-specific and in Del markers sets for ninerice blast resistance genes,Theor.Appl.Genet.,113(2):251-260
He X.Y.,Wang L.,Wu W.H.,Chen Z.M.,Lin F.,Cheng Y.S.,LiuW.,Chen Y.H.,and Liao Y.P.,2014,The progress of mapping,isolation of the genes resisting to blast and their breeding application in rice,Zhongguo Nongxue Tongbao(Chinese Agricultural Science Bulletin),30(6):1-12(何秀英,王玲,吴伟怀,陈钊明,林菲,程永盛,刘维,陈粤汉,廖耀平,2014,水稻稻瘟病抗性基因的定位、克隆及育种应用研究进展,中国农学通报,30(6):1-12)
Jeung J.U.,Kim B.R.,Cho Y.C.,Han S.S.,Moon H.P.,Lee Y.T.,and Jena K.K.,2007,A novel gene,Pi40(t),linked to the DNAmarkers derived from NBS-LRR motifs confers broad spectrum of blast resistance in rice,Theor.Appl.Genet,115(8):1163-1177
Li S.,Li Q.,Fan L.L.,Sha S.,Zeng Q.C.,and Luo Q.,2014,Identification of the rice blast resistance genes by pseudogenizationmolecular markers in plateau japonica variety Ziyu44,FenziZhiwu Yuzhong(Molecular Plant Breeding),12(2):219-225(李书,李权,樊琳琳,沙莎,曾千春,罗琼,2014,高原粳稻子预44中三个稻瘟病抗性基因的假基因化分子标记鉴定,分子植物育种,12(2):219-225)
Liu G.,Lu G.,Zeng L.,and Wang G.L.,2002,Two broad-spectrum blast resistance genes,Pi9(t)and Pi2(t),are physicallylinked on rice chromosome 6,Mol.Genet.Genomics,267(4):472-480
Liu J.L.,Wang X.J.,Thomas M.,Hu Y.J.,Liu X.L.,Dai L.Y.,andWang G.L.,2010,Recent progress and understanding of themolecular mechanism of the rice-Magnaporthe oryzae interaction,Mol.Plant Pathol.,11(3):419-427
Liu X.Q.,Yang Q.Z.,Lin F.,Hua L.X.,Wang C.T.,Wang L.,andPan Q.H.,2007,Identification and fine mapping of Pi39(t),amajor gene conferring the broad-spectrum resistance to Magnaporthe oryzae,Mol.Genet.Genomics.,278(4):403-410
Lv Q.M.,Xu X.,Shang J.J.,Jiang G.H.,Pang Z.Q.,Zhou Z.Z.,Wang J.,Liu Y.,Li T.,Li X.B.,Xu J.C.,Cheng Z.K.,Zhao X.F.,Li S.G.,and Zhu L.H.,2013,Functional analysis of Pid3-A4,an ortholog of rice blast resistance gene Pid3 revealed byallele mining in common wild rice,Phytopathology,103(6):594-599
Pan Q.H.,Wang L.,Ikehashi H.,and Tanisaha T.,1996,Identification of a new blast resistance gene in the india rice cultivarKasalath using Japanese differential cultivars and isozymemarkers,Phytopathology,86:1071-1075
Pan Q.H.,Wang L.,Ikehashi H.,Yamagata H.,and Tanisaha T.,1998,Identification of two new genes conferring resistancerice blast in the Chinese native cultivar‘Maowangu’,PlantBreeding,117(1):27-31
Pari S.,and Sarah J.G.,2009,Against the grain:safeguarding ricefrom rice blast disease,Cell,27(3):141-150
Qu S.H.,Liu G.F.,Zhou B.,Maria B.,Zeng L.R.,Dai L.Y.,Han B.,and Wang G.L.,2006,The broad-spectrum blast resistancegene Pi9 encodes a nucleotide-binding site-leucine-rich repeat protein and a member of a multigene family in rice,Genetics,172(3):1901-1914
Ramkumar G.,Madhav M.S.,Rama Devi S.J.S.,Manimaran P.,Mohan K.M.,Prasad M.S.,Balachandran S.M.,Neeraja C.N.,Sundaram R.M.,and Viraktamath B.C.,2014,Nucleotide diversity of Pita,a major blast resistance gene and identificationof its minimal promoter,Elsevier B.V.,546(2):250-256
