Fine-Mapping of qTGW1.2a, a Quantitative Trait Locus for 1000-Grain Weight in Rice
|
摘要
Thousand-grain weight(TGW) is a key component of grain yield in rice. This study was conducted to validate and fine-map qTGW1.2a, a quantitative trait locus for grain weight and grain size previously located in a 933.6-kb region on the long arm of rice chromosome 1. Firstly, three residual heterozygotes(RHs) were selected from a BC_2F_(11) population of the indica rice cross Zhenshan 97(ZS97)///ZS97//ZS97/Milyang 46. The heterozygous segments in these RHs were arranged successively in physical positions, forming one set of sequential residual heterozygotes(SeqRHs). In each of the populations derived, non-recombinant homozygotes were identified to produce near isogenic lines(NILs)comprising the two homozygous genotypes. The NILs were tested for grain weight, grain length and grain width. QTL analyses for the three traits were performed. Then, the updated QTL location was followed for a new run of SeqRHs identification-NIL development-QTL mapping. Altogether, 11 NIL populations derived from four sets of SeqRHs were developed and used. qTGW1.2a was finally delimitated into a77.5-kb region containing 13 annotated genes. In the six populations segregating this QTL, which were in four generations and were tested across four years, the allelic direction of qTGW1.2a remained consistent and the genetic effects were stable. For TGW, the additive effects ranged from 0.23 to 0.38 g and the proportions of phenotypic variance explained ranged from 26.15% to 41.65%. These results provide a good foundation for the cloning and functional analysis of qTGW1.2a.
Thousand-grain weight(TGW) is a key component of grain yield in rice. This study was conducted to validate and fine-map qTGW1.2a, a quantitative trait locus for grain weight and grain size previously located in a 933.6-kb region on the long arm of rice chromosome 1. Firstly, three residual heterozygotes(RHs) were selected from a BC_2F_(11) population of the indica rice cross Zhenshan 97(ZS97)///ZS97//ZS97/Milyang 46. The heterozygous segments in these RHs were arranged successively in physical positions, forming one set of sequential residual heterozygotes(SeqRHs). In each of the populations derived, non-recombinant homozygotes were identified to produce near isogenic lines(NILs)comprising the two homozygous genotypes. The NILs were tested for grain weight, grain length and grain width. QTL analyses for the three traits were performed. Then, the updated QTL location was followed for a new run of SeqRHs identification-NIL development-QTL mapping. Altogether, 11 NIL populations derived from four sets of SeqRHs were developed and used. qTGW1.2a was finally delimitated into a77.5-kb region containing 13 annotated genes. In the six populations segregating this QTL, which were in four generations and were tested across four years, the allelic direction of qTGW1.2a remained consistent and the genetic effects were stable. For TGW, the additive effects ranged from 0.23 to 0.38 g and the proportions of phenotypic variance explained ranged from 26.15% to 41.65%. These results provide a good foundation for the cloning and functional analysis of qTGW1.2a.
Thousand-grain weight(TGW) is a key component of grain yield in rice. This study was conducted to validate and fine-map qTGW1.2a, a quantitative trait locus for grain weight and grain size previously located in a 933.6-kb region on the long arm of rice chromosome 1. Firstly, three residual heterozygotes(RHs) were selected from a BC_2F_(11) population of the indica rice cross Zhenshan 97(ZS97)///ZS97//ZS97/Milyang 46. The heterozygous segments in these RHs were arranged successively in physical positions, forming one set of sequential residual heterozygotes(SeqRHs). In each of the populations derived, non-recombinant homozygotes were identified to produce near isogenic lines(NILs)comprising the two homozygous genotypes. The NILs were tested for grain weight, grain length and grain width. QTL analyses for the three traits were performed. Then, the updated QTL location was followed for a new run of SeqRHs identification-NIL development-QTL mapping. Altogether, 11 NIL populations derived from four sets of SeqRHs were developed and used. qTGW1.2a was finally delimitated into a77.5-kb region containing 13 annotated genes. In the six populations segregating this QTL, which were in four generations and were tested across four years, the allelic direction of qTGW1.2a remained consistent and the genetic effects were stable. For TGW, the additive effects ranged from 0.23 to 0.38 g and the proportions of phenotypic variance explained ranged from 26.15% to 41.65%. These results provide a good foundation for the cloning and functional analysis of qTGW1.2a.
