Advances in Molecular Mechanisms Regulating Panicle Development in Rice

doi:10.13560/j.cnki.biotech.bull.1985.2024-1218

Abstract

Abstract:

Rice is one of staple food in the world, supplying more than half of the world’s population. Effective panicle number, grain number per panicle and 1 000-grain weight are three factors controlling rice yield. The development of rice panicle is an important procedure for number of grains per panicle, which is closely related to rice yield. Therefore, the research on rice panicle development and its underlying molecular mechanism provides foundation and theoretical basis for high-yield breeding in rice. The genetic mechanism and molecular regulation of rice panicle development have been concerned by breeders. On the basis of the research progress on genes associated with rice panicle development and their regulation mechanism by domestic and international specialists, including the newly isolated genes involved in rice panicle development in recent years, we discuss the process of meristem activity maintenance, identity transition from inflorescence meristem to spikelet meristem, flower organ formation and genetic regulation in rice. We also introduce the molecular mechanism of these genes regulating panicle development and the potential interactions. Meanwhile, we also summarize how these genes related to panicle development participate in plant hormone signaling pathways such as cytokinin, auxin, gibberellin and brassinolide. We further discuss the essential role of environmental factors, such as temperature, light, water and nutrients, in the development of rice panicle. Based on description of molecular mechanisms and genetic regulatory networks of rice panicle development, we systematically analyze the urgent issues in the study of rice panicle development and the corresponding solutions, and prospects about the key issues and avenues in this area in the future.

Key words: rice, panicle development, phytohormone, environmental factors, molecular mechanism

LIU Yuan-yuan, CHEN Xi-feng, QIAN Qian, GAO Zhen-yu. Advances in Molecular Mechanisms Regulating Panicle Development in Rice[J]. Biotechnology Bulletin, 2025, 41(5): 1-13.

Figures/Tables 3

References 83

1	Zuo JR, Li JY. Molecular genetic dissection of quantitative trait loci regulating rice grain size [J]. Annu Rev Genet, 2014, 48: 99-118.
2	Li GL, Zhang HL, Li JJ, et al. Genetic control of panicle architecture in rice [J]. Crop J, 2021, 9(3): 590-597.
3	卢寰, 时振英. 水稻穗发育的分子生物学研究进展 [J]. 植物生理学报, 2013, 49(2): 111-121.
	Lu H, Shi ZY. Molecular research progress of rice panicle development [J]. Plant Physiol J, 2013, 49(2): 111-121.
4	Ikeda K, Sunohara H, Nagato Y. Developmental course of inflorescence and spikelet in rice [J]. Breed Sci, 2004, 54(2): 147-156.
5	Itoh JI, Nonomura KI, Ikeda K, et al. Rice plant development: from zygote to spikelet [J]. Plant Cell Physiol, 2005, 46(1): 23-47.
6	Moon S, Jung KH, Lee DE, et al. The rice FON1 gene controls vegetative and reproductive development by regulating shoot apical meristem size [J]. Mol Cells, 2006, 21(1): 147-152.
7	Suzaki T, Toriba T, Fujimoto M, et al. Conservation and diversification of meristem maintenance mechanism in Oryza sativa: Function of the FLORAL ORGAN NUMBER2 gene [J]. Plant Cell Physiol, 2006, 47(12): 1591-1602.
8	Chu HW, Qian Q, Liang WQ, et al. The floral organ number4 gene encoding a putative ortholog of Arabidopsis CLAVATA3 regulates apical meristem size in rice [J]. Plant Physiol, 2006, 142(3): 1039-1052.
9	Ren DY, Xu QK, Qiu ZN, et al. FON4 prevents the multi-floret spikelet in rice [J]. Plant Biotechnol J, 2019, 17(6): 1007-1009.
10	Ohmori Y, Tanaka W, Kojima M, et al. WUSCHEL-RELATED HOMEOBOX4 is involved in meristem maintenance and is negatively regulated by the CLE gene FCP1 in rice [J]. Plant Cell, 2013, 25(1): 229-241.
11	Suzaki T, Ohneda M, Toriba T, et al. FON2 SPARE1 redundantly regulates floral meristem maintenance with FLORAL ORGAN NUMBER2 in rice [J]. PLoS Genet, 2009, 5(10): e1000693.
12	Ikeda-Kawakatsu K, Yasuno N, Oikawa T, et al. Expression level of ABERRANT PANICLE ORGANIZATION1 determines rice inflorescence form through control of cell proliferation in the meristem [J]. Plant Physiol, 2009, 150(2): 736-747.
