水稻颖壳不闭合基因SG5的鉴定与克隆

doi:10.16819/j.1001-7216.2026.250310

中国水稻科学 ›› 2026, Vol. 40 ›› Issue (2): 210-222.DOI: 10.16819/j.1001-7216.2026.250310

水稻颖壳不闭合基因SG5的鉴定与克隆

薛炮¹^,², 王友霜¹, 何弯弯¹^,², 黄晨博³, 张涵³, 丁震乾¹, 陈秋丽¹, 范运新¹, 丁成伟¹, 孙廉平³^,^*(), 胡婷婷¹^,²^,^*()

¹江苏徐淮地区徐州农业科学研究所, 江苏徐州 221131
²生物育种钟山实验室, 南京 210014
³中国水稻研究所国家水稻改良中心/水稻生物育种全国重点实验室, 杭州 311401

收稿日期:2025-03-12 修回日期:2025-06-05 出版日期:2026-03-10 发布日期:2026-03-17
通讯作者: * email: htt713@163.com; sunlianping@caas.cn
基金资助:
徐州市基础研究计划-青年科技人才项目(KC23042);生物育种钟山实验室项目(ZSBBL-KY2023-01-04);徐州市农业科学院科研基金项目(高层次人才类)(GC2023004)

Identification and Cloning of SG5 in Rice

XUE Pao¹^,², WANG Youshuang¹, HE Wanwan¹^,², HUANG Chenbo³, ZHANG Han³, DING Zhenqian¹, CHEN Qiuli¹, FAN Yunxin¹, DING Chengwei¹, SUN Lianping³^,^*(), HU Tingting¹^,²^,^*()

¹Xuzhou Agricultural Sciences Research Institute in Jiangsu Xuhuai Area, Xuzhou 221131, China
²Zhongshan Biological Breeding Laboratory, Nanjing 210014, China
³State Key Laboratory of Rice Biology and Breeding/National Center of Rice Improvement, China National Rice Research Institute, Hangzhou 311401, China

Received:2025-03-12 Revised:2025-06-05 Online:2026-03-10 Published:2026-03-17
Contact: * email: htt713@163.com; sunlianping@caas.cn

摘要/Abstract

摘要：

【目的】水稻颖壳发育直接影响稻米产量与品质，挖掘颖壳发育基因并探究其分子功能，可为水稻优质高产遗传改良提供更多的基因资源。【方法】利用EMS诱变徐稻3号获得一个颖壳开裂后不闭合的突变体sg5 (split glume 5)；对其农艺性状和不同开花时间进行浆片表型和细胞学观察；通过图位克隆与BSA-seq相结合的方法鉴定SG5基因；结合野生型各组织表达量检测及GUS转基因株系各组织染色情况，分析SG5组织表达特异性；利用激光共聚焦显微镜观察SG5蛋白在原生质体中的亚细胞定位；通过进化树分析，探讨了SG5在进化过程中的选择情况。【结果】突变体sg5在开花前颖壳发育正常，开花后颖壳保持开裂状态直至成熟。与野生型相比，突变体株高和有效穗数无显著变化，抽穗期显著提前，穗长增加，粒长、粒宽降低，结实率极显著降低。体视显微镜观察结果显示，突变体颖壳开裂是浆片保持膨大状态导致的。SG5基因定位在第5染色体InDel5- 11−InDel5-13区间内，物理间距为3.72 Mb，图位克隆表明sg5突变体中LOC_Os05g50890的第3内含子和第4外显子连接处发生G-A突变导致发生可变剪接，蛋白翻译提前终止。敲除实验证实SG5基因即为目的基因。SG5基因为组成型表达，在根、茎、叶中表达水平较高。SG5蛋白定位在细胞质和细胞核中。SG5在进化过程中发生籼稻、粳稻分化。【结论】本研究鉴定了OsJar1基因一个新的等位突变，发现SG5在自然选择过程中发生了籼、粳分化。本研究为后续深入研究颖壳调控分子机制提供了新的线索，并为水稻设计育种提供新的基因资源。

关键词: 水稻, 颖壳开裂, 浆片, sg5突变体

Abstract:

【Objective】 Rice glume development directly affects both grain yield and quality. Identifying genes that control glume development and characterizing their functions can provide valuable genetic resources for breeding. 【Method】 The split glume 5 (sg5) mutant was isolated from a mutant library generated by ethyl methanesulfonate (EMS) mutagenesis of the rice variety Xudao 3 (XU3). Phenotypic investigation and analysis were performed on both the wild type and the sg5 mutant. SG5 was identified using a combination of map-based cloning and bulked segregant analysis sequencing (BSA-seq). The temporal and spatial expression patterns of SG5 were examined by qRT PCR and GUS staining. Subcellular localization of SG5 was observed in rice protoplasts. An evolutionary tree was constructed to analyze the natural selection of SG5. 【Result】 The mutant exhibited normal glume development before flowering, but the glumes remained split until maturity. Compared with the wild type, the mutant showed earlier heading, longer panicles, reduced grain length and width, and a significantly lower seed-setting rate, while plant height and effective panicle number remained unchanged. Microscopic observation indicated that the split glume phenotype was caused by persistent swelling of the lodicules. SG5 was mapped to a 3.72 Mb physical interval between InDel5-11 and InDel5-13 on chromosome 5. In the sg5 mutant, a G-to-A mutation at the junction of intron 3 and exon 4 of LOC_Os05g50890 led to alternative splicing and premature protein termination. Knockout experiments confirmed that SG5 is the target gene. SG5 is constitutively expressed, with higher transcript levels in roots, stems, and leaves. The SG5 protein localizes to both the cytoplasm and nucleus. Evolutionary analysis revealed that SG5 has undergone differentiation between indica and japonica subspecies during evolution. 【Conclusion】 This study identifies a novel allele of OsJar1 and reveals that SG5 has diverged between indica and japonica under natural selection. These findings provide new clues for further dissection of the molecular regulatory network controlling glume development and offer valuable genetic resources for precision breeding in rice.

Key words: rice, glume cracking, lodicule, sg5 mutant

薛炮, 王友霜, 何弯弯, 黄晨博, 张涵, 丁震乾, 陈秋丽, 范运新, 丁成伟, 孙廉平, 胡婷婷. 水稻颖壳不闭合基因SG5的鉴定与克隆[J]. 中国水稻科学, 2026, 40(2): 210-222.

XUE Pao, WANG Youshuang, HE Wanwan, HUANG Chenbo, ZHANG Han, DING Zhenqian, CHEN Qiuli, FAN Yunxin, DING Chengwei, SUN Lianping, HU Tingting. Identification and Cloning of SG5 in Rice[J]. Chinese Journal OF Rice Science, 2026, 40(2): 210-222.

图/表 7

表1 本研究所用的引物序列

Table 1. Primer sequences used in this study

引物名称 Primer name	引物序列（5'‎→3'） Primer sequence (5'→3')	功能 Function
InDel 5-11F	GGAGTTCTGGTGGCTTTCC	基因定位 Gene mapping
InDel 5-11R	CAGTTCCCCATTTCCCTCTA
InDel 5-13F	TGAGTTTCCGGTGTTCCATA
InDel 5-13R	AAGGCAAAGTCGTTCAGCTT
SG5-Cas9-F	ggcaATATTGAGCCCTATATCCAG	基因编辑 Gene editing
SG5- Cas9-R	aaacCTGGATATAGGGCTCAATAT	基因编辑 Gene editing
SG5-1305-GUS-F	catgattacgaattcATCACCTCTGCCCAAAACCA	GUS载体构建 Construction of GUS vector
SG5-1305-GUS-R	tcagatctaccatggTGTAATGGTAGAATCCTGGCTTTACCA	GUS载体构建 Construction of GUS vector
SG5-1305-GFP-F	aagtccggagctagctctagaATGACGATCTGCAGCTGTGAGG	亚细胞定位 Subcellular localization
SG5-1305-GFP-R	ggtcctcgagacgtctctagaAAATCCATAGGCAGTACTGAAATAACTTTGGG	亚细胞定位 Subcellular localization
Hyg-F	ACGGTGTCGTCCATCACAGTTTGCC	潮霉素基因检测 Detection of hygromycin resistance gene
Hyg-R	TTCCGGAAGTGCTTGACATTGGGGA	潮霉素基因检测 Detection of hygromycin resistance gene
Cas9-F	CGTGGAAGATCGGTTCAACGC	阳性株检测 Sequencing of positive plants
Cas9-R	CTGCCGGCCAGATTGGCA	阳性株检测 Sequencing of positive plants
CR-SG5-F	TGATGCCAACTCCTACTGCAC	突变位点检测 Mutation site detection
CR-SG5-R	ACTTGGGCTTTCCGTGTGTT	突变位点检测 Mutation site detection
UBQ10-qPCR-F	TGGTCAGTAATCAGCCAGTTTGG	qRT-PCR
UBQ10-qPCR-R	GCACCACAAATACTTGACGAACAG
SG5-qPCR-F	GATACCTCACCAGTGGTCACTG
SG5-qPCR-R	TCCTAAACGCGTAAGATGTACGG