引文
Bai X F,Wu B,Xing Y Z.2012.Yield-related QTLs and their applications in rice genetic improvement.J Integr Plant Biol,54(5):300-311.
Chen J Y,Zhang H W,Zhang H L,Ying J Z,Ma L Y,Zhuang J Y.2018.Natural variation at q Hd1 affects heading date acceleration at high temperatures with pleiotropism for yield traits in rice.BMC Plant Biol,18:112.
Chen X,Temnykh S,Xu Y,Cho Y G,Mc Couch S R.1997.Development of a microsatellite framework map providing genome-wide coverage in rice(Oryza sativa L.).Theor Appl Genet,95(4):553-567.
Dai W M,Zhang K Q,Wu J R,Wang L,Duan B W,Zheng K L,Cai R,Zhuang J Y.2008.Validating a segment on the short arm of chromosome 6 responsible for genetic variation in the hull silicon content and yield traits of rice.Euphytica,160(3):317-324.
Dong Q,Zhang Z H,Wang L L,Zhu Y J,Fan Y Y,Mou T M,Ma LY,Zhuang J Y.2018.Dissection and fine-mapping of two QTLfor grain size linked in a 460-kb region on chromosome 1 of rice.Rice,11(1):44.
Duan P G,Ni S,Wang J M,Zhang B L,Xu R,Wang Y X,Chen HQ,Zhu X D,Li Y H.2015.Regulation of Os GRF4 by OsmiR396controls grain size and yield in rice.Nat Plants,2:15203.
Duan P G,Xu J S,Zeng D L,Zhang B L,Geng M F,Zhang G Z,Huang K,Huang L J,Xu R,Ge S,Qian Q,Li Y H.2017.Natural variation in the promoter of GSE5 contributes to grain size diversity in rice.Mol Plant,10(5):685-694.
Fan C C,Xing Y Z,Mao H L,Lu T T,Han B,Xu C G,Li X H,Zhang Q F.2006.GS3,a major QTL for grain length and weight and minor QTL for grain width and thickness in rice,encodes a putative transmembrane protein.Theor Appl Genet,112(6):1164-1171.
Hu J,Wang Y X,Fang Y X,Zeng L J,Xu J,Yu H P,Shi Z Y,Pan JJ,Zhang D,Kang S J,Zhu L,Dong G J,Guo L B,Zeng D L,Zhang G H,Xie L H,Xiong G S,Li J Y,Qian Q.2015.A rare allele of GS2 enhances grain size and grain yield in rice.Mol Plant,8(10):1455-1465.
Hu Z J,Lu S J,Wang M J,He H H,Sun L,Wang H R,Liu X H,Jiang L,Sun J L,Xin X Y,Kong W,Chu C C,Xue H W,Yang JS,Luo X J,Liu J X.2018.A novel QTL qTGW3 encodes the GSK3/SHAGGY-like kinase OsGSK5/OsSK41 that interacts with OsARF4 to negatively regulate grain size and weight in rice.Mol Plant,11(5):736-749.
Huang R Y,Jiang L R,Zheng J S,Wang T S,Wang H C,Huang YM,Hong Z L.2013.Genetic bases of rice grain shape:So many genes,so little known.Trends Plant Sci,18(4):218-226.
Ishimaru K,Hirotsu N,Madoka Y,Murakami N,Hara N,Onodera H,Kashiwagi T,Ujiie K,Shimizu B,Onishi A,Miyagawa H,Katoh E.2013.Loss of function of the IAA-glucose hydrolase gene TGW6 enhances rice grain weight and increases yield.Nat Genet,45(6):707-711.
Li N,Xu R,Duan P G,Li Y H.2018.Control of grain size in rice.Plant Reprod,31(3):237-251.
Li Y B,Fan C C,Xing Y Z,Jiang Y H,Luo L J,Sun L,Shao D,Xu C J,Li X H,Xiao J H,He Y Q,Zhang Q F.2011.Natural variation in GS5 plays an important role in regulating grain size and yield in rice.Nat Genet,43(12):1266-1269.