13	Ikeda-Kawakatsu K, Maekawa M, Izawa T, et al. ABERRANT PANICLE ORGANIZATION 2/RFL, the rice ortholog of Arabidopsis LEAFY, suppresses the transition from inflorescence meristem to floral meristem through interaction with APO1 [J]. Plant J, 2012, 69(1): 168-180.
14	Huang LJ, Hua K, Xu R, et al. The LARGE2-APO1/APO2 regulatory module controls panicle size and grain number in rice [J]. Plant Cell, 2021, 33(4): 1212-1228.
15	Yoshida A, Sasao M, Yasuno N, et al. TAWAWA1, a regulator of rice inflorescence architecture, functions through the suppression of meristem phase transition [J]. Proc Natl Acad Sci USA, 2013, 110(2): 767-772.
16	Nakagawa M, Shimamoto K, Kyozuka J. Overexpression of RCN1 and RCN2, rice TERMINAL FLOWER 1/CENTRORADIALIS homologs, confers delay of phase transition and altered panicle morphology in rice [J]. Plant J, 2002, 29(6): 743-750.
17	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.
18	Li F, Liu WB, Tang JY, et al. Rice dense and erect panicle 2 is essential for determining panicle outgrowth and elongation [J]. Cell Res, 2010, 20(7): 838-849.
19	Matin MN, Kang SG. Genetic and phenotypic analysis of lax1-6, a mutant allele of LAX PANICLE1 in rice [J]. J Plant Biol, 2012, 55(1): 50-63.
20	Tabuchi H, Zhang Y, Hattori S, et al. LAX PANICLE2 of rice encodes a novel nuclear protein and regulates the formation of axillary meristems [J]. Plant Cell, 2011, 23(9): 3276-3287.
21	Wu HM, Xie DJ, Tang ZS, et al. PINOID regulates floral organ development by modulating auxin transport and interacts with MADS16 in rice [J]. Plant Biotechnol J, 2020, 18(8): 1778-1795.
22	Han ML, Lv QY, Zhang J, et al. Decreasing nitrogen assimilation under drought stress by suppressing DST-mediated activation of Nitrate Reductase 1.2 in rice [J]. Mol Plant, 2022, 15(1): 167-178.
23	Bai XF, Huang Y, Mao DH, et al. Regulatory role of FZP in the determination of panicle branching and spikelet formation in rice [J]. Sci Rep, 2016, 6: 19022.
24	Bai XF, Huang Y, Hu Y, et al. Duplication of an upstream silencer of FZP increases grain yield in rice [J]. Nat Plants, 2017, 3(11): 885-893.
25	Jiang LY, Ma X, Zhao SS, et al. The APETALA2-like transcription factor SUPERNUMERARY BRACT controls rice seed shattering and seed size [J]. Plant Cell, 2019, 31(1): 17-36.
26	Lee DY, An G. Two AP2 family genes, supernumerary bract (SNB) and Osindeterminate spikelet 1 (OsIDS1), synergistically control inflorescence architecture and floral meristem establishment in rice [J]. Plant J, 2012, 69(3): 445-461.
27	Lu JY, Wang CL, Wang HY, et al. OsMFS1/OsHOP2 complex participates in rice male and female development [J]. Front Plant Sci, 2020, 11: 518.
28	Ren DY, Yu HP, Rao YC, et al. 'Two-floret spikelet' as a novel resource has the potential to increase rice yield [J]. Plant Biotechnol J, 2018, 16(2): 351-353.
29	Kobayashi K, Maekawa M, Miyao A, et al. PANICLE PHYTOMER2 (PAP2), encoding a SEPALLATA subfamily MADS-box protein, positively controls spikelet meristem identity in rice [J]. Plant Cell Physiol, 2010, 51(1): 47-57.
30	Zhu WW, Yang L, Wu D, et al. Rice SEPALLATA genes OsMADS5 and OsMADS34 cooperate to limit inflorescence branching by repressing the TERMINAL FLOWER1-like gene RCN4 [J]. New Phytol, 2022, 233(4): 1682-1700.
31	Chung YY, Kim SR, Finkel D, et al. Early flowering and reduced apical dominance result from ectopic expression of a rice MADS box gene [J]. Plant Mol Biol, 1994, 26(2): 657-665.
32	Jeon JS, Jang S, Lee S, et al. Leafy hull sterile1 is a homeotic mutation in a rice MADS box gene affecting rice flower development [J]. Plant Cell, 2000, 12(6): 871-884.
