生物钟与光温环境信号互作网络研究进展

doi:10.11983/CBB24174

植物学报 ›› 2025, Vol. 60 ›› Issue (3): 315-341.DOI: 10.11983/CBB24174 cstr: 32102.14.CBB24174

• 特邀综述 • 下一篇

生物钟与光温环境信号互作网络研究进展

苏晨¹^,²^,^†, 牛钰凡¹^,²^,^†, 徐航¹^,², 王希岭¹, 于英俊¹, 何雨晴¹^,^*(), 王雷¹^,²^,³^,^*()

¹中国科学院植物研究所, 饲草种质高效设计与利用全国重点实验室, 北京 100093
²中国科学院大学, 北京 100049
³ 国家植物园, 北京 100093

收稿日期:2024-11-18 接受日期:2024-12-26 出版日期:2025-05-10 发布日期:2024-12-27
通讯作者: *E-mail: heyuqing@ibcas.ac.cn; wanglei@ibcas.ac.cn
作者简介:
†共同第一作者
基金资助:
国家自然科学基金青年基金(32300294);国家自然科学基金(32370307);中国博士后第73批面上项目(2023M733729)

Advances of Plant Circadian Clock Response to Light and Temperature Signals

Su Chen¹^,²^,^†, Niu Yufan¹^,²^,^†, Xu Hang¹^,², Wang Xiling¹, Yu Yingjun¹, He Yuqing¹^,^*(), Wang Lei¹^,²^,³^,^*()

¹State Key Laboratory of Forage Breeding-by-Design and Utilization, Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China
²University of Chinese Academy of Sciences, Beijing 100049, China
³ China National Botanical Garden, Beijing 100093, China

Received:2024-11-18 Accepted:2024-12-26 Online:2025-05-10 Published:2024-12-27
Contact: *E-mail: heyuqing@ibcas.ac.cn; wanglei@ibcas.ac.cn
About author:
†These authors contributed equally to this work

摘要/Abstract

摘要： 随着全球气候的急剧变化, 植物生长发育所处的生态环境日益恶劣, 生物钟与光、温受体互作协同传递环境信号并调控下游生长发育的应答机制开始受到科研人员的广泛关注。生物钟作为植物内源计时器, 其核心由多个耦联的转录-翻译反馈环(TTFL)组成, 在转录、转录后、翻译、翻译后和表观遗传层面受到多层级精细调控。这些精密的调控机制保证了生物钟能不断被外界环境信号驯化和重置, 使内源节律与外界环境相匹配, 从而赋予植物优化资源利用和趋向最适生长的能力, 对于指导农作物遗传改良和引种驯化具有重要意义。该综述总结了生物钟核心振荡器的多层级调控机制以及生物钟同源基因在农作物中的分子功能, 详述了生物钟与光、温环境信号通路间的互作网络, 展望了以此为基础的作物分子育种, 为提高农作物的环境适应性和优化改良农艺性状提供了新思路。

关键词: 生物钟, 光温信号, TTFL, 环境适应性, 分子育种

Abstract: With the sharp change of the global climate, the ecological environment for plant is becoming increasingly harsh, therefore the molecular mechanisms underlying how circadian synergistically interacts with light or temperature receptors to transmit environmental signals and rhythmically regulate various growth and development process received widespread attention. As an endogenous timer of plants, the core oscillator of circadian clock is composed of multiple coupled transcriptional-translational feedback loops (TTFL), and it is modified from transcription, post-transcription, translation, post-translation to epigenetic levels. These multi-precise regulatory mechanisms ensure that the circadian clock can be synchronized and reset by external signals, so that the endogenous rhythm matches with external cycles, thereby endowing plants with the ability to optimize resource utilization and tend towards the optimal growth, which also has an important significance for guiding the genetic improvement and domestication of crops. In this review, we summarized the multi-level of regulatory mechanisms of core oscillator as well as the molecular function of circadian homologous genes in crops, thoroughly described the interaction network between the circadian clock and the light and temperature signal pathways and give prospects for molecular breeding based on the opinion, which provides new ideas for expanding the environmental adaptability and optimizing agronomic traits of crops.

Key words: circadian clock, light and temperature signals, TTFL, environmental adaptability, molecular breeding

苏晨, 牛钰凡, 徐航, 王希岭, 于英俊, 何雨晴, 王雷. 生物钟与光温环境信号互作网络研究进展. 植物学报, 2025, 60(3): 315-341.

