水稻抽穗期调控的分子机理及育种应用进展

doi:10.16819/j.1001-7216.2026.250706

摘要/Abstract

摘要：

抽穗期是决定水稻生产季节与地域适应性的关键农艺性状，了解水稻抽穗期调控分子机理对于培育适应特定生态环境的高产水稻品种具有重要意义。近年来，大量水稻抽穗期相关基因相继被克隆，其功能逐渐明晰，抽穗期调控的分子网络也日趋完善。本文系统综述了以光周期调控网络为核心的水稻抽穗期调控分子机理，及其在育种实践中的应用进展，以期为水稻抽穗期遗传改良和生态适应性育种提供理论依据。

关键词: 水稻, 抽穗期, 分子机理, 育种

Abstract:

Heading date, a pivotal agronomic trait determining rice production seasons and regional adaptability, is fundamental for breeding high-yield cultivars tailored to specific ecological environments. In recent years, numerous genes controlling heading date in rice have been cloned, and their functional mechanisms have been increasingly elucidated. Consequently, the molecular regulatory network governing heading date has become progressively refined. This review systematically summarizes the molecular mechanisms regulating rice heading date, with a focus on the photoperiodic regulatory network, and discusses its applications in breeding practices. It aims to provide a theoretical foundation for the genetic improvement of heading date and ecological adaptability breeding in rice.

Key words: rice, heading date, molecular mechanism, breeding

杨大兵, 杜雪树, 李进波, 夏明元, 胡亮, 石桓, 万丙良. 水稻抽穗期调控的分子机理及育种应用进展[J]. 中国水稻科学, 2026, 40(2): 145-154.

YANG Dabing, DU Xueshu, LI Jinbo, XIA Mingyuan, HU Liang, SHI Huan, WAN Bingliang. Advances in Molecular Mechanism and Breeding Application of Heading Date Regulation in Rice[J]. Chinese Journal OF Rice Science, 2026, 40(2): 145-154.