Liu J F,Chen J,Zheng X M,Wu F Q,Lin Q B,Heng Y Q,Tian P,Cheng Z J,Yu X W,Zhou K N,Zhang X,Guo X P,Wang J L,Wang H Y,Wan J M.2017.GW5 acts in the brassinosteroid signalling pathway to regulate grain width and weight in rice.Nat Plants,3:17043.
Liu Q,Han R X,Wu K,Zhang J Q,Ye Y F,Wang S S,Chen J F,Pan Y J,Li Q,Xu X P,Zhou J W,Tao D Y,Wu Y J,Fu X D.2018.G-proteinβγsubunits determine grain size through interaction with MADS-domain transcription factors in rice.Nat Commun,9:852.
Mackay T F C,Stone E A,Ayroles J F.2009.The genetics of quantitative traits:Challenges and prospects.Nat Rev Genet,10(8):565-577.
Matsubara K,Ogiso-Tanaka E,Hori K,Ebana K,Ando T,Yano M.2012.Natural variation in Hd17,a homolog of Arabidopsis ELF3 that is involved in rice photoperiodic flowering.Plant Cell Physiol,53(4):709-716.
Qi L,Ding Y B,Zheng X M,Xu R,Zhang L Z,Wang Y Y,Wang XN,Zhang L F,Cheng Y L,Yang Q W.2018.Fine mapping and identification of a novel locus q GL12.2 control grain length in wild rice(Oryza rufipogon Griff.).Theor Appl Genet,131(7):1497-1508.
Qi P,Lin Y S,Song X J,Shen J B,Huang W,Shan J X,Zhu M Z,Jiang L,Gao J P,Lin H X.2012.The novel quantitative trait locus GL3.1 controls rice grain size and yield by regulating Cyclin-T1;3.Cell Res,22(12):1666-1680.
SAS Institute Inc.1999.SAS/STAT User’s Guide.Cary:SASInstitute.
Shao G N,Tang S Q,Luo J,Jiao G A,Wei X J,Tang A,Wu J L,Zhuang J Y,Hu P S.2010.Mapping of q GL7-2,a grain length QTL on chromosome 7 of rice.J Genet Genom,37(8):523-531.
Si L Z,Chen J Y,Huang X H,Gong H,Luo J H,Hou Q Q,Zhou TY,Lu T T,Zhu J J,Shangguan Y Y,Chen E W,Gong G X,Zhao Q,Jing Y F,Zhao Y,Li Y,Cui L L,Fan D L,Lu Y Q,Weng Q J,Wang Y C,Zhan Q L,Liu K Y,Wei X H,An K,An G,Han B.2016.OsSPL13 controls grain size in cultivated rice.Nat Genet,48:447-456.
Song X J,Huang W,Shi M,Zhu M Z,Lin H X.2007.A QTL for rice grain width and weight encodes a previously unknown RING-type E3 ubiquitin ligase.Nat Genet,39:623-630.
Song X J,Kuroha T,Ayano M,Furuta T,Nagai K,Komeda N,Segami S,Miura K,Ogawa D,Kamura T,Suzuki T,Higashiyama T,Yamasaki M,Mori H,Inukai Y,Wu J Z,Kitano H,Sakakibara H,Jacobsen S E,Ashikari M.2015.Rare allele of a previously unidentified histone H4 acetyltransferase enhances grain weight,yield,and plant biomass in rice.Proc Natl Acad Sci USA,112(1):76-81.
Wang L L,Chen Y Y,Guo L,Zhang H W,Fan Y Y,Zhuang J Y.2015.Dissection of qTGW1.2 to three QTLs for grain weight and grain size in rice(Oryza sativa L.).Euphytica,202(1):119-127.
Wang S K,Wu K,Yuan Q B,Liu X Y,Liu Z B,Lin X,Y Zeng R Z,Zhu H T,Dong G J,Qian Q,Zhang G Q,Fu X D.2012.Control of grain size,shape and quality by OsSPL16 in rice.Nat Genet,44(8):950-954.
Wang S K,Li S,Liu Q,Wu K,Zhang J Q,Wang S S,Wang Y,Chen X B,Zhang Y,Gao C X,Wang F,Huang H X,Fu X D.2015.The OsSPL16-GW7 regulatory module determines grain shape and simultaneously improves rice yield and grain quality.Nat Genet,47(8):949-954.
