Research Progress on the Mechanism and Application of Brassinosteroid in Regulating Plant Height

doi:10.16420/j.issn.0513-353x.2025-0519

Acta Horticulturae Sinica ›› 2026, Vol. 53 ›› Issue (5): 1535-1552.doi: 10.16420/j.issn.0513-353x.2025-0519

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Research Progress on the Mechanism and Application of Brassinosteroid in Regulating Plant Height

LÜ Xinyang¹, GUO Yue¹, FENG Minghan¹, ZHU Pengfang¹^,², FENG Xin¹^,²^,^*()

¹ College of Forestry， Shenyang Agricultural University， Shenyang 110086， China
² Key Laboratory of Forest Tree Genetics， Breeding and Cultivation of Liaoning Province， Shenyang 110086， China

Received:2025-09-22 Revised:2026-03-17 Online:2026-05-26 Published:2026-05-26
Contact: FENG Xin

Abstract

Abstract:

Plant height is a key trait that determines plant architecture，lodging resistance，and yield. Brassinosteroid（BR）plays a crucial role in regulating plant height. This review summarizes recent advances in understanding how BR regulates plant height. At the physiological level，BR modulates cell elongation and division by affecting water uptake，cell wall modification，and microtubule stability. At the molecular level，BR precisely controls plant height through key biosynthesis genes such as DWF4，CPD，DET2，ROT3，CYP90D1，and CYP85A1/2，as well as signal transduction genes including BRI1，BAK1，BSU1，BIN2，BZR1，and BES1. During their functioning，BR crosstalks with gibberellin，auxin，jasmonate，abscisic acid，ethylene，and strigolactone，forming a complex regulatory network for plant height. The application of BR in dwarf breeding is also discussed，BR-related dwarf germplasm demonstrates significant potential in improving plant architecture and increasing yield. Finally，the problems existing in the current research are analyzed and prospects are put forward. The key research directions highlighted in this review will provide a a reference for the in-depth study on the regulatory mechanism of plant height and the practice of dwarf breeding.

Key words: brassinosteroid, plant height, regulatory mechanisms, dwarf breeding

LÜ Xinyang, GUO Yue, FENG Minghan, ZHU Pengfang, FENG Xin. Research Progress on the Mechanism and Application of Brassinosteroid in Regulating Plant Height[J]. Acta Horticulturae Sinica, 2026, 53(5): 1535-1552.

