Advances in Molecular and Physiological Mechanisms of Cold Tolerance Regulation of Rice at the Booting Stage

doi:10.16819/j.1001-7216.2025.231208

Abstract

Abstract:

Cold tolerance at the booting stage is crucial for the reproductive function of rice (Oryza sativa L.), as it affects anther development and can lead to a significant decrease in seed setting rate and overall yield. The levels of certain growth regulatory substances, such as sugars, reactive oxygen species (ROS), and plant endogenous hormones like abscisic acid (ABA) and gibberellin (GA), are closely related to cold tolerance during the booting stage of rice. In this review, we systematically summarize the effects of cold stress on anther development in rice at the booting stage, the changes in growth regulatory substances, and the functional analysis of quantitative trait loci (QTL) and cold tolerance regulatory genes. This review aims to provide theoretical guidance for the breeding and improvement of cold-tolerant rice varieties and the development of materials at the booting stage.

Key words: rice, cold tolerance, booting stage, quantitative trait locus, cold tolerance gene

摘要：

水稻孕穗期低温胁迫会破坏生殖器官功能，使花药发育受损、花粉育性降低，最终导致水稻结实率下降、产量降低。部分生长调节物质，如可溶性糖、活性氧、植物内源激素（脱落酸和赤霉素）的含量都与水稻孕穗期耐冷性密切相关。本文系统总结了水稻孕穗期低温胁迫对水稻花药发育的影响，生长调节物质的变化，以及水稻孕穗期耐冷QTL和克隆的孕穗期耐冷基因的功能解析，以期为水稻孕穗期耐冷品种育种改良和材料创制提供理论指导。

关键词: 水稻, 耐冷性, 孕穗期, 数量性状基因座, 耐冷基因

SUI Jingjing, ZHAO Guilong, JIN Xin, BU Qingyun, TANG Jiaqi. Advances in Molecular and Physiological Mechanisms of Cold Tolerance Regulation of Rice at the Booting Stage[J]. Chinese Journal OF Rice Science, 2025, 39(1): 1-10.

随晶晶, 赵桂龙, 金欣, 卜庆云, 唐佳琦. 水稻孕穗期耐冷调控的分子及生理机制研究进展[J]. 中国水稻科学, 2025, 39(1): 1-10.

