Research Progress on the Mechanism of Herbicide Resistance in Rice (Oryza sativa) and Molecular Breeding
ZHANG Jin-Xia1, HU Xiu-Ming1, SUN Jian-Quan1, WANG He-Le1, ZHANG Qian-Qian1, LIU He-Mei1, YIN Chun-Yuan1, SANG Shi-Fei2, LI Feng-Mei3, NIU Jia-Yu3, HUANG Jin-Hua1,*
1 Xinxiang Academy of Agricultural Sciences, Xinxiang 453002, China; 2 College of Life Sciences, Henan Normal University, Xinxiang 453007, China; 3 Xinxiang University, Xinxiang 453003, China
Abstract：As the most widely cultivated grain crop, the safe production of rice (Oryza sativa) is vital to the staple food source for nearly half of the global population. With the promotion of the simplified cultivation model, the planting area of direct seeding rice has continued to expand. However, the outbreak of drug-resistant weed populations such as Leptochloa chinensis and Echinochloa crus-galli has become a key biological stress factor restricting the improvement of rice yield and quality. Chemical weed control, as the primary method of management, faces issues such as limited selectivity, evolution of weed resistance, and ecological residue. This review summarized the molecular mechanisms of herbicide resistance in rice and recent advances in molecular breeding for herbicide-resistant rice varieties, while also analyzing technical bottlenecks and challenges in current herbicide application. The aim is to provide insights for the development and industrialization of herbicide-resistant rice.
张金霞, 胡秀明, 孙建权, 王和乐, 张倩倩, 刘贺梅, 殷春渊, 桑世飞, 李凤梅, 牛佳宇, 黄金华. 水稻抗除草剂作用机制及分子育种研究进展[J]. 农业生物技术学报, 2026, 34(7): 1553-1568.
ZHANG Jin-Xia, HU Xiu-Ming, SUN Jian-Quan, WANG He-Le, ZHANG Qian-Qian, LIU He-Mei, YIN Chun-Yuan, SANG Shi-Fei, LI Feng-Mei, NIU Jia-Yu, HUANG Jin-Hua. Research Progress on the Mechanism of Herbicide Resistance in Rice (Oryza sativa) and Molecular Breeding. 农业生物技术学报, 2026, 34(7): 1553-1568.
[1] 毕亚玲, 李君君, 戴玲玲, 等.2020. 杂草对除草剂非靶标抗性机理研究进展[J].植物保护, 46(5): 1-5. (Bi Y L, Li J J, Dai L L, et al.2020. Research advances in non-target resistance mechanisms of weeds to herbicides[J].Plant Protection, 46(5): 1-5.) [2] 陈东亮, 崔翠, 任义英, 等. 2018. 草铵膦胁迫下油菜苗期叶片药害相关性状的全基因组关联分析[J]. 作物学报, 44(4): 542-553. (Chen D L, Cui C, Ren Y Y, et al.2018. Genome-wide association analysis of some phytotoxicity related traits at seedling stage in rapeseed under glufosinate stress[J].Acta Agronomica Sinica, 44(4): 542-553.) [3] 陈海伟, 张鲁华, 陈德富,等. 2012. 除草剂及抗除草剂作物的应用现状与展望[J]. 生物技术通报, (10): 35-40. (Chen H W, Zhang L H, Chen D F, et al. 2012. Current utilization status and future prospect of herbicide and herbicide resistant crops[J]. Biotechnology Bulletin, (10): 35-40.) [4] 陈竹锋, 王承旭, 柳威, 等. 2013. 水稻抗除草剂蛋白及其在植物育种中的应用[P]. CN201210037789.9. (Chen Z F, Wang C X, Liu W, et al.2013. Herbicide-resistant protein in rice and its application in plant breeding[P]. CN201210037789.9.) [5] 崔永祯, 杨晓云, 李绍祥, 等. 2022. 