植物开花时间的遗传调控通路研究进展

doi:10.17520/biods.2020370

摘要/Abstract

摘要：

开花时间对植物的繁殖成功至关重要。广泛分布的物种经常发生开花时间的分化, 从而能够更好地适应不同的环境条件。为了探索植物开花行为发生适应性分化的分子机制, 首先要明确调控开花行为的遗传通路。本文梳理了植物各类群调控开花时间的遗传通路, 以期为开花时间适应性分化的分子机制研究提供依据。植物从营养生长向繁殖转变时, 其开花行为主要受到光照、温度、水分等外界环境因子和赤霉素等内在因素的影响。通过对模式植物拟南芥(Arabidopsis thaliana)和其他类群的研究, 总结出了调控植物开花时间的6条通路, 包括日照长度和光质影响开花的光依赖通路, 长时间冷暴露后促进植物开花的春化通路, 高温或低温环境影响开花的温度通路, 以及赤霉素通路、年龄通路和自主通路3条内部调节过程。植物开花时间调控的6条上游通路信号传递到下游的开花整合基因FT(FLOWERING LOCUS T)和SOC1(SUPPRESSOR OF OVEREXPRESSION OF CONSTANS 1), 整合基因将这些复杂的调节因子整合后进一步传递到下游花分生组织, 从而启动开花。此外, 非编码RNA、转座子对开花时间的调控也具有重要作用。部分遗传通路被证实在植物适应环境的过程中起到了重要作用。目前对植物开花调控的研究已经有一百多年历史, 理论相对成熟。然而, 仍然存在许多具有争议和未解决的问题, 如开花基因的表达方式、开花行为的特殊调控机制、开花时间变异的适应性意义等等, 需要更进一步的研究。

关键词: 开花基因, 开花时间, 适合度, 遗传调控, 遗传通路

Abstract

Aims: Flowering time is critical for plant reproductive success. Plant species, especially which are widely distributed, are observed to exhibit significant differentiation in flowering time to maximize their fitness. In order to explore the molecular mechanism of adaptive differentiation of flowering time in plants, it is essential to clarify the regulating pathways of flowering. This paper summarizes the genetic pathways that regulate flowering time in various groups of plants, which can provide important reference for molecular mechanism research of adaptive differentiation of flowering time.

Progresses: As a vital transition from vegetative growth to reproduction in plants, the flowering time has been reported to be regulated by both extrinsic and intrinsic factors. Based on researches of the model plant Arabidopsis thaliana and other taxa, six pathways that regulate flowering time have been identified. The light-dependent pathway, vernalization pathway, and temperature pathway refer to the regulation of flowering time in response to day length or quality of light, a long period of cold exposure, and external high or low temperature, respectively. While the gibberellin pathway, the age pathway and the autonomic pathway are the endogenous pathways. Signals from the six upstream flowering regulation pathways are transmitted to the downstream flowering integratorsFT (FLOWERING LOCUS T) andSOC1(SUPPRESSOR OF OVEREXPRESSION OF CONSTANS 1), assembled by these two integrators, and transmitted to the downstream flowering meristem where flowering initiates. In addition, most recent studies have shown that non-coding RNA and transposable elements also play important roles in the regulation of flowering time.

Prospect: The researches on the regulation of plant flowering have studied more than a hundred years ago, forming many mature theories. However, there are still many controversial and unresolved problems, such as the expression ways of flowering genes, the special regulatory mechanism of flowering, and the adaptive significance of differentiated flowering time, which require further study.

Key words: flowering gene, flowering time, fitness, genetic regulation, genetic pathway

杨小凤, 李小蒙, 廖万金 (2021) 植物开花时间的遗传调控通路研究进展. 生物多样性, 29, 825-842. DOI: 10.17520/biods.2020370.

Xiaofeng Yang, Xiaomeng Li, Wanjin Liao (2021) Advances in the genetic regulating pathways of plant flowering time. Biodiversity Science, 29, 825-842. DOI: 10.17520/biods.2020370.

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链接本文: https://www.biodiversity-science.net/CN/10.17520/biods.2020370

https://www.biodiversity-science.net/CN/Y2021/V29/I6/825

图/表 2

图1 调控开花行为的遗传通路(以拟南芥为例)。图中自上而下分为三部分: 上游调控通路、下游开花整合基因和花分生组织基因。虚线方框内代表拟南芥调控开花的6个通路以及相关的基因。方框内的大写字母代表基因的简称, 椭圆框内的字母代表蛋白的简称。带箭头的实线代表促进作用, 平端的虚线代表抑制作用。改自Banta和Purugganan (2011)。

Fig. 1 The genetic pathways regulating flowering (take Arabidopsis thaliana as an example). The figure includes three parts from top to bottom: upstream regulatory pathway, downstream floral integrators and floral meristem identity genes. Six pathways and related genes that regulate flowering in Arabidopsis thaliana are enclosed in the dashed boxes separately. The letters in the boxes stand for abbreviation of genes, and the letters in the ovals stand for abbreviation of proteins. Lines with arrows indicate activation, and broken lines with blunt ends indicate repression. Modified from Banta & Purugganan ( 2011).

图2 拟南芥开花调控各通路的详细图解。(A)光周期通路; (B)春化通路; (C)温度通路; (D)赤霉素通路; (E)年龄通路。黑色填充方框代表外界调节因子。方框内的字母代表基因的简称, 椭圆框内的字母代表蛋白的简称。带箭头实线代表促进作用, 平端实线代表抑制作用。双向箭头代表相互作用。在图(A)中, 圆角矩形框指开花素转运到的位置, 带箭头的虚线代表转运过程。

Fig. 2 Detailed diagrams of the regulation pathways of flowering in Arabidopsis thaliana. (A) The photoperiodic pathway; (B) The vernalization pathway; (C) The temperature pathway; (D) The gibberellin pathway; (E) The age pathway. The black-filled box represents external regulating factors. The letters in the boxes stand for abbreviation of genes, and the letters in the ovals stand for abbreviation of proteins. Lines with arrows and blunt ends indicate activation and repression, respectively. The two-way arrows represent interactions. In figure (A), rectangle with rounded corners indicates the position where the florigen to be transferred, and the broken lines with arrows represent the transport process.

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