水稻短圆粒突变体srg10的基因克隆及其功能分析

doi:10.16819/j.1001-7216.2026.250707

摘要/Abstract

摘要：

【目的】短圆粒突变体是研究水稻粒形调控机制的理想材料，可为水稻产量育种提供新的靶标和育种材料。【方法】利用BSA-Seq技术对水稻短圆粒突变体srg10的突变基因进行了定位，并利用CRISPR/Cas9敲除技术对候选基因进行了功能验证，鉴定了srg10对水稻主要产量性状的效应，并分析了其影响粒形的机制。【结果】突变体srg10每穗粒数、粒重等主要农艺性状均受到显著影响，产量有明显降低趋势，同时蒸煮食味品质等也受到影响。Os10g0363700基因上的一个单核苷酸多态性(single nucleotide polymorphism, SNP)，即由G突变为A是srg10产生短圆粒表型的原因。突变体srg10幼穗中出现292个差异表达基因，GO富集分析显示，这些差异表达基因主要与细胞发育生长、信号响应等生物学过程相关。已知粒形基因表达分析显示srg10突变可能涉及细胞周期、转录因子、RNA剪接、植物激素等调控通路，实现对幼穗发育的调控。【结论】Os10g0363700基因突变会产生短圆粒等不利产量的表型，RNA-Seq分析表明该突变基因可能与细胞微丝形成有关，同时也可能通过已知粒形基因直接或间接作用调控粒形。

关键词: 水稻, 粒形, BSA-Seq, 短圆粒突变体, SRG10

Abstract:

【Objective】Small and round grain mutants serve as ideal genetic materials for dissecting the regulation mechanism of rice grain size and can provide new breeding material for yield improvement.【Method】BSA-Seq was employed to delimit the causal locus in the small and round grain mutant srg10. Candidate gene was functionally verified by CRISPR/Cas9-mediated knockout. The phenotypic effects of srg10 on the main yield-related traits were identified, and the underlying mechanism modulating grain size was investigated.【Result】The srg10 mutation significantly reduced grain number per panicle and grain weight, resulting in a pronounced yield penalty. A single nucleotide substitution (SNP, G>A) within the Os10g0363700 gene was identified as the cause for the small and round grain phenotype. Comparative transcriptome profiling of young panicles from srg10 and wild-type plants revealed 292 differentially expressed genes (DEGs). GO enrichment analysis showed that these DEGs are predominantly associated with biological processes such as cell development, cell growth, and signal transduction. Expression profiling of known grain size genes further suggested that the srg10 mutation may involve multiple regulatory pathways such as cell cycle, transcriptional regulation, RNA splicing, and plant hormones, thereby modulating panicle development.【Conclusion】Mutation of Os10g0363700 produces pleiotropic, yield-adverse phenotypes characterized by small and round grains. RNA-Seq analysis showed that the mutant gene may be related to cell microfilament organization, and may also directly or indirectly regulate grain size via known grain size genes.

Key words: rice, grain size, BSA-Seq, small and round grain, SRG10

赵冬生, 李羽帆, 赵贝贝, 张涛, 李钱峰, 刘巧泉. 水稻短圆粒突变体srg10的基因克隆及其功能分析[J]. 中国水稻科学, 2026, 40(6): 756-766.

ZHAO Dongsheng, LI Yufan, ZHAO Beibei, ZHANG Tao, LI Qianfeng, LIU Qiaoquan. Gene Cloning and Function Analysis of Small and Round Grain Mutant srg10 in Rice[J]. Chinese Journal OF Rice Science, 2026, 40(6): 756-766.