Figures/Tables 3

References 132

[1]	Azpiroz R, Wu Y, LoCascio J C, Feldmann K A. 1998. An Arabidopsis brassinosteroid-dependent mutant is blocked in cell elongation. The Plant Cell, 10 (2):219-230. doi: 10.1105/tpc.10.2.219 URL
[2]	Bao F, Shen J, Brady S R, Muday G K, Asami T, Yang Z. 2004. Brassinosteroids interact with auxin to promote lateral root development in Arabidopsis Plant Physiology, 134 (4):1624-1631. doi: 10.1104/pp.103.036897 URL
[3]	Best N B, Hartwig T, Budka J, Fujioka S, Johal G, Schulz B, Dilkes B P. 2016. nana plant2 encodes a maize ortholog of the Arabidopsis brassinosteroid biosynthesis gene DWARF1,identifying developmental interactions between brassinosteroids and gibberellins. Plant Physiology, 171 (4):2633-2647. doi: 10.1104/pp.16.00399 URL
[4]	Blanco-Touriñán N, Rana S, Nolan T M, Li K, Vukašinović N, Hsu C-W, Russinova E, Hardtke C S. 2024. The brassinosteroid receptor gene BRI1 safeguards cell-autonomous brassinosteroid signaling across tissues. Science Advances, 10 (39):eadq3352.
[5]	Castorina G, Consonni G. 2020. The role of brassinosteroids in controlling plant height in Poaceae:a genetic perspective. International Journal of Molecular Sciences, 21 (4):1191. doi: 10.3390/ijms21041191 URL
[6]	Chai G, Qi G, Wang D, Zhuang Y, Xu H, Bai Z, Bai M-Y, Hu R, Wang Z, Zhou G, Kong Y. 2022. The CCCH zinc finger protein C3H 15 negatively regulates cell elongation by inhibiting brassinosteroid signaling. Plant Physiology, 189 (1):285-300. doi: 10.1093/plphys/kiac046 URL
[7]	Chaudhary A, Hsiao Y C, Yeh F L J, Župunski M, Zhang H, Aizezi Y, Malkovskiy A, Grossmann G, Wu H, Cheung A Y, Xu S, Wang Z. 2025. FERONIA signaling maintains cell wall integrity during brassinosteroid-induced cell expansion in Arabidopsis Molecular Plant, 18 (4):603-618. doi: 10.1016/j.molp.2025.02.001 URL
[8]	Cheon J, Park S Y, Schulz B, Choe S. 2010. Arabidopsis brassinosteroid biosynthetic mutant dwarf7-1 exhibits slower rates of cell division and shoot induction. BMC Plant Biology, 10 (1):270. doi: 10.1186/1471-2229-10-270
[9]	Choe S, Dilkes B P, Fujioka S, Takatsuto S, Sakurai A, Feldmann K A. 1998. The DWF4 gene of Arabidopsis encodes a cytochrome P450 that mediates multiple 22α-hydroxylation steps in brassinosteroid biosynthesis. The Plant Cell, 10 (2):231-243.
[10]	Choe S, Fujioka S, Noguchi T, Takatsuto S, Yoshida S, Feldmann K A. 2001. Overexpression of DWARF4 in the brassinosteroid biosynthetic pathway results in increased vegetative growth and seed yield in Arabidopsis The Plant Journal, 26 (6):573-582. doi: 10.1046/j.1365-313x.2001.01055.x URL
[11]	Chono M, Honda I, Zeniya H, Yoneyama K, Saisho D, Takeda K, Takatsuto S, Hoshino T, Watanabe Y. 2003. A semidwarf phenotype of barley uzu results from a nucleotide substitution in the gene encoding a putative brassinosteroid receptor. Plant Physiology, 133 (3):1209-1219. doi: 10.1104/pp.103.026195 pmid: 14551335
[12]	Chory J, Nagpal P, Peto C A. 1991. Phenotypic and genetic analysis of det2,a new mutant that affects light-regulated seedling development in Arabidopsis The Plant Cell, 3 (5):445-459. doi: 10.1105/tpc.3.5.445 pmid: 12324600
[13]	Chung Y, Maharjan P M, Lee O, Fujioka S, Jang S, Kim B, Takatsuto S, Tsujimoto M, Kim H, Cho S, Park T, Cho H, Hwang I, Choe S. 2011. Auxin stimulates DWARF4 expression and brassinosteroid biosynthesis in Arabidopsis. The Plant Journal, 66 (4):564-578. doi: 10.1111/tpj.2011.66.issue-4 URL
[14]	Clouse S D. 1996. Plant hormones:brassinosteroids in the spotlight. Current Biology, 6 (6):658-661. pmid: 8793287
[15]	Clouse S D, Sasse J M. 1998. Brassinosteroids:essential regulators of plant growth and development. Annual Review of Plant Physiology and Plant Molecular Biology, 49 (1):427-451. doi: 10.1146/arplant.1998.49.issue-1 URL
[16]	Davière J M, Achard P. 2013. Gibberellin signaling in plants. Development, 140 (6):1147-1151. doi: 10.1242/dev.087650 URL
[17]	Delesalle C, Montiel-Jorda A, Aiba R, Spielmann J, Neveu J, Fujita S, Vert G. 2026. Arabidopsis microtubule-BRI1-associated proteins negatively regulate hypocotyl elongation by controlling brassinosteroid-dependent cortical microtubule reorientation. Plant Communications, 7 (2):101637. doi: 10.1016/j.xplc.2025.101637 URL
[18]	Dong L L, Yang C X, Wang J, Li J J, Zhao M, Li D L, Qiu Z Y, Ma C H, Cui Z H. 2024. Insights into the dwarfing mechanism of pear(Pyrus betulaefolia)based on anatomical and structural analysis using X‐ray scanning, Horticultural Plant Journal, 10 (2):355-366. doi: 10.1016/j.hpj.2023.03.013 URL
[19]	Dong W, Wu D, Wang C, Liu Y, Wu D. 2021. Characterization of the molecular mechanism underlying the dwarfism of dsh mutant watermelon plants. Plant Science, 313:111074. doi: 10.1016/j.plantsci.2021.111074 URL
[20]	Fernandez M G S, Becraft P W, Yin Y, Lübberstedt T. 2009. From dwarves to giants? Plant height manipulation for biomass yield. Trends in Plant Science, 14 (8):454-461. doi: 10.1016/j.tplants.2009.06.005 pmid: 19616467
[21]	Foster T M, McAtee P A, Waite C N, Boldingh H L, McGhie T K. 2017. Apple dwarfing rootstocks exhibit an imbalance in carbohydrate allocation and reduced cell growth and metabolism. Horticulture Research, 4:17009. doi: 10.1038/hortres.2017.9 pmid: 28435686
[22]	Fujita S, Ohnishi T, Watanabe B, Yokota T, Takatsuto S, Fujioka S, Yoshida S, Sakata K, Mizutani M. 2006. Arabidopsis CYP90B1 catalyses the early C-22 hydroxylation of C27,C28 and C29 sterols. The Plant Journal, 45 (5):765-774. doi: 10.1111/tpj.2006.45.issue-5 URL
[23]	Gallego-Bartolomé J, Minguet E G, Grau-Enguix F, Abbas M, Locascio A, Thomas S G, Alabadí D, Blázquez M A. 2012. Molecular mechanism for the interaction between gibberellin and brassinosteroid signaling pathways in Arabidopsis. Proceedings of the National Academy of Sciences, 109 (33):13446-13451.
[24]	Goda H, Sawa S, Asami T, Fujioka S, Shimada Y, Yoshida S. 2004. Comprehensive comparison of auxin-regulated and brassinosteroid-regulated genes in Arabidopsis Plant Physiology, 134 (4):1555-1573. doi: 10.1104/pp.103.034736 URL
[25]	Gui J, Zheng S, Liu C, Shen J, Li J, Li L. 2016. OsREM4.1 interacts with OsSERK 1 to coordinate the interlinking between abscisic acid and brassinosteroid signaling in rice. Developmental Cell, 38 (2):201-213. doi: 10.1016/j.devcel.2016.06.011 URL
[26]	He G, Liu J, Dong H, Sun J. 2019. The blue-light receptor CRY1 interacts with BZR1 and BIN2 to modulate the phosphorylation and nuclear function of BZR1 in repressing BR signaling in Arabidopsis. Molecular Plant, 12 (5):689-703. doi: 10.1016/j.molp.2019.02.001 URL
[27]	He J, Gendron J M, Yang Y, Li J, Wang Z. 2002. The GSK3-like kinase BIN2 phosphorylates and destabilizes BZR1,a positive regulator of the brassinosteroid signaling pathway in Arabidopsis. Proceedings of the National Academy of Sciences, 99 (15):10185-10190.
[28]	Hedden P. 2003. The genes of the green revolution. Trends in Genetics, 19 (1):5-9. doi: 10.1016/s0168-9525(02)00009-4 pmid: 12493241
[29]	Hong Z, Ueguchi-Tanaka M, Shimizu-Sato S, Inukai Y, Fujioka S, Shimada Y, Takatsuto S, Agetsuma M, Yoshida S, Watanabe Y, Uozu S, Kitano H, Ashikari M, Matsuoka M. 2002. Loss-of-function of a rice brassinosteroid biosynthetic enzyme,C-6 oxidase,prevents the organized arrangement and polar elongation of cells in the leaves and stem. The Plant Journal, 32 (4):495-508. doi: 10.1046/j.1365-313X.2002.01438.x URL
[30]	Hou Leiping, Li Meilan. 2001. Progress of studies on the plant growth promoting mechanism of brassinolide(BR). Chinese Bulletin of Botany, 18 (5):560-566. (in Chinese).
	侯雷平, 李梅兰. 2001. 油菜素内酯(BR)促进植物生长机理研究进展. 植物学通报, 18 (5):560-566.
[31]	Hou S, Niu H, Tao Q, Wang S, Gong Z, Li S, Weng Y, Li Z. 2017. A mutant in the CsDET2 gene leads to a systemic brassinosteriod deficiency and super compact phenotype in cucumber(Cucumis sativus L.). Theoretical and Applied Genetics, 130 (8):1693-1703. doi: 10.1007/s00122-017-2919-z URL
[32]	Hu Y, Bao F, Li J. 2000. Promotive effect of brassinosteroids on cell division involves a distinct CycD3-induction pathway in Arabidopsis. The Plant Journal, 24 (5):693-701. doi: 10.1046/j.1365-313x.2000.00915.x URL
[33]	Huang S, Zheng C, Zhao Y, Li Q, Liu J, Deng R, Lei T, Wang S, Wang X. 2021. RNA interference knockdown of the brassinosteroid receptor BRI1 in potato(Solanum tuberosum L.) reveals novel functions for brassinosteroid signaling in controlling tuberization. Scientia Horticulturae, 290:110516. doi: 10.1016/j.scienta.2021.110516 URL
[34]	Ibañez C, Delker C, Martinez C, Bürstenbinder K, Janitza P, Lippmann R, Ludwig W, Sun H, James G V, Klecker M, Grossjohann A, Schneeberger K, Prat S, Quint M. 2018. Brassinosteroids dominate hormonal regulation of plant thermomorphogenesis via BZR1. Current Biology, 28 (2):303-310. doi: S0960-9822(17)31602-0 pmid: 29337075
[35]	Jia D, Gong X, Li M, Li C, Sun T, Ma F. 2018. Overexpression of a novel apple NAC transcription factor gene,MdNAC1,confers the dwarf phenotype in transgenic apple(Malus domestica). Genes, 9 (5):229. doi: 10.3390/genes9050229 URL
[36]	Kim E J, Lee S H, Park C H, Kim T W. 2018. Functional role of BSL 1 subcellular localization in brassinosteroid signaling. Journal of Plant Biology, 61 (1):40-49. doi: 10.1007/s12374-017-0363-x
[37]	Kim G, Fujioka S, Kozuka T, Tax F E, Takatsuto S, Yoshida S, Tsukaya H. 2005a. CYP90C1 and CYP90D1 are involved in different steps in the brassinosteroid biosynthesis pathway in Arabidopsis thaliana The Plant Journal, 41 (5):710-721. doi: 10.1111/tpj.2005.41.issue-5 URL
[38]	Kim G T, Tsukaya H, Saito Y, Uchimiya H. 1999. Changes in the shapes of leaves and flowers upon overexpression of cytochrome P450 in Arabidopsis. Proceedings of the National Academy of Sciences, 96 (16):9433-9437.
[39]	Kim T W, Guan S, Burlingame A L, Wang Z Y. 2011. The CDG1 kinase mediates brassinosteroid signal transduction from BRI1 receptor kinase to BSU1 phosphatase and GSK3-like kinase BIN2. Molecular Cell, 43 (4):561-571. doi: 10.1016/j.molcel.2011.05.037 URL
[40]	Kim T W, Guan S, Sun Y, Deng Z, Tang W, Shang J, Sun Y, Burlingame A L, Wang Z. 2009. Brassinosteroid signal transduction from cell-surface receptor kinases to nuclear transcription factors. Nature Cell Biology, 11 (10):1254-1260. doi: 10.1038/ncb1970
[41]	Kim T W, Hwang J Y, Kim Y S, Joo S H, Chang S C, Lee J S, Takatsuto S, Kim S K. 2005b. Arabidopsis CYP85A2,a cytochrome P450,mediates the baeyer-villiger oxidation of castasterone to brassinolide in brassinosteroid biosynthesis. The Plant Cell, 17 (8):2397-2412. doi: 10.1105/tpc.105.033738 URL
[42]	Kir G, Ye H, Nelissen H, Neelakandan A K, Kusnandar A S, Luo A, Inzé D, Sylvester A W, Yin Y, Becraft P W. 2015. RNA interference knockdown of BRASSINOSTEROID INSENSITIVE 1 in maize reveals novel functions for brassinosteroid signaling in controlling plant architecture. Plant Physiology, 169 (1):826-839. doi: 10.1104/pp.15.00367 URL
[43]	Koka C V, Cerny R E, Gardner R G, Noguchi T, Fujioka S, Takatsuto S, Yoshida S, Clouse S D. 2000. A putative role for the tomato genes DUMPY and CURL-3 in brassinosteroid biosynthesis and response. Plant Physiology, 122 (1):85-98. doi: 10.1104/pp.122.1.85 pmid: 10631252
