水稻赤霉素信号负调控因子SLR1的生物学功能及其调控网络

doi:10.13560/j.cnki.biotech.bull.1985.2025-0643

摘要/Abstract

摘要：

赤霉素（GAs）是植物中一类重要的调控激素，广泛参与植物生长发育与逆境响应等多种生命过程。GA的合成及信号通路调控促成了作物育种的第一次绿色革命。SLR1是水稻中唯一的DELLA蛋白，是GA信号转导途径中关键的负调控因子，阻遏GA下游信号转导。SLR1还参与了脱落酸（ABA）、茉莉酸（JA）、油菜素内酯（BR）、独脚金内酯（SL）等激素途径，充当植物激素互作的“分子桥梁”。然而，SLR1蛋白不具备典型的DNA结合域，目前仍无证据表明SLR1能直接结合DNA序列，其主要通过与其他转录因子互作抑制或激活下游基因的表达来发挥调控功能。此外，SLR1自身的表达与功能也受到多种调控。在转录水平，SLR1基因受到OsYABBY4、OsWRKY36等转录因子的负调控；在蛋白水平，SLR1受到泛素化、糖基化、SUMO化、磷酸化等修饰，以及与其他蛋白互作对自身稳定性与活性的调控。SLR1不仅广泛调控水稻多种生长发育过程，还参与水稻响应多种生物与非生物胁迫，在水稻的整个生长周期中扮演了重要的角色。最新研究揭示SLR1在育种中的巨大潜力，通过调控SLR1在水稻中的蛋白丰度并使其维持中等含量，对提高水稻耐碱‒热性具有重要意义。本文综述了水稻SLR1分子结构和作用机制、在各植物激素途径中的串联作用、SLR1自身的蛋白修饰与调控以及具体的生物学功能，以期进一步发掘SLR1蛋白关联组分和调控网络，为水稻分子设计育种提供参考。

关键词: 水稻, SLR1蛋白, 赤霉素, 激素互作, 蛋白修饰, 调控网络, 生物学功能, 逆境胁迫

Abstract:

Gibberellins (GAs) are a class of important regulatory hormones in plants, widely involved in various life processes such as plant growth and development, and responses to stress. The regulation of GA biosynthesis and signaling pathways contributed to the first green revolution in crop breeding. SLR1 is the only DELLA protein in rice, serving as a key negative regulator in the GA signal transduction pathway that represses downstream GA signal transduction. SLR1 also participates in hormone pathways such as abscisic acid (ABA), jasmonic acid (JA), brassinosteroid (BR), and strigolactone (SL), acting as a "molecular bridge" for plant hormone crosstalk. However, the SLR1 protein lacks a typical DNA-binding domain, and there is currently no evidence indicating its ability to directly bind to DNA sequences. It mainly exerts its modulatory functions by interacting with other transcription factors to inhibit or activate the expressions of downstream genes. In addition, the expression and function of SLR1 itself are also subject to multiple regulations. At the transcriptional level, the SLR1 gene is negatively regulated by transcription factors such as OsYABBY4 and OsWRKY36; at the protein level, SLR1 undergoes modifications including ubiquitination, glycosylation, SUMOylation, and phosphorylation, as well as regulation of its own stability and activity through interactions with other proteins. SLR1 not only extensively regulates various growth and development processes but also participates responses to multiple biotic and abiotic stresses in rice, playing an important role throughout the entire growth cycle of rice. Recent research has revealed the great potential of SLR1 in breeding: regulating the protein abundance of SLR1 in rice and maintaining it at a moderate level is of great significance for improving rice tolerance to alkali and heat stress. This article mainly reviews the molecular structure and mechanism of action of SLR1, its crosstalk role in various plant hormone pathways, the protein modifications and regulations of SLR1 itself, as well as its specific biological functions. It aims to further explore SLR1-associated components and regulatory networks, providing references for rice molecular design breeding.

Key words: rice, SLR1 protein, gibberellin, hormone crosstalk, protein modification, regulatory network, biological function, abiotic stress

费思恬, 侯鹰翔, 李兰, 张超. 水稻赤霉素信号负调控因子SLR1的生物学功能及其调控网络[J]. 生物技术通报, 2026, 42(1): 13-30.

FEI Si-tian, HOU Ying-xiang, LI Lan, ZHANG Chao. Biological Functions and Regulatory Network of SLR1， a Negative Regulator of Gibberellin Signaling in Rice[J]. Biotechnology Bulletin, 2026, 42(1): 13-30.