33	Agrawal GK, Abe K, Yamazaki M, et al. Conservation of the E-function for floral organ identity in rice revealed by the analysis of tissue culture-induced loss-of-function mutants of the OsMADS1 gene [J]. Plant Mol Biol, 2005, 59(1): 125-135.
34	Wang L, Zeng XQ, Zhuang H, et al. Ectopic expression of OsMADS1 caused dwarfism and spikelet alteration in rice [J]. Plant Growth Regul, 2017, 81(3): 433-442.
35	Xiao H, Wang Y, Liu DF, et al. Functional analysis of the rice AP3 homologue OsMADS16 by RNA interference [J]. Plant Mol Biol, 2003, 52(5): 957-966.
36	Dreni L, Jacchia S, Fornara F, et al. The D-lineage MADS-box gene OsMADS13 controls ovule identity in rice [J]. Plant J, 2007, 52(4): 690-699.
37	Cui RF, Han JK, Zhao SZ, et al. Functional conservation and diversification of class E floral homeotic genes in rice (Oryza sativa) [J]. Plant J, 2010, 61(5): 767-781.
38	Pelucchi N, Fornara F, Favalli C, et al. Comparative analysis of rice MADS-box genes expressed during flower development [J]. Sex Plant Reprod, 2002, 15(3): 113-122.
39	Wang KJ, Tang D, Hong LL, et al. DEP and AFO regulate reproductive habit in rice [J]. PLoS Genet, 2010, 6(1): e1000818.
40	Sang XC, Li YF, Luo ZK, et al. CHIMERIC FLORAL ORGANS1, encoding a monocot-specific MADS box protein, regulates floral organ identity in rice[J]. Plant Physiol, 2012, 160(2): 788-807.
41	Hu Y, Wang L, Jia R, et al. Rice transcription factor MADS32 regulates floral patterning through interactions with multiple floral homeotic genes [J]. J Exp Bot, 2021, 72(7): 2434-2449.
42	Rashotte AM. The evolution of cytokinin signaling and its role in development before Angiosperms [J]. Semin Cell Dev Biol, 2021, 109: 31-38.
43	Kurakawa T, Ueda N, Maekawa M, et al. Direct control of shoot meristem activity by a cytokinin-activating enzyme [J]. Nature, 2007, 445(7128): 652-655.
44	Li GL, Xu BX, Zhang YP, et al. RGN1 controls grain number and shapes panicle architecture in rice [J]. Plant Biotechnol J, 2022, 20(1): 158-167.
45	Gu BG, Zhou TY, Luo JH, et al. An-2 encodes a cytokinin synthesis enzyme that regulates awn length and grain production in rice [J]. Mol Plant, 2015, 8(11): 1635-1650.
46	Ashikari M, Sakakibara H, Lin SY, et al. Cytokinin oxidase regulates rice grain production [J]. Science, 2005, 309(5735): 741-745.
47	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.
48	Li M, Tang D, Wang KJ, et al. Mutations in the F-box gene LARGER PANICLE improve the panicle architecture and enhance the grain yield in rice [J]. Plant Biotechnol J, 2011, 9(9): 1002-1013.
49	Huang Y, Bai XF, Luo MF, et al. Short Panicle 3 controls panicle architecture by upregulating APO2/RFL and increasing cytokinin content in rice [J]. J Integr Plant Biol, 2019, 61(9): 987-999.
50	Wu Y, Wang Y, Mi XF, et al. The QTL GNP1 encodes GA20ox1, which increases grain number and yield by increasing cytokinin activity in rice panicle meristems [J]. PLoS Genet, 2016, 12(10): e1006386.
51	Rong CY, Liu YX, Chang ZY, et al. Cytokinin oxidase/dehydrogenase family genes exhibit functional divergence and overlap in rice growth and development, especially in control of tillering [J]. J Exp Bot, 2022, 73(11): 3552-3568.
52	Guo T, Lu ZQ, Shan JX, et al. ERECTA1 acts upstream of the OsMKKK10-OsMKK4-OsMPK6 cascade to control spikelet number by regulating cytokinin metabolism in rice [J]. Plant Cell, 2020, 32(9): 2763-2779.
53	Zhang JH, Lin QB, Wang X, et al. The dense and erect panicle1-grain number associated module enhances rice yield by repressing cytokinin oxidase 2 expression [J]. Plant Cell, 2024, 37(1): koae309.
54	淳雁, 李学勇. 水稻穗型的遗传调控研究进展 [J]. 植物学报, 2017, 52(1): 19-29.
