Hormonal 'Code' for the Number of Maize Tassel Branches

doi:10.11983/CBB25003

Abstract

Abstract: Maize is a main cereal crop in China. The number of tassel branches significantly affects density-tolerant plant architecture, which is the key to increasing yield. Currently, multiple genes associated with tassel branch number, many of which are closely related to plant hormone pathways, have been cloned and identified. This review summarizes the roles and signaling pathways of auxin, cytokinin, strigolactone, gibberellin and abscisic acid in regulating maize tassel branch number, preliminarily investigates the interaction mechanisms among hormones and the impact of environmental factors on hormonal regulatory mechanisms, and outlines the hormonal regulation network for tassel branch number in maize. These findings suggest that plant hormones integrate environmental factors to regulate the tassel branch number. This review aims to provide a theoretical reference for revealing the molecular mechanisms of phytohormone regulation of maize tassel branch number and to lay a scientific foundation for novel cultivars of the maize ideotype through plant hormones.

Key words: maize, tassel branch number, plant hormone, environmental factor, genetic regulation

Miao Lin. Hormonal 'Code' for the Number of Maize Tassel Branches[J]. Chinese Bulletin of Botany, 2026, 61(1): 157-169.

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URL: https://www.chinbullbotany.com/EN/10.11983/CBB25003

https://www.chinbullbotany.com/EN/Y2026/V61/I1/157

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References 0

	Alberti S (2020). The plant response to heat requires phase separation. Nature 585, 191–192
	Bensen RJ, Johal GS, Crane VC, Tossberg JT, Schnable PS, Meeley RB, Briggs SP (1995). Cloning and characterization of the maize An1 gene. Plant Cell 7, 75–84
	Bolduc N, Hake S (2009). The maize transcription factor KNOTTED1 directly regulates the gibberellin catabolism gene ga2ox1. Plant Cell 21, 1647–1658
	Bolduc N, Yilmaz A, Mejia-Guerra MK, Morohashi K, O’Connor D, Grotewold E, Hake S (2012). Unraveling the KNOTTED1 regulatory network in maize meristems. Genes Dev 26, 1685–1690
	Carraro N, Forestan C, Canova S, Traas J, Varotto S (2006). ZmPIN1a and ZmPIN1b encode two novel putative candidates for polar auxin transport and plant architecture determination of maize. Plant Physiol 142, 254–264
	Chen FQ, Zhang JQ, Ma HL (2024). Progress of research on hormone regulation of branching or tillering in plants. Acta Pratacul Sin 33, 212–225. (in Chinese)
	陈奋奇, 张金青, 马晖玲 (2024). 激素调控植物分枝/分蘖的研究进展. 草业学报 33, 212–225
	Chen Y, Hou MM, Liu LJ, Wu S, Shen Y, Ishiyama K, Kobayashi M, McCarty DR, Tan BC (2014). The maize DWARF1 encodes a gibberellin 3-oxidase and is dual localized to the nucleus and cytosol. Plant Physiol 166, 2028–2039
