蛋白质泛素化介导的植物低温胁迫反应

doi:10.13560/j.cnki.biotech.bull.1985.2020-1389

摘要/Abstract

摘要：

低温胁迫限制植物的生长发育及其地理分布。不同物种在低温胁迫反应过程中发生了重大的转录组重新编程,许多蛋白质被认为是这种适应性反应的重要因子。泛素化是一种翻译后修饰,能够调控蛋白的丰度、活性、亚细胞区隔和转运,并参与低温应答过程。E3是泛素-蛋白酶体系统的主要组成部分,识别目标蛋白,并将泛素从E2上转移到目标蛋白上。由于E3具有底物识别的特异性,从而调控植物多种低温胁迫反应相关的信号通路。以此为基础,总结了参与泛素-蛋白酶体系统的组成成分,并详细阐述了E3在低温胁迫反应中的作用。

关键词: 低温胁迫, 泛素化, 泛素连接酶, 信号转导, 抗冻性

Abstract:

Low-temperature stress restricts plant growth,development,and geographical distribution. A large transcriptome reprogramming occurs as a response to low-temperature stress in diverse species,and a number of proteins have been identified as important factors in this adaptive response. Ubiquitination is a post-translational modification that regulates abundance,activities,subcellular compartmentalization and transport,and involved in the low-temperature stress response. The E3 is a major component of the ubiquitin-proteasome system that recognizes the target protein and transfer of ubiquitin from the E2 to the target protein. Owing to its substrate recognition specificity,the E3 regulates various signaling pathways involved in the low-temperature stress response in plants. Therefore,components involved in the ubiquitin-proteasome system are summarized in the present study,and the role of E3 in the low-temperature stress response is described in detail.

Key words: low temperature stress, ubiquitination, E3 ubiquitin ligases, signal transduction, freezing tolerance

乌凤章, 王贺新. 蛋白质泛素化介导的植物低温胁迫反应[J]. 生物技术通报, 2021, 37(6): 225-235.

WU Feng-zhang, WANG He-xin. Low Temperature Stress Response Mediated by Protein Ubiquitination in Plant[J]. Biotechnology Bulletin, 2021, 37(6): 225-235.

图/表 3

表1 参与植物低温胁迫反应的E3连接酶的一般性功能描述

Table 1 General functional descriptions of the E3 ligases involved low temperature stress responses in plant

类型 Type	泛素连接酶 Ub-ligase	靶目标 Target(s)	物种 Species	一般性功能描述 General function description	参考文献 References
RING	HOS1	ICE1	拟南芥 Arabidopsis thaliana	负调控抗冻性 Negatively regulate freezing resistance	[22]
	MaSINA1	MaICE1	香蕉 Musa acuminata	负调控耐冷性 Negatively regulate chilling tolerance	[32]
	MdMIEL1	MdMYB308L	苹果 Malus domestica	负调控抗冻性 Negatively regulate freezing resistance	[31]
	SAP9	Rad23d	拟南芥 Arabidopsis thaliana	正调控抗寒性 Positively regulates cold resistance	[61]
U-box	CaPUB1	RPN6	水稻 Oryza sativa	正调控耐冷性 Positively regulates chilling tolerance	[56]
	PUB25/26	MYB15	拟南芥 Arabidopsis thaliana	正调控抗冻性 Positively regulates freezing resistance	[29]
	AtCHIP	PP2A	拟南芥 Arabidopsis thaliana	负调控耐冷性 Negatively regulate chilling tolerance	[60]
SCF	EBF1/2	PIF3	拟南芥 Arabidopsis thaliana	正调控抗冻性 Positively regulates freezing resistance	[30]
	EBF1/2	EIN3/EIL1	拟南芥 Arabidopsis thaliana	正调控抗冻性 Positively regulates freezing resistance	[49-50]
	FBP7	LOS1?	拟南芥 Arabidopsis thaliana	负调控耐冷性 Negatively regulate chilling tolerance	[55]
	COI1	JAZs	拟南芥 Arabidopsis thaliana	正调控抗冻性 Positively regulates freezing resistance	[51,53]
DDB	COP1	HY5	拟南芥 Arabidopsis thaliana	负调控抗冻性 Negatively regulate freezing resistance	[34]
BTB	ETO1/EOL1/2	ACS5	拟南芥 Arabidopsis thaliana	负调控抗冻性 Negatively regulate freezing resistance	[45]

图1 蛋白质泛素化修饰调节CBF依赖的低温信号途径 CBFs转录因子负责调控其启动子中含有CRT/DRE(CCGAC)基序的COR基因。CBFs被ICE1和CAMTA转录因子激活,而被MYB15抑制。HOS1和SIZ1分别编码RING E3和SUMO E3,负调控ICE1蛋白的丰度。OST1是一种正调控ICE1蛋白丰度的激酶。弯箭头表示翻译后调节,实箭头表示促进作用,T形线表示抑制作用。下同

Fig. 1 Protein ubiquitination regulates the CBF-dependent low temperature signaling pathway CBFs transcription factors are responsible for the regulation of COR genes containing CRT/DRE(CCGAC)motifs in their promoters. CBFs are activated by ICE1 and CAMTA transcription factors,whereas repressed by MYB15. HOS1 and SIZ1 encode RING E3 ligase and SUMO E3 ligase,respectively,which antagonistically regulate the abundance of ICE1 protein. OST1 is a kinase that positively regulates the abundance of the ICE1 protein. Crooked arrows indicate post-translational regulation,and solid arrows indicate activation. T-shaped bars represent repression;the solid line represent direct interaction. The same below

图2 E3s参与乙烯途径调控拟南芥抗冻性示意图在乙烯生物合成方面,E3s ETO1和EOL1/2与ACS5相互作用并通过蛋白酶体依赖途径降解而抑制其活性,介导乙烯生物合成。在乙烯生物合信号转导方面,在没有乙烯的情况下,高度磷酸化的乙烯受体激活CTR1激酶活性,进而磷酸化EIN2,可能导致由F-box蛋白EPT1和EPT2介导的EIN2的降解。同时,EIN3/EILs也受到F-box蛋白EBF1和EBF2介导的蛋白酶体降解。在乙烯存在的情况下,乙烯与受体的结合抑制受体的磷酸化,从而使受体失活,结果导致CTR1失活。去磷酸化的EIN2因此被剪切,其C端结构域转移到细胞核中,导致EIN3/EILs和下游转录级联的激活,最终调控抗冻性反应

Fig. 2 Schematic diagram depicts the involvement of E3s in regulation of freezing stress tolerance by the ethylene(ET)pathway in Arabidopsis In the biosynthesis of ethylene,the ETO1,EOL1 and EOL2 modulate ET biosynthesis by interacting with the ET biosynthetic enzyme ACS5 and inhibiting its activity by proteasome-dependent degradation. In the signal transductions of ethylene,in the absence of ethylene,the highly phosphorylated ethylene receptors activate CTR1 kinase activity,which in turn phosphorylates EIN2,likely causing the degradation of EIN2 by F-box proteins EPT1 and EPT2. Meanwhile,EIN3/EILs are also subjected to proteasomal degradation mediated by F-box proteins EBF1 and EBF2. In the presence of ethylene,ethylene binding inactivates the receptors by suppressing its phosphorylation,which consequently leads to deactivation of CTR1. The un-phosphorylated EIN2 is thus cleaved and its C-terminal domain is translocated into the nucleus,resulting in activation of EIN3/EILs and downstream transcriptional cascades. Finally,they regulate the freezing stress tolerance response

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