植物CBL调控逆境胁迫响应的作用机制

doi:10.13560/j.cnki.biotech.bull.1985.2024-1265

摘要/Abstract

摘要：

钙调磷酸酶B蛋白（calcineurin B like protein, CBL）是一类植物特有的Ca²⁺传感器，在植物生长发育和逆境胁迫响应中发挥重要作用。CBL由4个典型的用于Ca²⁺结合的手性延伸因子（elongation factor hands, EF-hands）结构域构成，并且每个EF-hand含有12个相对保守氨基酸组成的α-螺旋-环-α-螺旋结构。CBL启动子区域的顺式调控作用元件（如W-box、MBS和G-BOX等）可与上游转录因子（transcription factors, TFs）结合，通过激活或抑制下游基因的表达，从而进行转录调控。CBL通过脱落酸（abscisic acid, ABA）、呼吸爆发氧化酶同源物（respiratory burst oxidases homolog, RBOH）以及活性氧（reactive oxygen species, ROS）等信号通路调控植物气孔运动，减少水分蒸发，以适应各种逆境胁迫。大量研究表明，当植物受到盐、干旱、极端温度、营养胁迫和病原菌等逆境胁迫时，CBL能够迅速感知细胞内瞬时Ca²⁺信号变化，不仅与CBL互作蛋白激酶（CBL-interacting protein kinase, CIPK）互作磷酸化下游靶标蛋白实现Ca²⁺信号传导，还与其他蛋白质（如高亲和力K⁺转运蛋白5、蛋白S-酰化转移酶10和2C类蛋白磷酸酶等）互作，从而正向或负向调控植物的抗逆性。此外，CBL介导植物器官和组织生长发育，通过调节糖信号促进果实成熟。CBL也与开花时间基因（gigantea, GI）结合以影响植物的开花时间。论文综述了植物CBL的发现、结构、分类、调控机制、生物学功能及其调控逆境胁迫响应的作用机制，并对其未来研究方向进行了展望，以期为农作物抗逆性遗传改良和生物育种提供基因资源和理论依据。

关键词: 钙调磷酸酶B样蛋白, 调控机制, 蛋白互作, 生长发育, 抗逆性, 顺式调控元件, 蛋白激酶

Abstract:

Calcineurin B like protein (CBL) is a kind of specific Ca²⁺ sensor in plants, which plays a crucial role in plant growth and development, and the response to environmental stresses. CBL is composed of four typical elongation factor hands (EF-hands) domains for Ca²⁺ binding. Each EF-hand is α-helix-loop-α-helix structures composed of 12 relatively conserved amino acids. The cis-acting regulatory elements (such as W-box, MBS and G-BOX) in the promoter regionof CBL can bind with upstream transcription factors (TFs) to regulate transcription by activating or inhibiting the expression of downstream genes. CBL is involved in the regulation of signaling pathways such as abscisic acid (ABA), hormones, respiratory burst oxidases homolog (RBOH), and reactive oxygen species (ROS), to reduce water evaporation, thereby adapting various abiotic stresses. There are evidences that CBL can rapidly perceive changes in the transient intracellular Ca²⁺ signal when plants were subjected to abiotic and biotic stresses such as salt, drought, extreme temperature, nutrient stress and pathogen. CBLs not only interact with CBL-interacting protein kinase CIPK to phosphorylate downstream target proteins for calcium signal transduction, but also interact with other proteins (such as high-affinity K⁺ transporter 5, protein S-acyl transferase10 and type 2C protein phosphatases), thereby positively or negatively regulating stress tolerance in plant. Additionally, CBL mediates the growth and development of plant organs and tissues and promotes fruit ripening by regulating sugar signals. CBL also interacts with gigantea (GI) to affect the flowering time of plants. In this review, recent research findings on the discovery, structure, classification, regulatory mechanisms and roles of plant CBL in regulating the response to adversity stresses are summarized, and its future research directions are prospected, aiming to provide the gene resources and the theoretical basis for improving tolerance of crops and biological breeding to stress.

Key words: calcineurin B like protein, regulatory mechanism, protein-protein interaction, growth and development, stress tolerance, cis-acting regulatory elements, protein kinase

王从欢, 伍国强, 魏明. 植物CBL调控逆境胁迫响应的作用机制[J]. 生物技术通报, 2025, 41(7): 1-16.