	Chun Y, Li XY. Research progress in genetic regulation of rice panicle architecture [J]. Chin Bull Bot, 2017, 52(1): 19-29.
55	Lu GW, Coneva V, Casaretto JA, et al. OsPIN5b modulates rice (Oryza sativa) plant architecture and yield by changing auxin homeostasis, transport and distribution [J]. Plant J, 2015, 83(5): 913-925.
56	Morita Y, Kyozuka J. Characterization of OsPID, the rice ortholog of PINOID, and its possible involvement in the control of polar auxin transport [J]. Plant Cell Physiol, 2007, 48(3): 540-549.
57	Zhao L, Tan LB, Zhu ZF, et al. PAY1 improves plant architecture and enhances grain yield in rice [J]. Plant J, 2015, 83(3): 528-536.
58	Tang YY, Liu HH, Guo SY, et al. OsmiR396d affects gibberellin and brassinosteroid signaling to regulate plant architecture in rice [J]. Plant Physiol, 2018, 176(1): 946-959.
59	Gao SP, Chu CC. Gibberellin metabolism and signaling: targets for improving agronomic performance of crops [J]. Plant Cell Physiol, 2020, 61(11): 1902-1911.
60	Agata A, Ando K, Ota S, et al. Diverse panicle architecture results from various combinations of Prl5/GA20ox4 and Pbl6/APO1 alleles [J]. Commun Biol, 2020, 3(1): 302.
61	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.
62	Qi WW, Sun F, Wang QJ, et al. Rice ethylene-response AP2/ERF factor OsEATB restricts internode elongation by down-regulating a gibberellin biosynthetic gene [J]. Plant Physiol, 2011, 157(1): 216-228.
63	Zou XH, Qin ZR, Zhang CY, et al. Over-expression of an S-domain receptor-like kinase extracellular domain improves panicle architecture and grain yield in rice [J]. J Exp Bot, 2015, 66(22): 7197-7209.
64	肖辉海. 水稻长穗颈隐性高秆突变体穗颈节间的细胞学观察 [J]. 西北农林科技大学学报: 自然科学版, 2008, 36(1): 131-136.
	Xiao HH. Cytological studies on uppermost internode of recessive tall stalk rice with eui gene [J]. J Northwest A F Univ Nat Sci Ed, 2008, 36(1): 131-136.
65	Hong Z, Ueguchi-Tanaka M, Shimizu-Sato S, et al. 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 [J]. Plant J, 2002, 32(4): 495-508.
66	Hong Z, Ueguchi-Tanaka M, Fujioka S, et al. The rice brassinosteroid-deficient dwarf2 mutant, defective in the rice homolog of Arabidopsis DIMINUTO/DWARF1, is rescued by the endogenously accumulated alternative bioactive brassinosteroid, dolichosterone [J]. Plant Cell, 2005, 17(8): 2243-2254.
67	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.
68	Fang N, Xu R, Huang LJ, et al. SMALL GRAIN 11 controls grain size, grain number and grain yield in rice [J]. Rice, 2016, 9(1): 64.
69	Chen XW, Zuo SM, Schwessinger B, et al. An XA21-associated kinase (OsSERK2) regulates immunity mediated by the XA21 and XA3 immune receptors [J]. Mol Plant, 2014, 7(5): 874-892.
70	Lin Y, Zhao ZG, Zhou SR, et al. Top Bending Panicle1 is involved in brassinosteroid signaling and regulates the plant architecture in rice [J]. Plant Physiol Biochem, 2017, 121: 1-13.
71	Zhang XX, Meng WJ, Liu DP, et al. Enhancing rice panicle branching and grain yield through tissue-specific brassinosteroid inhibition [J]. Science, 2024, 383(6687): eadk8838.
72	Zhang P, Zhu WW, He Y, et al. THERMOSENSITIVE BARREN PANICLE (TAP) is required for rice panicle and spikelet development at high ambient temperature [J]. New Phytol, 2023, 237(3): 855-869.
73	成勤勤. 关于水稻幼穗分化发育的研究 [D]. 扬州: 扬州大学, 2006.
	Cheng QQ. Study on the differentiation and development of young panicles in rice [D]. Yangzhou: Yangzhou University, 2006.
74	Zhang WY, Chen YJ, Wang ZQ, et al. Polyamines and ethylene in rice young panicles in response to soil drought during panicle differentiation [J]. Plant Growth Regul, 2017, 82(3): 491-503.
