相分离调控植物胁迫感知和应答的研究进展

doi:10.13560/j.cnki.biotech.bull.1985.2023-0720

摘要/Abstract

摘要：

生物大分子凝聚体（biomolecular condensates）是由相分离（phase separation）驱动形成的具有特定功能的无膜细胞器（MLOs），为特定的生化反应提供微环境，对细胞生命活动进行精细的时空调控。相分离是一种高度动态的过程，对温度、pH、盐浓度等理化因素的变化非常敏感。因此，相分离可以快速响应外界刺激，作为生物感受器感知压力信号，参与胁迫应答。近年来，相分离在植物中的应用受到越来越多的关注，特别是在胁迫感知和逆境响应方面。本文基于近期关于相分离响应胁迫的研究，探讨了相分离在胁迫信号感知和应答中的机制，综述了相分离在植物响应胁迫的研究成果，旨在为进一步研究植物胁迫感知和逆境响应中相分离的作用提供参考依据。

关键词: 生物大分子凝聚体, 相分离, 胁迫感受器, 植物, 胁迫响应

Abstract:

The formation of biomolecular condensates via phase separation has emerged as the key way of organization inside cells. Biomolecular condensates possess specific functions and provide a microenvironment for specific biochemical reactions, and finely regulates cellular activities in space and time. Phase separation is a highly dynamic process that is extremely sensitive to changes in temperature, pH, salt concentration, and other physicochemical factors. Consequently, phase separation can rapidly respond to external stimuli, serving as a stress sensor to perceive and transduce stress signals. In recent years, the involvement of phase separation in plants has gained increasing attention, particularly in the context of stress sensing and response. This article summarizes recent research on the roles of phase separation in stress signal perception and response, and provides an overview of the research progress regarding the role of phase separation in plant stress responses. The aim is to provide insights for further studies on the roles of phase separation in stress perception and adaptation in plants.

Key words: biomolecular condensate, phase separation, stress sensor, plant, stress response

张红红, 方晓峰. 相分离调控植物胁迫感知和应答的研究进展[J]. 生物技术通报, 2023, 39(11): 44-53.

ZHANG Hong-hong, FANG Xiao-feng. Advances in the Regulation of Stress Sensing and Responses by Phase Separation in Plants[J]. Biotechnology Bulletin, 2023, 39(11): 44-53.

图/表 1

参考文献 81

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蛋白质 Protein	胁迫信号 Stress signal	分子机制 Molecular mechanism	相分离功能 Functions of phase separation	文献 Reference
FLOE1	水势	由DS和QPS这两个IDR施加的相反力调控LLPS	介导水分胁迫响应，调控种子萌发	[60]
ELF3	热胁迫	PrD驱动LLPS，温度敏感性受PrD的PolyQ重复序列和ELF4的水平调节	高温胁迫感受器，调控植物开花	[42]
PhyB	光照和温度	NTE的无序性和C末端结构寡聚化共同驱动LLPS，NTE直接感知温度信号	通过可逆构象转变（光控）与LLPS（温控），同时感知光照与温度	[48]
SEU	渗透胁迫	LLPS取决于IDR1，IDR1的2个α-螺旋的构象变化感知大分子拥挤	渗透胁迫感受器，增强植物抗逆性	[54]
RBGD2/4	热胁迫	LCD富集酪氨酸提供疏水作用驱动LLPS，促进耐热相关的蛋白和转录本招募于SGs	响应热胁迫，增强拟南芥耐热性	[44]
NPR1	SA	IDRs的保守半胱氨酸簇介导LLPS，形成SINCs，富集细胞死亡调控和应激响应蛋白，在各种胁迫下促进细胞存活	响应SA，调控细胞程序性死亡和防御反应	[76]
GBPL3	生物胁迫	病原菌侵染诱导GBPL3的LLPS，促进GBPL3与防御相关基因启动子结合，并招募Mediator和RNAPII的组分，实现宿主防御	响应免疫信号，调控植物防御相关基因转录，增强抗病性。	[49]
TMF	氧化胁迫	半胱氨酸的巯基氧化，生成二硫键和IDR共同驱动LLPS	响应氧化信号，调控番茄茎尖分生组织的成熟和开花转换	[59]
ALBA4/5/6	热胁迫	热胁迫诱导ALBA的LLPS，协助SGs和P-bodies招募HSFmRNAs，抑制HSF mRNAs的降解	响应热胁迫，稳定HSF mRNAs，调控植物耐热性	[45]
FRI	低温	低温信号诱导FRI核凝聚体形成，使FRI从FLC基因上脱离，FLC反义RNA COOLAIR促进FRI凝聚体积累，稳定FRI蛋白，以快速响应温度变化	响应冷信号，抑制FLC表达，调控植物响应季节性温度变化	[70]
NUP62/58/54	免疫信号	N端IDR富含芳香族氨基酸驱动LLPS，决定核孔复合体选择性和渗透性，调控MPK3蛋白的核转运，参与植物抵御多种生物胁迫	正向调控核孔复合体中央屏障的选择性运输，控植物对病虫害的防御作用	[79]
MED19a	缺氮胁迫	在缺氮条件下，MED19a赖氨酸乙酰化水平降低调控LLPS，增强与转录因子ORE1互作以激活缺氮衰老反应基因	响应缺氮胁迫，调控衰老反应基因	[80]
HEM1	免疫信号	LCD驱动LLPS，形成凝聚体，隔离翻译因子，抑制免疫基因翻译	响应ETI，参与植物防御反应调控	[74]
STM	盐胁迫	PrLD驱动LLPS，与BELL蛋白和MED8互作且共相分离	响应盐胁迫，增强自身转录调控活性，促进分生组织活性和耐盐性	[81]