图/表 8

表1 本研究用到的引物序列信息

Table 1. Information of primer sequences used in the study

引物名称 Primer	引物序列 Sequence (5'-3')	用途 Function
SRG10-SNP-F	ACCTCACGACCTCACCTGC	检测P9516背景中EMS诱变产生的SNP位点
SRG10-SNP-R	CCTGAAGAAGCCGTGTTG	检测P9516背景中EMS诱变产生的SNP位点
gRNA-F	GGCAGTGCCGAAGTATCCTCGCTA	构建CRISPR/Cas9基因敲除载体
gRNA-R	AAACTAGCGAGGATACTTCGGCAC	构建CRISPR/Cas9基因敲除载体
SRG10-1F	ATGTCAAGGGTACCTAAATGGAAGA	PCR扩增并测序鉴定SRG10的转录产物
SRG10-1R	TCAGTGTTTAATGTGGACTGTTTATGTAA
SRG10-2F	TATGCAAACTGTCTCTGCATTGC
SRG10-2R	CTATGATCTCATGGGTGATCGGT
SRG10-KO-F	ACAAGGGAGCAATGAGTGG	检测基因敲除产生的靶位点序列变异
SRG10-KO-F	GCAGCATTTGTGGGTGGA	检测基因敲除产生的靶位点序列变异
Actin-RT-F	CCAAGGCCAATCGTGAGAAGA	基因表达量检测
Actin-RT-F	AATCAGTGAGATCACGCCCAG
LOC_Os01g09200-F	AGCGCAAGGAGAGGCGCTTT
LOC_Os01g09200-R	GCCGCATGTACTCTGTTTCC
LOC_Os02g56140-F	AGTCTTCGCGTTCCAATCTT
LOC_Os02g56140-R	AACCGACTTCTCATCGGTTC
LOC_Os03g25770-F	TATGATTATGGTCCATCACCGC
LOC_Os03g25770-R	AATGGTCTTCGTAGATCATCAGG
LOC_Os04g39430-F	GTGAAAAGGGCGGAAGGGA
LOC_Os04g39430-R	TCTTGCTTGTCCCTTGGCTTG
LOC_Os05g27730-F	AACGGCTGCTCCATGAAGAA
LOC_Os05g27730-R	TTGTGTGCGCCCTTGTAGAC
LOC_Os05g37470-F	GCGTCGTCCTCTTTTGTTTGG
LOC_Os05g37470-R	GGGCCACCTTCCTAGTTAGCAT
LOC_Os06g15620-F	CATTGGTGTGTCACTGCCTC
LOC_Os06g15620-R	TCCTCTCTCGCACACATTCC
LOC_Os07g39920-F	AATGGTCTTCGTAGATCATCAGG
LOC_Os07g39920-R	TCTTCAGGATTTCCACGCAC
LOC_Os07g41200-F	CCCCTAGCATCGACACCAAG
LOC_Os07g41200-R	CGGGTTCCAGCACTCCTCT

图1 野生型P9516与突变体srg10表型比较 A: 野生型P9516与突变体srg10的植株对比；比例尺为10 cm；B: 野生型P9516与突变体srg10的穗型对比；比例尺为1 cm；C: 野生型P9516与突变体srg10的粒形对比；比例尺为10 mm；D~M: 野生型P9516与突变体srg10的农艺性状。**表示在P<0.01 水平上差异极显著；NS表示没有显著差异。

Fig. 1. Comparison between wild type P9516 and mutant srg10 A, Comparison of plant between P9516 and srg10; Scale=10 cm; B, Comparison of panicle type between P9516 and srg10; Scale=1 cm; C, Comparison of grain size between P9516 and srg10; Scale=10 mm; D-M, Agronomic traits of P9516 and srg10. ** indicate extremely significant difference at P<0.01 level; NS indicate no significant difference.

图2 基于BSA-Seq定位srg10基因 A: 基于BSA-Seq获得的SNP的ΔSNP-index关联分析图，散点图为原始值，黑色曲线为拟合值，蓝色和红色横线为置信区间边界位置或指定阈值位置；B: 连锁区域内srg10候选突变位点分析；C: Sanger测序鉴定分离群体中各个单株在该候选突变位点处的SNP基因分型；D~F: 该候选SNP位点不同基因分型所对应的粒长、粒宽、籽粒长宽比。

Fig. 2. Mapping the srg10 gene based on SNP of BSA-Seq A, ΔSNP-index association analysis chart based on SNP of BSA-seq. The scatter plot is the original value, the black curve is the fitted value, and the blue and red horizontal lines are the positions of the confidence interval boundaries or the specified thresholds; B, Analysis of candidate SNP within the linkage interval for srg10 mutant; C, Genotype of plants at the SNP site. D-F, Analysis of grain length, grain width, and grain length-width ratio corresponding to different genotyping of this SNP site.

