Genetic Basis and Breeding Applications of Grain Number per Panicle in Rice

doi:10.16819/j.1001-7216.2026.251208

Abstract

Abstract:

Rice grain number per panicle is a key component determining rice yield. Genetic studies and breeding practices have demonstrated that increasing grain number per panicle can effectively enhance grain yield per unit area. The formation of grain number per panicle involves complex physiological processes, including the differentiation and degeneration of the panicle meristem, source-sink allocation and assimilate balance, and the coordinated action of multiple phytohormone signaling pathways. Adequate assimilate supply, a well-developed vascular system, and suitable environmental conditions together constitute the physiological basis for grain number formation. To date, many genes controlling grain number per panicle have been cloned and characterized, which regulate grain number by influencing panicle architecture, meristem activity, branch differentiation, floral organ development, and hormone metabolism. More importantly, these genes form intricate regulatory networks, within which many genes exhibit pleiotropic effects on other agronomic traits, such as tiller number, grain shape, and heading date etc., resulting in trade-offs and synergistic effects. Based on the elucidation of the functional mechanisms and regulatory networks of these genes, breeders have applied grain number-related genes in rice genetic improvement through molecular design breeding, pyramiding of elite alleles, and genome editing approaches. This review systematically summarizes recent advances in the physiological and genetic bases of grain number per panicle formation in rice, outlines the application of grain number-controlling genes in rice variety improvement in China, and discusses future research directions and prospects for molecular breeding.

Key words: rice, grain number per panicle, genetic basis, functional genes, regulatory network, molecular breeding

摘要：

水稻每穗粒数是决定水稻产量的关键要素。遗传研究和育种实践表明提高穗粒数可有效提升水稻单位面积产量。穗粒数的形成涉及复杂的生理过程，包括穗分生组织的分化与退化、养分的源库分配和平衡以及多种激素信号的协同作用等。充足的同化物供应、健全的维管系统以及适宜的环境条件共同构成穗粒数形成的基础。目前已经克隆和鉴定了众多控制穗粒数的基因，这些基因之间形成了纵横交织的调控网络，通过影响穗部形态建成、分生组织活性、枝梗分化、花器官发育及激素代谢等途径调控水稻每穗粒数。控制穗粒数基因大多具有多效性，参与调控分蘖数、粒型、开花期等其他重要农艺性状，从而产生协同和拮抗效应。基于这些基因的作用机理和调控网络，通过分子设计育种聚合优良等位基因以及基因编辑等手段将穗粒数基因应用于水稻遗传改良。本文综述了水稻穗粒数形成的生理和遗传基础的最新研究进展，总结了控制穗粒数基因在我国水稻品种改良中的应用情况，并对未来的研究方向和分子育种前景进行了展望。

关键词: 水稻, 穗粒数, 遗传基础, 功能基因, 调控网络, 分子育种

LI Hongzhou, CHEN Jiale, Muhammad ZUBAIR, ZHANG Jian, YANG Longwei, YING Jiezheng. Genetic Basis and Breeding Applications of Grain Number per Panicle in Rice[J]. Chinese Journal OF Rice Science, 2026, 40(6): 726-741.

李洪洲, 陈家乐, Muhammad ZUBAIR, 张健, 杨隆维, 应杰政. 水稻穗粒数的遗传基础和育种利用[J]. 中国水稻科学, 2026, 40(6): 726-741.

