Identification of New Resistance Loci Against Sheath Blight Disease in Rice Through Genome-Wide Association Study
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摘要
Sheath blight(SB) caused by the soil borne pathogen Rhizoctonia solani is one of the most serious global rice diseases. Breeding resistant cultivar is the most economical and effective strategy to control the disease. However, no rice varieties are completely resistant to SB, and only a few reliable quantitative trait loci(QTLs) linked with SB resistance have been identified to date. In this study, we conducted a genome-wide association study(GWAS) of SB resistance using 299 varieties from the rice diversity panel 1(RDP1) that were genotyped using 44 000 high-density single nucleotide polymorphism(SNP) markers. Through artificial inoculation, we found that only 36.5% of the tested varieties displayed resistance or moderate resistance to SB. In particular, the aromatic and aus sub-populations displayed higher SB resistance than the tropical japonica(TRJ), indica and temperate japonica sub-populations. Seven varieties showed similar resistance levels to the resistant control YSBR1. GWAS identified at least 11 SNP loci significantly associated with SB resistance in the three independent trials, leading to the identification of two reliable QTLs, qSB-3 and qSB-6, on chromosomes 3 and 6. Using favorable alleles or haplotypes of significantly associated SNP loci, we estimated that both QTLs had obvious effects on reducing SB disease severity and can be used for enhancing SB resistance, especially in improving SB resistance of TRJ sub-population rice varieties. These results provided important information and genetic materials for developing SB resistant varieties through breeding.
Sheath blight(SB) caused by the soil borne pathogen Rhizoctonia solani is one of the most serious global rice diseases. Breeding resistant cultivar is the most economical and effective strategy to control the disease. However, no rice varieties are completely resistant to SB, and only a few reliable quantitative trait loci(QTLs) linked with SB resistance have been identified to date. In this study, we conducted a genome-wide association study(GWAS) of SB resistance using 299 varieties from the rice diversity panel 1(RDP1) that were genotyped using 44 000 high-density single nucleotide polymorphism(SNP) markers. Through artificial inoculation, we found that only 36.5% of the tested varieties displayed resistance or moderate resistance to SB. In particular, the aromatic and aus sub-populations displayed higher SB resistance than the tropical japonica(TRJ), indica and temperate japonica sub-populations. Seven varieties showed similar resistance levels to the resistant control YSBR1. GWAS identified at least 11 SNP loci significantly associated with SB resistance in the three independent trials, leading to the identification of two reliable QTLs, qSB-3 and qSB-6, on chromosomes 3 and 6. Using favorable alleles or haplotypes of significantly associated SNP loci, we estimated that both QTLs had obvious effects on reducing SB disease severity and can be used for enhancing SB resistance, especially in improving SB resistance of TRJ sub-population rice varieties. These results provided important information and genetic materials for developing SB resistant varieties through breeding.
Sheath blight(SB) caused by the soil borne pathogen Rhizoctonia solani is one of the most serious global rice diseases. Breeding resistant cultivar is the most economical and effective strategy to control the disease. However, no rice varieties are completely resistant to SB, and only a few reliable quantitative trait loci(QTLs) linked with SB resistance have been identified to date. In this study, we conducted a genome-wide association study(GWAS) of SB resistance using 299 varieties from the rice diversity panel 1(RDP1) that were genotyped using 44 000 high-density single nucleotide polymorphism(SNP) markers. Through artificial inoculation, we found that only 36.5% of the tested varieties displayed resistance or moderate resistance to SB. In particular, the aromatic and aus sub-populations displayed higher SB resistance than the tropical japonica(TRJ), indica and temperate japonica sub-populations. Seven varieties showed similar resistance levels to the resistant control YSBR1. GWAS identified at least 11 SNP loci significantly associated with SB resistance in the three independent trials, leading to the identification of two reliable QTLs, qSB-3 and qSB-6, on chromosomes 3 and 6. Using favorable alleles or haplotypes of significantly associated SNP loci, we estimated that both QTLs had obvious effects on reducing SB disease severity and can be used for enhancing SB resistance, especially in improving SB resistance of TRJ sub-population rice varieties. These results provided important information and genetic materials for developing SB resistant varieties through breeding.
引文
Arun S,Annam M C,Shannonrm P,Josephl K,Arobert S,Rodantee T,Robert F.2009.Genetic mapping of sheath blight resistance QTLs within tropical japonica rice cultivars.Crop Sci,49(1):256-264.
Brachi B,Morris G P,Borevitz J O.2011.Genome-wide association studies in plants:The missing heritability is in the field.Genome Biol,12(10):232.
Bradbury P J,Zhang Z,Kroon D E,Casstevens T M,Ramdoss Y,Buckler E S.2007.TASSEL:Software for association mapping of complex traits in diverse samples.Bioinformatics,23(19):2633-2635.