转基因技术与基因编辑技术在抗除草剂农作物上的应用[J]. 中国种业, (4): 19-22. (Cui Y Z, Yang X Y, Li S X, et al. 2022. Application of transgenic technology and gene editing technology in herbicide-resistant crops[J]. China Seed Industry, (4): 19-22.) [6] 戴玲玲, 梁蓉, 谢广乐, 等. 2024. 抗除草剂作物的研发与应用进展[J]. 杂草学报, 42(01): 1-16. (Dai L L, Liang R, Xie G L, et al.2024. Progress in development and application of herbicide-resistant crops[J]. Journal of Weed Science, 42(01): 1-16.) [7] 戴燚, 赵德刚. 2018. 抗草甘膦水稻突变体osgr-1 EPSPS基因克隆及生物信息学分析[J]. 种子, 37(3): 1-6, 11. (Dai Y, Zhao D G.2018. Bioinformatic analysis of EPSPS gene from the rice resistant mutant osgr-1 of glyphosate[J]. Seed, 37(3): 1-6,11.) [8] 邓力华, 邓晓湘, 魏岁军, 等. 2014. 抗虫抗除草剂转基因水稻B1C893的获得与鉴定[J]. 杂交水稻, 29(01): 67-71+75. (Deng L H, Deng X X, Wei S J, et al.2014. Development and identification of herbicide and insect resistant transgenic plant B1C893 in rice[J]. Hybrid Rice, 29(01):67-71+75.) [9] 丁佳宁, 周利斌. 2024. 我国水稻辐射诱变育种研究进展[J]. 辐射研究与辐射工艺学报, 42(02): 4-20. (Ding J N, Zhou L B.2024. Progress on radiation-induced mutation breeding of rice in China[J]. Journal of Radiation Research and Radiation Processing, 42(02): 4-20.) [10] 董世凤, 张梦园, 邢凌云, 等. 2025. 作物抗除草剂的遗传研究与应用[J]. 分子植物育种, 23(22): 7496-7504. (Dong S F, Zhang M Y, Xing L Y, et al.2025. Genetic research and application of crop herbicide resistance[J]. Molecular Plant Breeding, 23(22): 7496-7504.) [11] 费云燕, 杨军, 景德道, 等. 2022. CRISPR/Cas技术在抗除草剂作物育种中的研究与应用进展[J]. 生物技术进展, 12(2): 189-197. (Fei Y Y, Yang J, Jing D D, et al.2022. Research and application progress of CRISPR/Cas technology in herbicide-resistant crops breeding[J].Current Biotechnology, 12(2): 189-197.) [12] 胡江博, 任正鹏, 丁翔, 等. 2023. 稻田除草剂应用现状与抗除草剂水稻育种研究进展[J]. 中国稻米, 29(04): 13-19. (Hu J B, Ren Z P, Ding X, et al.2023. Application of herbicides in rice fields and research progress on herbicide-resistant rice varieties breeding[J]. China Rice, 29(04): 13-19.) [13] 黄世霞, 王庆亚, 董立尧, 等. 2003. 乙酰辅酶A羧化酶抑制剂类除草剂与杂草的抗药性[J]. 杂草科学, 2: 1-5. (Huang S X, Wang Q Y, Dong L Y, et al.2003. ACCase-inhibiting herbicides and weed resistance[J]. Weed Science, 2: 1-5.) [14] 蒋显斌, 陈玉冲, 邓力华, 等. 2015. 转基因抗除草剂水稻 Bar68-1的遗传稳定性分析[J]. 西南农业学报, 28(3): 925-929. (Jiang X B, Chen Y C, Deng L H, et al.2015. Analysis on genetic stability of genetically modified herbicide-tolerant (GMHT) rice Bar68-1[J]. Southwest China Journal of Agricultural Sciences, 28(3): 925-929.) [15] 李关羽, 付颖, 叶非. 2012. 靶标酶在除草剂研究与开发中的作用[J]. 农药科学与管理, 33(9): 20-24. (Li G Y, Fu Y, Ye F.2012. The role of target enzymes in the research and development of herbicides[J]. Pesticide Science And Administration, 33(9): 20-24.) [16] 李慧冬. 2020. 莠去津等多种农药残留风险评估及莠去津在水/沉积物体系中降解研究[D]. 博士学位论文, 华中农业大学, 导师: 朱端卫, pp. 87-89. (Li H D.2020. Research on multy-rediue risk assessment of pesticides including atrazine and degradation of atrazine in the system of sediment and water[D]. Thesis for Ph. D., Huazhong Agricultural University, Supervisor: Zhu D W, pp. 87-89.) [17] 李杰华. 2018. 