中国农业科学 ›› 2015, Vol. 48 ›› Issue (4): 646-660.doi: 10.3864/j.issn.0578-1752.2015.04.03

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

种子活力与萌发的生理与分子机制研究进展

李振华1,2,王建华1   

  1. 1中国农业大学农学与生物技术学院,北京100193
    2贵州省烟草科学研究院/烟草行业分子遗传研究重点实验室,贵阳 550081
  • 收稿日期:2014-08-27 出版日期:2015-02-16 发布日期:2015-02-16
  • 通讯作者: 王建华,Tel:010-62732263;E-mail:Wangjh63@cau.edu.cn
  • 作者简介:李振华,Tel/Fax:0851-4117096;E-mail:lixing_19841014@126.com
  • 基金资助:
    国家公益性行业(农业)科研专项(201303002)、贵州省地方标准制修订项目(201243)

Advances in Research of Physiological and Molecular Mechanism in Seed Vigor and Germination

LI Zhen-hua1,2, WANG Jian-hua1   

  1. 1College of Agronomy and Biotechnology, China Agricultural University, Beijing 100193
    2Guizhou Academy of Tobacco Science Molecular Genetics Key Laboratory of China Tobacco, Guiyang 550081
  • Received:2014-08-27 Online:2015-02-16 Published:2015-02-16

摘要: 种子活力形成于种子发育的脱水阶段,与以往理解不同,近来研究表明种子脱水不仅仅是一个简单的失水过程,脱水与萌发存在一些相同的基因表达和代谢特征。萌发始于吸水膨胀,种子代谢恢复,胚根突破胚乳和种皮等外围包被组织完成萌发。种子萌发的质量关键在于储藏的mRNA的质量,另外,蛋白质稳定性和DNA完整性影响萌发的表型。激素作为一种信号小分子物质,浓度极小甚至是趋于零时对种子萌发仍具有非常重要的作用,近年来研究表明ABA/GAs的阈值范围调控种子休眠与萌发,其中,ABA几乎作用于种子萌发的全部过程,GA的作用并不像ABA那样广泛,主要在胚根突出时发生作用,且ABA和GA彼此抑制对方的合成与分解代谢基因,它们均可调控α-淀粉酶基因的转录。除ABA和GA外,近来发现AUX参与调控种子的休眠与萌发,其对胚根突出的调控比对子叶开展更加精细,AUX信号与ABA信号存在交叉,AUX通过其响应蛋白ARF10/16间接调控ABA信号通路中ABI3的稳定性来调控拟南芥种子休眠与萌发。与光照条件下相比,在土壤中萌发的幼苗将形成一个特异性的组织“顶钩”,它的主要作用是在幼苗“顶土”时保护顶端分生组织,“顶钩”形成与生长素的不均匀分布有关。为了提高种子活力,引发技术被运用于生产,引发的关键在于控制种子“萌而未发”,在“回干窗口”内及时脱水,引发过程中种子储藏的mRNAs和蛋白已经执行功能,回干后这种分子机制被“牢记”,再吸胀的种子可以迅速整齐的萌发。除毒害分子外,ROS还作为信号分子参与调控种子休眠释放、胚乳松弛和贮藏物动员,且其与激素分子ABA和GA等存在交互作用,ROS还参与蛋白的翻译和翻译后修饰调控种子萌发吸水和贮藏物动员。在种子萌发过程中,甲硫氨酸代谢是代谢核心,其代谢产物广泛参与调控种子萌发的一些生理生化反应,如:DNA的合成,蛋白质的稳定性,染色体结构的形成和重塑,生物素的合成等,另外还与激素分子ABA、GA、ETH和CTK及活性氧活性(氮)存在交互作用。近来还发现甲硫氨酸亚砜还原酶决定种子的寿命,它可能是种子活力的一个新的分子标记。本文将围绕上述内容对国内外研究现状进行综述,并对今后的研究热点,种子“提早”收获的感官依据,高活力种子田间出苗差异的分子机制,生长素在胚根突出时的重要作用,甲硫氨酸代谢,种子活力检测方法的选择等内容进行了展望。

关键词: 种子活力, 萌发, 激素, 活性氧, 甲硫氨酸代谢, 组学

Abstract: Seed acquired vigor during desiccation stage in seed development, different from the previous opinions, new evidence insisted that seed desiccation is not only involved in water loss process, but also as a significant proportion of the gene expression and metabolic signatures of which resemble those characterized seed germination, implied that the preparation of the seeds for germination began already during seed desiccation. The germination of seeds initiated from water-uptake, accompanied by metabolic recovery, then radicle breaked through endosperm and seed coat and other peripheral completed germination. The main contributor of seed germination success is the quality of the messenger RNAs stored during embryo maturation in the mother plant. In addition, proteostasis and DNA integrity play a major role in the germination phenotype. Plant hormone, as a signal, concentration extremely fewer even approximate to zero, was also important for seed dormancy release and germination. Recently, more and more ideas have considened that the ABA/GA ratio regulats the metabolic transition required for germination. GAs, although required for the completion of germination, are not directly involved in many processes taking place during germination like ABA, which occurred at a stage coinciding with or very close to radicle emergence. It appears that reciprocal downregulation of the respective metabolic pathways accounts for a significant part of GA and ABA interplay, and the α-amylase gene, is transcriptionally regulated by both ABA and GA. In addition to the ABA and GA, recently research found IAA finely regulated radicle emerge that more strictly than cotyledons open during seed germination. Auxin action in seed dormancy and germination requires the auxin response factors 10 and 16 to indirectly control the expression of ABI3. Compared with germination in light condition, seedlings germinated in soil will form a specific organization ‘Apical hook’, its main role was to protect the ‘SAM’. Auxin, accumulating at the concave side, was critical for the formation and maintenance of the hook structure, whereas a release of the auxin maximum correlates with hook opening. To improve the vigor, seed were primed before sowing, the key of which was to control seeds very close to radicle emergence but redried before it in time. In fact, seed stored mRNAs and proteins began to perform its function during priming, and also the molecular biological mechanism was "memorized" at the later redried state, so re-imbibed seeds could quickly germinate. Except as toxic molecules, ROS is also involved in the mobilization of storage and endosperm loose as signaling molecules during seed germination, and it is always interacted with hormone molecule ABA and GA, that also controls seed germination via translation and posttranslational modifications. Methionine metabolism is the metabolism core in seed germination and its metabolites widely regulate the physiological and biochemical reaction of seed germination, such as DNA synthesis, protein stability, chromosome structure formation and remodeling, biotin synthesis, and is also interacted with hormone molecule ABA, GA, ETH, and active oxygen or nitrogen. Recently new evidence insisits on the methionine sulfoxide reductase took part in repair system in plant seed longevity, which also might play a role in seed vigor. This article focused on the domestic and foreign research status on seed vigor, and also provided an outlook of the future research hotspots, such as the senses traits in seeds "preharvest", the molecular mechanism about higher vigor seeds but less field emergence, the important role of auxin in radicle breakout, methionine metabolism, and the methods about seed vigor testing, etc.

Key words: seed vigor, germination, hormone, reactive oxygen species, methionine metabolism, -omics