Channamallikarjuna V,Sonah H,Prasad M,Rao G J N,Chand S,Upreti H C,Singh N K,Sharma T R.2010.Identification of major quantitative trait loci q SBR11-1 for sheath blight resistance in rice.Mol Breeding,25(1):155-166.
Che K P,Zhan Q C,Xing Q H,Wang Z P,Jin D M,He D J,Wang B.2003.Tagging and mapping of rice sheath blight resistant gene.Theor Appl Genet,106(2):293-297.
Chen X J,Wang L,Zuo S M,Wang Z B,Chen Z X,Zhang Y F,Lu G D,Zhou E X,Guo Z J,Huang S W,Pan X B.2009.Screening of varieties and isolates for identifying interaction between host and pathogen of rice sheath blight.Acta Phytopathol Sin,39:514-520.(in Chinese with English abstract)
Chen Z X,Zou J H,Xu J Y,Tong Y H,Tang S Z,Wang Z B,Jiang R M,Ling B,Tang J,Pan X B.2000.A preliminary study on resources of resistance to rice sheath blight.Chin J Rice Sci,14(1):15-18.(in Chinese with English abstract)
Chen Z X,Feng Z M,Wang L P,Feng F,Zhang Y F,Ma Y Y,Pan X B,Zuo S M.2017.Breeding potential of rice TAC1 gene for tiller angle.Chin J Rice Sci,31(6):590-598.(in Chinese with English abstract)
Christiansen T,Foy B D,Wall L,Orwant J.2012.Programming Perl:Unmatched power for text processing and scripting.Covers Version 5.14.Sebastopol,CA,USA:O’Reilly.
Eizenga G C,Ali M L,Bryant R J,Yeater K M,McClung A M,Mc Couch S R.2014.Registration of the rice diversity panel 1for genome-wide association studies.J Plant Reg,8(1):109.
Eizenga G C,Jia M H,Pinson S R,Gasore E R,Prasad B.2015.Exploring sheath blight quantitative trait loci in a Lemont/O.meridionalis advanced backcross population.Mol Breeding,35(6):1-19.
Famoso A N,Zhao K,Clark R T,Tung C W,Wright M H,Bustamante C,Kochian L V,Mc Couch S R.2011.Genetic architecture of aluminum tolerance in rice(Oryza sativa)determined through genome-wide association analysis and QTLmapping.PLo S Genet,7(8):e1002221.
Han Y P,Xing Y Z,Chen Z X,Gu S L,Pan X B,Chen X L,Zhang Q F.2002.Mapping QTLs for horizontal resistance to sheath blight in an elite rice restorer line,Minghui 63.Acta Genet Sin,29(7):622-626.(in Chinese with English abstract)
Han Y P,Xing Y Z,Gu S L,Chen Z X,Pan X B,Chen X L.2003.Effect of morphological traits on sheath blight resistance in rice.Acta Bot Sin,45(7):825-831.(in Chinese with English abstract)
Henderson C R.1975.Best linear unbiased estimation and prediction under a selection model.Biometrics,31(2):423-447.
Huang X H,Wei X H,Sang T,Zhao Q,Feng Q,Zhao Y,Li C Y,Zhu C R,Lu T T,Zhang Z W,Li M,Fan D L,Guo Y L,Wang A H,Wang L,Deng L W,Li W J,Lu Y Q,Weng Q J,Liu K Y,Huang T,Zhou T Y,Jing Y F,Li W,Lin Z,Buckler E S,Qian Q,Zhang Q F,Li J Y,Han B.2010.Genome-wide association studies of 14 agronomic traits in rice landraces.Nat Genet,42:961-967.
Jia L M,Yan W G,Zhu C S,Agrama H A,Jackson A,Yeater K,Li X,Huang B H,Hu B L,McClung A.2012.Allelic analysis of sheath blight resistance with association mapping in rice.PLoSOne,7(3):e32703.
Jia Y,Correa-Victoria F,McClung A,Zhu L,Liu G,Wamishe Y,Xie J,Marchetti M A,Pinson S R M,Rutger J N.2007.Rapid determination of rice cultivar responses to the sheath blight pathogen Rhizoctonia solani using a micro-chamber screening method.Plant Dis,91(5):485-489.
Jia Y,Liu G,Correa-Victoria F J,McClung A M,Oard J H,Bryant R J,Jia M H,Correll J C.2012.Registration of four rice germplasm lines with improved resistance to sheath blight and blast diseases.J Plant Reg,6(1):95-100.
Jia Y L,Liu G J,Costanzo S,Lee S,Dai Y T.2009.Current progress on genetic interactions of rice with rice blast and sheath blight fungi.Front Agric China,3(3):231-239.