	Chuck G, Cigan AM, Saeteurn K, Hake S (2007). The heterochronic maize mutant Corngrass1 results from overexpression of a tandem microRNA. Nat Genet 39, 544–549
	Chuck GS, Brown P, Meeley R, Hake S (2014). Maize SBP-box transcription factors unbranched2 and unbranched3 affect yield traits by regulating the rate of lateral primordia initiation. Proc Natl Acad Sci USA 111, 18775–18780
	Ding ST, Sun Q, Luo HB (2014). Research progress on differentiation and development of tassel in maize. Crop Res 28, 97–102. (in Chinese)
	丁帅涛, 孙琴, 罗红兵 (2014). 玉米雄穗分化发育研究进展. 作物研究 28, 97–102
	Dong ZB, Hu GY, Chen QY, Shemyakina EA, Chau G, Whipple CJ, Fletcher JC, Chuck G (2024). A regulatory network controlling developmental boundaries and meristem fates contributed to maize domestication. Nat Genet 56, 2528–2537
	Dong ZB, Jiang C, Chen XY, Zhang T, Ding L, Song WB, Luo HB, Lai JS, Chen HB, Liu RY, Zhang XL, Jin WW (2013). Maize LAZY1 mediates shoot gravitropism and inflorescence development through regulating auxin transport, auxin signaling, and light response. Plant Physiol 163, 1306–1322
	Du YF, Liu L, Li MF, Fang S, Shen XM, Chu JF, Zhang ZX (2017). UNBRANCHED3 regulates branching by modulating cytokinin biosynthesis and signaling in maize and rice. New Phytol 214, 721–733
	Du YF, Liu L, Peng Y, Li MF, Li YF, Liu D, Li XW, Zhang ZX (2020). UNBRANCHED3 expression and inflorescence development is mediated by UNBRANCHED2 and the distal enhancer, KRN4, in maize. PLoS Genet 16, e1008764
	Gallavotti A, Barazesh S, Malcomber S, Hall D, Jackson D, Schmidt RJ, McSteen P (2008a). sparse inflorescence1 encodes a monocot-specific YUCCA-like gene required for vegetative and reproductive development in maize. Proc Natl Acad Sci USA 105, 15196–15201
	Gallavotti A, Malcomber S, Gaines C, Stanfield S, Whipple C, Kellogg E, Schmidt RJ (2011). BARREN STALK FASTIGIATE1 is an AT-hook protein required for the formation of maize ears. Plant Cell 23, 1756–1771
	Gallavotti A, Yang Y, Schmidt RJ, Jackson D (2008b). The relationship between auxin transport and maize branching. Plant Physiol 147, 1913–1923
	Gallavotti A, Zhao Q, Kyozuka J, Meeley RB, Ritter MK, Doebley JF, Enrico Pè M, Schmidt RJ (2004). The role of barren stalk1 in the architecture of maize. Nature 432, 630–635
	Galli M, Liu QJ, Moss BL, Malcomber S, Li W, Gaines C, Federici S, Roshkovan J, Meeley R, Nemhauser JL, Gallavotti A (2015). Auxin signaling modules regulate maize inflorescence architecture. Proc Natl Acad Sci USA 112, 13372–13377
	Guan HH, Chen XJ, Wang KL, Liu XY, Zhang DF, Li YX, Song YC, Shi YS, Wang TY, Li CH, Li Y (2022). Genetic variation in ZmPAT7 contributes to tassel branch number in maize. Int J Mol Sci 23, 2586
	Guo HL, Tian MZ, Ri X, Chen YF (2025). Phosphorus acquisition, translocation, and redistribution in maize. J Genet Genom 52, 287–296
