远红光介导茉莉酸调控番茄种子萌发的分子机制

Molecular Mechanism of Far-Red Light-Mediated Jasmonate Regulation of Tomato Seed Germination

摘要

种子萌发是植物生长发育的关键生理阶段，对农业生产具有重要影响。番茄（ Solanum lycopersicum）作为全球主要蔬菜作物之一，其种子休眠与萌发的调控机制的解析，不仅有助于提升番茄产业效益，也为理解植物种子萌发的分子机理提供了新的视角和参考依据。本研究以番茄栽培品种以及茉莉酸合成和信号传导途径的突变体作为试验材料，探究茉莉酸和远红光对种子萌发的影响。试验结果显示：外施低浓度（1 μM）茉莉酸可促进番茄种子萌发，而高浓度（5-100 μM）则显著抑制；通过茉莉酸合成和信号传导途径遗传材料表明，茉莉酸合成途径和信号途径均有抑制番茄种子萌发的作用；远红光抑制番茄种子萌发，可通过抑制茉莉酸生物合成进而提高脱落酸（ABA）积累水平，最终实现对种子萌发的调控。进一步的试验表明，该过程依赖于茉莉酸信号途径核心转录因子MYC2的介导，在 MYC2-RNAi种子中，远红光处理和黑暗条件下，ABA含量差异消失，同时远红光对 SlDOG1（番茄萌发延迟蛋白基因）、 SlNCED1（番茄ABA生物合成关键基因）和 SlGAI（番茄赤霉素不敏感基因）等关键基因的显著上调作用也不复存在。本研究揭示了番茄种子萌发的光信号调控，依赖于茉莉酸信号途径对ABA合成的调控，为作物遗传改良和种子处理技术提供了理论支撑。

Abstract

Seed germination is a critical physiological stage in plant growth and development, with significant implications for agricultural production. Tomato ( Solanum lycopersicum), as a globally important vegetable crop, elucidating the regulatory mechanisms governing its seed dormancy and germination not only enhances tomato industry efficiency but also provides new perspectives and references for understanding the molecular basis of seed germination in plants. This study utilizes cultivated tomato varieties and mutants in jasmonic acid (JA) biosynthesis and signaling pathways as experimental materials to investigate the effects of JA and far-red light (FR) on seed germination. Experimental results demonstrate that exogenous application of low-concentration JA (1 μM) promotes tomato seed germination, whereas higher concentrations (5-100 μM) significantly inhibit it. Genetic evidence from JA biosynthesis and signaling mutants reveal that both pathways exert inhibitory effects on seed germination. Far-red light suppresses tomato seed germination through downregulation of JA biosynthesis, which subsequently elevates abscisic acid (ABA) accumulation, ultimately regulating germination. Further experiments show that this regulatory process depends on MYC2, a core transcription factor in JA signaling. In MYC2-RNAi seeds, the differential ABA content between far-red light treatment and dark conditions disappear, accompanied by the elimination of far-red light-induced upregulation of key genes including SlDOG1 (a tomato delayed germination protein gene), SlNCED1 (a pivotal ABA biosynthesis gene), and SlGAI (a gibberellin-insensitive gene). This study uncovers that the photoregulation of tomato seed germination is mediated through JA signaling pathway-dependent modulation of ABA biosynthesis, providing a theoretical foundation for crop genetic improvement and seed treatment technologies.

关键词

番茄 / 种子萌发 / 远红光 / 茉莉酸

Key words

Tomato / seed germination / far-red light / jasmonic acid

参考文献

[1] Nuruddin M M, Madramootoo C A, Dodds G T . Effects of water stress at different growth stages on greenhouse tomato yield and quality[J]. HortScience, 2003, 38(7): 1389-1393.

[2] Sun X, Tian R, Zhao M, et al. MtCIR2 negatively regulates seed germination to salt stress by disrupting metabolisms and signaling of abscisic acid and gibberellins[J]. Plant Physiology and Biochemistry, 2025, 220: 109493.

[3] 李飞飞, 刘杨, 徐飞. 种子萌发的奥秘:生理、生态与分子的综合解析[J]. 生命科学, 2024, 36(12): 1470-1477.