抗广谱性除草剂转基因油菜创制及抗性评价[D]. 硕士学位论文, 华中农业大学, 导师:周永明, pp. 8-10. (Li J H.2018. Development and evaluation of transgenic rapeseed with broad-spectrum herbicide resistant gene[D]. Thesis for M. S., Huazhong Agricultural University, Supervisor: Zhou Y M, pp. 8-10.) [18] 李香菊. 2023. 我国耐除草剂转基因作物研发与产业化应用前景[J]. 植物保护, 49(05): 316-324. (Li X J.2023. Development of herbicide tolerant crops and their commercialization in China[J]. Plant Protection, 49(05): 316-324.) [19] 李燕敏, 祁显涛, 刘昌林, 等. 2017. 除草剂抗性农作物育种研究进展[J]. 作物杂志, (2): 1-6. (Li Y M, Qi X T, Liu C L, et al. 2017. Progress of crop breeding on resistance to herbicides[J]. Crops, (2): 1-6.) [20] 林清晨. 2023. 水稻HPPD抑制剂类除草剂抗性基因的功能分析和挖掘研究[D]. 硕士学位论文, 南京农业大学, 导师: 张保龙, pp. 3-5. (Lin Q C.2023. Functional analysis and excavation of resistance genes of HPPD inhibitor herbicides in rice[D]. Thesis for M. S., Nanjing:Nanjing Agricultural University, Supervisor: Zhang B L, pp. 3-5.) [21] 刘明宏. 2012. 耐二种除草剂转基因水稻研究[D]. 博士学位论文, 浙江大学, 导师: 沈志成, pp. 65-69. (Liu M H.2012. Development of transgenic rice tolerant to two herbicides[D]. Thesis for Ph. D., Zhejiang University, Supervisor: Shen Z C, pp. 65-69.) [22] 刘洋. 2017. HPPD抑制剂类除草剂在水稻上的开发与应用[J]. 农药市场信息, 4: 24-27. (Liu Y.2017. Development and application of HPPD-inhibiting herbicides on rice[J]. Pesticide Market Information, 4: 24-27.) [23] 阮燕晔, 薛寥莎. 2012. 植物抗除草剂基因研究进展[J]. 南方农业学报, 43(04): 462-466. (Ruan Y Y, Xue L S.2012. Advances in plant anti-herbicide gene[J]. Journal of Southern Agriculture, 43(4): 462-466.) [24] 韶也. 2021. 咪唑啉酮与草甘膦除草剂抗性水稻种质创新研究[D]. 博士学位论文, 湖南农业大学, 导师: 刘雄伦, pp. 64-65. (Shao Y.2021. Studies on germplasm innovation of imidazolinones and glyphosate herbicide resistant rice[D]. Thesis for Ph. D., Hunan Agricultural University, Supervisor: Liu X L, pp. 64-65.) [25] 苏少泉. 2002. 生物技术与抗除草剂作物[M]. 北京: 化学工业出版社, pp.102-110. (Su S Q.2002. Biotechnology and Herbicide-resistant Crops[M]. Beijing: Chemical Industry Press, pp.102-110.) [26] 苏旺苍, 孙兰兰, 张强, 等. 2013. 甲咪唑烟酸在土壤中的残留对后茬小麦幼苗生长和光合作用的影响[J]. 麦类作物学报, 33(6): 1226-1231. (Su W C, Sun L L, Zhang Q, et al.2013. Effects of imazapic residues on the growth and photosynthetic parameters of wheat seedlings as succeeding crop[J]. Acta Agronomica Sinica, 33(6): 1226-1231.) [27] 孙丙耀, 倪丕冲, 沈桂芳. 2002. 农作物抗除草剂基因工程[J]. 生物技术通报, (6): 5-10. (Sun B Y, Ni P H, Shen G F. 2002. Advances of genetic engineering for herbicide resistance in plant crops[J]. Biotechnology Bulletin, (6): 5-10.) [28] 唐晓艳, 邓兴旺, 周君莉, 等. 2018. 除草剂抗性突变体及其应用[P]. 201810138888.3. (Tang X Y, Deng X W, Zhou J L, et al.2018. Herbicide-resistant mutant and application[P]. 201810138888.3.) [29] 唐旭. 2020. 基于CRISPR-Cas的高效水稻基因组编辑系统构建及应用[D]. 博士学位论文, 电子科技大学, 导师: 张勇, pp. 62-73. (Tang X.2020. Construction and application of efficient CRISPR-Cas genome editing systems in rice[D]. Thesis for Ph. D., University of Electronic Science and Technology of China, Supervisor:Zhang Y, pp.62-73.) [30] 万丽丽, 王转茸, 辛强, 等. 2016. GOX-CP4EPSP转化甘蓝型油菜的抗草甘膦研究[J]. 