水稻低温、低氧发芽力的QTL定位
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摘要
本研究鉴定了不同水稻品种种子的低溫、低氧发芽力，并检测了种子低温萌发过程中内源激素GA和ABA含量、淀粉酶类型的变化；利用RIL群体和分子标记技术检测了种子低温、低氧发芽力QTL；同时以USSR5/N22 F_2为作图群体，构建了包含114个SSR标记的连锁图谱，并利用该图谱对种子低温、低氧发芽力进行了QTL分析。结果如下：
1．水稻品种资源低温、低氧发芽力的鉴定
本研究鉴定了521份水稻品种资源的低温、低氧发芽力，结果表明低温、低氧发芽力在籼、粳稻品种间差异极显著。籼稻的低温发芽力高于粳稻品种，而粳稻品种的低氧发芽力高于籼稻。
水稻种子低温、低氧发芽力都存在地域性差异．部分云南品种的低氧发芽力强，华南品种的低温发芽力高。
低温、低氧发芽力的相关系数在0.05水平上不显著，在0.1水平上显著，说明两者之间相关性较小。
2．水稻种子低温萌发生理机制的初步研究
低温萌发时，种子一直保持较高的ABA水平，与低溫发芽力强的品种USSR5相比，低温发芽力弱的品种N22对ABA敏感度高。外源ABA、GA处理结果也说明了这一点。
淀粉酶电泳结果说明温度没有影响淀粉酶类型，只是使其表达量减小，且表达延迟。电泳图谱中淀粉酶带3的变化与种子的萌发密切相关。
3．水稻低温、低氧发芽力的QTL定位
对Kinmaze/DV85 RIL群体进行低温、低氧发芽力QTL分析。在第2、6、7、11、12染色体上的X67、X386、R1440、G1465、X148等5个标记处分别检测到5个低温发芽力QTL，贡献率在8.8％-27.1％之间。在第3、5、7染色体上检测到两对互作位点。
在第1、2、5、7染色体上检测到5个低氧发芽力QTL，第5染色体上分布有两个低氧发芽力QTL，分别位于G260、X105标记附近。5个QTLs的贡献率在
硕士学位论文水稻低温、低氧发芽力的QTL定位
12.05%一19.56%之间。所检测到的3对互作位点分别位于第2、3、5、11染色体上。
4.SSR连锁图谱的构建及水稻低温、低氧发芽力的QTL定位
本研究利用“USSRS闪22”的F:为作图群体，构建了一个包含114个分子标记，
覆盖基因组总长度为2115.4cM的SSR遗传连锁图。标记间平均距离为18.56cM。
对逐日检测的F:群体的低温发芽率进行QTL分析，在12个染色体区域检测到低
温发芽力QTL。低温萌发早期:7一9天，位于第7、8、10、12染色体上的QTL起到
了重要作用，其贡献率在7.1%一70.2%之间。位于第3、4、5、9、n染色体上的QTL
是低温萌发过程中稳定表达的QTL。
对F:群体在30℃、0.0035889/lo鲍水的氧浓度下黑暗萌发5天的芽长进行低氧发
芽力QTL分析，共检测到4个低氧发芽力QTLs，分别位于第3、5、11染色体上。
LOD值为2.78一4.89。贡献率在6.70%一14.50%之间。
不同群体中控制种子低温、低氧发芽力的QTL大多不同，但主效QTL的遗传是稳
定的。在第H染色体上存在控制种子低温发芽力主效QTL，在第3、5染色体上存在
控制种子低氧发芽力主效QTL。对种子低温、低氧发芽力QTL定位结果进行比较分析，
两直播性状相关，但不密切，与表型鉴定结果同。
LTG (low temperature germinability) and AG (anoxia germinability) in 521 rice varieties were evaluated. The mechanism of low temperature germination was discussed about content of endogenous ABA and GA, and the type of amylase during germination at low temperature. Mapping of quantitative trait loci (QTLs) controlling LTG and AG had been carried out in a RIL population using molecular markers. A linkage map including 114 SSR markers was constructed using USSR5/N22 F2 as a mapping population, and QTLs for LTG and AG were identified. The results were as follows:
1. The evaluation of LTG and AG in rice varieties
In this study, seed LTG and AG in 521 varieties were detected. The result showed there was significant difference in LTG and AG between japonica and indica. LTG in indica cultivars was higher than that in japonica. While AG in japonica cultivars was stronger than that in indica.