图3 候选基因鉴定及功能验证 A: 候选基因Os10g0363700的基因结构图；SNP: P9516背景中EMS诱变产生的srg10变异位点；P9516和srg10：诱变产生的SNP变异位点影响转录本序列变化情况；gRNA: CRISPR/Cas9基因敲除的靶位点及ZH11与SRG10-KO的靶位点序列变异情况；B: 基于水稻基因组注释数据库预测的转录本序列，PCR扩增测序鉴定该SNP位点导致的Os10g0363700的转录产物差异；SRG10-1F/1R和SRG10-2F/2R是用于扩增SRG10基因的引物；M: DNA 标记；C：P9516和srg10的Os10g0363700基因推测蛋白序列比对分析；D: 野生型ZH11与敲除系SRG10-KO的植株比较；比例尺为10 cm；E: 野生型ZH11与敲除系SRG10-KO的穗型比较；比例尺为1 cm；F: 野生型ZH11与敲除系SRG10-KO的粒形比较；比例尺为10 mm；G~P: 野生型ZH11与敲除系SRG10-KO的农艺性状。**表示在P<0.01水平上差异极显著，NS表示没有显著差异。

Fig. 3. Identification and function analysis of candidate genes A, The transcript of Os10g0363700 in young panicle of P9516. SNP, srg10 mutation site generated by EMS mutagenesis in P9516 background; gRNA, Target site of CRISPR/Cas9 knockout, and sequence variation of ZH11 and SRG10-KO at the target site. B, PCR product analysis by agarose gel electrophoresis. M, DNA marker; C, Comparison of predicted protein sequences of Os10g0363700 of P9516 and srg10. D, Comparison of plants between ZH11 and SRG10-KO; Scale=10 cm; E, Comparison of panicles between ZH11 and SRG10-KO; Scale=1 cm; F, Comparison of grain size between ZH11 and SRG10-KO; Scale=10 mm; G-P, Agronomic traits of ZH11 and SRG10-KO. ** indicate extremely significant difference at P<0.01 level; NS indicate no significant difference.

图4 srg10突变体稻米外观和蒸煮食味品质 A~B: 野生型与srg10突变的精米外观对比；比例尺为1 cm；C~D: 野生型与srg10突变的稻米直链淀粉含量对比；E~F: 野生型与srg10突变的黏度对比。*表示在P<0.05水平上显著差异; **表示在P<0.01水平上极显著差异。

Fig. 4. Rice grain appearance, cooking and eating qualities of srg10 A and B, Comparison of appearance of polished rice between wild type and srg10 mutation; Scale=1 cm; C and D, Comparison of amylose content between wild type and srg10 mutation; E and F, Comparison of viscosity between wild type and srg10 mutation. * indicate significant difference at P<0.05 level; ** indicate extremely significant difference at P<0.01 level.

图5 差异表达基因GO 富集分析三大类中前10条条目

Fig. 5. Top 10 terms in each of the three categories of GO enrichment for differentially expressed genes

表2 G10-KO和ZH11之间幼穗组织中存在显著差异表达的籽粒大小相关基因

Table 2. Significantly differentially expressed grain size genes in young panicle between SRG10-KO and ZH11

