水稻抽穗扬花期耐热性QTL(qHTH10)的鉴定及遗传效应分析
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摘要
为了深入挖掘和利用元江野生稻(Oryza rufipogon Griff.)的优异等位基因,构建了一个元江普通野生稻‘荷花塘4号’为供体,籼稻‘天丰B’(O. sativa subsp. indica)为受体的BC5F3群体,以抽穗扬花期耐热性为目标表型性状进行考察。在利用SSR标记分析检测后,结合单标记相关分析的方法,在10号染色体上发现标记RM25216与抽穗扬花期耐热性相关极显著。为进一步分析目标片段,将含有元江野生稻目标渐渗片段的近等基因系YJ01-201 (BC5F3)与轮回亲本‘天丰B’回交,构建了BC6F2和BC6F3次级分离群体,通过连续2年的大田高温胁迫及人工气候室模拟高温胁迫试验,证实了目的 QTL存在于目标片段上。同时,在BC6F2、BC6F3世代群体中,通过QTL分析发现目标QTL (qHTH10)在标记区间(RM25216～RM25228)遗传效应稳定,物理图距为277.1 kb,两个世代分别解释的表型变异值为7.4%和15.2%,增效作用来自元江普通野生稻。本研究为水稻耐热育种提供了新资源,也为进一步精细定位和克隆q HTH10提供了科学依据。
To deep excavate and utili ze the excellent alleles in Yuanjiang Oryza rufipogon, a BC5 F3 population was developed using common Yuanjiang wild rice 'Hehuatang No.4' as the donor and Tianfeng B(O. sativa subsp.indica) as the receptor. The investigation was carried out using the heat tolerance at heading and flowering stage as the target phenotypic trait. After SSR marker analysis and single marker correlation analysis, it was found that the marker RM25216 on chromosome 10 was highly correlated with heat tolerance at heading and flowering stage. To further analyze the target fragment, a secondary isolation population of BC6 F2 and BC6 F3 was constructed by backcrossing near isogenic line YJ01-201(BC5 F3) containing target introgressive fragments of Yuanjiang wild rice with recurrent parent 'Tianfeng B'. Through two consecutive years of field high temperature stress and simulated high temperature stress in artificial climate chamber, it was confirmed that the target QTL existed on the target fragment. At the same time, in BC6 F2 and BC6 F3 generations, the genetic effects of the target QTL(qHTH10) were stable in the marker interval(RM25216-RM25228), and the physical map spacing was 277.1 kb. The phenotypic variances explained by the two generations were 7.4% and 15.2%, respectively. The synergies were from common Yuanjiang wild rice. This study could provide new resources for rice breeding with heat tolerance and scientific reference for fine mapping and cloning of qHTH10.
To deep excavate and utili ze the excellent alleles in Yuanjiang Oryza rufipogon, a BC5 F3 population was developed using common Yuanjiang wild rice 'Hehuatang No.4' as the donor and Tianfeng B(O. sativa subsp.indica) as the receptor. The investigation was carried out using the heat tolerance at heading and flowering stage as the target phenotypic trait. After SSR marker analysis and single marker correlation analysis, it was found that the marker RM25216 on chromosome 10 was highly correlated with heat tolerance at heading and flowering stage. To further analyze the target fragment, a secondary isolation population of BC6 F2 and BC6 F3 was constructed by backcrossing near isogenic line YJ01-201(BC5 F3) containing target introgressive fragments of Yuanjiang wild rice with recurrent parent 'Tianfeng B'. Through two consecutive years of field high temperature stress and simulated high temperature stress in artificial climate chamber, it was confirmed that the target QTL existed on the target fragment. At the same time, in BC6 F2 and BC6 F3 generations, the genetic effects of the target QTL(qHTH10) were stable in the marker interval(RM25216-RM25228), and the physical map spacing was 277.1 kb. The phenotypic variances explained by the two generations were 7.4% and 15.2%, respectively. The synergies were from common Yuanjiang wild rice. This study could provide new resources for rice breeding with heat tolerance and scientific reference for fine mapping and cloning of qHTH10.