	Hao SL (2010). Effect of emasculation on physiological index of maize plant. Rain Fed Crops 30, 342–343. (in Chinese)
	郝淑丽 (2010). 去雄对玉米植株各项生理指标的影响. 杂粮作物 30, 342–343
	He XQ, Liu D (2009). Occurrence regularity and control methods of corn brown spot and small spot diseases. Mod Rural Technol (18), 29. (in Chinese)
	何晓庆, 刘冬 (2009). 玉米褐斑病小斑病发生规律及防治方法. 现代农村科技 (18), 29
	Herrero MP, Johnson RR (1981). Drought stress and its effects on maize reproductive system. Crop Sci 21, 105–110
	Hou ZW, Wang D (2021). Function of wheat tiller suppressor gene and its application prospect in population regulation. Jiangsu Agric Sci 49 (17), 53–57. (in Chinese)
	侯振伟, 王东 (2021). 小麦分蘖抑制基因的功能及其在群体调控中的应用前景. 江苏农业科学 49 (17), 53–57
	Huang P, Jiang H, Zhu CM, Barry K, Jenkins J, Sandor L, Schmutz J, Box MS, Kellogg EA, Brutnell TP (2017). Sparse panicle1 is required for inflorescence development in Setaria viridis and maize. Nat Plants 3, 17054
	Jafari F, Wang BB, Wang HY, Zou JJ (2024). Breeding maize of ideal plant architecture for high-density planting tolerance through modulating shade avoidance response and beyond. J Integr Plant Biol 66, 849–864
	Jia SF, Li CF, Dong ST, Zhang JW (2010). Physiological mechanism of shading stress on photosynthetic efficiency in summer maize (Zea mays). Chin J Plant Ecol 34, 1439–1447. (in Chinese)
	贾士芳, 李从锋, 董树亭, 张吉旺 (2010). 弱光胁迫影响夏玉米光合效率的生理机制初探. 植物生态学报 34, 1439–1447
	Jiang L, Liu X, Xiong GS, Liu HH, Chen FL, Wang L, Meng XB, Liu GF, Yu H, Yuan YD, Yi W, Zhao LH, Ma HL, He YZ, Wu ZS, Melcher K, Qian Q, Xu HE, Wang YH, Li JY (2013). DWARF 53 acts as a repressor of strigolactone signaling in rice. Nature 504, 401–405
	Jiao YQ, Wang YH, Xue DW, Wang J, Yan MX, Liu GF, Dong GJ, Zeng DL, Lu ZF, Zhu XD, Qian Q, Li JY (2010). Regulation of OsSPL14 by OsmiR156 defines ideal plant architecture in rice. Nat Genet 42, 541–544
	Jing HW, Korasick DA, Emenecker RJ, Morffy N, Wilkinson EG, Powers SK, Strader LC (2022). Regulation of AUXIN RESPONSE FACTOR condensation and nucleo-cytoplasmic partitioning. Nat Commun 13, 4015
	Kerstetter RA, Laudencia-Chingcuanco D, Smith LG, Hake S (1997). Loss-of-function mutations in the maize homeobox gene, knotted1, are defective in shoot meristem maintenance. Development 124, 3045–3054
	Kleine-Vehn J, Huang F, Naramoto S, Zhang J, Michniewicz M, Offringa R, Friml J (2009). PIN auxin efflux carrier polarity is regulated by PINOID kinase-mediated recruitment into GNOM-independent trafficking in Arabidopsis. Plant Cell 21, 3839–3849
	Kong DX, Li CY, Xue WC, Wei HB, Ding H, Hu GZ, Zhang XM, Zhang GS, Zou T, Xian YT, Wang BB, Zhao YP, Liu YT, Xie YR, Xu MY, Wu H, Liu Q, Wang HY (2023). UB2/UB3/TSH4-anchored transcriptional networks regulate early maize inflorescence development in response to simulated shade. Plant Cell 35, 717–737
	Lambert RJ, Johnson RR (1978). Leaf angle, tassel morphology, and the performance of maize hybrids. Crop Sci 18, 499–502