Figures/Tables 3

References 73

[1]	Roy P, Orikasa T, Okadome H, Nakamura N, Shiina T. Processing conditions, rice properties, health and environment[J]. International Journal of Environmental Research and Public Health, 2011, 8(6): 1957-1976.
[2]	Khush G S. Origin, dispersal, cultivation and variation of rice[J]. Plant Molecular Biology, 1997, 35: 25-34.
[3]	罗玉坤, 朱智伟, 陈能, 段彬伍, 章林平. 中国主要稻米的粒型及其品质特性[J]. 中国水稻科学, 2004, 18(2): 49-53.
	Luo Y K, Zhu Z W, Chen N, Duan B W, Zhang L P. Grain types and related characteristics of rice in China[J]. Chinese Journal of Rice Science, 2004, 18(2): 49-53. (in Chinese with English abstract)
[4]	Shi S J, Wang E T, Li C X, Zhou H, Cai M L, Cao C G, Jiang Y. Comprehensive evaluation of 17 qualities of 84 types of rice based on principal component analysis[J]. Foods, 2021, 10(11): 2883.
[5]	胡培松, 翟虎渠, 万建民. 中国水稻生产新特点与稻米品质改良[J]. 中国农业科技导报, 2002(4): 33-39.
	Hu P S, Zhai H Q, Wan J M. New characteristics of rice production and quality improvement in China[J]. Journal of Agricultural Science and Technology, 2002(4): 33-39. (in Chinese with English abstract)
[6]	国家统计局. 中国统计年鉴[M]. 北京: 中国统计出版社, 2020.
	National Bureau of Statistics. China Statistical Yearbook[M]. Beijing: China Statistics Press, 2020. (in Chinese)
[7]	Gough N R. Rice that tolerates a chill[J]. Science Signaling, 2015, 8(370): ec76-ec76.
[8]	杨荣教, 郭梦逸, 王白昌, 余选礼, 王白, 徐津. 不同海拔条件下籼、粳稻生长发育差异分析[J/OL]. 分子植物育种, https://link.cnki.net/urlid/46.1068.S.20230831.2009.014
	Yang R J, Guo M Y, Wang B C, Yu X L, Wang B, Xu J. Analysis of growth and development differences between indica and japonica rice under different altitude conditions[J/OL]. Molecular Plant Breeding, https://link.cnki.net/urlid/46.1068.S.20230831.2009.014. (in Chinese with English abstract)
[9]	杨荣教, 雷娟, 王白昌, 余选礼, 王白, 徐津. 不同海拔条件下籼粳稻产量和品质性状的差异分析[J/OL]. 分子植物育种, https://link.cnki.net/urlid/46.1068.S.20230811.1332.002.
	Yang R J, Lei J, Wang B C, Yu X L, Wang B, Xu J. Difference analysis of yield and quality traits of indica and japonica rice under different altitude conditions[J/OL]. Molecular Plant Breeding, https://link.cnki.net/urlid/46.1068.S.20230811.1332.002. (in Chinese with English abstract)
[10]	李文枫, 毕洪文, 黄峰华, 李晓晨, 李金霞, 张妍, 刘艳霞. 黑龙江省水稻产业发展现状及展望[J]. 农业展望, 2020, 16(12): 48-53.
	Li W F, Bi H W, Huang F H, Li X C, Li J X, Zhang Y, Liu Y X. Current status and prospects of rice industry development in Heilongjiang Province[J]. Agricultural Outlook, 2020, 16 (12): 48-53. (in Chinese)
[11]	国家统计局. 中国统计年鉴[M]. 北京: 中国统计出版社, 2021.
	National Bureau of Statistics. China Statistical Yearbook[M]. Beijing: China Statistics Press, 2021. (in Chinese)
[12]	Liu Z X, Deng H B. Development of genetic and QTLs analysis for cold tolerance in rice[J]. Chinese Agricultural Science Bulletin, 2009, 25: 45-50.
[13]	马建勇, 许吟隆, 潘婕. 东北地区农业气象灾害的趋势变化及其对粮食产量的影响[J]. 中国农业气象, 2012, 33(2): 283-288.
	Ma J Y, Xu Y L, Pan J. Analysis of agro-meteorological disasters tendency variation and the impacts on grain yield over Northeast China[J]. Chinese Journal of Agrometeorology, 2012, 33(2): 283-288. (in Chinese with English abstract)
[14]	张养才, 何维勋, 李世奎. 中国农业气象灾害概论[M]. 北京: 气象出版社, 1991: 25-30.
	Zhang Y C, He W X, Li S K. Introduction to agricultural meteorological disasters in China[M]. Beijing: Meteorological Press, 1991: 25-30. (in Chinese)
[15]	王书裕. 作物低温冷害研究[M]. 北京: 气象出版社, 1995, 116-120.
	Wang S Y. Research on low-temperature chilling damage to crops[M]. Beijing: Meteorological Press, 1995: 116-120. (in Chinese)
[16]	王绍武, 马树庆, 陈莉, 黄建斌. 低温冷害[M]. 北京: 气象出版社, 2009: 23-40.
	Wang S W, Ma S Q, Chen L, Huang J B. Cold damage[M]. Beijing: Meteorological Press, 2009: 23-40. (in Chinese)
[17]	张莉萍, 黄少锋, 王丽萍, 刘颖, 沈巧梅. 2002年黑龙江省东部水稻冷害解析[J]. 黑龙江农业科学, 2004(1): 39-42.
	Zhang L P, Huang S F, Wang L P, Liu Y, Shen Q M. Analysis of chilling damage to rice in eastern Heilongjiang Province in 2002[J]. Heilongjiang Agricultural Sciences, 2004(1): 39-42. (in Chinese with English abstract)
[18]	Mamun E A, Cantrill L C, Overall R L, Sutton B G. Mechanism of low-temperature-induced pollen failure in rice[J]. Cell Biology International, 2010, 34(5): 469-476.
[19]	Xu L M, Zhou L, Zeng Y W, Wang F M, Zhang H L, Shen S Q, Li Z C. Identification and mapping of quantitative trait loci for cold tolerance at the booting stage in a japonica rice near-isogenic line[J]. Plant Science, 2008, 174(3): 340-347.