图1 野生型徐稻3号与sg5表型特征 A: 抽穗时野生型及突变体单株表型，比例尺=20 cm；B: 野生型及突变体颖壳及穗部形态，比例尺=5 cm；C: 野生型及突变体籽粒性状，比例尺=1 cm；D: 抽穗期、株高、有效穗数、穗长、结实率、粒长、粒宽表型差异。数据表示为平均值±标准差（n≥20），学生氏t-检验计算差异显著性。*突变体与野生型间的表型差异在P < 0.05水平上显著；**突变体与野生型间的表型差异在P < 0.01水平上极显著。

Fig. 1. Phenotypic characterization of wildtype (Xudao 3, XU3) and sg5 A, Characterization of plant type in wild type and sg5. Scale bar=20 cm. B, Glume morphology and panicle morphology in wild type and sg5. Scale bar=5 cm. C, Comparison of grain size in wild type and sg5. Scale bar=10 mm. D, Comparison of heading date, plant height, panicle number, panicle length, seed setting rate, grain length and grain width in wild type and sg5. Data are given as Mean ± SD (n≥20). Student’s t-test was used to generate P value. *, P < 0.05; **, P < 0.01.

图2 野生型（XU3）和sg5在开花前后不同时间颖壳及浆片的发育形态变化野生型及突变体未开花前穗部状态(A)、颖壳形态(B)、开花后穗部状态(C)和开花后颖壳形态(D)，比例尺=5 cm；E: 野生型及突变体前期长势一致的颖花，在开花前后不同时间的颖壳及浆片形态变化，比例尺=20μm。白框中是相对应的颖花。

Fig. 2. Developmental morphological changes of glumes and lodicules in wild type (XU3) and sg5 at various time points around flowering A, Characterization of panicle before flowering(A), glume before flowering(B), panicle after flowering(C), glume after flowering(D) in the wild type and sg5(Scale bar=5 cm). E, Florets of consistent early growth from wild type and sg5, showing morphological changes of glumes and lodicules at various time points around flowering. Scale bar = 20 μm. The white boxes indicate the corresponding florets.

图3 SG5图位克隆及功能验证 A：SG5图位克隆；B: 野生型、突变体DNA序列及蛋白序列分析；下划线表示野生型SG5第4外显子前端序列，其他序列代表第3内含子后半部分；虚线红框表示剪接突变位点及新的剪接位点。黑色竖线表示茉莉酮酸酯位点；绿色竖线表示L-α-氨基酸位点。C：SG5基因编辑及纯合突变株序列；D: 野生型、突变体、SG5#CR-1、SG5#CR-2抽穗时单株表型，比例尺=20 cm, E: 野生型、突变体、SG5#CR-1、SG5#CR-2受精后颖壳形态；F: 野生型、突变体、SG5#CR-1、SG5#CR-2成熟期穗部形态，比例尺=5 cm；G：野生型、突变体、SG5#CR-1每穗实粒数；红点：正常籽粒；黑点：发霉籽粒。H：抽穗期、株高、有效穗数、穗长、结实率、粒长、粒宽表型差异。红点：单株数目。数据表示为平均值±标准差（n≥10），学生氏t检验计算差异显著性。*表示表型在P < 0.05水平显著；**表示表型在P < 0.01水平极显著。

