豆科植物种子休眠的生理及分子机制研究进展

doi:10.11686/cyxb2025337

摘要/Abstract

摘要：

种子休眠是植物在长期进化过程中形成的一种适应性特征，使植物能够在逆境条件下存活。豆科植物作为全球农业生产的核心作物类群（涵盖粮食、饲料、绿肥等），其种子休眠特性直接影响播种质量、田间出苗率及产量稳定性。豆科植物种子休眠的分子调控主要以脱落酸（ABA）和赤霉素（GA）含量平衡为核心，通过种皮发育基因、转录因子网络、表观修饰及环境信号整合的复杂系统，该机制既保证种子在不利环境中存活，又能在适宜条件下精准萌发。生理休眠是种子休眠的主要因素，ABA与GA并非直接相互作用，而是通过其拮抗效应分别促进“休眠维持”和“萌发启动”，ABA下游发芽抑制因子ABI3有许多靶基因在拟南芥中已被报道，包括ABI5、DELLA和bHLH转录因子，但在豆科植物中相应的调控网络尚不明确，ABI3在ABA信号传导通路与其他因子的相互作用仍待研究。因此，基于近年来的相关研究成果，系统梳理了豆科植物种子休眠的类型、种子休眠破除技术及休眠的分子调控网络，重点总结豆科植物种子特有的休眠特征。此外，基于模式植物种子休眠的分子调控研究进展，提出豆科植物种子休眠的潜在分子调控假设模型，为豆科植物种子休眠分子调控机制的进一步研究提供理论指导，也为豆科植物栽培实践和后续品种改良提供理论支撑。

关键词: 豆科植物种子, 物理休眠, 生理休眠, 种子休眠解除, 分子调控机制

Abstract:

Seed dormancy is an adaptive trait that developed during long-term evolution， enabling plants to survive under adverse conditions. Legumes are a core crop group in global agricultural production （encompassing food， forage， and green manure）， and the dormancy characteristics of their seeds directly affect sowing quality， field emergence rate， and yield stability. The molecular regulation of seed dormancy in legumes primarily revolves around the balance between abscisic acid （ABA） and gibberellin （GA） levels， involving a complex system that integrates the expression of genes involved in seed coat development and transcription factor networks， as well as epigenetic modifications and environmental signals. This mechanism ensures seed survival under unfavorable conditions while enabling precise germination under suitable circumstances. Physiological dormancy is the primary factor in seed dormancy， where ABA and GA do not directly interact but instead exhibit antagonistic effects-ABA promotes “dormancy maintenance” while GA triggers “germination initiation”. In Arabidopsis thaliana， many downstream targets of the ABA-responsive germination inhibitor ABI3 have been identified， including the ABI5， DELLA， and bHLH transcription factors. However， the corresponding regulatory network in legumes remains unclear， and further research is required to explore the interactions between ABI3 and other factors in the ABA signaling pathway. Based on recent research， this paper systematically reviews the types of seed dormancy in legumes， dormancy-breaking techniques， and molecular regulatory networks， with a focus on legume-specific dormancy traits. Furthermore， drawing on advances in research on the molecular regulation of seed dormancy in model plants， we propose a hypothetical model for the molecular regulation of seed dormancy in legumes. This review provides a theoretical foundation for understanding the molecular mechanisms of seed dormancy in legumes while offering insights for cultivation practices and varietal improvement.

Key words: legume seeds, physical dormancy, physiological dormancy, seed dormancy release, molecular regulatory mechanisms

刘昊臻, 赵士钦, 冯树蓉, 王成, 张景鈜, 孙守江. 豆科植物种子休眠的生理及分子机制研究进展[J]. 草业学报, 2026, 35(8): 207-220.

Hao-zhen LIU, Shi-qin ZHAO, Shu-rong FENG, Cheng WANG, Jing-hong ZHANG, Shou-jiang SUN. Progress in research on the physiological and molecular mechanisms of seed dormancy in legumes[J]. Acta Prataculturae Sinica, 2026, 35(8): 207-220.

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类型Type	种子休眠原因Reasons for seed dormancy	文献References
PD	生理休眠一般由激素平衡来解释，其中脱落酸作为抑制剂，赤霉素作为促进剂。Physiological dormancy is typically elucidated through the lens of hormonal equilibrium， wherein abscisic acid （ABA） functions as an inhibitory regulator， while gibberellins （GA） serve as promotive agents.	［23］
PY	种子的不透水性导致种子不能吸收水分，因此不能发芽。The impermeability of the seed coat prevents water absorption， thereby inhibiting germination.	［33］
PY+PD	种子同时存在两种或多种休眠类型，既具有生理休眠的特点，内部含有抑制萌发的物质，又有物理休眠，种皮阻碍了种子与外界环境的物质交换。Seeds may exhibit multiple types of dormancy concurrently， characterized by both physiological dormancy， involving internal substances that inhibit germination， and physical dormancy， where the seed coat impedes material exchange with the external environment.	［37］