[20]	Li S C, Li W B, Huang B, Cao X M, Zhou X Y, Ye S M, Li C B, Gao F Y, Zou T, Xie K L, Ren Y, Ai P, Tang Y F, Li X M, Deng Q M, Wang S Q, Zheng A P, Zhu J, Liu H N, Wang L X, Li P. Natural variation in PTB1 regulates rice seed setting rate by controlling pollen tube growth[J]. Nature Communications, 2013, 4: 2793.
[21]	Nishiyama I. Male sterility caused by cooling treatment at the meiotic stage in rice plants: IV. Respiratory activity of anthers following cooling treatments at the meiotic stage[J]. Japanese Journal of Crop Science, 1970, 39: 65-70.
[22]	Satake T. Male sterility caused by cooling treatment at the young microspore stage in rice plants[J]. Japanese Journal of Crop Science, 1991, 60: 523-528.
[23]	Yamamori K, Ogasawara K, Ishiguro S, Koide Y, Takamure I, Fujino K, Sato Y, Kishima Y. Revision of the relationship between anther morphology and pollen sterility by cold stress at the booting stage in rice[J]. Annals of Botany, 2021, 128(5): 559-575.
[24]	Saito K, Miura K, Nagano K, Hayano-Saito Y, Araki H, Kato A. Identification of two closely linked quantitative trait loci for cold tolerance on chromosome 4 of rice and their association with anther length[J]. Theoretical and Applied Genetics, 2001, 103: 862-868.
[25]	Shi Y, Guo E, Cheng X, Wang L, Jiang S, Yang X, Ma H, Zhang T, Li T, Yang X. Effects of chilling at different growth stages on rice photosynthesis, plant growth, and yield[J]. Environmental and Experimental Botany, 2022, 203: 105045.
[26]	Gothandam K M, Kim E S, Chung Y Y. Ultrastructural study of rice tapetum under low-temperature stress[J]. Journal of Plant Biology, 2007, 50: 396-402.
[27]	Liang Z M, Luo J, Wei B, Liao Y C, Liu Y. Trehalose can alleviate decreases in grain number per spike caused by low-temperature stress at the booting stage by promoting floret fertility in wheat[J]. Journal of Agronomy and Crop Science, 2021, 207(4): 717-732.
[28]	Wu L B, Eom J S, Isoda R, Li C H, Char S N, Luo D P, Schepler-Luu V, Nakamura M, Yang B, Frommer W B. OsSWEET11b, a potential sixth leaf blight susceptibility gene involved in sugar transport-dependent male fertility[J]. New Phytologist, 2022, 234(3): 975-989.
[29]	Oliver S N, van Dongen J T, Alfred S C, Mamun E A, Zhao X C, Saini H S, Fernandes S F, Blanchard C L, Sutton B G, Geigenberger P, Dennis E S, Dolferus R. Cold-induced repression of the rice anther-specific cell wall invertase gene OSINV4 is correlated with sucrose accumulation and pollen sterility[J]. Plant Cell and Environment, 2005, 28(12): 1534-1551.
[30]	Mamun E A, Alfred S, Cantrill L C, Overall R L, Sutton B G. Effects of chilling on male gametophyte development in rice[J]. Cell Biology International, 2006, 30(7): 583-591.
[31]	Oliver S N, Dennis E S, Dolferus R. ABA regulates apoplastic sugar transport and is a potential signal for cold-induced pollen sterility in rice[J]. Plant and Cell Physiology, 2007, 48(9): 1319-1330.
[32]	Sakata T, Oda S, Tsunaga Y, Shomura H, Kawagishi- Kobayashi M, Aya K, Saeki K, Endo T, Nagano K, Kojima M, Sakakibara H, Watanabe M, Matsuoka M, Higashitani A. Reduction of gibberellin by low temperature disrupts pollen development in rice[J]. Plant Physiology, 2014, 164(4): 2011-2019.
[33]	Liu C T, Schläppi M R, Mao B G, Wang W, Wang A J, Chu C C. The bZIP73 transcription factor controls rice cold tolerance at the reproductive stage[J]. Plant Biotechnology Journal, 2019, 17(9): 1834-1849.
[34]	Mittal D, Madhyastha D A, Grover A. Genome-wide transcriptional profiles during temperature and oxidative stress reveal coordinated expression patterns and overlapping regulons[J]. PLoS One, 2012, 7: e40899.
[35]	Chakraborty A, Bhattacharjee S. Differential competence of redox-regulatory mechanism under extremes of temperature determines growth performances and cross tolerance in two indica rice cultivars[J]. Journal of Plant Physiology, 2015, 176: 65-77.
[36]	El-Esawi M A, Alayafi A A. Overexpression of rice Rab7 gene improves drought and heat tolerance and increases grain yield in rice (Oryza sativa L.)[J]. Genes (Basel), 2019, 10(1): 56.
[37]	刘次桃, 王威, 毛毕刚, 储成才. 水稻耐低温逆境研究:分子生理机制及育种展望[J]. 遗传, 2018, 40(3): 171-185.
