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作物学报 ›› 2019, Vol. 45 ›› Issue (7): 969-981.doi: 10.3724/SP.J.1006.2019.84175

• 综述 •    下一篇

乙烯的生物合成与信号及其对种子萌发和休眠的调控

宋松泉1,3,*(),刘军2,徐恒恒2,张琪2,黄荟3,伍贤进3   

  1. 1 中国科学院植物研究所, 北京100093
    2 广东省农业科学院农业生物基因研究中心, 广东广州 510640
    3 怀化学院民族药用植物资源研究与利用湖南省重点实验室 / 生物与食品工程学院, 湖南怀化 418008
  • 收稿日期:2018-11-22 接受日期:2019-01-19 出版日期:2019-07-12 网络出版日期:2019-04-09
  • 通讯作者: 宋松泉
  • 基金资助:
    本研究由国家科技支撑计划项目(2012BAC01B05);国家自然科学基金项目(31371715);国家自然科学基金项目(31640059);广东省科技计划项目(2016B030303007)

Biosynthesis and signaling of ethylene and their regulation on seed germination and dormancy

SONG Song-Quan1,3,*(),LIU Jun2,XU Heng-Heng2,ZHANG Qi2,HUANG Hui3,WU Xian-Jin3   

  1. 1 Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China
    2 Agro-Biological Gene Research Center, Guangdong Academy of Agricultural Sciences, Guangzhou 510640, Guangdong, China
    3 Key Laboratory of Research and Utilization of Ethnomedicinal Plant Resources of Hunan Province / College of Biological and Food Engineering, Huaihua University, Huaihua 418008, Hunan, China
  • Received:2018-11-22 Accepted:2019-01-19 Published:2019-07-12 Published online:2019-04-09
  • Contact: Song-Quan SONG
  • Supported by:
    This study was supported by the National Science and Technology Support Program(2012BAC01B05);the National Natural Science Foundation of China(31371715);the National Natural Science Foundation of China(31640059);the Guangdong Science and Technology Program(2016B030303007)

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摘要：

种子萌发是一种关键的生态和农业性状, 由调控种子休眠状态和萌发潜势的内在和外部信息所决定, 在植物随后的生长发育和产量中起着极其重要的作用。休眠是指种子在合适的条件下暂时不能萌发。乙烯是一种简单的具有多种功能的气体植物激素, 在分子、细胞和整体植物水平调节植物的代谢。在适宜和逆境条件下, 乙烯通过与其他信号分子的相互作用影响植物的行为。本文主要综述乙烯的生物合成与信号、乙烯在种子萌发和休眠释放中的作用以及乙烯与植物激素脱落酸和赤霉素的相互作用; 并提出了需要进一步研究的科学问题, 试图为解释乙烯调控种子萌发与休眠的分子机制提供新的研究思想。

关键词: 脱落酸, 生物合成与信号, 交叉反应, 乙烯, 赤霉素, 种子萌发和休眠

Abstract:

Seed germination, a key ecological and agronomic trait, is determined by both internal and external cues that regulate the dormancy status and the potential for germination in seeds, and plays a critical role during the subsequent growth, development and production of plants. Dormancy is the temporary failure of seed germination under favorable conditions. Ethylene is a simple gaseous phytohormone with multiple roles in regulation of metabolism at molecular, cellular, and whole plant levels. It influences performance of plants under optimal and stressful environments by interacting with other signaling molecules. In the present paper, we mainly summarize ethylene biosynthesis and signaling, the role of ethylene in seed germination and dormancy release, and the interaction of ethylene with phytohormone abscisic acid and gibberellin, and propose some scientific problems to be required to investigate further in order to provide an idea for explaining the molecular mechanism of seed germination and dormancy regulated by ethylene.

Key words: abscisic acid, biosynthesis and signaling, crosstalk, ethylene, gibberellin, seed germination and dormancy

图1

乙烯生物合成途径 S-腺苷甲硫氨酸(S-AdoMet)合成酶催化从甲硫氨酸形成S-AdoMet, 合成1分子的S-AdoMet消耗1分子的ATP(1)。ACC合酶催化S-AdoMet转化成为ACC是乙烯合成的限速步骤(2)。随着ACC的合成, 甲硫腺苷(MTA)是ACC合酶产生的副产物。MTA回到甲硫氨酸的再循环保存了甲硫基, 能够维持细胞中恒定的甲硫氨酸浓度。ACC丙二酰化作用成为丙二酰-ACC 使ACC库枯竭并减少乙烯的产生。ACC氧化酶利用ACC作为底物, 催化乙烯合成的最后步骤, 同时产生二氧化碳和氰化物(3)。氰化物被β-氰丙氨酸合酶代谢产生无毒的物质。ACC合酶和ACC氧化酶被同源异构蛋白、发育和环境信息的转录调节用虚线箭头表示。引自Lin等[19]。"