图/表 2

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基因符号 Gene symbol	RAP位点 RAP locus	功能 Function	参考文献 Reference
LHD3/OsphyC/PHYC	Os03g0752100	光敏色素基因，与PHYB与PHYA协同感应红光和远红光。	[6]
PHYB	Os03g0309200	光敏色素基因，红光受体。调节Hd1介导的水稻成花素Hd3a的表达和临界日长。	[7]
PHYA	Os03g0719800	光敏色素基因，远红光受体。与PHYB 、PHYC协同调控水稻抽穗期。	[7]
OsCRY2	Os02g0625000	隐花色素基因，蓝光受体。	[8]
OsPRR73/OsCCT11	Os03g0284100	生物钟核心转录因子，通过结合Ehd1启动子，参与水稻昼夜节律钟的反馈回路，并连接光周期开花途径。	[53]
PCL1/OsLUX	Os01g0971800	生物钟核心元件，通过招募OsELF3-1和OsELF4s组成三元抑制蛋白复合物OsEC1，抑制Hd1和Ghd7基因表达。	[13]
OsCCA1/OsLHY/Nhd1	Os08g0157600	生物钟核心元件，通过OsGI-Hd1路径精准调控抽穗期的日长临界点，也是氮素介导的抽穗开花因子。	[9,51,54]
Hd17/OsELF3/ OsELF3-1	Os06g0142600	生物钟元件，与OsLUX和OsELF4s形成OsEC1复合物，通过结合Hd1和Ghd7启动子，抑制基因表达。	[13,55]
ELF4A	Os11g0621500	生物钟元件，与OsELF3-1和OsLUX组成三元蛋白复合物OsEC1，抑制Hd1和Ghd7的表达。	[13]
OsGI	Os01g0182600	生物钟元件，长日照条件下抑制抽穗，受三元抑制蛋白复合物OsEC1的直接调控。	[56]
Hd1	Os06g0275000	抽穗期调控核心基因，在长日照条件下，与Ghd7形成复合体共同抑制Ehd1基因的表达，从而延迟抽穗；在短日照条件下促进抽穗。	[1,15]
Ehd1	Os10g0463400	抽穗期调控核心基因，短日照条件下正向调控Hd3a/RFT1，促进抽穗。	[2]
Ghd7	Os07g0261200	抽穗期调控核心基因，每穗粒数、株高和抽穗期多效性控制基因。长日照条件下，增强表达能推迟抽穗、增加株高和每穗粒数。	[3]
Ghd8/DTH8/Hd5/ OsHAP3H/LHD1	Os08g0174500	籽粒产量、株高和抽穗期多效性控制基因，开花抑制因子。长日照条件下，DTH8协同Ghd7抑制Ehd1、Hd3a/RFT1表达，从而延迟开花时间。	[4]
Hd3a/FTL2	Os06g0157700	成花素基因，将上游光周期开花信号传递给下游开花基因。主要在短日照条件下起作用。	[20]
RFT1/FTL3	Os06g0157500	成花素基因，将上游光周期开花信号传递给下游开花基因。长日照条件下的主要开花激活因子。	[21]
OsC2DP5/OsFTIP9	Os01g0587300	成花素互作蛋白基因，介导Hd3a/RFT1从叶片向茎端分生组织(SAM)的运输。	[57]
OsFTIP1	Os06g0614000	成花素互作蛋白基因，负责将成花素RFT1从叶片向茎尖分生组织(SAM)的转运。	[25]
RIFLA/OsMADS56	Os10g0536150	MADS盒基因，与OsMADS50相互作用形成复合物，调控下游基因OsLFL1-Ehd1的表达。	[32]
OsMADS51	Os01g0922800	MADS盒基因，开花促进因子，在短日照下，作用于OsGI 下游，参与将OsGI的信号传递至Ehd1。	[18]
OsMADS14	Os03g0752800	MADS盒基因，受成花素激活复合体FAC调控，促进抽穗。	[18]
OsMADS50/OsSOC1/ DTH3	Os03g0122600	MADS盒基因，与其他MADS-box基因如OsMADS56相互作用，形成复合物，协同调控下游基因OsLFL1-Ehd1。	[58]
Ehd3	Os08g0105000	早抽穗基因，长日照下，Ehd3通过抑制Ghd7诱导水稻抽穗，同时，又能通过不依赖Ghd7的方式上调Ehd1的表达促进抽穗。	[31]