	Liu C T, Wang W, Mao B G, Chu C C. Cold stress tolerance in rice: Physiological changes, molecular mechanism, and future prospects[J]. Hereditas(Beijing) 2018, 40(3): 171-185. (in Chinese with English abstract)
[38]	Sato Y, Masuta Y, Saito K, Murayama S, Ozawa K. Enhanced chilling tolerance at the booting stage in rice by transgenic overexpression of the ascorbate peroxidase gene, OsAPXa[J]. Plant Cell Reports, 2011, 30(3): 399-406.
[39]	Suzuki K, Aoki N, Matsumura H, Okamura M, Ohsugi R, Shimono H. Cooling water before panicle initiation increases chilling-induced male sterility and disables chilling-induced expression of genes encoding OsFKBP65 and heat shock proteins in rice spikelets[J]. Plant Cell and Environment, 2015, 38(7): 1255-1274.
[40]	Tovuu A, Zulfugarov I S, Wu G X, Kang I S, Kim C, Moon B Y, An G, Lee C H. Rice mutants deficient in ω-3 fatty acid desaturase (FAD8) fail to acclimate to cold temperatures[J]. Plant Physiology and Biochemistry, 2016, 109: 525-535.
[41]	Luo D P, Xu H, Liu Z L, Guo J X, Li H Y, Chen L T, Fang C, Zhang Q Y, Bai M, Yao N, Wu H, Wu H, Ji C H, Zheng H Q, Chen Y L, Ye S, Li X Y, Zhao X C, Li R Q, Liu Y G. A detrimental mitochondrial-nuclear interaction causes cytoplasmic male sterility in rice[J]. Nature Genetics, 2013, 45(5): 573-577.
[42]	Ko S S, Li M J, Ho Y C, Yu C P, Yang T T, Lin Y J, Hsing H C, Chen T K, Jhong C M, Li W H, Sun-Ben Ku M. Rice transcription factor GAMYB modulates bHLH142 and is homeostatically regulated by TDR during anther tapetal and pollen development[J]. Journal of Experimental Botany, 2021, 72(13): 4888-4903.
[43]	Choudhury F K, Rivero R M, Blumwald E, Mittler R. Reactive oxygen species, abiotic stress and stress combination[J]. The Plant Journal, 2017, 90(5): 856-867.
[44]	Dreyer A, Dietz K J. Reactive oxygen species and the redox-regulatory network in cold stress acclimation[J]. Antioxidants (Basel), 2018, 7(11): 169.
[45]	Mittler R. ROS are good[J]. Trends in Plant Science, 2017, 22(1): 11-19.
[46]	Aya K, Ueguchi-Tanaka M, Kondo M, Hamada K, Yano K, Nishimura M, Matsuoka M. Gibberellin modulates anther development in rice via the transcriptional regulation of GAMYB[J]. The Plant Cell, 2009, 21(5): 1453-1472.
[47]	Khan D R, Mackill D J, Vergara B S. Selection for tolerance to low temperature-induced spikelet sterility at anthesis in rice[J]. Crop Science, 1986, 26(4): 694-698.
[48]	Yang T F, Zhang S H, Zhao J L, Huang Z H, Zhang G Q, Liu B. Meta-analysis of QTLs underlying cold tolerance in rice (Oryza sativa L.)[J]. Molecular Plant Breeding, 2015, 13: 1-15.
[49]	Li J H, Zhang Z Y, Chong K, Xu Y Y. Chilling tolerance in rice: Past and present[J]. Journal of Plant Physiology, 2022, 268: 153576.
[50]	Kuroki M, Saito K, Matsuba S, Yokogami N, Shimizu H, Ando I, Sato Y. A quantitative trait locus for cold tolerance at the booting stage on rice chromosome 8[J]. Theoretical and Applied Genetics, 2007, 115: 593-600.
[51]	Shirasawa S, Endo T, Nakagomi K, Yamaguchi M, Nishio T. Delimitation of a QTL region controlling cold tolerance at booting stage of a cultivar, ‘Lijiangxintuanheigu’, in rice, Oryza sativa L[J]. Theoretical and Applied Genetics, 2012, 124: 937-946.
[52]	Zhu Y J, Chen K, Mi X F, Chen T X, Ali J, Ye G Y, Xu J L, Li Z K. Identification and fine mapping of a stably expressed QTL for cold tolerance at the booting stage using an interconnected breeding population in rice[J]. PLoS One, 2015, 10(12): e0145704.
[53]	Li J L, Pan Y H, Guo H F, Zhou L, Yang S M, Zhang Z Y, Yang J Z, Zhang H L, Li J J, Zeng Y W, Li Z C. Fine mapping of QTL qCTB10-2 that confers cold tolerance at the booting stage in rice[J]. Theoretical and Applied Genetics, 2018, 131: 157-166.
[54]	Zhou L, Zeng Y W, Zheng W W, Tang B, Yang S M, Zhang H L, Li J J, Li Z C. Fine mapping a QTL qCTB7 for cold tolerance at the booting stage on rice chromosome 7 using a near-isogenic line[J]. Theoretical and Applied Genetics, 2010, 121: 895-905.
[55]	Takeuchi Y, Hayasaka H, Chiba B, Tanaka Isao, Shimano T, Yamagishi M, Nagano K, Sasaki T, Yano M. Mapping quantitative trait loci controlling cool-temperature tolerance at booting stage in temperate japonica rice[J]. Breeding Science, 2001, 51(3): 191-197.