湖北农业科学, 55(10): 2661-2666. (Wan L L, Wang Z R, Xin Q, et al.2016. Study on glyphosate-resistant GOX-CP4EPSP transformation of Brassica napus L.[J]. Hubei Agricultural Sciences, 55(10): 2661-2666.) [31] 王芳权, 杨杰, 范方军, 等. 2018. 水稻抗咪唑啉酮类除草剂基因ALS功能标记的开发与应用[J]. 作物学报, 44(3): 324-331. (Wang F Q, Yang J, Fan F J, et al.2018. Development and application of the functional marker for imidazolinone herbicides resistant ALS gene in rice[J]. Acta Agronomica Sinica, 44(3): 324-331.) [32] 王丰玲, 张英锋, 李阳, 等. 2009. 转基因抗除草剂作物的研究进展[J]. 渤海大学学报(自然科学版), 30(1): 13-16. (Wang F L, Zhang Y F, Li Y, et al.2009. Research progress on transgenic herbicide-resistant crops[J]. Journal of Bohai University (Natural Science Edition), 30(1): 13-16.) [33] 王柯. 2023. 转基因抗除草剂油菜的抗性特征研究[D]. 硕士学位论文, 长江大学, 导师: 危文亮, pp. 43-45. (Wang K.2023. Study on resistance characteristics of transgenic Brassica napus with herbicide-resistance[D]. Thesis for M. S., Yangtze University, Supervisor: Wei W L, pp. 43-45.) [34] 王云鹏, 马景勇, 马瑞, 等. 2014. 土壤宏基因组中抗草甘膦新基因的克隆与转化水稻的研究[J]. 作物学报, 40(7):1190-1196. (Wang Y P, Ma J Y, Ma R, et al.2014. Cloning of new herbicide resistant gene in soil metagenomicsand generation of transgenic rice plants[J]. Acta Agronomica Sinica, 40(7): 1190-1196.) [35] 翁绿水, 蒋利平, 肖国樱. 2013. 抗虫抗除草剂转基因水稻恢复系B2A68的培育[J]. 杂交水稻, 28(1): 63-67. (Weng L S, Jiang L P, Xiao G Y.2013. Development of an insect-resistant and herbicide-resistant transgenic restorer line B2A68 in rice[J]. Hybrid Rice, 28(1): 63-67.) [36] 吴元龙, 惠凤娇, 潘振远, 等. 2023. 后基因组时代发展抗除草剂作物的机遇及挑战[J]. 中国农业科学, 56(17):3285-3301. (Wu Y L, Hui F J, Pan Z Y, et al.2023. Opportunities and challenges for developing herbicide-resistance crops in the post-genomic era[J]. Scientia Agricultura Sinica, 56(17):3285-3301.) [37] 项星淳, 芋鑫鑫, 符晶晶, 等. 2024. 抗除草剂基因及其在作物中的应用研究进展[J]. 植物生理学报, 60(07): 1068-1078. (Xiang X C, Yu X X, Fu J J, et al.2024. Progress in the study of herbicide-resistant genes and their application in crops[J]. Plant Physiology Journal, 60(07):1068-1078.) [38] 谢小艳, 郝培媛, 谭杰, 等.2025. 水稻原卟啉原Ⅸ氧化酶编码基因OsPPO1的功能及除草剂抗性分析[J/OL]. 分子植物育种, https://link.cnki.net/urlid/46.1068.S.20250314.1918.013. (Xie X Y, Hao P Y, Tan J,et al.2025. Functional analysis of the rice protoporphyrinogen ix oxidase encoding gene osppo1 and herbicide resistance[J/OL]. Molecular Plant Breeding, https://link-cnki-net-s.webvpn.hnagri.org.cn/urlid/46.1068.S.20250314.1918.013.) [39] 徐海英, 何峰, 初宇, 等. 2016. 一种来源于恶臭假单胞菌的抗HPPD抑制剂基因及其应用[P]. CN105567706A. (Xu H Y, He F, Chu Y, et al.2016. An anti-HPPD inhibitor gene derived from Pseudomonas putida and application[P]. CN105567706A.) [40] 尹丽颖, 张元野, 李荣田, 等. 2022. 利用CRISPR/Cas9技术创制高效抗除草剂水稻[J]. 中国水稻科学, 36(5): 459-466. (Yin L Y, Zhang Y Y, Li R T, et al.2022. Improvement of herbicide resistance in rice by using CRISPR/Cas9 system[J].China Journal of Rice Science, 36(5): 459-466.) [41] 尹言言, 刘靖, 郑炳松, 等. 2025. 