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作物学报 ›› 2026, Vol. 52 ›› Issue (11): 3221-3231.doi: 10.3724/SP.J.1006.2026.62009

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

一个新的水稻窄叶突变体nal24的基因定位及克隆

赵汭彤1,2,李毅林1,周皖峰1,2,方成1,张东杨1,尚丽君1,程维民1,陶亮之1,吴跃进1,武健东2,张从合3,叶亚峰1,*,刘斌美1,*   

  1. 1 中国科学院合肥物质科学研究院, 安徽合肥 230031; 2 安徽农业大学, 安徽合肥 230036; 3 安徽荃银高科种业股份有限公司, 安徽合肥 230088
  • 收稿日期:2026-03-22 修回日期:2026-08-21 接受日期:2026-08-21 出版日期:2026-11-12 网络出版日期:2026-09-01
  • 通讯作者: 刘斌美, E-mail: liubm@ipp.ac.cn; 叶亚峰, E-mail: yyfeng@ipp.ac.cn
  • 基金资助:
    本研究由安徽省重点研究与开发计划项目(2023n06020006),安徽省级科技创新攻坚计划项目(202523anull050002),长三角科技创新共同体联合攻关(基础研究)项目(2024CSJZN01100),国家自然科学基金青年项目C (32401738)和中国科学院青年促进会项目(2022454)资助。

Gene mapping and cloning of a novel narrow-leaf mutant nal24 in rice

Zhao Rui-Tong1,2,Li Yi-Lin1,Zhou Wan-Feng1,2,Fang Cheng1,Zhang Dong-Yang1,Shang Li-Jun1,Cheng Wei-Min1,Tao Liang-Zhi1,Wu Yue-Jin1,Wu Jian-Dong2,Zhang Cong-He3,Ye Ya-Feng1,*,Liu Bin-Mei1,*   

  1. 1 Hefei Institutes of Physical Science, Chinese Academy of Sciences, Hefei 230031, Anhui, China; 2 Anhui Agricultural University, Hefei 230036, Anhui, China; 3 Winall Hi-tech Seed Co., Ltd., Hefei 230088, Anhui, China
  • Received:2026-03-22 Revised:2026-08-21 Accepted:2026-08-21 Published:2026-11-12 Published online:2026-09-01
  • Supported by:
    This study was supported by the Key R&D Program of Anhui Province (2023n06020006), the Anhui Provincial Science and Technology Innovation Tackling Plan (202523anull050002), the Yangtze River Delta Science and Technology Innovation Community Joint Research (Basic Research) Project (2024CSJZN01100), the Youth Project C of the National Natural Science Foundation of China (32401738), and the Youth Promotion Association Program of the Chinese Academy of Sciences (2022454).

摘要: 水稻叶片形态是影响光合效率与群体产量的关键农艺性状。本研究利用重离子束辐射诱变粳稻品种武运粳7号(wyj7),获得一个稳定遗传的窄叶突变体nal24。与野生型相比,nal24突变体的叶片宽度极显著降低,且株高变矮、分蘖数增多。组织学分析表明,其窄叶表型源于大、小叶脉数量减少以及小叶脉间距缩小。光合特性测定显示,nal24突变体在不同光强下的净光合速率均显著低于野生型,表明其光合作用能力降低。遗传分析表明,该性状受单隐性核基因控制。利用图位克隆技术,将目的基因nal24精细定位于1号染色体上77.6 kb的物理区间内,测序发现该区间内编码磷脂酰丝氨酸合酶的基因LOC_Os01g02890发生了单碱基替换(T→A),导致提前形成终止密码子。通过CRISPR/Cas9技术在野生型中敲除该基因,植株表现出窄叶表型,回补材料也恢复了野生型叶宽。表达分析显示,nal24突变体中多个NAL家族基因的表达显著下调,而叶片宽度负调控基因WL1显著上调。本研究克隆了调控水稻叶脉发育与叶片形态建成的新基因NAL24,为解析水稻叶形建成的分子机制提供了新的理论依据与基因资源。

