Molecular Mechanisms of Rice Grain Size Regulation Related to Plant Hormone Signaling Pathways

doi:10.13560/j.cnki.biotech.bull.1985.2023-0273

Abstract

Abstract:

Rice is a major food crop for humans, and how to effectively improve its yield and quality is a major scientific concern. Rice grain size is one of the main factors affecting yield, and research on the regulation of rice grain development is an important guide for using molecular design breeding to improve yield and quality. Grain size is determined by a combination of the length, width and thickness of the grain; it is a quantitative trait regulated by multiple genes, and is one of the crucial determinants of rice yield and quality. In recent years, a large number of quantitative trait loci（QTLs）related to grain size have been identified through the study of mutants with seed development defects in rice, and some related genes have been cloned and identified, and the complex signalling pathways regulating grain size in rice are gradually being elucidated. Several studies have found that functional genes regulating rice seed development are involved in the synthesis, catabolism and transport of plant hormones as well as the signal transduction pathways of plant hormones. This review outlines the basic process of rice endosperm development, summarizes the overall the overall understanding of the dynamic changes of plant hormones during endosperm development, focuses on the current research status of QTLs related to phytohormone signalling pathways associated with rice grain size, summarizes and analyses the relationship between the pathways related to cytokinin, brassinosteroid, growth hormone, gibberellin, ethylene and jasmonic acid and grain size regulation, and further sorts out the regulation network of rice grain shape-related phytohormone signaling. It is aimed to provide a reference for identifying and analyzing the molecular mechanisms of phytohormone regulation of rice grain size, and to provide new ideas for rice molecular design breeding.

Key words: rice, grain size, gene, plant hormone signal

YAO Sha-sha, WANG Jing-jing, WANG Jun-jie, LIANG Wei-hong. Molecular Mechanisms of Rice Grain Size Regulation Related to Plant Hormone Signaling Pathways[J]. Biotechnology Bulletin, 2023, 39(8): 80-90.

Figures/Tables 3

References 77

[1]	Fan YW, Li YB. Molecular, cellular and Yin-Yang regulation of grain size and number in rice[J]. Mol Breeding, 2019, 39(12): 163. doi: 10.1007/s11032-019-1078-0
[2]	Xing YZ, Zhang QF. Genetic and molecular bases of rice yield[J]. Annu Rev Plant Biol, 2010, 61: 421-442. doi: 10.1146/annurev-arplant-042809-112209 pmid: 20192739
[3]	Li GM, Tang JY, Zheng JK, et al. Exploration of rice yield potential: Decoding agronomic and physiological traits[J]. Crop J, 2021, 9(3): 577-589. doi: 10.1016/j.cj.2021.03.014
[4]	Zhao DS, Zhang CQ, Li QF, et al. Genetic control of grain appearance quality in rice[J]. Biotechnol Adv, 2022, 60: 108014. doi: 10.1016/j.biotechadv.2022.108014 URL
[5]	刘迪, 冯连杰, 梁卫红. 水稻粒型调控相关信号通路的鉴定与解析[J]. 中国生物化学与分子生物学报, 2023, 39(2): 212-221.
	Liu D, Feng LJ, Liang WH. Identification and analysis of grain shape related regulation signal pathways in rice[J]. Chin J Biochem Mol Biol, 2023, 39(2): 212-221.