Fig. 3. Map-based cloning and functional verification of SG5 A, Map-based cloning of SG5. B, Characterization of DNA and protein sequences in Xudao 3(XU3) and sg5. Underlined nucleotides are the sequence of the ﬁrst half of exons 4 in XU3, others are at the terminal of the intron 3. Red boxes are the splicing site mutation and new splicing sites. Black vertical line, the site of jasmonate. Green vertical line, the site of L-alpha-amino acid. C, Gene editing and homozygous mutant sequences. D, Characterization of plant type in XU3, sg5 mutant, SG5#CR-1, and SG5#CR-2. Scale bar=20 cm. E, Glume morphology of XU3, sg5 mutant, SG5#CR-1, and SG5#CR-2; F, Panicle morphology of wild type, sg5 mutant, SG5#CR-1, and SG5#CR-2. Scale bar=5 cm. G, Number of grains per panicle in XU3, sg5 mutant, and SG5#CR-1; Red dot, Normal seeds. Black dot, Moldy seeds. H, Comparison of heading date, plant height, panicle length, panicle number, and seed setting rate in XU3, SG5#CR-1, and SG5#CR-2. Red dot, Number of plants investigated. Data are given as Mean ± SD (n≥10). Student’s t-test was used to generate P value. * P < 0.05; ** P < 0.01.

表2 BSA分析检测的差异位点汇总

Table 2. Summary of differentially expressed genes detected using BSA analysis

差异种类 Category	差异基因数目 Number of differential genes
基因上游 Upstream	638
功能获得型 Stop gain	6
功能缺失型 Stop loss	2
5' UTR	10
3' UTR	14
保守结构缺失/插入 Conservative inframe deletion / insertion	4
散乱结构缺失/插入 Disruptive inframe deletion / insertion	3
移码突变 Frameshift variant	23
错义突变 Missense variant	119
同义突变 Synonymous variant	98
剪接受体变异 Splice acceptor variant & intron variant	4
内含子变异 Intronic	60
基因间变异 Intergenic region	72
基因下游 Downstream	212
合计 Total	1265

图4 SG5时空表达模式及亚细胞定位 A：SG5基因的组织表达模式分析，数据表示为平均值±标准差（n=3）；B：SG5启动子启动GUS蛋白在各个组织部位中的染色情况(比例尺=10 mm)。C：SG5在原生质体中的亚细胞定位，35S::GFP空载 (比例尺=50 μm)；35S::SG5-GFP载体(比例尺=5 μm)。

Fig. 4. Expression pattern analysis of SG5 and subcellular localization A, qRT-PCR analysis of SG5 gene expression in various tissues. Data are given as Mean ± SD (n=3). B, Histochemical analysis of GUS activity in various tissues.( Scale bar=10 mm). C, Subcellular localization of SG5 in rice protoplast. 35S::GFP(scale bar=50 μm). 35S::SG5-GFP(scale bar=5 μm).

图5 SG5进化树分析及蛋白多序列比对 A：SG5蛋白进化树分析。红框：具有GH3-5结构域的种质资源；红色箭头指向：SG5目的蛋白。B：SG5蛋白多序列对分析。灰色框：GH3-5结构域；绿色框：茉莉酮酸酯；红色框：变异位点氨基酸；蓝色框：L-α-氨基酸位点。

Fig. 5. Phylogenetic tree and multiple sequence alignment analysis of SG5 in rice A, Phylogenetic tree analysis of SG5 in rice. Red frame, Rice resources with the GH3-5 domain. Red arrow, SG5 protein. B, Multiple sequence alignment of SG5 in rice. Gray frame, The domain of GH3-5. Green frame, The site of jasmonate. Blue frame, The site of L-alpha-amino acid.

