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作物学报 ›› 2024, Vol. 50 ›› Issue (1): 1-15.doi: 10.3724/SP.J.1006.2024.34012

• 综述 •    下一篇

种子休眠与萌发调控的研究进展

宋松泉1,3,*(), 唐翠芳2,3, 雷华平3, 姜孝成2, 王伟青1, 程红焱1   

  1. 1中国科学院植物研究所, 北京 100093
    2湖南师范大学生命科学学院, 湖南长沙 410081
    3湘南学院南岭现代种业研究院, 湖南郴州 423099
  • 收稿日期:2022-12-18 接受日期:2023-09-08 出版日期:2024-01-12 网络出版日期:2023-09-08
  • 通讯作者: *宋松泉, E-mail: sqsong2019@163.com
  • 基金资助:
    国家科技支撑计划项目(2012BAC01B05);郴州国家可持续发展议程创新示范区建设省级专项(2022sfq06);郴州市人民政府和湘南学院人才项目资助

Research progress of seed dormancy and germination regulation

SONG Song-Quan1,3,*(), TANG Cui-Fang2,3, LEI Hua-Ping3, JIANG Xiao-Cheng2, WANG Wei-Qing1, CHENG Hong-Yan1   

  1. 1Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China
    2College of Life Sciences, Hunan Normal University, Changsha 410081, Hunan, China
    3Nanling Research Institute for Modern Seed Industry, Xiangnan University, Chenzhou 423099, Hunan, China
  • Received:2022-12-18 Accepted:2023-09-08 Published:2024-01-12 Published online:2023-09-08
  • Contact: *E-mail: sqsong2019@163.com
  • Supported by:
    National Key Technology Support Program of China(2012BAC01B05);Provincial Special Project of Chenzhou National Sustainable Development Agenda Innovation Demonstration Zone Construction(2022sfq06);Chenzhou Municipal People’s Government and Xiangnan University Talent Project

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摘要：

休眠使植物种子能够为萌发安排时间, 直到环境条件变得有利于幼苗的存活与生长。种子的休眠特性具有重要的生态适应性意义和显著的农业价值。植物激素脱落酸(ABA)和赤霉素(GA)是种子休眠与萌发的关键因素, 处于休眠状态的成熟种子含有高水平的ABA和低水平的GA; ABA诱导和维持种子休眠, 而GA拮抗ABA的作用和促进种子萌发。萌发延迟-1 (DOG1)是种子休眠的主要调控因子, 与ABA协同作用延迟萌发。DOG1通过与PP2C ABA过敏感萌发(AHG1/AHG3)结合以增强ABA的信号转导, 以及抑制AHG1的作用以增加ABA的敏感性和强加种子休眠。印记基因在受精前后受到表观遗传机制的调控, 与种子休眠的建立与释放密切相关。近年来, 种子休眠的调控研究取得了显著的进展。本文综述了植物激素ABA和GA对种子休眠与萌发的控制, DOG1调控种子休眠的作用机制, 以及种子休眠与萌发的表观遗传调控, 并提出了在本领域需要进一步研究的科学问题, 以期为深入理解种子休眠与萌发的分子机制、抗穗萌发育种以及促进休眠种子的萌发提供参考。

关键词: DOG1, 表观遗传, 分子机制, 植物激素, 种子休眠

Abstract:

Dormancy enables plant seeds to time germination until environmental conditions become favorable for seedling survival and growth. The dormancy characteristics of seeds are of important ecological adaptive significance and notable agricultural value. Phytohormone abscisic acid (ABA) and gibberellin (GA) are the key factors for seed dormancy and germination. Mature seeds in dormancy state contain high levels of ABA and low levels of GA. ABA induces and maintains seed dormancy, while GA antagonizes ABA and promotes seed germination. DELAY OF GERMINATION-1 (DOG1) is a major regulator of seed dormancy and had a synergistic effect with ABA to delay germination. DOG1 enhances ABA signal transduction by combining with PP2C ABA hypersensitive germination (AHG1/AHG3), and inhibits the action of AHG1 to increase ABA sensitivity and impose seed dormancy. Imprinted genes are regulated by epigenetic mechanisms before and after fertilization, and are closely related to the establishment and release of seed dormancy. In recent years, remarkable progress has been made in the regulation of seed dormancy. In the present paper, we reviewed the effects of phytohormones ABA and GA on seed dormancy and germination, the action mechanism regulating seed dormancy by DOG1, and the epigenetic regulation of seed dormancy and germination. In addition, we also propose some scientific issues that need to be further investigated in this field to provide some information for understanding the molecular mechanism of seed dormancy and germination, breeding in anti-preharvest sprouting in crop plants, and promoting the germination of dormant seeds.

