水稻开花习性及其在粳型三系不育系选育中的应用

doi:10.16819/j.1001-7216.2025.241120

中国水稻科学 ›› 2025, Vol. 39 ›› Issue (6): 731-743.DOI: 10.16819/j.1001-7216.2025.241120

水稻开花习性及其在粳型三系不育系选育中的应用

陈玲¹^,⁵, 林文英³, 梁丽梅⁴, 欧阳由男¹, 叶胜海²^,^*(), 季芝娟¹^,⁵^,^*()

¹中国水稻研究所稻作技术研究与发展中心/水稻生物育种全国重点实验室, 杭州 311401
²浙江省农业科学院作物与核技术利用研究所, 杭州 310021
³松阳县农业技术推广中心, 浙江丽水 323400
⁴松阳县农业农村局，浙江丽水 323400
⁵中国农业科学院国家南繁研究院，海南三亚 572025

收稿日期:2024-11-29 修回日期:2025-12-25 出版日期:2025-11-10 发布日期:2025-11-19
通讯作者: * email:yesh@zaas.ac.cn;zhijuanji@126.com
基金资助:
浙江省“尖兵”“领雁”研发攻关计划资源项目(2023C04024);海南省重点研发项目(ZDYF2023XDNY086);三亚中国农业科学院国家南繁研究院“南繁专项”(YDLH2302)

Flowering Habits of Rice and Its Application in Breeding japonica Cytoplasmic Male Sterile Lines

CHEN Ling¹^,⁵, LIN Wenying³, LIANG Limei⁴, OUYANG Younan¹, YE Shenghai²^,^*(), JI Zhijuan¹^,⁵^,^*()

¹Research and Development Center of Rice Cropping Technology/State Key Laboratory for Rice Biology and Breeding, China National Rice Research Institute, Hangzhou 311401, China
²Institute of Crop and Nuclear Technology Utilization, Zhejiang Academy of Agricultural Sciences, Hangzhou 310021, China
³Songyang County Agricultural Technology Extension Center, Lishui 323400, China
⁴Agricultural and Rural Bureau of Songyang County, Lishui 323400, China
⁵National Nanfan Research Institute (Sanya), Chinese Academy of Agricultural Sciences, Sanya 572025, China

Received:2024-11-29 Revised:2025-12-25 Online:2025-11-10 Published:2025-11-19
Contact: * email:yesh@zaas.ac.cn;zhijuanji@126.com

摘要/Abstract

摘要：

杂种优势的利用大幅提高了水稻的产量。籼粳亚种间亲缘关系较远，其杂种优势通常强于亚种内杂交，因而具备更大的产量潜力，是进一步提高水稻单产的重要途径。目前，籼粳杂交稻以粳型不育系与籼型恢复系组合为主。粳不籼恢三系杂交稻因长势清秀、株型理想、不早衰以及具备超高产优势，在生产上受到了广泛关注。然而，粳型三系不育系因柱头外露率低、开花时间偏晚等开花习性上的局限，在制种过程中存在父母本花时不遇、异交结实率低等瓶颈问题，导致制种产量不高、不稳，限制了籼粳杂交稻的进一步推广应用。本文从水稻开花习性的影响因素和遗传机制等方面综述了其在粳型三系不育系选育中的应用，可为促进籼粳亚种间杂交稻的培育、生产应用和杂交水稻的可持续发展提供参考。

关键词: 籼粳杂交稻, 粳型三系不育系, 柱头外露, 开花时间, 开颖特性, 制种产量

Abstract:

The utilization of heterosis has significantly increased the yield of rice. Hybridization between indica and japonica subspecies exhibits stronger heterosis due to their distant genetic relationships. Consequently, indica-japonica hybrid rice has greater yield potential. Inter-subspecies hybridization between indica and japonica is an important way to enhance rice yields. Currently, indica-japonica hybrid rice mainly refers to the combinations derived from crossing a japonica cytoplasmic male sterility (CMS) line with an indica restorer line. These combinations have received widespread attention in production because of their vigorous growth, ideal plant architecture, resistance to premature senescence, and super high yield advantages. However, the japonica CMS line is characterized by its flowering habits, such as low stigma exsertion rate and late flowering time. These habits lead to asynchronous flowering time between the indica and japonica parental lines and low outcrossing rates in the seed production process. Thus, the yield of seed production is low and unstable, which limits further application of indica-japonica hybrid rice. In this work, we review the influencing factors and genetic mechanisms of rice flowering habits. We also summarize and analyze the application research on these mechanisms in japonica CMS lines. This review provides a reference for promoting the breeding, production, and application of indica-japonica hybrid rice and the sustainable development of hybrid rice.