脱落酸调控植物非生物胁迫研究进展[J]. 核农学报, 39(09): 1916-1927. (Yin Y Y, L iu J, Zheng B S, et al.2025. Research progress of abscisic acid in regulating plant abiotic stress[J]. Journal of Nuclear Agricultural Sciences, 39(09): 1916-1927.) [42] 俞蕴馨, 伏建国, 李井干, 等. 2020. 全球抗草甘膦杂草的发生概况与检疫思考[J]. 植物检疫, 34(5): 21-27. (Yu Y X, Fu J G, Li J G, et al.2020. Global occurrence of glyphosate-resistant weeds and quarantine thoughts[J]. Plant Quarantine, 34(5): 21-27. [43] 张保龙, 王金彦, 凌溪铁. 2018. 使植物具有除草剂抗性的ACCase突变型蛋白及其应用[P]. CN108369646A. (Zhang B L, Wang J Y, Ling X T.2018. ACCase mutant protein conferring herbicide resistance to plants and application[P]. CN108369646A.) [44] 张保龙, 陈天子, 王金彦, 等. 2017. 粳稻ALS突变型基因及其蛋白在抗除草剂方面的应用[P]. CN106868027A. (Zhang B L, Chen T Z, Wang J Y, et al.2017. Application of Japonica rice ALS mutant gene and its protein in herbicide resistance[P]. CN106868027A.) [45] 张彬, 金燕, 张自常, 等. 2014. 解草啶浸种减轻丙草胺对水稻药害的机制[J]. 江苏农业学报, 30(06): 1345-1349. (Zhang B, Jin Y, Zhang Z C, et al.2014. Mechanism of fenclorim reducing the phytotoxicity of pretilachlor to rice by seed soaking[J]. Jiangsu Journal of Agricultural Sciences, 30(06): 1345-1349.) [46] 张洪程, 龚金龙. 2014. 中国水稻种植机械化高产农艺研究现状及发展探讨[J]. 中国农业科学, 47(7): 1273-1289. (Zhang H C, Gong J L.2014. Research status and development discussion on high-yielding agronomy of mechanized planting rice in China[J]. Scientia Agricultura Sinica,47(7): 1273-1289.) [47] 张怡, 陈丽萍, 徐笔奇, 等. 2020. 浙江稻区千金子对氰氟草酯和噁唑酰草胺的抗药性及其分子机制研究[J]. 农药学学报, 22(3): 447-453. (Zhang Y, Chen L P, Xu B Q, et al.2020. Resistance of Leptochloa chinensis (L.) Nees tocyhalofop-butyl and metamifop in rice fields of Zhejiang province and involved molecular mechanism[J]. Chinese Journal of Pesticide Science, 22(3): 447-453.) [48] 周延彪, 秦鹏, 赵新辉, 等. 2019. 抗除草剂水稻种质资源研究进展[J]. 杂交水稻, 34(01): 1-5. (Zhou Y B, Qin P, Zhao X H, et al.2019. Research progress of herbicide-resistant rice genetic resources[J]. Hybrid Rice, 34(01): 1-5.) [49] 周蕴赟, 李正名. 2013. HPPD抑制剂类除草剂作用机制和研究进展[J]. 世界农药, 35(1): 1-7. (Zhou Y Y, Li Z M, 2013. Recent advances in 4-Hydroxyphenylpyruvate dioxygenase (HPPD) inhibitors[J]. World Pesticide, 35(1):1-7.) [50] 左平春, 纪明山, 臧晓霞, 等. 2017. 稻田稗草对唑酰草胺的抗药性水平和ACCase活性[J]. 植物保护学报, 44(6):1040-1045. (Zuo P C, Ji M S, Zang X X, et al.2017. The resistance level and acetyl co-enzyme a carboxylase activity of echinochloa phyllopogon populations to metamifop in paddies[J]. Journal of Plant Protection, 44(6):1040-1045.) [51] Agostinetto D, Benemann D P, Cechin J, et al.2019. Gene expression related to oxidative stress induced by herbicides in rice[J]. Agronomy Journal, 111(3): 1239-1246. [52] Akbar N, Ehsanullah, Jabran K, et al.2011. Weed management improves yield and quality of direct seeded rice[J]. Australian Journal of Crop Science, 5(6): 688-694. [53] Antralina