Jiang S F,Wang C J Z,Shu C W,Zhou E X.2018.Cloning and expression analysis of RsPhm gene in Rhizoctonia solani AG-1IA of rice sheath blight pathogen.Chin J Rice Sci,32(2):111-118.(in Chinese with English abstract)
Kang H M,Sul J H,Service S K,Zaitlen N A,Kong S,Freimer NB,Sabatti C,Eskin E.2010.Variance component model to account for sample structure in genome-wide association studies.Nat Genet,42(4):348-354.
Kang H X,Wang Y,Peng S S,Zhang Y L,Xiao Y H,Wang D,Qu S H,Li Z Q,Yan S Y,Wang Z L,Liu W D,Ning Y S,Korniliev P,Leung H,Mezey J,Mc Couch S R,Wang G L.2016.Dissection of the genetic architecture of rice resistance to the blast fungus Magnaporthe oryzae.Mol Plant Pathol,17(6):959-972.
Kunihiro Y,Qian Q,Sato H,Teng S,Zeng D L,Fujimoto K,Zhu L H.2002.QTL analysis of sheath blight resistance in rice(Oryza sativa L.).Acta Genet Sin,29(1):50-55.(in Chinese with English abstract)
Lee F N,Rush M C.1983.Rice sheath blight:A major rice disease.Plant Dis,67(7):829-832.
Li H,Song C Y,Cong W B,Wang G L.2000.Evaluation and screening of resistance in keng rice varieties to sheath blight.Plant Prot,26(1):19-21.(in Chinese with English abstract)
Li Y,Xiao J H,Chen L L,Huang X H,Cheng Z K,Han B,Zhang Q F,Wu C Y.2018.Rice functional genomics research:Past decade and future.Mol Plant,11(3):359-380.
Li Z K,Pinson S R M,Marchetti M A,Stansel J W,Park W D.1995.Characterization of quantitative trait loci(QTLs)in cultivated rice contributing to field resistance to sheath blight(Rhizoctonia solani).Theor Appl Genet,91:382-388.
Liu G,Jia Y,Correa-Victoria F J,Prado G A,Yeater K M,Mcclung A,Correll J C.2009.Mapping quantitative trait loci responsible for resistance to sheath blight in rice.Phytopathology,99(9):1078-1084.
Liu W Z,Maccaferri M,Chen X m,Laghetti G,Pignone D,Pumphrey M,Tuberosa R.2017.Genome-wide association mapping reveals a rich genetic architecture of stripe rust resistance loci in emmer wheat(Triticum turgidum ssp.dicoccum).Theor Appl Genet,130(11):2249-2270.
Loan L C,Du P V,Li Z K.2004.Molecular dissection of quantitative resistance of sheath blight in rice(Oryza sativa L.).Omonrice,12:1-12.
Marchetti M A,Bollich C N.1991.Quantification of the relationship between sheath blight severity and yield loss in rice.Plant Dis,75(8):773.
Mc Couch S R,Zhao K Y,Wright M,Tung C W,Ebana K,Thomson M,Reynolds A,Wang D,Declerck G,Ali M L,Mc Clung A,Eizenga G,Bustamante C.2010.Development of genome-wide SNP assays for rice.Breeding Sci,60(5):524-535.
Meena B,Ramamoorthy V,Banu J G,Thangavelu R,Muthusamy M.2000.Screening of rice genotypes against sheath blight disease.J Ecobiol,12(2):103-109.
Morris G P,Ramu P,Deshpande S P,Hash C T,Shah T,Upadhyaya H D,Riera-Lizarazu O,Brown P J,Acharya C B,Mitchell S E,Harriman J,Glaubitz J C,Buckler E S,Kresovich S.2013.Population genomic and genome-wide association studies of agroclimatic traits in sorghum.Prol Natl Acad Sci USA,110(2):453-458.
Norton G J,Douglas A,Lahner B,Yakubova E,Guerinot M L,Pinson S R M,Tarpley L,Eizenga G C,Mcgrath S P,Zhao F J,Islam S,Duan G,Zhu Y G,Salt D E,Meharg A A,Price A H.2014.Genome wide association mapping of grain arsenic,copper,molybdenum and zinc in rice(Oryza sativa L.)grown at four international field sites.PLoS One,9(2):e89685.
Pan X B,Rush M C,Sha X Y,Xie Q J,Linscombe S D,Stetina SR,Oard J H.1999a.Major gene,nonallelic sheath blight resistance from the rice cultivars Jasmine 85 and Teqing.Crop Sci,39(2):338-346.