[6]	Azizi P, Osman M, Hanafi MM, et al. Molecular insights into the regulation of rice kernel elongation[J]. Crit Rev Biotechnol, 2019, 39(7): 904-923. doi: 10.1080/07388551.2019.1632257 pmid: 31303070
[7]	Ren DY, Ding CQ, Qian Q. Molecular bases of rice grain size and quality for optimized productivity[J]. Sci Bull, 2023, 68(3): 314-350. doi: 10.1016/j.scib.2023.01.026 URL
[8]	Li N, Xu R, Duan PG, et al. Control of grain size in rice[J]. Plant Reprod, 2018, 31(3): 237-251. doi: 10.1007/s00497-018-0333-6 pmid: 29523952
[9]	Basunia MA, Nonhebel HM. Hormonal regulation of cereal endosperm development with a focus on rice(Oryza sativa)[J]. Funct Plant Biol, 2019, 46(6): 493-506. doi: 10.1071/FP18323 pmid: 30955506
[10]	Zhang XF, Tong JH, Bai AN, et al. Phytohormone dynamics in developing endosperm influence rice grain shape and quality[J]. J Integr Plant Biol, 2020, 62(10): 1625-1637. doi: 10.1111/jipb.12927
[11]	Li N, Xu R, Li YH. Molecular networks of seed size control in plants[J]. Annu Rev Plant Biol, 2019, 70: 435-463. doi: 10.1146/annurev-arplant-050718-095851 pmid: 30795704
[12]	Wu XB, Liu JX, Li DQ, et al. Rice caryopsis development I: Dynamic changes in different cell layers[J]. J Integr Plant Biol, 2016, 58(9): 772-785. doi: 10.1111/jipb.12440
[13]	Wu XB, Liu JX, Li DQ, et al. Rice caryopsis development II: Dynamic changes in the endosperm[J]. J Integr Plant Biol, 2016, 58(9): 786-798. doi: 10.1111/jipb.12488
[14]	Weier DA, Thiel J, Kohl S, et al. Gibberellin-to-abscisic acid balances govern development and differentiation of the nucellar projection of barley grains[J]. J Exp Bot, 2014, 65(18): 5291-5304. doi: 10.1093/jxb/eru289 pmid: 25024168
[15]	Yang JC, Zhang JH, Wang ZQ, et al. Post-anthesis development of inferior and superior spikelets in rice in relation to abscisic acid and ethylene[J]. J Exp Bot, 2006, 57(1): 149-160. pmid: 16330527
[16]	Liu K, Li MJ, Zhang B, et al. Poaceae orthologs of rice OsSGL, DUF1645 domain-containing genes, positively regulate drought tolerance, grain length and weight in rice[J]. Rice Sci, 2022, 29(3): 257-267. doi: 10.1016/j.rsci.2021.11.001
[17]	Wang ML, Lu XD, Xu GY, et al. OsSGL, a novel pleiotropic stress-related gene enhances grain length and yield in rice[J]. Sci Rep, 2016, 6: 38157. doi: 10.1038/srep38157 pmid: 27917884
[18]	Jameson PE, Song JC. Cytokinin: A key driver of seed yield[J]. J Exp Bot, 2016, 67(3): 593-606. doi: 10.1093/jxb/erv461 pmid: 26525061
[19]	Li SY, Zhao BR, Yuan DY, et al. Rice zinc finger protein DST enhances grain production through controlling Gn1a/OsCKX2 expression[J]. Proc Natl Acad Sci USA, 2013, 110(8): 3167-3172. doi: 10.1073/pnas.1300359110 URL
[20]	Xiao YH, Liu DP, Zhang GX, et al. Big Grain3, encoding a purine permease, regulates grain size via modulating cytokinin transport in rice[J]. J Integr Plant Biol, 2019, 61(5): 581-597. doi: 10.1111/jipb.v61.5 URL
[21]	Makarevitch I, Thompson A, Muehlbauer GJ, et al. Brd1 gene in maize encodes a brassinosteroid C-6 oxidase[J]. Plos One, 2012, 7(1): e30798. doi: 10.1371/journal.pone.0030798 URL
[22]	Huang JP, Chen ZM, Lin JJ, et al. Natural variation of the BRD2 allele affects plant height and grain size in rice[J]. Planta, 2022, 256(2): 27. doi: 10.1007/s00425-022-03939-7
[23]	Hong Z, Ueguchi-Tanaka M, Umemura K, et al. A rice brassinosteroid-deficient mutant, ebisu dwarf(d2), is caused by a loss of function of a new member of cytochrome P450[J]. Plant Cell, 2003, 15(12): 2900-2910. doi: 10.1105/tpc.014712 pmid: 14615594
[24]	Tanabe S, Ashikari M, Fujioka S, et al. A novel cytochrome P450 is implicated in brassinosteroid biosynthesis via the characterization of a rice dwarf mutant, dwarf11, with reduced seed length[J]. Plant Cell, 2005, 17(3): 776-790. doi: 10.1105/tpc.104.024950 URL
[25]	Wu YZ, Fu YC, Zhao SS, et al. CLUSTERED PRIMARY BRANCH 1, a new allele of DWARF11, controls panicle architecture and seed size in rice[J]. Plant Biotechnol J, 2016, 14(1): 377-386. doi: 10.1111/pbi.12391 URL
[26]	Zhou Y, Tao YJ, Zhu JY, et al. GNS4, a novel allele of DWARF11, regulates grain number and grain size in a high-yield rice variety[J]. Rice(NY), 2017, 10(1): 34.