Figures/Tables 2

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[86]	Lu Y, Chuan M, Wang H, Chen R, Tao T, Zhou Y, Xu Y, Li P, Yao Y, Xu C, Yang Z. Genetic and molecular factors in determining grain number per panicle of rice[J]. Frontiers in Plant Science, 2022, 13: 964246.
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基因 Gene	基因编号 Gene ID	蛋白类型 Protein type	功能要点 Functional description	文献 Reference
NOG1	LOC_Os01g54860	烯醇辅酶水合酶 enoyl-CoA hydratase	促进小穗分化并增加每穗粒数，改善脂肪酸和能量代谢 Promote spikelet differentiation and increase grain number per panicle, improve fatty acid and energy metabolism	[2,60]
Ghd7	LOC_Os07g15770	转录因子 CCT motif family protein	延迟抽穗时间，维持穗分生组织活动，显著提高穗粒数并改善维管发育 Delay heading time, maintain panicle meristem activity, significantly increase grain number per panicle and improve vascular development	[10]
NAL1	LOC_Os04g52479	调控蛋白 Regulating proteins	影响叶片和穗颈维管束结构，增强生长素极性运输，增加产量和穗粒数 Affects the structure of vascular bundles in leaves and panicle neck, enhances the polar transport of auxin, and increases yield and grain number per panicle	[11]
OsCOMT	LOC_Os08g06100	O-甲基转移酶 O-methyltransferase	提高褪黑素合成，增强维管发育并延缓衰老，增加穗粒数 Increase melatonin synthesis, enhance vascular development and delay aging, increase the number of grains per panicle	[12]
PAY1	LOC_Os08g31470	株型调控蛋白 Plant type regulatory protein	调节极性生长素运输，降低穗部生长素浓度，增加分枝数和穗粒数 Regulating polar auxin transport, reducing auxin concentration in panicle, increasing branch number and grain number per panicle	[14]
ETR2	LOC_Os04g08740	乙烯受体 Ethylene receptor	延迟营养生长向生殖生长转变，高水平乙烯导致小穗退化 Delay the transition from vegetative growth to reproductive growth, and high levels of ethylene lead to spikelet degeneration	[16]
GNP3	LOC_Os03g49640	乙烯相关转录因子 Ethylene-related transcription factors	减少乙烯合成/信号传递，提升穗粒数 Reduce ethylene synthesis/signal transmission and increase grain number per panicle	[17]
BRD3	LOC_Os06g39880	染色质识别因子 Chromatin recognition factor	靶向增强BR分解，延缓小穗退化，形成簇生小穗并提升单穗粒数 Targeted enhancement of BR decomposition, delaying spikelet degradation, forming clustered spikelets and increasing the number of grains per panicle	[19]
GNP2	LOC_Os01g10840	GSK3样激酶 GSK3-like kinase	BR信号负调控因子，增强穗分生组织发育和分枝形成，提高穗粒数 BR signal negative regulators, enhanced panicle meristem development and branching formation, increased grain number per panicle	[20]
NSP2	LOC_Os03g15680	信号转导蛋白 Signal transducin	低磷条件下促进独脚金内酯(SL)生物合成，调控株型和产量 Promote the biosynthesis of strigolactone (SL), regulate plant type and yield under low phosphorus conditions	[22]
OsJAZ11	LOC_Os03g08320	JAZ家族蛋白 ZIM domain containing protein	参与茉莉酸信号精细平衡，影响小穗及花器官发育与结实 Involved in the fine balance of jasmonic acid signals and affects the development and fruiting of spikelets and floral organs	[23,24]
OsPRMT6a	LOC_Os10g34740	精氨酸甲基转移酶 Arginine methyltransferase	维持茉莉酸(JA)响应，保障小穗正常发育并提高高温适应性 Maintaining jasmonic acid (JA) response, ensuring normal development of spikelets and improving high temperature adaptability	[23]
Gn1a (OsCKX2)	LOC_Os01g10110	细胞分裂素氧化酶 Cytokinin oxidase	降解细胞分裂素；功能缺失可提高分生组织CK含量，增加分枝和小穗 Degrading cytokinin; loss of function can increase CK content in meristem, increase branches and spikelets	[27]