关键词: 窄叶, NAL24, 基因定位, 水稻, 叶脉发育

Abstract: Rice leaf morphology is a key agronomic trait affecting photosynthetic efficiency and population yield. In this study, a stably inherited narrow-leaf mutant nal24 was obtained by heavy-ion beam irradiation mutagenesis of the japonica rice cultivar Wuyunjing 7 (wyj7). Compared with the wild type, the nal24 mutant exhibited a highly significant reduction in leaf width, accompanied by decreased plant height and increased tiller number. Histological analysis revealed that the narrow-leaf phenotype was caused by reduced numbers of large and small veins as well as reduced distances between small veins. Measurements of photosynthetic characteristics showed that the net photosynthetic rate of nal24 was markedly lower than that of the wild type under different light intensities, indicating reduced photosynthetic capacity. Genetic analysis demonstrated that this trait was controlled by a single recessive nuclear gene. The target gene NAL24 was finely mapped to a 77.6 kb physical interval on chromosome 1 via map-based cloning. Sequencing results identified a single base substitution (T→A) in LOC_Os01g02890, which encodes a phosphatidylserine synthase within this interval, resulting in a premature termination codon. Knockout of this gene in wild-type plants using the CRISPR/Cas9 system produced a narrow-leaf phenotype, and genetic complementation restored wild-type leaf width. Transcriptional analysis indicated that the expression levels of multiple NAL family genes were significantly downregulated in the nal24 mutant, whereas the negative regulator of leaf width, WL1, was significantly upregulated. In conclusion, this study cloned the novel gene NAL24 that regulates rice vein development and leaf morphogenesis. The identification of this gene provides a theoretical basis and genetic resources for elucidating the molecular mechanisms underlying rice leaf shape formation.

Key words: narrow leaf, NAL24, gene mapping, rice, leaf vein development

[1] Cho S H, Kang K, Lee S H, et al. OsWOX3A is involved in negative feedback regulation of the gibberellic acid biosynthetic pathway in rice (Oryza sativa). J Exp Bot, 2016, 67: 1677–1687.

[2] Lin L H, Zhao Y F, Liu F, et al. Narrow leaf 1 (NAL1) regulates leaf shape by affecting cell expansion in rice (Oryza sativa L.). Biochem Biophys Res Commun, 2019, 516: 957–962.

[3] Fujino K, Matsuda Y, Ozawa K, et al. NARROW LEAF 7 controls leaf shape mediated by auxin in rice. Mol Genet Genom, 2008, 279: 499–507.

[4] Zhang Y, Wei M, You J, et al. WL1 regulates grain width development in rice. Plant J, 2025, 123: e70418.

[5] Liu J J, Shi X A, Chang Z Y, et al. Auxin efflux transporters OsPIN1c and OsPIN1d function redundantly in regulating rice (Oryza sativa L.) panicle development. Plant Cell Physiol, 2022, 63: 305–316.

[6] 龙海馨, 邱海阳, Muhammad UZAIR, 等. 水稻窄叶突变体nal20的表型分析与基因定位. 作物学报, 2018, 44: 1301–1310.
Long H X, Qiu H Y, Uzair M, et al. Phenotypic analysis and gene mapping of the rice narrow-leaf mutant nal20. Acta Agron Sin, 2018, 44: 1301–1310 (in Chinese with English abstract).

[7] 朱忠林, 文月, 周棋, 等. 水稻OsCNGC10基因抗倒伏性以及抗旱性功能研究. 作物学报, 2024, 50: 1351–1360.
Zhu Z L, Wen Y, Zhou Q, et al. Mechanism of loding residence and drought tolerance of OsCNGC10 gene in rice. Acta Agron Sin, 2024, 50: 1351–1360 (in Chinese with English abstract).

[8] 韩操, 张丽娜, 颜南, 等. BMP-2/bFGF双基因转染的骨髓间充质干细胞复合生物陶瓷骨促进兔桡骨缺损修复. 中国医科大学学报, 2022, 51(2): 125–130.
Han C, Zhang L N, Yan N, et al. Repair efficacy of bioceramic bone composited with BMP-2/bFGF double gene-transfected bone marrow stem cells on radial defects in rabbits. J China Med Univ, 2022, 51(2): 125–130 (in Chinese with English abstract).

[9] Gong X T, Chen J, Chen Y X, et al. Advancements in rice leaf development research. Plants, 2024, 13: 904.

[10] 杨晔, 孙琦, 形欣欣, 等. 包穗突变体sui1-5鉴定及OsPSS1互作蛋白筛选. 作物学报, 2023, 49: 597–607.
Yang Y, Sun Q, Xing X X, et al. Identification of sheathed panicle mutant sui1-5 and screening of OsPPS1 interaction protein in rice. Acta Agron Sin, 2023, 49: 597–607 (in Chinese with English abstract).

[11] 李文浩, 朱晓彤, 罗会超, 等. 水稻黄叶少分蘖突变体yllt10的遗传与定位. 生物工程学报, 2024, 40: 3471–3484.
Li W H, Zhu X T, Luo H C, et al. Heredity and fine mapping of a yellow leaf and less tillering mutant yllt10 in rice. Chin J Biotechnol, 2024, 40: 3471–3484 (in Chinese with English abstract).

[12] 廉院训. 水稻斑马叶基因ZL7的基因克隆和功能分析. 浙江师范大学硕士学位论文, 浙江金华, 2023.
Lian Y X. Gene Cloning and Functional Analysis of Zebra Leaf Gene ZL7 in Rice. MS Thesis of Zhejiang Normal University, Jinhua, Zhejiang, China, 2023 (in Chinese with English abstract).