75	Ding CQ, You J, Chen L, et al. Nitrogen fertilizer increases spikelet number per panicle by enhancing cytokinin synthesis in rice [J]. Plant Cell Rep, 2014, 33(2): 363-371.
76	Chen Q, Tian FA, Cheng TT, et al. Translational repression of FZP mediated by CU-rich element/OsPTB interactions modulates panicle development in rice [J]. Plant J, 2022, 110(5): 1319-1331.
77	Wu LH, Hu M, Lyu SW, et al. A 48-bp deletion upstream of LIGULELESS 1 alters rice panicle architecture [J]. Crop J, 2024, 12(2): 354-363.
78	Wu XW, Liang Y, Gao HB, et al. Enhancing rice grain production by manipulating the naturally evolved cis-regulatory element-containing inverted repeat sequence of OsREM20 [J]. Mol Plant, 2021, 14(6): 997-1011.
79	Xue WY, Xing YZ, Weng XY, et al. Natural variation in Ghd7 is an important regulator of heading date and yield potential in rice [J]. Nat Genet, 2008, 40(6): 761-767.
80	Si LZ, Chen JY, Huang XH, et al. OsSPL13 controls grain size in cultivated rice [J]. Nat Genet, 2016, 48(4): 447-456.
81	Peng YL, Gao ZY, Zhang B, et al. Fine mapping and candidate gene analysis of a major QTL for panicle structure in rice [J]. Plant Cell Rep, 2014, 33(11): 1843-1850.
82	Liu EB, Liu Y, Wu GC, et al. Identification of a candidate gene for panicle length in rice (Oryza sativa L.) via association and linkage analysis [J]. Front Plant Sci, 2016, 7: 596.
83	Yang Y, Zhang Y, Li J, et al. Three QTL from Oryza meridionalis could improve panicle architecture in Asian cultivated rice [J]. Rice, 2023, 16(1): 22.

基因/QTL Gene/QTL	基因登录号 Loc number	蛋白类型 Protein type	功能 Function	参考文献 Reference
FON1	LOC_Os06g50340	富亮氨酸重复受体激酶	调节分生组织大小来控制营养和生殖发育	［6］
FON4	LOC_Os11g38270	含CLE功能域的分泌蛋白	调节水稻茎尖分生组织大小和花分生组织的确定性	［8］
FCP1	LOC_Os04g39770	含CLE结构域的分泌蛋白	调节茎尖分生组织和根顶端分生组织分生活性的维持	［10］
FOS1	LOC_Os02g21890	含CLE结构域的分泌蛋白	控制水稻花器官数目	［11］
APO1	LOC_Os06g45460	F-box蛋白	参与分生组织的调控，正调节一级枝梗数目和小穗数目	［12］
APO2	LOC_Os04g51000	拟南芥RFL同源蛋白	穗分枝形成的正调控因子，参与枝梗分生组织命运维持	［13］
TAW1	LOC_Os10g33780	ALOG家族蛋白	编码功能未知核蛋白，参与调控枝梗分生组织	［15］
DEP1	LOC_Os09g26999	异三聚体G蛋白γ亚基	参与细胞分裂调控枝梗数和穗粒数	［17］
LAX1	LOC_Os01g61480	bHLH转录因子	控制水稻腋芽原基形成和小穗枝梗原基的分化	［19］
LAX2	LOC_Os04g32510	核定位蛋白	含植物特异保守结构域，参与调控水稻腋生分生组织的形成	［20］