引文
Buu B.C.,Ha P.T.T.,Tam B.P.,Nien T.T.,Hieu V.N.,Phuoc N.T.,the Minh L.,Ciang L.H.,and Lang N.T.,2014,Quantitative trait loci associated with heat tolerance in rice(Oryza sativa L.),Plant Breed.Biotech.,2(1):14-24
Cao L.Y.,Zhao J.G.,Zhan X.D.,Li D.L.,and He L.B.,2003,Mapping QTLs for heat tolerance and correlation between heat tolerance and photosynthetic rate in rice,Zhongguo Shuidao Kexue(Chinese Journal of Rice Science),17(3):223-227(曹立勇,赵建根,占小登,李登楼,何立斌,2003,水稻耐热性的QTL定位及耐热性与光合速率的相关性,中国水稻科学,17(3):223-227)
Hirabayashi H.,Sasaki K.,Kambe T.,Gannaban R.B.,Miras M.A.,Mendioro M.S.,Simon E.V.,Lumanglas P.D.,Fujita D.,Takemoto-Kuno Y.,Takeuchi Y.,Kaji R.,Kondo M.,Kobayashi N.,Ogawa T.,Ando I.,Jagadish K.S.V.,and Takeuchi Y.,2014,q EMF3,a novel QTL for the early-morning flowering trait from wild rice,Oryza officinalis,to mitigate heat stress damage at flowering in rice,O.sativa,J.Exp.Bot.,66(5):1227-1236
Jagadish S.V.K.,Cairns J.,Lafitte R.,Wheeler T.R.,Price A.H.,and Craufurd P.Q.,2010a,Genetic analysis of heat tolerance at anthesis in rice,Crop Sci.,50(5):1633-1641
Jagadish S.V.K.,Muthurajan R.,Oane R.,Wheeler T.R.,Heuer S.,Bennett J.,and Craufurd Q.P.,2010b,Physiological and proteomic approaches to address heat tolerance during anthesis in rice,J.Exp.Bot.,61(1):143-156
Jagadish S.V.K.,Craufurd P.Q.,and Wheeler T.R.,2007,High temperature stress and spikelet fertility in rice(Oryza sativa L.),J.Exp.Bot.,58(7):1627-1635
Kui L.M.,Tang L.B.,Tu J.,Lu Y.X.,and Sun C.Q.,2008,Identification of QTLs associated with heat tolerance of Yuanjiang common wild rice(Oryza rufipogon Griff.)at flowering stage,Nongye Shengwu Jishu Xuebao(Journal of Agricultural Biotechnology),16(3):461-464(奎丽梅,谭禄宾,涂建,卢义宣,孙传清,2008,云南元江野生稻抽穗开花期耐热QTL定位,农业生物技术学报,16(3):461-464)
Liao J.L.,Zhang H.Y.,Shao X.L.,Zhong P.A.,and Huang Y.J.,2011,Identification on heat tolerance of backcross recombinant rice lines and screening of backcross introgression lines with heat tolerance at milky stage,Rice Science,18(4):279-286
Liao J.L.,Zhou H.W.,Zhang H.Y.,Zhong P.A.,and Huang Y.J.,2013,Comparative proteomic analysis of differentially expressed proteins in the early milky stage of rice grains during high temperature stress,J.Exp.Bot.,65(2):655-671
Liu Q.,Yang T.,Yu T.,Zhang S.H.,Mao X.X.,Zhao J.L.,Wang X.F.,Dong J.F.,and Liu B.,2017,Integrating small RNAsequencing with QTL mapping for identification of mi RNAs and their target genes associated with heat tolerance at the flowering stage in rice,Front.Plant Sci.,8:43
Matsui T.,and Omasa K.,2002,Rice cultivars tolerant to high temperature at flowering:anther characteristics,Ann.Bot.,89(6):683-687