Key words: DOG1, epigenetic, molecular mechanism, phytohormone, seed dormancy

图1

种子休眠与萌发的植物激素调控网络 3种主要的植物激素, 包括脱落酸(ABA)、赤霉素(GA)和生长素(auxin), 是种子休眠与萌发的关键因素。成熟种子处于休眠状态, 含有高水平的ABA和低水平的GA。几种转录因子(ABI4、DDF1、OsAP2-39、AP2和CHO1)通过正调控(+) ABA的积累和降低GA的含量参与种子的休眠。当种子休眠被打破时, 种子就变为非休眠状态, 能够开始萌发。在此阶段, ABA/GA的平衡通过几乎所有其他植物激素的正和负调控信号被维持, 包括乙烯(ET)、油菜素内酯(BR)、茉莉酸(JA)、水杨酸(SA)、细胞分裂素(CTK)和独脚金内酯(SL)。转录因子(包括ARF、MYB96、ABI3、ABI4和ABI5)通过与CYP707A1和CYP7072相互作用来调控ABA的生物合成, 而GA负调控(-)是通过DELLA基因实现的。这种平衡被持续保持直到种子出苗。引自Sohn等[8]。"

图2

拟南芥种子休眠的DOG1基因(启动子为红色, 编码区为绿色)的转录调控模型 GBL1顺式元件的下游为INDEL (插入缺失标记), 一个285 bp的序列。INDEL存在于Ler-0材料(弱休眠)的DOG1启动子中。然而, INDEL在Cvi-0生态型(强休眠)中是缺陷的。INDEL在体内直接影响bZIP67反式激活DOG1的能力, 以及可以解释在生态型之间观察到的DOG1转录本水平和休眠的变化。这种与DOG1表达的可塑性相偶联的DOG1自然等位基因的变异在田间具有重要的适应性意义, 因为季节性环境因素可能变化很大, 而种子萌发的最适时间是原始的。ETR1: 乙烯反应-1; DRE/CRT: 脱水反应元件/C-重复序列; TPL1: Topless-1; ERF12: 乙烯反应转录因子-12; GBL1和GBL2: G-box-like-1和2; SOM: Somnus。引自Carrillo-Barral等[17]。"

图3

描述亲本等位基因的不同表观遗传修饰组合如何影响胚乳基因表达和控制种子休眠的模型 其母系等位基因被FIS-PRC2介导的H3K27me3抑制的基因编码诱导萌发必需的乙烯途径基因。这些基因的父系等位基因可能被非典型的RdDM途径建立的CHHm所抑制。冷胁迫诱导的AGO6也可能靶定母系等位基因并通过非典型的RdDM途径涉及冷胁迫反应的CHHm。REF6与CTCTGYTY基序(Y = C或者T)结合以激活靶基因。在休眠种子中, 母系等位基因被H3K27me3去甲基化酶REF6激活, 以及这些基因表现出母系偏向的表达模式。在非休眠种子中, 这些基因是双等位基因表达的, 可能是因为RdDM途径在休眠诱导条件下特别活跃。FIS-PRC2、SUVH家族蛋白和CMT3分别在PEG的母系等位基因上建立H3K27me3、H3K9me2和CHGm。因为REF6结合位点上的CHGm抑制REF6的结合活性, 因此母系等位基因被持续沉默, 这些基因在整个胚乳发育过程中(包括萌发)表现出父系偏向的基因表达模式。ABI3是一个建立种子休眠的重要转录因子, 属于这种类型。母系等位基因上具有单一H3K9me2的基因在整个发育过程中被完全沉默。这些母系专一的表观遗传修饰在授粉后4 d的发育胚乳中被检测到。引自Sato和K?hler[2]。"

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