[27]	Li Y, Li XM, Fu DB, et al. Panicle Morphology Mutant 1(PMM1)determines the inflorescence architecture of rice by controlling brassinosteroid biosynthesis[J]. BMC Plant Biol, 2018, 18(1): 348. doi: 10.1186/s12870-018-1577-x
[28]	Zhan HD, Lu MM, Luo Q, et al. OsCPD1 and OsCPD2 are functional brassinosteroid biosynthesis genes in rice[J]. Plant Sci, 2022, 325: 111482. doi: 10.1016/j.plantsci.2022.111482 URL
[29]	Tong HN, Liu LC, Jin Y, et al. DWARF AND LOW-TILLERING acts as a direct downstream target of a GSK3/SHAGGY-Like kinase to mediate brassinosteroid responses in rice[J]. Plant Cell, 2012, 24(6): 2562-2577. doi: 10.1105/tpc.112.097394 URL
[30]	Tian XJ, He ML, Mei EY, et al. WRKY53 integrates classic brassinosteroid signaling and the mitogen-activated protein kinase pathway to regulate rice architecture and seed size[J]. Plant Cell, 2021, 33(8): 2753-2775. doi: 10.1093/plcell/koab137 URL
[31]	Xiao YH, Liu DP, Zhang GX, et al. Brassinosteroids regulate OFP1, a DLT interacting protein, to modulate plant architecture and grain morphology in rice[J]. Front Plant Sci, 2017, 8: 1698. doi: 10.3389/fpls.2017.01698 pmid: 29021808
[32]	Xiao Y, Zhang G, Liu D, et al. GSK2 stabilizes OFP3 to suppress brassinosteroid responses in rice[J]. Plant J, 2020, 102(6): 1187-1201. doi: 10.1111/tpj.v102.6 URL
[33]	Yang C, Ma Y, He YM, et al. OsOFP19 modulates plant architecture by integrating the cell division pattern and brassinosteroid signaling[J]. Plant J, 2018, 93(3): 489-501. doi: 10.1111/tpj.2018.93.issue-3 URL
[34]	Chen XL, Jiang LR, Zheng JS, et al. A missense mutation in Large Grain Size 1 increases grain size and enhances cold tolerance in rice[J]. J Exp Bot, 2019, 70(15): 3851-3866. doi: 10.1093/jxb/erz192 URL
[35]	Zhao DS, Li QF, Zhang CQ, et al. GS9 acts as a transcriptional activator to regulate rice grain shape and appearance quality[J]. Nat Commun, 2018, 9(1): 1240. doi: 10.1038/s41467-018-03616-y
[36]	Huang YS, Dong H, Mou CL, et al. Ribonuclease H-like gene SMALL GRAIN2 regulates grain size in rice through brassinosteroid signaling pathway[J]. J Integr Plant Biol, 2022, 64(10): 1883-1900. doi: 10.1111/jipb.v64.10 URL
[37]	Gong JY, Miao JS, Zhao Y, et al. Dissecting the genetic basis of grain shape and chalkiness traits in hybrid rice using multiple collaborative populations[J]. Mol Plant, 2017, 10(10): 1353-1356. doi: S1674-2052(17)30231-9 pmid: 28803900
