LEA蛋白在逆境胁迫下的研究进展

doi:10.16035/j.issn.1001-7283.2025.04.001

摘要/Abstract

摘要：

植物在生长发育过程中不可避免地面临逆境胁迫，在应对胁迫的过程中，进化出一类响应逆境胁迫的功能性蛋白，如胚胎晚期发育丰富蛋白（late embryogenesis abundant，LEA）。LEA蛋白是一种高度亲水的富含甘氨酸的蛋白，广泛存在于植物中，其内在的无序性和高亲水性促进了LEA蛋白保护功能机制的多样性。LEA蛋白在响应逆境胁迫、激素信号和参与免疫应答等方面发挥重要作用。本文主要综述了LEA蛋白的结构分类、功能分析及逆境胁迫下LEA蛋白的功能研究，为深入了解LEA蛋白的抗逆机制以及通过基因工程提高生物的耐逆性提供参考。

关键词: LEA蛋白, 逆境胁迫, AhLEA1

Abstract:

Plants inevitably face stress during growth and development. In response to stress, a class of functional proteins has evolved, such as late embryogenesis abundant proteins (LEA). LEA proteins are highly hydrophilic glycine-rich proteins that are widely found in plants. Their intrinsic disorder and high hydrophilicity facilitate a variety of mechanisms for the protective functions of LEA proteins. LEA proteins have been found to play important roles in stress responses, hormone signaling and participation in immune responses. This paper primarily reviews the structural classification, functional analyses, as well as the functional research of LEA proteins under stress conditions. This provides insights into the resistance mechanisms of LEA proteins and serves as a reference for enhancing organism resistance through genetic engineering.

Key words: LEA protein, Stress condition, AhLEA1

黄若兰, 李帅, 蔡兆琴, 陈会鲜, 肖冬. LEA蛋白在逆境胁迫下的研究进展[J]. 作物杂志, 2025 (4): 1–8

Huang Ruolan, Li Shuai, Cai Zhaoqin, Chen Huixian, Xiao Dong. Research Progress of LEA Proteins under Stress Conditions[J]. Crops, 2025(4): 1–8