蛋白质 Protein	胁迫信号 Stress signal	分子机制 Molecular mechanism	相分离功能 Functions of phase separation	文献 Reference
FLOE1	水势	由DS和QPS这两个IDR施加的相反力调控LLPS	介导水分胁迫响应，调控种子萌发	[60]
ELF3	热胁迫	PrD驱动LLPS，温度敏感性受PrD的PolyQ重复序列和ELF4的水平调节	高温胁迫感受器，调控植物开花	[42]
PhyB	光照和温度	NTE的无序性和C末端结构寡聚化共同驱动LLPS，NTE直接感知温度信号	通过可逆构象转变（光控）与LLPS（温控），同时感知光照与温度	[48]
SEU	渗透胁迫	LLPS取决于IDR1，IDR1的2个α-螺旋的构象变化感知大分子拥挤	渗透胁迫感受器，增强植物抗逆性	[54]
RBGD2/4	热胁迫	LCD富集酪氨酸提供疏水作用驱动LLPS，促进耐热相关的蛋白和转录本招募于SGs	响应热胁迫，增强拟南芥耐热性	[44]
NPR1	SA	IDRs的保守半胱氨酸簇介导LLPS，形成SINCs，富集细胞死亡调控和应激响应蛋白，在各种胁迫下促进细胞存活	响应SA，调控细胞程序性死亡和防御反应	[76]
GBPL3	生物胁迫	病原菌侵染诱导GBPL3的LLPS，促进GBPL3与防御相关基因启动子结合，并招募Mediator和RNAPII的组分，实现宿主防御	响应免疫信号，调控植物防御相关基因转录，增强抗病性。	[49]
TMF	氧化胁迫	半胱氨酸的巯基氧化，生成二硫键和IDR共同驱动LLPS	响应氧化信号，调控番茄茎尖分生组织的成熟和开花转换	[59]
ALBA4/5/6	热胁迫	热胁迫诱导ALBA的LLPS，协助SGs和P-bodies招募HSFmRNAs，抑制HSF mRNAs的降解	响应热胁迫，稳定HSF mRNAs，调控植物耐热性	[45]
FRI	低温	低温信号诱导FRI核凝聚体形成，使FRI从FLC基因上脱离，FLC反义RNA COOLAIR促进FRI凝聚体积累，稳定FRI蛋白，以快速响应温度变化	响应冷信号，抑制FLC表达，调控植物响应季节性温度变化	[70]
NUP62/58/54	免疫信号	N端IDR富含芳香族氨基酸驱动LLPS，决定核孔复合体选择性和渗透性，调控MPK3蛋白的核转运，参与植物抵御多种生物胁迫	正向调控核孔复合体中央屏障的选择性运输，控植物对病虫害的防御作用	[79]
MED19a	缺氮胁迫	在缺氮条件下，MED19a赖氨酸乙酰化水平降低调控LLPS，增强与转录因子ORE1互作以激活缺氮衰老反应基因	响应缺氮胁迫，调控衰老反应基因	[80]
HEM1	免疫信号	LCD驱动LLPS，形成凝聚体，隔离翻译因子，抑制免疫基因翻译	响应ETI，参与植物防御反应调控	[74]
STM	盐胁迫	PrLD驱动LLPS，与BELL蛋白和MED8互作且共相分离	响应盐胁迫，增强自身转录调控活性，促进分生组织活性和耐盐性	[81]