WANG Cong-huan, WU Guo-qiang, WEI Ming. Functional Mechanism of Plant CBL in Regulating the Responses to Abiotic and Biotic Stresses[J]. Biotechnology Bulletin, 2025, 41(7): 1-16.

图/表 6

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物种 Species	基因名称 Gene name	基因数量 Number of genes	种类 Class	生物功能 Biological function	参考文献 Reference
拟南芥Arabidopsis thaliana	AtCBL	10	十字花科 Brassicaceae	生长发育、高盐、干旱、低温、营养平衡 Growth and development， high-salt， drought， low temperature， and nutrition balance	［19］
雷蒙德氏棉 Gossypium raimondii	GrCBL	13	锦葵科 Malvaceae	/	［20］
木薯 Manihot esculenta	MeCBL	8	大戟科 Euphorbiaceae	高温、营养平衡 High-temperature， and nutrition balance	［21］
菠萝 Ananas comosus	AcCBL	8	凤梨科 Bromeliaceae	高盐、极端温度、病害 High salt， extreme temperature， and disease	［8］
辣椒 Capsicum annuum	CaCBL	9	茄科 Solanaceae	高盐 High salt	［25］
甜橙 Citrus sinensis	CsCBL	8	芸香科 Rutaceae	干旱、病害 Drought， and disease	［26］
金银花Lonicera japonica	LjCBL	6	忍冬科 Caprifoliaceae	生长和发育、高盐、干旱 Growth and development， high salt， and drought	［27］
紫花苜蓿Medicago sativa	MsCBL	10	豆科 Leguminosae	生长和发育、高盐、干旱 Growth and development， high salt， and drought	［28］
蒺藜苜蓿Medicago truncatula	MtCBL	13	豆科 Leguminosae	高盐 High salt	［29］
甜根子草Saccharum spontaneum	SsCBL	10	禾本科 Graminoceae	营养平衡 Nutrition balance	［23］
甜菜 Beta vulgaris	BvCBL	7	藜科 Chenopodiaceae	干旱 Drought	［22］
藜麦Chenopodium quinoa	CqCBL	16	藜科 Chenopodiaceae	胚胎发育、干旱、低温 Embryonic development， drought stress， and low temperature	［30］
白菜 Brassica rapa	BraCBL	18	十字花科 Brassicaceae	高盐、低温、营养平衡 High salt， low temperature， and nutrition balance	［24］
桃 Prunus persica	PprCBL	8	蔷薇科 Rosaceae	糖平衡、干旱 Sugar balance， and drought	［31］
甜樱桃 Prunus avium	PavCBL	7	蔷薇科 Rosaceae	高盐 High salt	［32］
烟草 Nicotiana tabacum	NtCBL	24	茄科 Solanaceae	高盐 High salt	［33］
大豆 Glycine max	GmCBL	15	豆科 Leguminosae	生长发育、高盐、干旱 Growth and development， high salt， and drought	［34］
丹参 Salvia miltiorrhiza	SmCBL	8	唇形科 Lamiaceae	高盐 High salt	［35］