	Lee BH, Johnston R, Yang Y, Gallavotti A, Kojima M, Travencolo BAN, da F Costa L, Sakakibara H, Jackson D (2009). Studies of aberrant phyllotaxy1 mutants of maize indicate complex interactions between auxin and cytokinin signaling in the shoot apical meristem. Plant Physiol 150, 205–216
	Leonard A, Holloway B, Guo M, Rupe M, Yu GX, Beatty M, Zastrow-Hayes G, Meeley R, Llaca V, Butler K, Stefani T, Jaqueth J, Li BL (2014). tassel-less1 encodes a boron channel protein required for inflorescence development in maize. Plant Cell Physiol 55, 1044–1054
	Li JF, Cao HR, Li SX, Dong XN, Zhao Z, Jia ZT, Yuan LX (2025). Genetic and molecular mechanisms underlying nitrogen use efficiency in maize. J Genet Genom 52, 276–286
	Li M, Tang D, Wang KJ, Wu XR, Lu LL, Yu HX, Gu MH, Yan CJ, Cheng ZK (2011). Mutations in the F-box gene LARGER PANICLE improve the panicle architecture and enhance the grain yield in rice. Plant Biotechnol J 9, 1002–1013
	Li SK, Zhao JR, Dong ST, Zhao M, Li CH, Cui YH, Liu YH, Gao JL, Xue JQ, Wang LC, Wang P, Lu WP, Wang JH, Yang QF, Wang ZM (2017). Advances and prospects of maize cultivation in China. Sci Agric Sin 50, 1941–1959. (in Chinese)
	李少昆, 赵久然, 董树亭, 赵明, 李潮海, 崔彦宏, 刘永红, 高聚林, 薛吉全, 王立春, 王璞, 陆卫平, 王俊河, 杨祁峰, 王子明 (2017). 中国玉米栽培研究进展与展望. 中国农业科学 50, 1941–1959
	Li ZL, Li BZ, Zhang JL, Wang HL, Wang M, Guo SY, Wang PT, Li Z, Galbraith DW, Li DD, Song CP (2023). GA Associated Dwarf 5 encodes an ent-kaurenoic acid oxidase required for maize gibberellin biosynthesis and morphogenesis. Crop J 11, 1742–1751
	Liu QJ, Galli M, Liu X, Federici S, Buck A, Cody J, Labra M, Gallavotti A (2019a). NEEDLE1 encodes a mitochondria localized ATP-dependent metalloprotease required for thermotolerant maize growth. Proc Natl Acad Sci USA 116, 19736–19742
	Liu X, Galli M, Camehl I, Gallavotti A (2019b). RAMOSA1 ENHANCER LOCUS2-mediated transcriptional repression regulates vegetative and reproductive architecture. Plant Physiol 179, 348–363
	Liu YT, Wu GX, Zhao YP, Wang HHL, Dai ZY, Xue WC, Yang J, Wei HB, Shen RX, Wang HY (2021). DWARF53 interacts with transcription factors UB2/UB3/TSH4 to regulate maize tillering and tassel branching. Plant Physiol 187, 947–962
	Liu ZF (2011). Effects of Population Light Transmission on Stalk Strength and Lodging-resistance of Maize. Master’s thesis. Yangling: Northwest A&F University. pp. 1–54. (in Chinese)
	刘仲发 (2011). 群体光分布对玉米茎秆强度及抗倒伏能力的影响. 硕士论文. 杨凌: 西北农林科技大学. pp. 1–54
	Lu ZF, Yu H, Xiong GS, Wang J, Jiao YQ, Liu GF, Jing YH, Meng XB, Hu XM, Qian Q, Fu XD, Wang YH, Li JY (2013). Genome-wide binding analysis of the transcription activator IDEAL PLANT ARCHITECTURE1 reveals a complex network regulating rice plant architecture. Plant Cell 25, 3743–3759
	McSteen P, Hake S (2001). barren inflorescence2 regulates axillary meristem development in the maize inflorescence. Development 128, 2881–2891