[44]	Lan D, Cao L, Liu M, Ma F, Yan P, Zhang X, Hu J, Niu F, He S, Cui J, Yuan X, Yang J, Wang Y, Luo X. 2023. The identification and characterization of a plant height and grain length related gene hfr131 in rice. Frontiers in Plant Science, 14:1152196. doi: 10.3389/fpls.2023.1152196 URL
[45]	Lee J, Shim D, Moon S, Kim H, Bae W, Kim K, Kim Y H, Rhee S K, Hong C P, Hong S Y, Lee Y J, Sung J, Ryu H. 2018. Genome-wide transcriptomic analysis of BR-deficient Micro-Tom reveals correlations between drought stress tolerance and brassinosteroid signaling in tomato. Plant Physiology and Biochemistry, 127:553-560. doi: S0981-9428(18)30189-X pmid: 29723826
[46]	Li J, Nagpal P, Vitart V, McMorris T C, Chory J. 1996. A role for brassinosteroids in light-dependent development of Arabidopsis. ,Science, 272 (5260):398-401.
[47]	Li J, Nam K H. 2002. Regulation of brassinosteroid signaling by a GSK3/SHAGGY-like kinase. Science, 295 (5558):1299-1301. doi: 10.1126/science.1065769 pmid: 11847343
[48]	Li J, Nam K H, Vafeados D, Chory J. 2001. BIN2,a new brassinosteroid-insensitive locus in Arabidopsis Plant physiology, 127 (1):14-22. doi: 10.1104/pp.127.1.14 pmid: 11553730
[49]	Li J, Wen J, Lease K A, Doke J T, Tax F E, Walker J C. 2002. BAK1,an Arabidopsis LRR receptor-like protein kinase,interacts with BRI1 and modulates brassinosteroid signaling. Cell, 110 (2):213-222. doi: 10.1016/S0092-8674(02)00812-7 URL
[50]	Li L, Yu X, Thompson A, Guo M, Yoshida S, Asami T, Chory J, Yin Y. 2009. Arabidopsis MYB30 is a direct target of BES1 and cooperates with BES1 to regulate brassinosteroid-induced gene expression. The Plant Journal, 58 (2):275-286. doi: 10.1111/tpj.2009.58.issue-2 URL
[51]	Li Q, He J. 2013. Mechanisms of signaling crosstalk between brassinosteroids and gibberellins. Plant Signaling & Behavior, 8 (7):e24686.
[52]	Li Q, Wang C, Jiang L, Li S, Sun S S M, He J. 2012. An interaction between BZR1 and DELLAs mediates direct signaling crosstalk between brassinosteroids and gibberellins in Arabidopsis Science Signaling, 5 (244):ra72.
[53]	Li X, Li J, Zabed H M, Li J, Xiong M, Shi H, Li J. 2025. Manipulating brassinosteroid signaling pathway to genetically improve horticultural plants. aBIOTECH, 6 (2):328-345. doi: 10.1007/s42994-025-00201-y pmid: 40641636
[54]	Li Zehua, Du Juan, He Xuejiao, Zhao Shutang, Liu Yingli, Lu Mengzhu. 2020. Cloning and characterization of brassinosteriod biosynthesis-related gene DET2 in poplar. Forest Research, 33 (2):85-92. (in Chinese)
	李泽华, 杜娟, 贺学娇, 赵树堂, 刘颖丽, 卢孟柱. 2020. 杨树油菜素内酯合成基因DET2的克隆与功能分析. 林业科学研究, 33 (2):85-92.
[55]	Lin Lu, Yu Lu, Xie Peng, Li Zhiqiang, Wang Hongning, Zhao Guoping, Niu Zimian. 2024. Influences of dwarfing intermediate stock SC 1 on apple photosynthetic characteristics and fruit quality. Acta Horticulturae Sinica, 51 (12):2871-2885. (in Chinese)
	林琭, 蔚露, 谢鹏, 李志强, 王红宁, 赵国平, 牛自勉. 2024. 矮化中间砧SC1对苹果光合特性及果实品质的影响. 园艺学报, 51 (12):2871-2885. doi: 10.16420/j.issn.0513-353x.2024-0143
[56]	Liu F, Wang P, Zhang X, Li X, Yan X, Fu D, Wu G. 2018a. The genetic and molecular basis of crop height based on a rice model. Planta, 247 (1):1-26. doi: 10.1007/s00425-017-2798-1 URL
[57]	Liu Gangyun, Duan Shixiang, Xu Nana, Guo Yaomiao, Dou Junling, Yang Sen, Niu Huanhuan, Liu Dongming, Yang Luming, Hu Jianbin, Zhu Huayu. 2024. Molecular markers assisted construction of Cmerecta near-isogenic line of melon dwarfing gene. Acta Horticulturae Sinica, 51 (9):2048-2062. (in Chinese)
	刘港运, 段世享, 许娜娜, 郭姚淼, 豆峻岭, 杨森, 牛欢欢, 刘东明, 杨路明, 胡建斌, 朱华玉. 2024. 分子标记辅助构建甜瓜矮化基因Cmerecta近等基因系. 园艺学报, 51 (9):2048-2062.
[58]	Liu X, Yang Q, Wang Y, Wang L, Fu Y, Wang X. 2018b. Brassinosteroids regulate pavement cell growth by mediating BIN2-induced microtubule stabilization. Journal of Experimental Botany, 69 (5):1037-1049. doi: 10.1093/jxb/erx467 URL
[59]	Ma Y, Ma C, Zhou P, Gao F, Tan W, Huang X, Bai Y, Li M, Wang Z, Hayat F, Shi T, Ni Z, Gao Z. 2024. PmLBD3 links auxin and brassinosteroid signalling pathways on dwarfism in Prunus mume. BMC Biology, 22 (1):184. doi: 10.1186/s12915-024-01985-z
[60]	Ma Y, Xue H, Zhang L, Zhang F, Ou C, Wang F, Zhang Z. 2016. Involvement of auxin and brassinosteroid in dwarfism of autotetraploid apple (Malus × domestica). Scientific Reports, 6 (1):1-14.
[61]	Mazzoni-Putman S M, Brumos J, Zhao C, Alonso J M, Stepanova A N. 2021. Auxin interactions with other hormones in plant development. Cold Spring Harbor Perspectives in Biology, 13 (10):a039990. doi: 10.1101/cshperspect.a039990 URL
[62]	Miao L, Wang X, Yu C, Ye C, Yan Y, Wang H. 2024. What factors control plant height? Journal of Integrative Agriculture, 23 (6):1803-1824. doi: 10.1016/j.jia.2024.03.058 URL
[63]	Miao R, Lin Q, Cao P, Zhou C, Feng M, Lan J, Luo S, Zhang F, Wu H, Hao Q, Zheng H, Ma T, Huang Y, Mou C, Nguyen T, Cheng Z, Guo X, Liu S, Jiang L, Wan J. 2025. SMALL AND ROUND GRAIN is involved in the brassinosteroid signaling pathway which regulates grain size in rice. Journal of Integrative Plant Biology, 67 (5):1290-1306. doi: 10.1111/jipb.v67.5 URL
[64]	Miao R, Wang M, Yuan W, Ren Y, Li Y, Zhang N, Zhang J, Kronzucker H J, Xu W. 2018. Comparative analysis of Arabidopsis ecotypes reveals a role for brassinosteroids in root hydrotropism. Plant Physiology, 176 (4):2720-2736. doi: 10.1104/pp.17.01563 pmid: 29439211
[65]	Minami A, Takahashi K, Inoue S, Tada Y, Kinoshita T. 2019. Brassinosteroid induces phosphorylation of the plasma membrane H⁺-ATPase during hypocotyl elongation in Arabidopsis thaliana. Plant and Cell Physiology, 60 (5):935-944. doi: 10.1093/pcp/pcz005 pmid: 30649552
[66]	Mitchell J W, Mandava N, Worley J F, Plimmer J R, Smith M V. 1970. Brassins—a new family of plant hormones from rape pollen. Nature, 225 (5237):1065-1066. doi: 10.1038/2251065a0
[67]	Miyazawa Y, Nakajima N, Abe T, Sakai A, Fujioka S, Kawano S, Yoshida S. 2003. Activation of cell proliferation by brassinolide application in tobacco BY-2 cells:effects of brassinolide on cell multiplication,cell-cycle-related gene expression,and organellar DNA contents. Journal of Experimental Botany, 54 (393):2669-2678. doi: 10.1093/jxb/erg312 pmid: 14623940
[68]	Morillon R, Catterou M, Sangwan R S, Sangwan B S, Lassalles J P. 2001. Brassinolide may control aquaporin activities in Arabidopsis thaliana. Planta, 212 (2):199-204. pmid: 11216840
[69]	Nam K H, Li J. 2002. BRI1/BAK1,a receptor kinase pair mediating brassinosteroid signaling. Cell, 110 (2):203-212. doi: 10.1016/S0092-8674(02)00814-0 URL
[70]	Nakajima N, Shida A, Toyama S. 1996. Effects of brassinosteroid on cell division and colony formation of Chinese cabbage mesophyll protoplasts. Japanese Journal of Crop Science, 65 (1):114-118. doi: 10.1626/jcs.65.114 URL
[71]	Nakamura A, Goda H, Shimada Y, Yoshida S. 2004. Brassinosteroid selectively regulates PIN gene expression in Arabidopsis. Bioscience Biotechnology and Biochemistry, 68 (4):952-954. pmid: 15118332
[72]	Nakaya M, Tsukaya H, Murakami N, Kato M. 2002. Brassinosteroids control the proliferation of leaf cells of Arabidopsis thaliana. Plant and Cell Physiology, 43 (2):239-244. doi: 10.1093/pcp/pcf024 URL
[73]	Nemhauser J L, Hong F, Chory J. 2006. Different plant hormones regulate similar processes through largely nonoverlapping transcriptional responses. Cell, 126 (3):467-475. doi: 10.1016/j.cell.2006.05.050 pmid: 16901781
[74]	Nemhauser J L, Mockler T C, Chory J. 2004. Interdependency of brassinosteroid and auxin signaling in Arabidopsis PLoS Biology, 2 (9):e258. doi: 10.1371/journal.pbio.0020258 pmid: 15328536
[75]	Nishitani K, Tominaga R. 1992. Endo-xyloglucan transferase,a novel class of glycosyltransferase that catalyzes transfer of a segment of xyloglucan molecule to another xyloglucan molecule. Journal of Biological Chemistry, 267 (29):21058-21064. pmid: 1400418
[76]	Noguchi T, Fujioka S, Takatsuto S, Sakurai A, Yoshida S, Li J, Chory J. 1999. Arabidopsis det2 is defective in the conversion of (24R)-24- methylcholest-4-en-3-one to (24R)-24-methyl-5α-cholestan-3-one in brassinosteroid biosynthesis1. Plant Physiology, 120 (3):833-840. doi: 10.1104/pp.120.3.833 pmid: 10398719
[77]	Nolan T M, Vukašinović N, Liu D, Russinova E, Yin Y. 2020. Brassinosteroids:multidimensional regulators of plant growth,development,and stress responses. The Plant Cell, 32 (2):295-318. doi: 10.1105/tpc.19.00335 URL
[78]	Nomura T, Jager C E, Kitasaka Y, Takeuchi K, Fukami M, Yoneyama K, Matsushita Y, Nyunoya H, Takatsuto S, Fujioka S, Smith J J, Kerckhoffs L H J, Reid J B, Yokota T. 2004. Brassinosteroid deficiency due to truncated steroid 5α-reductase causes dwarfism in the lk mutant of pea. Plant Physiology, 135 (4):2220-2229. doi: 10.1104/pp.104.043786 URL
[79]	Nomura T, Kushiro T, Yokota T, Kamiya Y, Bishop G J, Yamaguchi S. 2005. The last reaction producing brassinolide is catalyzed by cytochrome P-450s,CYP85A3 in tomato and CYP85A2 in Arabidopsis. Journal of Biological Chemistry, 280 (18):17873-17879. doi: 10.1074/jbc.M414592200 URL
[80]	Oh E, Zhu J, Bai M, Arenhart R A, Sun Y, Wang Z. 2014. Cell elongation is regulated through a central circuit of interacting transcription factors in the Arabidopsis hypocotyl. eLife, 3:e03031. doi: 10.7554/eLife.03031 URL
[81]	Oh M H, Clouse S D. 1998. Brassinolide affects the rate of cell division in isolated leaf protoplasts of Petunia hybrida. Plant Cell Report, 17 (12):921-924.
[82]	Oh M H, Honey S H, Tax F E. 2020. The control of cell expansion,cell division,and vascular development by brassinosteroids:a historical perspective. International Journal of Molecular Sciences, 21 (5):1743-1743. doi: 10.3390/ijms21051743 URL
[83]	Ohnishi T, Godza B, Watanabe B, Fujioka S, Hategan L, Ide K, Shibata K, Yokota T, Szekeres M, Mizutani M. 2012. CYP90A1/CPD,a brassinosteroid biosynthetic cytochrome P 450 of Arabidopsis,catalyzes C-3 oxidation. Journal of Biological Chemistry, 287 (37):31551-31560. doi: 10.1074/jbc.M112.392720 URL
[84]	Ohnishi T, Szatmari A-M, Watanabe B, Fujita S, Bancos S, Koncz C, Lafos M, Shibata K, Yokota T, Sakata K, Szekeres M, Mizutani M. 2006. C-23 hydroxylation by Arabidopsis CYP90C1 and CYP90D 1 reveals a novel shortcut in brassinosteroid biosynthesis. The Plant Cell, 18 (11):3275-3288. doi: 10.1105/tpc.106.045443 URL
[85]	Peng J, Richards D E, Hartley N M, Murphy G P, Devos K M, Flintham J E, Beales J, Fish L J, Worland A J, Pelica F, Sudhakar D, Christou P, Snape J W, Gale M D, Harberd N P. 1999. ‘Green revolution’genes encode mutant gibberellin response modulators. Nature, 400 (6741):256-261. doi: 10.1038/22307
[86]	Perrot-Rechenmann C. 2010. Cellular responses to auxin:division versus expansion. Cold Spring Harbor Perspectives in Biology, 2 (5):a001446.
[87]	Qiao S, Sun S, Wang L, Wu Z, Li C, Li X, Wang T, Leng L, Tian W, Lu T, Wang X. 2017. The RLA1/SMOS 1 transcription factor functions with OsBZR1 to regulate brassinosteroid signaling and rice architecture. The Plant Cell, 29 (2):292-309. doi: 10.1105/tpc.16.00611 URL
[88]	Ryu H, Kim K, Cho H, Hwang I. 2010. Predominant actions of cytosolic BSU1 and nuclear BIN2 regulate subcellular localization of BES1 in brassinosteroid signaling. Molecules and Cells, 29 (3):291-296. doi: 10.1007/s10059-010-0034-y pmid: 20387035
