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Article(id=1311625838920036956, tenantId=1146029695717560320, journalId=1306273872116547672, issueId=1311625833022845462, articleNumber=null, orderNo=null, doi=10.16819/j.1001-7216.2026.251212, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=null, receivedDate=1766764800000, receivedDateStr=2025-12-27, revisedDate=1773676800000, revisedDateStr=2026-03-17, acceptedDate=null, acceptedDateStr=null, onlineDate=1790645568827, onlineDateStr=2026-09-29, pubDate=1788969600000, pubDateStr=2026-09-10, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1790645568827, onlineIssueDateStr=2026-09-29, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1790645568827, creator=13701087609, updateTime=1790645568827, updator=13701087609, issue=Issue{id=1311625833022845462, tenantId=1146029695717560320, journalId=1306273872116547672, year='2026', volume='40', issue='5', pageStart='569', pageEnd='704', issueExtLink='null', onlineDate='null', pubDate='1788969600000', pubDateStr='2026-09-10', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=1, specialIssue=null, createTime=1790645567421, creator='13701087609', updateTime=1790746157512, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1312047738519384984, tenantId=1146029695717560320, journalId=1306273872116547672, issueId=1311625833022845462, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1312047738519384985, tenantId=1146029695717560320, journalId=1306273872116547672, issueId=1311625833022845462, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=584, endPage=597, ext={EN=ArticleExt(id=1311625839188472413, articleId=1311625838920036956, tenantId=1146029695717560320, journalId=1306273872116547672, language=EN, title=Cloning and Haplotype Analysis of GSN4, a Gene for Grain Size and Grain Number in Rice, columnId=1307407385149071973, journalTitle=Chinese Journal of Rice Science, columnName=Research Papers, runingTitle=null, highlight=null, articleAbstract=
Objective

Grain size is a critical determinant of rice yield. This study aims to explore novel genes regulating grain size and expand the current genetic regulatory network, and provide valuable genetic resources for molecular design breeding of high-yield rice varieties.

Method

The large-grain mutant gsn4, generated by EMS mutagenesis of the japonica rice variety W7, was used as experimental material. A combination of systematic phenotypic identification, cytological observation, gene mapping, functional verification, subcellular localization, and haplotype analysis was performed to clone and functionally characterize the candidate gene.

Result

Phenotypic identification revealed that the gsn4 mutant exhibited pleiotropic effects. Compared with the wild type, its grain length, grain width, and 1000-grain weight were significantly increased, whereas plant height, panicle length, and grain number per panicle were significantly decreased. Cytological analysis indicated that GSN4 regulates rice grain size by modulating both cell proliferation and expansion in glumes. Map-based cloning, together with CRISPR/Cas9 gene-editing validation, confirmed LOC_Os04g46450 as the candidate gene underlying GSN4. This gene encodes a nuclear-localized E3 ubiquitin ligase protein, which acts as a negative regulator of rice grain size and modulates plant growth and development through the brassinosteroid (BR) biosynthesis pathway. Haplotype analysis further identified Hap1 as the elite haplotype conferring the most favorable effects on grain width and 1000-grain weight.

Conclusion

Collectively, this study elucidates the function and regulatory mechanism of GSN4, enriches the genetic regulatory network governing rice grain size, and provides important genetic resources and a theoretical basis for molecular design breeding of high-yield rice varieties.

, authors=Bingze CHAI1, Junge WANG1, Kaixiong WU1, Jialong LIU1, Yue WU1, Huisen WANG1, Lin TAN1, Deyong REN1, Zhenyu GAO1, Li ZHU1, Qiang ZHANG1, 2, Guangheng ZHANG1, 2, Qian QIAN1, 2, Jiang HU1, 2, *, authorsList=Bingze CHAI, Junge WANG, Kaixiong WU, Jialong LIU, Yue WU, Huisen WANG, Lin TAN, Deyong REN, Zhenyu GAO, Li ZHU, Qiang ZHANG, Guangheng ZHANG, Qian QIAN, Jiang HU, authorCompany=null, correspAuthors=Qian QIAN, Jiang HU, authorNote=null, correspAuthorsNote=