M, Istina I N, Yuwariah Y, et al.2015. Effect of different weed control methods to yield of lowland rice in the SOBARI[J]. Procedia Food Science, 3: 323-329. [54] Busconi M, Rossi D, Lorenzoni C, et al.2012. Spread of herbicide-resistant weedy rice (red rice, Oryza sativa L.) after 5 years of clear field rice cultivation in Italy[J]. Plant Biology, 14: 751-759. [55] Castle L A, Siehl D L, Gorton R, et al.2004. Discovery and directed evolution of a glyphosate tolerance gene[J]. Science, 304(5674): 1151-1154. [56] Dong Y F,Jin X,Tang Q L,et al.2017. Development and event-specific detection of transgenic glyphosate-resistant rice expressing the G2-EPSPS gene[J]. Frontiers in Plant Science, 8: 885. [57] Fritze I M,Linden L,Freigang J, et al.2004. The crystal structures of Zea mays and Arabidopsis 4-hydroxyphenylpyruvate dioxygenase[J]. Plant Physiology, 134(4): 1388-1400. [58] Ge X, D'Avignon D A, Ackerman J J H, et al.2012. Vacuolar glyphosate-sequestration correlates with glyphosate resistance in Ryegrass (Lolium spp.) from Australia, South America, and Europe: A 31P NMR investigation[J]. Journal of Agricultural and Food Chemistry, 60(5): 1243-1250. [59] Han H P, Yu Q, Purba E, et al.2012. A novel amino acid substitution Ala-22-Tyr in ALS confers high-level and broad resistance across ALS-inhibiting herbicides[J]. Pest Management Science, 68(8): 1164-1170. [60] Hartmut A, Gudrun L, Thomas M, et al.2013. ChemInform abstract: 4‐hydroxyphenylpyruvate dioxygenase inhibitors in combination with safeners: Solutions for modern and sustainable agriculture[J]. ChemInform, 52(36): 9388-9398. [61] Kawahigashi H, Hirose S, Inui H, et al.2005. Enhanced herbicide cross-tolerance in transgenic rice plants co-expressing human CYP1A1, CYP2B6 and CYP2C19[J]. Plant Science, 168(3): 773-781. [62] Kuang Y, Li S, Ren B, et al.2020. Base-editing-mediated artificial evolution of OsALS1 in planta to develop novel herbicide tolerant rice germplasms[J]. Molecular Plant, 13(4): 565-572. [63] Li C, Zong Y, Wang Y, et al.2018. Expanded base editing in rice and wheat using a Cas9-adenosine deaminase fusion[J]. Genome Biology, 19(1): 1-9. [64] Li H, Li J, Chen J, et al.2020. Precise modifications of both exogenous and endogenous genes in rice by prime editing[J]. Molecular Plant, 13(5): 671-674. [65] Li J, Meng X B, Zong Y, et al.2016. Gene replacements and insertions in rice by intron targeting using CRISPR-Cas9[J]. Nature Plants, 2(10): 16139. [66] Liu L, Kuang Y, Yan F, et al.2021. Developing a novel artificial rice germplasm for dinitroaniline herbicide resistance by baseediting of OsTubA2[J]. Plant Biotechnology, 19(1): 5-7. [67] Luo Q, Liu Y G.2025. Breeding herbicide-resistant rice using CRISPR-Cas gene editing and other technologies[J]. Plant Communications, 6(2): 101172. [68] Maeda H, Murata K, Sakuma N, et al.2019. A rice gene that confers broad-spectrum