图2

拟南芥中乙烯信号途径的最近模型 乙烯由受体蛋白ETR1、ERS1、ETR2、ERS2和EIN4 (绿色表示)感受, 受体是乙烯信号的负调控因子。受体通过它们的GAF结构域(在受体的细胞质区域用五边形表示)与其他的受体相互作用, 并在ER膜中形成更高层次的复合物。铜(一种乙烯结合的辅因子, 红色圆圈)由铜转运体RAN1 (橙色表示)传递给受体。RTE1 (粉红色)与ETR1相联系, 介导受体信号输出。(A)在乙烯缺乏时, 受体激活CTR1 (黄色)。CTR1通过直接磷酸化EIN2的C-末端(蓝色圆圈)使其失活(紫色)。EIN2能够直接与受体的激酶结构域(在受体的细胞质区域在五边形下较大的椭圆)相互作用。EIN2的水平通过26S蛋白酶体(灰色)被F-box蛋白ETP1和ETP2(绿色星状物)负调控。在细胞核中, 转录因子EIN3/EIL1 (红色)通过蛋白酶体被另外2个F-box蛋白EBF1/2(蓝色星状物)降解。在EIN3/EIL1缺乏时, 乙烯反应基因的转录被关闭。(B)在乙烯存在时, 受体与激素结合并失去活性, 依次关闭CTR1。这种失活阻止正调控因子EIN2的磷酸化。EIN2的C-末端被一种未知的机制所剪切, 并移动到细胞核, 在细胞核中使EIN3/EIL1稳定和诱导EBF1/2的降解。转录因子EIN3/EIL1形成二聚体, 激活乙烯靶基因的表达, 包括F-box基因EBF2 (深蓝色卷曲线, 它产生抑制乙烯途径活性的负反馈环)或者转录因子基因ERF1(淡蓝色线, 它依次始启一个转录级联, 导致数百个乙烯调控基因的活化和抑制)。在乙烯反应基因中有受体基因ETR2 (绿色线), 它的mRNA被乙烯上调, 以及被翻译成为一批新的没有与乙烯结合的受体分子; 这些受体分子然后活化负调控因子CTR1, 从而提供了在不添加乙烯的情况下向下调节乙烯信号的手段。途径中的其他调控节点是核糖核酸外切酶EIN5 (淡橙色, 它控制EBF2 的mRNA水平)以及F-box蛋白ETP1和ETP2 (绿色星状物, 在乙烯存在时, 它们被降解, 导致EIN2的稳定)。正箭头和负箭头分别表示激活和下调这个过程。颜色变浅表示的分子(在‘没有乙烯’中的EIN3/EIL1, 或者在‘乙烯’中的ETP1/2和EBF1/2)相应于蛋白酶体介导降解的被标记的不稳定蛋白。卷曲线表示特定的mRNA, 它们的颜色与相应的蛋白质颜色相一致。引自Merchante等[51]。"

表1

乙烯、乙烯利或者1-氨基环丙烷-1-羧酸打破种子休眠的物种(引自Corbuneau et al.[16])"

初生休眠 Primary dormancy 次生休眠 Secondary dormancy 热休眠 Thermo dormancy
尾穗苋 Amaranthus caudatus 尾穗苋 Amaranthus caudatus 莴苣 Lactuca sativa
反枝苋 Amaranthus retroflexus 繁穗苋 Amaranthus paniculatus
拟南芥 Arabidopsis thaliana 向日葵 Helianthus annuus
花生 Arachis hypogaea 莴苣 Lactuca sativa
藜 Chenopodium album 皱叶酸模 Rumex crispus
欧洲水青冈 Fagus sylvatica 苣头苍耳 Xanthium pennsylvanicum
向日葵 Helianthus annuus
苹果 Malus pumila
南欧盐肤木 Rhus coriaria
皱叶酸模 Rumex crispus
柱花草 Stylosanthes humilis
地三叶 Trifolium subterraneum
苣头苍耳 Xanthium pennsylvanicum

图3

乙烯、脱落酸和赤霉素在种子萌发和休眠调控中的相互作用 该方案是基于正文中引证的种子对乙烯、脱落酸或者GA响应的遗传分析、芯片数据和生理研究。乙烯通过抑制ABA的合成和促进它的失活或者分解代谢下调ABA的积累, 也负调控ABA信号。ABA通过ACS和ACO的活性抑制乙烯的生物合成。乙烯也增强GA的代谢和信号, 反过来也一样。“→”和“┤”分别表示信号级联的不同元素之间的正、负相互作用。根据Corbineau等[16]重绘。"

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