Wang Y X,Xiong G S,Hu J,Jiang L,Yu H,Xu J,Fang Y X,Zeng L J,Xu E B,Xu J,Ye W J,Meng X B,Liu R F,Chen H Q,Jing Y H,Wang Y H,Zhu X D,Li J Y,Qian Q.2015.Copy number variation at the GL7 locus contributes to grain size diversity in rice.Nat Genet,47(8):944-948.
Wu W G,Liu X Y,Wang M H,Meyer R S,Luo X J,Ndjiondjop MN,Tan L B,Zhang J W,Wu J Z,Cai H W,Sun C Q,Wang X K,Wing R A,Zhu Z F.2017.A single-nucleotide polymorphism causes smaller grain size and loss of seed shattering during African rice domestication.Nat Plants,3:17064.
Wu W X,Zheng X M,Lu G W,Zhong Z Z,Gao H,Chen L P,Wu C Y,Wang H J,Wang Q,Zhou K N,Wang J L,Wu F Q,Zhang X,Guo X P,Cheng Z J,Lei C L,Lin Q B,Jiang L,Wang H Y,Ge S,Wan J M.2013.Association of functional nucleotide polymorphisms at DTH2 with the northward expansion of rice cultivation in Asia.Proc Natl Acad Sci USA,110(8):2775-2780.
Xia D,Zhou H,Liu R J,Dan W H,Li P B,Wu B,Chen J X,Wang L Q,Gao G J,Zhang Q L,He Y Q.2018.GL3.3,a novel QTLencoding a GSK3/SHAGGY-like kinase,epistatically interacts with GS3 to form extra-long grains in rice.Mol Plant,11(5):754-756.
Xu C J,Liu Y,Li Y B,Xu X D,Xu C G,Li X H,Xiao J H,Zhang Q F.2015.Differential expression of GS5 regulates grain size in rice.J Exp Bot,66(9):2611-2623.
Ying J Z,Ma M,Bai C,Huang X H,Liu J L,Fang Y Y,Song X J.2018.TGW3,a major QTL that negatively modulates grain length and weight in rice.Mol Plant,11(5):750-753.
Yu J P,Xiong H Y,Zhu X Y,Zhang H L,Li H H,Miao J L,Wang W S,Tang Z S,Zhang Z Y,Yao G X,Zhang Q,Pan Y H,Wang X,Rashid M A R,Li J J,Gao Y M,Li Z K,Yang W C,Fu X D,Li Z C.2017.OsLG3 contributing to rice grain length and yield was mined by Ho-LAMap.BMC Biol,15:28.
Yu J P,Miao J L,Zhang Z Y,Xiong H Y,Zhu X Y,Sun X M,Pan YH,Liang Y T,Zhang Q,Rashid M A R,Li J J,Zhang H L,Li ZC.2018.Alternative splicing of OsLG3b controls grain length and yield in japonica rice.Plant Biotechnol J,16(9):1667-1678.
Zhang H W,Fan Y Y,Zhu Y J,Chen J Y,Yu S B,Zhuang J Y.2016.Dissection of the qTGW1.1 region into two tightly-linked minor QTLs having stable effects for grain weight in rice.BMC Genet,17(1):98.
Zhang X J,Wang J F,Huang J,Lan H X,Wang C L,Yin C F,Wu YY,Tang H J,Qian Q,Li J Y,Zhang H S.2012.Rare allele of OsPPKL1 associated with grain length causes extra-large grain and a significant yield increase in rice.Proc Natl Acad Sci USA,109:21534-21539.
Zhao D S,Li Q F,Zhang C Q,Zhang C,Yang Q Q,Pan L X,Ren X Y,Lu J,Gu M H,Liu Q Q.2018.GS9 acts as a transcriptional activator to regulate rice grain shape and appearance quality.Nat Commun,9:1240.
Zheng K L,Huang N,Bennett J,Khush G S.1995.PCR-ased marker-assisted selection in rice breeding.IRRI Discussion Paper Series No.12.Los Banos,the Phillipines:International Rice Research Institute.
Zhu Y J,Fan Y Y,Wang K,Huang D R,Liu W Z,Ying J Z,Zhuang J Y.2017.Rice Flowering Locus T1 plays an important role in heading date influencing yield traits in rice.Sci Rep,7:4918.