[89]	Sadura I, Libik-Konieczny M, Jurczyk B, Gruszka D, Janeczko A. 2020. Plasma membrane ATPase and the aquaporin HvPIP1 in barley brassinosteroid mutants acclimated to high and low temperature. Journal of Plant Physiology, 244:153090. doi: 10.1016/j.jplph.2019.153090 URL
[90]	Sakamoto T, Morinaka Y, Inukai Y, Kitano H, Fujioka S. 2013. Auxin signal transcription factor regulates expression of the brassinosteroid receptor gene in rice. The Plant Journal, 73 (4):676-688. doi: 10.1111/tpj.12071 pmid: 23146214
[91]	Sakamoto T, Morinaka Y, Ohnishi T, Sunohara H, Fujioka S, Ueguchi-Tanaka M, Mizutani M, Sakata K, Takatsuto S, Yoshida S, Tanaka H, Kitano H, Matsuoka M. 2006. Erect leaves caused by brassinosteroid deficiency increase biomass production and grain yield in rice. Nature Biotechnology, 24 (1):105-109. doi: 10.1038/nbt1173 pmid: 16369540
[92]	Shimada Y, Fujioka S, Miyauchi N, Kushiro M, Takatsuto S, Nomura T, Yokota T, Kamiya Y, Bishop G J, Yoshida S. 2001. Brassinosteroid- 6-oxidases from Arabidopsis and tomato catalyze multiple C-6 oxidations in brassinosteroid biosynthesis. Plant Physiology, 126 (2):770-779. doi: 10.1104/pp.126.2.770 pmid: 11402205
[93]	Song L, Liu J, Cao B, Liu B, Zhang X, Chen Z, Dong C, Liu X, Zhang Z, Wang W, Chai L, Liu J, Zhu J, Cui S, He F, Peng H, Hu Z, Su Z, Guo W, Xin M, Yao Y, Yan Y, Song Y, Bai G, Sun Q, Ni Z. 2023. Reducing brassinosteroid signalling enhances grain yield in semi-dwarf wheat. Nature, 617 (7959):118-124. doi: 10.1038/s41586-023-06023-6
[94]	Song S, Liu G, Ma F, Bao Z. 2022. Brassinazole represses tomato hypocotyl elongation via inhibition of cell division. Plant Growth Regulation, 96 (3):463-472. doi: 10.1007/s10725-022-00798-w
[95]	Sun C, Liu Y, Li G, Chen Y, Li M, Yang R, Qin Y, Chen Y, Cheng J, Tang J, Fu Z. 2024a. ZmCYP90D1 regulates maize internode development by modulating brassinosteroid-mediated cell division and growth. The Crop Journal, 12 (1):58-67. doi: 10.1016/j.cj.2023.11.002 URL
[96]	Sun P, Zhao H, Cao L, Zhang T, Zhang H, Yang T, Zhao B, Jiang Y, Dong J, Chen T, Jiang B, Li Z, Shen J. 2024b. A DUF 21 domain-containing protein regulates plant dwarfing in watermelon. Plant Physiology, 196 (4):3091-3104. doi: 10.1093/plphys/kiae486 URL
[97]	Sun S, Wang T, Wang L, Li X, Jia Y, Liu C, Huang X, Xie W, Wang X. 2018. Natural selection of a GSK3 determines rice mesocotyl domestication by coordinating strigolactone and brassinosteroid signaling. Nature Communications, 9 (1):2523. doi: 10.1038/s41467-018-04952-9
[98]	Szekeres M, Németh K, Koncz-Kálmán Z, Mathur J, Kauschmann A, Altmann T, Rédei GP, Nagy F, Schell J, Koncz C. 1996. Brassinosteroids rescue the deficiency of CYP90,a cytochrome P450,controlling cell elongation and de-etiolation in Arabidopsis. Cell, 85 (2):171-182. doi: 10.1016/s0092-8674(00)81094-6 pmid: 8612270
[99]	Tang Y, Liu H, Guo S, Wang B, Li Z, Chong K, Xu Y. 2018. OsmiR396d affects gibberellin and brassinosteroid signaling to regulate plant architecture in rice. Plant Physiology, 176 (1):946-959. doi: 10.1104/pp.17.00964 pmid: 29180380
[100]	Tang Z, Sun M, Li J, Song B, Liu Y, Tian Y, Wang C, Wu J. 2022. Comparative transcriptome analysis provides insights into the mechanism of pear dwarfing. Journal of Integrative Agriculture, 21 (7):1952-1967. doi: 10.1016/S2095-3119(21)63774-7
[101]	Tian H, Lv B, Ding T, Bai M, Ding Z. 2018. Auxin-BR interaction regulates plant growth and development. Frontiers in Plant Science, 8:2256. doi: 10.3389/fpls.2017.02256 URL
[102]	Tong H, Xiao Y, Liu D, Gao S, Liu L, Yin Y, Jin Y, Qian Q, Chu C. 2014. Brassinosteroid regulates cell elongation by modulating gibberellin metabolism in rice. The Plant Cell, 26 (11):4376-4393. doi: 10.1105/tpc.114.132092 pmid: 25371548
[103]	Unterholzner S J, Rozhon W, Papacek M, Ciomas J, Lange T, Kugler K G, Mayer K F, Sieberer T, Poppenberger B. 2015. Brassinosteroids are master regulators of gibberellin biosynthesis in Arabidopsis. The Plant Cell, 27(8):2261-2272. doi: 10.1105/tpc.15.00433 pmid: 26243314
[104]	Vert G, Chory J. 2006. Downstream nuclear events in brassinosteroid signalling. Nature, 441 (7089):96-100. doi: 10.1038/nature04681
[105]	Vert G, Walcher C L, Chory J, Nemhauser J L. 2008. Integration of auxin and brassinosteroid pathways by auxin response factor 2. Proceedings of the National Academy of Sciences, 105 (28):9829-9834.
[106]	Wang B, Wang Z, Tang Y, Zhong N, Wu J. 2024a. Cotton BOP1 mediates SUMOylation of GhBES 1 to regulate fibre development and plant architecture. Plant Biotechnology Journal, 22 (11):3054-3067. doi: 10.1111/pbi.v22.11 URL
[107]	Wang Donglei, Wang Zhiquan, Li Qing, Ouyang Suying, Yang Bozhi, Zhang Zhishuo, Liu Feng, Hu Bowen, Zou Xuexiao. 2021. Research progress of brassinosteroid-related genes regulating plant dwarfing. Journal of Plant Genetic Resources, 22 (4):921-929. (in Chinese). doi: 10.13430/j.cnki.jpgr.20200828002
	王东磊, 王志权, 李青, 欧阳素影, 杨博智, 张志硕, 刘峰, 胡博文, 邹学校. 2021. 油菜素类固醇相关基因调控植株矮化研究进展. 植物遗传资源学报, 22 (4):921-929. doi: 10.13430/j.cnki.jpgr.20200828002
[108]	Wang H, Tang J, Liu J, Hu J, Liu J, Chen Y, Cai Z, Wang X. 2018. Abscisic acid signaling inhibits brassinosteroid signaling through dampening the dephosphorylation of BIN2 by ABI1 and ABI2. Molecular Plant, 11 (2):315-325. doi: S1674-2052(17)30383-0 pmid: 29275167
[109]	Wang H, Li W, Qin Y, Pan Y, Wang X, Weng Y, Chen P, Li Y. 2017. The cytochrome P450 gene CsCYP85A1 is a putative candidate for Super Compact-1(Scp-1)plant architecture mutation in cucumber(Cucumis sativus L.). Frontiers in Plant Science,8:266.
[110]	Wang S, Wang K, Li Z, Li Y, He J, Li H, Wang B, Xin T, Tian H, Tian J, Zhang G, Li H, Huang S, Yang X. 2022. Architecture design of cucurbit crops for enhanced productivity by a natural allele. Nature Plants, 8 (12):1394-1407. doi: 10.1038/s41477-022-01297-6 pmid: 36509843
[111]	Wang T, Cosgrove D J, Arteca R N. 1993. Brassinosteroid stimulation of hypocotyl elongation and wall relaxation in pakchoi(Brassica chinensis cv. Lei-Choi). Plant Physiology, 101 (3):965-968.
[112]	Wang X, Ren Z, Xie S, Li Z, Zhou Y, Duan L. 2024b. Jasmonate mimic modulates cell elongation by regulating antagonistic bHLH transcription factors via brassinosteroid signaling. Plant Physiology, 195 (4):2712-2726. doi: 10.1093/plphys/kiae217 URL
[113]	Wang Y, Li J. 2008. Molecular basis of plant architecture. Annual Review of Plant Biology, 59 (1):253-279. doi: 10.1146/arplant.2008.59.issue-1 URL
[114]	Wang Z, Nakano T, Gendron J, He J, Chen M, Vafeados D, Yang Y, Fujioka S, Yoshida S, Asami T, Chory J. 2002. Nuclear-localized BZR1 mediates brassinosteroid-induced growth and feedback suppression of brassinosteroid biosynthesis. Developmental Cell, 2 (4):505-513. doi: 10.1016/s1534-5807(02)00153-3 pmid: 11970900
[115]	Wang Z, Seto H, Fujioka S, Yoshida S, Chory J. 2001. BRI 1 is a critical component of a plasma-membrane receptor for plant steroids. Nature, 410 (6826):380-383. doi: 10.1038/35066597 URL
[116]	Wei Z, Li J. 2020. Regulation of brassinosteroid homeostasis in higher plants. Frontiers in Plant Science, 11:583622. doi: 10.3389/fpls.2020.583622 URL
[117]	Yang Y, Chu C, Qian Q, Tong H. 2024. Leveraging brassinosteroids towards the next green revolution. Trends in Plant Science, 29 (1):86-98. doi: 10.1016/j.tplants.2023.09.005 URL
[118]	Yin W, Dong N, Li X, Yang Y, Lu Z, Zhou W, Qian Q, Chu C, Tong H. 2025. Understanding brassinosteroid-centric phytohormone interactions for crop improvement. Journal of Integrative Plant Biology, 67 (3):563-581. doi: 10.1111/jipb.13849
[119]	Yin Y, Vafeados D, Tao Y, Yoshida S, Asami T, Chory J. 2005. A new class of transcription factors mediates brassinosteroid-regulated gene expression in Arabidopsis. Cell, 120 (2):249-259. doi: 10.1016/j.cell.2004.11.044 URL
[120]	Yin Y, Wang Z Y, Mora-Garcia S, Li J, Yoshida S, Asami T, Chory J. 2002. BES 1 accumulates in the nucleus in response to brassinosteroids to regulate gene expression and promote stem elongation. Cell, 109 (2):181-191. doi: 10.1016/S0092-8674(02)00721-3 URL
[121]	Yu Z, Ma J, Zhang M, Li X, Sun Y, Zhang M, Ding Z. 2023. Auxin promotes hypocotyl elongation by enhancing BZR1 nuclear accumulation in Arabidopsis. Science Advances, 9 (1):eade2493.
[122]	Zhan H, Lu M, Luo Q, Tan F, Zhao Z, Liu M, He Y. 2022. OsCPD1 and OsCPD2 are functional brassinosteroid biosynthesis genes in rice. Plant Science, 325:111482. doi: 10.1016/j.plantsci.2022.111482 URL
[123]	Zhang Haili, Gao Jing, Zhang Hao, Li Shenghui, Xing Jihong, Wang Fengru, Dong Jingao. 2015. The regulation of brassinosteroid(BR)on elongation and division of rice(Oryza sativa)cells. Journal of Agricultural Biotechnology, 23 (1):71-79. (in Chinese).
	张海丽, 高静, 张昊, 李生辉, 邢继红, 王凤茹, 董金皋. 2015. 油菜素内酯对水稻细胞伸长和分裂的调控. 农业生物技术学报, 23 (1):71-79.
[124]	Zhang S, Deng R, Liu J, Luo D, Hu M, Huang S, Jiang M, Du J, Jin T, Liu D, Li Y, Khan M, Wang S, Wang X. 2024. Phosphorylation of the transcription factor SlBIML 1 by SlBIN2 kinases delays flowering in tomato. Plant Physiology, 196 (4):2583-2598. doi: 10.1093/plphys/kiae489 URL
[125]	Zhang T, Xu P, Wang W, Wang S, Caruana J C, Yang H-Q, Lian H. 2018. Arabidopsis G-protein β subunit AGB1 interacts with BES1 to regulate brassinosteroid signaling and cell elongation. Frontiers in Plant Science,8:2225.
[126]	Zhao N, Zhao M, Tian Y, Wang Y, Han C, Fan M, Guo H, Bai M. 2021. Interaction between BZR1 and EIN3 mediates signalling crosstalk between brassinosteroids and ethylene. New Phytologist, 232 (6):2308-2323. doi: 10.1111/nph.17694 pmid: 34449890
[127]	Zheng X, Li Y, Ma C, Chen B, Sun Z, Tian Y, Wang C. 2022. A mutation in the promoter of the arabinogalactan protein 7-like gene PcAGP7-1 affects cell morphogenesis and brassinolide content in pear(Pyrus communis L.)stems. The Plant Journal, 109 (1):47-63. doi: 10.1111/tpj.v109.1 URL
[128]	Zhiponova M K, Vanhoutte I, Boudolf V, Betti C, Dhondt S, Coppens F, Mylle E, Maes S, González-García M, Caño-Delgado A I, Inzé D, Beemster G T S, De Veylder L, Russinova E. 2013. Brassinosteroid production and signaling differentially control cell division and expansion in the leaf. New Phytologist, 197 (2):490-502. doi: 10.1111/nph.12036 pmid: 23253334
[129]	Zhou X, Song L, Xue H. 2013. Brassinosteroids regulate the differential growth of Arabidopsis hypocotyls through auxin signaling components IAA19 and ARF7. Molecular Plant, 6 (3):887-904. doi: 10.1093/mp/sss123 URL
[130]	Zhu Q, Wang M, Luo H, Zhang L, Lu H, Liu J. 2025. The HECT-family protein UPL 3 suppresses thermomorphogenesis by modulating BZR1 accumulation. The Plant Journal, 122 (3):e70204. doi: 10.1111/tpj.v122.3 URL
[131]	Zhu Z, Liang H, Chen G, Tang B, Tian S, Hu Z. 2019. Isolation of the brassinosteroid receptor genes and recharacterization of dwarf plants by silencing of SlBRI1 in tomato. Plant Growth Regulation, 89 (1):59-71. doi: 10.1007/s10725-019-00524-z
[132]	Zurek DM, Clouse S D. 1994. Molecular cloning and characterization of a brassinosteroid-regulated gene from elongating soybean(Glycine max L.) epicotyls. Plant Physiol, 104 (1):161-170.