Pan X B,Zou J H,Chen Z X,Lu J F,Yu H X,Li H T,Wang Z B,Rush M C,Li Z.1999b.Molecular marker localization of the main QTLs of rice variety Jasmine 85 resistant to sheath blight.Chin Sci Bull,44:1629-1635.(in Chinese with English abstract)
Pan X B,Zou J H,Chen Z X,Lu J F,Yu H X,Li H T,Wang Z B,Pan X Y,Rush M C,Zhu L H.1999c.Tagging major quantitative trait loci for sheath blight resistance in a rice variety,Jasmine 85.Chin Sci Bull,44(19):1783-1789.(in Chinese with English abstract)
Pinson S R M,Capdevielle F M,Oard J H.2005.Confirming QTLs and finding additional loci conditioning sheath blight resistance in rice using recombinant inbred lines.Crop Sci,45(2):503-510.
Pinson S R M,Oard J H,Groth D,Miller R,Marchetti M A,Shank A R,Jia M H,Jia Y,Fjellstrom R G,Li Z.2008.Registration of TIL:455,TIL:514,and TIL:642,three rice germplasm lines containing introgressed sheath blight resistance alleles.J Plant Reg,2(3):251-254.
Prasad B,Eizenga G C.2008.Rice sheath blight disease resistance identified in Oryza spp.varieties.Plant Dis,92(11):1503-1509.
Sato H,Ideta O,Ando I,Kunihiro Y,Hirabayashi H,Iwano M,Miyasaka A,Nemoto H,Imbe T.2004.Mapping QTLs for sheath blight resistance in the rice line WSS2.Breeding Sci,54(3):265-271.
Rush M C.1999.Tagging major quantitative trait loci for sheath blight resistance in a rice variety,Jasmine 85.Chin Sci Bull,44:1783-1789.(in Chinese with English abstract).
Srinivasachary,Willocquet L,Savary S.2011.Resistance to rice sheath blight(Rhizoctonia solani Kühn)[(teleomorph:Thanatephorus cucumeris(A.B.Frank)Donk.]disease:Current status and perspectives.Euphytica,178(1):1-22.
Sun X T,Lu D D,Ou-yang L J,Hu L F,Bian J M,Peng X S,Chen X R,Fu J R,He X P,He H H,Zhu C L.2014.Association mapping and resistant alleles analysis for sheath blight resistance in rice.Acta Agron Sin,40(5):779.(in Chinese with English abstract)
Taguchi-Shiobara F,Ozaki H,Sato H,Maeda H,Kojima Y,Ebitani T,Yano M.2013.Mapping and validation of QTLs for rice sheath blight resistance.Breeding Sci,63(3):301-308.
Tan C X,Ji X M,Yang Y,Pan X Y,Zuo S M,Zhang Y F,Zou J H,Chen Z X,Zhu L H,Pan X B.2005.Identification and markerassisted selection of two major quantitative genes controlling rice sheath blight resistance in backcross generations.Acta Genet Sin,32(4):399-405.(in Chinese with English abstract)
Tung C W,Zhao K,Wright M H,Ali M L,Jung J,Kimball J,Tyagi W,Thomson M J,Mcnally K,Leung H,Kim H Ahn S N,Reynolds A,Scheffler B,Eizenga G,McClung A,Bustamante C,Mc Couch A R.2010.Development of a research platform for dissecting phenotype-genotype associations in rice(Oryza spp.).Rice,3(4):205-217.
Ueda Y,Frimpong F,Qi Y,Matthus E,Wu L,H?ller S,Kraska T,Frei M.2015.Genetic dissection of ozone tolerance in rice(Oryza sativa L.)by a genome-wide association study.J Exp Bot,66(1):293-306.
Wamishe Y A,Jia Y,Singh P,Cartwright R D.2007.Identification of field isolates of Rhizoctonia solani to detect quantitative resistance in rice under greenhouse conditions.Front Agric China,1(4):361-367.
Wang L,Huang W W,Liu L M,Fu Q,Huang S W.2011.Evaluation of resistance to sheath blight(Rhizoctonia solani)in some indica hybrid rice from southern China.Acta Agron Sin,37(2):263-270.(in Chinese with English abstract)
Wang Y,Prison S R M,Fjellstrom R G,Tabien R E.2012.Phenotypic gain from introgression of two QTL,q SB9-2 and qSB12-1,for rice sheath blight resistance.Mol Breeding,30(1):293-303.