LOG	LOC_Os01g40630	细胞分裂素激活酶 Cytokinin activated enzyme	负责将前体转化为活性细胞分裂素，促进穗分生组织发育 Responsible for converting precursors into active cytokinins, promoting the development of panicle meristems	[29]
An-2	LOC_Os04g43840	细胞分裂素合成酶 Cytokinin synthase	参与CK合成，控制芒长度，功能缺失导致每穗粒数降低 Participate in CK synthesis, control the length of Miscanthus sinensis, and functional deficiency leads to a reduction in the number of grains per panicle	[30]
DST (LHD1)	LOC_Os03g57240	C₂H₂锌指转录因子 C₂H₂ zinc finger transcription factor	促进Gn1a表达，降低CK含量；突变可显著增加穗粒数 Promote the expression of Gn1a and reduce CK content; mutations can significantly increase the number of grains per panicle	[31]
OsER1	LOC_Os06g10230	跨膜受体蛋白激酶 Transmembrane receptor protein kinase	促进细胞分裂素分解途径（上调CKX），负调控穗粒数 Promoting cytokinin degradation pathway (upregulating CKX), negatively regulating grain number per panicle	[32]
GSN1	LOC_Os05g02500	MAPK磷酸酶 MAPK phosphatase	使OsMPK6失活，提高CK水平，增加穗粒数（与粒型粒重拮抗） Inactivate OsMPK6, increase CK levels, increase grain number per panicle (antagonistic to grain type and weight)	[33]
OsMED25	LOC_Os09g13610	转录介体复合体亚基 Transcription mediator complex subunit	与DST互作并募集RNA聚合酶II至Gn1a启动子，负调控穗粒数 Interacts with DST and recruits RNA polymerase II to the Gn1a promoter, negatively regulating grain number per panicle	[35]
OsCYP71D8L	LOC_Os02g09220	细胞色素P450酶 Cytochrome P450 enzyme	促进GA生物合成，抑制穗的分枝伸长并降低穗粒数 Promotes GA biosynthesis, inhibits panicle branching elongation and reduces grain number	[37]
sped1-D	LOC_Os06g39650	PPR蛋白 PPR protein	缺陷导致多个GA受体下调，表现为穗轴缩短，二次枝梗变少 Defects lead to down-regulation of multiple GA receptors, manifesting as shortened panicle axis and fewer secondary rachis branches	[37]
GNP1	LOC_Os03g63970	GA-20-氧化酶 GA-20-oxidase	增加穗部GA含量并反馈上调CK水平，促进穗粒数 Increases panicle GA content and feedback upregulates CK levels, promoting grain number	[38]
DEP1 (qPE9-1)	LOC_Os09g26999	G蛋白γ亚基 G protein gamma subunit	突变导致G蛋白信号持续激活，使穗形紧凑、着粒密度显著提高 Mutation causes continuous activation of G protein signaling, resulting in compact panicle shape and significantly increased grain density	[39]
RGB1	LOC_Os03g46650	G蛋白β/α亚基 G protein beta/alpha subunit	参与异源三聚体G蛋白通路，与转录因子互作调控穗粒数与粒型 Participates in the heterotrimeric G protein pathway, interacting with transcription factors to regulate grain number and grain shape	[40,63]
RGA1	LOC_Os05g26890	G蛋白β/α亚基 G protein beta/alpha subunit	参与异源三聚体G蛋白通路，与转录因子互作调控穗粒数与粒型 Participates in the heterotrimeric G protein pathway, interacting with transcription factors to regulate grain number and grain shape	[40,63]
EP3	LOC_Os02g15950	调控蛋白 Regulatory protein	影响穗分枝角度和排列，功能缺失导致穗缩短、粒数增加 Affects panicle branching angle and arrangement, loss of function results in shortened panicle and increased grain number	[41]
DEP3	LOC_Os06g46350	含Patatin样磷脂酶域蛋白 Patatin-like protein	影响维管束发育与穗形态，突变致穗紧凑并增加粒数 Affects vascular bundle development and panicle morphology, mutation leads to compact panicle and increased grain number	[42]
SP1	LOC_Os11g12740	肽类转运蛋白 (PTR) Peptide transporter (PTR)	主要在维管束表达，促进穗轴伸长、分枝变长，增加分枝数和粒数 Mainly expressed in vascular bundles, promotes panicle axis elongation, branching lengthening, and increases branching number and grain number	[43]
MOC1	LOC_Os06g40780	GRAS转录因子 GRAS transcription factor	侧芽和侧生分生组织起始必需因子；缺失则几乎无分蘖和二次枝梗 Essential factor for lateral bud and lateral meristem initiation; loss results in almost no tillering and secondary rachis branches	[44]