通路 Pathway	拟南芥 Arabidopsis thaliana		水稻 Oryza sativa		玉米 Zea mays		番茄 Solanum lycopersicum
通路 Pathway	基因名 Gene name	基因ID Gene ID	基因名 Gene name	基因ID Gene ID	基因名 Gene name	基因ID Gene ID	基因名 Gene name	基因ID Gene ID
BR合成 BR biosynthesis	AtDWF4	AT3G50660	OsDWARF4	Os03g0227700	ZmDWF4	GRMZM2G065635	SlCYP90B3	Solyc02g085360
	AtDWF4	AT3G50660	OsD11	Os04g0469800	ZmDWF4	GRMZM2G065635	SlCYP90B3	Solyc02g085360
	AtCPD	AT5G05690	OsCPD1	Os11g0143200			SlCPD	Solyc06g051750
	AtCPD	AT5G05690	OsCPD2	Os12g0139300			SlCPD	Solyc06g051750
	AtDET2	AT2G38050	OsDET2	Os01g0851600	ZmDET2	GRMZM2G449033	SlDET2	Solyc10g086500
	AtDET2	AT2G38050	OsDET2	Os11g0184100	ZmDET2	GRMZM2G449033	SlDET2	Solyc10g086500
	AtROT3	AT4G36380					SlROT3	Solyc02g084740
	AtCYP90D1	AT3G13730					SlCYP90D1	Solyc03g121510
	AtCYP85A1	AT5G38970	OsCYP85A1	Os03g0602300	ZmBRD1	GRMZM2G103773	SlDWARF	Solyc02g089160
	AtCYP85A2	AT3G30180
BR信号转导 BR signaling	AtBRI1	AT4G39400	OsBRI1	Os01g0718300	ZmBRI1a	GRMZM2G048294	SlBRI1	Solyc04g051510
	AtBRI1	AT4G39400	OsBRI1	Os01g0718300	ZmBRI1b	GRMZM2G449830	SlBRI1	Solyc04g051510
	AtBAK1	AT4G33430	OsBAK1	Os08g0174700	ZmBAK1	GRMZM2G145720	SlBAK1	Solyc10g047140
	AtBAK1	AT4G33430	OsBAK1	Os08g0174700	ZmBAK1	GRMZM2G145720	SlBAK1	Solyc01g104970
	AtBSU1	AT1G03445	OsPPKL1	Os03g0646900			SlBSU1	Solyc09g074320
	AtBSL1	AT4G03080	OsPPKL2	Os05g0144400
	AtBSL2	AT1G08420	OsPPKL1	Os03g0646900
	AtBSL3	AT2G27210	OsPPKL3	Os12g0617900
BR信号转导 BR signaling	AtBIN2	AT4G18710	OsGSK1	Os01g0205700	ZmGSK2	GRMZM2G472625	SlBIN2.1	Solyc02g072300
			OsGSK2	Os05g0207500			SlBIN2.2	Solyc07g055200
			OsGSK2	Os05g0207500			SlBIN2.3	Solyc03g006070
	AtBZR1	AT1G75080	OsBZR1	Os07g0580500	ZmBZR1	GRMZM5G852801	SlBZR1	Solyc12g089040
	AtBES1	AT1G19350	OsBZR2	Os01g0203000	ZmBES1	GRMZM2G102514	SlBES1	Solyc04g079980