Sallaud C.,Lorieux M.,Roumen E.,Tharreau D.,Berruyer R.,Svestasrani P.,Garsmeur O.,Ghesquiere A.,and Notteghem J.L.,2003,Identification of five new blast resistance genes inthe highly blast-resistant rice variety IR64 using a QTL mapping strategy,Theor.Appl.Genet.,106(5):794-803
Savary S.,Willocquet L.,Elazegui F.A.,Castilla N.P.,and Teng P.S.,2000,Rice pest constraints in tropical Asia:quantificationof yield losses due to rice pests in a range of production situations,Plant Disease,84(3):357-369
Shang J.J.,Tao Y.,Chen X.W.,Zou Y.,Lei C.L.,Wang J.,Li X.B.,Zhao X.F.,Zhang M.J.,Lu Z.K.,Xu J.C.,Cheng Z.K.,Wan J.M.,and Zhu L.H.,2009,Identification of a new rice blast resistance gene,Pid3,by genomewide comparison of pairednucleotide-binding site-leucine-rich repeat genes and theirpseudogene alleles between the two sequenced rice genomes,Genetics,182(4):1303-1311
Sun G.C.,Chai R.Y.,Du X.F.,Tao R.X.,and Sun S.Y.,1999,Thedirectional selection and stabilizing selection from the interaction between rice variety and blast fungus Magnaporthegrisea,Zhiwu Bingli Xuebao(Acta Phytopathologica Sinica),29(1):45-49(孙国昌,柴荣耀,杜新法,陶荣祥,孙漱沅,1999,水稻品种与稻瘟病群体互作的选择作用研究,植物病理学报,29(1):45-49)
Sun S.Y.,and Sun G.C.,1996,Status and prospect of China's riceblast,Zhibao Jishu Yu Tuiguang(Plant Protection Technology and Extension),16(3):39-40(孙漱沅,孙国昌,1996,我国稻瘟病研究的现状和展望,植保技术与推广,16(3):39-40)
Tabien R.E.,Li Z.,Paterson A.H.,Marchetti M.A.,Stansel J.W.,Pinson S.R.M.,and Park W.D.,2000,Mapping of four majorrice blast resistance genes from‘Lemont’and‘Teqing’andevaluation of their combinatorial effect for field resistance,Theor.Appl.Genet,101(8):1215-1225
Wang G.L.,Mackill D.J.,Bonman J.M.,Mc Couch S.R.,Champoux M.C.,and Nelson R.J.,1994,RFLP mapping of genesconferring complete and partial resistance to blast in adurably resistant rice cultivar,Genetics,136(4):1421-1434
Wu J.,Liu X.L.,Dai L.Y.,and Wang G.L.,2007,Advances on theidentification and characterization of broad spectrum blastresistance genes in rice,Shengming Kexue(Chinese Bulletinof Life Sciences),19(2):233-238(吴俊,刘雄伦,戴良英,王国梁,2007,水稻广谱抗稻瘟病基因研究进展,生命科学,19(2):233-238)
Wu J.L.,Fan Y.Y.,Li D.B.,Zheng K.L.,Leung H.,and Zhuang J.Y.,2005,Genetic control of rice blast resistance in the durablyresistance cultivar Gumei 2 against multiple isolates,Theor.Appl.Genet.,111(1):50-56
Xu X.,Lv Q.M.,Shang J.J.,Pang Z.Q.,Zhou Z.Z.,Wang J.,JiangG.H.,Tao Y.,Xu Q.,Li X.B.,Li S.G.,Xu J.C.,and Zhu L.H.,2014,Excavation of Pid3 orthologs with differential resistance spectra to Magnaporthe oryzae in rice resource,PLo SOne,9(3):e9327
Zhang J.W.,Hong R.K.,Zhang Y.Y.,Fan J.H.,Zeng Q.C.,and LuoQ.,2011,Analysis of broad spectrum and persistent rice blastresistance in Yunnan local rice variety,Xinan Nongye Kexue(Southwest China Journal of Agriculture),24(4):1323-1326(张锦文,洪汝科,范静华,张祎颖,曾千春,罗琼,2011,一份云南地方稻广谱持久抗稻瘟病初步分析,西南农业学报,24(4):1323-1326)