Morita S.,Yonemaru J.I.,and Takanashi J.I.,2005,Grain growth and endosperm cell size under high night temperatures in rice(Oryza sativa L.),Ann.Bot.,95(4):695-701
Pachauri R.K.,and Reisinger A.,2007,Climate change and its impacts in the near and long term under different scenarios,In:Climate Change:Synthesis Report,IPCC(Intergovernmental Panel on Climate Change)Geneva,Switzerland,pp.43-54
Pan Y.,Luo L.H.,Deng H.B.,Zhang G.L.,Tang W.B.,Chen L.Y.,and Xiao Y.H.,2011,Quantitative trait loci associated with pollen fertility under high temperature stress at flowering stage in rice,Zhongguo Shuidao Kexue(Chinese Journal of Rice Science),25(1):99-102(盘毅,罗丽华,邓化冰,张桂莲,唐文邦,陈立云,肖应辉,2011,水稻开花期高温胁迫下的花粉活力QTL定位,中国水稻科学,25(1):99-102)
Peng S.,Huang J.,Sheehy J.E.,Laza R.C.,Visperas R.M.,Zhong X.,Centeno G.S.,Khush G.S.,and Cassman K.G.,2004,Rice yield decline with higher night temperature from global warming,Proc.Natl.Acad.Sci.USA,101(27):9971-9975
Poli Y.,Basava R.K.,Panigrahy M.,Vinukonda V.P.,Dokula N.R.,Voleti S.R.,Desiraju S.,and Neelamraju S.,2013,Char acterization of a Nagina 22 rice mutant for heat tolerance and mapping of yield traits,Rice,6(1):36
Prasad P.V.V.,Boote K.J.,Allen L.H.,Sheehy J.E.,and Thomas J.M.G.,2006,Species,ecotype and cultivar differences in spikelet fertility and harvest index of rice in response to high temperature stress,Field Crops Res.,95(2):398-411
Prasanth V.V.,Basava K.R.,Babu M.S.,V.G.N.V.T.,Devi S.J.S.R.,Mangrauthia S.K.,Voleti S.R.,and Sarla N.,2016,Field level evaluation of rice introgression lines for heat tolerance and validation of markers linked to spikelet fertility,Physiol.Mol.Biol.Plants,22(2):179-192
Qin B.X.,Liu L.L.,Han F.Y.,Chen L.,Li N.,Huang M.,and Li R.B.,2015,Study on cold tolerance of chromosome segment substitution lines derived from wild rice(O.rufipogon)as donor at booting stage,Jiyinzuxue Yu Yingyong Shengwuxue(Genomics and Applied Biology),34(6):1283-1289(覃宝祥,刘立龙,韩飞怡,陈霖,李楠,黄敏,李容柏,2015,普通野生稻染色体片段代换系的孕穗期耐冷性研究,基因组学与应用生物学,34(6):1283-1289)
Roger S.O.,and Bendich A.J.,1994,Extraction of total cellular DNA from plants,algae and fungi,In:Plant Molecular Biology Manual,Kluwer Academic Publishers,Dordrecht,pp.183-190
Shanmugavadivel P.S.,SV A.M.,Prakash C.,MK R..,Tiwari R.,Mohapatra T.,and Singh N.K.,2017,High resolution mapping of QTLs for heat tolerance in rice using a 5K SNP array,Rice,10(1):28
Shi W.,Ishimaru T.,Gannaban R.B.,Oane W.,and Jagadish S.V.K.,2015,Popular rice(Oryza sativa L.)cultivars show contrasting responses to heat stress at gametogenesis and anthesis,Crop Sci.,55:589-596
Stocker T.,ed.,2013,Climate change 2013:the physical science basis,contribution of working group I to the fifth assessment report of the Intergovernmental Panel on Climate Change,U-niversity of Cambridge,Cambridge,United Kingdom,pp.5-14