DST	LOC_Os03g57240	锌指转录因子	参与水稻穗部形态建成，负调控穗粒数	［22］
FZP	LOC_Os07g47330	ERF转录因子	决定穗分枝向小穗形成的转化	［23］
SNB	LOC_Os07g13170	植物特有的AP2转录因子	参与小穗分生组织向花分生组织转变及花器官模式形成	［25］
OsIDS1	LOC_Os03g60430	AP2/ERF型转录抑制因子	调控水稻小穗发育和花器官特征	［26］
OsMFS1	LOC_Os09g10850	减数分裂螺旋蛋白	主要在花药中表达，与小穗原基向花原基的转化有关	［27］
OsMADS34	LOC_Os03g54170	属于SEP亚家族转录因子	调控水稻从营养生长向生殖生长转化及花序分生组织建成	［29］
OsMADS5	LOC_Os06g06750	属于SEP亚家族转录因子	调控水稻穗发育，限制穗分枝及促进小穗分生组织特性转变	［30］
OsMADS1	LOC_Os03g11614	编码由257个氨基酸组成的蛋白	参与花器官形成并抑制小穗分生组织的逆转	［31］
OsMADS16	LOC_Os06g49840	含MADS框225个氨基酸组成的蛋白	控制水稻花器官的发育，调控浆片和雄蕊的发育	［35］
LOG	LOC_Os01g40630	细胞分裂素激活酶	与分生组织活性维持有关	［43］
An-1	LOC_Os04g28280	bHLH蛋白	调控水稻芒发育、籽粒大小和籽粒数	［45］
Gn1a	LOC_Os01g10110	降解细胞分裂素的酶	使细胞分裂素积累在花序分生组织中影响穗粒数	［46］
LP	LOC_Os02g15950	富含Kelch的F-box蛋白	参与植物组织的细胞分裂素水平的调节	［48］
SP3	LOC_Os03g55610	Dof转录因子	通过增加细胞分裂素含量控制水稻穗型	［49］
OsER1	LOC_Os06g10230	类受体蛋白激酶	负调控穗粒数	［52］
OsPIN5b	LOC_Os08g41720	PIN蛋白	改变生长素稳态、运输和分布，调控水稻株型和产量	［55］
OsPID	LOC_Os12g42020	含有激酶结构域蛋白	负调控穗粒数	［56］
PAY1	LOC_Os08g31470	包含肽酶S64结构域蛋白	通过影响生长素极性运输改变穗粒数和产量	［57］
OsGRF6	LOC_Os03g51970	生长调节因子	正调控生长素信号通路，促进花序发育，增加穗粒数	［58］
OsLSK1	LOC_Os01g47900	典型的Ⅲ型S-结构域类受体激酶	影响株高和一次枝梗数和枝梗着粒数	［63］
EUI1	LOC_Os05g40384	577个氨基酸组成的蛋白	促进细胞生长和细胞增殖来调节籽粒大小和剑叶夹角	［64］
OsBRD1	LOC_Os03g40540	BR生物合成关键酶	通过BR合成途径调控水稻穗和籽粒发育	［65］
SMG11	LOC_Os01g10040	参与BR的生物合成的细胞色素	控制水稻籽粒大小、籽粒数目和谷物产量	［68］
OsTBP1	LOC_Os08g07760	BR信号受体BRI1的激酶	调节株高、叶夹角、穗粒数和籽粒大小	［70］
OsBRD3	LOC_Os06g39880	编码BR代谢酶	参与调控二次枝梗分生组织的转变和二次枝梗的数目	［71］
TAP	LOC_Os02g18370	水稻转座酶衍生的转录因子蛋白	高温条件下能够介导调控水稻的花序维持和小穗正常发育	［72］
Ghd7	LOC_Os07g15770	含CCT结构域的转录抑制因子	能同时控制水稻穗粒数、株高和抽穗期3个性状的主效QTL	［79］

基因/QTL Gene/QTL	基因登录号 Loc number	蛋白类型 Protein type	功能 Function	参考文献 Reference
FON1	LOC_Os06g50340	富亮氨酸重复受体激酶	调节分生组织大小来控制营养和生殖发育	［6］
FON4	LOC_Os11g38270	含CLE功能域的分泌蛋白	调节水稻茎尖分生组织大小和花分生组织的确定性	［8］
FCP1	LOC_Os04g39770	含CLE结构域的分泌蛋白	调节茎尖分生组织和根顶端分生组织分生活性的维持	［10］
FOS1	LOC_Os02g21890	含CLE结构域的分泌蛋白	控制水稻花器官数目	［11］
APO1	LOC_Os06g45460	F-box蛋白	参与分生组织的调控，正调节一级枝梗数目和小穗数目	［12］
APO2	LOC_Os04g51000	拟南芥RFL同源蛋白	穗分枝形成的正调控因子，参与枝梗分生组织命运维持	［13］
TAW1	LOC_Os10g33780	ALOG家族蛋白	编码功能未知核蛋白，参与调控枝梗分生组织	［15］
DEP1	LOC_Os09g26999	异三聚体G蛋白γ亚基	参与细胞分裂调控枝梗数和穗粒数	［17］
LAX1	LOC_Os01g61480	bHLH转录因子	控制水稻腋芽原基形成和小穗枝梗原基的分化	［19］
LAX2	LOC_Os04g32510	核定位蛋白	含植物特异保守结构域，参与调控水稻腋生分生组织的形成	［20］