Su Chen, Niu Yufan, Xu Hang, Wang Xiling, Yu Yingjun, He Yuqing, Wang Lei. Advances of Plant Circadian Clock Response to Light and Temperature Signals. Chinese Bulletin of Botany, 2025, 60(3): 315-341.

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图/表 8

图1 拟南芥生物钟核心振荡器转录-翻译反馈环模式图箭头表示直接或间接激活作用, 钝线表示直接或间接抑制作用。

Figure 1 Transcription-translation feedback loop of Arabidopsis circadian core oscillator Lines with arrows mean direct or indirect activation while blunt lines mean direct or indirect repression.

图2 生物钟核心振荡器组分的转录后修饰 (A) 剪接体组分介导拟南芥生物钟基因的可变剪接; (B) 温度调控CCA1的可变剪接; (C) m6A甲基转移酶FIO1和MTA复合体促进CCA1 mRNA的m6A修饰。

Figure 2 Post-transcriptional modification of the circadian core oscillator components (A) Components of spliceosome mediate alternative splicing of circadian genes in Arabidopsis; (B) Temperature regulates the splicing isoforms of CCA1; (C) m6A methyltransferases FIO1 and the MTA complex promotes m6A installation on CCA1 mRNA.

图3 生物钟核心振荡器组分的染色质修饰水平调控模式图 (A) CCA1/LHY或TOC1与LDL1/2-HDA6复合物结合抑制彼此的乙酰化(AC)和三甲基化(Me3)修饰; (B) PRR5/7/9招募TPL/TPR和HDA6去除靶基因启动子区域的乙酰化; (C) HAF2结合靶基因的启动子区域, 促进H3AC沉积, 激活基因的表达; (D) 组蛋白去甲基化酶LSD1、JmjC、JMJD5/JMD30及甲基转移酶HMT、SDG2/ATXR3直接或间接调控生物钟基因启动子区域的甲基化水平; (E) CCA1介导MLK3和MLK4调控GI启动子区域的H2AS95磷酸化(P)和H4乙酰化; (F) HUB1参与调控靶基因启动子区域的H2B单泛素化(Ub)和H3甲基化修饰。

Figure 3 A diagram showing chromatin modifications regulation of the circadian core oscillators (A) CCA1/LHY or TOC1 interacts with LDL1/2-HDA6 complex to inhibit the acetylation (AC) and trimethylation (Me3) modifications of each other; (B) PRR5/7/9 recruits TPL/TPR and HDA6 to remove the acetylation modification of target genes; (C) The histone acetyltransferase HAF2 binds to the promoter region of the target genes to promote the deposition of H3AC and activate gene expression; (D) The histone demethylase LSD1, JmjC, JMJD5/JMD30, and the methyltransferase HMT, SDG2/ATXR3 directly or indirectly regulate the methylation level of the promoter region of clock genes; (E) CCA1 mediates the regulation of MLK3 and MLK4 on the phosphorylation (P) of H2AS95 and the acetylation of H4 in the GI promoter region; (F) HUB1 is involved in regulating the monoubiquitination (Ub) of H2B and the methylation modification of H3 in the promoter regions of target genes.

图4 拟南芥生物钟调控光周期依赖的开花模式图 (A) 生物钟组分通过调控CO-FT模块介导开花的简易模型(黑线代表转录调控, 红线代表蛋白调控); (B) 不同光周期下生物钟组分差异调控CO-FT表达的时相示意图。?: 未知

Figure 4 The diagrams showing photoperiodic flowering regulated by the circadian clock in Arabidopsis (A) A simplified model of flowering mediated by the clock components through regulation of CO-FT modules (the black lines represent transcriptional regulation, the red Lines represent protein regulation); (B) Schematic diagrams of how the clock differentially regulate the expression of CO-FT under different photoperiods. ?: Unknown

图5 拟南芥生物钟调控光周期依赖的下胚轴生长模式图 (A) 不同光周期下生物钟差异调控PIFs转录水平的模式图(箭头代表转录激活, 钝线代表转录抑制); (B) 昼夜环境下, 生物钟调控PIFs蛋白水平和转录活性示意图。“x”代表干扰转录激活活性, “p”代表磷酸化, 多个同色小圆代表该蛋白降解。?: 未知