Wissuwa M.,Wegner J.,Ae N.,and Yano M.,2002,Substitution mapping of Pup1:a major QTL increasing phosphorus uptake of rice from a phosphorus-deficient soil,Theor.Appl.Genet.,105(6-7):890-897
Xiao Y.H.,Pan Y.,Luo L.H.,Zhang G.L.,Deng H.B.,Dai L.Y.,Tang W.B.,Chen L.Y.,and Wang G.L.,2011,Quantitative trait loci associated with seed set under high temperature stress at the flowering stage in rice(Oryza sativa L.),Euphytica,178(3):331-338
Yang T.F.,Zhang S.H.,Wang X.F.,Huang Z.H.,Zhao J.L.,Zhang G.Q.,and Liu B.,2012,Screening for germplasm with heat tolerance at flowering stage in Oryza sativa,Huanan Nongye Daxue Xuebao(Journal of South China Agricultural University),33(4):585-588(杨梯丰,张少红,王晓飞,黄章慧,赵均良,张桂权,刘斌,2012,水稻抽穗开花期耐热种质资源的筛选鉴定,华南农业大学学报,33(4):585-588)
Ye C.,Argayoso M.A.,Redon E.D.,Sierra S.N.,Laza M.A.,Dilla C.J.,Mo Y.,Thomson M.J.,Chin J.,Delavin C.B.,Diaz G.Q.,and Hernandez J.E.,2011,Mapping QTL for heat tolerance at flowering stage in rice using SNP markers,Plant Breeding,131(1):33-41
Ye C.,Tenorio F.A.,Redo觡a E.D.,Morales-Cortezano P.S.,Cabrega G.A.,Jagadish K.S.V.,and Gregorio G.B.,2015,Finemapping and validating qHTSF4.1 to increase spikelet fertility under heat stress at flowering in rice,Theor.Appl.Genet.,128(8):1507-1517
Zhang G.L.,Chen L.Y.,Xiao G.Y.,Xiao Y.H.,Chen X.B.,and Zhang S.T.,2009,Bulked segregant analysis to detect QTLrelated to heat tolerance in rice(Oryza sativa L.)using SSRmarkers,Agricultural Sciences in China,8(4):482-487
Zhang T.,Yang L.,Jiang K.F.,Huang M.,Sun Q.,Chen W.F.,and Zheng J.K.,2008,QTL mapping for heat tolerance of the tassel period of rice,Fenzi Zhiwu Yuzhong(Molecular Plant Breeding),6(5):867-873(张涛,杨莉,蒋开锋,黄敏,孙群,陈温福,郑家奎,2008,水稻抽穗扬花期耐热性的QTL分析,分子植物育种,6(5):867-873)
Zhao L.,Lei J.G.,Huang Y.J.,Zhu S.,Chen H.P.,Huang R.L.,Peng Z.Q.,Tu Q.H.,Shen X.H.,and Yan S.,2016,Mapping quantitative trait loci for heat tolerance at anthesis in rice using chromosomal segment substitution lines,Breeding Sci.,66(3):358-366
Zhao Z.G.,Jiang L.,Xiao Y.H.,Zhang W.W.,Zhai H.Q.,and Wan J.M.,2006,Identification of QTLs for heat tolerance at the booting stage in rice(Oryza sativa L.),Zuowu Xuebao(Acta Agronomica Sinica),32(5):640-644(赵志刚,江玲,肖应辉,张文伟,翟虎渠,万建民,2006,水稻孕穗期耐热性QTLs分析,作物学报,32(5):640-644)
Zhou W.H.,Xue D.W.,and Zhang G.P.,2012,Identification and physiological characterization of thermo-tolerant rice genotypes,Zhejiang Daxue Xuebao(Journal of Zhejiang University(Agriculture and Life Sciences)),38(1):1-9(周伟辉,薛大伟,张国平,2012,耐热水稻基因型的鉴定及相关生理学特征分析,浙江大学学报(农业与生命科学版),38(1):1-9)
Zhu S.,Huang R.L.,Wai H.P.,Xiong H.L.,Shen X.H.,He H.H.,and Yan S.,2017,Mapping quantitative trait loci for heat tolerance at the booting stage using chromosomal segment substitution lines in rice,Physiol.Mol.Biol.Plants,23(4):817-825