图/表 7

参考文献 103

[1]	Hedden P, Proebsting WM. Genetic analysis of gibberellin biosynthesis [J]. Plant Physiol, 1999, 119(2): 365-370.
[2]	Olszewski N, Sun TP, Gubler F. Gibberellin signaling: biosynthesis, catabolism, and response pathways [J]. Plant Cell, 2002, 14(): S61-S80.
[3]	Richards DE, King KE, Ait-Ali T, et al. How gibberellin regulates plant growth and development: a molecular genetic analysis of gibberellin signaling [J]. Annu Rev Plant Physiol Plant Mol Biol, 2001, 52: 67-88.
[4]	Khush GS. Green revolution: preparing for the 21st century [J]. Genome, 1999, 42(4): 646-655.
[5]	Pingali PL. Green revolution: impacts, limits, and the path ahead [J]. Proc Natl Acad Sci USA, 2012, 109(31): 12302-12308.
[6]	Monna L, Kitazawa N, Yoshino R, et al. Positional cloning of rice semidwarfing gene, sd-1: rice "green revolution gene" encodes a mutant enzyme involved in gibberellin synthesis [J]. DNA Res, 2002, 9(1): 11-17.
[7]	Spielmeyer W, Ellis MH, Chandler PM. Semidwarf (sd-1), "green revolution" rice, contains a defective gibberellin 20-oxidase gene [J]. Proc Natl Acad Sci USA, 2002, 99(13): 9043-9048.
[8]	Hedden P. The genes of the green revolution [J]. Trends Genet, 2003, 19(1): 5-9.
[9]	Sun TP, Gubler F. Molecular mechanism of gibberellin signaling in plants [J]. Annu Rev Plant Biol, 2004, 55: 197-223.
[10]	Pysh LD, Wysocka-Diller JW, Camilleri C, et al. The GRAS gene family in Arabidopsis: sequence characterization and basic expression analysis of the SCARECROW-LIKE genes [J]. Plant J, 1999, 18(1): 111-119.
[11]	Ogawa M, Kusano T, Katsumi M, et al. Rice gibberellin-insensitive gene homolog, OsGAI, encodes a nuclear-localized protein capable of gene activation at transcriptional level [J]. Gene, 2000, 245(1): 21-29.
[12]	Fu XD, Richards DE, Ait-Ali T, et al. Gibberellin-mediated proteasome-dependent degradation of the barley DELLA protein SLN1 repressor [J]. Plant Cell, 2002, 14(12): 3191-3200.
[13]	Jiang CF, Fu XD. GA action: turning on de-DELLA repressing signaling [J]. Curr Opin Plant Biol, 2007, 10(5): 461-465.
[14]	Ikeda A, Ueguchi-Tanaka M, Sonoda Y, et al. Slender rice, a constitutive gibberellin response mutant, is caused by a null mutation of the SLR1 gene, an ortholog of the height-regulating gene GAI/RGA/RHT/D8 [J]. Plant Cell, 2001, 13(5): 999-1010.
[15]	Willige BC, Ghosh S, Nill C, et al. The DELLA domain of GA INSENSITIVE mediates the interaction with the GA INSENSITIVE DWARF1A gibberellin receptor of Arabidopsis [J]. Plant Cell, 2007, 19(4): 1209-1220.
[16]	Ueguchi-Tanaka M, Nakajima M, Katoh E, et al. Molecular interactions of a soluble gibberellin receptor, GID1, with a rice DELLA protein, SLR1, and gibberellin [J]. Plant Cell, 2007, 19(7): 2140-2155.
[17]	Van De Velde K, Ruelens P, Geuten K, et al. Exploiting DELLA signaling in cereals [J]. Trends Plant Sci, 2017, 22(10): 880-893.
[18]	Li MZ, An FY, Li WY, et al. DELLA proteins interact with FLC to repress flowering transition [J]. J Integr Plant Biol, 2016, 58(7): 642-655.
[19]	Gomi K, Sasaki A, Itoh H, et al. GID2, an F-box subunit of the SCF E3 complex, specifically interacts with phosphorylated SLR1 protein and regulates the gibberellin-dependent degradation of SLR1 in rice [J]. Plant J, 2004, 37(4): 626-634.
[20]	Itoh H, Ueguchi-Tanaka M, Sato Y, et al. The gibberellin signaling pathway is regulated by the appearance and disappearance of SLENDER RICE1 in nuclei [J]. Plant Cell, 2002, 14(1): 57-70.
[21]	白云赫, 朱旭东, 樊秀彩, 等. 植物DELLA蛋白及其应答赤霉素信号调控植物生长发育的研究进展 [J]. 分子植物育种, 2019, 17(8): 2509-2516.
	Bai YH, Zhu XD, Fan XC, et al. Reseach progress of plant DELLA proteins and its response to gibberellin signal regulating plant growth and development [J]. Mol Plant Breed, 2019, 17(8): 2509-2516.
