中国水稻科学 ›› 2026, Vol. 40 ›› Issue (5): 584-597.DOI: 10.16819/j.1001-7216.2026.251212
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,*()
收稿日期:2025-12-27
修回日期:2026-03-17
出版日期:2026-09-10
发布日期:2026-09-16
通讯作者:
*email: qianqian188@hotmail.com;基金资助:
CHAI Bingze1, WANG Junge1, WU Kaixiong1, LIU Jialong1, WU Yue1, WANG Huisen1, TAN Lin1, REN Deyong1, GAO Zhenyu1, ZHU Li1, ZHANG Qiang1,2, ZHANG Guangheng1,2, QIAN Qian1,2,*(), HU Jiang1,2,*()
Received:2025-12-27
Revised:2026-03-17
Online:2026-09-10
Published:2026-09-16
摘要：
【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。
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[J]. 中国水稻科学, 2026, 40(5): 584-597.
CHAI Bingze, WANG Junge, WU Kaixiong, LIU Jialong, WU Yue, WANG Huisen, TAN Lin, REN Deyong, GAO Zhenyu, ZHU Li, ZHANG Qiang, ZHANG Guangheng, QIAN Qian, HU Jiang. 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.
图1 gsn4的表型分析 A：W7与gsn4突变体的植株形态，标尺为10 cm；B~D：两者的穗部形态，标尺均为1 cm；C：两者的叶夹角，标尺为 1 cm；E~F：两者的籽粒性状（粒长、粒宽），标尺均为1 cm。G~P：W7及gsn4表型分析。数据以均值±标准差表示（n=20）。****表示 P < 0.0001，ns表示不显著。
Fig. 1. Phenotypic analysis of gsn4 A, Plant morphologies of W7 and gsn4, bar = 10 cm; B−D, Panicle morphologies of the two materials, bars = 1 cm; C, Leaf angle phenotypes of the two materials, bar = 1 cm; E−F, Grain traits (grain length, grain width) of the two materials, bars = 1 cm；G−P, Phenotypic analysis of W7 and gsn4. Data are presented as mean ± standard deviation (n=20). ****, P<0.0001; ns, Not significant.
图2 W7及gsn4颖壳细胞大小分析 A：W7及gsn4的颖壳，标尺为5 mm。B：W7和gsn4颖壳的横切面，标尺为100 μm。C图为B图中横切面局部放大，标尺为100 μm。D~E：W7及gsn4颖壳外表皮细胞(D)及内表皮细胞(E)，标尺为100 μm。F~K：W7及gsn4颖壳细胞性状统计。数据以均值±标准差表示（n=20）。****表示 P < 0.0001， ***表示 P < 0.001， **表示 P < 0.01， *表示 P < 0.05，ns表示不显著。
Fig. 2. Analysis of the cell size of W7 and gsn4 glumes A, Glumes of W7 and gsn4 mutant, bar = 5 mm. B, Cross-sections of glumes of W7 and gsn4, bar = 100 μm. C, Partial magnification of the cross-section in B, bar = 100 μm. D-E, Outer epidermal cells (D) and inner epidermal cells (E) of glumes of W7 and gsn4, bars = 100 μm. F−K, Statistics of glume cell traits of W7 and gsn4. Data are expressed as mean ± standard deviation (n=20). ****, P<0.0001; ***, P<0.001; **, P<0.01; *, P<0.05; ns, Not significant.
图3 GSN4基因的定位与敲除株系表型验证 A~B：GSN4的定位。C：GSN4的突变位点。D：GSN4的测序分析。E：植株形态，标尺为10 cm。F：株高调查。数据以均值±标准差表示（n=10）。 t 检验，****表示 P < 0.0001，ns表示不显著。
Fig. 3. Mapping of GSN4 locus and the knockout of GSN4 A−B, Mapping of GSN4; C, Mutation site of GSN4; D, Sequencing analysis of GSN4; E, Plant morphology, bar = 10 cm; F, Investigation of plant height. Data are expressed as mean ± standard deviation (n=10). ****, P<0.0001; ns, Not significant.
