水稻穗发育调控的分子机制研究进展

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

摘要/Abstract

摘要：

水稻是全球主要的粮食作物之一，为全球超过半数人口的基本食粮。有效穗数、穗粒数和千粒重是水稻产量的三大要素，水稻穗发育又是决定穗粒数的重要过程，并与产量的形成有着密切关系。因此，研究水稻穗发育及其分子调控机制可为水稻高产育种实践提供理论依据和指导。水稻穗发育的遗传机制和分子调控一直以来就受到育种家们的关注。本文基于国内外对水稻穗发育相关基因及其作用机理的研究进展以及近年来新发现的水稻穗发育调控基因，从水稻分生组织活性维持、花序分生组织向小穗分生组织转换、花器官形成过程和遗传调控方面展开论述。分类介绍了这些基因调控穗发育的分子机制及可能存在的相互作用关系，同时归纳了细胞分裂素、生长素、赤霉素、油菜素内酯等植物激素的信号途径中这些基因如何参与调控水稻穗发育，还进一步探讨了温度、光照、水分和营养元素等环境因素在水稻穗发育过程中的重要作用。在对水稻穗发育的分子调控机制和遗传调控网络展开全面综述的基础上，系统梳理和分析了现阶段水稻穗发育研究存在的棘手问题和相应的解决策略，并就未来水稻穗发育研究领域的关键问题和核心手段作了展望。

关键词: 水稻, 穗发育, 植物激素, 环境因素, 分子机制

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

刘园园, 陈析丰, 钱前, 高振宇. 水稻穗发育调控的分子机制研究进展[J]. 生物技术通报, 2025, 41(5): 1-13.

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.

图/表 3

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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］

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