Key words: indica-japonica hybrid rice, japonica CMS line, stigma exsertion, flowering time, glume opening characteristics, seed production yield

陈玲, 林文英, 梁丽梅, 欧阳由男, 叶胜海, 季芝娟. 水稻开花习性及其在粳型三系不育系选育中的应用[J]. 中国水稻科学, 2025, 39(6): 731-743.

CHEN Ling, LIN Wenying, LIANG Limei, OUYANG Younan, YE Shenghai, JI Zhijuan. Flowering Habits of Rice and Its Application in Breeding japonica Cytoplasmic Male Sterile Lines[J]. Chinese Journal OF Rice Science, 2025, 39(6): 731-743.

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关键基因/主效QTL Key gene/major QTL	基因/QTL的定位群体或研究材料 Mapping population or research material for gene/QTL	处理/研究方法 Treatment/research method	控制的性状 Controlled trait	参考文献 Reference
qTSE4	F₂(02428/ ZH464)	分子标记辅助选择 Molecular marker-assisted selection	柱头外露 Stigma exposure	[41]
GS3, GW8, GS9	单、双和三敲除的中花11突变体 Single, double and triple knockout mutants of Zhonghua 11	同时敲除 Knocking out simultaneously	柱头外露 Stigma exposure	[39]
GW5gs3	GW5、GS3转基因材料 Transgenic materials with GW5 and GS3 genes	导入基因 Gene introgression	柱头外露和花数 Stigma exposure and flower number	[42]
tg1	浙粳22物理诱变的突变体tg1 Mutant tg1 obtained by physical mutagenesis of Zhejing 22	导入基因 Gene introgression	颖花开放和柱头外露 Glumes opening and stigma exsertion	[43]
qSER-7	F₂(沪旱1B/II-32B) F₂(Huhan 1B / II-32B)	构建近等基因系 Near-isogenic line construction	柱头外露 Stigma exsertion	[22]
qSSE-4-1, qDSE-4, qTSE-4-1	RIL(贵9B/热研2号) RIL(Gui 9B / Reyan 2)	QTL定位 QTL mapping	柱头外露 Stigma exsertion	[44]
qSER-2a, qSER-2b, qSER-3a, qSER-3b	SSSLs(华粳籼74) SSSLs(Huajingxian 74)	替换作图 Substitution mapping	柱头外露 Stigma exsertion	[45]
LESR	F₂(115S柱头突变体lesr / 93S) F₂(stigma mutant lesr from 115S / 93S)	遗传分析 Genetic analysis	柱头外露 Stigma exsertion	[19]
qSTGL8.0	作图群体(IRGC110404 / IR64) Mapping population(IRGC110404/IR64)	导入QTL QTL introgression	柱头外露 Stigma exsertion	[46]
OsMYC2	中花11的OsMYC2-OE过表达系 Overexpression line of OsMYC2-OE in Zhonghua 11	过表达 Overexpression	每日开花时间 DFT	[47]
OsMYB8	天丰B和中花11 Tianfeng B and Zhonghua 11	敲除/转基因 Knocking out/ transgene	每日开花时间 DFT	[33]
OsOPR7	中花11的OsOPR7过表达系 Overexpression line of OsOPR7 in Zhonghua 11	过表达 Overexpression	每日开花时间 DFT	[48]
OsHAN1	中花11的OsHAN1过表达系和敲除系 Overexpression line and knockout line of OsHAN1 in Zhonghua 11	敲除 Knocking out	每日开花时间 DFT	[48]
FT4	典型籼粳稻及其衍生群体 Typical indica-japonica rice and its derived populations	克隆 Cloning	开花时间 Flowering time	[6]
OsFTL12	日本晴 Nipponbare	过表达 Overexpression	开花时间 Flowering time	[49]
EMF1	宜香1B背景缺失突变体emf1 Background deletion mutant emf1 of Yixiang 1B	敲除 Knocking out	开花时间 Flowering time	[50]
OsJAG^ogl	突变体ogl Mutant ogl	点突变 Point mutation	颖花开放 Glume opening	[51]
OsOAT	黄华占突变体Osoat-1和盐稻8号突变体Osoat-2 Mutant Osoat-1 of Huanghuazhan and mutant Osoat-2 of Yandao 8	辐射/基因编辑 Radiation / gene editing	颖花开放 Glume opening	[52]
EHD6	从Dongjin转基因品系后代中获得的突变体ehd6 Mutant ehd6 obtained from the offspring of Dongjin transgenic line	批量分离分析鉴定 Batch separation analysis and identification	开花 Flowering	[53]
FTL10	日本晴 Nipponbare	过表达 Overexpression	开花 Flowering	[38]
Hd16	NIL(日本晴/越光) NIL(Nipponbare/Koshihikari)	表达及生化特性分析 Expression and biochemical characteristics analysis	开花 Flowering	[54]

