光信号和昼夜节律调控植物感知冷胁迫的研究进展

doi:10.13560/j.cnki.biotech.bull.1985.2024-0234

摘要/Abstract

摘要：

温度和光是调节植物生长发育的重要环境因子，植物感知温度变化并通过改变基因表达模式等响应低温环境，这些响应受到光信号和昼夜节律等因素的影响。然而，光信号诱导昼夜节律调控植物响应冷胁迫的分子调控网络尚不清楚。本文聚焦光信号和昼夜节律在植物感知冷胁迫中的作用。光信号参与冷胁迫主要通过光敏色素诱导CBF基因途径激活冷基因的表达，这主要有两种途径，一是光敏色素受体通过直接调控CBF和COR基因表达而调节植株抗冷性；二是光依赖性信号转导的正调控因子HY5激活冷驯化COR基因。昼夜节律参与冷胁迫主要是通过昼夜节律的组分CCA1/LHY和RVE4/RVE8介导DREB1下游基因在冷胁迫下的表达。明确植物中光信号和昼夜节律在冷信号感知及传导途径中的作用，不仅有助于更好地理解植物抗冷的机制和功能，还有助于植物生长与温度胁迫反应之间的权衡，为提升植物应对昼夜温差变化提供理论基础。

关键词: 冷感知, 抗冷基因, 光信号, 昼夜节律

Abstract:

Temperature and light are important environmental factors in regulating plant growth and development, and plants perceive temperature changes and respond to cold stress by altering gene expression patterns and other responses. The responses are affected by light signal and circadian rhythms. However, the molecular regulatory networks involved in plant response to cold stress are not well understood. This review focuses the roles of light signal and circadian rhythms in the perception of plants to cold stress. Light signal is involved in cold stress mainly through photopigment-induced CBF gene pathways to activate the expressions of cold genes. It includes two main pathways: one is that photosensitive pigments directly regulate cold tolerance by modulating CBF and COR gene expression; the other is the activation of COR（COLD REGULATED）genes by HY5, a positive regulator factor. Circadian rhythms are involved in cold stress mainly through their components, CCA1/LHY and RVE4/RVE8, regulating the expression of DREB1 downstream gene. Clarifying the roles of light signal and circadian rhythms in cold perception and conduction pathways not only contributes to a better understanding of the mechanisms of cold resistance in plants, but also helps in the trade-offs between plant growth and stress responses. It provides a theoretical basis for enhancing the response sensitivities of plants to diurnal and nocturnal temperature variations.

Key words: cold perception, cold resistance genes, light signal, circadian rhythm

李文萃, 彭羽佳, 刘勇波. 光信号和昼夜节律调控植物感知冷胁迫的研究进展[J]. 生物技术通报, 2024, 40(8): 1-12.

LI Wen-cui, PENG Yu-jia, LIU Yong-bo. Research Progress in Light Signal and Circadian Rhythm Regulating the Perception of Plants to Cold Stress[J]. Biotechnology Bulletin, 2024, 40(8): 1-12.

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基因/蛋白 Genes/Proteins	调控机制 Regulation mechanisms	物种 Species	参考文献 Reference
光敏色素A（phyA）Phytochrome A	远红光激活phyA诱导ABA信号传导，触发JA信号传导，激活CBF途径	拟南芥Arabidopsis thaliana 番茄S. lycopersicum	[46] [47]
光敏色素B（phyB）Phytochrome B	phyB与PIF4和PIF7相互作用，负调控CBF表达和冷冻耐受性	拟南芥A. thaliana 番茄S. lycopersicum 水稻Oryza sativa	[36] [35] [48]
光敏色素D（phyD）Phytochrome D	介导CBF调控的表达，phyD突变体耐冷性增加	拟南芥A. thaliana	[25]
向光素（PHOTs）Phototropins	蓝光传感器，在蓝光光激发时自磷酸化，激活C-末端激酶结构域，调控其他蛋白质	地钱 Marchantia polymorpha	[49]
F-box 蛋白 Zeitlupe（ZTL）Zeitlupe	ZTL属于LOV结构域光感受器，与热休克伴侣蛋白HSP 90相互作用从而正调控CBF表达	拟南芥A. thaliana	[27]
隐花色素1（CRY1）Cryptochrome 1	蓝光下CRY1对温度响应对下胚轴伸长具有强烈的抑制作用，同时也抑制PIF4的转录	拟南芥A. thaliana	[50]
隐花色素2（CRY1）Cryptochrome 2	CRY2-COP1相互作用减弱COP1与HY5的相互作用，从而增强HY5在冷胁迫下的光稳定性	拟南芥A. thaliana 水稻O. sativa	[51] [52]
UVR8感受器（UVR8） UV resistance locus 8	UVR8单体通过降低PIF4表达水平和抑制PIF4蛋白的转录活性来抑制下胚轴伸长	拟南芥A. thaliana	[53]
光敏色素相互作用因子3（PIF3） Phytochrome interacting factor 3	抑制CBF表达负调控抗冷性	拟南芥A. thaliana	[54]
光敏色素相互作用因子4（PIF4） Phytochrome interacting factor 4	长日照下，PIF4和PIF7与CBF基因启动子结合抑制其表达	拟南芥A. thaliana 番茄S. lycopersicum	[55] [56]
光敏色素相互作用因子7（PIF7） Phytochrome interacting factor 7	长日照下，PIF4和PIF7与CBF基因启动子结合抑制其表达	拟南芥A. thaliana	[57]
昼夜节律核心蛋白（CCA1） Circadian clock-associated 1	CCA1与CBF启动子结合，通过选择性剪接机制增强冷适应	拟南芥A. thaliana	[29] [58]
昼夜节律核心蛋白（LHY） Late elongated hypocotyl	正调控CBFs及其下游COR基因的表达和植物的抗冷性	拟南芥A. thaliana 草莓Fragaria×ananassa 茶树Camellia sinensis	[31] [59] [30]
伪应答调控蛋白5/7/9（PRR5/7/9） Pseudo response regulator 5/7/9	CBF途径的负调控因子	拟南芥A. thaliana	[60]
昼夜节律因子（TOC1） Timing of cab expression	与CBF2转录阻遏物的phyB一起发挥作用	拟南芥A. thaliana	[61]
伸长下胚轴5（HY5） Elongated hypocotyl 5	HY5通过Z-box/LTRE正向调控冷诱导基因表达	拟南芥A. thaliana 番茄S. lycopersicum 油菜 Brassica napus	[62] [63] [43]
核心时钟蛋白（GI）Gigantea	通过CBF非依赖性途径正调控抗冷性	拟南芥A. thaliana	[64]
生物钟核心基因（LUX）Lux arrhythmo	LUX被CBF1正调控，表达上调，正向调控植物抗冷性	拟南芥A. thaliana	[65]
生物钟组分RVE4/RVE8	RVE4/RVE8迅速从细胞质转移到细胞核中的冷应激反应，并作为DREB 1表达的直接转录激活因子	拟南芥A. thaliana	[66]
生物钟组分LNK1/2	LNK1/2是转录辅激活因子，与RVE4和RVE8相互作用	拟南芥A. thaliana	[67]