	McSteen P, Malcomber S, Skirpan A, Lunde C, Wu XT, Kellogg E, Hake S (2007). barren inflorescence2 encodes a co-ortholog of the PINOID serine/threonine kinase and is required for organogenesis during inflorescence and vegetative development in maize. Plant Physiol 144, 1000–1011
	Moore S, Jervis G, Topping JF, Chen CL, Liu JL, Lindsey K (2024). A predictive model for ethylene-mediated auxin and cytokinin patterning in the Arabidopsis root. Plant Commun 5, 100886
	Mudgett M, Shen ZX, Dai XH, Briggs SP, Zhao YD (2023). Suppression of pinoid mutant phenotypes by mutations in PIN-FORMED 1 and PIN1-GFP fusion. Proc Natl Acad Sci USA 120, 2312918120
	Nielsen RL (2020). Tassel emergence & pollen shed. Corny news network. Purdue University. https://www.agry.purdue.edu/ext/corn/news/timeless/Tassels.html. 2025.12.23
	Patil SB, Barbier FF, Zhao JF, Zafar SA, Uzair M, Sun YL, Fang JJ, Perez-Garcia MD, Bertheloot J, Sakr S, Fichtner F, Chabikwa TG, Yuan SJ, Beveridge CA, Li XY (2022). Sucrose promotes D53 accumulation and tillering in rice. New Phytol 234, 122–136
	Phillips KA, Skirpan AL, Liu X, Christensen A, Slewinski TL, Hudson C, Barazesh S, Cohen JD, Malcomber S, McSteen P (2011). vanishing tassel2 encodes a grass-specific tryptophan aminotransferase required for vegetative and reproductive development in maize. Plant Cell 23, 550–566
	Qin XE, Tian SK, Zhang WL, Dong X, Ma CX, Wang Y, Yan JB, Yue B (2021). Q ^Dtbn1, an F-box gene affecting maize tassel branch number by a dominant model. Plant Biotechnol J 19, 1183–1194
	Schwarz S, Grande AV, Bujdoso N, Saedler H, Huijser P (2008). The microRNA regulated SBP-box genes SPL9 and SPL15 control shoot maturation in Arabidopsis. Plant Mol Biol 67, 183–195
	Skirpan A, Culler AH, Gallavotti A, Jackson D, Cohen JD, McSteen P (2009). BARREN INFLORESCENCE2 interaction with ZmPIN1a suggests a role in auxin transport during maize inflorescence development. Plant Cell Physiol 50, 652–657
	Skirpan A, Wu XT, McSteen P (2008). Genetic and physical interaction suggest that BARREN STALK1 is a target of BARREN INFLORESCENCE2 in maize inflorescence development. Plant J 55, 787–797
	Song FB, Dai JY (2005). Response and adaptability of maize to drought stress II. Responses of ear and tassel growth and development of maize to drought stress. J Jilin Agric Univ 27, 1–5, 10. (in Chinese)
	宋凤斌, 戴俊英 (2005). 玉米对干旱胁迫的反应和适应性II. 玉米雌穗和雄穗生长发育对干旱胁迫的反应. 吉林农业大学学报 27, 1–5, 10
	Song XG, Lu ZF, Yu H, Shao GN, Xiong JS, Meng XB, Jing YH, Liu GF, Xiong GS, Duan JB, Yao XF, Liu CM, Li HQ, Wang YH, Li JY (2017). IPA1 functions as a downstream transcription factor repressed by D53 in strigolactone signaling in rice. Cell Res 27, 1128–1141