Wang Z B,Zuo S M,Li G,Chen X J,Chen Z X,Zhang Y F,Pan XB.2009.Rapid identification technology of resistance to rice sheath blight in seedling stage.Acta Phytopathol Sin,39(2):174-182.(in Chinese with English abstract)
Wang Z B.2013.New exploration of inoculation identification system for rice sheath blight and analysis of resistance to sheath blight in new germplasm YSBR1[Ph D Thesis].Yangzhou:Yangzhou University.(in Chinese)
Zeng Y X,Li X M,Ma L Y,Ji Z J,Yang C D.2010.Research progress on mapping of gene conferring resistance to sheath blight and exploitation of resistance resources in rice.Chin JRice Sci,24(5):544-550.(in Chinese with English abstract)
Zhao K,Tung C W,Eizenga G C,Wright M H,Ali M L,Price A H,Norton G J,Islam M R,Reynolds A,Mezey J,McClung A M,Bustamante C D,McCouch S R.2011.Genome-wide association mapping reveals a rich genetic architecture of complex traits in Oryza sativa.Nat Commun,2:467.
Zhu D,Kang H,Li Z,Liu M,Zhu X,Wang Y,Wang D,Wang Z,Liu W,Wang G L.2016.A genome-wide association study of field resistance to Magnaporthe oryzae in rice.Rice,9(1):44.
Zou J H,Pan X B,Chen Z X,Xu J Y,Lu J F,Zhai W X,Zhu L H.2000.Mapping quantitative trait loci controlling sheath blight resistance in two rice cultivars(Oryza sativa L.).Theor Appl Genet,101:569-573.
Zuo S M,Wang Z B,Chen X J,Gu F,Zhan Y F,Chen Z X,Pan XB,Pan C H.2009,Evaluation of resistance of a novel rice germplasm YSBR1 to sheath blight.Acta Agron Sin,35(4):608-614.(in Chinese with English abstract)
Zuo S M,Zhang Y F,Chen Z X,Chen X J,Pan X B.2010.Current progress in genetics and breeding in resistance to rice sheath blight.Sci Sin,40:1014-1023.(in Chinese with English abstract)
Zuo S M,Yin Y J,Pan C H,Chen Z X,Zhang Y F,Gu S L,Zhu LH,Pan X B.2013.Fine mapping of q SB-11LE,the QTL that confers partial resistance to rice sheath blight.Theor Appl Genet,126(5):1257-1272.
Zuo S M,Zhang Y F,Yin Y J,Li G Z,Zhang G W,Wang H,Chen Z X,Pan X B.2014.Fine-mapping of q SB-9TQ,a gene conferring major quantitative resistance to rice sheath blight.Mol Breeding,34(4):2191-2203.
Brachi B,Morris G P,Borevitz J O.2011.Genome-wide association studies in plants:The missing heritability is in the field.Genome Biol,12(10):232.
Bradbury P J,Zhang Z,Kroon D E,Casstevens T M,Ramdoss Y,Buckler E S.2007.TASSEL:Software for association mapping of complex traits in diverse samples.Bioinformatics,23(19):2633-2635.
Channamallikarjuna V,Sonah H,Prasad M,Rao G J N,Chand S,Upreti H C,Singh N K,Sharma T R.2010.Identification of major quantitative trait loci q SBR11-1 for sheath blight resistance in rice.Mol Breeding,25(1):155-166.
Che K P,Zhan Q C,Xing Q H,Wang Z P,Jin D M,He D J,Wang B.2003.Tagging and mapping of rice sheath blight resistant gene.Theor Appl Genet,106(2):293-297.
Chen X J,Wang L,Zuo S M,Wang Z B,Chen Z X,Zhang Y F,Lu G D,Zhou E X,Guo Z J,Huang S W,Pan X B.2009.Screening of varieties and isolates for identifying interaction between host and pathogen of rice sheath blight.Acta Phytopathol Sin,39:514-520.(in Chinese with English abstract)
Chen Z X,Zou J H,Xu J Y,Tong Y H,Tang S Z,Wang Z B,Jiang R M,Ling B,Tang J,Pan X B.2000.A preliminary study on resources of resistance to rice sheath blight.Chin J Rice Sci,14(1):15-18.(in Chinese with English abstract)
Chen Z X,Feng Z M,Wang L P,Feng F,Zhang Y F,Ma Y Y,Pan X B,Zuo S M.2017.Breeding potential of rice TAC1 gene for tiller angle.Chin J Rice Sci,31(6):590-598.(in Chinese with English abstract)
Christiansen T,Foy B D,Wall L,Orwant J.2012.Programming Perl:Unmatched power for text processing and scripting.Covers Version 5.14.Sebastopol,CA,USA:O’Reilly.
Eizenga G C,Ali M L,Bryant R J,Yeater K M,McClung A M,Mc Couch S R.2014.Registration of the rice diversity panel 1for genome-wide association studies.J Plant Reg,8(1):109.
Eizenga G C,Jia M H,Pinson S R,Gasore E R,Prasad B.2015.Exploring sheath blight quantitative trait loci in a Lemont/O.meridionalis advanced backcross population.Mol Breeding,35(6):1-19.