[38]	Liu JF, Chen J, Zheng XM, et al. GW5 acts in the brassinosteroid signalling pathway to regulate grain width and weight in rice[J]. Nat Plants, 2017, 3: 17043. doi: 10.1038/nplants.2017.43 URL
[39]	Li YB, Fan CC, Xing YZ, et al. Natural variation in GS5 plays an important role in regulating grain size and yield in rice[J]. Nat Genet, 2011, 43(12): 1266-1269. doi: 10.1038/ng.977
[40]	Xu CJ, Liu Y, Li YB, et al. Differential expression of GS5 regulates grain size in rice[J]. J Exp Bot, 2015, 66(9): 2611-2623. doi: 10.1093/jxb/erv058 URL
[41]	Weijers D, Wagner D. Transcriptional responses to the auxin hormone[J]. Annu Rev Plant Biol, 2016, 67: 539-574. doi: 10.1146/annurev-arplant-043015-112122 pmid: 26905654
[42]	Powers SK, Strader LC. Regulation of auxin transcriptional responses[J]. Dev Dyn, 2020, 249(4): 483-495. doi: 10.1002/dvdy.v249.4 URL
[43]	Hu ZJ, Lu SJ, Wang MJ, et al. A novel QTL qTGW3 encodes the GSK3/SHAGGY-like kinase OsGSK5/OsSK41 that interacts with OsARF4 to negatively regulate grain size and weight in rice[J]. Mol Plant, 2018, 11(5): 736-749. doi: 10.1016/j.molp.2018.03.005 URL
[44]	Qiao JY, Jiang HZ, Lin YQ, et al. A novel miR167a-OsARF6-OsAUX3 module regulates grain length and weight in rice[J]. Mol Plant, 2021, 14(10): 1683-1698. doi: 10.1016/j.molp.2021.06.023 URL
[45]	Zhang ZY, Li JJ, Tang ZS, et al. Gnp4/LAX2, a RAWUL protein, interferes with the OsIAA3-OsARF25 interaction to regulate grain length via the auxin signaling pathway in rice[J]. J Exp Bot, 2018, 69(20): 4723-4737. doi: 10.1093/jxb/ery256 pmid: 30295905
[46]	Zhang SY, Zhu LM, Shen CB, et al. Natural allelic variation in a modulator of auxin homeostasis improves grain yield and nitrogen use efficiency in rice[J]. Plant Cell, 2021, 33(3): 566-580. doi: 10.1093/plcell/koaa037 URL
[47]	Guo T, Chen K, Dong NQ, et al. Tillering and small grain 1 dominates the tryptophan aminotransferase family required for local auxin biosynthesis in rice[J]. J Integr Plant Biol, 2020, 62(5): 581-600. doi: 10.1111/jipb.v62.5 URL
[48]	Dong NQ, Sun YW, Guo T, et al. UDP-glucosyltransferase regulates grain size and abiotic stress tolerance associated with metabolic flux redirection in rice[J]. Nat Commu, 2020, 11(1): 2629.