[22]	Hirano K, Asano K, Tsuji H, et al. Characterization of the molecular mechanism underlying gibberellin perception complex formation in rice [J]. Plant Cell, 2010, 22(8): 2680-2696.
[23]	Hellens RP, Allan AC, Friel EN, et al. Transient expression vectors for functional genomics, quantification of promoter activity and RNA silencing in plants [J]. Plant Methods, 2005, 1: 13.
[24]	Hauvermale AL, Ariizumi T, Steber CM. Gibberellin signaling: a theme and variations on DELLA repression [J]. Plant Physiol, 2012, 160(1): 83-92.
[25]	Ueguchi-Tanaka M. Gibberellin metabolism and signaling [J]. Biosci Biotechnol Biochem, 2023, 87(10): 1093-1101.
[26]	Phokas A, Coates JC. Evolution of DELLA function and signaling in land plants [J]. Evol Dev, 2021, 23(3): 137-154.
[27]	Yano K, Aya K, Hirano K, et al. Comprehensive gene expression analysis of rice aleurone cells: probing the existence of an alternative gibberellin receptor [J]. Plant Physiol, 2015, 167(2): 531-544.
[28]	Sun HW, Guo XL, Zhu XL, et al. Strigolactone and gibberellin signaling coordinately regulate metabolic adaptations to changes in nitrogen availability in rice [J]. Mol Plant, 2023, 16(3): 588-598.
[29]	Zhao Y, Zhou DX. Epigenomic modification and epigenetic regulation in rice [J]. J Genet Genom, 2012, 39(7): 307-315.
[30]	Chen XS, Zhou DX. Rice epigenomics and epigenetics: challenges and opportunities [J]. Curr Opin Plant Biol, 2013, 16(2): 164-169.
[31]	Li TT, Chen XS, Zhong XC, et al. Jumonji C domain protein JMJ705-mediated removal of histone H3 lysine 27 trimethylation is involved in defense-related gene activation in rice [J]. Plant Cell, 2013, 25(11): 4725-4736.
[32]	Li JJ, Li Q, Wang WT, et al. DELLA-mediated gene repression is maintained by chromatin modification in rice [J]. EMBO J, 2023, 42(21): e114220.
[33]	Itoh H, Matsuoka M, Steber CM. A role for the ubiquitin-26S-proteasome pathway in gibberellin signaling [J]. Trends Plant Sci, 2003, 8(10): 492-497.
[34]	Mashita O, Koishihara H, Fukui K, et al. Discovery and identification of 2-methoxy-1-naphthaldehyde as a novel strigolactone-signaling inhibitor [J]. J Pestic Sci, 2016, 41(3): 71-78.
[35]	Han SN, Liu YX, Bao A, et al. OsCSN1 regulates the growth of rice seedlings through the GA signaling pathway in blue light [J]. J Plant Physiol, 2023, 280: 153904.
[36]	Fernandes T, Gonçalves NM, Matiolli CC, et al. SUMOylation of rice DELLA SLR1 modulates transcriptional responses and improves yield under salt stress [J]. Planta, 2024, 260(6): 136.
[37]	Itoh H, Sasaki A, Ueguchi-Tanaka M, et al. Dissection of the phosphorylation of rice DELLA protein, SLENDER RICE1 [J]. Plant Cell Physiol, 2005, 46(8): 1392-1399.
[38]	Dai C, Xue HW. Rice early flowering1, a CKI, phosphorylates DELLA protein SLR1 to negatively regulate gibberellin signalling [J]. EMBO J, 2010, 29(11): 1916-1927.
[39]	Shimada A, Ueguchi-Tanaka M, Sakamoto T, et al. The rice SPINDLY gene functions as a negative regulator of gibberellin signaling by controlling the suppressive function of the DELLA protein, SLR1 and modulating brassinosteroid synthesis [J]. Plant J, 2006, 48(3): 390-402.
[40]	Yano K, Morinaka Y, Wang FM, et al. GWAS with principal component analysis identifies a gene comprehensively controlling rice architecture [J]. Proc Natl Acad Sci USA, 2019, 116(42): 21262-21267.
[41]	Yang C, Ma YM, Li JX. The rice YABBY4 gene regulates plant growth and development through modulating the gibberellin pathway [J]. J Exp Bot, 2016, 67(18): 5545-5556.
[42]	Lan J, Lin QB, Zhou CL, et al. Small grain and semi-dwarf 3, a WRKY transcription factor, negatively regulates plant height and grain size by stabilizing SLR1 expression in rice [J]. Plant Mol Biol, 2020, 104(4/5): 429-450.