图4 GSN4蛋白的系统发育分析 A：12种GSN4同源物的氨基酸序列比对。完全或部分保守的氨基酸分别用蓝色和绿色阴影表示。B：GSN4的系统树及其同源物。代表性蛋白序列的登录号如下：水稻(Oryza sativa L., CAD41603.3)、海雀稗(Paspalum vaginatum, KAJ1260653.1)、玉米(Zea mays, NP 001333816.1)、糜子(Panicum miliaceum, RLN12872.1)、谷子(Setaria italica, RCV42415.1)、菰(Zizania latifolia, KAL5214521.1)、黑麦草(Lolium perenne, XP 051193481.1)、大麦(Hordeum vulgare subsp. vulgare, XP 044977658.1)、二穗短柄草(Brachypodium distachyon, XP 003577829.1)、芦苇(Phragmites australis, XP 062184470.1)、弯叶画眉草(Eragrostis curvula, TVU13814.1)、穇(Eleusine coracana subsp. coracana, KAK3143048.1)。
Fig. 4. Phylogenetic analysis of GSN4 protein A, Amino acid sequence alignment of twelve GSN4 homologs. Fully or partially conserved amino acids are shaded in blue and green, respectively. B, Phylogenetic tree of GSN4 and its homologs. The accession numbers of representative protein sequences are as follows: Oryza sativa L. (CAD41603.3), Paspalum vaginatum (KAJ1260653.1), Zea mays (NP_001333816. 1), Panicum miliaceum (RLN12872.1), Setaria italica (RCV42415.1), Zizania latifolia (KAL5214521.1), Lolium perenne (XP_051193481.1), Hordeum vulgare subsp. Vulgare (XP_044977658.1), Brachypodium distachyon (XP_003577829.1), Phragmites australis (XP_062184470.1), Eragrostis curvula (TVU13814.1), Eleusine coracana subsp. Coracana (KAK3143048.1).
图5 GSN4的亚细胞定位和表达模式分析 A：GSN4的水稻原生质体亚细胞定位分析。绿色荧光显示GFP，红色荧光显示叶绿体自发荧光，比例尺=10 µm。B：GSN4的烟草亚细胞定位分析，绿色荧光显示GFP，比例尺=40 µm。C：GSN4在水稻组织的表达模式。数据以均值±标准差表示（n=3）。****表示 P < 0.0001 ， ***表示 P < 0.001， **表示 P < 0.01，ns表示不显著。
Fig. 5. Subcellular localization and expression pattern analysis of GSN4 A, Subcellular localization analysis of GSN4 in rice protoplasts. Green fluorescence indicates GFP, and red fluorescence indicates chloroplast autofluorescence; scale bar = 10 μm. B, Subcellular localization analysis of GSN4 in tobacco. Green fluorescence indicates GFP; scale bar = 40 μm. C, Expression pattern of GSN4 in rice tissues. Data are expressed as mean ± standard deviation (n=3). ****, P<0.0001; ***, P<0.001; **, P<0.01; ns, Not significant.
图6 野生型W7及突变体gsn4的转录组分析 A：差异表达基因火山图；B：KEGG富集分析图。
Fig. 6. RNA-seq analysis of wild-type W7 and mutant gsn4 A, Volcanogram of differentially expressed genes. B, KEGG enrichment analysis diagram.
图7 野生型W7及突变体gsn4差异基因表达热图分析 图中颜色代表基因的相对表达量，红色代表高表达，蓝色代表低表达。W7为野生型，gsn4为突变体。
Fig. 7. Cluster analysis of differential gene expression patterns in wild-type W7 and mutant gsn4 The colors indicates the relative expression level of genes, with red for high expression and blue for low expression. W7 indicates the wild type and gsn4 indicates the mutant.
图8 W7和gsn4突变体幼穗中细胞增殖、细胞扩张以及粒形相关基因表达量分析 A：W7和gsn4幼穗中细胞增殖相关基因表达分析；B：W7和gsn4幼穗中细胞扩张相关基因表达分析；C：W7和gsn4幼穗中粒形相关基因表达分析。数据以均值±标准差表示（n=3）。****表示 P < 0.0001 ， ***表示 P < 0.001， **表示 P < 0.01， *表示 P < 0.05，NS 表示不显著。
Fig. 8. Expression analysis of genes related to cell proliferation, cell expansion, and grain size in young panicles of W7 and gsn4 A, Gene expression analysis of cell proliferation-related genes in young panicles of W7 and gsn4; B, Gene expression analysis of cell expansion-related genes in young panicles of W7 and gsn4; C, Grain size-related gene expression analysis in young panicles of W7 and gsn4. Data are expressed as mean ± standard deviation (n=3). ****, P<0.0001; ***, P<0.001; **, P<0.01; *, P<0.05; NS, not significant.