[49]	Davière JM, Achard P. Gibberellin signaling in plants[J]. Development, 2013, 140(6): 1147-1151. doi: 10.1242/dev.087650 URL
[50]	Kumar A, Singh A, Kumar P, et al. Giberellic acid-stimulated transcript proteins evolved through successive conjugation of novel motifs and their subfunctionalization[J]. Plant Physiol, 2019, 180(2): 998-1012. doi: 10.1104/pp.19.00305 pmid: 30971449
[51]	Shi CL, Dong NQ, Guo T, et al. A quantitative trait locus GW6 controls rice grain size and yield through the gibberellin pathway[J]. Plant J, 2020, 103(3): 1174-1188. doi: 10.1111/tpj.v103.3 URL
[52]	Li XB, Shi SY, Tao QD, et al. OsGASR9 positively regulates grain size and yield in rice(Oryza sativa)[J]. Plant Sci, 2019, 286: 17-27. doi: 10.1016/j.plantsci.2019.03.008 URL
[53]	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): 429-450. doi: 10.1007/s11103-020-01049-0
[54]	Chen WW, Cheng ZJ, Liu LL, et al. Small Grain and Dwarf 2, encoding an HD-Zip II family transcription factor, regulates plant development by modulating gibberellin biosynthesis in rice[J]. Plant Sci, 2019, 288: 110208. doi: 10.1016/j.plantsci.2019.110208 URL
[55]	Jang S, Cho JY, Do GR, et al. Modulation of rice leaf angle and grain size by expressing OsBCL1 and OsBCL2 under the control of OsBUL1 promoter[J]. Int J Mol Sci, 2021, 22(15): 7792. doi: 10.3390/ijms22157792 URL
[56]	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. doi: 10.3390/ijms23158715 URL
[57]	Yun P, Li YB, Wu B, et al. OsHXK3 encodes a hexokinase-like protein that positively regulates grain size in rice[J]. Theor Appl Genet, 2022, 135(10): 3417-3431. doi: 10.1007/s00122-022-04189-7
[58]	Hussain S, Huang J, Zhu CQ, et al. Pyridoxal 5'-phosphate enhances the growth and morpho-physiological characteristics of rice cultivars by mitigating the ethylene accumulation under salinity stress[J]. Plant Physiol Biochem, 2020, 154: 782-795. doi: 10.1016/j.plaphy.2020.05.035 URL
[59]	Tao YJ, Wang J, Miao J, et al. The spermine synthase OsSPMS1 regulates seed germination, grain size, and yield[J]. Plant Physiol, 2018, 178(4): 1522-1536. doi: 10.1104/pp.18.00877 pmid: 30190417
[60]	Zhao H, Yin CC, Ma B, et al. Ethylene signaling in rice and arabidopsis: new regulators and mechanisms[J]. J Integr Plant Biol, 2021, 63(1): 102-125. doi: 10.1111/jipb.v63.1 URL
[61]	Liu C, Ma T, Yuan DY, et al. The OsEIL1-OsERF115-target gene regulatory module controls grain size and weight in rice[J]. Plant Biotechnol J, 2022, 20(8): 1470-1486. doi: 10.1111/pbi.v20.8 URL
[62]	Uji Y, Kashihara K, Kiyama H, et al. Jasmonic acid-induced VQ-motif-containing protein OsVQ13 influences the OsWRKY45 signaling pathway and grain size by associating with OsMPK6 in rice[J]. Int J Mol Sci, 2019, 20(12): 2917. doi: 10.3390/ijms20122917 URL
[63]	Callens C, Tucker MR, Zhang DB, et al. Dissecting the role of MADS-box genes in monocot floral development and diversity[J]. J Exp Bot, 2018, 69(10): 2435-2459. doi: 10.1093/jxb/ery086 pmid: 29718461
[64]	Chongloi GL, Prakash S, Vijayraghavan U. Regulation of meristem maintenance and organ identity during rice reproductive development[J]. J Exp Bot, 2019, 70(6): 1719-1736. doi: 10.1093/jxb/erz046 pmid: 30753578
[65]	Mehra P, Pandey BK, Verma L, et al. OsJAZ11 regulates spikelet and seed development in rice[J]. Plant Direct, 2022, 6(5): e401. doi: 10.1002/pld3.v6.5 URL
[66]	Kato T, Sakurai N, Kuraishi S. The changes of endogenous abscisic acid in developing grain of two rice cultivars with different grain size[J]. Japanese Journal of Crop Science, 1993, 62(3): 456-461. doi: 10.1626/jcs.62.456 URL
[67]	Chen Y, Xiang ZP, Liu M, et al. ABA biosynthesis gene OsNCED3 contributes to preharvest sprouting resistance and grain development in rice[J]. Plant, Cell Environ, 2023, 46(4): 1384-1401. doi: 10.1111/pce.14480 URL
[68]	Liu DP, Zhao H, Xiao YH, et al. A cryptic inhibitor of cytokinin phosphorelay controls rice grain size[J]. Mol Plant, 2022, 15(2): 293-307. doi: 10.1016/j.molp.2021.09.010 URL
[69]	Gao XY, Zhang JQ, Zhang XJ, et al. Rice qGL3/OsPPKL1 functions with the GSK3/SHAGGY-like kinase OsGSK3 to modulate brassinosteroid signaling[J]. Plant Cell, 2019, 31(5): 1077-1093. doi: 10.1105/tpc.18.00836 URL
[70]	Tian P, Liu JF, Yan BH, et al. OsBSK3 positively regulates grain length and weight by inhibiting the phosphatase activity of OsPPKL1[J]. Plants(Basel), 2022, 11(12): 1586.