[43]	Zhu N, Cheng SF, Liu XY, et al. The R2R3-type MYB gene OsMYB91 has a function in coordinating plant growth and salt stress tolerance in rice [J]. Plant Sci, 2015, 236: 146-156.
[44]	Zhang YH, Liu K, Zhu XM, et al. Rice tocopherol deficiency 1 encodes a homogentisate phytyltransferase essential for tocopherol biosynthesis and plant development in rice [J]. Plant Cell Rep, 2018, 37(5): 775-787.
[45]	Yang D, Liu X, Yin XM, et al. Rice non-specific phospholipase C6 is involved in mesocotyl elongation [J]. Plant Cell Physiol, 2021, 62(6): 985-1000.
[46]	Chen JS, Wang ST, Jiang SQ, et al. Overexpression of Calcineurin B-like interacting protein kinase 31 promotes lodging and sheath blight resistance in rice [J]. Plants, 2024, 13(10): 1306.
[47]	Davière JM, Achard P. A pivotal role of DELLAs in regulating multiple hormone signals [J]. Mol Plant, 2016, 9(1): 10-20.
[48]	赵春丽, 王晓, 陈家兰, 等. 植物DELLA蛋白家族研究进展 [J]. 应用与环境生物学报, 2020, 26(5): 1299-1308.
	Zhao CL, Wang X, Chen JL, et al. Progress in research on plant DELLA family proteins [J]. Chin J Appl Environ Biol, 2020, 26(5): 1299-1308.
[49]	Eckardt NA. GA perception and signal transduction: molecular interactions of the GA receptor GID1 with GA and the DELLA protein SLR1 in rice [J]. Plant Cell, 2007, 19(7): 2095-2097.
[50]	Asano K, Hirano K, Ueguchi-Tanaka M, et al. Isolation and characterization of dominant dwarf mutants, Slr1-d, in rice [J]. Mol Genet Genomics, 2009, 281(2): 223-231.
[51]	Jung YJ, Kim JH, Lee HJ, et al. Generation and transcriptome profiling of Slr1-d7 and Slr1-d8 mutant lines with a new semi-dominant dwarf allele of SLR1 using the CRISPR/Cas9 system in rice [J]. Int J Mol Sci, 2020, 21(15): 5492.
[52]	Zentella R, Zhang ZL, Park M, et al. Global analysis of DELLA direct targets in early gibberellin signaling in Arabidopsis [J]. Plant Cell, 2007, 19(10): 3037-3057.
[53]	Tsuji H, Aya K, Ueguchi-Tanaka M, et al. GAMYB controls different sets of genes and is differentially regulated by microRNA in aleurone cells and anthers [J]. Plant J, 2006, 47(3): 427-444.
[54]	Gao J, Chen H, Yang HF, et al. A brassinosteroid responsive miRNA-target module regulates gibberellin biosynthesis and plant development [J]. New Phytol, 2018, 220(2): 488-501.
[55]	Hou XL, Lee LYC, Xia KF, et al. DELLAs modulate jasmonate signaling via competitive binding to JAZs [J]. Dev Cell, 2010, 19(6): 884-894.
[56]	Um TY, Lee HY, Lee S, et al. Jasmonate zim-domain protein 9 interacts with slender rice 1 to mediate the antagonistic interaction between jasmonic and gibberellic acid signals in rice [J]. Front Plant Sci, 2018, 9: 1866.
[57]	Li LL, Zhang HH, Yang ZH, et al. Independently evolved viral effectors convergently suppress DELLA protein SLR1-mediated broad-spectrum antiviral immunity in rice [J]. Nat Commun, 2022, 13(1): 6920.
[58]	Nakamura H, Xue YL, Miyakawa T, et al. Molecular mechanism of strigolactone perception by DWARF14 [J]. Nat Commun, 2013, 4: 2613.
[59]	Liao ZG, Zhang YC, Yu Q, et al. Coordination of growth and drought responses by GA-ABA signaling in rice [J]. New Phytol, 2023, 240(3): 1149-1161.
[60]	Liao ZG, Yu H, Duan JB, et al. SLR1 inhibits MOC1 degradation to coordinate tiller number and plant height in rice [J]. Nat Commun, 2019, 10(1): 2738.
[61]	Wu K, Wang SS, Song WZ, et al. Enhanced sustainable green revolution yield via nitrogen-responsive chromatin modulation in rice [J]. Science, 2020, 367(6478): eaaz2046.
[62]	Li XX, Xie ZZ, Qin T, et al. The SLR1-OsMADS23-D14 module mediates the crosstalk between strigolactone and gibberellin signaling to control rice tillering [J]. New Phytol, 2025, 246(5): 2137-2154.