[56]	Dai L Y, Lin X H, Ye C R, Ise K, Saito K J, Kato A, Xu F R, Yu T Q, Zhang D P. Identification of quantitative trait loci controlling cold tolerance at the reproductive stage in Yunnan landrace of rice, Kunmingxiaobaigu[J]. Breeding Science, 2004, 54(3): 253-258.
[57]	Yang L M, Lei L, Wang J G, Zheng H L, Xin W, Liu H L, Zou D T. qCTB7 positively regulates cold tolerance at booting stage in rice[J]. Theoretical and Applied Genetics, 2023, 136(6): 135.
[58]	Saito K, Hayano-Saito Y, Kuroki M, Sato Y. Map-based cloning of the rice cold tolerance gene Ctb1[J]. Plant Science, 2010, 179(1-2): 97-102.
[59]	Zhang Z Y, Li J J, Pan Y H, Li J L, Zhou L, Shi H L, Zeng Y W, Guo H F, Yang S M, Zheng W W, Yu J P, Sun X M, Li G L, Ding Y L, Ma L, Shen S Q, Dai L Y, Zhang H L, Yang S H, Guo Y, Li Z C. Natural variation in CTB4a enhances rice adaptation to cold habitats[J]. Nature Communications, 2017, 8: 14788.
[60]	Li J L, Zeng Y M, Pan Y H, Zhang Z Y, Guo H F, Luo Q J, Shui G H, Zhang H L, Yang S H, Guo Y, Ge S, Li J J, Li Z C. Stepwise selection of natural variations at CTB2 and CTB4a improves cold adaptation during domestication of japonica rice[J]. New Phytologist, 2021, 231(3): 1056-1072.
[61]	Lou Q J, Guo H F, Li J, Han S C, Khan N U, Gu Y S, Zhao W T, Zhang Z Y, Zhang H L, Li Z C, Li J J. Cold-adaptive evolution at the reproductive stage in Geng/japonica subspecies reveals the role of OsMAPK3 and OsLEA9[J]. Plant Journal, 2022, 111(4): 1032-1051.
[62]	Liu C T, Ou S J, Mao B G, Tang J Y, Wang W, Wang H R, Cao S Y, Schläppi M R, Zhao B R, Xiao G Y, Wang X P, Chu C C. Early selection of bZIP73 facilitated adaptation of japonica rice to cold climates[J]. Nature Communications, 2018, 9(1): 3302.
[63]	Xu Y F, Wang R C, Wang Y M, Zhang L, Yao S G. A point mutation in LTT1 enhances cold tolerance at the booting stage in rice[J]. Plant Cell and Environment, 2020, 43(4): 992-1007.
[64]	Tang J Q, Tian X J, Mei E Y, He M L, Gao J W, Yu J, Xu M, Liu J L, Song L, Li X F, Wang Z Y, Guan Q J, Zhao Z G, Wang C M, Bu Q Y. WRKY53 negatively regulates rice cold tolerance at the booting stage by fine-tuning anther gibberellin levels[J]. The Plant Cell, 2022, 34(11): 4495-4515.
[65]	Mei E Y, Tang J Q, He M L, Liu Z Q, Tian X J, Bu Q Y. OsMKKK70 negatively regulates cold tolerance at booting stage in rice[J]. International Journal of Molecular Sciences, 2022, 23(22): 14472.
[66]	Liu D F, Luo S T, Li Z C, Liang G H, Guo Y L, Xu Y Y, Chong K. COG3 confers the chilling tolerance to mediate OsFtsH2-D1 module in rice[J]. New Phytologist, 2024, 241: 2143-2157.
[67]	Saito K, Hayano-Saito Y, Maruyama-Funatsuki W, Sato Y, Kato A. Physical mapping and putative candidate gene identification of a quantitative trait locus Ctb1 for cold tolerance at the booting stage of rice[J]. Theoretical and Applied Genetics, 2004, 109: 515-522.
[68]	Xu L M, Zhou L, Zeng Y W, Wang F M, Zhang H L, Shen S Q, Li Z C. Identification and mapping of quantitative trait loci for cold tolerance at the booting stage in a japonica rice near-isogenic line[J]. Plant Science, 2008, 174: 340-347.
[69]	Zhou L, Zeng Y W, Hu G G, Pan Y H, Yang S M, You A Q, Zhang H L, Li J J, Li Z C. Characterization and identification of cold tolerant near-isogenic lines in rice[J]. Breed Science, 2012, 62(2): 196-201.
[70]	Zhang Z Y, Li J H, Li F, Liu H H, Yang W S, Chong K, Xu Y Y. OsMAPK3 Phosphorylates OsbHLH002/OsICE1 and inhibits its ubiquitination to activate OsTPP1 and enhances rice chilling tolerance[J]. Developmental Cell, 2017, 43(6): 731-743.e5.
[71]	Hu L F, Ye M, Li R, Zhang T F, Zhou G X, Wang Q, Lu J, Lou Y G. The rice transcription factor WRKY53 suppresses herbivore-induced defenses by acting as a negative feedback modulator of mitogen-activated protein kinase activity[J]. Plant Physiology, 2015, 169(4): 2907-2921.
[72]	Gao Y, Xue C Y, Liu J M, He Y, Mei Q, Wei S H, Xuan Y H. Sheath blight resistance in rice is negatively regulated by WRKY53 via SWEET2a activation[J]. Biochemical and Biophysical Research Communications, 2021, 585: 117-123.
[73]	Xie W Y, Ke Y G, Cao J B, Wang S P, Yuan M. Knock out of transcription factor WRKY53 thickens sclerenchyma cell walls, confers bacterial blight resistance[J]. Plant Physiology, 2021, 187(3): 1746-1761.