[71]	Zhang JQ, Gao XY, Cai G, et al. An adenylate kinase OsAK3 involves brassinosteroid signaling and grain length in rice(Oryza sativa L.)[J]. Rice(NY), 2021, 14(1): 105.
[72]	Tao YJ, Miao J, Wang J, et al. RGG1, involved in the cytokinin regulatory pathway, controls grain size in rice[J]. Rice(NY), 2020, 13(1): 76.
[73]	Zhang DP, Zhang MY, Liang JS. RGB1 regulates grain development and starch accumulation through its effect on OsYUC11-mediated auxin biosynthesis in rice endosperm cells[J]. Front Plant Sci, 2021, 12: 585174. doi: 10.3389/fpls.2021.585174 URL
[74]	Huang XZ, Qian Q, Liu ZB, et al. Natural variation at the DEP1 locus enhances grain yield in rice[J]. Nat Genet, 2009, 41(4): 494-497. doi: 10.1038/ng.352
[75]	Zhang DP, Zhang MY, Zhou Y, et al. The rice G protein γ subunit DEP1/qPE9-1 positively regulates grain-filling process by increasing auxin and cytokinin content in rice grains[J]. Rice(NY), 2019, 12(1): 91.
[76]	Jin SK, Zhang MQ, Leng YJ, et al. OsNAC129 regulates seed development and plant growth and participates in the brassinosteroid signaling pathway[J]. Front Plant Sci, 2022, 13: 905148. doi: 10.3389/fpls.2022.905148 URL
[77]	Chen HY, Yu H, Jiang WZ, et al. Overexpression of ovate family protein 22 confers multiple morphological changes and represses gibberellin and brassinosteroid signalings in transgenic rice[J]. Plant Sci, 2021, 304: 110734. doi: 10.1016/j.plantsci.2020.110734 URL

植物激素 Plant hormone	基因 Gene	基因登录号 Accession No.	蛋白类型 Protein category	正/负调控 Positive（+）/ negative（-）regulator	参考文献 References
细胞分裂素 Cytokinin	OsSGL	LOC_Os02g04130	DUF1645蛋白	粒长（+）粒宽（-）	[16-17]
	DST	Os03g0786400	锌指转录因子	粒重（-）	[19]
	BG3/OsPUP4	Os01g0680200	嘌呤渗透酶	粒长（-）粒宽（-）粒重（-）	[20]
油菜素内酯Brassinolide	GS5	Os05g0158500	丝氨酸羧肽酶	粒宽（+）粒重（+）	[40]
	BRI1	Os01g0718300	受体激酶	粒宽（-）	[3]
	BAK1	Os03g0440900	BRI1相关激酶	粒长（+）粒宽（+）粒重（+）	[3]