图/表 2

表1

表2

不同胁迫下LEA蛋白的功能

胁迫 Stress	物种 Species	基因/蛋白 Gene/protein	功能 Function	参考文献 Reference
干旱 Drought	腊梅	CpLEA	抗旱性与基因CpLEAs的过表达量呈正相关。	[27]
	玉米	ZmLEA34、ZmNHL1	胁迫条件下，基因ZmLEA34的表达量增加；ZmNHL1通过提高ROS清除能力和维持细胞膜通透性，促进了35S::ZmNHL1转基因植株对干旱胁迫的耐受性。	[28-29]
	油菜	BnaCPK5	BnaCPK5在一定程度上通过磷酸化2个核心ABA信号元件来调节RD29B的表达，从而充当耐旱性的正向调节因子。	[30]
	棉花	Gh_A08G0694	在四倍体棉花中敲除Gh_A08G0694后，棉花植株对盐碱和干旱胁迫的敏感性显著增加，该基因被发现与关键的非生物胁迫耐受基因、电压依赖性阴离子通道1（VDAC1）和甘油醛-3-磷酸脱氢酶A（gapA）有很强的相互作用。	[31]
	哺乳动物/黑腹果蝇	AfrLEA	哺乳动物细胞系和昆虫细胞系在短期暴露于干燥和高渗压力后的存活率明显高于未表达AfrLEA蛋白的细胞系。	[32]
	链格绿藻	LEA	LEA在藻类植物陆生化过程中的干燥耐受性发挥重要作用。	[33]
低温 Low temperature	枳	PtrLEA1、PtrLEA23	PtrLEA1、PtrLEA23受低温胁迫后高表达。	[34]
	大白菜	BrLEAs	BrLEAs受低温胁迫后高表达。	[35]
	茶树	CsLEA1	CsLEA1提高了大肠杆菌和酵母对冷胁迫的耐受性。	[36]
	梨树	PcLEA14	PcLEA14低温耐受机制与脱水反应元件结合蛋白/重复结合因子1（DREB1）有关。	[37]
	早花百子莲	ApY2SK2 DHN	ApY2SK2 DHN通过螯合金属离子（Fe³⁺、Cu²⁺）影响ROS代谢，从而激活抗氧化系统，降低低温保存过程中的膜脂过氧化，提高细胞复苏后的存活率。	[38]
	荷花	NnRab18	NnRab18可在低温保存过程中直接保护酶活性。	[39]
高温 High temperature	绿豆	VrLEA-2、VrLEA-40、 VrLEA-47、VrLEA-55	热胁迫下，VrLEAs表达显著上调。	[40]
	大豆	GmLEA4	钙调蛋白GmCaM1和GmLEA4互作，提高了种子清除ROS的能力，改善种子在高温高湿胁迫下的活力。	[41]
	豇豆	VuDREB2A	VuDREB2A与启动子中的DRE结合激活下游应激反应基因，热胁迫条件下表现出更强的渗透调节能力和光合作用活性。	[42]
盐Salt	纤枝短月藓	BeLEA5-1	BeLEA5-1蛋白的表达提高了大肠杆菌对盐胁迫的耐受力。	[43]
	黄麻	LEA14	LEA14在叶片和根部的表达量均在盐处理48 h时达到高峰。	[44]
	水稻	OsLEA5	OsLEA5参与了ABA介导的抗氧化防御，在水稻的干旱和盐胁迫响应中发挥作用。	[45]
	大豆	GsPM30	耐盐的正向调节因子GsCBRLK与GsPM30相互作用，提高了拟南芥幼苗期和成苗期的耐盐和耐旱性。	[46]
	紫花苜蓿	MsDIUP1	MsDIUP1可通过调节胁迫信号传导、抗氧化防御、离子平衡、渗透调节和光合作用来提高植物的耐盐性。	[47]
重金属 Heavy metal	玉米	ZmDHN13	ZmDHN13过表达烟草通过结合金属和减少ROS的积累，赋予烟草对铜胁迫的耐受性。	[48]
	耐辐射奇球菌	DrLEA3	定位于细胞膜上的DrLEA3蛋白纯化后，能够结合特定的金属离子，维持细胞内Mn²⁺、Fe³⁺等离子的动态平衡。	[49]
	水稻	OsLea14-A	过表达OsLea14-A降低了转基因植株中Na、Cr和Cu的积累，但增加了Hg的积累。	[15]
	花生	AhLEAs	在铝胁迫下，过表达AhLEAs增强了酿酒酵母的活力。	[50]
病虫害 Pests and diseases	谷子	SiLEA14	过表达SiLEA14的拟南芥植株对灰霉病的抗性增强。	[51]
	胡桃	JrLEAs	抗炭疽病品种中JrLEAs的表达水平相对较高。	[52]
	苏云金芽孢杆菌	Bt-LEA-Ⅱ	一种基于表达载体pHT01的LEA-Ⅱ多肽共表达系统，该系统具有强sigma A依赖性启动子，在IPTG诱导多肽12 h后，Bt晶体蛋白的产量提高了3倍。	[53]
	小麦	TdLEA3	过表达TdLEA3的拟南芥转基因植株对禾本科镰刀菌、灰霉病菌和黑曲霉具有耐受性。	[54]

表2

参考文献 59

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LEA蛋白分类 Classification of LEA proteins	结构特点 Structural feature	参考文献 Reference
LEA_1	C段序列保守性较差，N段有1个由74~78个氨基酸组成的保守序列。遭受胁迫时，保守序列由无序形成α-螺旋结构。	[11]
LEA_2	无明显保守片段，比其他LEA蛋白同源性低，含有高比例的疏水氨基酸，疏水性强。	[12]
LEA_3	含有大量的疏水氨基酸，热稳定性较差，沸水条件下不能维持水溶状态。	[13]
LEA_4	包括11个氨基酸的重复基序，其保守序列为TAQAAKEKAXE，推测含有2个亲水性α-螺旋。可以形成二聚体。其C端富含甘氨酸和组氨酸残基，能够结合金属离子，N端区域具有类似伴侣的活性。	[14]
LEA_5	保守结构域较少，缺少高特异性和显著的基团，疏水残基的含量高，被认为是一种非典型LEA蛋白。	[15]
LEA_6/PvLEA18	分子量小，只有7~14 kDa，不含半胱氨酸和色氨酸，具有较高的亲水性与热稳定性，暴露于高温时不会凝结。	[16]
Dehydrin	氨基酸组成为82~575个不等，不同成员间相对分子量在9~200 kDa，含有3个高度保守域：K、Y、S，K片段（[E/K/R]KKG[I/L]MDKIKEKLPG）（位于C末端）富含赖氨酸，Y片段（[V/T]D[E/Q]YGNP）（位于N末端）富含酪氨酸，S片段（SSSSSSSD）富含丝氨酸，可被蛋白激酶磷酸化。部分Dehydrin存在一段富含甘氨酸和极性氨基酸的φ片段，保守性较差。	[17]
SMP	分子量为5.3~37.4 kDa，与其他LEA亚族同源性较低，保守性强，亲水性和热稳定性高。	[7]