Zhang J.W.,Tan Y.L.,Hong R.K.,Fan J.H.,Luo Q.,and Zeng Q.C.,2009,Genetic analysis and gene mapping of rice blast resistance in japonica variety Ziyu44,Zhongguo Shuidao Kexue(Chinese Journal of Rice Science),23(1):31-35(张锦文,谭亚玲,洪汝科,范静华,罗琼,曾千春,2009,高原粳稻子预44抗稻瘟病基因遗传分析和定位,中国水稻科学,23(1):31-35)
Zhang J.Y.,Xie G.Q.,and Han Z.L.,2009,Analysis of the conditions of occurrence of rice blast in Yunnan Province in 2008,Yunnan Nongye Keji(Yunnan Agricultural Science and Technology),(1):50-51(张加云,谢国清,韩忠良,2009,2008年云南省水稻稻瘟病发生条件分析,云南农业科技,(1):50-51)
Zhi G.Y.,Zhang G.M.,Lei C.L.,Xin A.H.,Chen P.,Ma J.T.,XiaoJ.L.,Liu Y.X.,and Ren Y.,2010,Heilongjiang Province in2007 The investigation and suggestions of rice blast,Heilongjiang Nongye Kexue(Heilongjiang Agricultural Sciences),(4):68-70(支庚银,张国民,雷财林,辛爱华,陈平,马军韬,肖家雷,刘迎雪,任洋,2010,黑龙江省2007年水稻稻瘟病生产调研及建议,黑龙江农业科学,(4):68-70)
Zhu X.Y.,Chen S.,Yang J.Y.,Zhou S.C.,Zeng L.X.,Han J.L.,SuJ.,Wang L.,and Pan Q.H.,2012,The identification of Pi50(t),a new member of the rice blast resistance Pi2/Pi9 multigenefamily,Theor.Appl.Genet.,124(7):1295-1304
Chen X.W.,Li S.G.,Xu J.C.,Zhai W.X.,Ling Z.Z.,Ma B.T.,WangY.P.,Wang W.M.,Cao G.,Ma Y.Q.,Shang J.J.,Zhao X.F.,Zhou K.D,and Zhu L.H.,2004,Identification of two blast resistance genes in a rice variety,Digu,J.Phytopathol.,152(2):77-85
Chen X.W.,Shang J.J.,Chen D.X.,Lei C.L.,Zou Y.,Zhai W.X.,Liu G.Z.,Xu J.C.,Ling Z.Z.,Cao G.,Ma B.T.,Wang Y.P.,Zhao X.F.,Li S.G.,and Zhu L.H.,2006,A B-lectin receptorkinase gene conferring rice blast resistance,Plant J.,46(5):794-804
Deng Y.W.,Zhu X.D.,Shen Y.,and He Z.H.,2006,Genetic characterization and fine mapping of the blast resistance locusPigm(t)tightly linked to Pi2 and Pi9 in a broad-spectrum resistant Chinese variety,Theor.Appl.Genet.,113(4):705-713
Hayashi K.,Yoshida H.,and Ashikawa I.,2006,Development ofPCR-based allele-specific and in Del markers sets for ninerice blast resistance genes,Theor.Appl.Genet.,113(2):251-260
He X.Y.,Wang L.,Wu W.H.,Chen Z.M.,Lin F.,Cheng Y.S.,LiuW.,Chen Y.H.,and Liao Y.P.,2014,The progress of mapping,isolation of the genes resisting to blast and their breeding application in rice,Zhongguo Nongxue Tongbao(Chinese Agricultural Science Bulletin),30(6):1-12(何秀英,王玲,吴伟怀,陈钊明,林菲,程永盛,刘维,陈粤汉,廖耀平,2014,水稻稻瘟病抗性基因的定位、克隆及育种应用研究进展,中国农学通报,30(6):1-12)
Jeung J.U.,Kim B.R.,Cho Y.C.,Han S.S.,Moon H.P.,Lee Y.T.,and Jena K.K.,2007,A novel gene,Pi40(t),linked to the DNAmarkers derived from NBS-LRR motifs confers broad spectrum of blast resistance in rice,Theor.Appl.Genet,115(8):1163-1177
Li S.,Li Q.,Fan L.L.,Sha S.,Zeng Q.C.,and Luo Q.,2014,Identification of the rice blast resistance genes by pseudogenizationmolecular markers in plateau japonica variety Ziyu44,FenziZhiwu Yuzhong(Molecular Plant Breeding),12(2):219-225(李书,李权,樊琳琳,沙莎,曾千春,罗琼,2014,高原粳稻子预44中三个稻瘟病抗性基因的假基因化分子标记鉴定,分子植物育种,12(2):219-225)