[4] Oh E, Yamaguchi S, Hu J, et al. PIL5, a phytochrome-interacting bHLH protein, regulates gibberellin responsiveness by binding directly to the GAI and RGA promoters in Arabidopsis seeds [J]. The Plant Cell, 2007, 19(4): 1192-1208.

[5] Kim W, Lee Y, Park J, et al. HONSU, a protein phosphatase 2C, regulates seed dormancy by inhibiting ABA signaling in Arabidopsis [J]. Plant and Cell Physiology, 2013, 54(4): 555-572.

[6] Jacobsen J V, Barrero J M, Hughes T, et al. Roles for blue light, jasmonate and nitric oxide in the regulation of dormancy and germination in wheat grain (Triticum aestivum L.) [J]. Planta, 2013, 238(1): 121-138.

[7] Zhiren C, Yuan D, Xi H . Plant responses to UV-B radiation: signaling, acclimation and stress tolerance [J]. Stress Biology, 2022, 2(1): 51- 51.

[8] Holdsworth M J, Bentsink L, Soppe W J . Molecular networks regulating Arabidopsis seed maturation, after-ripening, dormancy and germination[J]. New Phytologist, 2008, 179(1): 33-54.

[9] Razzaq K, Du J . Phytohormonal regulation of plant development in response to fluctuating light conditions[J]. Journal of Plant Growth Regulation, 2024, (11): 1-34.

[10] Chen L, Lu B, Liu L, et al. Melatonin promotes seed germination under salt stress by regulating ABA and GA3 in cotton (Gossypium hirsutum L.) [J]. Plant Physiology and Biochemistry, 2021, 162: 506-516.

[11] Vaistij F E, Gan Y, Penfield S, et al. Differential control of seed primary dormancy in Arabidopsis ecotypes by the transcription factor SPATULA [J]. Proceedings of the National Academy of Sciences of the United States of America, 2013, 110(26): 10866-10871.

[12] Frey A, Effroy D, Lefebvre V, et al. Epoxycarotenoid cleavage by NCED5 fine-tunes ABA accumulation and affects seed dormancy and drought tolerance with other NCED family members[J]. The Plant Journal, 2012, 70(3): 501-512.

[13] Wang Y, Fan Y, Fan D, et al. The noncoding RNA HIDDEN TREASURE 1 promotes phytochrome B-dependent seed germination by repressing abscisic acid biosynthesis[J]. The Plant Cell, 2023, 35(2): 700-716.

[14] Yu H, Wang Y, Wang Y, et al. The Arabidopsis RING‐Type E3 ligase TEAR4 controls seed germination by targeting RGA for degradation[J]. Advanced Science, 2025, 12(4): 2400116.

[15] Panda D, Mohanty S, Das S, et al. The role of phytochrome-mediated gibberellic acid signaling in the modulation of seed germination under low light stress in rice (O. sativa L.) [J]. Physiology and Molecular Biology of Plants, 2022, 28(3): 585-605.

[16] 刘慧娜, 宾金华. 茉莉酸甲酯对花生种子萌发的抑制[J]. 嘉应大学学报, 1999(06): 51-54.

[17] 宾金华, 黄胜琴, 何树春, 等. 茉莉酸甲酯对水稻种子萌发和贮藏物质降解的影响[J]. 植物学报, 2001, 43(06): 578-585.

[18] Nambara E, Okamoto M, Tatematsu K, et al. Abscisic acid and the control of seed dormancy and germination [J]. Seed Science Research, 2010, 20(2): 55-67.

[19] 胡海英, 贺海明, 梁新华, 等. 茉莉酸甲酯对乌拉尔甘草种子萌发和过氧化物酶活性的影响[J]. 时珍国医国药, 2011, 22(07): 1579-1580.

[20] 邹燕, 王瑞雪, 沈亮余. 渗透胁迫下外源茉莉酸甲酯对油菜种子生理特性的影响[J]. 西北植物学报, 2011(03): 564-568.