	BZR1	Os07g0580500	BZR转录因子	粒长（+）粒宽（+）粒重（+）	[3]
	GW5	Os05g0187500	钙调素结合蛋白	粒宽（-）	[37-38]
	OFP19	Os05g0324600	OVATE家族蛋白	粒长（-）粒宽（+）	[33]
	GS2/OsGRF4	Os02g0701300	转录因子	粒长（+）粒宽（+）粒重（+）	[34]
	OsWRKY53	Os05g0343400	WRKY 转录因子	粒长（+）粒宽（+）	[30]
	GS9	Os09g0448500	转录激活因子	粒长（-）粒宽（+）	[35]
	DLT	Os06g0127600	GRAS家族蛋白	粒长（+）粒宽（-）粒重（-）	[29]
	D11	Os04g0469800	细胞色素P450蛋白	粒长（+）粒宽（+）	[24]
	D2	LOC_Os01g10040	细胞色素P450蛋白	粒长（+）粒宽（+）	[23]
	BRD1	OS03g0602300	生物合成酶	粒长（+）粒宽（+）	[21]
	BRD2	OS10g0397400	生物合成酶	粒长（+）粒宽（+）	[22]
生长素Auxin	OsARF4	Os01g0927600	生长素应答因子	粒长（-）粒宽（-）粒重（-）	[43]
	OsARF6	Os02g0164900	生长素应答因子	粒长（-）粒重（-）	[44]
	OsAUX3	Os05g0447200	生长素转运载体	粒长（-）粒重（-）	[44]
	Gnp4/LAX2	Os04g0396500	RAWUL蛋白	粒长（+）粒重（+）	[45]
	OsIAA3	Os12g0601400	Aux/IAA 蛋白	粒长（-）	[45]
	OsARF25	Os12g0613700	转录因子	粒长（+）粒宽（+）	[45]
	GSA1	Os03g0757500	UDP葡萄糖基转移酶	粒长（+）粒宽（+）粒重（+）	[48]
	TSG1	LOC_Os01g07500	色氨酸氨基转移酶	粒长（+）粒宽（+）粒重（+）	[47]
	DNR1	LOC_Os01g08270	生长素应答因子	粒重（+）	[46]
赤霉素Gibberellin	GW6	Os06g0266800	GAST 家族蛋白	粒长（+）粒宽（+）粒重（+）	[51]
	OsGASR9	Os07g0592000	GAST 家族蛋白	粒长（+）粒宽（+）粒重（+）	[52]
	OsWRKY36	Os04g0545000	WRKY转录因子	粒长（-）粒宽（-）粒重（-）	[53]
	SGD2	Os01g0643600	HD-Zip转录因子	粒长（+）粒宽（+）粒重（+）	[54]
	OsBC1	Os09g0510500	bHLH转录因子	粒长（+）粒宽（-）粒重（+）	[55]
	SNG1	Os01g0940100	己糖激酶样蛋白	粒长（+）粒宽（+）粒重（+）	[57]
乙烯Ethylene	OsERF115	Os08g0521600	ERF转录因子	粒宽（+）	[61]
茉莉酸Jasmonic acid	OsJAZ11	Os03g0180900	JAZ蛋白	粒长（+）粒宽（+）粒重（+）	[63-64]
脱落酸Abscisic acid	OsNCED3	Os03g0645900	9-顺式-环氧类胡萝卜素双加氧酶	粒长（+）粒宽（+）粒重（+）	[67]
其他调控因子Other regulators	RGB1	Os03g0669200	Gβ 亚基	粒长（+）	[72]
	qPE9-1/DEP1	Os09g0441900	Gγ亚基	粒长（+）	[74-75]
	qGL3/OsPPKL1	Os03g0646900	丝氨酸/苏氨酸磷酸酶	粒长（-）	[68-69]
	OsYUC11	Os12g0189500	黄素单加氧酶	粒重（+）	[73]

植物激素 Plant hormone	基因 Gene	基因登录号 Accession No.	蛋白类型 Protein category	正/负调控 Positive（+）/ negative（-）regulator	参考文献 References
细胞分裂素 Cytokinin	OsSGL	LOC_Os02g04130	DUF1645蛋白	粒长（+）粒宽（-）	[16-17]