LAX1	LOC_Os01g61480	bHLH转录因子 bHLH transcription factor	维持分枝分生组织的形成和持续分裂，突变减少二次枝梗和小穗 Maintains the formation and continuous division of branching meristems, mutation reduces secondary rachis branches and spikelets	[45]
LAX2 (Gnp4)	LOC_Os04g32510	核定位蛋白 Nuclear localization protein	协同LAX1调控侧生分生组织，维持穗分枝；可被GNP2磷酸化稳定 Collaborates with LAX1 to regulate lateral meristems, maintaining panicle branching; can be stabilized by GNP2 phosphorylation	[46]
PAP2	LOC_Os03g54170	MADS-box转录因子 MADS-box transcription factor	延长穗分生组织活动期，推迟花分生组织转变，增加分枝和小穗 Extends the activity period of panicle meristematic tissue, delays the transition to floral meristematic tissue, and increases branching and spikelets	[47]
FON4	LOC_Os11g38270	CLE肽类配体 CLE peptide ligand	控制穗顶端分生组织规模，限制分生组织大小以避免畸形多花小穗 Controls the scale of panicle apical meristematic tissue, limits meristematic tissue size to avoid malformed multiple-flowered spikelets	[48]
APO2 (RFL)	LOC_Os04g51000	LEAFY同源转录因子 LEAFY homologous transcription factor	与APO1互作，共同抑制穗分生组织过早进入花分化阶段 Interacts with APO1 to jointly inhibit premature entry of panicle meristematic tissue into the floral differentiation stage	[49]
LARGE2	LOC_Os12g24080	E3泛素连接酶 E3 ubiquitin ligase	专门介导APO1和APO2蛋白降解，负向调节穗分枝特性维持 Specifically mediates the degradation of APO1 and APO2 proteins, negatively regulates the maintenance of panicle branching characteristics	[50]
APO1 (SCM2)	LOC_Os06g45460	F-box蛋白 F-box protein	维持分枝分生组织特性更长时间，延缓成花，增加穗分枝和粒数 Maintains branching meristem characteristics for a longer time, delays flowering, and increases panicle branching and grain number	[51]
FZP	LOC_Os07g47330	AP2家族转录因子 AP2 family transcription factor	终止新分枝分化并促使转为花分生组织；抑制其表达可增加分枝和粒数 Terminates new branching differentiation and promotes transition to floral meristematic tissue; inhibiting its expression can increase branching and grain number	[52,54]
TAW1	LOC_Os10g33780	TCP转录因子 TCP transcription factor	延缓小穗分化、维持分枝分生组织活性，使穗分枝延长并增加枝梗 Delays spikelet differentiation, maintains branching meristem activity, extends panicle branching and increases rachis branches	[55]
RCN1	LOC_Os11g05470	TFL1同源蛋白 (PEBP) TFL1 homologous protein (PEBP)	延迟穗分生组织向花转变，增加分枝级次和粒数 Delays the transition of panicle meristematic tissue to floral tissue, increases branching order and grain number	[56]
RCN2	LOC_Os02g32950	TFL1同源蛋白 (PEBP) TFL1 homologous protein (PEBP)	延迟穗分生组织向花转变，增加分枝级次和粒数 Delays the transition of panicle meristematic tissue to floral tissue, increases branching order and grain number	[56]
MFS1	LOC_Os05g41760	AP2类转录因子 AP2-like transcription factor	控制小穗分生组织命运，突变形成一穗多粒的多花小穗表型 Controls the fate of spikelet meristematic tissue, mutation results in a multi-grain multi-flowered spikelet phenotype	[57]
MFS2	LOC_Os04g47890	MYB转录因子 MYB transcription factor	抑制小穗过早确定，功能缺失导致多器官和多花小穗 Suppresses premature determination of spikelets, loss of function leads to multi-organ and multi-flowered spikelets	[58]
OsMADS15	LOC_Os07g01820	MADS-box转录因子 MADS-box transcription factor	参与花序发育，过量表达会扰乱花器官，造成颖花败育、粒数降低 Participates in inflorescence development, overexpression disrupts floral organs, causing glume abortion and reduced grain number	[59]