*

email:
, copyrightStatement=null, copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=null, magXml=null, pdfUrl=null, pdf=null, pdfFileSize=null, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=null, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=null, mapNumber=null, fund=null), CN=ArticleExt(id=1311625839431742046, articleId=1311625838920036956, tenantId=1146029695717560320, journalId=1306273872116547672, language=CN, title=水稻粒形和穗粒数基因GSN4的克隆与单倍型分析, columnId=1307407385430090343, journalTitle=中国水稻科学, columnName=研究报告, runingTitle=null, highlight=null, articleAbstract=
目的

粒形是决定水稻产量的重要因素之一，挖掘水稻粒形调控新基因，完善水稻粒形遗传调控网络，为高产水稻分子设计育种提供基因资源。

方法

以EMS诱变粳稻品种武运粳7号（W7）获得的大粒突变体gsn4为试验材料，系统开展表型鉴定、细胞学观察、基因定位、功能验证、亚细胞定位及单倍型分析等对候选基因进行克隆和功能分析。

结果

表型鉴定表明，gsn4突变体表现出一因多效性，与野生型相比，其粒长、粒宽及千粒重显著增加，而株高、穗长及每穗粒数显著降低；细胞学分析显示，gsn4通过影响颖壳细胞的增殖与扩张过程调控水稻籽粒大小。图位克隆结合CRISPR/Cas9基因编辑验证，明确LOC_Os04g46450为GSN4的候选基因，该基因编码核定位的E3泛素连接酶蛋白，属于水稻粒形负调控因子，通过参与油菜素内酯（BR）生物合成通路调控水稻生长发育进程。单倍型分析结果筛选出对粒宽和千粒重调控效果最优的优异单倍型Hap1。

结论

本研究明确了GSN4的功能及调控机制，丰富了水稻粒形遗传调控网络，为高产水稻品种的分子设计育种提供了重要的基因资源和理论依据。

, authors=柴炳泽1, 王俊格1, 吴凯雄1, 刘嘉龙1, 吴越1, 王荟森1, 谭霖1, 任德勇1, 高振宇1, 朱丽1, 张强1, 2, 张光恒1, 2, 钱前1, 2, *, *, 胡江1, 2, *, authorsList=柴炳泽, 王俊格, 吴凯雄, 刘嘉龙, 吴越, 王荟森, 谭霖, 任德勇, 高振宇, 朱丽, 张强, 张光恒, 钱前, 胡江, authorCompany=null, correspAuthors=钱前, 胡江, authorNote=null, correspAuthorsNote=