Zuo J R,Li J Y.2014.Molecular genetic dissection of quantitative trait loci regulating rice grain size.Annu Rev Genet,48:99-118.
Chen J Y,Zhang H W,Zhang H L,Ying J Z,Ma L Y,Zhuang J Y.2018.Natural variation at q Hd1 affects heading date acceleration at high temperatures with pleiotropism for yield traits in rice.BMC Plant Biol,18:112.
Chen X,Temnykh S,Xu Y,Cho Y G,Mc Couch S R.1997.Development of a microsatellite framework map providing genome-wide coverage in rice(Oryza sativa L.).Theor Appl Genet,95(4):553-567.
Dai W M,Zhang K Q,Wu J R,Wang L,Duan B W,Zheng K L,Cai R,Zhuang J Y.2008.Validating a segment on the short arm of chromosome 6 responsible for genetic variation in the hull silicon content and yield traits of rice.Euphytica,160(3):317-324.
Dong Q,Zhang Z H,Wang L L,Zhu Y J,Fan Y Y,Mou T M,Ma LY,Zhuang J Y.2018.Dissection and fine-mapping of two QTLfor grain size linked in a 460-kb region on chromosome 1 of rice.Rice,11(1):44.
Duan P G,Ni S,Wang J M,Zhang B L,Xu R,Wang Y X,Chen HQ,Zhu X D,Li Y H.2015.Regulation of Os GRF4 by OsmiR396controls grain size and yield in rice.Nat Plants,2:15203.
Duan P G,Xu J S,Zeng D L,Zhang B L,Geng M F,Zhang G Z,Huang K,Huang L J,Xu R,Ge S,Qian Q,Li Y H.2017.Natural variation in the promoter of GSE5 contributes to grain size diversity in rice.Mol Plant,10(5):685-694.
Fan C C,Xing Y Z,Mao H L,Lu T T,Han B,Xu C G,Li X H,Zhang Q F.2006.GS3,a major QTL for grain length and weight and minor QTL for grain width and thickness in rice,encodes a putative transmembrane protein.Theor Appl Genet,112(6):1164-1171.
Hu J,Wang Y X,Fang Y X,Zeng L J,Xu J,Yu H P,Shi Z Y,Pan JJ,Zhang D,Kang S J,Zhu L,Dong G J,Guo L B,Zeng D L,Zhang G H,Xie L H,Xiong G S,Li J Y,Qian Q.2015.A rare allele of GS2 enhances grain size and grain yield in rice.Mol Plant,8(10):1455-1465.
Hu Z J,Lu S J,Wang M J,He H H,Sun L,Wang H R,Liu X H,Jiang L,Sun J L,Xin X Y,Kong W,Chu C C,Xue H W,Yang JS,Luo X J,Liu J X.2018.A novel QTL qTGW3 encodes the GSK3/SHAGGY-like kinase OsGSK5/OsSK41 that interacts with OsARF4 to negatively regulate grain size and weight in rice.Mol Plant,11(5):736-749.
Huang R Y,Jiang L R,Zheng J S,Wang T S,Wang H C,Huang YM,Hong Z L.2013.Genetic bases of rice grain shape:So many genes,so little known.Trends Plant Sci,18(4):218-226.
Ishimaru K,Hirotsu N,Madoka Y,Murakami N,Hara N,Onodera H,Kashiwagi T,Ujiie K,Shimizu B,Onishi A,Miyagawa H,Katoh E.2013.Loss of function of the IAA-glucose hydrolase gene TGW6 enhances rice grain weight and increases yield.Nat Genet,45(6):707-711.
Li N,Xu R,Duan P G,Li Y H.2018.Control of grain size in rice.Plant Reprod,31(3):237-251.
Li Y B,Fan C C,Xing Y Z,Jiang Y H,Luo L J,Sun L,Shao D,Xu C J,Li X H,Xiao J H,He Y Q,Zhang Q F.2011.Natural variation in GS5 plays an important role in regulating grain size and yield in rice.Nat Genet,43(12):1266-1269.
Liu J F,Chen J,Zheng X M,Wu F Q,Lin Q B,Heng Y Q,Tian P,Cheng Z J,Yu X W,Zhou K N,Zhang X,Guo X P,Wang J L,Wang H Y,Wan J M.2017.GW5 acts in the brassinosteroid signalling pathway to regulate grain width and weight in rice.Nat Plants,3:17043.