基因 Gene	基因编号 Gene ID	蛋白类型 Protein type	功能要点 Functional description	文献 Reference
NOG1	LOC_Os01g54860	烯醇辅酶水合酶 enoyl-CoA hydratase	促进小穗分化并增加每穗粒数，改善脂肪酸和能量代谢 Promote spikelet differentiation and increase grain number per panicle, improve fatty acid and energy metabolism	[2,60]
Ghd7	LOC_Os07g15770	转录因子 CCT motif family protein	延迟抽穗时间，维持穗分生组织活动，显著提高穗粒数并改善维管发育 Delay heading time, maintain panicle meristem activity, significantly increase grain number per panicle and improve vascular development	[10]
NAL1	LOC_Os04g52479	调控蛋白 Regulating proteins	影响叶片和穗颈维管束结构，增强生长素极性运输，增加产量和穗粒数 Affects the structure of vascular bundles in leaves and panicle neck, enhances the polar transport of auxin, and increases yield and grain number per panicle	[11]
OsCOMT	LOC_Os08g06100	O-甲基转移酶 O-methyltransferase	提高褪黑素合成，增强维管发育并延缓衰老，增加穗粒数 Increase melatonin synthesis, enhance vascular development and delay aging, increase the number of grains per panicle	[12]
PAY1	LOC_Os08g31470	株型调控蛋白 Plant type regulatory protein	调节极性生长素运输，降低穗部生长素浓度，增加分枝数和穗粒数 Regulating polar auxin transport, reducing auxin concentration in panicle, increasing branch number and grain number per panicle	[14]
ETR2	LOC_Os04g08740	乙烯受体 Ethylene receptor	延迟营养生长向生殖生长转变，高水平乙烯导致小穗退化 Delay the transition from vegetative growth to reproductive growth, and high levels of ethylene lead to spikelet degeneration	[16]
GNP3	LOC_Os03g49640	乙烯相关转录因子 Ethylene-related transcription factors	减少乙烯合成/信号传递，提升穗粒数 Reduce ethylene synthesis/signal transmission and increase grain number per panicle	[17]
BRD3	LOC_Os06g39880	染色质识别因子 Chromatin recognition factor	靶向增强BR分解，延缓小穗退化，形成簇生小穗并提升单穗粒数 Targeted enhancement of BR decomposition, delaying spikelet degradation, forming clustered spikelets and increasing the number of grains per panicle	[19]
GNP2	LOC_Os01g10840	GSK3样激酶 GSK3-like kinase	BR信号负调控因子，增强穗分生组织发育和分枝形成，提高穗粒数 BR signal negative regulators, enhanced panicle meristem development and branching formation, increased grain number per panicle	[20]
NSP2	LOC_Os03g15680	信号转导蛋白 Signal transducin	低磷条件下促进独脚金内酯(SL)生物合成，调控株型和产量 Promote the biosynthesis of strigolactone (SL), regulate plant type and yield under low phosphorus conditions	[22]
OsJAZ11	LOC_Os03g08320	JAZ家族蛋白 ZIM domain containing protein	参与茉莉酸信号精细平衡，影响小穗及花器官发育与结实 Involved in the fine balance of jasmonic acid signals and affects the development and fruiting of spikelets and floral organs	[23,24]
OsPRMT6a	LOC_Os10g34740	精氨酸甲基转移酶 Arginine methyltransferase	维持茉莉酸(JA)响应，保障小穗正常发育并提高高温适应性 Maintaining jasmonic acid (JA) response, ensuring normal development of spikelets and improving high temperature adaptability	[23]
Gn1a (OsCKX2)	LOC_Os01g10110	细胞分裂素氧化酶 Cytokinin oxidase	降解细胞分裂素；功能缺失可提高分生组织CK含量，增加分枝和小穗 Degrading cytokinin; loss of function can increase CK content in meristem, increase branches and spikelets	[27]
LOG	LOC_Os01g40630	细胞分裂素激活酶 Cytokinin activated enzyme	负责将前体转化为活性细胞分裂素，促进穗分生组织发育 Responsible for converting precursors into active cytokinins, promoting the development of panicle meristems	[29]