参考文献 55

[1]	Xing Y Z, Zhang Q F. Genetic and molecular bases of rice yield[J]. Annual Review of Plant Biology, 2010, 61: 421-442.
[2]	Ren D Y, Ding C Q, Qian Q. Molecular bases of rice grain size and quality for optimized productivity[J]. Science Bulletin, 2023, 68(3): 314-350.
[3]	Theissen G, Melzer R. Molecular mechanisms underlying origin and diversification of the Angiosperm flower[J]. Annals of Botany, 2007, 100(3): 603-619.
[4]	Soltis D E, Chanderbali A S, Kim S, Buzgo M, Soltis P S. The ABC model and its applicability to basal angiosperms[J]. Annals of Botany, 2007, 100(2): 155-163.
[5]	Maoileidigh D S O, Graciet E, Wellmer F. Gene networks controlling Arabidopsis thaliana flower development[J]. New Phytologist, 2014, 201: 16-30.
[6]	Krizek B A, Fletcher J C. Molecular mechanisms of flower development: An armchair guide[J]. Nature Reviews Genetics, 2005, 6(9): 688-698.
[7]	Causier B, Schwarz-Sommer Z, Davies B. Floral organ identity: 20 years of ABCs[J]. Seminars in Cell & Developmental Biology, 2010, 21(1): 73-79.
[8]	郭爽, 李云峰, 任德勇, 王增, 杜青, 何光华. 水稻内稃扭曲突变体palea distortion 1 (pd1) 的鉴定与基因定位[J]. 分子植物育种, 2011, 9(3): 256-260.
	Guo S, Li Y F, Ren D Y, Wang Z, Du Q, He G H. Identification and gene mapping of a palea distortion 1 (pd1) mutant in rice (Oryza sativa L.)[J]. Molecular Plant Breeding, 2011, 9(3): 256-260. (in Chinese with English abstract)
[9]	Jin Y, Luo Q, Tong H N, Wang A J, Cheng Z J, Tang J F, Li D Y, Zhao X F, Li X B, Wan J M, Jiao Y L, Chu C C, Zhu L H. An AT-hook gene is required for palea formation and floral organ number control in rice[J]. Developmental Biology, 2011, 359(2): 277-288.
[10]	陈东, 毛毕刚, 潘银林, 彭彦, 韶也, 吴天昊, 赵炳然. 水稻颖花畸形突变体(gm(t))的遗传分析与基因定位[J]. 分子植物育种, 2019, 17(15): 5026-5031.
	Chen D, Mao B G, Pan Y L, Peng Y, Shao Y, Wu T H, Zhao B R. Genetic analysis and gene mapping of glume malformation mutant(gm(t)) in rice[J]. Molecular Plant Breeding, 2019, 17(15): 5026-5031. (in Chinese with English abstract)
[11]	薛大伟. 两个水稻花器官突变体的遗传分析[D]. 北京: 中国农业科学院, 2006.
	Xue D W. The genetic analysis of two floral organ mutants from rice[D]. Chinese Academy of Agricultural Sciences, 2006. Beijing: Chinese Academy of Agricultural Sciences, 2006. (in Chinese with English abstract)
[12]	Jeon J S, Jang S, Lee S, Nam J, Kim C, Lee S H, Chung Y Y, Kim S R, Lee Y H, Cho Y G, An G. leafy hull sterile1 is a homeotic mutation in a rice MADS box gene affecting rice flower development[J]. The Plant Cell, 2000, 12(6): 871-884.
[13]	Agrawal G K, Abe K, Yamazaki M, Miyao A, Hirochika H. Conservation of the E-function for floral organ identity in rice revealed by the analysis of tissue culture-induced loss-of-function mutants of the OsMADS1 gene[J]. Plant Molecular Biology, 2005, 59(1): 125-135.
[14]	Prasad K, Parameswaran S, Vijayraghavan U. OsMADS1, a rice MADS‐box factor, controls differentiation of specific cell types in the lemma and palea and is an early‐acting regulator of inner floral organs[J]. The Plant Journal, 2005, 43(6): 915-928.
[15]	Hu Y, Liang W Q, Yin C S, Yang X L, Ping B Z, Li A X, Jia R, Chen M J, Luo Z J, Cai Q, Zhao X X, Zhang D B, Yuan Z. Interactions of OsMADS1 with floral homeotic genes in rice flower development[J]. Molecular plant, 2015, 8(9): 1366-1384.