通路 Pathway	拟南芥 Arabidopsis thaliana		水稻 Oryza sativa		玉米 Zea mays		番茄 Solanum lycopersicum
通路 Pathway	基因名 Gene name	基因ID Gene ID	基因名 Gene name	基因ID Gene ID	基因名 Gene name	基因ID Gene ID	基因名 Gene name	基因ID Gene ID
BR合成 BR biosynthesis	AtDWF4	AT3G50660	OsDWARF4	Os03g0227700	ZmDWF4	GRMZM2G065635	SlCYP90B3	Solyc02g085360
	AtDWF4	AT3G50660	OsD11	Os04g0469800	ZmDWF4	GRMZM2G065635	SlCYP90B3	Solyc02g085360
	AtCPD	AT5G05690	OsCPD1	Os11g0143200			SlCPD	Solyc06g051750
	AtCPD	AT5G05690	OsCPD2	Os12g0139300			SlCPD	Solyc06g051750
	AtDET2	AT2G38050	OsDET2	Os01g0851600	ZmDET2	GRMZM2G449033	SlDET2	Solyc10g086500
	AtDET2	AT2G38050	OsDET2	Os11g0184100	ZmDET2	GRMZM2G449033	SlDET2	Solyc10g086500
	AtROT3	AT4G36380					SlROT3	Solyc02g084740
	AtCYP90D1	AT3G13730					SlCYP90D1	Solyc03g121510
	AtCYP85A1	AT5G38970	OsCYP85A1	Os03g0602300	ZmBRD1	GRMZM2G103773	SlDWARF	Solyc02g089160
	AtCYP85A2	AT3G30180
BR信号转导 BR signaling	AtBRI1	AT4G39400	OsBRI1	Os01g0718300	ZmBRI1a	GRMZM2G048294	SlBRI1	Solyc04g051510
	AtBRI1	AT4G39400	OsBRI1	Os01g0718300	ZmBRI1b	GRMZM2G449830	SlBRI1	Solyc04g051510
	AtBAK1	AT4G33430	OsBAK1	Os08g0174700	ZmBAK1	GRMZM2G145720	SlBAK1	Solyc10g047140
	AtBAK1	AT4G33430	OsBAK1	Os08g0174700	ZmBAK1	GRMZM2G145720	SlBAK1	Solyc01g104970
	AtBSU1	AT1G03445	OsPPKL1	Os03g0646900			SlBSU1	Solyc09g074320
	AtBSL1	AT4G03080	OsPPKL2	Os05g0144400
	AtBSL2	AT1G08420	OsPPKL1	Os03g0646900
	AtBSL3	AT2G27210	OsPPKL3	Os12g0617900
BR信号转导 BR signaling	AtBIN2	AT4G18710	OsGSK1	Os01g0205700	ZmGSK2	GRMZM2G472625	SlBIN2.1	Solyc02g072300
			OsGSK2	Os05g0207500			SlBIN2.2	Solyc07g055200
			OsGSK2	Os05g0207500			SlBIN2.3	Solyc03g006070
	AtBZR1	AT1G75080	OsBZR1	Os07g0580500	ZmBZR1	GRMZM5G852801	SlBZR1	Solyc12g089040
	AtBES1	AT1G19350	OsBZR2	Os01g0203000	ZmBES1	GRMZM2G102514	SlBES1	Solyc04g079980