Cao L.Y.,Zhao J.G.,Zhan X.D.,Li D.L.,and He L.B.,2003,Mapping QTLs for heat tolerance and correlation between heat tolerance and photosynthetic rate in rice,Zhongguo Shuidao Kexue(Chinese Journal of Rice Science),17(3):223-227(曹立勇,赵建根,占小登,李登楼,何立斌,2003,水稻耐热性的QTL定位及耐热性与光合速率的相关性,中国水稻科学,17(3):223-227)
Hirabayashi H.,Sasaki K.,Kambe T.,Gannaban R.B.,Miras M.A.,Mendioro M.S.,Simon E.V.,Lumanglas P.D.,Fujita D.,Takemoto-Kuno Y.,Takeuchi Y.,Kaji R.,Kondo M.,Kobayashi N.,Ogawa T.,Ando I.,Jagadish K.S.V.,and Takeuchi Y.,2014,q EMF3,a novel QTL for the early-morning flowering trait from wild rice,Oryza officinalis,to mitigate heat stress damage at flowering in rice,O.sativa,J.Exp.Bot.,66(5):1227-1236
Jagadish S.V.K.,Cairns J.,Lafitte R.,Wheeler T.R.,Price A.H.,and Craufurd P.Q.,2010a,Genetic analysis of heat tolerance at anthesis in rice,Crop Sci.,50(5):1633-1641
Jagadish S.V.K.,Muthurajan R.,Oane R.,Wheeler T.R.,Heuer S.,Bennett J.,and Craufurd Q.P.,2010b,Physiological and proteomic approaches to address heat tolerance during anthesis in rice,J.Exp.Bot.,61(1):143-156
Jagadish S.V.K.,Craufurd P.Q.,and Wheeler T.R.,2007,High temperature stress and spikelet fertility in rice(Oryza sativa L.),J.Exp.Bot.,58(7):1627-1635
Kui L.M.,Tang L.B.,Tu J.,Lu Y.X.,and Sun C.Q.,2008,Identification of QTLs associated with heat tolerance of Yuanjiang common wild rice(Oryza rufipogon Griff.)at flowering stage,Nongye Shengwu Jishu Xuebao(Journal of Agricultural Biotechnology),16(3):461-464(奎丽梅,谭禄宾,涂建,卢义宣,孙传清,2008,云南元江野生稻抽穗开花期耐热QTL定位,农业生物技术学报,16(3):461-464)
Liao J.L.,Zhang H.Y.,Shao X.L.,Zhong P.A.,and Huang Y.J.,2011,Identification on heat tolerance of backcross recombinant rice lines and screening of backcross introgression lines with heat tolerance at milky stage,Rice Science,18(4):279-286
Liao J.L.,Zhou H.W.,Zhang H.Y.,Zhong P.A.,and Huang Y.J.,2013,Comparative proteomic analysis of differentially expressed proteins in the early milky stage of rice grains during high temperature stress,J.Exp.Bot.,65(2):655-671
Liu Q.,Yang T.,Yu T.,Zhang S.H.,Mao X.X.,Zhao J.L.,Wang X.F.,Dong J.F.,and Liu B.,2017,Integrating small RNAsequencing with QTL mapping for identification of mi RNAs and their target genes associated with heat tolerance at the flowering stage in rice,Front.Plant Sci.,8:43
Matsui T.,and Omasa K.,2002,Rice cultivars tolerant to high temperature at flowering:anther characteristics,Ann.Bot.,89(6):683-687
Morita S.,Yonemaru J.I.,and Takanashi J.I.,2005,Grain growth and endosperm cell size under high night temperatures in rice(Oryza sativa L.),Ann.Bot.,95(4):695-701
