植物CBL-CIPK信号系统响应非生物胁迫的调控机制

doi:10.11983/CBB21024

摘要/Abstract

摘要： 钙调磷酸酶B蛋白(CBLs)及其互作蛋白激酶(CIPKs)组成的信号系统是非生物逆境响应的重要调控网络。CBL-CIPK系统通过磷酸化感应并解码Ca²⁺信号, 参与植物对非生物胁迫的应答调控。该文综述了CBLs和CIPKs结构、CBLs-CIPKs对不同底物磷酸化及其响应非生物胁迫调控机制的研究进展, 并展望了未来的研究方向, 以期为作物抗逆性遗传改良提供思路。

关键词: 非生物胁迫, 钙信号, CBL-CIPK, 离子通道, 磷酸化

Abstract: Calcineurin B-like proteins (CBLs) and their CBL-interacting protein kinases (CIPKs) are important regulatory network in response to abiotic stresses. The CBL-CIPK system senses and decodes Ca²⁺-signals through phosphorylation to regulate plant response to abiotic stresses. In this review, the basic structures of CBLs and CIPKs, and their phosphorylation on different substrates, as well as regulatory mechanisms of plants in response to abiotic stresses were summarized. We also put forward a perspective on the future research directions of CBLs and CIPKs, as well as their potential applications in genetic improvement of crops for stress tolerance.

Key words: abiotic stresses, calcium signaling, CBL-CIPK, ion channels, phosphorylation

谢玲玲, 王金龙, 伍国强. 植物CBL-CIPK信号系统响应非生物胁迫的调控机制. 植物学报, 2021, 56(5): 614-626.

Lingling Xie, Jinlong Wang, Guoqiang Wu. Regulatory Mechanisms of the Plant CBL-CIPK Signaling System in Response to Abiotic Stress. Chinese Bulletin of Botany, 2021, 56(5): 614-626.

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链接本文: https://www.chinbullbotany.com/CN/10.11983/CBB21024

https://www.chinbullbotany.com/CN/Y2021/V56/I5/614

图/表 4

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物种	CBLs数量	CIPKs数量	参考文献
拟南芥(Arabidopsis thaliana)	10	26	Kolukisaoglu et al., 2004
油菜(Brassica napus)	7	23	Zhang et al., 2014a
杨树(Populus trichocarpa)	10	25	Weinl and Kudla, 2009
辣椒(Capsicum annuum)	9	26	Ma et al., 2019
水稻(Oryza sativa)	10	31	Weinl and Kudla, 2009; Piao et al., 2010
茶(Camellia sinensis)	7	18	Liu et al., 2019
小麦(Triticum aestivum)	24	79	Sun et al., 2015; Liu et al., 2018
芜菁(B. rapa var. rapa)	19	51	Yin et al., 2017
木薯(Manihot esculenta)	8	26	Hu et al., 2015; Mo et al., 2018
茄子(Solanum melongena)	5	15	Li et al., 2016
菠萝(Ananas comosus)	8	21	Aslam et al., 2019
葡萄(Vitis vinifera)	8	20	Xi et al., 2017
玉米(Zea mays)	8	43	Chen et al., 2011
甜菜(Beta vulgaris)	7	20	未发表数据

物种	CBLs数量	CIPKs数量	参考文献
拟南芥(Arabidopsis thaliana)	10	26	Kolukisaoglu et al., 2004
油菜(Brassica napus)	7	23	Zhang et al., 2014a
杨树(Populus trichocarpa)	10	25	Weinl and Kudla, 2009
辣椒(Capsicum annuum)	9	26	Ma et al., 2019
水稻(Oryza sativa)	10	31	Weinl and Kudla, 2009; Piao et al., 2010
茶(Camellia sinensis)	7	18	Liu et al., 2019
小麦(Triticum aestivum)	24	79	Sun et al., 2015; Liu et al., 2018
芜菁(B. rapa var. rapa)	19	51	Yin et al., 2017
木薯(Manihot esculenta)	8	26	Hu et al., 2015; Mo et al., 2018
茄子(Solanum melongena)	5	15	Li et al., 2016
菠萝(Ananas comosus)	8	21	Aslam et al., 2019
葡萄(Vitis vinifera)	8	20	Xi et al., 2017
玉米(Zea mays)	8	43	Chen et al., 2011
甜菜(Beta vulgaris)	7	20	未发表数据