*

email:
, copyrightStatement=作者同意论文在本刊发表,论文的数字化复制权、发行权、汇编权及信息网络传播权即转让本刊编辑部。论文的著作权转让费在本刊稿酬中一次性支付。, copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=A7eDThXfY1MLfwU6VGQqeg==, magXml=KiRvpELqwXsT20Juz8o0/w==, pdfUrl=null, pdf=VB0m9vDiZc2pXTn1+E+Nbg==, pdfFileSize=3358943, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=null, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=CdsKyOuUkBf6cu1/uKzqtQ==, mapNumber=null, fund=null)}, authors=[Author(id=1312069797253308842, tenantId=1146029695717560320, journalId=1306273872116547672, articleId=1311625838920036956, orderNo=0, firstName=null, middleName=null, lastName=null, nameCn=null, orcid=null, stid=null, country=null, authorPic=null, dead=0, email=null, emailSecond=null, emailThird=null, correspondingAuthor=0, authorType=1, ext={EN=AuthorExt(id=1312069797333000620, tenantId=1146029695717560320, journalId=1306273872116547672, 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水稻粒形和穗粒数基因GSN4的克隆与单倍型分析
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柴炳泽 1 , 王俊格 1 , 吴凯雄 1 , 刘嘉龙 1 , 吴越 1 , 王荟森 1 , 谭霖 1 , 任德勇 1 , 高振宇 1 , 朱丽 1 , 张强 1, 2 , 张光恒 1, 2 , 钱前 1, 2, *, * , 胡江 1, 2, *
中国水稻科学 | 研究报告 2026,40(5): 584-597
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中国水稻科学 |研究报告 2026 , 40 (5) : 584 -597
水稻粒形和穗粒数基因GSN4的克隆与单倍型分析
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柴炳泽1, 王俊格1, 吴凯雄1, 刘嘉龙1, 吴越1, 王荟森1, 谭霖1, 任德勇1, 高振宇1, 朱丽1, 张强1, 2, 张光恒1, 2, 钱前1, 2, *, * , 胡江1, 2, *
作者信息
  • 1中国水稻研究所 水稻生物育种全国重点实验室，杭州 310006
  • 2三亚中国农业科学院国家南繁研究院院士工作站，三亚 572024
通讯作者:
Cloning and Haplotype Analysis of GSN4, a Gene for Grain Size and Grain Number in Rice
Bingze CHAI1, Junge WANG1, Kaixiong WU1, Jialong LIU1, Yue WU1, Huisen WANG1, Lin TAN1, Deyong REN1, Zhenyu GAO1, Li ZHU1, Qiang ZHANG1, 2, Guangheng ZHANG1, 2, Qian QIAN1, 2 , Jiang HU1, 2, *
Affiliations
  • 1 State Key Laboratory of Rice Biology and Breeding, China National Rice Research Institute, Hangzhou 310006, China
  • 2Academician Workstation, National Nanfan Research Institute (Sanya), Chinese Academy of Agricultural Sciences, Sanya 572024, China
    • Corresponding Author:

    *

    email:
出版时间: 2026-09-10 doi: 10.16819/j.1001-7216.2026.251212
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摘要
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目的

粒形是决定水稻产量的重要因素之一，挖掘水稻粒形调控新基因，完善水稻粒形遗传调控网络，为高产水稻分子设计育种提供基因资源。

方法

以EMS诱变粳稻品种武运粳7号（W7）获得的大粒突变体gsn4为试验材料，系统开展表型鉴定、细胞学观察、基因定位、功能验证、亚细胞定位及单倍型分析等对候选基因进行克隆和功能分析。

结果

表型鉴定表明，gsn4突变体表现出一因多效性，与野生型相比，其粒长、粒宽及千粒重显著增加，而株高、穗长及每穗粒数显著降低；细胞学分析显示，gsn4通过影响颖壳细胞的增殖与扩张过程调控水稻籽粒大小。图位克隆结合CRISPR/Cas9基因编辑验证，明确LOC_Os04g46450为GSN4的候选基因，该基因编码核定位的E3泛素连接酶蛋白，属于水稻粒形负调控因子，通过参与油菜素内酯（BR）生物合成通路调控水稻生长发育进程。单倍型分析结果筛选出对粒宽和千粒重调控效果最优的优异单倍型Hap1。

结论

本研究明确了GSN4的功能及调控机制，丰富了水稻粒形遗传调控网络，为高产水稻品种的分子设计育种提供了重要的基因资源和理论依据。

粒形  /  GSN4  /  细胞周期  /  细胞扩张  /  水稻  /  单倍型  /  油菜素内酯
Objective

Grain size is a critical determinant of rice yield. This study aims to explore novel genes regulating grain size and expand the current genetic regulatory network, and provide valuable genetic resources for molecular design breeding of high-yield rice varieties.

Method

The large-grain mutant gsn4, generated by EMS mutagenesis of the japonica rice variety W7, was used as experimental material. A combination of systematic phenotypic identification, cytological observation, gene mapping, functional verification, subcellular localization, and haplotype analysis was performed to clone and functionally characterize the candidate gene.