DST	LOC_Os03g57240	锌指转录因子	参与水稻穗部形态建成，负调控穗粒数	［22］
FZP	LOC_Os07g47330	ERF转录因子	决定穗分枝向小穗形成的转化	［23］
SNB	LOC_Os07g13170	植物特有的AP2转录因子	参与小穗分生组织向花分生组织转变及花器官模式形成	［25］
OsIDS1	LOC_Os03g60430	AP2/ERF型转录抑制因子	调控水稻小穗发育和花器官特征	［26］
OsMFS1	LOC_Os09g10850	减数分裂螺旋蛋白	主要在花药中表达，与小穗原基向花原基的转化有关	［27］
OsMADS34	LOC_Os03g54170	属于SEP亚家族转录因子	调控水稻从营养生长向生殖生长转化及花序分生组织建成	［29］
OsMADS5	LOC_Os06g06750	属于SEP亚家族转录因子	调控水稻穗发育，限制穗分枝及促进小穗分生组织特性转变	［30］
OsMADS1	LOC_Os03g11614	编码由257个氨基酸组成的蛋白	参与花器官形成并抑制小穗分生组织的逆转	［31］
OsMADS16	LOC_Os06g49840	含MADS框225个氨基酸组成的蛋白	控制水稻花器官的发育，调控浆片和雄蕊的发育	［35］
LOG	LOC_Os01g40630	细胞分裂素激活酶	与分生组织活性维持有关	［43］
An-1	LOC_Os04g28280	bHLH蛋白	调控水稻芒发育、籽粒大小和籽粒数	［45］
Gn1a	LOC_Os01g10110	降解细胞分裂素的酶	使细胞分裂素积累在花序分生组织中影响穗粒数	［46］
LP	LOC_Os02g15950	富含Kelch的F-box蛋白	参与植物组织的细胞分裂素水平的调节	［48］
SP3	LOC_Os03g55610	Dof转录因子	通过增加细胞分裂素含量控制水稻穗型	［49］
OsER1	LOC_Os06g10230	类受体蛋白激酶	负调控穗粒数	［52］
OsPIN5b	LOC_Os08g41720	PIN蛋白	改变生长素稳态、运输和分布，调控水稻株型和产量	［55］
OsPID	LOC_Os12g42020	含有激酶结构域蛋白	负调控穗粒数	［56］
PAY1	LOC_Os08g31470	包含肽酶S64结构域蛋白	通过影响生长素极性运输改变穗粒数和产量	［57］
OsGRF6	LOC_Os03g51970	生长调节因子	正调控生长素信号通路，促进花序发育，增加穗粒数	［58］
OsLSK1	LOC_Os01g47900	典型的Ⅲ型S-结构域类受体激酶	影响株高和一次枝梗数和枝梗着粒数	［63］
EUI1	LOC_Os05g40384	577个氨基酸组成的蛋白	促进细胞生长和细胞增殖来调节籽粒大小和剑叶夹角	［64］
OsBRD1	LOC_Os03g40540	BR生物合成关键酶	通过BR合成途径调控水稻穗和籽粒发育	［65］
SMG11	LOC_Os01g10040	参与BR的生物合成的细胞色素	控制水稻籽粒大小、籽粒数目和谷物产量	［68］
OsTBP1	LOC_Os08g07760	BR信号受体BRI1的激酶	调节株高、叶夹角、穗粒数和籽粒大小	［70］
OsBRD3	LOC_Os06g39880	编码BR代谢酶	参与调控二次枝梗分生组织的转变和二次枝梗的数目	［71］
TAP	LOC_Os02g18370	水稻转座酶衍生的转录因子蛋白	高温条件下能够介导调控水稻的花序维持和小穗正常发育	［72］
Ghd7	LOC_Os07g15770	含CCT结构域的转录抑制因子	能同时控制水稻穗粒数、株高和抽穗期3个性状的主效QTL	［79］

[1]	DU Liang-heng, TANG Huang-lei, ZHANG Zhi-guo. Map-based Cloning of Light-responsive Gene ELM1 in Rice [J]. Biotechnology Bulletin, 2025, 41(5): 82-89.
[2]	CHEN Xiao-jun, HUI Jian, MA Hong-wen, BAI Hai-Bo, ZHONG Nan, LI Jia-run, FAN Yun-fang. Creating Rice Gerplasm Resources OsALS Rsistant to Herbicide through Single Base Gene Editing Technology [J]. Biotechnology Bulletin, 2025, 41(4): 106-114.
[3]	LI Xin-peng, ZHANG Wu-han, ZHANG Li, SHU Fu, HE Qiang, GUO Yang, DENG Hua-feng, WANG Yue, SUN Ping-yong. Creation of Rice Mutant by Gamma-ray and Its Molecular Identification [J]. Biotechnology Bulletin, 2025, 41(3): 35-43.