Regional difference existed in LTG and AG in rice varieties. Some varieties from Yunnan had stronger AG. LTG of Varieties from Southern China was stronger than that from other regions.
The correlate coefficient between LTG and AG was significant at 0.1 level, which indicate correlation degree of two traits was low.
2. The physiological mechanism of low temperature germination
Germinating at low temperature, the level of ABA content in rice seeds kept high. Comparing with strong LTG variety, USSR5, poor LTG variety, N22 was more sensitive to ABA. Moreover, the result of exogenous ABA treatment also supports this conclusion.
The results of amylase electrophoresis indicated that the type of amylase have no variation in germinating seeds at low temperature, but its expression lowered and delayed.
3. Identification of QTL controlling LTG and AG in rice
81 Rl lines from Kinmaze/DV85 were analyzed genetically using 137markers. Five putative LTG QTLs, qLTG-2, qLTG-6, qLTG-7, qLTG-11 and qLTG-12 were detected on chromosome 2, 6, 7, 11 and 12, respectively. PVE ranged from 8.8% to 27.1%. Two pairs of epistatic QTLs were detected on chromosome 3, 5 and 7 with significant epistatic effects.
Five putative QTLs for AG linking with XR2635, R418, G260, XI05, X3 79 , whose PVE ranged from 12.05% to 19.56% , were detected on chromosome 1 , 2, 5, 7. There were two QTLs for LTG on chromosome 5. Three pairs of epistatic QTLs were detected on chromosome 2, 3, 5 and 11 with significant epistatic effects.
4. Construction of a SSR linkage map and QTL analysis of LTG and AG
A linkage map including 114 SSR markers, which covered 2115.4cM of 12 rice chromosomes, was constructed using 189USSR5/N22 p2 plants as a mapping population. The average distance between two markers was 18.56cM. Using this map, QTLs controlling LTG and AG were identified.
LTG was evaluated day by day were analyze genetically. 12QTLs controlling LTG were detected. QTLs for LTG on chromosome 7, 8, 10, 12 played an important role at early germinating stage. PVE ranged from 7.1% to 70.2%. QTLs on chromosome 3, 4, 5, 9, 11 were expressed stably during the whole germination stage.
AG was evaluated at 30 C and 0.003588g/100g H2O for 5days. Four putative QTLs for AG on chromosome 3, 5 , 11, respectively, were identified, and the LOD value ranged from 2.78 to 4.89, and PVE ranged from 6.70% to 14.50%.
QTLs for LTG and AG detected in different population always changed, but main QTL was stable. Main QTL for LTG existed on chromosome 11. Main QTL for AG was found on chromosome 3, 5. The result of phenotype evaluation and QTL mapping showed LTG correlated with AG at some extent.
1．水稻品种资源低温、低氧发芽力的鉴定
本研究鉴定了521份水稻品种资源的低温、低氧发芽力，结果表明低温、低氧发芽力在籼、粳稻品种间差异极显著。籼稻的低温发芽力高于粳稻品种，而粳稻品种的低氧发芽力高于籼稻。
水稻种子低温、低氧发芽力都存在地域性差异．部分云南品种的低氧发芽力强，华南品种的低温发芽力高。
低温、低氧发芽力的相关系数在0.05水平上不显著，在0.1水平上显著，说明两者之间相关性较小。
2．水稻种子低温萌发生理机制的初步研究
低温萌发时，种子一直保持较高的ABA水平，与低溫发芽力强的品种USSR5相比，低温发芽力弱的品种N22对ABA敏感度高。外源ABA、GA处理结果也说明了这一点。
淀粉酶电泳结果说明温度没有影响淀粉酶类型，只是使其表达量减小，且表达延迟。电泳图谱中淀粉酶带3的变化与种子的萌发密切相关。
3．水稻低温、低氧发芽力的QTL定位
对Kinmaze/DV85 RIL群体进行低温、低氧发芽力QTL分析。在第2、6、7、11、12染色体上的X67、X386、R1440、G1465、X148等5个标记处分别检测到5个低温发芽力QTL，贡献率在8.8％-27.1％之间。在第3、5、7染色体上检测到两对互作位点。