物种 Species	基因名称 Gene name	基因数量 Number of genes	种类 Class	生物功能 Biological function	参考文献 Reference
拟南芥Arabidopsis thaliana	AtCBL	10	十字花科 Brassicaceae	生长发育、高盐、干旱、低温、营养平衡 Growth and development， high-salt， drought， low temperature， and nutrition balance	［19］
雷蒙德氏棉 Gossypium raimondii	GrCBL	13	锦葵科 Malvaceae	/	［20］
木薯 Manihot esculenta	MeCBL	8	大戟科 Euphorbiaceae	高温、营养平衡 High-temperature， and nutrition balance	［21］
菠萝 Ananas comosus	AcCBL	8	凤梨科 Bromeliaceae	高盐、极端温度、病害 High salt， extreme temperature， and disease	［8］
辣椒 Capsicum annuum	CaCBL	9	茄科 Solanaceae	高盐 High salt	［25］
甜橙 Citrus sinensis	CsCBL	8	芸香科 Rutaceae	干旱、病害 Drought， and disease	［26］
金银花Lonicera japonica	LjCBL	6	忍冬科 Caprifoliaceae	生长和发育、高盐、干旱 Growth and development， high salt， and drought	［27］
紫花苜蓿Medicago sativa	MsCBL	10	豆科 Leguminosae	生长和发育、高盐、干旱 Growth and development， high salt， and drought	［28］
蒺藜苜蓿Medicago truncatula	MtCBL	13	豆科 Leguminosae	高盐 High salt	［29］
甜根子草Saccharum spontaneum	SsCBL	10	禾本科 Graminoceae	营养平衡 Nutrition balance	［23］
甜菜 Beta vulgaris	BvCBL	7	藜科 Chenopodiaceae	干旱 Drought	［22］
藜麦Chenopodium quinoa	CqCBL	16	藜科 Chenopodiaceae	胚胎发育、干旱、低温 Embryonic development， drought stress， and low temperature	［30］
白菜 Brassica rapa	BraCBL	18	十字花科 Brassicaceae	高盐、低温、营养平衡 High salt， low temperature， and nutrition balance	［24］
桃 Prunus persica	PprCBL	8	蔷薇科 Rosaceae	糖平衡、干旱 Sugar balance， and drought	［31］
甜樱桃 Prunus avium	PavCBL	7	蔷薇科 Rosaceae	高盐 High salt	［32］
烟草 Nicotiana tabacum	NtCBL	24	茄科 Solanaceae	高盐 High salt	［33］
大豆 Glycine max	GmCBL	15	豆科 Leguminosae	生长发育、高盐、干旱 Growth and development， high salt， and drought	［34］
丹参 Salvia miltiorrhiza	SmCBL	8	唇形科 Lamiaceae	高盐 High salt	［35］