基因号 Gene ID	基因符号 Gene symbol	基因名称 Gene name	涉及调控通路 Regulatory pathway	表达丰度 FPKM_ZH11	表达丰度 FPKM_srg10	差异倍数 Fold change	q值 q value	上调/下调 Up/Down
LOC_Os01g09200	OsSTK38; TINY1	丝氨酸/苏氨酸蛋白激酶	细胞周期	46.60	41.39	0.84	1.66E-02	下调Down
LOC_Os01g43230	OsTCL2	MYB转录因子		29.05	40.74	1.33	2.17E-03	上调Up
LOC_Os01g48960	OsGOGAT1; NADH-GOGAT1	依赖于NADH的谷氨酸合酶; 谷氨酸合成酶		85.66	69.98	0.78	2.34E-03	下调Down
LOC_Os01g62760	OsPP2C09	2C型蛋白磷酸酶		16.27	12.98	0.76	2.70E-03	下调Down
LOC_Os02g56140	OsbHLH107; lgs1	bHLH转录因子; 大粒	细胞周期	14.39	11.65	0.77	4.50E-02	下调Down
LOC_Os02g58214	OsmiR396f	微RNA	GRF转录因子	41.25	54.34	1.25	7.29E-03	上调Up
LOC_Os03g20790	MHZ6; OsEIL1	猫胡子基因; 乙烯信号调控因子		12.33	16.36	1.26	1.93E-02	上调Up
LOC_Os03g25770	OsSCL26	剪接因子	RNA剪接因子-GW6/OsLG3	22.89	15.88	0.66	7.04E-04	下调Down
LOC_Os04g18090	OsHis1.1	组蛋白H1		237.45	484.91	1.94	2.49E-10	上调Up
LOC_Os04g39430	d11; CYP724B1; CPB1; sg4; cl4; NBG4; GNS4	矮秆; 细胞色素P450;小粒; 簇生穗; 腹切米; 粒数和籽粒大小	油菜素内酯BR	16.45	9.73	0.56	1.32E-04	下调Down
LOC_Os04g52540	OsAGO2	AGO蛋白		27.70	20.27	0.69	6.15E-06	下调Down
LOC_Os05g03040	RSR1	AP2/EREBP转录因子; 淀粉调节因子		7.97	12.73	1.52	1.51E-04	上调Up
LOC_Os05g27730	OsWRKY53	WRKY转录因子	MAPK-BR	6.26	9.36	1.42	1.48E-02	上调Up
LOC_Os05g37470	OsAUX3; qGL5	生长素内向转运载体; 生长素输入载体; 生长素流入载体	生长素 Auxin	18.13	28.16	1.48	2.77E-02	上调Up
LOC_Os06g15620	OsGSR1; GW6; OsGASR7	赤霉素刺激转录基因; 粒宽; 粒长	赤霉素 GA	23.11	18.48	0.76	2.44E-02	下调Down
LOC_Os07g39920	DRT1; OsFH13	成蛋白	微丝形成	12.52	8.48	0.64	5.57E-05	下调Down
LOC_Os07g41200	GL7; GW7; SLG7	粒长; 粒宽; 籽粒长宽比	SPL转录因子	12.88	17.35	1.28	1.03E-03	上调Up
LOC_Os07g44950	OsbZIP60; O3; OPAQUE3	bZIP转录因子		22.54	14.78	0.62	5.11E-04	下调Down
LOC_Os07g49400	OsAPX2; APXb	抗坏血酸过氧化物酶		706.35	615.03	0.83	3.68E-02	下调Down
LOC_Os08g07010	OsABCG18	ABC转运蛋白; ATP结合盒转运蛋白G亚组蛋白		9.29	12.98	1.33	8.98E-03	上调Up
LOC_Os09g26970	qGL9.1	细胞色素P450		0.02	0.24	12.11	1.82E-07	上调Up

表2 G10-KO和ZH11之间幼穗组织中存在显著差异表达的籽粒大小相关基因