[63]	Zhuang H, Li YF. Strigolactone and gibberellin crosstalk: the role of the SLR1-OsMADS23-D14 module in regulating rice tiller development [J]. New Phytol, 2025, 246(5): 1893-1895.
[64]	Lu YZ, Feng Z, Meng YL, et al. SLENDER RICE1 and Oryza sativa INDETERMINATE DOMAIN2 regulating OsmiR396 are involved in stem elongation [J]. Plant Physiol, 2020, 182(4): 2213-2227.
[65]	Huang DB, Wang SG, Zhang BC, et al. A gibberellin-mediated DELLA-NAC signaling cascade regulates cellulose synthesis in rice [J]. Plant Cell, 2015, 27(6): 1681-1696.
[66]	Chen Y, Qi HY, Yang LJ, et al. The OsbHLH002/OsICE1-OSH1 module orchestrates secondary cell wall formation in rice [J]. Cell Rep, 2023, 42(7): 112702.
[67]	Ye YF, Liu BM, Zhao M, et al. CEF1/OsMYB103L is involved in GA-mediated regulation of secondary wall biosynthesis in rice [J]. Plant Mol Biol, 2015, 89(4/5): 385-401.
[68]	Liu YF, Wu Q, Qin ZL, et al. Transcription factor OsNAC055 regulates GA-mediated lignin biosynthesis in rice straw [J]. Plant Sci, 2022, 325: 111455.
[69]	Aya K, Ueguchi-Tanaka M, Kondo M, et al. Gibberellin modulates anther development in rice via the transcriptional regulation of GAMYB [J]. Plant Cell, 2009, 21(5): 1453-1472.
[70]	宋欣玥. SLR1调控水稻雄蕊发育的功能研究 [D]. 上海: 上海师范大学, 2023.
	Song XY. Study on the function of SLR1 in regulating stamen development in rice [D]. Shanghai: Shanghai Normal University, 2023.
[71]	Tang JQ, Tian XJ, Mei EY, et al. WRKY53 negatively regulates rice cold tolerance at the booting stage by fine-tuning anther gibberellin levels [J]. Plant Cell, 2022, 34(11): 4495-4515.
[72]	Jin Y, Song XY, Chang HZ, et al. The GA-DELLA-OsMS188 module controls male reproductive development in rice [J]. New Phytol, 2022, 233(6): 2629-2642.
[73]	Su S, Hong J, Chen XF, et al. Gibberellins orchestrate panicle architecture mediated by DELLA-KNOX signalling in rice [J]. Plant Biotechnol J, 2021, 19(11): 2304-2318.
[74]	Zhang S, Deng L, Cheng R, et al. RID1 sets rice heading date by balancing its binding with SLR1 and SDG722 [J]. J Integr Plant Biol, 2022, 64(1): 149-165.
[75]	Liu RJ, Feng QF, Li PB, et al. GLW7.1, a strong functional allele of Ghd7, enhances grain size in rice [J]. Int J Mol Sci, 2022, 23(15): 8715.
[76]	Wu H, He Q, He B, et al. Gibberellin signaling regulates lignin biosynthesis to modulate rice seed shattering [J]. Plant Cell, 2023, 35(12): 4383-4404.
[77]	Xie ZZ, Jin L, Sun Y, et al. OsNAC120 balances plant growth and drought tolerance by integrating GA and ABA signaling in rice [J]. Plant Commun, 2024, 5(3): 100782.
[78]	Huang JS, Xie B, Xian FJ, et al. Gibberellin signalling mediates nucleocytoplasmic trafficking of Sucrose Synthase 1 to regulate the drought tolerance in rice [J]. Plant Biotechnol J, 2025, 23(6): 1909-1926.
[79]	Lyu YS, Dong XL, Niu SP, et al. An orchestrated ethylene-gibberellin signaling cascade contributes to mesocotyl elongation and emergence of rice direct seeding [J]. J Integr Plant Biol, 2024, 66(7): 1427-1439.
[80]	Mo WP, Tang WJ, Du YX, et al. PHYTOCHROME-INTERACTING FACTOR-LIKE14 and SLENDER RICE1 interaction controls seedling growth under salt stress [J]. Plant Physiol, 2020, 184(1): 506-517.
[81]	Jin XK, Zhang YF, Li XX, et al. OsNF-YA3 regulates plant growth and osmotic stress tolerance by interacting with SLR1 and SAPK9 in rice [J]. Plant J, 2023, 114(4): 914-933.
[82]	Li Z, Chen H, Guan QJ, et al. Gibberellic acid signaling promotes resistance to saline-alkaline stress by increasing the uptake of ammonium in rice [J]. Plant Physiol Biochem, 2024, 207: 108424.
[83]	Li ZT, Wang B, Zhang ZY, et al. OsGRF6 interacts with SLR1 to regulate OsGA2ox1 expression for coordinating chilling tolerance and growth in rice [J]. J Plant Physiol, 2021, 260: 153406.