胁迫响应 Response to stress	基因名称 Gene name	物种 Species	功能 Function	参考文献References
盐胁迫 Salinity stress	AtCBL1/9/10	拟南芥 A. thaliana	提高基因表达量；SOS途径调节离子平衡；清除ROS减少细胞氧化损伤；依赖ABA和SA途径提高植物耐盐性 Increasing gene expression； SOS pathway regulates ion balance； eliminating ROS and reducing oxidative damage of cells； improving salt tolerance of plants by ABA and SA	［76-78］
	LjCBL2/4	金银花 L. japonica		［27］
	MsCBL04.1/12.1	紫花苜蓿 M. sativa		［28］
	PeCBL4	胡杨 Populus euphratica		［79］
	NtCBL5A	烟草 N. tabacum		［80］
	PavCBL4	甜樱桃 P. avium		［32］
	AcCBL1	菠萝 A. comosus		［8］
	GmCBL1	大豆 G. max		［34］
	BnCBL1	甘蓝性油菜Brassica napus		［81］
	MdCBL10.1	苹果 M. domestica		［82］
	DoCBL5/8	油柿 Diospyros oleifera		［83］
	MtCBL10	蒺藜苜蓿Medicago truncatula		［29］
	DcCBL1/4/5/6/10	铁皮石斛 D. catenatum		［44］
	BraCBL1.2/3.3/8/9.1/9.2	白菜 B. rapa		［24］
	OsCBL4/8	水稻 O. sativa		［50］
	CaCBL2/3/9	辣椒 C. annuum		［25］
	SmCBL3	丹参 S. miltiorrhiza		［35］
干旱胁迫 Drought stress	AtCBL1/5/9	拟南芥 A. thaliana	提高基因表达量；通过AKT1信号途径吸收K⁺导致植物叶片气孔关闭；介导ABA依赖的气孔运动，提高抗旱 Increasing gene expression； absorption of K⁺ through AKT1 signal pathway leads to stomatal closure of plant leaves； mediating ABA-dependent stomatal movement and improving drought resistance	［84］
	LjCBL2/3	金银花 L. japonica		［27］
	BvCBL3	甜菜 B. vulgaris		［22］
	GmCBL1	大豆 G. max		［34］
	DcCBL4	铁皮石斛 D. catenatum		［44］
	MsCBL04.1/12.1	紫花苜蓿 M. sativa		［28］
	OsCBL8	水稻 O. sativa		［50］
	CcCBL1	鸽豆 Cajanus cajan		［85-86］
	PprCBL1/2/5/6	桃 P. persica		［31］
	CqCBL13	藜麦 C. quinoa		［30］
	DoCBL5/8	油柿 Diospyros oleifera		［83］
	CsCBL1/7/8	甜橙 C. sinensis		［26］
冷胁迫 Cold stress	AcCBL1	菠萝 A. comosus	参与ABA和SA途径提高耐寒性；与CBF途径相互作用，参与基因调控和ROS平衡；与MAPK网络协同作用调控TFs Participating in ABA and SA pathway to improve cold tolerance； interacting with CBF pathway， participating in gene regulation and ROS balance； synergistic effect with MAPK network to regulate transcription factors	［8］
	BraCBL2.2/3.1/4.3/6/10.3	白菜 B. rapa		［24］
	KoCBL	秋茄树Kandelia obovate		［13］
	DcCBL1/4	铁皮石斛 D. catenatum		［44］
	VaCBL	山葡萄 Vitis amurensis		［87］
	AtCBL1	拟南芥 A. thaliana		［88］
	CqCBL8	藜麦 C. quinoa		［74］
	CuCBL5/6/8	柑橘 C. unshiu		［89］
高温胁迫 Heat stress	AcCBL1	菠萝 A. comosus	上调基因表达量；参与热应激；调控TFs抵抗高温 Up-regulating gene expression； involved in heat stress； combining transcription factors to resist high temperatures	［8］
	CsCBL4	茶树 Camellia sinensis		［90］
	MeCBL5	木薯 M. esculenta		［21］
	OsCBL8	水稻 O. sativa		［50］
	DcCBL6/10	铁皮石斛 D. catenatum		［44］
营养胁迫 Nutritional stress	AtCBL1/2/3/4/9	拟南芥 A. thaliana	参与多种信号网络；磷酸化相关蛋白和TFs削弱下游基因的表达 Participating in various signal networks； phosphorylation-related proteins and transcription factors weaken the expression of downstream genes	［7，47-48，75，91-95］
	BnCBL1	甘蓝性油菜Brassica napus		［81］
	MeCBL1/9	木薯 M. esculenta		［96］
	SsCBL01	甜根子草Saccharum spontanum		［23］
	DoCBL5	油柿 Diospyros oleifera		［83］
生物胁迫 Biotic stress	OsCBL4	水稻 O. sativa	提高基因表达量；与下游蛋白RBOHB相互作用，促进ROS增加；靶向NAC77和JAMYB以赋予对病原体的抗性 Increasing gene expression； interacting with downstream protein RBOHB to promote the increase of ROS； targeting NAC77 and JAMYB to confer resistance to pathogens	［50］
	TaCBL4	小麦 Triticum aestivum		［97］
	AcCBL1	菠萝 A. comosus		［8］
	StCBL4	马铃薯Solanum tuberosum		［98］
	CsCBLs	甜橙 C. sinensis		［26，89］
	ScCBL	甘蔗 Saccharum spp.		［99］