Famoso A N,Zhao K,Clark R T,Tung C W,Wright M H,Bustamante C,Kochian L V,Mc Couch S R.2011.Genetic architecture of aluminum tolerance in rice(Oryza sativa)determined through genome-wide association analysis and QTLmapping.PLo S Genet,7(8):e1002221.
Han Y P,Xing Y Z,Chen Z X,Gu S L,Pan X B,Chen X L,Zhang Q F.2002.Mapping QTLs for horizontal resistance to sheath blight in an elite rice restorer line,Minghui 63.Acta Genet Sin,29(7):622-626.(in Chinese with English abstract)
Han Y P,Xing Y Z,Gu S L,Chen Z X,Pan X B,Chen X L.2003.Effect of morphological traits on sheath blight resistance in rice.Acta Bot Sin,45(7):825-831.(in Chinese with English abstract)
Henderson C R.1975.Best linear unbiased estimation and prediction under a selection model.Biometrics,31(2):423-447.
Huang X H,Wei X H,Sang T,Zhao Q,Feng Q,Zhao Y,Li C Y,Zhu C R,Lu T T,Zhang Z W,Li M,Fan D L,Guo Y L,Wang A H,Wang L,Deng L W,Li W J,Lu Y Q,Weng Q J,Liu K Y,Huang T,Zhou T Y,Jing Y F,Li W,Lin Z,Buckler E S,Qian Q,Zhang Q F,Li J Y,Han B.2010.Genome-wide association studies of 14 agronomic traits in rice landraces.Nat Genet,42:961-967.
Jia L M,Yan W G,Zhu C S,Agrama H A,Jackson A,Yeater K,Li X,Huang B H,Hu B L,McClung A.2012.Allelic analysis of sheath blight resistance with association mapping in rice.PLoSOne,7(3):e32703.
Jia Y,Correa-Victoria F,McClung A,Zhu L,Liu G,Wamishe Y,Xie J,Marchetti M A,Pinson S R M,Rutger J N.2007.Rapid determination of rice cultivar responses to the sheath blight pathogen Rhizoctonia solani using a micro-chamber screening method.Plant Dis,91(5):485-489.
Jia Y,Liu G,Correa-Victoria F J,McClung A M,Oard J H,Bryant R J,Jia M H,Correll J C.2012.Registration of four rice germplasm lines with improved resistance to sheath blight and blast diseases.J Plant Reg,6(1):95-100.
Jia Y L,Liu G J,Costanzo S,Lee S,Dai Y T.2009.Current progress on genetic interactions of rice with rice blast and sheath blight fungi.Front Agric China,3(3):231-239.
Jiang S F,Wang C J Z,Shu C W,Zhou E X.2018.Cloning and expression analysis of RsPhm gene in Rhizoctonia solani AG-1IA of rice sheath blight pathogen.Chin J Rice Sci,32(2):111-118.(in Chinese with English abstract)
Kang H M,Sul J H,Service S K,Zaitlen N A,Kong S,Freimer NB,Sabatti C,Eskin E.2010.Variance component model to account for sample structure in genome-wide association studies.Nat Genet,42(4):348-354.
Kang H X,Wang Y,Peng S S,Zhang Y L,Xiao Y H,Wang D,Qu S H,Li Z Q,Yan S Y,Wang Z L,Liu W D,Ning Y S,Korniliev P,Leung H,Mezey J,Mc Couch S R,Wang G L.2016.Dissection of the genetic architecture of rice resistance to the blast fungus Magnaporthe oryzae.Mol Plant Pathol,17(6):959-972.
Kunihiro Y,Qian Q,Sato H,Teng S,Zeng D L,Fujimoto K,Zhu L H.2002.QTL analysis of sheath blight resistance in rice(Oryza sativa L.).Acta Genet Sin,29(1):50-55.(in Chinese with English abstract)
Lee F N,Rush M C.1983.Rice sheath blight:A major rice disease.Plant Dis,67(7):829-832.
Li H,Song C Y,Cong W B,Wang G L.2000.Evaluation and screening of resistance in keng rice varieties to sheath blight.Plant Prot,26(1):19-21.(in Chinese with English abstract)
Li Y,Xiao J H,Chen L L,Huang X H,Cheng Z K,Han B,Zhang Q F,Wu C Y.2018.Rice functional genomics research:Past decade and future.Mol Plant,11(3):359-380.
Li Z K,Pinson S R M,Marchetti M A,Stansel J W,Park W D.1995.Characterization of quantitative trait loci(QTLs)in cultivated rice contributing to field resistance to sheath blight(Rhizoctonia solani).Theor Appl Genet,91:382-388.
Liu G,Jia Y,Correa-Victoria F J,Prado G A,Yeater K M,Mcclung A,Correll J C.2009.Mapping quantitative trait loci responsible for resistance to sheath blight in rice.Phytopathology,99(9):1078-1084.