Table 2. Significantly differentially expressed grain size genes in young panicle between SRG10-KO and ZH11

基因号 Gene ID	基因符号 Gene symbol	基因名称 Gene name	涉及调控通路 Regulatory pathway	表达丰度 FPKM_ZH11	表达丰度 FPKM_srg10	差异倍数 Fold change	q值 q value	上调/下调 Up/Down
LOC_Os01g09200	OsSTK38; TINY1	丝氨酸/苏氨酸蛋白激酶	细胞周期	46.60	41.39	0.84	1.66E-02	下调Down
LOC_Os01g43230	OsTCL2	MYB转录因子		29.05	40.74	1.33	2.17E-03	上调Up
LOC_Os01g48960	OsGOGAT1; NADH-GOGAT1	依赖于NADH的谷氨酸合酶; 谷氨酸合成酶		85.66	69.98	0.78	2.34E-03	下调Down
LOC_Os01g62760	OsPP2C09	2C型蛋白磷酸酶		16.27	12.98	0.76	2.70E-03	下调Down
LOC_Os02g56140	OsbHLH107; lgs1	bHLH转录因子; 大粒	细胞周期	14.39	11.65	0.77	4.50E-02	下调Down
LOC_Os02g58214	OsmiR396f	微RNA	GRF转录因子	41.25	54.34	1.25	7.29E-03	上调Up
LOC_Os03g20790	MHZ6; OsEIL1	猫胡子基因; 乙烯信号调控因子		12.33	16.36	1.26	1.93E-02	上调Up
LOC_Os03g25770	OsSCL26	剪接因子	RNA剪接因子-GW6/OsLG3	22.89	15.88	0.66	7.04E-04	下调Down
LOC_Os04g18090	OsHis1.1	组蛋白H1		237.45	484.91	1.94	2.49E-10	上调Up
LOC_Os04g39430	d11; CYP724B1; CPB1; sg4; cl4; NBG4; GNS4	矮秆; 细胞色素P450;小粒; 簇生穗; 腹切米; 粒数和籽粒大小	油菜素内酯BR	16.45	9.73	0.56	1.32E-04	下调Down
LOC_Os04g52540	OsAGO2	AGO蛋白		27.70	20.27	0.69	6.15E-06	下调Down
LOC_Os05g03040	RSR1	AP2/EREBP转录因子; 淀粉调节因子		7.97	12.73	1.52	1.51E-04	上调Up
LOC_Os05g27730	OsWRKY53	WRKY转录因子	MAPK-BR	6.26	9.36	1.42	1.48E-02	上调Up
LOC_Os05g37470	OsAUX3; qGL5	生长素内向转运载体; 生长素输入载体; 生长素流入载体	生长素 Auxin	18.13	28.16	1.48	2.77E-02	上调Up
LOC_Os06g15620	OsGSR1; GW6; OsGASR7	赤霉素刺激转录基因; 粒宽; 粒长	赤霉素 GA	23.11	18.48	0.76	2.44E-02	下调Down
LOC_Os07g39920	DRT1; OsFH13	成蛋白	微丝形成	12.52	8.48	0.64	5.57E-05	下调Down
LOC_Os07g41200	GL7; GW7; SLG7	粒长; 粒宽; 籽粒长宽比	SPL转录因子	12.88	17.35	1.28	1.03E-03	上调Up
LOC_Os07g44950	OsbZIP60; O3; OPAQUE3	bZIP转录因子		22.54	14.78	0.62	5.11E-04	下调Down
LOC_Os07g49400	OsAPX2; APXb	抗坏血酸过氧化物酶		706.35	615.03	0.83	3.68E-02	下调Down
LOC_Os08g07010	OsABCG18	ABC转运蛋白; ATP结合盒转运蛋白G亚组蛋白		9.29	12.98	1.33	8.98E-03	上调Up
LOC_Os09g26970	qGL9.1	细胞色素P450		0.02	0.24	12.11	1.82E-07	上调Up