Liu G.,Lu G.,Zeng L.,and Wang G.L.,2002,Two broad-spectrum blast resistance genes,Pi9(t)and Pi2(t),are physicallylinked on rice chromosome 6,Mol.Genet.Genomics,267(4):472-480
Liu J.L.,Wang X.J.,Thomas M.,Hu Y.J.,Liu X.L.,Dai L.Y.,andWang G.L.,2010,Recent progress and understanding of themolecular mechanism of the rice-Magnaporthe oryzae interaction,Mol.Plant Pathol.,11(3):419-427
Liu X.Q.,Yang Q.Z.,Lin F.,Hua L.X.,Wang C.T.,Wang L.,andPan Q.H.,2007,Identification and fine mapping of Pi39(t),amajor gene conferring the broad-spectrum resistance to Magnaporthe oryzae,Mol.Genet.Genomics.,278(4):403-410
Lv Q.M.,Xu X.,Shang J.J.,Jiang G.H.,Pang Z.Q.,Zhou Z.Z.,Wang J.,Liu Y.,Li T.,Li X.B.,Xu J.C.,Cheng Z.K.,Zhao X.F.,Li S.G.,and Zhu L.H.,2013,Functional analysis of Pid3-A4,an ortholog of rice blast resistance gene Pid3 revealed byallele mining in common wild rice,Phytopathology,103(6):594-599
Pan Q.H.,Wang L.,Ikehashi H.,and Tanisaha T.,1996,Identification of a new blast resistance gene in the india rice cultivarKasalath using Japanese differential cultivars and isozymemarkers,Phytopathology,86:1071-1075
Pan Q.H.,Wang L.,Ikehashi H.,Yamagata H.,and Tanisaha T.,1998,Identification of two new genes conferring resistancerice blast in the Chinese native cultivar‘Maowangu’,PlantBreeding,117(1):27-31
Pari S.,and Sarah J.G.,2009,Against the grain:safeguarding ricefrom rice blast disease,Cell,27(3):141-150
Qu S.H.,Liu G.F.,Zhou B.,Maria B.,Zeng L.R.,Dai L.Y.,Han B.,and Wang G.L.,2006,The broad-spectrum blast resistancegene Pi9 encodes a nucleotide-binding site-leucine-rich repeat protein and a member of a multigene family in rice,Genetics,172(3):1901-1914
Ramkumar G.,Madhav M.S.,Rama Devi S.J.S.,Manimaran P.,Mohan K.M.,Prasad M.S.,Balachandran S.M.,Neeraja C.N.,Sundaram R.M.,and Viraktamath B.C.,2014,Nucleotide diversity of Pita,a major blast resistance gene and identificationof its minimal promoter,Elsevier B.V.,546(2):250-256
Sallaud C.,Lorieux M.,Roumen E.,Tharreau D.,Berruyer R.,Svestasrani P.,Garsmeur O.,Ghesquiere A.,and Notteghem J.L.,2003,Identification of five new blast resistance genes inthe highly blast-resistant rice variety IR64 using a QTL mapping strategy,Theor.Appl.Genet.,106(5):794-803
Savary S.,Willocquet L.,Elazegui F.A.,Castilla N.P.,and Teng P.S.,2000,Rice pest constraints in tropical Asia:quantificationof yield losses due to rice pests in a range of production situations,Plant Disease,84(3):357-369
Shang J.J.,Tao Y.,Chen X.W.,Zou Y.,Lei C.L.,Wang J.,Li X.B.,Zhao X.F.,Zhang M.J.,Lu Z.K.,Xu J.C.,Cheng Z.K.,Wan J.M.,and Zhu L.H.,2009,Identification of a new rice blast resistance gene,Pid3,by genomewide comparison of pairednucleotide-binding site-leucine-rich repeat genes and theirpseudogene alleles between the two sequenced rice genomes,Genetics,182(4):1303-1311
Sun G.C.,Chai R.Y.,Du X.F.,Tao R.X.,and Sun S.Y.,1999,Thedirectional selection and stabilizing selection from the interaction between rice variety and blast fungus Magnaporthegrisea,Zhiwu Bingli Xuebao(Acta Phytopathologica Sinica),29(1):45-49(孙国昌,柴荣耀,杜新法,陶荣祥,孙漱沅,1999,水稻品种与稻瘟病群体互作的选择作用研究,植物病理学报,29(1):45-49)