	Song XG, Meng XB, Guo HY, Cheng Q, Jing YH, Chen MJ, Liu GF, Wang B, Wang YH, Li JY, Yu H (2022). Targeting a gene regulatory mixed enhances rice grain yield by decoupling panicle number and size. Nat Biotechnol 40, 1403–1411
	Spielmeyer W, Ellis MH, Chandler PM (2002). Semidwarf (sd-1), “green revolution” rice, contains a defective gibberellin 20-oxidase gene. Proc Natl Acad Sci USA 99, 9043–9048
	Su S, Hong J, Chen XF, Zhang CQ, Chen MJ, Luo ZJ, Chang SW, Bai SX, Liang WQ, Liu QQ, Zhang DB (2021). Gibberellins orchestrate panicle architecture mediated by DELLA-KNOX signaling in rice. Plant Biotechnol J 19, 2304–2318
	Tang SY, Xie Q (2019). Sorghum—a small crop with numerous uses. Biotechnol Bull 35 (5), 1. (in Chinese)
	唐三元, 谢旗 (2019). 高粱——小作物大用途. 生物技术通报 35 (5), 1
	Vukašinović N, Hsu CW, Marconi M, Li SP, Zachary C, Shahan R, Szekley P, Aardening Z, Vanhoutte I, Ma Q, Pinto L, Krupař P, German N, Zhang JY, Simon-Wezo C, Perez-Sancho J, Quijada PC, Zhou QZ, Lee LR, Cai JH, Bayer EM, Fendrych M, Truernit E, Zhou Y, Savaldi-Goldstein S, Wabnik K, Nolan TM, Russinova E (2025). Polarity-guided uneven mitotic divisions control brassinosteroid activity in proliferating plant root cells. Cell 188, 2063–2080
	Wang B, Li JY, Wang YH (2006). Advances in understanding the roles of auxin involved in modulating plant architecture. Chin Bull Bot 23, 443–458. (in Chinese with Enqlish Abstract)
	王冰, 李家洋, 王永红 (2006). 生长素调控植物株型形成的研究进展. 植物学报 23, 443–458
	Wang BB, Wang HY (2023). Molecular design of ideal plant architecture for high-density tolerance of maize plant. Biotechnol Bull 39 (8), 11–30. (in Chinese)
	王宝宝, 王海洋 (2023). 理想株型塑造之于玉米耐密改良. 生物技术通报 39 (8), 11–30
	Wang L, Wang B, Jiang L, Liu XL, Li XF, Lu Z, Meng XB, Wang YH, Smith SM, Li JY (2015). Strigolactone signaling in Arabidopsis regulates shoot development by targeting D53-like SMXL repressor proteins for ubiquitination and degradation. Plant Cell 27, 3128–3142
	Wang X, Li J, Han LQ, Liang CY, Li JX, Shang XY, Miao XX, Luo Z, Zhu WC, Li Z, Li TH, Qi YW, Li HH, Lu XD, Li L (2023). QTG-Miner aids rapid dissection of the genetic base of tassel branch number in maize. Nat Commun 14, 5232
	Wang YB, Luo Y, Guo X, Li YF, Yan JL, Shao WW, Wei WJ, Wei XF, Yang T, Chen J, Chen LH, Ding Q, Bai MJ, Zhuo L, Li L, Jackson D, Zhang ZX, Xu X, Yan JB, Liu H, Liu L, Yang N (2024). A spatial transcriptome map of the developing maize ear. Nat Plants 10, 815–827
	Wang YJ, Deng DX, Ding HD, Xu XM, Zhang R, Wang SX, Bian YL, Yin ZT, Chen Y (2013). Gibberellin biosynthetic deficiency is responsible for maize dominant Dwarf11 (D11) mutant phenotype: physiological and transcriptomic evidence. PLoS One 8, e66466
	Wu HM, Xie DJ, Tang ZS, Shi DQ, Yang WC (2020). PINOID regulates floral organ development by modulating auxin transport and interacts with MADS16 in rice. Plant Biotechnol J 18, 1778–1795