QTL名称 QTL name	染色体 Chromosome	遗传功能验证 Genetic function verification	物理位置 Physical position (bp)	候选基因 Candidate gene	参考文献Reference
qCTB8	8	无 No	2900000―3100000	LOC_Os08g44340	[50]
qLTB3	3	无 No	33540013―34653353	LOC_Os03g57680 LOC_Os03g59200	[51]
qCT-3-2	3	无 No	1823958―2096096	无 No	[52]
qCTB10-2	10	无 No	4892513―8990159	LOC_Os10g11730 LOC_Os10g11770 LOC_Os10g11810	[53]
qCTB7	7	有 Yes	3866978―391073	LOC_Os07g07690	[57]

QTL名称 QTL name	染色体 Chromosome	遗传功能验证 Genetic function verification	物理位置 Physical position (bp)	候选基因 Candidate gene	参考文献Reference
qCTB8	8	无 No	2900000―3100000	LOC_Os08g44340	[50]
qLTB3	3	无 No	33540013―34653353	LOC_Os03g57680 LOC_Os03g59200	[51]
qCT-3-2	3	无 No	1823958―2096096	无 No	[52]
qCTB10-2	10	无 No	4892513―8990159	LOC_Os10g11730 LOC_Os10g11770 LOC_Os10g11810	[53]
qCTB7	7	有 Yes	3866978―391073	LOC_Os07g07690	[57]