在第1、2、5、7染色体上检测到5个低氧发芽力QTL，第5染色体上分布有两个低氧发芽力QTL，分别位于G260、X105标记附近。5个QTLs的贡献率在
硕士学位论文水稻低温、低氧发芽力的QTL定位
12.05%一19.56%之间。所检测到的3对互作位点分别位于第2、3、5、11染色体上。
4.SSR连锁图谱的构建及水稻低温、低氧发芽力的QTL定位
本研究利用“USSRS闪22”的F:为作图群体，构建了一个包含114个分子标记，
覆盖基因组总长度为2115.4cM的SSR遗传连锁图。标记间平均距离为18.56cM。
对逐日检测的F:群体的低温发芽率进行QTL分析，在12个染色体区域检测到低
温发芽力QTL。低温萌发早期:7一9天，位于第7、8、10、12染色体上的QTL起到
了重要作用，其贡献率在7.1%一70.2%之间。位于第3、4、5、9、n染色体上的QTL
是低温萌发过程中稳定表达的QTL。
对F:群体在30℃、0.0035889/lo鲍水的氧浓度下黑暗萌发5天的芽长进行低氧发
芽力QTL分析，共检测到4个低氧发芽力QTLs，分别位于第3、5、11染色体上。
LOD值为2.78一4.89。贡献率在6.70%一14.50%之间。
不同群体中控制种子低温、低氧发芽力的QTL大多不同，但主效QTL的遗传是稳
定的。在第H染色体上存在控制种子低温发芽力主效QTL，在第3、5染色体上存在
控制种子低氧发芽力主效QTL。对种子低温、低氧发芽力QTL定位结果进行比较分析，
两直播性状相关，但不密切，与表型鉴定结果同。
LTG (low temperature germinability) and AG (anoxia germinability) in 521 rice varieties were evaluated. The mechanism of low temperature germination was discussed about content of endogenous ABA and GA, and the type of amylase during germination at low temperature. Mapping of quantitative trait loci (QTLs) controlling LTG and AG had been carried out in a RIL population using molecular markers. A linkage map including 114 SSR markers was constructed using USSR5/N22 F2 as a mapping population, and QTLs for LTG and AG were identified. The results were as follows:
1. The evaluation of LTG and AG in rice varieties
In this study, seed LTG and AG in 521 varieties were detected. The result showed there was significant difference in LTG and AG between japonica and indica. LTG in indica cultivars was higher than that in japonica. While AG in japonica cultivars was stronger than that in indica.
Regional difference existed in LTG and AG in rice varieties. Some varieties from Yunnan had stronger AG. LTG of Varieties from Southern China was stronger than that from other regions.
The correlate coefficient between LTG and AG was significant at 0.1 level, which indicate correlation degree of two traits was low.
2. The physiological mechanism of low temperature germination
Germinating at low temperature, the level of ABA content in rice seeds kept high. Comparing with strong LTG variety, USSR5, poor LTG variety, N22 was more sensitive to ABA. Moreover, the result of exogenous ABA treatment also supports this conclusion.
The results of amylase electrophoresis indicated that the type of amylase have no variation in germinating seeds at low temperature, but its expression lowered and delayed.
3. Identification of QTL controlling LTG and AG in rice
81 Rl lines from Kinmaze/DV85 were analyzed genetically using 137markers. Five putative LTG QTLs, qLTG-2, qLTG-6, qLTG-7, qLTG-11 and qLTG-12 were detected on chromosome 2, 6, 7, 11 and 12, respectively. PVE ranged from 8.8% to 27.1%. Two pairs of epistatic QTLs were detected on chromosome 3, 5 and 7 with significant epistatic effects.
Five putative QTLs for AG linking with XR2635, R418, G260, XI05, X3 79 , whose PVE ranged from 12.05% to 19.56% , were detected on chromosome 1 , 2, 5, 7. There were two QTLs for LTG on chromosome 5. Three pairs of epistatic QTLs were detected on chromosome 2, 3, 5 and 11 with significant epistatic effects.