关键基因/主效QTL Key gene/major QTL	基因/QTL的定位群体或研究材料 Mapping population or research material for gene/QTL	处理/研究方法 Treatment/research method	控制的性状 Controlled trait	参考文献 Reference
qTSE4	F₂(02428/ ZH464)	分子标记辅助选择 Molecular marker-assisted selection	柱头外露 Stigma exposure	[41]
GS3, GW8, GS9	单、双和三敲除的中花11突变体 Single, double and triple knockout mutants of Zhonghua 11	同时敲除 Knocking out simultaneously	柱头外露 Stigma exposure	[39]
GW5gs3	GW5、GS3转基因材料 Transgenic materials with GW5 and GS3 genes	导入基因 Gene introgression	柱头外露和花数 Stigma exposure and flower number	[42]
tg1	浙粳22物理诱变的突变体tg1 Mutant tg1 obtained by physical mutagenesis of Zhejing 22	导入基因 Gene introgression	颖花开放和柱头外露 Glumes opening and stigma exsertion	[43]
qSER-7	F₂(沪旱1B/II-32B) F₂(Huhan 1B / II-32B)	构建近等基因系 Near-isogenic line construction	柱头外露 Stigma exsertion	[22]
qSSE-4-1, qDSE-4, qTSE-4-1	RIL(贵9B/热研2号) RIL(Gui 9B / Reyan 2)	QTL定位 QTL mapping	柱头外露 Stigma exsertion	[44]
qSER-2a, qSER-2b, qSER-3a, qSER-3b	SSSLs(华粳籼74) SSSLs(Huajingxian 74)	替换作图 Substitution mapping	柱头外露 Stigma exsertion	[45]
LESR	F₂(115S柱头突变体lesr / 93S) F₂(stigma mutant lesr from 115S / 93S)	遗传分析 Genetic analysis	柱头外露 Stigma exsertion	[19]
qSTGL8.0	作图群体(IRGC110404 / IR64) Mapping population(IRGC110404/IR64)	导入QTL QTL introgression	柱头外露 Stigma exsertion	[46]
OsMYC2	中花11的OsMYC2-OE过表达系 Overexpression line of OsMYC2-OE in Zhonghua 11	过表达 Overexpression	每日开花时间 DFT	[47]
OsMYB8	天丰B和中花11 Tianfeng B and Zhonghua 11	敲除/转基因 Knocking out/ transgene	每日开花时间 DFT	[33]
OsOPR7	中花11的OsOPR7过表达系 Overexpression line of OsOPR7 in Zhonghua 11	过表达 Overexpression	每日开花时间 DFT	[48]
OsHAN1	中花11的OsHAN1过表达系和敲除系 Overexpression line and knockout line of OsHAN1 in Zhonghua 11	敲除 Knocking out	每日开花时间 DFT	[48]
FT4	典型籼粳稻及其衍生群体 Typical indica-japonica rice and its derived populations	克隆 Cloning	开花时间 Flowering time	[6]
OsFTL12	日本晴 Nipponbare	过表达 Overexpression	开花时间 Flowering time	[49]
EMF1	宜香1B背景缺失突变体emf1 Background deletion mutant emf1 of Yixiang 1B	敲除 Knocking out	开花时间 Flowering time	[50]
OsJAG^ogl	突变体ogl Mutant ogl	点突变 Point mutation	颖花开放 Glume opening	[51]
OsOAT	黄华占突变体Osoat-1和盐稻8号突变体Osoat-2 Mutant Osoat-1 of Huanghuazhan and mutant Osoat-2 of Yandao 8	辐射/基因编辑 Radiation / gene editing	颖花开放 Glume opening	[52]
EHD6	从Dongjin转基因品系后代中获得的突变体ehd6 Mutant ehd6 obtained from the offspring of Dongjin transgenic line	批量分离分析鉴定 Batch separation analysis and identification	开花 Flowering	[53]
FTL10	日本晴 Nipponbare	过表达 Overexpression	开花 Flowering	[38]
Hd16	NIL(日本晴/越光) NIL(Nipponbare/Koshihikari)	表达及生化特性分析 Expression and biochemical characteristics analysis	开花 Flowering	[54]