图9 野生型W7和突变体gsn4对油菜素内酯的敏感性及BR合成相关基因表达分析 A~B：在不同浓度表油菜素内酯(2,4-e BL)及油菜素唑(BRZ)的处理下，野生型W7和突变体gsn4第二片叶的叶夹角变化图，标尺为1 cm。C~D：叶夹角变化统计学分析。数据以均值±标准差表示（n=10）。E：BR生物合成基因的表达量。数据以均值±标准差表示（n=3）。****表示 P < 0.0001，ns 表示不显著。
Fig. 9. Brassinosteroid sensitivity and expression analysis of BR biosynthetic-related genes in wild-type W7 and mutant gsn4. A-B, Scans of leaf angle changes in the second leaf of wild-type W7 and mutant gsn4 under different concentrations of 2,4-e BL and BRZ treatment, Bar=1 cm. C-D, Statistical analysis of leaf angle change. Data are expressed as mean ± standard deviation (n=10). E, Expression of BR biosynthetic gene. Data are expressed as mean ± standard deviation (n=3). ****, P<0.0001; ns, Not significant.
| 位置 Location | 27551959 | 27552410 | 27552420 | 27554431 | 27554638 | 27555074 | 27555189 | 27555943 | 27556520 | 27556903 | 27557628 |
|---|---|---|---|---|---|---|---|---|---|---|---|
| SNP | G/A | C/A | C/T | A/G | C/A | A/G | A/G | T/C | G/A | A/T | G/A |
| Hap1 | G | C | C | A | C | A | A | T | G | A | G |
| Hap2 | A | A | T | G | A | G | G | C | A | A | A |
| Hap3 | G | A | C | G | C | G | G | C | A | T | G |
表1 GSN4基因Hap1-Hap3的核心多态性位点基因型
Table 1. Genotypes of core polymorphic sites of Hap1-Hap3 in GSN4
| 位置 Location | 27551959 | 27552410 | 27552420 | 27554431 | 27554638 | 27555074 | 27555189 | 27555943 | 27556520 | 27556903 | 27557628 |
|---|---|---|---|---|---|---|---|---|---|---|---|
| SNP | G/A | C/A | C/T | A/G | C/A | A/G | A/G | T/C | G/A | A/T | G/A |
| Hap1 | G | C | C | A | C | A | A | T | G | A | G |
| Hap2 | A | A | T | G | A | G | G | C | A | A | A |
| Hap3 | G | A | C | G | C | G | G | C | A | T | G |
| 单倍型 Haplotype | 亚群 Subgroup | 粒长 Grain length(mm) | 粒宽 Grain width(mm) | 千粒重 1000-grain weight(g) |
|---|---|---|---|---|
| Hap1 | XI: 245；admix: 40；Aus: 43；GJ: 778；Bas: 61；na: 1 | 8.566 | 3.178 | 26.032 |
| Hap2 | XI: 880；admix: 14；Aus: 1 | 8.597 | 2.893 | 24.688 |
| Hap3 | XI: 415；admix: 14；Aus: 138；GJ: 1；Bas: 2 | 8.654 | 2.905 | 24.231 |
表2 GSN4基因Hap1-Hap3的亚群分布及粒形性状
Table 2. Subgroup distribution and grain traits of Hap1-Hap3 in GSN4
| 单倍型 Haplotype | 亚群 Subgroup | 粒长 Grain length(mm) | 粒宽 Grain width(mm) | 千粒重 1000-grain weight(g) |
|---|---|---|---|---|
| Hap1 | XI: 245；admix: 40；Aus: 43；GJ: 778；Bas: 61；na: 1 | 8.566 | 3.178 | 26.032 |
| Hap2 | XI: 880；admix: 14；Aus: 1 | 8.597 | 2.893 | 24.688 |
| Hap3 | XI: 415；admix: 14；Aus: 138；GJ: 1；Bas: 2 | 8.654 | 2.905 | 24.231 |
图10 GSN4的优势单倍型的粒形性状分布 Hap1-Hap3 单倍型对应粒长(GL)、粒宽(GW)、千粒重(TGW)的小提琴图，图中箱线代表四分位数，小提琴轮廓反映表型分布密度。
Fig. 10. Grain trait distribution of dominant haplotypes of GSN4 Violin plots of grain length (GL), grain width (GW), and thousand-grain weight (TGW) corresponding to Hap1-Hap3, respectively. The boxplot in the figure represents quartiles, and the violin outline reflects the phenotypic distribution density.
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