[16]	Pelucchi N, Fornara F, Favalli C, Masiero S, Lago C, P E M, Colombo L, Kater M M. Comparative analysis of rice MADS-box genes expressed during flower development[J]. Sexual Plant Reproduction, 2002, 15(3): 113-122.
[17]	Wang K J, Tang D, Hong L L, Xu W Y, Huang J, Li M, Gu M H, Xue Y B, Cheng Z K. DEP and AFO regulate reproductive habit in rice[J]. PLoS Genetics, 2010, 6(1): e1000818.
[18]	Kobayashi K, Yasuno N, Sato Y, Yoda M, Yamazaki R, Kimizu M, Yoshida H, Nagamura Y, Kyozuka J. Inflorescence meristem identity in rice is specified by overlapping functions of three AP1/FUL-like MADS box genes and PAP2 a SEPALLATA MADS box gene[J]. The Plant Cell, 2012, 24(5): 1848-1859.
[19]	Yadav S R, Prasad K, Vijayraghavan U. Divergent regulatory OsMADS2 functions control size, shape and differentiation of the highly derived rice floret second-whorl organ[J]. Genetics, 2007, 176(1): 283-294.
[20]	Yao S G, Ohmori S, Kimizu M, Yoshida H. Unequal genetic redundancy of rice PISTILLATA orthologs, OsMADS2 and OsMADS4, in lodicule and stamen development[J]. Plant & Cell Physiology, 2008, 49(5): 853-857.
[21]	Yu X Q, Xia S S, Xu Q K, Cui Y J, Gong M, Zeng D L, Zhang Q, Shen L, Jiao G A, Gao Z Y, Hu J, Zhang G H, Zhu L, Guo L B, Ren D Y, Qian Q. ABNORMAL FLOWER AND GRAIN 1 encodes OsMADS6 and determines palea identity and affects rice grain yield and quality[J]. Science China Life Sciences, 2020, 63(2): 228-238.
[22]	Ohmori S, Kimizu M, Sugita M, Miyao A, Hirochika H, Uchida E, Nagato Y, Yoshida H. MOSAIC FLORAL ORGANS1, an AGL6 -like MADS box gene, regulates floral organ identity and meristem fate in rice[J]. The Plant Cell, 2009, 21(10): 3008-3025.
[23]	Zhang J, Nallamilli B R, Mujahid H, Peng Z H. OsMADS6 plays an essential role in endosperm nutrient accumulation and is subject to epigenetic regulation in rice (Oryza sativa)[J]. The Plant Journal, 2010, 64(4): 604-617.
[24]	Duan Y L, Xing Z, Diao Z J, Xu W Y, Li S P, Du X Q, Wu G H, Wang C L, Lan T, Meng Z, Liu H Q, Wang F, Wu W R, Xue Y B. Characterization of Osmads6-5, a null allele, reveals that OsMADS6 is a critical regulator for early flower development in rice (Oryza sativa L.)[J]. Plant Molecular Biology, 2012, 80(4-5): 429-442.
[25]	Wang H H, Zhang L, Cai Q, Hu Y, Jin Z M, Zhao X X, Fan W, Huang Q M, Luo Z J, Chen M J, Zhang D B, Yuan Z. OsMADS32 interacts with PI-like proteins and regulates rice flower development[J]. Journal of Integrative Plant Biology, 2015, 57(5): 504-513.
[26]	Luo Q, Zhou K D, Zhao X F, Zeng Q C, Xia H A, Zhai W X, Xu J C, Wu X J, Yang H S, Zhu L H. Identification and fine mapping of a mutant gene for palealess spikelet in rice[J]. Planta, 2005, 221(2): 222-230.
[27]	Xiao H, Tang J F, Li Y F, Wang W M, Li X B, Jin L, Xie R, Luo H F, Zhao X F, Meng Z, He G H, Zhu L H. STAMENLESS 1, encoding a single C₂H₂ zinc finger protein, regulates floral organ identity in rice[J]. The Plant Journal, 2009, 59(5): 789-801.
[28]	Yan D W, Zhang X M, Zhang L, Ye S H, Zeng L J, Liu J Y, Li Q, He Z H. CURVED CHIMERIC PALEA 1 encoding an EMF1-like protein maintains epigenetic repression of OsMADS58in rice palea development[J]. The Plant Journal, 2015, 82(1): 12-24.