	DST	Os03g0786400	锌指转录因子	粒重（-）	[19]
	BG3/OsPUP4	Os01g0680200	嘌呤渗透酶	粒长（-）粒宽（-）粒重（-）	[20]
油菜素内酯Brassinolide	GS5	Os05g0158500	丝氨酸羧肽酶	粒宽（+）粒重（+）	[40]
	BRI1	Os01g0718300	受体激酶	粒宽（-）	[3]
	BAK1	Os03g0440900	BRI1相关激酶	粒长（+）粒宽（+）粒重（+）	[3]
	BZR1	Os07g0580500	BZR转录因子	粒长（+）粒宽（+）粒重（+）	[3]
	GW5	Os05g0187500	钙调素结合蛋白	粒宽（-）	[37-38]
	OFP19	Os05g0324600	OVATE家族蛋白	粒长（-）粒宽（+）	[33]
	GS2/OsGRF4	Os02g0701300	转录因子	粒长（+）粒宽（+）粒重（+）	[34]
	OsWRKY53	Os05g0343400	WRKY 转录因子	粒长（+）粒宽（+）	[30]
	GS9	Os09g0448500	转录激活因子	粒长（-）粒宽（+）	[35]
	DLT	Os06g0127600	GRAS家族蛋白	粒长（+）粒宽（-）粒重（-）	[29]
	D11	Os04g0469800	细胞色素P450蛋白	粒长（+）粒宽（+）	[24]
	D2	LOC_Os01g10040	细胞色素P450蛋白	粒长（+）粒宽（+）	[23]
	BRD1	OS03g0602300	生物合成酶	粒长（+）粒宽（+）	[21]
	BRD2	OS10g0397400	生物合成酶	粒长（+）粒宽（+）	[22]
生长素Auxin	OsARF4	Os01g0927600	生长素应答因子	粒长（-）粒宽（-）粒重（-）	[43]
	OsARF6	Os02g0164900	生长素应答因子	粒长（-）粒重（-）	[44]
	OsAUX3	Os05g0447200	生长素转运载体	粒长（-）粒重（-）	[44]
	Gnp4/LAX2	Os04g0396500	RAWUL蛋白	粒长（+）粒重（+）	[45]
	OsIAA3	Os12g0601400	Aux/IAA 蛋白	粒长（-）	[45]
	OsARF25	Os12g0613700	转录因子	粒长（+）粒宽（+）	[45]
	GSA1	Os03g0757500	UDP葡萄糖基转移酶	粒长（+）粒宽（+）粒重（+）	[48]
	TSG1	LOC_Os01g07500	色氨酸氨基转移酶	粒长（+）粒宽（+）粒重（+）	[47]
	DNR1	LOC_Os01g08270	生长素应答因子	粒重（+）	[46]
赤霉素Gibberellin	GW6	Os06g0266800	GAST 家族蛋白	粒长（+）粒宽（+）粒重（+）	[51]
	OsGASR9	Os07g0592000	GAST 家族蛋白	粒长（+）粒宽（+）粒重（+）	[52]
	OsWRKY36	Os04g0545000	WRKY转录因子	粒长（-）粒宽（-）粒重（-）	[53]
	SGD2	Os01g0643600	HD-Zip转录因子	粒长（+）粒宽（+）粒重（+）	[54]
	OsBC1	Os09g0510500	bHLH转录因子	粒长（+）粒宽（-）粒重（+）	[55]
	SNG1	Os01g0940100	己糖激酶样蛋白	粒长（+）粒宽（+）粒重（+）	[57]
乙烯Ethylene	OsERF115	Os08g0521600	ERF转录因子	粒宽（+）	[61]
茉莉酸Jasmonic acid	OsJAZ11	Os03g0180900	JAZ蛋白	粒长（+）粒宽（+）粒重（+）	[63-64]
脱落酸Abscisic acid	OsNCED3	Os03g0645900	9-顺式-环氧类胡萝卜素双加氧酶	粒长（+）粒宽（+）粒重（+）	[67]
其他调控因子Other regulators	RGB1	Os03g0669200	Gβ 亚基	粒长（+）	[72]
	qPE9-1/DEP1	Os09g0441900	Gγ亚基	粒长（+）	[74-75]
	qGL3/OsPPKL1	Os03g0646900	丝氨酸/苏氨酸磷酸酶	粒长（-）	[68-69]
	OsYUC11	Os12g0189500	黄素单加氧酶	粒重（+）	[73]