Liu Q,Han R X,Wu K,Zhang J Q,Ye Y F,Wang S S,Chen J F,Pan Y J,Li Q,Xu X P,Zhou J W,Tao D Y,Wu Y J,Fu X D.2018.G-proteinβγsubunits determine grain size through interaction with MADS-domain transcription factors in rice.Nat Commun,9:852.
Mackay T F C,Stone E A,Ayroles J F.2009.The genetics of quantitative traits:Challenges and prospects.Nat Rev Genet,10(8):565-577.
Matsubara K,Ogiso-Tanaka E,Hori K,Ebana K,Ando T,Yano M.2012.Natural variation in Hd17,a homolog of Arabidopsis ELF3 that is involved in rice photoperiodic flowering.Plant Cell Physiol,53(4):709-716.
Qi L,Ding Y B,Zheng X M,Xu R,Zhang L Z,Wang Y Y,Wang XN,Zhang L F,Cheng Y L,Yang Q W.2018.Fine mapping and identification of a novel locus q GL12.2 control grain length in wild rice(Oryza rufipogon Griff.).Theor Appl Genet,131(7):1497-1508.
Qi P,Lin Y S,Song X J,Shen J B,Huang W,Shan J X,Zhu M Z,Jiang L,Gao J P,Lin H X.2012.The novel quantitative trait locus GL3.1 controls rice grain size and yield by regulating Cyclin-T1;3.Cell Res,22(12):1666-1680.
SAS Institute Inc.1999.SAS/STAT User’s Guide.Cary:SASInstitute.
Shao G N,Tang S Q,Luo J,Jiao G A,Wei X J,Tang A,Wu J L,Zhuang J Y,Hu P S.2010.Mapping of q GL7-2,a grain length QTL on chromosome 7 of rice.J Genet Genom,37(8):523-531.
Si L Z,Chen J Y,Huang X H,Gong H,Luo J H,Hou Q Q,Zhou TY,Lu T T,Zhu J J,Shangguan Y Y,Chen E W,Gong G X,Zhao Q,Jing Y F,Zhao Y,Li Y,Cui L L,Fan D L,Lu Y Q,Weng Q J,Wang Y C,Zhan Q L,Liu K Y,Wei X H,An K,An G,Han B.2016.OsSPL13 controls grain size in cultivated rice.Nat Genet,48:447-456.
Song X J,Huang W,Shi M,Zhu M Z,Lin H X.2007.A QTL for rice grain width and weight encodes a previously unknown RING-type E3 ubiquitin ligase.Nat Genet,39:623-630.
Song X J,Kuroha T,Ayano M,Furuta T,Nagai K,Komeda N,Segami S,Miura K,Ogawa D,Kamura T,Suzuki T,Higashiyama T,Yamasaki M,Mori H,Inukai Y,Wu J Z,Kitano H,Sakakibara H,Jacobsen S E,Ashikari M.2015.Rare allele of a previously unidentified histone H4 acetyltransferase enhances grain weight,yield,and plant biomass in rice.Proc Natl Acad Sci USA,112(1):76-81.
Wang L L,Chen Y Y,Guo L,Zhang H W,Fan Y Y,Zhuang J Y.2015.Dissection of qTGW1.2 to three QTLs for grain weight and grain size in rice(Oryza sativa L.).Euphytica,202(1):119-127.
Wang S K,Wu K,Yuan Q B,Liu X Y,Liu Z B,Lin X,Y Zeng R Z,Zhu H T,Dong G J,Qian Q,Zhang G Q,Fu X D.2012.Control of grain size,shape and quality by OsSPL16 in rice.Nat Genet,44(8):950-954.
Wang S K,Li S,Liu Q,Wu K,Zhang J Q,Wang S S,Wang Y,Chen X B,Zhang Y,Gao C X,Wang F,Huang H X,Fu X D.2015.The OsSPL16-GW7 regulatory module determines grain shape and simultaneously improves rice yield and grain quality.Nat Genet,47(8):949-954.