[84]	Ge Q, Zhang YY, Xu YY, et al. Cyclophilin OsCYP20-2 with a novel variant integrates defense and cell elongation for chilling response in rice [J]. New Phytol, 2020, 225(6): 2453-2467.
[85]	Mittal L, Tayyeba S, Sinha AK. Finding a breather for Oryza sativa: Understanding hormone signalling pathways involved in rice plants to submergence stress [J]. Plant Cell Environ, 2022, 45(2): 279-295.
[86]	Fukao T, Bailey-Serres J. Submergence tolerance conferred by Sub1A is mediated by SLR1 and SLRL1 restriction of gibberellin responses in rice [J]. Proc Natl Acad Sci USA, 2008, 105(43): 16814-16819.
[87]	Schmitz AJ, Folsom JJ, Jikamaru Y, et al. SUB1A-mediated submergence tolerance response in rice involves differential regulation of the brassinosteroid pathway [J]. New Phytol, 2013, 198(4): 1060-1070.
[88]	Li S, Tian YH, Wu K, et al. Modulating plant growth-metabolism coordination for sustainable agriculture [J]. Nature, 2018, 560(7720): 595-600.
[89]	Mirza Z, Gupta M. Iron reprogrammes the root system architecture by regulating OsWRKY71 in arsenic-stressed rice (Oryza sativa L.) [J]. Plant Mol Biol, 2024, 114(1): 11.
[90]	De Vleesschauwer D, Seifi HS, Filipe O, et al. The DELLA protein SLR1 integrates and amplifies salicylic acid- and jasmonic acid-dependent innate immunity in rice [J]. Plant Physiol, 2016, 170(3): 1831-1847.
[91]	Liu MM, Shi ZY, Zhang XH, et al. Inducible overexpression of Ideal Plant Architecture1 improves both yield and disease resistance in rice [J]. Nat Plants, 2019, 5(4): 389-400.
[92]	Sun Q, Li TY, Li DD, et al. Overexpression of Loose Plant Architecture 1 increases planting density and resistance to sheath blight disease via activation of PIN-FORMED 1a in rice [J]. Plant Biotechnol J, 2019, 17(5): 855-857.
[93]	Sakata T, Oda S, Tsunaga Y, et al. Reduction of gibberellin by low temperature disrupts pollen development in rice [J]. Plant Physiol, 2014, 164(4): 2011-2019.
[94]	Guo SQ, Chen YX, Ju YL, et al. Fine-tuning gibberellin improves rice alkali-thermal tolerance and yield [J]. Nature, 2025, 639(8053): 162-171.
[95]	Lu L, Chen XY, Tan QY, et al. Gibberellin-mediated sensitivity of rice roots to aluminum stress [J]. Plants, 2024, 13(4): 543.
[96]	Hui SG, Ke YG, Chen D, et al. Rice microRNA156/529- SQUAMOSA PROMOTER BINDING PROTEIN-LIKE7/14/17 modules regulate defenses against bacteria [J]. Plant Physiol, 2023, 192(3): 2537-2553.
[97]	Zhu HY, Chen H, Kantharaj V, et al. SLR1-LPA1 signal regulates sheath blight resistance and lamina joint angle in rice [J]. Plant Physiol Biochem, 2025, 222: 109689.
[98]	De Vleesschauwer D, Van Buyten E, Satoh K, et al. Brassinosteroids antagonize gibberellin- and salicylate-mediated root immunity in rice [J]. Plant Physiol, 2012, 158(4): 1833-1846.
[99]	Zhang J, Luo T, Wang WW, et al. Silencing OsSLR1 enhances the resistance of rice to the brown planthopper Nilaparvata lugens [J]. Plant Cell Environ, 2017, 40(10): 2147-2159.
[100]	Yimer HZ, Nahar K, Kyndt T, et al. Gibberellin antagonizes jasmonate-induced defense against Meloidogyne graminicola in rice [J]. New Phytol, 2018, 218(2): 646-660.
[101]	Itoh H, Shimada A, Ueguchi-Tanaka M, et al. Overexpression of a GRAS protein lacking the DELLA domain confers altered gibberellin responses in rice [J]. Plant J, 2005, 44(4): 669-679.
[102]	Liu T, Gu JY, Xu CJ, et al. Overproduction of OsSLRL2 alters the development of transgenic Arabidopsis plants [J]. Biochem Biophys Res Commun, 2007, 358(4): 983-989.
[103]	Wang JD, Wang J, Huang LC, et al. ABA-mediated regulation of rice grain quality and seed dormancy via the NF-YB1-SLRL2-bHLH144 Module [J]. Nat Commun, 2024, 15(1): 4493.