Liu W Z,Maccaferri M,Chen X m,Laghetti G,Pignone D,Pumphrey M,Tuberosa R.2017.Genome-wide association mapping reveals a rich genetic architecture of stripe rust resistance loci in emmer wheat(Triticum turgidum ssp.dicoccum).Theor Appl Genet,130(11):2249-2270.
Loan L C,Du P V,Li Z K.2004.Molecular dissection of quantitative resistance of sheath blight in rice(Oryza sativa L.).Omonrice,12:1-12.
Marchetti M A,Bollich C N.1991.Quantification of the relationship between sheath blight severity and yield loss in rice.Plant Dis,75(8):773.
Mc Couch S R,Zhao K Y,Wright M,Tung C W,Ebana K,Thomson M,Reynolds A,Wang D,Declerck G,Ali M L,Mc Clung A,Eizenga G,Bustamante C.2010.Development of genome-wide SNP assays for rice.Breeding Sci,60(5):524-535.
Meena B,Ramamoorthy V,Banu J G,Thangavelu R,Muthusamy M.2000.Screening of rice genotypes against sheath blight disease.J Ecobiol,12(2):103-109.
Morris G P,Ramu P,Deshpande S P,Hash C T,Shah T,Upadhyaya H D,Riera-Lizarazu O,Brown P J,Acharya C B,Mitchell S E,Harriman J,Glaubitz J C,Buckler E S,Kresovich S.2013.Population genomic and genome-wide association studies of agroclimatic traits in sorghum.Prol Natl Acad Sci USA,110(2):453-458.
Norton G J,Douglas A,Lahner B,Yakubova E,Guerinot M L,Pinson S R M,Tarpley L,Eizenga G C,Mcgrath S P,Zhao F J,Islam S,Duan G,Zhu Y G,Salt D E,Meharg A A,Price A H.2014.Genome wide association mapping of grain arsenic,copper,molybdenum and zinc in rice(Oryza sativa L.)grown at four international field sites.PLoS One,9(2):e89685.
Pan X B,Rush M C,Sha X Y,Xie Q J,Linscombe S D,Stetina SR,Oard J H.1999a.Major gene,nonallelic sheath blight resistance from the rice cultivars Jasmine 85 and Teqing.Crop Sci,39(2):338-346.
Pan X B,Zou J H,Chen Z X,Lu J F,Yu H X,Li H T,Wang Z B,Rush M C,Li Z.1999b.Molecular marker localization of the main QTLs of rice variety Jasmine 85 resistant to sheath blight.Chin Sci Bull,44:1629-1635.(in Chinese with English abstract)
Pan X B,Zou J H,Chen Z X,Lu J F,Yu H X,Li H T,Wang Z B,Pan X Y,Rush M C,Zhu L H.1999c.Tagging major quantitative trait loci for sheath blight resistance in a rice variety,Jasmine 85.Chin Sci Bull,44(19):1783-1789.(in Chinese with English abstract)
Pinson S R M,Capdevielle F M,Oard J H.2005.Confirming QTLs and finding additional loci conditioning sheath blight resistance in rice using recombinant inbred lines.Crop Sci,45(2):503-510.
Pinson S R M,Oard J H,Groth D,Miller R,Marchetti M A,Shank A R,Jia M H,Jia Y,Fjellstrom R G,Li Z.2008.Registration of TIL:455,TIL:514,and TIL:642,three rice germplasm lines containing introgressed sheath blight resistance alleles.J Plant Reg,2(3):251-254.
Prasad B,Eizenga G C.2008.Rice sheath blight disease resistance identified in Oryza spp.varieties.Plant Dis,92(11):1503-1509.
Sato H,Ideta O,Ando I,Kunihiro Y,Hirabayashi H,Iwano M,Miyasaka A,Nemoto H,Imbe T.2004.Mapping QTLs for sheath blight resistance in the rice line WSS2.Breeding Sci,54(3):265-271.
Rush M C.1999.Tagging major quantitative trait loci for sheath blight resistance in a rice variety,Jasmine 85.Chin Sci Bull,44:1783-1789.(in Chinese with English abstract).
Srinivasachary,Willocquet L,Savary S.2011.Resistance to rice sheath blight(Rhizoctonia solani Kühn)[(teleomorph:Thanatephorus cucumeris(A.B.Frank)Donk.]disease:Current status and perspectives.Euphytica,178(1):1-22.
Sun X T,Lu D D,Ou-yang L J,Hu L F,Bian J M,Peng X S,Chen X R,Fu J R,He X P,He H H,Zhu C L.2014.Association mapping and resistant alleles analysis for sheath blight resistance in rice.Acta Agron Sin,40(5):779.(in Chinese with English abstract)
Taguchi-Shiobara F,Ozaki H,Sato H,Maeda H,Kojima Y,Ebitani T,Yano M.2013.Mapping and validation of QTLs for rice sheath blight resistance.Breeding Sci,63(3):301-308.