胁迫响应 Response to stress	基因名称 Gene name	物种 Species	功能 Function	参考文献References
盐胁迫 Salinity stress	AtCBL1/9/10	拟南芥 A. thaliana	提高基因表达量；SOS途径调节离子平衡；清除ROS减少细胞氧化损伤；依赖ABA和SA途径提高植物耐盐性 Increasing gene expression； SOS pathway regulates ion balance； eliminating ROS and reducing oxidative damage of cells； improving salt tolerance of plants by ABA and SA	［76-78］
	LjCBL2/4	金银花 L. japonica		［27］
	MsCBL04.1/12.1	紫花苜蓿 M. sativa		［28］
	PeCBL4	胡杨 Populus euphratica		［79］
	NtCBL5A	烟草 N. tabacum		［80］
	PavCBL4	甜樱桃 P. avium		［32］
	AcCBL1	菠萝 A. comosus		［8］
	GmCBL1	大豆 G. max		［34］
	BnCBL1	甘蓝性油菜Brassica napus		［81］
	MdCBL10.1	苹果 M. domestica		［82］
	DoCBL5/8	油柿 Diospyros oleifera		［83］
	MtCBL10	蒺藜苜蓿Medicago truncatula		［29］
	DcCBL1/4/5/6/10	铁皮石斛 D. catenatum		［44］
	BraCBL1.2/3.3/8/9.1/9.2	白菜 B. rapa		［24］
	OsCBL4/8	水稻 O. sativa		［50］
	CaCBL2/3/9	辣椒 C. annuum		［25］
	SmCBL3	丹参 S. miltiorrhiza		［35］
干旱胁迫 Drought stress	AtCBL1/5/9	拟南芥 A. thaliana	提高基因表达量；通过AKT1信号途径吸收K⁺导致植物叶片气孔关闭；介导ABA依赖的气孔运动，提高抗旱 Increasing gene expression； absorption of K⁺ through AKT1 signal pathway leads to stomatal closure of plant leaves； mediating ABA-dependent stomatal movement and improving drought resistance	［84］
	LjCBL2/3	金银花 L. japonica		［27］
	BvCBL3	甜菜 B. vulgaris		［22］
	GmCBL1	大豆 G. max		［34］
	DcCBL4	铁皮石斛 D. catenatum		［44］
	MsCBL04.1/12.1	紫花苜蓿 M. sativa		［28］
	OsCBL8	水稻 O. sativa		［50］
	CcCBL1	鸽豆 Cajanus cajan		［85-86］
	PprCBL1/2/5/6	桃 P. persica		［31］
	CqCBL13	藜麦 C. quinoa		［30］
	DoCBL5/8	油柿 Diospyros oleifera		［83］
	CsCBL1/7/8	甜橙 C. sinensis		［26］
冷胁迫 Cold stress	AcCBL1	菠萝 A. comosus	参与ABA和SA途径提高耐寒性；与CBF途径相互作用，参与基因调控和ROS平衡；与MAPK网络协同作用调控TFs Participating in ABA and SA pathway to improve cold tolerance； interacting with CBF pathway， participating in gene regulation and ROS balance； synergistic effect with MAPK network to regulate transcription factors	［8］
	BraCBL2.2/3.1/4.3/6/10.3	白菜 B. rapa		［24］
	KoCBL	秋茄树Kandelia obovate		［13］
	DcCBL1/4	铁皮石斛 D. catenatum		［44］
	VaCBL	山葡萄 Vitis amurensis		［87］
	AtCBL1	拟南芥 A. thaliana		［88］
	CqCBL8	藜麦 C. quinoa		［74］
	CuCBL5/6/8	柑橘 C. unshiu		［89］
高温胁迫 Heat stress	AcCBL1	菠萝 A. comosus	上调基因表达量；参与热应激；调控TFs抵抗高温 Up-regulating gene expression； involved in heat stress； combining transcription factors to resist high temperatures	［8］
	CsCBL4	茶树 Camellia sinensis		［90］
	MeCBL5	木薯 M. esculenta		［21］
	OsCBL8	水稻 O. sativa		［50］
	DcCBL6/10	铁皮石斛 D. catenatum		［44］
营养胁迫 Nutritional stress	AtCBL1/2/3/4/9	拟南芥 A. thaliana	参与多种信号网络；磷酸化相关蛋白和TFs削弱下游基因的表达 Participating in various signal networks； phosphorylation-related proteins and transcription factors weaken the expression of downstream genes	［7，47-48，75，91-95］
	BnCBL1	甘蓝性油菜Brassica napus		［81］
	MeCBL1/9	木薯 M. esculenta		［96］
	SsCBL01	甜根子草Saccharum spontanum		［23］
	DoCBL5	油柿 Diospyros oleifera		［83］
生物胁迫 Biotic stress	OsCBL4	水稻 O. sativa	提高基因表达量；与下游蛋白RBOHB相互作用，促进ROS增加；靶向NAC77和JAMYB以赋予对病原体的抗性 Increasing gene expression； interacting with downstream protein RBOHB to promote the increase of ROS； targeting NAC77 and JAMYB to confer resistance to pathogens	［50］
	TaCBL4	小麦 Triticum aestivum		［97］
	AcCBL1	菠萝 A. comosus		［8］
	StCBL4	马铃薯Solanum tuberosum		［98］
	CsCBLs	甜橙 C. sinensis		［26，89］
	ScCBL	甘蔗 Saccharum spp.		［99］