4. Construction of a SSR linkage map and QTL analysis of LTG and AG
A linkage map including 114 SSR markers, which covered 2115.4cM of 12 rice chromosomes, was constructed using 189USSR5/N22 p2 plants as a mapping population. The average distance between two markers was 18.56cM. Using this map, QTLs controlling LTG and AG were identified.
LTG was evaluated day by day were analyze genetically. 12QTLs controlling LTG were detected. QTLs for LTG on chromosome 7, 8, 10, 12 played an important role at early germinating stage. PVE ranged from 7.1% to 70.2%. QTLs on chromosome 3, 4, 5, 9, 11 were expressed stably during the whole germination stage.
AG was evaluated at 30 C and 0.003588g/100g H2O for 5days. Four putative QTLs for AG on chromosome 3, 5 , 11, respectively, were identified, and the LOD value ranged from 2.78 to 4.89, and PVE ranged from 6.70% to 14.50%.
QTLs for LTG and AG detected in different population always changed, but main QTL was stable. Main QTL for LTG existed on chromosome 11. Main QTL for AG was found on chromosome 3, 5. The result of phenotype evaluation and QTL mapping showed LTG correlated with AG at some extent.
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63．严长杰，李欣，程祝宽，于恒秀，顾铭洪，朱立煌。利用分子标记定位水稻芽期耐冷性基因．中国水稻科学，1999，13(3)：134-138
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61. Cui K.H., et al. Molecular dissection of seedling-vigor and associated physiological traits in rice. Theor Appl Genet, 2002, 105: 745-753.
62. Wan J., Nakazaki T., Kawaura K., Ikehashi H.. Identification of marker loci for seed dormancy in rice (Oryza sativa L.),Crop Sci. 1997,37:1759-1763.
63．严长杰，李欣，程祝宽，于恒秀，顾铭洪，朱立煌。利用分子标记定位水稻芽期耐冷性基因．中国水稻科学，1999，13(3)：134-138
64．詹庆才，曾曙珍，熊伏星，齐腾浩二，加藤明。水稻苗期耐冷性QTLs的分子定位。湖南农业大学学报(自然科学版)，2003，29(1)：7-11
65．钱前等．水稻籼粳交DH群体苗期的耐冷性QTLS分析．科学通报，1999，44(22)：2402-2407
66. Ye C.R., Kato A., Saito K., Dai L.Y., Yang Q.Z.. QTL analysis of cold tolerance at the booting stage in Yunnan rice variety Chongtui. Chinese J Rice Sci, 2001, 15(3):13-16.
67. Andaya V.C., Mackill D.J.. QTLs conferring cold tolerance at the booting stage of rice using recombinant inbred lines from a japonica×indica cross. Theor Appl Genet, 2003, 106: 1084-1090.
68. Kearsey M.J., Farquhar A.G.L.. QTL analysis in plants: where are we now. Heredity, 1998, 80: 137-142.
69．吴为人，唐定中，李维明。数量性状的遗传剖析和分子剖析。作物学报，2000，26(4)：501-507。
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71. Wright S. The genetic structure of population. Ann. Eugen, 1951, 15: 323-354.
72. Allard RW. Formulas and tables to facilitate the calculation of recombination values in heredity. Hilgrardia, 1956, 24: 235-278
73. Tanksley SD, Hewitt JD. Use of molecular markers in breeding for soluble solids in tomato: a re-examination. Theor Appl Gener, 1988, 75: 811-823.
74. Doebley J., et al.. Teosinte branchedl and the origin of maize: evidence for epistasis and the evolution of dominance. Genetics, 1995, 141: 333-346.
75. Lark KG, et al.. Interactions between quantitative trait loci in soybean in which trait variation at one locus is conditioned upon a specific allele at another. Proc Natl Acad Sci USA. 1995, 92: 4656-4660.
76. Li Z.K., Pinson S.R.M., Park W.D., Paterson A.H., Stansel J.W.. Epistasis for three Grain Yield Components in rice (Oryza sativa I.)[J].Genetics, 1997a, 145: 453—465.
77. Li ZK, et al.. Genetics of hybrid sterility and hybrid breakdown in an inter-subspecific rice(Oryza sativa L.) population. Genetics, 1997b, 145: 1139-1148.