[29]	Zheng M, Wang Y H, Wang Y L, Wang C M, Ren Y L, Lv J, Peng C, Wu T, Liu K, Zhao S L, Liu X, Guo X P, Jiang L, Terzaghi W, Wan J M. DEFORMED FLORAL ORGAN1 (DFO1) regulates floral organ identity by epigenetically repressing the expression of OsMADS58 in rice (Oryza sativa)[J]. The New Phytologist, 2015, 206(4): 1476-1490.
[30]	Yuan Z, Gao S, Xue D W, Luo D, Li L T, Ding S Y, Yao X, Wilson Z A, Qian Q, Zhang D B. RETARDED PALEA1 controls palea development and floral zygomorphy in rice[J]. Plant Physiology, 2009, 149(1): 235-244.
[31]	Zeng D D, Qin R, Alamin M, Liang R, Yang C C, Jin X L, Shi C H. DBOP specifies palea development by suppressing the expansion of the margin of palea in rice[J]. Genes and Genomics, 2016, 38(11): 1095-1103.
[32]	Lee D Y, Lee J, Moon S, Park S Y, An G. The rice heterochronic gene SUPERNUMERARY BRACT regulates the transition from spikelet meristem to floral meristem[J]. The Plant Journal, 2007, 49(1): 64-78.
[33]	Ren D Y, Li Y F, Zhao F M, Sang X C, Shi J Q, Wang N, Guo S, Ling Y H, Zhang C W, Yang Z L, He G H. MULTI-FLORET SPIKELET1, which encodes an AP2/ERF protein, determines spikelet meristem fate and sterile lemma identity in rice[J]. Plant Physiology, 2013, 162(2): 872-884.
[34]	Ren D Y, Rao Y C, Wu L W, Xu Q K, Li Z Z, Yu H P, Zhang Y, Leng Y J, Hu J, Zhu L, Gao Z Y, Dong G J, Zhang G H, Guo L B, Zeng D L, Qian Q. The pleiotropic ABNORMAL FLOWER AND DWARF1 affects plant height, floral development and grain yield in rice[J]. Journal of Integrative Plant Biology, 2016, 58(6): 529-539.
[35]	Yan D W, Zhou Y, Ye S H, Zeng L J, Zhang X M, He Z H. BEAK-SHAPED GRAIN 1/TRIANGULAR HULL 1, a DUF640 gene, is associated with grain shape, size and weight in rice[J]. Science China Life Sciences, 2013, 56(3): 275-283.
[36]	Livak K J, Schmittgen T D. Analysis of relative gene expression data using real-time quantitative PCR and the 2^-ΔΔCTmethod[J]. Methods, 2001, 25(4): 402-408.
[37]	王忠, 顾蕴洁, 高煜珠. 水稻开颖机理的探讨: Ⅲ.浆片的结构及其在开颖过程中内含物的变化[J]. 作物学报, 1991, 17(2): 96-101, 161-162.
	Wang Z, Gu Y J, Gao Y Z. Studies on the mechanism of the anthesis of rice Ⅲ.Structure of the lodicule and changes of its contents during flowering[J]. Acta Agronomica Sinica, 1991, 17(2): 96-101, 161-162. (in Chinese with English abstract)
[38]	秦鹏. 水稻颖壳发育基因hzp和ug1的研究[D]. 长沙: 湖南农业大学, 2021.
	Qin P. The study on genes hzp and ug1 controlling rice glume development[D]. Changsha: Hunan Agricultural University, 2021. (in Chinese with English abstract)
[39]	Terol J, Domingo C, Talón M. The GH3 family in plants: Genome wide analysis in rice and evolutionary history based on EST analysis[J]. Gene, 2006, 371(2): 279-290.
[40]	Riemann M, Riemann M, Takano M. Rice JASMONATE RESISTANT 1 is involved in phytochrome and jasmonate signalling[J]. Plant, Cell & Environment, 2008, 31(6): 783-792.
[41]	Xiao Y G, Chen Y, Charnikhova T, Mulder P P J, Heijmans J, Hoogenboom A, Agalou A, Michel C, Morel J B, Dreni L, Kater M M, Bouwmeester H, Wang M, Zhu Z, Ouwerkerk P B F. OsJAR1 is required for JA-regulated floret opening and anther dehiscence in rice[J]. Plant Molecular Biology, 2014, 86(1/2): 19-33.
[42]	Jain M, Kaur N, Tyagi A K, Khurana J P. The auxin-responsive GH3 gene family in rice (Oryza sativa)[J]. Functional & Integrative Genomics, 2006, 6(1): 36-46.
[43]	Zhao S Q, Xiang J J, Xue H W. Studies on the rice LEAF INCLINATION1 (LC1), an IAA-amido synthetase, reveal the effects of auxin in leaf inclination control[J]. Molecular Plant, 2013, 6(1): 174-187.