OsTOC1/ OsPRR1	Os11g0157600	转录抑制因子，作用于生物钟钟核心元件，如OsLHY和OsGI，抑制其表达。	[59]
OsRR1	Os04g0442300	A型反应调节因子，与Ehd1结合形成异二聚体，抑制Ehd1的活性，延迟开花。	[60]
OsSGI1/OsHBP1	Os04g0489600	bHLH转录因子，激活Hd1表达，促进抽穗。	[61]
OsFD1/OsbZIP77	Os09g0540800	b-ZIP转录因子，与成花素Hd3a/RFT1与14-3-3蛋白形成成花素复合物，诱导OsMADS15/OsMADS14转录，促进抽穗。	[24]
OsbZIP69/OsFD4	Os08g0549600	bZIP转录因子，与Gf14s、RFT1互作，形成成花素激活复合物，促进成花转变。	[62]
OsbZIP1/OsRE1	Os01g0174000	bZIP转录因子，抑制Ehd1表达，微调抽穗期。	[27]
OsbZIP42/HBF1	Os05g0489700	开花抑制因子，抑制Ehd1表达，延迟抽穗。	[63]
Ehd2/RID1/OsId1/ Ghd10	Os10g0419200	锌指转录因子，通过上调Ehd1的表达来促进抽穗。	[16,64]
Ghd2	Os02g0731700	控制抽穗期、株高和每穗颖花数的多效性基因，通过抑制Ehd1途径延迟抽穗。	[65]
Ehd4	Os03g0112700	CCH类锌指蛋白基因，通过上调Ehd1的表达来促进Hd3a和RFT1的表达。	[17]
OsCO3	Os09g0240200	CONSTANS基因，通过抑制Ehd1、Hd3a和RFT1的转录来负向调节开花。	[34]
DHD4	Os02g0110100	CONSTANS转录因子，与OsFD1相互作用，影响Hd3a-14-3-3-OsFD1三蛋白FAC复合体的形成，延迟抽穗。	[30]
OsCOL4	Os02g0610500	CONSTANS转录因子，组成型的开花抑制因子，作用于Ehd1上游。	[66]
OsCOL15	Os08g0536300	CONSTANS转录因子，通过上调Ghd7和下调RID1来抑制抽穗。	[67]
HD6/CK2α	Os03g0762000	酪蛋白激酶CK2α亚基，通过调控Hd1基因，在长日照条件下延迟抽穗。	[68]
Hd16/CKI/EL1	Os03g0793500	酪蛋白激酶CKI，长日条件下磷酸化Ghd7，增强其功能，延迟抽穗。	[41]
OsFTL12	Os06g0552900	FT家族蛋白，与GF14b和OsFD1相互作用，形成成花素抑制复合体FRC，调控OsMADS14和OsMADS15，延迟抽穗。	[26]
IDD4/SID1	Os02g0672100	IDD转录因子，作用于Hd3a和RFT1的启动子区域，启动水稻开花转变。	[69]
OsRIP1	Os04g0540200	OsRE1相互作用蛋白，在依赖OsRE1的方式下抑制Ehd1的转录表达调控抽穗期。	[27]
OsVIL1	Os12g0533500	与OsVIL2互作，形成PRC2复合体，短日照条件下通过抑制OsLFL1表达诱导抽穗，而长日照条件下通过提高Ghd7表达延迟抽穗。	[70]
Ehd5	Os08g0493900	WD40结构域蛋白，与Roc4和Ghd8相互作用，影响Ghd7-Ghd8复合物的形成，促进开花相关基因的表达。	[71]
DTH7/Ghd7.1/ OsPRR37/Hd2	Os07g0695100	PRR蛋白，长日条件下，抑制Ehd1表达延迟抽穗；短日条件下，根据互作基因的不同，既可抑制也可促进抽穗。	[72]
SDG724/OsSET34	Os09g0307800	组蛋白H3K36甲基转移酶，介导MADS50和RFT1的H3K36me2/3沉积，促进水稻开花。	[73]
Hd18	Os08g0143400	组蛋白乙酰转移酶基因，通过提高Ehd1的转录水平促进抽穗。	[74]
OsSUF4	Os09g0560900	锌指转录因子，促进H3K36me3对RFT1和Hd3a的修饰，从而促进抽穗。	[75]
OsHUB2/ FRRP1	Os10g0565600	E3泛素连接酶，对Ehd1的组蛋白H2B进行泛素化，抑制抽穗。	[76]
HAF1	Os04g0648800	E3泛素连接酶，作用于Hd1并将其降解，精细调控Hd1昼夜节律的积累。	[77]