Tan C X,Ji X M,Yang Y,Pan X Y,Zuo S M,Zhang Y F,Zou J H,Chen Z X,Zhu L H,Pan X B.2005.Identification and markerassisted selection of two major quantitative genes controlling rice sheath blight resistance in backcross generations.Acta Genet Sin,32(4):399-405.(in Chinese with English abstract)
Tung C W,Zhao K,Wright M H,Ali M L,Jung J,Kimball J,Tyagi W,Thomson M J,Mcnally K,Leung H,Kim H Ahn S N,Reynolds A,Scheffler B,Eizenga G,McClung A,Bustamante C,Mc Couch A R.2010.Development of a research platform for dissecting phenotype-genotype associations in rice(Oryza spp.).Rice,3(4):205-217.
Ueda Y,Frimpong F,Qi Y,Matthus E,Wu L,H?ller S,Kraska T,Frei M.2015.Genetic dissection of ozone tolerance in rice(Oryza sativa L.)by a genome-wide association study.J Exp Bot,66(1):293-306.
Wamishe Y A,Jia Y,Singh P,Cartwright R D.2007.Identification of field isolates of Rhizoctonia solani to detect quantitative resistance in rice under greenhouse conditions.Front Agric China,1(4):361-367.
Wang L,Huang W W,Liu L M,Fu Q,Huang S W.2011.Evaluation of resistance to sheath blight(Rhizoctonia solani)in some indica hybrid rice from southern China.Acta Agron Sin,37(2):263-270.(in Chinese with English abstract)
Wang Y,Prison S R M,Fjellstrom R G,Tabien R E.2012.Phenotypic gain from introgression of two QTL,q SB9-2 and qSB12-1,for rice sheath blight resistance.Mol Breeding,30(1):293-303.
Wang Z B,Zuo S M,Li G,Chen X J,Chen Z X,Zhang Y F,Pan XB.2009.Rapid identification technology of resistance to rice sheath blight in seedling stage.Acta Phytopathol Sin,39(2):174-182.(in Chinese with English abstract)
Wang Z B.2013.New exploration of inoculation identification system for rice sheath blight and analysis of resistance to sheath blight in new germplasm YSBR1[Ph D Thesis].Yangzhou:Yangzhou University.(in Chinese)
Zeng Y X,Li X M,Ma L Y,Ji Z J,Yang C D.2010.Research progress on mapping of gene conferring resistance to sheath blight and exploitation of resistance resources in rice.Chin JRice Sci,24(5):544-550.(in Chinese with English abstract)
Zhao K,Tung C W,Eizenga G C,Wright M H,Ali M L,Price A H,Norton G J,Islam M R,Reynolds A,Mezey J,McClung A M,Bustamante C D,McCouch S R.2011.Genome-wide association mapping reveals a rich genetic architecture of complex traits in Oryza sativa.Nat Commun,2:467.
Zhu D,Kang H,Li Z,Liu M,Zhu X,Wang Y,Wang D,Wang Z,Liu W,Wang G L.2016.A genome-wide association study of field resistance to Magnaporthe oryzae in rice.Rice,9(1):44.
Zou J H,Pan X B,Chen Z X,Xu J Y,Lu J F,Zhai W X,Zhu L H.2000.Mapping quantitative trait loci controlling sheath blight resistance in two rice cultivars(Oryza sativa L.).Theor Appl Genet,101:569-573.
Zuo S M,Wang Z B,Chen X J,Gu F,Zhan Y F,Chen Z X,Pan XB,Pan C H.2009,Evaluation of resistance of a novel rice germplasm YSBR1 to sheath blight.Acta Agron Sin,35(4):608-614.(in Chinese with English abstract)
Zuo S M,Zhang Y F,Chen Z X,Chen X J,Pan X B.2010.Current progress in genetics and breeding in resistance to rice sheath blight.Sci Sin,40:1014-1023.(in Chinese with English abstract)
Zuo S M,Yin Y J,Pan C H,Chen Z X,Zhang Y F,Gu S L,Zhu LH,Pan X B.2013.Fine mapping of q SB-11LE,the QTL that confers partial resistance to rice sheath blight.Theor Appl Genet,126(5):1257-1272.
Zuo S M,Zhang Y F,Yin Y J,Li G Z,Zhang G W,Wang H,Chen Z X,Pan X B.2014.Fine-mapping of q SB-9TQ,a gene conferring major quantitative resistance to rice sheath blight.Mol Breeding,34(4):2191-2203.