[4]	KUANG Jian-hua, CHENG Zhi-peng, ZHAO Yong-jing, YANG Jie, CHEN Run-qiao, CHEN Long-qing, HU Hui-zhen. Expression Analysis of the GH3 Gene Family in Nelumbo nucifera underHormonal and Abiotic Stresses [J]. Biotechnology Bulletin, 2025, 41(2): 221-233.
[5]	FANG Hui-min, GU Yi-shu, ZHANG Jing, ZHANG Long. Isolation and Physicochemical Properties Analysis of Starch from Rice Leaves [J]. Biotechnology Bulletin, 2025, 41(2): 51-57.
[6]	JIN Su-kui, GUO Qian-qian, LIU Qiao-quan, GAO Ji-ping. A Simplified Method for Extracting Genomic DNA from Rice Leaves [J]. Biotechnology Bulletin, 2025, 41(1): 74-84.
[7]	LIU Wen-zhi, HE Dan, LI Peng, FU Ying-lin, ZHANG Yi-xin, WEN Hua-jie, YU Wen-qing. Paenibacillus polymyxa New Strain X-11 and Its Growth-promoting Effects on Tomato and Rice [J]. Biotechnology Bulletin, 2024, 40(9): 249-259.
[8]	LI Qing-mao, PENG Cong-gui, QI Xiao-han, LIU Xing-lei, LI Zhen-yuan, LI Qin-yan, HUANG Li-yu. Screening and Identification of Excellent Strains of Endophytic Bacteria Promoting Rice Iron Absorption from Wild Rice [J]. Biotechnology Bulletin, 2024, 40(8): 255-263.
[9]	SUN Zhi-yong, DU Huai-dong, LIU Yang, MA Jia-xin, YU Xue-ran, MA Wei, YAO Xin-jie, WANG Min, LI Pei-fu. Genome-wide Association Analysis of γ-aminobutyric Acid in Rice Grains [J]. Biotechnology Bulletin, 2024, 40(8): 53-62.
[10]	DU Zhong-yang, YANG Ze, LIANG Meng-jing, LIU Yi-zhen, CUI Hong-li, SHI Da-ming, XUE Jin-ai, SUN Yan, ZHANG Chun-hui, JI Chun-li, LI Run-zhi. Effect of Nano-selenium（SeNPs）in Alleviating Lead Stress and Promoting Growth of Tobacco Seedlings [J]. Biotechnology Bulletin, 2024, 40(7): 183-196.
[11]	PANG Meng-zhen, XU Han-qin, LIU Hai-yan, SONG Juan, WANG Jia-han, SUN Li-na, JI Pei-mei, YIN Ze-zhi, HU You-chuan, ZHAO Xiao-meng, LIANG Shan-shan, ZHANG Si-ju, LUAN Wei-jiang. Gene Identification and Functional Analysis of Yellowish and Early Heading Mutant hz1 in Rice [J]. Biotechnology Bulletin, 2024, 40(7): 125-136.
[12]	TIAN Sheng-ni, ZHANG Qin, DONG Yu-fei, DING Zhou, YE Ai-hua, ZHANG Ming-zhu. Effects of Acid Mine Drainage on Physicochemical Factors and Nitrogen-fixing Microorganisms in the Root Zone of Mature Rice [J]. Biotechnology Bulletin, 2024, 40(6): 271-280.
[13]	LIU Rong, TIAN Min-yu, LI Guang-ze, TAN Cheng-fang, RUAN Ying, LIU Chun-lin. Identification and Induced-expression Analysis of REVEILLE Family in Brassica napus L. [J]. Biotechnology Bulletin, 2024, 40(6): 161-171.
[14]	KONG De-ting, QI Xiao-han, LIU Xing-lei, LI Li-ping, HU Feng-yi, HUANG Li-yu, QIN Shi-wen. Comparison and Analysis of Endophytic Bacterial Communities in Different Perennial Rice Varieties [J]. Biotechnology Bulletin, 2024, 40(5): 225-236.
[15]	YANG Qi, WEI Zi-di, SONG Juan, TONG Kun, YANG Liu, WANG Jia-han, LIU Hai-yan, LUAN Wei-jiang, MA Xuan. Construction and Transcriptomic Analysis of Rice Histone H1 Triple Mutant [J]. Biotechnology Bulletin, 2024, 40(4): 85-96.