图6 RT-qPCR分析差异表达基因的表达水平

Fig. 6. RT-qPCR analysis of the expression level of DEGs

参考文献 23

[1]	Sakamoto T, Matsuoka M. Identifying and exploiting grain yield genes in rice[J]. Current Opinion in Plant Biology, 2008, 11: 209-214.
[2]	Fan Y, Li Y. Molecular, cellular and Yin-Yang regulation of grain size and number in rice[J]. Molecular Breeding, 2019, 39: 163.
[3]	Huang R, Jiang L, Zheng J, Wang T, Wang H, Huang Y, Hong Z. Genetic bases of rice grain shape: So many genes, so little known[J]. Trends in Plant Science, 2013, 18(4): 218-226.
[4]	Li N, Xu R, Li Y. Molecular networks of seed size control in plants[J]. Annual Review of Plant Biology, 2019, 70: 11. 1-11.30.
[5]	Lu X, Li F, Xiao Y, Wang F, Zhang G, Deng H, Tang W. Grain shape genes: Shaping the future of rice breeding[J]. Rice Science, 2023, 30: 379-404.
[6]	Zhao D, Zhang C, Li Q, Liu Q. Genetic control of grain appearance quality in rice[J]. Biotechnology Advances, 2022, 60: 108014.
[7]	Sun S, Wang L, Mao H, Shao L, Li X, Xiao J, Ouyang Y, Zhang Q. A G-protein pathway determines grain size in rice[J]. Nature Communications, 2018, 9(1): 851.
[8]	Guo T, Chen K, Dong N, Shi C, Ye W, Gao J, Shan J, Lin H. GRAIN SIZE AND NUMBER1 negatively regulates the OsMKKK10-OsMKK4-OsMPK6 cascade to coordinate the trade-off between grain number per panicle and grain size in rice[J]. Plant Cell, 2018, 30: 871-888.
[9]	Xu R, Duan P, Yu H, Zhou Z, Zhang B, Wang R, Li J, Zhang G, Zhuang S, Lyu J, Li N, Chai T, Tian Z, Yao S, Li Y. Control of grain size and weight by the OsMKKK10-OsMKK4-OsMAPK6 signaling pathway in rice[J]. Molecular Plant, 2018, 11: 860-873.
[10]	Si L, Chen J, Huang X, Gong H, Luo J, Hou Q, Zhou T, Lu T, Zhu J, Shangguan Y, Chen E, Gong C, Zhao Q, Jing Y, Zhao Y, Li Y, Cui L, Fan D, Lu Y, Weng Q, Wang Y, Zhan Q, Liu K, Wei X, An K, An G, Han B. OsSPL13 controls grain size in cultivated rice[J]. Nature Genetics, 2016, 48: 447-456.
[11]	Wang S, Li S, Liu Q, Wu K, Zhang J, Wang S, Wang Y, Chen X, Zhang Y, Gao C, Wang F, Huang H, Fu X. Control of grain size, shape and quality by OsSPL16 in rice[J]. Nature Genetics, 2012, 44: 950-954.
[12]	Che R, Tong H, Shi B, Liu Y, Fang S, Liu D, Xiao Y, Hu B, Liu L, Wang H, Zhao M, Chu C. Control of grain size and rice yield by GL2-mediated brassinosteroid responses[J]. Nature Plants, 2015, 2: 15195.
[13]	Duan P, Ni S, Wang J, Zhang B, Xu R, Wang Y, Chen H, Zhu X, Li Y. Regulation of OsGRF4 by OsmiR396 controls grain size and yield in rice[J]. Nature Plants, 2015, 2: 15203.
[14]	Hu J, Wang Y, Fang Y, Zeng L, Xu J, Yu H, Shi Z, Pan J, Zhang D, Kang S, Zhu L, Dong G, Guo L, Zeng D, Zhang G, Xie L, Xiong G, Li J, Qian Q. A rare allele of GS2 enhances grain size and grain yield in rice[J]. Molecular Plant, 2015, 8: 1455-1465.
[15]	李堂, 曹华盛, 熊亮, 王福军, 李曙光, 顾海永, 罗文永, 何高, 梁世胡. 水稻粒型调控基因功能研究进展[J]. 广东农业科学, 2023, 50(12): 12-28.
	Li T, Cao H, Xiong L, Wang F, Li S, Gu H, Luo W, He G, Liang S H. Research progress on the function of rice grain type genes[J]. Guangdong Agricultural Sciences, 2023, 50(12): 12-28. (in Chinese with English abstract)
[16]	Ren D, Ding C, Qiao Q. Molecular bases of rice grain size and quality for optimized productivity[J]. Science Bulletin, 2023, 68: 314-350.
[17]	Luo Y, Chen Y, Xue P, Wang B, Kang Y, Zhang Y, Chen D, Hong Y, Wu W, Liu Q, Zhan X, Lin Y, Cheng S, Zhang Y, Cao L. Modulation of rice grain shape and appearance by the GS10-encoded long coiled-coil protein[J]. Crop Journal, 2025, 13: 158-169.
[18]	Liu H, Ni J, Zhang Y, Chen Y, Luo Y, Wang Y, Shang F, Yang Y, Xu R, Cao L, Hong L, Xu J, Yang Y, Zhou M. GLGW10 controls grain size associated with the lignin content in rice[J]. Journal of Genetics and Genomics, 2025, doi.org/10.1016/j.jgg.2025.02.009.
[19]	Wang C, Shen L, Fu Y, Yan C, Wang K. A simple CRISPR/Cas9 system for multiplex genome editing in rice[J]. Journal of Genetics and Genomics, 2015, 42: 703-706.
[20]	Zeng D, Tian Z, Rao Y, Dong G, Yang Y, Huang L, Leng Y, Xu J, Sun C, Zhang G, Hu J, Zhu L, Gao Z, Hu X, Guo L, Xiong G, Wang Y, Li J, Qian Q. Rational design of high-yield and superior-quality rice[J]. Nature Plants, 2017, 3: 17031.
[21]	Zhang Y, Dong G, Wu L, Wang X, Chen F, Xiong E, Xiong G, Zhou Y, Kong Z, Fu Y, Zeng D, Ma D, Qian Q, Yu Y. Formin protein DRT1 affects gross morphology and chloroplast relocation in rice[J]. Plant Physiology, 2023, 191(1): 280-298.
[22]	Zhang T, Wang Z, Liu Q, Zhao D. Genetic improvement of rice grain size using the CRISPR/Cas9 system[J]. Rice, 2025, 18: 3.
[23]	Song X, Meng X, Guo H, Cheng Q, Jing Y, Chen M, Liu G, Wang B, Wang Y, Li J, Yu H. Targeting a gene regulatory element enhances rice grain yield by decoupling panicle number and size[J]. Nature Biotechnology, 2022, 40: 1403-1411.