An-2	LOC_Os04g43840	细胞分裂素合成酶 Cytokinin synthase	参与CK合成，控制芒长度，功能缺失导致每穗粒数降低 Participate in CK synthesis, control the length of Miscanthus sinensis, and functional deficiency leads to a reduction in the number of grains per panicle	[30]
DST (LHD1)	LOC_Os03g57240	C₂H₂锌指转录因子 C₂H₂ zinc finger transcription factor	促进Gn1a表达，降低CK含量；突变可显著增加穗粒数 Promote the expression of Gn1a and reduce CK content; mutations can significantly increase the number of grains per panicle	[31]
OsER1	LOC_Os06g10230	跨膜受体蛋白激酶 Transmembrane receptor protein kinase	促进细胞分裂素分解途径（上调CKX），负调控穗粒数 Promoting cytokinin degradation pathway (upregulating CKX), negatively regulating grain number per panicle	[32]
GSN1	LOC_Os05g02500	MAPK磷酸酶 MAPK phosphatase	使OsMPK6失活，提高CK水平，增加穗粒数（与粒型粒重拮抗） Inactivate OsMPK6, increase CK levels, increase grain number per panicle (antagonistic to grain type and weight)	[33]
OsMED25	LOC_Os09g13610	转录介体复合体亚基 Transcription mediator complex subunit	与DST互作并募集RNA聚合酶II至Gn1a启动子，负调控穗粒数 Interacts with DST and recruits RNA polymerase II to the Gn1a promoter, negatively regulating grain number per panicle	[35]
OsCYP71D8L	LOC_Os02g09220	细胞色素P450酶 Cytochrome P450 enzyme	促进GA生物合成，抑制穗的分枝伸长并降低穗粒数 Promotes GA biosynthesis, inhibits panicle branching elongation and reduces grain number	[37]
sped1-D	LOC_Os06g39650	PPR蛋白 PPR protein	缺陷导致多个GA受体下调，表现为穗轴缩短，二次枝梗变少 Defects lead to down-regulation of multiple GA receptors, manifesting as shortened panicle axis and fewer secondary rachis branches	[37]
GNP1	LOC_Os03g63970	GA-20-氧化酶 GA-20-oxidase	增加穗部GA含量并反馈上调CK水平，促进穗粒数 Increases panicle GA content and feedback upregulates CK levels, promoting grain number	[38]
DEP1 (qPE9-1)	LOC_Os09g26999	G蛋白γ亚基 G protein gamma subunit	突变导致G蛋白信号持续激活，使穗形紧凑、着粒密度显著提高 Mutation causes continuous activation of G protein signaling, resulting in compact panicle shape and significantly increased grain density	[39]
RGB1	LOC_Os03g46650	G蛋白β/α亚基 G protein beta/alpha subunit	参与异源三聚体G蛋白通路，与转录因子互作调控穗粒数与粒型 Participates in the heterotrimeric G protein pathway, interacting with transcription factors to regulate grain number and grain shape	[40,63]
RGA1	LOC_Os05g26890	G蛋白β/α亚基 G protein beta/alpha subunit	参与异源三聚体G蛋白通路，与转录因子互作调控穗粒数与粒型 Participates in the heterotrimeric G protein pathway, interacting with transcription factors to regulate grain number and grain shape	[40,63]
EP3	LOC_Os02g15950	调控蛋白 Regulatory protein	影响穗分枝角度和排列，功能缺失导致穗缩短、粒数增加 Affects panicle branching angle and arrangement, loss of function results in shortened panicle and increased grain number	[41]
DEP3	LOC_Os06g46350	含Patatin样磷脂酶域蛋白 Patatin-like protein	影响维管束发育与穗形态，突变致穗紧凑并增加粒数 Affects vascular bundle development and panicle morphology, mutation leads to compact panicle and increased grain number	[42]
SP1	LOC_Os11g12740	肽类转运蛋白 (PTR) Peptide transporter (PTR)	主要在维管束表达，促进穗轴伸长、分枝变长，增加分枝数和粒数 Mainly expressed in vascular bundles, promotes panicle axis elongation, branching lengthening, and increases branching number and grain number	[43]
MOC1	LOC_Os06g40780	GRAS转录因子 GRAS transcription factor	侧芽和侧生分生组织起始必需因子；缺失则几乎无分蘖和二次枝梗 Essential factor for lateral bud and lateral meristem initiation; loss results in almost no tillering and secondary rachis branches	[44]