Pachauri R.K.,and Reisinger A.,2007,Climate change and its impacts in the near and long term under different scenarios,In:Climate Change:Synthesis Report,IPCC(Intergovernmental Panel on Climate Change)Geneva,Switzerland,pp.43-54
Pan Y.,Luo L.H.,Deng H.B.,Zhang G.L.,Tang W.B.,Chen L.Y.,and Xiao Y.H.,2011,Quantitative trait loci associated with pollen fertility under high temperature stress at flowering stage in rice,Zhongguo Shuidao Kexue(Chinese Journal of Rice Science),25(1):99-102(盘毅,罗丽华,邓化冰,张桂莲,唐文邦,陈立云,肖应辉,2011,水稻开花期高温胁迫下的花粉活力QTL定位,中国水稻科学,25(1):99-102)
Peng S.,Huang J.,Sheehy J.E.,Laza R.C.,Visperas R.M.,Zhong X.,Centeno G.S.,Khush G.S.,and Cassman K.G.,2004,Rice yield decline with higher night temperature from global warming,Proc.Natl.Acad.Sci.USA,101(27):9971-9975
Poli Y.,Basava R.K.,Panigrahy M.,Vinukonda V.P.,Dokula N.R.,Voleti S.R.,Desiraju S.,and Neelamraju S.,2013,Char acterization of a Nagina 22 rice mutant for heat tolerance and mapping of yield traits,Rice,6(1):36
Prasad P.V.V.,Boote K.J.,Allen L.H.,Sheehy J.E.,and Thomas J.M.G.,2006,Species,ecotype and cultivar differences in spikelet fertility and harvest index of rice in response to high temperature stress,Field Crops Res.,95(2):398-411
Prasanth V.V.,Basava K.R.,Babu M.S.,V.G.N.V.T.,Devi S.J.S.R.,Mangrauthia S.K.,Voleti S.R.,and Sarla N.,2016,Field level evaluation of rice introgression lines for heat tolerance and validation of markers linked to spikelet fertility,Physiol.Mol.Biol.Plants,22(2):179-192
Qin B.X.,Liu L.L.,Han F.Y.,Chen L.,Li N.,Huang M.,and Li R.B.,2015,Study on cold tolerance of chromosome segment substitution lines derived from wild rice(O.rufipogon)as donor at booting stage,Jiyinzuxue Yu Yingyong Shengwuxue(Genomics and Applied Biology),34(6):1283-1289(覃宝祥,刘立龙,韩飞怡,陈霖,李楠,黄敏,李容柏,2015,普通野生稻染色体片段代换系的孕穗期耐冷性研究,基因组学与应用生物学,34(6):1283-1289)
Roger S.O.,and Bendich A.J.,1994,Extraction of total cellular DNA from plants,algae and fungi,In:Plant Molecular Biology Manual,Kluwer Academic Publishers,Dordrecht,pp.183-190
Shanmugavadivel P.S.,SV A.M.,Prakash C.,MK R..,Tiwari R.,Mohapatra T.,and Singh N.K.,2017,High resolution mapping of QTLs for heat tolerance in rice using a 5K SNP array,Rice,10(1):28
Shi W.,Ishimaru T.,Gannaban R.B.,Oane W.,and Jagadish S.V.K.,2015,Popular rice(Oryza sativa L.)cultivars show contrasting responses to heat stress at gametogenesis and anthesis,Crop Sci.,55:589-596
Stocker T.,ed.,2013,Climate change 2013:the physical science basis,contribution of working group I to the fifth assessment report of the Intergovernmental Panel on Climate Change,U-niversity of Cambridge,Cambridge,United Kingdom,pp.5-14