	Xie SY, Luo HB, Huang YM, Wang YX, Ru W, Shi YL, Huang W, Wang H, Dong ZB, Jin WW (2020a). A missense mutation in a large subunit of ribonucleotide reductase confers temperature-gated tassel formation. Plant Physiol 184, 1979–1997
	Xie YR, Liu Y, Ma MD, Zhou Q, Zhao YP, Zhao BB, Wang BB, Wei HB, Wang HY (2020b). Arabidopsis FHY3 and FAR1 integrate light and strigolactone signaling to regulate branching. Nat Commun 11, 1955
	Xie YR, Zhao YP, Chen LH, Wang YL, Xue WC, Kong DX, Li CY, Zhou LY, Li HR, Zhao YF, Wang BB, Xu MY, Zhao BB, Bilska-Kos A, Wang HY (2024). ZmELF3.1 integrates the RA2-TSH4 module to repress maize tassel branching. New Phytol 241, 490–503
	Xu HW, Song FB, Tong SY, Zhu XC (2012). Study on carbon and nitrogen metabolism of tassel in maize. Hubei Agric Sci 51, 2918–2920. (in Chinese)
	徐洪文, 宋凤斌, 童淑媛, 朱先灿 (2012). 玉米雄穗碳氮代谢研究. 湖北农业科学 51, 2918–2920
	Yan JB, Zhao JR (2023). Dense planting for high yield—the key goal of maize breeding in China. Biotechnol Bull 39 (8), 1–3. (in Chinese)
	严建兵, 赵久然 (2023). 密植高产——我国玉米育种的最核心目标. 生物技术通报 39 (8), 1–3
	Yang T, Lian YK, Kang JH, Bian ZY, Xuan LJ, Gao ZS, Wang XY, Deng JM, Wang CY (2020). The SUPPRESSOR of MAX21 (SMAX1)-like SMXL6, SMXL7 and SMXL8 act as negative regulators in response to drought stress in Arabidopsis. Plant Cell Physiol 61, 1477–1492
	Yao H, Skirpan A, Wardell B, Matthes MS, Best NB, McCubbin T, Durbak A, Smith T, Malcomber S, McSteen P (2019). The barren stalk2 gene is required for axillary meristem development in maize. Mol Plant 12, 374–389
	Yoshida A, Ohmori Y, Kitano H, Taguchi-Shiobara F, Hirano HY (2012). ABERRANT SPIKELET AND PANICLE1, encoding a TOPLESS-related transcriptional co-repressor is involved in the regulation of meristem fate in rice. Plant J 70, 327–339
	Yuan K, Zhang H, Yu CJ, Luo N, Yan JJ, Zheng S, Hu QL, Zhang DH, Kou LQ, Meng XB, Jing YH, Chen MJ, Ban XW, Yan ZY, Lu ZF, Wu J, Zhao Y, Liang Y, Wang YH, Xiong GS, Chu JF, Wang ET, Li J, Wang B (2023). Low phosphorus promotes NSP1-NSP2 heterodimerization to enhance strigolactone biosynthesis and regulate shoot and root architecture in rice. Mol Plant 16, 1811–1831
	Zhang D, Sun W, Singh R, Zheng YY, Cao Z, Li MF, Lunde C, Hake S, Zhang ZX (2018). GRF-interacting factor1 regulates shoot architecture and meristem determinacy in maize. Plant Cell 30, 360–374
	Zhang JF, Zhang YY, Chen JG, Xu MF, Guan XY, Wu C, Zhang SN, Qu HY, Chu JF, Xu YF, Gu M, Liu Y, Xu GH (2024). Sugar transporter modulates nitrogen-determined tillering and yield formation in rice. Nat Commun 15, 9233
	Zhang JW, Dong ST, Wang KJ, Hu CH, Liu P (2006). Effects of shading on the growth, development and grain yield of summer maize. Chin J Appl Ecol 17, 657–662. (in Chinese)
	张吉旺, 董树亭, 王空军, 胡昌浩, 刘鹏 (2006). 遮荫对夏玉米产量及生长发育的影响. 应用生态学报 17, 657–662
	Zhu CM, Box MS, Thiruppathi D, Hu H, Yu YQ, Martin C, Doust AN, McSteen P, Kellogg EA (2022). Pleiotropic and nonredundant effects of an auxin importer in setaria and maize. Plant Physiol 189, 715–734