基因符号 Gene symbol	RAP位点 RAP locus	功能 Function	参考文献 Reference
LHD3/OsphyC/PHYC	Os03g0752100	光敏色素基因，与PHYB与PHYA协同感应红光和远红光。	[6]
PHYB	Os03g0309200	光敏色素基因，红光受体。调节Hd1介导的水稻成花素Hd3a的表达和临界日长。	[7]
PHYA	Os03g0719800	光敏色素基因，远红光受体。与PHYB 、PHYC协同调控水稻抽穗期。	[7]
OsCRY2	Os02g0625000	隐花色素基因，蓝光受体。	[8]
OsPRR73/OsCCT11	Os03g0284100	生物钟核心转录因子，通过结合Ehd1启动子，参与水稻昼夜节律钟的反馈回路，并连接光周期开花途径。	[53]
PCL1/OsLUX	Os01g0971800	生物钟核心元件，通过招募OsELF3-1和OsELF4s组成三元抑制蛋白复合物OsEC1，抑制Hd1和Ghd7基因表达。	[13]
OsCCA1/OsLHY/Nhd1	Os08g0157600	生物钟核心元件，通过OsGI-Hd1路径精准调控抽穗期的日长临界点，也是氮素介导的抽穗开花因子。	[9,51,54]
Hd17/OsELF3/ OsELF3-1	Os06g0142600	生物钟元件，与OsLUX和OsELF4s形成OsEC1复合物，通过结合Hd1和Ghd7启动子，抑制基因表达。	[13,55]
ELF4A	Os11g0621500	生物钟元件，与OsELF3-1和OsLUX组成三元蛋白复合物OsEC1，抑制Hd1和Ghd7的表达。	[13]
OsGI	Os01g0182600	生物钟元件，长日照条件下抑制抽穗，受三元抑制蛋白复合物OsEC1的直接调控。	[56]
Hd1	Os06g0275000	抽穗期调控核心基因，在长日照条件下，与Ghd7形成复合体共同抑制Ehd1基因的表达，从而延迟抽穗；在短日照条件下促进抽穗。	[1,15]
Ehd1	Os10g0463400	抽穗期调控核心基因，短日照条件下正向调控Hd3a/RFT1，促进抽穗。	[2]
Ghd7	Os07g0261200	抽穗期调控核心基因，每穗粒数、株高和抽穗期多效性控制基因。长日照条件下，增强表达能推迟抽穗、增加株高和每穗粒数。	[3]
Ghd8/DTH8/Hd5/ OsHAP3H/LHD1	Os08g0174500	籽粒产量、株高和抽穗期多效性控制基因，开花抑制因子。长日照条件下，DTH8协同Ghd7抑制Ehd1、Hd3a/RFT1表达，从而延迟开花时间。	[4]
Hd3a/FTL2	Os06g0157700	成花素基因，将上游光周期开花信号传递给下游开花基因。主要在短日照条件下起作用。	[20]
RFT1/FTL3	Os06g0157500	成花素基因，将上游光周期开花信号传递给下游开花基因。长日照条件下的主要开花激活因子。	[21]
OsC2DP5/OsFTIP9	Os01g0587300	成花素互作蛋白基因，介导Hd3a/RFT1从叶片向茎端分生组织(SAM)的运输。	[57]
OsFTIP1	Os06g0614000	成花素互作蛋白基因，负责将成花素RFT1从叶片向茎尖分生组织(SAM)的转运。	[25]
RIFLA/OsMADS56	Os10g0536150	MADS盒基因，与OsMADS50相互作用形成复合物，调控下游基因OsLFL1-Ehd1的表达。	[32]
OsMADS51	Os01g0922800	MADS盒基因，开花促进因子，在短日照下，作用于OsGI 下游，参与将OsGI的信号传递至Ehd1。	[18]
OsMADS14	Os03g0752800	MADS盒基因，受成花素激活复合体FAC调控，促进抽穗。	[18]
OsMADS50/OsSOC1/ DTH3	Os03g0122600	MADS盒基因，与其他MADS-box基因如OsMADS56相互作用，形成复合物，协同调控下游基因OsLFL1-Ehd1。	[58]
Ehd3	Os08g0105000	早抽穗基因，长日照下，Ehd3通过抑制Ghd7诱导水稻抽穗，同时，又能通过不依赖Ghd7的方式上调Ehd1的表达促进抽穗。	[31]
OsTOC1/ OsPRR1	Os11g0157600	转录抑制因子，作用于生物钟钟核心元件，如OsLHY和OsGI，抑制其表达。	[59]