LAX1	LOC_Os01g61480	bHLH转录因子 bHLH transcription factor	维持分枝分生组织的形成和持续分裂，突变减少二次枝梗和小穗 Maintains the formation and continuous division of branching meristems, mutation reduces secondary rachis branches and spikelets	[45]
LAX2 (Gnp4)	LOC_Os04g32510	核定位蛋白 Nuclear localization protein	协同LAX1调控侧生分生组织，维持穗分枝；可被GNP2磷酸化稳定 Collaborates with LAX1 to regulate lateral meristems, maintaining panicle branching; can be stabilized by GNP2 phosphorylation	[46]
PAP2	LOC_Os03g54170	MADS-box转录因子 MADS-box transcription factor	延长穗分生组织活动期，推迟花分生组织转变，增加分枝和小穗 Extends the activity period of panicle meristematic tissue, delays the transition to floral meristematic tissue, and increases branching and spikelets	[47]
FON4	LOC_Os11g38270	CLE肽类配体 CLE peptide ligand	控制穗顶端分生组织规模，限制分生组织大小以避免畸形多花小穗 Controls the scale of panicle apical meristematic tissue, limits meristematic tissue size to avoid malformed multiple-flowered spikelets	[48]
APO2 (RFL)	LOC_Os04g51000	LEAFY同源转录因子 LEAFY homologous transcription factor	与APO1互作，共同抑制穗分生组织过早进入花分化阶段 Interacts with APO1 to jointly inhibit premature entry of panicle meristematic tissue into the floral differentiation stage	[49]
LARGE2	LOC_Os12g24080	E3泛素连接酶 E3 ubiquitin ligase	专门介导APO1和APO2蛋白降解，负向调节穗分枝特性维持 Specifically mediates the degradation of APO1 and APO2 proteins, negatively regulates the maintenance of panicle branching characteristics	[50]
APO1 (SCM2)	LOC_Os06g45460	F-box蛋白 F-box protein	维持分枝分生组织特性更长时间，延缓成花，增加穗分枝和粒数 Maintains branching meristem characteristics for a longer time, delays flowering, and increases panicle branching and grain number	[51]
FZP	LOC_Os07g47330	AP2家族转录因子 AP2 family transcription factor	终止新分枝分化并促使转为花分生组织；抑制其表达可增加分枝和粒数 Terminates new branching differentiation and promotes transition to floral meristematic tissue; inhibiting its expression can increase branching and grain number	[52,54]
TAW1	LOC_Os10g33780	TCP转录因子 TCP transcription factor	延缓小穗分化、维持分枝分生组织活性，使穗分枝延长并增加枝梗 Delays spikelet differentiation, maintains branching meristem activity, extends panicle branching and increases rachis branches	[55]
RCN1	LOC_Os11g05470	TFL1同源蛋白 (PEBP) TFL1 homologous protein (PEBP)	延迟穗分生组织向花转变，增加分枝级次和粒数 Delays the transition of panicle meristematic tissue to floral tissue, increases branching order and grain number	[56]
RCN2	LOC_Os02g32950	TFL1同源蛋白 (PEBP) TFL1 homologous protein (PEBP)	延迟穗分生组织向花转变，增加分枝级次和粒数 Delays the transition of panicle meristematic tissue to floral tissue, increases branching order and grain number	[56]
MFS1	LOC_Os05g41760	AP2类转录因子 AP2-like transcription factor	控制小穗分生组织命运，突变形成一穗多粒的多花小穗表型 Controls the fate of spikelet meristematic tissue, mutation results in a multi-grain multi-flowered spikelet phenotype	[57]
MFS2	LOC_Os04g47890	MYB转录因子 MYB transcription factor	抑制小穗过早确定，功能缺失导致多器官和多花小穗 Suppresses premature determination of spikelets, loss of function leads to multi-organ and multi-flowered spikelets	[58]
OsMADS15	LOC_Os07g01820	MADS-box转录因子 MADS-box transcription factor	参与花序发育，过量表达会扰乱花器官，造成颖花败育、粒数降低 Participates in inflorescence development, overexpression disrupts floral organs, causing glume abortion and reduced grain number	[59]