基因/蛋白 Genes/Proteins	调控机制 Regulation mechanisms	物种 Species	参考文献 Reference
光敏色素A（phyA）Phytochrome A	远红光激活phyA诱导ABA信号传导，触发JA信号传导，激活CBF途径	拟南芥Arabidopsis thaliana 番茄S. lycopersicum	[46] [47]
光敏色素B（phyB）Phytochrome B	phyB与PIF4和PIF7相互作用，负调控CBF表达和冷冻耐受性	拟南芥A. thaliana 番茄S. lycopersicum 水稻Oryza sativa	[36] [35] [48]
光敏色素D（phyD）Phytochrome D	介导CBF调控的表达，phyD突变体耐冷性增加	拟南芥A. thaliana	[25]
向光素（PHOTs）Phototropins	蓝光传感器，在蓝光光激发时自磷酸化，激活C-末端激酶结构域，调控其他蛋白质	地钱 Marchantia polymorpha	[49]
F-box 蛋白 Zeitlupe（ZTL）Zeitlupe	ZTL属于LOV结构域光感受器，与热休克伴侣蛋白HSP 90相互作用从而正调控CBF表达	拟南芥A. thaliana	[27]
隐花色素1（CRY1）Cryptochrome 1	蓝光下CRY1对温度响应对下胚轴伸长具有强烈的抑制作用，同时也抑制PIF4的转录	拟南芥A. thaliana	[50]
隐花色素2（CRY1）Cryptochrome 2	CRY2-COP1相互作用减弱COP1与HY5的相互作用，从而增强HY5在冷胁迫下的光稳定性	拟南芥A. thaliana 水稻O. sativa	[51] [52]
UVR8感受器（UVR8） UV resistance locus 8	UVR8单体通过降低PIF4表达水平和抑制PIF4蛋白的转录活性来抑制下胚轴伸长	拟南芥A. thaliana	[53]
光敏色素相互作用因子3（PIF3） Phytochrome interacting factor 3	抑制CBF表达负调控抗冷性	拟南芥A. thaliana	[54]
光敏色素相互作用因子4（PIF4） Phytochrome interacting factor 4	长日照下，PIF4和PIF7与CBF基因启动子结合抑制其表达	拟南芥A. thaliana 番茄S. lycopersicum	[55] [56]
光敏色素相互作用因子7（PIF7） Phytochrome interacting factor 7	长日照下，PIF4和PIF7与CBF基因启动子结合抑制其表达	拟南芥A. thaliana	[57]
昼夜节律核心蛋白（CCA1） Circadian clock-associated 1	CCA1与CBF启动子结合，通过选择性剪接机制增强冷适应	拟南芥A. thaliana	[29] [58]
昼夜节律核心蛋白（LHY） Late elongated hypocotyl	正调控CBFs及其下游COR基因的表达和植物的抗冷性	拟南芥A. thaliana 草莓Fragaria×ananassa 茶树Camellia sinensis	[31] [59] [30]
伪应答调控蛋白5/7/9（PRR5/7/9） Pseudo response regulator 5/7/9	CBF途径的负调控因子	拟南芥A. thaliana	[60]
昼夜节律因子（TOC1） Timing of cab expression	与CBF2转录阻遏物的phyB一起发挥作用	拟南芥A. thaliana	[61]
伸长下胚轴5（HY5） Elongated hypocotyl 5	HY5通过Z-box/LTRE正向调控冷诱导基因表达	拟南芥A. thaliana 番茄S. lycopersicum 油菜 Brassica napus	[62] [63] [43]
核心时钟蛋白（GI）Gigantea	通过CBF非依赖性途径正调控抗冷性	拟南芥A. thaliana	[64]
生物钟核心基因（LUX）Lux arrhythmo	LUX被CBF1正调控，表达上调，正向调控植物抗冷性	拟南芥A. thaliana	[65]
生物钟组分RVE4/RVE8	RVE4/RVE8迅速从细胞质转移到细胞核中的冷应激反应，并作为DREB 1表达的直接转录激活因子	拟南芥A. thaliana	[66]
生物钟组分LNK1/2	LNK1/2是转录辅激活因子，与RVE4和RVE8相互作用	拟南芥A. thaliana	[67]