Figure 5 The diagrams showing photoperiodic hypocotyl growth regulated by the circadian clock in Arabidopsis (A) Schematic models of the clock differentially regulate the expression of PIFs transcriptionally under different photoperiods (the arrows represent transcriptional activation, and the lines represent transcriptional inhibition); (B) A drawing of the clock regulation of PIFs protein levels and transcriptional activity in diurnal condition. “x” represent interference of transcriptional activation ability, “p” represent phosphorylation, and multiple small circles of the same color represent the degradation of the protein. ?: Unknown

表1 水稻生物钟基因的功能总结

Table 1 Functional summary of circadian genes in rice

基因名称	拟南芥同源基因	参与的生理过程	参考文献
OsCCA1	CCA1	开花时间调控和盐胁迫响应	Lee et al., 2022; Li et al., 2022; Wei et al., 2022
OsPRR1	PRR1/TOC1	开花时间调控	Li et al., 2023a
OsPRR37	PRR3/PRR7	开花时间调控	Kwon et al., 2015; Hu et al., 2021
OsPRR73	PRR3/PRR7	盐胁迫响应和开花时间调控	Liang et al., 2021; Wei et al., 2021
OsPRR59	PRR5/PRR9	光合碳固定、开花时间调控、脱落酸和盐胁迫响应	Chen et al., 2022; Wang et al., 2022b
OsPRR95	PRR5/PRR9	光合碳固定、开花时间调控、脱落酸和盐胁迫响应	Chen et al., 2022; Wang et al., 2022b
OsGI	GI	盐胁迫响应和开花时间调控	Izawa et al., 2011; Wang et al., 2021b
OsELF3-1	ELF3	盐胁迫响应和开花时间调控	Zhao et al., 2012; Wang et al., 2021b
OsELF3-2	ELF3	稻瘟病免疫响应、盐胁迫响应和开花时间调控	Zhao et al., 2012; Ning et al., 2015
OsELF4a	ELF4	盐胁迫响应、开花时间调控和免疫调控	Wang et al., 2021b; Xu et al., 2023
OsELF4b
OsELF4c
OsLUX	LUX	盐胁迫响应和开花时间调控	Wang et al., 2021b

表2 大豆生物钟基因的功能总结

Table 2 Functional summary of circadian genes in soybean

基因名称	拟南芥同源基因	参与的生理过程	参考文献
E2	GI	开花时间调控和产量形成	Dong et al., 2022; Wang et al., 2023
E2la
E2lb
GmLHY1a	LHY	干旱胁迫响应和开花时间调控	Marcolino-Gomes et al., 2014; Li et al., 2019; Wang et al., 2020b; Dong et al., 2021; Yuan et al., 2021b
GmLHY1b		干旱胁迫响应和开花时间调控
GmLHY2a		开花时间调控
GmLHY2b		开花时间调控
GmLUX1	LUX	开花时间调控	Bu et al., 2021
GmLUX2	LUX	开花时间调控	Bu et al., 2021
GmELF3/J	ELF3	开花时间调控和盐胁迫响应	Lu et al., 2017; Cheng et al., 2020
GmPRR3a	PRR3	干旱胁迫响应和开花时间调控	Syed et al., 2015; Lu et al., 2020
GmPRR3b	PRR3	干旱胁迫响应和开花时间调控	Syed et al., 2015; Lu et al., 2020

表3 玉米生物钟基因的功能总结

Table 3 Functional summary of circadian genes in maize

基因名称	拟南芥同源基因	参与的生理过程	参考文献
ZmGI1	GI	开花时间调控和株高控制	Bendix et al., 2013; Li et al., 2023b
ZmGI2	GI	开花时间调控	Bendix et al., 2013; Li et al., 2023b
ZmPRR1-2	PRR1/TOC1	开花时间调控	董柯清等, 2023
ZmELF3.1	ELF3	开花时间调控	Zhao et al., 2023
ZmELF3.2	ELF3	开花时间调控	Zhao et al., 2023
ZmELF4.1	ELF4	开花时间调控	Zhao et al., 2023
ZmELF4.2	ELF4	开花时间调控	Zhao et al., 2023
ZmLUX1	LUX	开花时间调控	Zhao et al., 2023
ZmLUX2	LUX	开花时间调控	Zhao et al., 2023
ZmPRR37a	PRR3/PRR7	开花时间调控	Zhao et al., 2023
ZmPRR73	PRR3/PRR7	开花时间调控	Zhao et al., 2023
ZmCCA1	CCA1	干旱胁迫响应	Tian et al., 2019

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生物钟与光温环境信号互作网络研究进展

Advances of Plant Circadian Clock Response to Light and Temperature Signals

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