基因名 Gene	基因登录号 Gene ID	优异功能位点 Excellent functional sites	克隆方式 Cloning method	参考文献 Reference
CTB1	LOC_Os04g52830	无 None	正向 Forward	[58]
APXa	LOC_Os03g17690	无 None	反向 Reverse	[38]
CTB4a	LOC_Os04g04330	SNP-2536(G> A) SNP-2511(C> T) SNP-1930(C> G)	正向 Forward	[59]
bZIP73	LOC_Os09g29820	SNP+511(G> A; Lys> Glu)	正向 Forward	[62]
LTT1	LOC_Os05g06660	无 None	正向 Forward	[63]
CTB2	LOC_Os04g04254	SNP+1222（G> A; Val> Ile）	正向 Forward	[60]
WRKY53	LOC_Os05g27730	无 None	反向 Reverse	[64]
MKKK70	LOC_Os01g50410	无 None	反向 Reverse	[65]
MAPK3	LOC_Os03g17700	SNP-1850(T> C) SNP-820(G> A) SNP-807(A> G)	正向 Forward	[61]
LEA9	LOC_Os01g21250	SNP-776(G> A)	正向 Forward	[61]
qCTB7	LOC_Os07g07690	SNP-1110(G> T) SNP-1074(G> A) SNP+4037(C> G; Pro> Arg)	正向 Forward	[57]
COG3	LOC_Os11g44680	SNP-793(C> A) SNP-663(A> G) SNP-379(-> G) SNP-63(G> A)	正向 Forward	[66]

基因名 Gene	基因登录号 Gene ID	优异功能位点 Excellent functional sites	克隆方式 Cloning method	参考文献 Reference
CTB1	LOC_Os04g52830	无 None	正向 Forward	[58]
APXa	LOC_Os03g17690	无 None	反向 Reverse	[38]
CTB4a	LOC_Os04g04330	SNP-2536(G> A) SNP-2511(C> T) SNP-1930(C> G)	正向 Forward	[59]
bZIP73	LOC_Os09g29820	SNP+511(G> A; Lys> Glu)	正向 Forward	[62]
LTT1	LOC_Os05g06660	无 None	正向 Forward	[63]
CTB2	LOC_Os04g04254	SNP+1222（G> A; Val> Ile）	正向 Forward	[60]
WRKY53	LOC_Os05g27730	无 None	反向 Reverse	[64]
MKKK70	LOC_Os01g50410	无 None	反向 Reverse	[65]
MAPK3	LOC_Os03g17700	SNP-1850(T> C) SNP-820(G> A) SNP-807(A> G)	正向 Forward	[61]
LEA9	LOC_Os01g21250	SNP-776(G> A)	正向 Forward	[61]
qCTB7	LOC_Os07g07690	SNP-1110(G> T) SNP-1074(G> A) SNP+4037(C> G; Pro> Arg)	正向 Forward	[57]
COG3	LOC_Os11g44680	SNP-793(C> A) SNP-663(A> G) SNP-379(-> G) SNP-63(G> A)	正向 Forward	[66]