相关基因或蛋白 Related genes or proteins	相关通路 Related pathways	生物学功能 Biological function	参考文献 Reference
GID1， GID2	GA	介导SLR1的降解，调控GA信号途径 Mediate the degradation of SLR1 and regulate the GA signaling pathway	［19-33］
D14	GA， SL	促进SLR1的降解，调控水稻分蘖 Promote the degradation of SLR1 and regulate rice tillering	［28］
D53	SL，氮素信号	高氮条件下，抑制D14介导SLR1的降解，调控氮素利用 Under high-nitrogen conditions， inhibit D14-mediated degradation of SLR1 to regulate nitrogen utilization	［28］
PRC2， HDA702	GA	介导染色质沉默 Mediate chromatin silencing	［32］
EL1	GA	促进SLR1磷酸化，调控GA信号途径 Promote the phosphorylation of SLR1， regulate the GA signaling pathway	［38］
OsSPY	GA	激活SLR1的抑制活性 Activate the inhibitory activity of SLR1	［39-40］
OsYABBY4	GA	调控小穗发育 Regulate spikelet development	［41］
OsWRKY36	GA	抑制GA信号和调控植株 Inhibit GA signaling and regulate plant height	［42］
OsMYB91	GA， ABA	平衡水稻生长和非生物胁迫抗性 Balance rice growth and abiotic stress resistance	［43］
RTD1	GA	负调控SLR1蛋白的转录和蛋白水平 Negatively regulate the transcription and protein level of SLR1 protein	［44］
OsNPC6	GA	调控中胚轴伸长 Regulate mesocotyl elongation	［45］
CIPK31	GA	抑制SLR1蛋白降解 Inhibit the degradation of SLR1 protein	［46］
OsGAMYBL2	GA， BR	调控GA合成和BR信号 Regulate GA biosynthesis and BR signaling	［54］
OsJAZ8， OsJAZ9	JA	调控水稻生长发育与抗逆 Regulate rice growth and development as well as stress resistance	［56］
MYC2/3	JA	调控JA响应基因的表达 Regulate the expression of JA-responsive genes	［57］
SP8， P2， M	GA， JA	介导广谱抗病毒防御反应 Mediate broad-spectrum antiviral defense response	［57］
TAD1	ABA	抑制ABA信号 Inhibit ABA signaling	［59］
MOC1	GA	调控株高和分蘖 Regulate plant height and tillering	［60］
NGR5	GA	提高氮素利用、促进分蘖 Enhance nitrogen utilization and promote tillering	［61］
OsMADS23	GA， SL	抑制D14基因转录，促进分蘖 Suppress the transcription of the D14 gene and promote tillering	［62-63］
OsIDD2	GA	调控细胞增殖 Regulate cell proliferation	［64］
OsNAC29/31	GA	调控纤维素合成 Regulate cellulose synthesis	［65］
OsKNAT7	GA	调控次生细胞壁的合成 Regulate the synthesis of secondary cell walls	［66］
OsMYB103L	GA	调控纤维素生物和次生细胞壁合成 Regulate cellulose and secondary cell wall biosynthesis	［67］
OsNAC055	木质素合成途径	调控木质素合成 Regulate lignin biosynthesis	［68］
GAMYB	GA	调控水稻育性 Regulate rice fertility	［69-70］
UDT1， TDR	GA	提高绒毡层发育相关基因的表达，调控水稻的育性 Enhance the expressions of tapetum development-related genes and regulate rice fertility	［71］
OsMS188	GA	调控孢粉素生物合成，促进花粉壁的形成 Regulate sporopollenin biosynthesis and promote pollen wall formation	［72］
OSH1	GA	调控次生细胞壁合成 Regulate secondary cell wall biosynthesis	［73］
RID1	GA	调控营养生长向生殖生长的转变 Regulate the transition from vegetative growth to reproductive growth	［74］
GHD7	GA	调控抽穗期和植株形态 Regulate rice heading date and plant morphology	［75］
qSH1， OSH15， SNB	GA	调控木质素含量和落粒性 Regulate lignin biosynthesis and seed shattering	［76］
OsNAC120	GA， ABA	调控干旱胁迫响应 Regulate the response to drought stress	［77］
OsBURP3， OsSUS1	GA	调控干旱胁迫响应 Regulate the response to drought stress	［78］
OsPIL13/14	GA，光信号	调控黑暗或盐胁迫下的幼苗生长 Regulate seedling growth under darkness or salt stress	［79-80］
OsNF-YA3	GA， ABA	调控生长发育与渗透胁迫耐受性 Regulate growth and development as well as osmotic stress tolerance	［81］
IDD10， bZIP23	GA，氮素信号	调控氮素吸收和盐碱耐受性 Regulate nitrogen absorption and saline-alkali tolerance	［82］
OsGRF6	GA	调控水稻生长和耐寒性 Regulate rice growth and cold tolerance	［83］
OsNuCYP20-2	GA	促进SLR1的降解，促进细胞伸长 Promote the degradation of SLR1 and cell elongation	［84］
Sub1A	GA， ET， ABA	促进SLR1的积累，抑制GA信号 Promote the accumulation of SLR1 and inhibit GA signaling	［85-87］
GRF4	GA， BR	调节根系代谢和氮素利用 Regulate root metabolism and nitrogen utilization	［88］
OsWRKY71	GA	调控逆境下的根系发育 Regulate root development under stress conditions	［89］
OsSPL7， OsSPL14	GA	调控Xoo的抗性 Regulate resistance to Xanthomonasoryzae pv. oryzae （Xoo）	［90-91］
LPA1	GA	对水稻纹枯病rice sheath blight（ShB）的抗性 Resistance to rice sheath blight （ShB）	［92］