Wissuwa M.,Wegner J.,Ae N.,and Yano M.,2002,Substitution mapping of Pup1:a major QTL increasing phosphorus uptake of rice from a phosphorus-deficient soil,Theor.Appl.Genet.,105(6-7):890-897
Xiao Y.H.,Pan Y.,Luo L.H.,Zhang G.L.,Deng H.B.,Dai L.Y.,Tang W.B.,Chen L.Y.,and Wang G.L.,2011,Quantitative trait loci associated with seed set under high temperature stress at the flowering stage in rice(Oryza sativa L.),Euphytica,178(3):331-338
Yang T.F.,Zhang S.H.,Wang X.F.,Huang Z.H.,Zhao J.L.,Zhang G.Q.,and Liu B.,2012,Screening for germplasm with heat tolerance at flowering stage in Oryza sativa,Huanan Nongye Daxue Xuebao(Journal of South China Agricultural University),33(4):585-588(杨梯丰,张少红,王晓飞,黄章慧,赵均良,张桂权,刘斌,2012,水稻抽穗开花期耐热种质资源的筛选鉴定,华南农业大学学报,33(4):585-588)
Ye C.,Argayoso M.A.,Redon E.D.,Sierra S.N.,Laza M.A.,Dilla C.J.,Mo Y.,Thomson M.J.,Chin J.,Delavin C.B.,Diaz G.Q.,and Hernandez J.E.,2011,Mapping QTL for heat tolerance at flowering stage in rice using SNP markers,Plant Breeding,131(1):33-41
Ye C.,Tenorio F.A.,Redo觡a E.D.,Morales-Cortezano P.S.,Cabrega G.A.,Jagadish K.S.V.,and Gregorio G.B.,2015,Finemapping and validating qHTSF4.1 to increase spikelet fertility under heat stress at flowering in rice,Theor.Appl.Genet.,128(8):1507-1517
Zhang G.L.,Chen L.Y.,Xiao G.Y.,Xiao Y.H.,Chen X.B.,and Zhang S.T.,2009,Bulked segregant analysis to detect QTLrelated to heat tolerance in rice(Oryza sativa L.)using SSRmarkers,Agricultural Sciences in China,8(4):482-487
Zhang T.,Yang L.,Jiang K.F.,Huang M.,Sun Q.,Chen W.F.,and Zheng J.K.,2008,QTL mapping for heat tolerance of the tassel period of rice,Fenzi Zhiwu Yuzhong(Molecular Plant Breeding),6(5):867-873(张涛,杨莉,蒋开锋,黄敏,孙群,陈温福,郑家奎,2008,水稻抽穗扬花期耐热性的QTL分析,分子植物育种,6(5):867-873)
Zhao L.,Lei J.G.,Huang Y.J.,Zhu S.,Chen H.P.,Huang R.L.,Peng Z.Q.,Tu Q.H.,Shen X.H.,and Yan S.,2016,Mapping quantitative trait loci for heat tolerance at anthesis in rice using chromosomal segment substitution lines,Breeding Sci.,66(3):358-366
Zhao Z.G.,Jiang L.,Xiao Y.H.,Zhang W.W.,Zhai H.Q.,and Wan J.M.,2006,Identification of QTLs for heat tolerance at the booting stage in rice(Oryza sativa L.),Zuowu Xuebao(Acta Agronomica Sinica),32(5):640-644(赵志刚,江玲,肖应辉,张文伟,翟虎渠,万建民,2006,水稻孕穗期耐热性QTLs分析,作物学报,32(5):640-644)
Zhou W.H.,Xue D.W.,and Zhang G.P.,2012,Identification and physiological characterization of thermo-tolerant rice genotypes,Zhejiang Daxue Xuebao(Journal of Zhejiang University(Agriculture and Life Sciences)),38(1):1-9(周伟辉,薛大伟,张国平,2012,耐热水稻基因型的鉴定及相关生理学特征分析,浙江大学学报(农业与生命科学版),38(1):1-9)
Zhu S.,Huang R.L.,Wai H.P.,Xiong H.L.,Shen X.H.,He H.H.,and Yan S.,2017,Mapping quantitative trait loci for heat tolerance at the booting stage using chromosomal segment substitution lines in rice,Physiol.Mol.Biol.Plants,23(4):817-825