NO调控植物种子休眠和萌发的研究进展

doi:10.13560/j.cnki.biotech.bull.1985.2025-1040

摘要/Abstract

摘要：

种子是农业的“芯片”，其萌发作为作物生命周期的起点，直接决定作物出苗质量及产量品质。一氧化氮（nitric oxide, NO）作为一种关键的气体信号分子，在调控种子休眠与萌发中的核心作用已成为植物生物学的研究前沿。本文系统综述了该领域的研究进展，阐释了植物中NO的主要合成与代谢途径及其稳态调控，重点剖析了其信号转导机制，尤其是其通过与激素、活性氧等信号网络的交叉对话以及蛋白质翻译后修饰，协同调控种子休眠与萌发关键节点的分子基础。尽管相关研究已取得显著进展，该领域仍存在若干重要科学问题亟待深入探索。未来在基础研究层面，需进一步解析NO合成、代谢与信号在种子不同组织和萌发阶段的时空动态调控网络；利用多组学技术系统鉴定其下游修饰靶点并阐明功能；揭示NO与光、温等环境信号整合的分子机制。在应用层面，可基于NO调控原理开发新型绿色种子处理技术，提升种子的逆境萌发能力与幼苗抗逆性；通过遗传或生物技术手段调控种子内源NO合成代谢通路，改良作物种子萌发特性，为培育出苗整齐、抗逆性强的新品种提供新策略。本综述通过系统梳理NO在植物种子休眠与萌发中的调控机制，旨在为深入理解种子生命起始的分子网络提供理论依据，并为作物种子品质提升与抗逆栽培技术创新提供新思路。

关键词: NO, 种子休眠, 种子萌发, 调控机制, 信号传导

Abstract:

Seeds are recognized as the “chips” of agriculture production, and their germination, as the starting point of a crop’s life cycle, directly determines seedling emergence quality as well as crop yield and quality. Nitric oxide (NO), a key gaseous signaling molecule, has emerged as a research frontier in plant biology due to its central role in regulating seed dormancy and germination. This review systematically summarizes the current progress in this field, elucidating the primary synthesis and metabolic pathways of NO in plants and their homeostatic regulation. It focuses on dissecting the signal transduction mechanisms of NO, particularly its molecular basis for coordinately regulating key nodes of seed dormancy and germination through crosstalk with hormonal and reactive oxygen species (ROS) signaling networks as well as through post-translational modifications. Although significant progress has been achieved, several important scientific issues in this area still remain to be deeply explored. At the basic research level, future efforts should further be on resolving the spatiotemporal specificity of NO synthesis and signaling within different seed tissues and germination stages, utilizing multi-omics technologies to systematically identify downstream modification targets of NO and clarify their functions. Additionally, elucidating the molecular mechanisms underlying the integration of NO signaling with environmental signals such as light and temperature is essential. At the applied level, novel green seed treatment technologies based on NO can be developed to enhance seed germination performance and seedling resistance to stress under adverse conditions. Furthermore, exploring genetic or biotechnological approaches to modulate endogenous NO synthesis and metabolism pathways in seeds could improve crop germination traits, offering new strategies for breeding crop varieties with uniform emergence and enhanced stress resistance. By systematically synthesizing the regulatory mechanisms of NO in seed dormancy and germination, this review aims to provide a theoretical foundation for a deeper understanding of the molecular regulatory network governing the initiation of seed life, while also offering new insights for the development of key technologies to enhance crop seed quality and innovate stress-resilient cultivation models.

Key words: NO, seed dormancy, seed germination, regulatory mechanism, signal transduction

王红阳, 邱艳红, 王德欣, 夏阳, 孟淑春, 徐秀兰, 张海军. NO调控植物种子休眠和萌发的研究进展[J]. 生物技术通报, doi: 10.13560/j.cnki.biotech.bull.1985.2025-1040.

WANG Hong-yang, QIU Yan-hong, WANG De-xin, XIA Yang, MENG Shu-chun, XU Xiu-lan, ZHANG Hai-jun. Research Progress in NO Regulating Seed Dormancy and Germination[J]. Biotechnology Bulletin, doi: 10.13560/j.cnki.biotech.bull.1985.2025-1040.

图/表 4

图1 NO的合成与清除虚线左侧为NO的合成过程，虚线右侧为NO的清除过程。NR：硝酸还原酶（例如拟南芥AtNIA1/2）；mETC：线粒体电子传递链；L-Citrulline：L-瓜氨酸；GSNO：S-亚硝基谷胱甘肽；NO2-FAs：硝基脂肪酸；NO-Mela：亚硝基褪黑素；Phytoglobins：植物血红蛋白；GSSG：谷胱甘二硫化物；ONOO-：过氧亚硝酸根离子

Fig. 1 Biosynthesis and scavenging of NOThe left side of the dashed line shows the synthesis process of NO, while the right side illustrates the scavenging process of NO. NR: Nitrate reductase (e.g., AtNIA1/2); mETC: mitochondrial electron transport chain; GSNO: S-nitrosoglutathione; NO2-FAs: nitro fatty acids; NO-Mela: N-nitrosomelatonin; GSSG: glutathione disulfide; ONOO-: peroxynitrite