[21] Ruan J, Zhou Y, Zhou M, et al. Jasmonic acid signaling pathway in plants[J]. International Journal of Molecular Sciences, 2019, 20(10): 2479.

[22] Kazan K, Manners J M . MYC2: The master in action [J]. Molecular Plant, 2013, 6(3): 686-703.

[23] Fernandez-Arbaizar A, Regalado J J, Lorenzo O . Isolation and characterization of novel mutant loci suppressing the ABA hypersensitivity of the Arabidopsis coronatine insensitive 1-16 (coi1-16) mutant during germination and seedling growth [J]. Plant and Cell Physiology, 2012, 53(1): 53-63.

[24] Ju L, Jing Y, Shi P, et al. JAZ proteins modulate seed germination through interaction with ABI 5 in bread wheat and Arabidopsis [J]. New Phytologist, 2019, 223(1): 246-260.

[25] Pan J, Hu Y, Wang H, et al. Molecular mechanism underlying the synergetic effect of jasmonate on abscisic acid signaling during seed germination in Arabidopsis [J]. The Plant Cell, 2020, 32(12): 3846-3865.

[26] Mei S, Zhang M, Ye J, et al. Auxin contributes to jasmonate-mediated regulation of abscisic acid signaling during seed germination in Arabidopsis [J]. The Plant Cell, 2023, 35(3): 1110-1133.

[27] Yi R, Yan J, Xie D . Light promotes jasmonate biosynthesis to regulate photomorphogenesis in Arabidopsis [J]. Science China Life Sciences, 2020, 63(7): 943-952.

[28] Chen H, Jones A D, Howe G A . Constitutive activation of the jasmonate signaling pathway enhances the production of secondary metabolites in tomato[J]. FEBS Letters, 2006, 580(11): 2540-2546.

[29] Wasternack C, Hause B . Jasmonates: biosynthesis, perception, signal transduction and action in plant stress response, growth and development[J]. An Update to the 2007 Review in Annals of Botany. Annals of Botany, 2013, 111(6): 1021-1058.

[30] Shu K, Zhang H, Wang S, et al. ABI4 regulates primary seed dormancy by regulating the biogenesis of abscisic acid and gibberellins in Arabidopsis [J]. PLoS Genetics., 2013, 9: e1003577.

[31] Wang Y, Fan Y, Fan D, et al. The noncoding RNA HIDDEN TREASURE1 promotes phytochrome B-dependent seed germination by repressing abscisic acid biosynthesis [J]. The Plant Cell, 2022, 35(2): 700-716.

[32] Shinomura T, Nagatani A, Hanzawa H, et al. Action spectra for phytochrome A- and B-specific photoinduction of seed germination in Arabidopsis thaliana [J]. Proceedings of the National Academy of Sciences of the United States of America, 1996, 93(15): 8129-8133.

[33] Pauwels L, Barbero G F, Geerinck J, et al. NINJA connects the co-repressor TOPLESS to jasmonate signaling[J]. Nature, 2010, 464(7289): 788-791.

[34] Tiancong Q, Susheng S, Qingcuo R, et al. The Jasmonate-ZIM-domain proteins interact with the WD-Repeat/bHLH/MYB complexes to regulate Jasmonate-mediated anthocyanin accumulation and trichome initiation in Arabidopsis thaliana . [J]. The Plant cell, 2011, 23(5): 1795-814.

[35] 钱双杰, 吴刘清, 邓雨琦, 等 . 赤霉素关键蛋白 DELLA 对植物雄蕊发育研究进展[J]. 上海师范大学学报 (自然科学版), 2023, 52(06): 736-742.

[36] Michael W, Jean-Michel D, Soizic C, et al. The Arabidopsis DELLA RGA-LIKE3 is a direct target of MYC2 and modulates jasmonate signaling responses[J]. The Plant cell, 2012, 24(8): 3307-19.

[37] 厉书豪. 远红光和赤霉素互作调控黄瓜幼苗节间伸长的机理研究[D].福州:福建农林大学, 2024.

基金资助

双一流建设专项 茉莉酸调控番茄种子休眠(20170921)