[44]	Du H, Wu N, Fu J, Wang S P, Li X H, Xiao J H, Xiong L Z. A GH3 family member, OsGH3-2, modulates auxin and abscisic acid levels and differentially affects drought and cold tolerance in rice[J]. Journal of Experimental Botany, 2012, 63(18): 6467-6480.
[45]	Fu J, Yu H H, Li X H, Xiao J H, Wang S P. Rice GH3 gene family: Regulators of growth and development[J]. Plant Signaling Behavior, 2011, 6(4): 570-574.
[46]	Kong W L, Zhong H, Deng X X, Gautam M, Gong Z Y, Zhang Y, Zhao G Q, Liu C, Li Y S. Evolutionary analysis of GH3 genes in six Oryza Species/Subspecies and their expression under salinity stress in Oryza sativa ssp. japonica[J]. Plants, 2019, 8(2): 30.
[47]	Mao C J, He J M, Liu L, Deng Q M, Yao X F, Liu C M, Qiao Y L, Li P, Ming F. OsNAC2 integrates auxin and cytokinin pathways to modulate rice root development[J]. Plant Biotechnology Journal, 2020, 18(2): 429-442.
[48]	Ding X H, Cao Y L, Huang L L, Zhao J, Xu C G, Li X H, Wang S P. Activation of the Indole-3-Acetic Acid-Amido synthetase GH3-8 suppresses expansin expression and promotes salicylate- and jasmonate-independent basal immunity in rice[J]. The Plant Cell, 2008, 20(1): 228-240.
[49]	Yadav S R, Khanday I, Majhi B B, Veluthambi K, Vijayraghavan U. Auxin-responsive OsMGH3, a common downstream target of OsMADS1 and OsMADS6, controls rice floret fertility[J]. Plant and Cell Physiology, 2011, 52(12): 2123-2135.
[50]	Fukumoto K, Alamgir K M, Yamashita Y, Mori I C, Matsuura H, Galis I. Response of rice to insect elicitors and the role of OsJAR1 in wound and herbivory-induced JA-Ile accumulation[J]. Journal of Integrative Plant Biology, 2013, 55(8): 775-784.
[51]	Song S S, Qi T C, Huang H, Ren Q C, Wu D W, Chang C Q, Peng W, Liu Y, Peng J R, Xie D X. The jasmonate-ZIM domain proteins interact with the R2R3-MYB transcription factors MYB21 and MYB24 to affect jasmonate-regulated stamen development in Arabidopsis[J]. The Plant Cell, 2011, 23(3): 1000-1013.
[52]	Wakuta S, Suzuki E, Saburi W, Matsuura H, Nabeta K, Imai R, Matsui H. OsJAR1 and OsJAR2 are jasmonyl-_L-isoleucine synthases involved in wound- and pathogen-induced Jasmonic acid signalling[J]. Biochemical and Biophysical Research Communications, 2011, 409: 634-639.
[53]	Zhang S N, Wang S K, Xu Y X, Yu C L, Shen C J, Qian Q, Geisler M, Jiang D A, Qi Y H. The auxin response factor, OsARF19, controls rice leaf angles through positively regulating OsGH3-5 and OsBRI1[J]. Plant, Cell and Environment, 2015, 38: 638-654.
[54]	Ding W Y, Gou Y J, Li Y J, Li J J, Fang Y D, Liu X P, Zhu X Y, Ye R J, Heng Y Q, Wang H Y, Shen R X. A jasmonate-mediated regulatory network modulates diurnal floret opening time in rice[J]. New phytologist, 2024, 244(1): 176-191.
[55]	王忠, 顾蕴洁, 高煜珠. 水稻开颖机理的探讨: V.不育系与可育系浆片和花丝结构的比较[J]. 作物学报, 1994, 20(1): 13-17, 129-131.
	Wang Z, Gu Y J, Gao Y Z. Studies on the mechanism of the anthesis of rice V. comparison of lodicule and filament structure between sterile line and fertile line[J]. Acta Agronomica Sinica, 1994, 20(1): 13-17, 129-131. (in Chinese with English abstract)

水稻颖壳不闭合基因SG5的鉴定与克隆

Identification and Cloning of SG5 in Rice

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