CBL	CIPK	靶标蛋白	定位	功能	参考文献
CBL4/SOS3	CIPK24/SOS2	SOS1	质膜	增强耐盐性	Zhu et al., 1998; Yin et al., 2020
CBL10	CIPK24	NHX7	质膜/液泡膜	增强耐盐性	Hu et al., 2015; Plasencia et al., 2021
CBL10	CIPK8	SOS1	质膜	增强耐盐性	Yin et al., 2020
CBL2/3	CIPK21	-	液泡膜	增强耐盐性	Sanyal et al., 2016
CBL2/7	CIPK11	H⁺-ATPase	质膜	维持pH平衡	Saito and Uozumi, 2020
CBL2/3	CIPK3/9/23/26	V-ATPase	液泡膜	维持pH平衡	Tang et al., 2012, 2015
CBL1	CIPK23	AMT1;1/AMT1;2	质膜	维持NH₄⁺平衡	Straub et al., 2017
-	CIPK8	NRT2.1	-	维持NO₃^-稳态	Hu et al., 2009
CBL1/9	CIPK23	CHL1/NPF6.3	质膜	维持NO₃^-稳态	Ho et al., 2009; Chu et al., 2021
CBL1/9	CIPK23	SLAH2/SLAH3	质膜	维持NO₃^-稳态	Léran et al., 2015
CBL7	-	NRT2.4/NRT2.5	-	维持NO₃^-稳态	Ma et al., 2015
CBL1/9	CIPK23	AKT1/HAK5	质膜	维持K⁺稳态	Lara et al., 2020
CBL4	CIPK6	AKT2	质膜	维持K⁺稳态	Saito and Uozumi, 2020
CBL2/3	CIPK3/9/23/26	TPK	液泡膜	维持K⁺稳态	Tang et al., 2020; Dong et al., 2021
CBL1/9	CIPK26	RBOHF	质膜	响应ROS信号	Zhang et al., 2014b
CBL1/9	CIPK23	SLAC1/SLAC3	质膜	响应ABA信号	Maierhofer et al., 2014a, 2014b
CBL5	CIPK11	SLAC1	质膜	响应ABA信号	Saito et al., 2018
CBL1	CIPK15	ABI1/ABI2	-	响应ABA信号	Guo et al., 2002
CBL9	CIPK3	ABR1	-	响应ABA信号	Sanyal et al., 2017; Wang et al., 2018
CBL2/3	CIPK9/17	PAT10	液泡膜	响应ABA信号	Song et al., 2018
-	CIPK26/11	ABI5	-	响应ABA信号	Zhou et al., 2015; Lyzenga et al., 2017
CBL2/3	CIPK3/9/23/26	Mg²⁺转运蛋白	液泡膜	维持Mg²⁺平衡	Tang et al., 2015

CBL	CIPK	靶标蛋白	定位	功能	参考文献
CBL4/SOS3	CIPK24/SOS2	SOS1	质膜	增强耐盐性	Zhu et al., 1998; Yin et al., 2020
CBL10	CIPK24	NHX7	质膜/液泡膜	增强耐盐性	Hu et al., 2015; Plasencia et al., 2021
CBL10	CIPK8	SOS1	质膜	增强耐盐性	Yin et al., 2020
CBL2/3	CIPK21	-	液泡膜	增强耐盐性	Sanyal et al., 2016
CBL2/7	CIPK11	H⁺-ATPase	质膜	维持pH平衡	Saito and Uozumi, 2020
CBL2/3	CIPK3/9/23/26	V-ATPase	液泡膜	维持pH平衡	Tang et al., 2012, 2015
CBL1	CIPK23	AMT1;1/AMT1;2	质膜	维持NH₄⁺平衡	Straub et al., 2017
-	CIPK8	NRT2.1	-	维持NO₃^-稳态	Hu et al., 2009
CBL1/9	CIPK23	CHL1/NPF6.3	质膜	维持NO₃^-稳态	Ho et al., 2009; Chu et al., 2021
CBL1/9	CIPK23	SLAH2/SLAH3	质膜	维持NO₃^-稳态	Léran et al., 2015
CBL7	-	NRT2.4/NRT2.5	-	维持NO₃^-稳态	Ma et al., 2015
CBL1/9	CIPK23	AKT1/HAK5	质膜	维持K⁺稳态	Lara et al., 2020
CBL4	CIPK6	AKT2	质膜	维持K⁺稳态	Saito and Uozumi, 2020
CBL2/3	CIPK3/9/23/26	TPK	液泡膜	维持K⁺稳态	Tang et al., 2020; Dong et al., 2021
CBL1/9	CIPK26	RBOHF	质膜	响应ROS信号	Zhang et al., 2014b
CBL1/9	CIPK23	SLAC1/SLAC3	质膜	响应ABA信号	Maierhofer et al., 2014a, 2014b
CBL5	CIPK11	SLAC1	质膜	响应ABA信号	Saito et al., 2018
CBL1	CIPK15	ABI1/ABI2	-	响应ABA信号	Guo et al., 2002
CBL9	CIPK3	ABR1	-	响应ABA信号	Sanyal et al., 2017; Wang et al., 2018
CBL2/3	CIPK9/17	PAT10	液泡膜	响应ABA信号	Song et al., 2018
-	CIPK26/11	ABI5	-	响应ABA信号	Zhou et al., 2015; Lyzenga et al., 2017
CBL2/3	CIPK3/9/23/26	Mg²⁺转运蛋白	液泡膜	维持Mg²⁺平衡	Tang et al., 2015