Wang Y X,Xiong G S,Hu J,Jiang L,Yu H,Xu J,Fang Y X,Zeng L J,Xu E B,Xu J,Ye W J,Meng X B,Liu R F,Chen H Q,Jing Y H,Wang Y H,Zhu X D,Li J Y,Qian Q.2015.Copy number variation at the GL7 locus contributes to grain size diversity in rice.Nat Genet,47(8):944-948.
Wu W G,Liu X Y,Wang M H,Meyer R S,Luo X J,Ndjiondjop MN,Tan L B,Zhang J W,Wu J Z,Cai H W,Sun C Q,Wang X K,Wing R A,Zhu Z F.2017.A single-nucleotide polymorphism causes smaller grain size and loss of seed shattering during African rice domestication.Nat Plants,3:17064.
Wu W X,Zheng X M,Lu G W,Zhong Z Z,Gao H,Chen L P,Wu C Y,Wang H J,Wang Q,Zhou K N,Wang J L,Wu F Q,Zhang X,Guo X P,Cheng Z J,Lei C L,Lin Q B,Jiang L,Wang H Y,Ge S,Wan J M.2013.Association of functional nucleotide polymorphisms at DTH2 with the northward expansion of rice cultivation in Asia.Proc Natl Acad Sci USA,110(8):2775-2780.
Xia D,Zhou H,Liu R J,Dan W H,Li P B,Wu B,Chen J X,Wang L Q,Gao G J,Zhang Q L,He Y Q.2018.GL3.3,a novel QTLencoding a GSK3/SHAGGY-like kinase,epistatically interacts with GS3 to form extra-long grains in rice.Mol Plant,11(5):754-756.
Xu C J,Liu Y,Li Y B,Xu X D,Xu C G,Li X H,Xiao J H,Zhang Q F.2015.Differential expression of GS5 regulates grain size in rice.J Exp Bot,66(9):2611-2623.
Ying J Z,Ma M,Bai C,Huang X H,Liu J L,Fang Y Y,Song X J.2018.TGW3,a major QTL that negatively modulates grain length and weight in rice.Mol Plant,11(5):750-753.
Yu J P,Xiong H Y,Zhu X Y,Zhang H L,Li H H,Miao J L,Wang W S,Tang Z S,Zhang Z Y,Yao G X,Zhang Q,Pan Y H,Wang X,Rashid M A R,Li J J,Gao Y M,Li Z K,Yang W C,Fu X D,Li Z C.2017.OsLG3 contributing to rice grain length and yield was mined by Ho-LAMap.BMC Biol,15:28.
Yu J P,Miao J L,Zhang Z Y,Xiong H Y,Zhu X Y,Sun X M,Pan YH,Liang Y T,Zhang Q,Rashid M A R,Li J J,Zhang H L,Li ZC.2018.Alternative splicing of OsLG3b controls grain length and yield in japonica rice.Plant Biotechnol J,16(9):1667-1678.
Zhang H W,Fan Y Y,Zhu Y J,Chen J Y,Yu S B,Zhuang J Y.2016.Dissection of the qTGW1.1 region into two tightly-linked minor QTLs having stable effects for grain weight in rice.BMC Genet,17(1):98.
Zhang X J,Wang J F,Huang J,Lan H X,Wang C L,Yin C F,Wu YY,Tang H J,Qian Q,Li J Y,Zhang H S.2012.Rare allele of OsPPKL1 associated with grain length causes extra-large grain and a significant yield increase in rice.Proc Natl Acad Sci USA,109:21534-21539.
Zhao D S,Li Q F,Zhang C Q,Zhang C,Yang Q Q,Pan L X,Ren X Y,Lu J,Gu M H,Liu Q Q.2018.GS9 acts as a transcriptional activator to regulate rice grain shape and appearance quality.Nat Commun,9:1240.
Zheng K L,Huang N,Bennett J,Khush G S.1995.PCR-ased marker-assisted selection in rice breeding.IRRI Discussion Paper Series No.12.Los Banos,the Phillipines:International Rice Research Institute.
Zhu Y J,Fan Y Y,Wang K,Huang D R,Liu W Z,Ying J Z,Zhuang J Y.2017.Rice Flowering Locus T1 plays an important role in heading date influencing yield traits in rice.Sci Rep,7:4918.
Zuo J R,Li J Y.2014.Molecular genetic dissection of quantitative trait loci regulating rice grain size.Annu Rev Genet,48:99-118.