相关基因或蛋白 Related genes or proteins	相关通路 Related pathways	生物学功能 Biological function	参考文献 Reference
GID1， GID2	GA	介导SLR1的降解，调控GA信号途径 Mediate the degradation of SLR1 and regulate the GA signaling pathway	［19-33］
D14	GA， SL	促进SLR1的降解，调控水稻分蘖 Promote the degradation of SLR1 and regulate rice tillering	［28］
D53	SL，氮素信号	高氮条件下，抑制D14介导SLR1的降解，调控氮素利用 Under high-nitrogen conditions， inhibit D14-mediated degradation of SLR1 to regulate nitrogen utilization	［28］
PRC2， HDA702	GA	介导染色质沉默 Mediate chromatin silencing	［32］
EL1	GA	促进SLR1磷酸化，调控GA信号途径 Promote the phosphorylation of SLR1， regulate the GA signaling pathway	［38］
OsSPY	GA	激活SLR1的抑制活性 Activate the inhibitory activity of SLR1	［39-40］
OsYABBY4	GA	调控小穗发育 Regulate spikelet development	［41］
OsWRKY36	GA	抑制GA信号和调控植株 Inhibit GA signaling and regulate plant height	［42］
OsMYB91	GA， ABA	平衡水稻生长和非生物胁迫抗性 Balance rice growth and abiotic stress resistance	［43］
RTD1	GA	负调控SLR1蛋白的转录和蛋白水平 Negatively regulate the transcription and protein level of SLR1 protein	［44］
OsNPC6	GA	调控中胚轴伸长 Regulate mesocotyl elongation	［45］
CIPK31	GA	抑制SLR1蛋白降解 Inhibit the degradation of SLR1 protein	［46］
OsGAMYBL2	GA， BR	调控GA合成和BR信号 Regulate GA biosynthesis and BR signaling	［54］
OsJAZ8， OsJAZ9	JA	调控水稻生长发育与抗逆 Regulate rice growth and development as well as stress resistance	［56］
MYC2/3	JA	调控JA响应基因的表达 Regulate the expression of JA-responsive genes	［57］
SP8， P2， M	GA， JA	介导广谱抗病毒防御反应 Mediate broad-spectrum antiviral defense response	［57］
TAD1	ABA	抑制ABA信号 Inhibit ABA signaling	［59］
MOC1	GA	调控株高和分蘖 Regulate plant height and tillering	［60］
NGR5	GA	提高氮素利用、促进分蘖 Enhance nitrogen utilization and promote tillering	［61］
OsMADS23	GA， SL	抑制D14基因转录，促进分蘖 Suppress the transcription of the D14 gene and promote tillering	［62-63］
OsIDD2	GA	调控细胞增殖 Regulate cell proliferation	［64］
OsNAC29/31	GA	调控纤维素合成 Regulate cellulose synthesis	［65］
OsKNAT7	GA	调控次生细胞壁的合成 Regulate the synthesis of secondary cell walls	［66］
OsMYB103L	GA	调控纤维素生物和次生细胞壁合成 Regulate cellulose and secondary cell wall biosynthesis	［67］
OsNAC055	木质素合成途径	调控木质素合成 Regulate lignin biosynthesis	［68］
GAMYB	GA	调控水稻育性 Regulate rice fertility	［69-70］
UDT1， TDR	GA	提高绒毡层发育相关基因的表达，调控水稻的育性 Enhance the expressions of tapetum development-related genes and regulate rice fertility	［71］
OsMS188	GA	调控孢粉素生物合成，促进花粉壁的形成 Regulate sporopollenin biosynthesis and promote pollen wall formation	［72］
OSH1	GA	调控次生细胞壁合成 Regulate secondary cell wall biosynthesis	［73］
RID1	GA	调控营养生长向生殖生长的转变 Regulate the transition from vegetative growth to reproductive growth	［74］
GHD7	GA	调控抽穗期和植株形态 Regulate rice heading date and plant morphology	［75］
qSH1， OSH15， SNB	GA	调控木质素含量和落粒性 Regulate lignin biosynthesis and seed shattering	［76］
OsNAC120	GA， ABA	调控干旱胁迫响应 Regulate the response to drought stress	［77］
OsBURP3， OsSUS1	GA	调控干旱胁迫响应 Regulate the response to drought stress	［78］
OsPIL13/14	GA，光信号	调控黑暗或盐胁迫下的幼苗生长 Regulate seedling growth under darkness or salt stress	［79-80］
OsNF-YA3	GA， ABA	调控生长发育与渗透胁迫耐受性 Regulate growth and development as well as osmotic stress tolerance	［81］
IDD10， bZIP23	GA，氮素信号	调控氮素吸收和盐碱耐受性 Regulate nitrogen absorption and saline-alkali tolerance	［82］
OsGRF6	GA	调控水稻生长和耐寒性 Regulate rice growth and cold tolerance	［83］
OsNuCYP20-2	GA	促进SLR1的降解，促进细胞伸长 Promote the degradation of SLR1 and cell elongation	［84］
Sub1A	GA， ET， ABA	促进SLR1的积累，抑制GA信号 Promote the accumulation of SLR1 and inhibit GA signaling	［85-87］
GRF4	GA， BR	调节根系代谢和氮素利用 Regulate root metabolism and nitrogen utilization	［88］
OsWRKY71	GA	调控逆境下的根系发育 Regulate root development under stress conditions	［89］
OsSPL7， OsSPL14	GA	调控Xoo的抗性 Regulate resistance to Xanthomonasoryzae pv. oryzae （Xoo）	［90-91］
LPA1	GA	对水稻纹枯病rice sheath blight（ShB）的抗性 Resistance to rice sheath blight （ShB）	［92］