Sun S.Y.,and Sun G.C.,1996,Status and prospect of China's riceblast,Zhibao Jishu Yu Tuiguang(Plant Protection Technology and Extension),16(3):39-40(孙漱沅,孙国昌,1996,我国稻瘟病研究的现状和展望,植保技术与推广,16(3):39-40)
Tabien R.E.,Li Z.,Paterson A.H.,Marchetti M.A.,Stansel J.W.,Pinson S.R.M.,and Park W.D.,2000,Mapping of four majorrice blast resistance genes from‘Lemont’and‘Teqing’andevaluation of their combinatorial effect for field resistance,Theor.Appl.Genet,101(8):1215-1225
Wang G.L.,Mackill D.J.,Bonman J.M.,Mc Couch S.R.,Champoux M.C.,and Nelson R.J.,1994,RFLP mapping of genesconferring complete and partial resistance to blast in adurably resistant rice cultivar,Genetics,136(4):1421-1434
Wu J.,Liu X.L.,Dai L.Y.,and Wang G.L.,2007,Advances on theidentification and characterization of broad spectrum blastresistance genes in rice,Shengming Kexue(Chinese Bulletinof Life Sciences),19(2):233-238(吴俊,刘雄伦,戴良英,王国梁,2007,水稻广谱抗稻瘟病基因研究进展,生命科学,19(2):233-238)
Wu J.L.,Fan Y.Y.,Li D.B.,Zheng K.L.,Leung H.,and Zhuang J.Y.,2005,Genetic control of rice blast resistance in the durablyresistance cultivar Gumei 2 against multiple isolates,Theor.Appl.Genet.,111(1):50-56
Xu X.,Lv Q.M.,Shang J.J.,Pang Z.Q.,Zhou Z.Z.,Wang J.,JiangG.H.,Tao Y.,Xu Q.,Li X.B.,Li S.G.,Xu J.C.,and Zhu L.H.,2014,Excavation of Pid3 orthologs with differential resistance spectra to Magnaporthe oryzae in rice resource,PLo SOne,9(3):e9327
Zhang J.W.,Hong R.K.,Zhang Y.Y.,Fan J.H.,Zeng Q.C.,and LuoQ.,2011,Analysis of broad spectrum and persistent rice blastresistance in Yunnan local rice variety,Xinan Nongye Kexue(Southwest China Journal of Agriculture),24(4):1323-1326(张锦文,洪汝科,范静华,张祎颖,曾千春,罗琼,2011,一份云南地方稻广谱持久抗稻瘟病初步分析,西南农业学报,24(4):1323-1326)
Zhang J.W.,Tan Y.L.,Hong R.K.,Fan J.H.,Luo Q.,and Zeng Q.C.,2009,Genetic analysis and gene mapping of rice blast resistance in japonica variety Ziyu44,Zhongguo Shuidao Kexue(Chinese Journal of Rice Science),23(1):31-35(张锦文,谭亚玲,洪汝科,范静华,罗琼,曾千春,2009,高原粳稻子预44抗稻瘟病基因遗传分析和定位,中国水稻科学,23(1):31-35)
Zhang J.Y.,Xie G.Q.,and Han Z.L.,2009,Analysis of the conditions of occurrence of rice blast in Yunnan Province in 2008,Yunnan Nongye Keji(Yunnan Agricultural Science and Technology),(1):50-51(张加云,谢国清,韩忠良,2009,2008年云南省水稻稻瘟病发生条件分析,云南农业科技,(1):50-51)
Zhi G.Y.,Zhang G.M.,Lei C.L.,Xin A.H.,Chen P.,Ma J.T.,XiaoJ.L.,Liu Y.X.,and Ren Y.,2010,Heilongjiang Province in2007 The investigation and suggestions of rice blast,Heilongjiang Nongye Kexue(Heilongjiang Agricultural Sciences),(4):68-70(支庚银,张国民,雷财林,辛爱华,陈平,马军韬,肖家雷,刘迎雪,任洋,2010,黑龙江省2007年水稻稻瘟病生产调研及建议,黑龙江农业科学,(4):68-70)
Zhu X.Y.,Chen S.,Yang J.Y.,Zhou S.C.,Zeng L.X.,Han J.L.,SuJ.,Wang L.,and Pan Q.H.,2012,The identification of Pi50(t),a new member of the rice blast resistance Pi2/Pi9 multigenefamily,Theor.Appl.Genet.,124(7):1295-1304