Result

Phenotypic identification revealed that the gsn4 mutant exhibited pleiotropic effects. Compared with the wild type, its grain length, grain width, and 1000-grain weight were significantly increased, whereas plant height, panicle length, and grain number per panicle were significantly decreased. Cytological analysis indicated that GSN4 regulates rice grain size by modulating both cell proliferation and expansion in glumes. Map-based cloning, together with CRISPR/Cas9 gene-editing validation, confirmed LOC_Os04g46450 as the candidate gene underlying GSN4. This gene encodes a nuclear-localized E3 ubiquitin ligase protein, which acts as a negative regulator of rice grain size and modulates plant growth and development through the brassinosteroid (BR) biosynthesis pathway. Haplotype analysis further identified Hap1 as the elite haplotype conferring the most favorable effects on grain width and 1000-grain weight.

Conclusion

Collectively, this study elucidates the function and regulatory mechanism of GSN4, enriches the genetic regulatory network governing rice grain size, and provides important genetic resources and a theoretical basis for molecular design breeding of high-yield rice varieties.

grain size  /  GSN4  /  cell cycle  /  cell expansion  /  rice  /  haplotype  /  BR
柴炳泽, 王俊格, 吴凯雄, 刘嘉龙, 吴越, 王荟森, 谭霖, 任德勇, 高振宇, 朱丽, 张强, 张光恒, 钱前, 胡江. 水稻粒形和穗粒数基因GSN4的克隆与单倍型分析. 中国水稻科学, 2026 , 40 (5) : 584 -597 . DOI: 10.16819/j.1001-7216.2026.251212
Bingze CHAI, Junge WANG, Kaixiong WU, Jialong LIU, Yue WU, Huisen WANG, Lin TAN, Deyong REN, Zhenyu GAO, Li ZHU, Qiang ZHANG, Guangheng ZHANG, Qian QIAN, Jiang HU. Cloning and Haplotype Analysis of GSN4, a Gene for Grain Size and Grain Number in Rice[J]. Chinese Journal of Rice Science, 2026 , 40 (5) : 584 -597 . DOI: 10.16819/j.1001-7216.2026.251212
基金
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  • 三亚中国农业科学院国家南繁研究院南繁专项(YBXM2526)
  • 国家自然科学基金资助项目(32272109; 32188102)
  • 海南省院士创新平台科研项目(YSPTZX202502)
  • 中国农业科学院农业科技创新计划资助项目(ASTIP; CAAS-ZDRW202401)
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文章信息
doi: 10.16819/j.1001-7216.2026.251212
  • 接收时间：2025-12-27
  • 首发时间：2026-09-29
  • 出版时间：2026-09-10
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出版历史
  • 收稿日期：2025-12-27
  • 修回日期：2026-03-17
基金
三亚中国农业科学院国家南繁研究院南繁专项(YBXM2526)
国家自然科学基金资助项目(32272109; 32188102)
海南省院士创新平台科研项目(YSPTZX202502)
中国农业科学院农业科技创新计划资助项目(ASTIP; CAAS-ZDRW202401)
作者信息
    1中国水稻研究所 水稻生物育种全国重点实验室，杭州 310006
    2三亚中国农业科学院国家南繁研究院院士工作站，三亚 572024

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https://castjournals.cast.org.cn/joweb/zgsdkx/CN/10.16819/j.1001-7216.2026.251212
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2种不同金属材料的力学参数
科
Family		 属数
Number of
genus		 种数
Number of
species		 占总种数比例
Percentage of
total species (%)		 属
Genus		 种数
Number of
species		 占总种数比例
Percentage of total
species (%)
鹅膏菌科Amanitaceae 2 11 5.26 鹅膏菌属 Amanita 10 4.78
小菇科 Mycenaceae 2 12 5.74 丝盖伞属 Inocybe 5 2.39
多孔菌科 Polyporaceae 8 14 6.70 蜡蘑属 Laccaria 5 2.39
红菇科 Russulaceae 3 23 11.00 小皮伞属 Marasmius 6 2.87
小菇属 Mycena 11 5.26
光柄菇属 Pluteus 5 2.39
红菇属 Russula 17 8.13
栓菌属 Trametes 5 2.39
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