类型Type	种子休眠原因Reasons for seed dormancy	文献References
PD	生理休眠一般由激素平衡来解释，其中脱落酸作为抑制剂，赤霉素作为促进剂。Physiological dormancy is typically elucidated through the lens of hormonal equilibrium， wherein abscisic acid （ABA） functions as an inhibitory regulator， while gibberellins （GA） serve as promotive agents.	［23］
PY	种子的不透水性导致种子不能吸收水分，因此不能发芽。The impermeability of the seed coat prevents water absorption， thereby inhibiting germination.	［33］
PY+PD	种子同时存在两种或多种休眠类型，既具有生理休眠的特点，内部含有抑制萌发的物质，又有物理休眠，种皮阻碍了种子与外界环境的物质交换。Seeds may exhibit multiple types of dormancy concurrently， characterized by both physiological dormancy， involving internal substances that inhibit germination， and physical dormancy， where the seed coat impedes material exchange with the external environment.	［37］

解除技术 Release technology	种子休眠解除原理 Seed dormancy release mechanisms	解除类型 Categories of dormancy release
层积处理Stratification treatment	软化或分解种皮、促使种子内部发生复杂的生理生化变化。此外，完成胚的发育。Softening or decomposing the seed coat， promoting complex physiological and biochemical changes within the seed， and in addition， completing the development of the embryo.	PD， PY+PD
机械处理Mechanical treatment	物理外力破坏种子坚硬或致密的外皮以解除其对水分、气体渗透或胚生长的阻碍，从而打破休眠。Physical external forces destroy the hard or dense outer coat of seeds to remove their obstruction to water and gas penetration or embryo growth， thereby breaking dormancy.	PY
热水处理Hot water treatment	高温使种皮蜡质层破裂或产生微裂纹，增强透水性，热激改变种子内源激素平衡。High temperature causes the waxy layer of the seed coat to crack or form microcracks， enhances water permeability， and heat shock alters the balance of endogenous hormones in seeds.	PY， PD
激素处理Hormonal treatment	抑制脱落酸作用，促进赤霉素合成、刺激胚发育。Inhibit the action of abscisic acid（ABA）， promote the synthesis of gibberellin（GA）， and stimulate embryo development.	PD
硫酸处理Sulfuric acid treatment	降低种壳机械强度，破坏种皮的一些局部结构、改善种皮的渗透性、急剧降温导致种皮开裂，进而提升种皮通透性。Reduce the mechanical strength of the seed coat， destroy some local structures of the seed coat， improve the permeability of the seed coat， and cause the seed coat to crack due to rapid cooling， thereby enhancing the permeability of the seed coat.	PY
液氮处理Liquid nitrogen treatment	使种皮产生微裂纹，或改变细胞膜透性及酶活性。Cause the seed coat to form microcracks， or alter the permeability of cell membranes and enzyme activity.	PD， PY
超声波处理Ultrasonic wave treatment	利用超声波在液体中产生空化气泡，气泡破裂时释放冲击波，使种皮产生微孔或剥离外层屏障。Ultrasonic waves are used to generate cavitation bubbles in liquids； when the bubbles collapse， they release shock waves， causing the seed coat to form micropores or peel off the outer barrier.	PY

解除技术 Release technology	种子休眠解除原理 Seed dormancy release mechanisms	解除类型 Categories of dormancy release
层积处理Stratification treatment	软化或分解种皮、促使种子内部发生复杂的生理生化变化。此外，完成胚的发育。Softening or decomposing the seed coat， promoting complex physiological and biochemical changes within the seed， and in addition， completing the development of the embryo.	PD， PY+PD
机械处理Mechanical treatment	物理外力破坏种子坚硬或致密的外皮以解除其对水分、气体渗透或胚生长的阻碍，从而打破休眠。Physical external forces destroy the hard or dense outer coat of seeds to remove their obstruction to water and gas penetration or embryo growth， thereby breaking dormancy.	PY
热水处理Hot water treatment	高温使种皮蜡质层破裂或产生微裂纹，增强透水性，热激改变种子内源激素平衡。High temperature causes the waxy layer of the seed coat to crack or form microcracks， enhances water permeability， and heat shock alters the balance of endogenous hormones in seeds.	PY， PD
激素处理Hormonal treatment	抑制脱落酸作用，促进赤霉素合成、刺激胚发育。Inhibit the action of abscisic acid（ABA）， promote the synthesis of gibberellin（GA）， and stimulate embryo development.	PD
硫酸处理Sulfuric acid treatment	降低种壳机械强度，破坏种皮的一些局部结构、改善种皮的渗透性、急剧降温导致种皮开裂，进而提升种皮通透性。Reduce the mechanical strength of the seed coat， destroy some local structures of the seed coat， improve the permeability of the seed coat， and cause the seed coat to crack due to rapid cooling， thereby enhancing the permeability of the seed coat.	PY
液氮处理Liquid nitrogen treatment	使种皮产生微裂纹，或改变细胞膜透性及酶活性。Cause the seed coat to form microcracks， or alter the permeability of cell membranes and enzyme activity.	PD， PY
超声波处理Ultrasonic wave treatment	利用超声波在液体中产生空化气泡，气泡破裂时释放冲击波，使种皮产生微孔或剥离外层屏障。Ultrasonic waves are used to generate cavitation bubbles in liquids； when the bubbles collapse， they release shock waves， causing the seed coat to form micropores or peel off the outer barrier.	PY