resistance to β-triketone herbicides[J]. Science, 365(64510): 393-396. [69] Maher M F, Nasti R A, Vollbrecht M, et al.2020. Plant gene editing through denovo induction of meristems[J]. Nature Biotechnology, 38(1): 84-89. [70] Pan L, Yu Q, Wang J, et al.2021. An ABCC-type transporter endowing glyphosate resistance in plants[J]. Proceedings of the National Academy of Sciences of the USA, 118(16): e2100136118. [71] Ren Y, Liu B M, Jiang H R, et al.2023. Precision editing of GLR1 confers glufosinate resistance without yield penalty in rice[J]. Plant Biotechnology Journal, 21(12): 2417-2419. [72] Saika H, Horita J, Taguchi-Shiobara F, et al.2014. A novel rice cytochrome P450 gene, CYP72A31, confers tolerance to acetolactate synthase-inhibiting herbicides in rice and Arabidopsis[J]. Plant Physiology, 166(3): 1232-1240. [73] Shimatani Z, Fujikura U, Ishii H, et al.2018. Herbicide tolerance-assisted multiplex targeted nucleotide substitution in rice[J]. Data Brief, 20: 1325-1331. [74] Tian X J, Wang Z Y, Li X F, et al.2015. Characterization and functional analysis of pyrabactin resistance-like abscisic acid receptor family in rice[J]. Rice, 8(1): 28. [75] Tian Y S, Xu J, Xing X J, et al.2015. Improved glyphosate resistance of 5-enolpyruvylshikimate-3-phosphate synthase from Vitis vinifera in transgenic Arabidopsis and rice by DNA shuffling[J]. Molecular Breeding, 35(7): 148. [76] Tseng T, Ou J, Wang C, et al.2013. Role of the ascorbate-glutathione cycle in paraquat tolerance of rice[J]. Weed Science, 61(3): 361-373. [77] Wang F Q, Xu Y, Li W Q, et al.2021. Creating a novel herbicidetolerance OsALS allele using CRISPR/Cas9-mediated gene editing[J]. The Crop Journal, 9(2): 305-312. [78] Wang S W, Shi L L, Sun Z X, et al.2005. Agrobacterium-mediated transformation of rice (Oryza sativa L.) with atrazine chlorohydrolase gene (atzA)[J]. Agricultural Sciences in China, 4(4): 241-246. [79] Wu Y, Xiao N, Cai Y, et al.2023. CRISPR/Cas9-mediated editing of OsHPPD 3'-UTR confers enhanced resistance to HPPD-inhibiting herbicide in rice[J]. Plant Communications, 4(5): 100605. [80] Xu W, Zhang C, Yang Y, et al.2020. Versatile nucleotides substitution in plant using an improved prime editing system[J]. Molecular Plant, 13(5): 675-678. [81] Yan D R, Ren B, Liu L, et al.2021. High-efficiency and multiplex adenine baseediting in plants using new TadA variants[J]. Molecular Plant, 14(5): 722-731. [82] Zhang C, Zhong X, Li S, et al.2023. Artificial evolution of OsEPSPS through an improved dual cytosine and adenine base editor generated a novel allele conferring rice glyphosate tolerance[J]. Journal of Integrative Plant Biology, 65(9): 2194-2203. [83] Zhao T,Lin C Y,Shen Z C.2011. Development of transgenic glyphosate-resistant rice with G6 gene encoding 5-enolpyruvylshikimate-3-phosphate synthase[J]. Agricultural Sciences in China, 10(9): 1307-1312.