水稻开花习性及其在粳型三系不育系选育中的应用

Flowering Habits of Rice and Its Application in Breeding japonica Cytoplasmic Male Sterile Lines

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[1]	陈明亮, 曾细华, 沈雨民, 罗世友, 胡兰香, 熊文涛, 熊焕金, 吴小燕, 肖叶青. 籼粳亚种间育性位点分型及籼粳杂交稻育性位点模式研究[J]. 中国水稻科学, 2024, 38(4): 386-396.
[2]	王亚梁, 朱德峰, 陈惠哲, 张玉屏, 向镜, 王志刚, 张义凯. 籼粳杂交稻精准条播育秧机插减氮增产的效应研究[J]. 中国水稻科学, 2021, 35(5): 495-502.
[3]	孟天瑶, 葛佳琳, 张徐彬, 韦还和, 周桂生, 戴其根. 甬优中熟籼粳杂交稻栽后植株磷素积累特征与模型分析[J]. 中国水稻科学, 2020, 34(3): 256-265.
[4]	曾研华, 张玉屏, 潘晓华, 朱德峰, 向镜, 陈惠哲, 张义凯, 曾勇军. 花后不同时段低温对籼粳杂交稻稻米品质性状的影响[J]. 中国水稻科学, 2017, 31(2): 166-174.
[5]	李威, 圣忠华, 朱子亮, 魏祥进, 石磊, 邬亚文, 唐绍清, 王建龙, 胡培松. 粳稻柱头外露率QTL定位[J]. 中国水稻科学, 2017, 31(1): 23-30.
[6]	曾研华, 张玉屏, 王亚梁, 向镜, 陈惠哲, 朱德峰. 甬优系列杂交稻组合开花期耐冷性评价[J]. 中国水稻科学, 2015, 29(3): 291-298.
[7]	宋昕蔚,林建荣,吴明国. 矮败型广亲和粳稻不育系的遗传改良及生物学特性研究[J]. 中国水稻科学, 2010, 24(6): 595-600 .
[8]	曹立勇,申宗坦. 籼粳杂交稻早熟性的研究[J]. 中国水稻科学, 1997, 11(3): 187-189 .
[9]	高勇, 滕利生, 申宗坦. 有关三系籼粳杂交稻的广亲和测验种的探讨[J]. 中国水稻科学, 1995, 9(1): 49-52 .
[10]	应存山, 章善庆. 对国际热带农业研究所收集的稻属种的柱头外露性状的研究（英文）[J]. 中国水稻科学, 1989, 3(2): 62-66 .