图2 NO与植物激素互作协同调控种子休眠及萌发NO通过抑制ABA合成及信号传递，促进GA、JA和乙烯的合成和信号传递，解除种子休眠并促进种子萌发。Nitration：硝化；S-nitrosylation：S-亚硝基化；CYP707A：细胞色素P450，707家族A亚家族；NCED：9-顺式-环氧类胡萝卜素双加氧酶；PYR/PYL/RCAR：ABA受体；PP2C：2C型蛋白磷酸酶；SnRKs：SNF1相关蛋白激酶；ABI5：脱落酸不敏感蛋白5；GAox：赤霉素氧化酶；RGL2：赤霉素类似物抑制因子2；ACS：S-腺苷-L-甲硫氨酸甲基硫腺苷裂解酶；ACO：ACC氧化酶；ETR：乙烯抗性蛋白1；CTR1：组成型三重反应蛋白1；EIN2：乙烯不敏感蛋白2；AOS：丙二烯氧化物合成酶。本图中的种子模型为示意图，所展示的调控机制在单子叶植物（如水稻）和双子叶植物（如拟南芥）中均存在

Fig. 2 Role of NO and phytohormones interaction in synergistically regulating seed dormancy and germinationNO promotes seed germination and breaks dormancy by antagonizing ABA biosynthesis and signaling, while concurrently activating the biosynthesis and signaling pathways of GA, JA, and ethylene. CYP707A: Cytochrome P450, Family 707, Subfamily A; NCED: nine-cis-epoxycarotenoid dioxygenase; PYR/PYL/RCAR: PYRABACTIN RESISTANCE1 (PYR1)/PYR1-LIKE (PYL)/REGULATORY COMPONENTS OF ABA RECEPTORS (RCAR) ; PP2C: type 2C protein phosphatases; SnRKs: SNF1-related protein kinase; ABI5: ABSCISIC ACID INSENSITIVE 5; GAox: GA oxidase; RGL2: REPRESSOR OF GA-LIKE2; ACS: S-adenosyl-L-methionine methylthioadenosine-lyase, ACO: ACC oxidase; ETR: ethylene resistant 1; CTR1: constitutive triple response 1; EIN2: ETHYLENE INSENSITIVE2; AOS: allene oxide synthase. The seed schematic represents a general model, and the illustrated NO-mediated mechanisms are operative in both monocot (e.g., rice) and dicot (e.g., Arabidopsis) species

表1 NO直接介导的蛋白翻译后修饰（PTMs）类型

Table 1 Types of protein post-translational modifications (PTMs) directly mediated by NO

PTM类型 Category	修饰蛋白 Protein	修饰位点 Modification sites	功能描述 Function description	参考文献 Reference
S-亚硝基化 S-nitrosylation	APX， GR， DHAR	—	激活抗坏血酸-谷胱甘肽循环中的抗氧化酶活性，提高顽拗型种子的脱水耐受性	［56］
	ABI5	Cys153	S-亚硝基化后，E3连接酶介导其降解，从而促进种子萌发	［57］
	SnRK2.2/ SnRK2.3	Cys137	抑制激酶活性，阻断ABA信号传递，促进种子萌发	［58］
	GSNOR1	—	高温胁迫诱导GSNOR1发生S-亚硝基化修饰并促使其降解，稳定ABI5蛋白导致种子热休眠	［62］
	GADPH	Cys154	诱导蛋白寡聚化，加速细胞死亡和种子老化	［60］
	HFR	Cys164	高温诱导HFR1 S-亚硝基化促使其降解，激活PIF1靶向的SOM基因表达，改变GA与ABA代谢平衡，抑制种子萌发	［61］
	MYB30	Cys49	NO介导MYB30 S-亚硝基化增强MYB30的转录活性，促进CYP707A2表达，降低ABA含量。打破种子休眠、促进萌发	［59］
酪氨酸硝化 Tyrosine nitration	储藏蛋白	—	促进储藏蛋白降解，为胚轴伸长提供必需的氨基酸与维生素，为种子萌发提供能量	［13］
酪氨酸硝化 Tyrosine nitration	PYR/PYL/RCAR	—	硝化使ABA受体失活，限制ABA信号传导	［64］

图3 NO的农业应用潜力在农业生产中，NO通过调控基因表达与蛋白质翻译后修饰，介导包含激素平衡重塑、氧化损伤缓解、细胞膜稳定及能量代谢调控在内的多重过程，最终实现打破种子休眠、促进萌发，显著提升种子的抗逆性，并维持其长期活力

Fig. 3 Agricultural application potential of NOIn agricultural practice, NO modulates gene expression and post-translational modifications, thereby reshaping hormonal balance, scavenging oxidative damage, maintaining membrane integrity, and reprogramming energy metabolism. These coordinated actions collectively break seed dormancy, promote germination, and ultimately enhance seed resistance to stress and maintain long-term vigor

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