[13] 陈春鹏. 水稻小粒突变体rgl2的基因定位与功能分析. 中国科学技术大学硕士学位论文, 安徽合肥, 2026.
Chen C P. Gene Mapping and Functional Analysis of the Small Grain Mutant rgl2 in Rice. MS Thesis of University of Science and Technology of China, Hefei, Anhui, China, 2026 (in Chinese with English abstract).

[14] Yao H Y, Xue H W. Phosphatidic acid plays key roles regulating plant development and stress responses. J Integr Plant Biol, 2018, 60: 851–863.

[15] 安国勇, 丁秀艳, 武桂丽, 等. ECS1参与调节CO2诱导的拟南芥气孔关闭和H2O2积累. 植物学报, 2012, 47: 209–216.
An G Y, Ding X Y, Wu G L, et al. ECS1 mediates CO2-induced stomatal closure and the production of H2O2 in Arabidopsis thaliana. Chin Bull Bot, 2012, 47: 209–216 (in Chinese with English abstract).

[16] Guo X P, Xiao X Y, Dong X X, et al. Heavy ion beam irradiation-induced mutational profiles in Saccharomyces cerevisiae and their dependencies on dose and intracellular state inform an enhanced microbial breeding strategy. J Biotechnol, 2026, 410: 236–257.

[17] 徐静, 王莉, 钱前, 等. 水稻叶片形态建成分子调控机制研究进展. 作物学报, 2013, 39: 767–774.
Xu J, Wang L, Qian Q, et al. Research adbance in molecule in regulation mechanism of leaf morphogenesis in rice, 2013, 39: 767–774 (in Chinese with English abstract).

[18] Li D Y, Chen H W, Vale G, et al. Molecular insights into human phosphatidylserine synthase 2 and its regulation of SREBP pathways. Proc Natl Acad Sci USA, 2025, 122: e2501177122.

[19] Shen W Q, Sun J J, Xiao Z, et al. Narrow and stripe leaf 2 regulates leaf width by modulating cell cycle progression in rice. Rice, 2023, 16: 20.

[20] 朱晓彤. 三个水稻类病变突变体的鉴定及其基因定位分析. 浙江师范大学硕士学位论文, 浙江金华, 2023.
Zhu X T. Identification and Gene Mapping Analysis of Three Rice Lesion Mimic Mutant. MS Thesis of Zhejiang Normal University, Jinhua, Zhejiang, China, 2023 (in Chinese with English abstract).

[21] Xu Y C, Yan S Y, Jiang S, et al. Identification of a rice leaf width gene narrow leaf 22 (NAL22) through genome-wide association study and gene editing technology. Int J Mol Sci, 2023, 24: 4073.

[22] Wang Y, Xu W X, Liu Y, et al. Identification and transcriptome analysis of a novel allelic mutant of NAL1 in rice. Genes, 2024, 15: 325.

[23] Yin H F, Gao P, Liu C W, et al. SUI-family genes encode phosphatidylserine synthases and regulate stem development in rice. Planta, 2013, 237: 15–27.

[24] Ma J, Cheng Z J, Chen J, et al. Phosphatidylserine synthase controls cell elongation especially in the uppermost internode in rice by regulation of exocytosis. PLoS One, 2016, 11: e0153119.

[25] 杨永义, 周学标, 姚方印, 等. 水稻窄叶突变体zy103突变基因鉴定. 核农学报, 2019, 33: 421–431.
Yang Y Y, Zhou X B, Yao F Y, et al. Gene identification of narrow leaf mutant zy103 in rice. J Nucl Agric Sci, 2019, 33: 421–431 (in Chinese with English abstract).

[26] Xie K Z, Niu F A, Hu P, et al. Phenotypic identification and fine-mapping of the rice narrow-leaf mutant nal25. Plants, 2025, 14: 2528.

[27] 贾彬, 陈可, 叶婵娟, 等. 水稻叶形态的建成及分子机制研究进展. 中国稻米, 2024, 30(5): 19–29.
Jia B, Chen K, Ye C J, et al. Progress in the establishment of rice leaf morphology and molecular mechanism. China Rice, 2024, 30(5): 19–29 (in Chinese with English abstract).

[28] Jiang D, Fang J J, Lou L M, et al. Characterization of a null allelic mutant of the rice NAL1 gene reveals its role in regulating cell division. PLoS One, 2015, 10: e0118169.

[29] 赫磊, 路凯, 赵春芳, 等. 水稻穗顶端退化突变体paa21的表型分析及基因克隆. 中国农业科学, 2022, 55: 4781–4792.
He L, Lu K, Zhao C F, et al. Phenotypic analysis and gene cloning of rice panicle apical abortion mutant paa21. Sci Agric Sin, 2022, 55: 4781–4792 (in Chinese with English abstract).

[30] Liu J J, Shi X A, Zhong T H, et al. PINOID and PIN-FORMED paralogous genes are required for leaf morphogenesis in rice. Plant Cell Physiol, 2023, 64: 1146–1158.

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