通路 Pathway	基因名称 Gene name	功能丧失型突变体 Loss-of- function mutant	功能丧失型突变体表型 Phenotype of the loss-of- function mutant	功能获得型突变体/过表达植株 Gain-of-function mutant/ Overexpression plant	功能获得型突变体/过表达植株表型 Phenotype of the gain-of-function mutant or overexpression plant	参考文献 Reference
BR合成 BR biosynthesis	AtDWF4	dwf4	矮化，深绿色，叶片小而圆，叶柄短，莲座叶数量增加，顶端优势减弱，雄性不育，开花延迟 Dwarf，dark green，small and rounded leaves，short petioles，increased number of rosette leaves，reduced apical dominance，male sterility，delayed flowering	AOD4	株高增加，分枝和角果数量增多，叶片和叶柄伸长 Increased plant height，increased branching and silique number，elongated leaves and petioles	Azpiroz et al.,1998； Choe et al.,2001
	AtCPD	cpd	矮化，叶片小而圆，花序茎和花器官缩短，雄性不育 Dwarf，small and rounded leaves，shortened inflorescence stems，and flower organs，male sterility	cpd comp.	将CPD cDNA在cpd 突变体中表达，突变体恢复了野生型的表型，甚至一部分比野生型叶片更大、分枝更多 Overexpression of CPD cDNA in the cpd mutant restored the wild-type phenotype；some lines exhibited even larger leaves and more branching than wild-type	Szekeres et al.,1996； Zhiponova et al.,2013
	AtDET2	det2	矮化，深绿色，叶片小，顶端优势减弱，雄性育性降低，开花延迟 Dwarf，dark green，small leaves，reduced apical dominance，reduced male fertility，delayed flowering			Chory et al.,1991； Li et al.,1996
	AtROT3	rot3-4 cyp90d1	严重矮化，下胚轴伸长缺陷 Severe dwarfism，defective hypocotyl elongation	A-1（ROT3 OE）	叶片和花器官伸长 Elongated leaves and flower organs	Kim et al.,1999； Kim et al.,2005a
	AtCYP90D1	rot3-4 cyp90d1	严重矮化，下胚轴伸长缺陷 Severe dwarfism，defective hypocotyl elongation			Kim et al.,1999； Kim et al.,2005a
	AtCYP85A1	cyp85a1-1/cyp85a2-2	严重矮化 Severe dwarfism	35S-CYP85A1	表型无变化 Do not cause any phenotypic changes	Kim et al.,2005b； Nomura et al.,2005
	AtCYP85A2	cyp85a1-1/cyp85a2-2	严重矮化 Severe dwarfism	35S-CYP85A2	莲座叶更大，叶柄更长，茎更高且多分枝，角果更大，花梗更长 Larger rosette leaves，longer petioles，taller stems with increased branching，larger siliques，longer pedicels	Kim et al.,2005b； Nomura et al.,2005
BR信号转导 BR signaling	AtBRI1	bri1	严重矮化，深绿色，叶片厚，顶端优势减弱，雄性不育 Severe dwarfism，dark green，thickened leaves，reduced apical dominance，male sterility	BRI1-GFP	叶柄伸长 Elongated petioles	Clouse,1996； Wang et al.,2001； Blanco- Touriñán et al.,2024
	AtBAK1	bak1-1	半矮化，莲座叶小而圆，叶柄短 Semi-dwarf，small and rounded rosette leaves，short petioles	bak1-1D	茎、叶和叶柄伸长 Elongated stems，leaves，and petioles	Li et al.,2002
	AtBSU1	bsu1;bsl1/BSL2,3-amiRNA	严重矮化 Severe dwarfism	BSU1-YFP/bri1-116	在bri1-116中过表达BSU1，部分挽救了其矮化表型 Overexpression of BSU1 in bri1-116 partially rescued the dwarf phenotype	Kim et al.,2009； Kim et al.,2018
	AtBSL1			BSL1-YFP/bri1-5	部分挽救了其矮化表型 Overexpression of BSL1 in bri1-5 partially rescued the dwarf phenotype
	AtBSL2			BSL2-YFP	叶柄伸长 Elongated petioles
	AtBSL3
	AtBIN2	Triple GSK3 mutant	下胚轴显著伸长 Significantly elongated hypocoty	bin2-1	矮化，深绿色，叶片卷曲，顶端优势减弱，雄性育性降低，开花延迟 Dwarf，dark green，curled leaves，reduced apical dominance，reduced male fertility，delayed flowering	Li et al.,2001； Vert & Chory,2006
	AtBZR1	BES1 RNAi	半矮化 Semi-dwarf	bzr1-1D	在黑暗中，下胚轴伸长；在光下，半矮化，呈深绿色，叶片宽，叶柄短，叶片数量增加，开花延迟 In darkness：elongated hypocotyl；In light：semi- dwarf，dark green，wide leaves，short petioles，increased leaf number，delayed flowering	Wang et al.,2002； Yin et al.,2005
	AtBES1	BES1 RNAi	半矮化 Semi-dwarf	bes1-D	完全挽救bri1的矮化表型，叶柄长而弯曲，叶片卷曲，过度的茎伸长 Fully rescued the dwarf phenotype of bri1，long and curved petioles，curled leaves，excessive stem elongation	Yin et al.,2002； Yin et al.,2005