OsRR1	Os04g0442300	A型反应调节因子，与Ehd1结合形成异二聚体，抑制Ehd1的活性，延迟开花。	[60]
OsSGI1/OsHBP1	Os04g0489600	bHLH转录因子，激活Hd1表达，促进抽穗。	[61]
OsFD1/OsbZIP77	Os09g0540800	b-ZIP转录因子，与成花素Hd3a/RFT1与14-3-3蛋白形成成花素复合物，诱导OsMADS15/OsMADS14转录，促进抽穗。	[24]
OsbZIP69/OsFD4	Os08g0549600	bZIP转录因子，与Gf14s、RFT1互作，形成成花素激活复合物，促进成花转变。	[62]
OsbZIP1/OsRE1	Os01g0174000	bZIP转录因子，抑制Ehd1表达，微调抽穗期。	[27]
OsbZIP42/HBF1	Os05g0489700	开花抑制因子，抑制Ehd1表达，延迟抽穗。	[63]
Ehd2/RID1/OsId1/ Ghd10	Os10g0419200	锌指转录因子，通过上调Ehd1的表达来促进抽穗。	[16,64]
Ghd2	Os02g0731700	控制抽穗期、株高和每穗颖花数的多效性基因，通过抑制Ehd1途径延迟抽穗。	[65]
Ehd4	Os03g0112700	CCH类锌指蛋白基因，通过上调Ehd1的表达来促进Hd3a和RFT1的表达。	[17]
OsCO3	Os09g0240200	CONSTANS基因，通过抑制Ehd1、Hd3a和RFT1的转录来负向调节开花。	[34]
DHD4	Os02g0110100	CONSTANS转录因子，与OsFD1相互作用，影响Hd3a-14-3-3-OsFD1三蛋白FAC复合体的形成，延迟抽穗。	[30]
OsCOL4	Os02g0610500	CONSTANS转录因子，组成型的开花抑制因子，作用于Ehd1上游。	[66]
OsCOL15	Os08g0536300	CONSTANS转录因子，通过上调Ghd7和下调RID1来抑制抽穗。	[67]
HD6/CK2α	Os03g0762000	酪蛋白激酶CK2α亚基，通过调控Hd1基因，在长日照条件下延迟抽穗。	[68]
Hd16/CKI/EL1	Os03g0793500	酪蛋白激酶CKI，长日条件下磷酸化Ghd7，增强其功能，延迟抽穗。	[41]
OsFTL12	Os06g0552900	FT家族蛋白，与GF14b和OsFD1相互作用，形成成花素抑制复合体FRC，调控OsMADS14和OsMADS15，延迟抽穗。	[26]
IDD4/SID1	Os02g0672100	IDD转录因子，作用于Hd3a和RFT1的启动子区域，启动水稻开花转变。	[69]
OsRIP1	Os04g0540200	OsRE1相互作用蛋白，在依赖OsRE1的方式下抑制Ehd1的转录表达调控抽穗期。	[27]
OsVIL1	Os12g0533500	与OsVIL2互作，形成PRC2复合体，短日照条件下通过抑制OsLFL1表达诱导抽穗，而长日照条件下通过提高Ghd7表达延迟抽穗。	[70]
Ehd5	Os08g0493900	WD40结构域蛋白，与Roc4和Ghd8相互作用，影响Ghd7-Ghd8复合物的形成，促进开花相关基因的表达。	[71]
DTH7/Ghd7.1/ OsPRR37/Hd2	Os07g0695100	PRR蛋白，长日条件下，抑制Ehd1表达延迟抽穗；短日条件下，根据互作基因的不同，既可抑制也可促进抽穗。	[72]
SDG724/OsSET34	Os09g0307800	组蛋白H3K36甲基转移酶，介导MADS50和RFT1的H3K36me2/3沉积，促进水稻开花。	[73]
Hd18	Os08g0143400	组蛋白乙酰转移酶基因，通过提高Ehd1的转录水平促进抽穗。	[74]
OsSUF4	Os09g0560900	锌指转录因子，促进H3K36me3对RFT1和Hd3a的修饰，从而促进抽穗。	[75]
OsHUB2/ FRRP1	Os10g0565600	E3泛素连接酶，对Ehd1的组蛋白H2B进行泛素化，抑制抽穗。	[76]
HAF1	Os04g0648800	E3泛素连接酶，作用于Hd1并将其降解，精细调控Hd1昼夜节律的积累。	[77]