	Zhu XH (2008). Selecting of Drought Resistant Resources and Its Analysis of Genetic Diversity in Foxtail Millet. Master’s thesis. Beijing: Chinese Academy of Agricultural Sciences. pp. 1−45. (in Chinese)
	朱学海 (2008). 谷子耐旱资源筛选及其遗传多样性分析. 硕士论文. 北京: 中国农业科学院. pp. 1−45
	Zong J, Wang L, Zhu L, Bian LL, Zhang B, Chen XF, Huang GQ, Zhang XL, Fan JY, Cao LM, Coupland G, Liang WQ, Zhang DB, Yuan Z (2022). A rice single cell transcriptomic atlas defines the developmental trajectories of rice floret and inflorescence meristems. New Phytol 234, 494−512

基因	基因编码产物	作用途径	生物学功能	参考文献
SPI1	黄素单加氧酶	参与生长素生物合成	通过正向调节分枝分生组织（BM）形成来正调控玉米雄穗分枝数	Gallavotti et al.，2008a
VT2	色氨酸转氨酶	参与生长素生物合成	通过正向调节BM形成来正调控玉米雄穗分枝数	Phillips et al.，2011
BIF2	丝氨酸/苏氨酸蛋白激酶	参与生长素运输	通过正向调节BM形成和维持来正调控玉米雄穗分枝数	McSteen and Hake，2001；McSteen et al.，2007
ZmPIN1a	生长素外排转运蛋白	参与生长素运输	通过正向调节BM形成来正调控玉米雄穗分枝数	Galli et al.，2015
ZmAUX1	生长素内流载体	参与生长素运输	正调控玉米雄穗分枝数	Zhu et al.，2022
BIF1&BIF4	AUX/IAA蛋白	参与生长素信号转导	通过正向调节BM形成来正调控玉米雄穗分枝数	Gallavotti et al.，2004
BA1	bHLH转录因子	生长素信号通路早期靶点	通过正向调节BM形成来正调控玉米雄穗分枝数	Gallavotti et al.，2004
ZmD53	SCF^D3泛素复合物底物	独脚金内酯（SL）信号通路核心抑制因子	通过正向调节BM形成来正调控玉米雄穗分枝数	Liu et al.，2021
UB3	SBP-box转录因子	间接参与细胞分裂素（CK）生物合成和降解反应	通过调节IM外侧细胞分化起始速率来调节分生组织活性，正调控玉米雄穗分枝数	Chuck et al.，2014；Du et al.，2017
Q^Dtbn1	含kelch结构域的F-box蛋白	间接参与脱落酸（ABA）信号转导	负调控玉米雄穗分枝数	Qin et al.，2021
D1	GA-3-氧化酶	参与赤霉素（GA）生物合成	正调控玉米雄穗分枝数	Chen et al.，2014
D5	细胞色素P450蛋白家族成员CYP88A1	参与GA生物合成	正调控玉米雄穗分枝数	Li et al.，2023
D11	未克隆，但与GA-20-氧化酶1的表达有关	参与GA生物合成	正调控玉米雄穗分枝数	Wang et al.，2013
AN1	催化GA第1个四环中间体合成的靶向叶绿体的植物环化酶	参与GA生物合成	正调控玉米雄穗分枝数	Bensen et al.，1995
KN1	同源域KNOTTED1转录因子	间接参与GA降解反应	正调控玉米雄穗分枝数	Kerstetter et al.，1997；Bolduc and Hake，2009

基因	基因编码产物	作用途径	生物学功能	参考文献
SPI1	黄素单加氧酶	参与生长素生物合成	通过正向调节分枝分生组织（BM）形成来正调控玉米雄穗分枝数	Gallavotti et al.，2008a
VT2	色氨酸转氨酶	参与生长素生物合成	通过正向调节BM形成来正调控玉米雄穗分枝数	Phillips et al.，2011
BIF2	丝氨酸/苏氨酸蛋白激酶	参与生长素运输	通过正向调节BM形成和维持来正调控玉米雄穗分枝数	McSteen and Hake，2001；McSteen et al.，2007
ZmPIN1a	生长素外排转运蛋白	参与生长素运输	通过正向调节BM形成来正调控玉米雄穗分枝数	Galli et al.，2015
ZmAUX1	生长素内流载体	参与生长素运输	正调控玉米雄穗分枝数	Zhu et al.，2022
BIF1&BIF4	AUX/IAA蛋白	参与生长素信号转导	通过正向调节BM形成来正调控玉米雄穗分枝数	Gallavotti et al.，2004
BA1	bHLH转录因子	生长素信号通路早期靶点	通过正向调节BM形成来正调控玉米雄穗分枝数	Gallavotti et al.，2004
ZmD53	SCF^D3泛素复合物底物	独脚金内酯（SL）信号通路核心抑制因子	通过正向调节BM形成来正调控玉米雄穗分枝数	Liu et al.，2021
UB3	SBP-box转录因子	间接参与细胞分裂素（CK）生物合成和降解反应	通过调节IM外侧细胞分化起始速率来调节分生组织活性，正调控玉米雄穗分枝数	Chuck et al.，2014；Du et al.，2017
Q^Dtbn1	含kelch结构域的F-box蛋白	间接参与脱落酸（ABA）信号转导	负调控玉米雄穗分枝数	Qin et al.，2021
D1	GA-3-氧化酶	参与赤霉素（GA）生物合成	正调控玉米雄穗分枝数	Chen et al.，2014
D5	细胞色素P450蛋白家族成员CYP88A1	参与GA生物合成	正调控玉米雄穗分枝数	Li et al.，2023
D11	未克隆，但与GA-20-氧化酶1的表达有关	参与GA生物合成	正调控玉米雄穗分枝数	Wang et al.，2013
AN1	催化GA第1个四环中间体合成的靶向叶绿体的植物环化酶	参与GA生物合成	正调控玉米雄穗分枝数	Bensen et al.，1995
KN1	同源域KNOTTED1转录因子	间接参与GA降解反应	正调控玉米雄穗分枝数	Kerstetter et al.，1997；Bolduc and Hake，2009

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