78. Yu S.B, et al.. Importance of epitasis as the genetic basis of heterosis in an elite rice hybrid. Proc. Natl. Acad. Sci. USA., 1997, 94: 9226-9231.
79. Li ZK. Molecular analysis of epistasis affecting complex traits. In: Paterson AH, editor. Molecular analysis of complex traits. Boca Rato, Fla.(USA): CRC Press, 2001: 119-130.
80. Wang D L, Zhu J, Li Z K, et al. Mapping QTLs with epistatic effects and QTL x environment interactions by mixed linear model approaches.[J]. Yheor Appl Genet, 1999, 99: 1255—1264.
81. Kao CH, et al.. Multiple interval mapping for quantitative trait loci. Genetics, 1999,152:1203-1216.
82. Yan J. Q., Zhu J., He C.X., Benmoussa M., Wu P., Molecular dissection of development behavior of plant height in rice(Oryza sativa L.). Genetics, 1998, 150: 1257-1265.
83．廖春燕，吴平，易可可，胡彬，倪俊健．不同遗传背景及环境中水稻(Oryza sativa L.)穗长的QTLs和上位性分析。遗传学报，2000，27(7)：599-607．
84. Lin H.X., Yamamoto T., Sasaki T., Yano M.. Characterization and detection of epistatic interactions of 3 QTLs, Hd1, Hd2, and Hd3, controlling heading date in rice using nearly isogenic lines. Theor. Appl. Gene., 2000, 101:1021-1028.
85. Kato-Noguchi H. Ethanol sensitivity of rice and oat coleoptiles. Physiol Plant, 2002 May, 115(1): 119-124
86. Mitsunori S. et al. Complex regulation of ABA biosynthesis in plants. Trends Plant Sci, 2002, 7(1): 41-48.
87．何忠效，张树政。电泳。科学出版社，1999，第8章．
88．王树文，植物生理与分子生物学，科学出版社，1992，318-320
89. Kaneko M., Itoh H., Tanaka M.U., Ashikari M., Matsuoka M.. The α-amylase induction in endosperm during rice seed germination is caused by gibberellin synthesized in epithelium. Plant Physiology, 2002, 128: 1264-1270
90. Miura K., Araki H.. Effect of temperature during the ripening period on the induction of secondary dormancy in rice seeds(Oryza sativa L.). Breeding Sci., 1999,49:7-10
91. Zeng Z.-B. Precision mapping of quantitative trait loci. Genetics, 1994, 136: 1457-1468
92. McCouch S R, Cho Y G, Yano M,et al. Report on QTL nomenclature. Rice Genet.Newslett, 1997,14: 11—13
93. Lin S.Y., Sasaki T., Yano M.. Mapping quantitative trait loci controlling seed dormancy and heading date in rice, Oryza Sativa L., using backcross inbred lines. Theor Appl Genet, 1998, 96: 997-1003
94. Chrisopher D.R. Pathways to abscisic acid-regulated gene expression. New phytol, 2000, 148: 357-396.
95. Ruth R. F., Srinivas S. L. Gampata et al.. Abscisic acid signaling in seeds and seedlings. The plant cell, 2002, supplement: 15-45.
96. Dellaporta S.L, Wood T, Hicks TB. A plant DNA mini preparation: version Ⅱ. Plant Mot Bioi Rep, 1983, 1: 19-21
97. Lander E., Green P., Abrahamson J., Barlow A., Daley M., Lincon S., Newburg L.. MAPMAKER: an interactive computer package for constructing primary genetic maps of experimental and natural populations. Genomics, 1987, 1:174-181
98. Lincoln S., Daley M., Lander E.. Constructing genetic maps with MAPMAKER/EXR3.0. Whitehead institute technical report, 3rd edn.. Whitehead institute, Cambridge, Mass. 1992a
99. Lincoln S., Daley M., Lander E.. Mapping genes controlling quantitative traits with MAPMAKER/OOTL 1.1. Whitehead institute technical report, 2nd edn., Whitehead institute, Cambridge, Mass. 1992b
100. Allard RW. Future direction in plant population genetics, evolution and breeding. In: International Symposium for Population Genetics and Germplasm Resources in Crop Improvement. Suppl, 1988: 9-18.
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