通路 Pathway	基因名称 Gene name	功能丧失型突变体 Loss-of- function mutant	功能丧失型突变体表型 Phenotype of the loss-of- function mutant	功能获得型突变体/过表达植株 Gain-of-function mutant/ Overexpression plant	功能获得型突变体/过表达植株表型 Phenotype of the gain-of-function mutant or overexpression plant	参考文献 Reference
BR合成 BR biosynthesis	AtDWF4	dwf4	矮化，深绿色，叶片小而圆，叶柄短，莲座叶数量增加，顶端优势减弱，雄性不育，开花延迟 Dwarf，dark green，small and rounded leaves，short petioles，increased number of rosette leaves，reduced apical dominance，male sterility，delayed flowering	AOD4	株高增加，分枝和角果数量增多，叶片和叶柄伸长 Increased plant height，increased branching and silique number，elongated leaves and petioles	Azpiroz et al.,1998； Choe et al.,2001
	AtCPD	cpd	矮化，叶片小而圆，花序茎和花器官缩短，雄性不育 Dwarf，small and rounded leaves，shortened inflorescence stems，and flower organs，male sterility	cpd comp.	将CPD cDNA在cpd 突变体中表达，突变体恢复了野生型的表型，甚至一部分比野生型叶片更大、分枝更多 Overexpression of CPD cDNA in the cpd mutant restored the wild-type phenotype；some lines exhibited even larger leaves and more branching than wild-type	Szekeres et al.,1996； Zhiponova et al.,2013
	AtDET2	det2	矮化，深绿色，叶片小，顶端优势减弱，雄性育性降低，开花延迟 Dwarf，dark green，small leaves，reduced apical dominance，reduced male fertility，delayed flowering			Chory et al.,1991； Li et al.,1996
	AtROT3	rot3-4 cyp90d1	严重矮化，下胚轴伸长缺陷 Severe dwarfism，defective hypocotyl elongation	A-1（ROT3 OE）	叶片和花器官伸长 Elongated leaves and flower organs	Kim et al.,1999； Kim et al.,2005a
	AtCYP90D1	rot3-4 cyp90d1	严重矮化，下胚轴伸长缺陷 Severe dwarfism，defective hypocotyl elongation			Kim et al.,1999； Kim et al.,2005a
	AtCYP85A1	cyp85a1-1/cyp85a2-2	严重矮化 Severe dwarfism	35S-CYP85A1	表型无变化 Do not cause any phenotypic changes	Kim et al.,2005b； Nomura et al.,2005
	AtCYP85A2	cyp85a1-1/cyp85a2-2	严重矮化 Severe dwarfism	35S-CYP85A2	莲座叶更大，叶柄更长，茎更高且多分枝，角果更大，花梗更长 Larger rosette leaves，longer petioles，taller stems with increased branching，larger siliques，longer pedicels	Kim et al.,2005b； Nomura et al.,2005
BR信号转导 BR signaling	AtBRI1	bri1	严重矮化，深绿色，叶片厚，顶端优势减弱，雄性不育 Severe dwarfism，dark green，thickened leaves，reduced apical dominance，male sterility	BRI1-GFP	叶柄伸长 Elongated petioles	Clouse,1996； Wang et al.,2001； Blanco- Touriñán et al.,2024
	AtBAK1	bak1-1	半矮化，莲座叶小而圆，叶柄短 Semi-dwarf，small and rounded rosette leaves，short petioles	bak1-1D	茎、叶和叶柄伸长 Elongated stems，leaves，and petioles	Li et al.,2002
	AtBSU1	bsu1;bsl1/BSL2,3-amiRNA	严重矮化 Severe dwarfism	BSU1-YFP/bri1-116	在bri1-116中过表达BSU1，部分挽救了其矮化表型 Overexpression of BSU1 in bri1-116 partially rescued the dwarf phenotype	Kim et al.,2009； Kim et al.,2018
	AtBSL1			BSL1-YFP/bri1-5	部分挽救了其矮化表型 Overexpression of BSL1 in bri1-5 partially rescued the dwarf phenotype
	AtBSL2			BSL2-YFP	叶柄伸长 Elongated petioles
	AtBSL3
	AtBIN2	Triple GSK3 mutant	下胚轴显著伸长 Significantly elongated hypocoty	bin2-1	矮化，深绿色，叶片卷曲，顶端优势减弱，雄性育性降低，开花延迟 Dwarf，dark green，curled leaves，reduced apical dominance，reduced male fertility，delayed flowering	Li et al.,2001； Vert & Chory,2006
	AtBZR1	BES1 RNAi	半矮化 Semi-dwarf	bzr1-1D	在黑暗中，下胚轴伸长；在光下，半矮化，呈深绿色，叶片宽，叶柄短，叶片数量增加，开花延迟 In darkness：elongated hypocotyl；In light：semi- dwarf，dark green，wide leaves，short petioles，increased leaf number，delayed flowering	Wang et al.,2002； Yin et al.,2005
	AtBES1	BES1 RNAi	半矮化 Semi-dwarf	bes1-D	完全挽救bri1的矮化表型，叶柄长而弯曲，叶片卷曲，过度的茎伸长 Fully rescued the dwarf phenotype of bri1，long and curved petioles，curled leaves，excessive stem elongation	Yin et al.,2002； Yin et al.,2005

Research Progress on the Mechanism and Application of Brassinosteroid in Regulating Plant Height

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