植物环核苷酸门控离子通道及其功能研究进展

doi:10.11983/CBB14146

植物学报 ›› 2015, Vol. 50 ›› Issue (6): 779-789.DOI: 10.11983/CBB14146

植物环核苷酸门控离子通道及其功能研究进展

刘海娇¹, 杜立群², 林金星^1,³, 李瑞丽^3,,A;^*()

¹中国科学院植物研究所, 北京 100093
²杭州师范大学生命与环境科学学院, 杭州 310036
³北京林业大学生物科学与技术学院, 北京 100083

收稿日期:2014-08-07 接受日期:2014-10-25 出版日期:2015-11-01 发布日期:2015-09-06
通讯作者: 李瑞丽
作者简介:
? 共同第一作者
基金资助:
国家重点基础研究发展规划(No.2011CB809103)、国家自然科学基金创新研究群体科学基金(No.31121065)、国家自然科学基金(No.31270224, No.31401149)、高等学校学科创新引智计划(No.111 project, B13007)、长江学者和创新团队发展计划(No;IRT13047)及中央高校基本科研业务费专项资金(No;BLX2014-19)

Recent Advances in Cyclic Nucleotide-gated Ion Channels with their Functions in Plants

Haijiao Liu¹, Liqun Du², Jinxing Lin^1,3, Ruili Li^3*

¹Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China
²College of Life and Environmental Sciences, Hangzhou Normal University, Hangzhou 310036, China
³College of Biological Sciences and Technology, Beijing Forestry University, Beijing 100083, China

Received:2014-08-07 Accepted:2014-10-25 Online:2015-11-01 Published:2015-09-06
Contact: Li Ruili
About author:
? These authors contributed equally to this paper

摘要/Abstract

摘要： 环核苷酸门控离子通道(CNGC)是非选择性的阳离子通道, 可以直接被细胞内信使小分子——环核苷酸(cAMP和cGMP)活化。该通道蛋白包含6个跨膜α-螺旋, C端各具一个交叠的环核苷酸与钙调蛋白结合区。CNGC广泛存在于各种植物细胞中。研究表明, 模式植物拟南芥(Arabidopsis thaliana)的CNGC家族有20个成员, 分为4个亚群, 它们在抗病、花粉管生长、对Ca²⁺响应、抵抗重金属离子毒害和抗盐等多种信号途径中发挥重要作用, 协助植物细胞应对各种生物与非生物胁迫。该文简要介绍了CNGC的结构、表达谱及其调控因子, 并着重总结了近年来CNGC生物学功能的研究进展, 以期为今后系统开展其功能研究提供理论依据。

Abstract: Cyclic nucleotide-gated channels (CNGCs) are non-selective cation channels that can be activated directly by cytosolic second messengers, such as the CNs cGMP and cAMP. CNGCs have six predicted transmembrane α-helices, followed by a cyclic nucleotide-binding domain (CNBD) and a highly conserved calmodulin-binding domain (CaMBD); the latter two domains partially overlap at the C-terminal cytosolic region. CNGCs exist widely in different kinds of plant cells. In the model plant Arabidopsis, CNGCs represent 20 members divided into four groups. A large body of evidence showed that the CNGCs play essential roles in various signaling pathways, including pathogen responses, pollen tube growth, Ca²⁺ signaling, tolerance to heavy metal poisoning and resistance to salinity and thus assist the plant cell response to various biological and abiotic stresses. This review focuses on the structure, expression profile and regulatory factors of CNGCs, with special emphasis on recent progress in biological functions, which will provide valuable information for further study on CNGCs in plants.

刘海娇, 杜立群, 林金星, 李瑞丽. 植物环核苷酸门控离子通道及其功能研究进展. 植物学报, 2015, 50(6): 779-789.

Haijiao Liu, Liqun Du, Jinxing Lin, Ruili Li. Recent Advances in Cyclic Nucleotide-gated Ion Channels with their Functions in Plants. Chinese Bulletin of Botany, 2015, 50(6): 779-789.

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链接本文: https://www.chinbullbotany.com/CN/10.11983/CBB14146

https://www.chinbullbotany.com/CN/Y2015/V50/I6/779

图/表 4

图1 动物(A)和植物(B)环核苷酸门控离子通道(CNGC)的膜拓扑结构示意图(Hua et al., 2003)。跨膜域由6个α-螺旋(S1-S6)构成, S5和S6之间形成孔状结构(P loop)。动植物CNGCs的N和C末端均延伸至细胞胞质侧。动物CNGCs的CaMBD在N端, CNBD在C端。植物CNGCs的结构与动物CNGCs基本一致(除CaMBD在C端外)。

Figure 1 Predicted membrane topology and domain structures of a cyclic nucleotide-gated ion channels (CNGC) subunit from animal cells (A) and plant cells (B) (Hua et al., 2003) .The bulk of CNGC are made of 6 transmembrane α-helices (S1-S6), a pore structure (P) is formed between S5 and S6. Both the N- and C-termini of plant and animal CNGCs extend into the cytosolic side of the plasma membrane. The CaMBD in animal CNGCs is located at the N-terminus and its CNBD is at the C-terminus. The structure of plant CNGC is similar to its counterparts from animals except that the CaMBD is at the C-terminus.

图2 用邻接树法构建的CNGCs无根进化树(Finka et al., 2012) 从进化树可以看出, 来自植物和人类的CNGCs分为4种明显的基因簇: 基因簇I (绿色)由苔藓特有的CNGCs构成; 基因簇II (蓝色)仅由植物特有的CNGCs构成; 基因簇III (黑色)包含苔藓和种子植物的CNGCs, 由此又被称为陆生植物所特有的CN- GCs; 基因簇IV (品红), 仅包含人类的CNGCs。

Figure 2 Unrooted phylogenetic tree generated using neigh- bor joining method (Finka et al., 2012).The phylogenetic tree shows CNGCs from plants and human could be grouped into four distinct clusters: cluster I (green) containing moss-specific CNGCs; cluster II (blue) are higher plant CNGCs; cluster III (black), comprise of both moss and seed plant CNGCs and is therefore inferred to be land plant specific; and cluster IV (magenta) containing only human CNGCs.

图3 植物固有免疫系统应对病原入侵的早期措施描述模式(Ali et al., 2007)(1) 细胞外PAMP或诱导子被植物细胞质膜上假设的受体识别; (2) 病原菌或者PAMP诱导子被受体识别激活CNGC2 (或者通过上调核苷酸三磷酸环化酶增加胞内活性配体的数量, 或者通过其它未知途径); (3) 内在CNGC2流的激活导致胞内暂时的Ca2+浓度增加; (4) Ca2+内流导致胞内Ca2+/CaM含量升高; (5) 胞内Ca2+/CaM含量升高抑制了CNGC2, 进而结束了短暂的胞质Ca2+飙升。

Figure 3 Model illustrating early events of plant innate immunity in response to avirulent pathogens (Ali et al., 2007) (1) The presence of extracellular PAMP/elicitor is recognized by a hypothetic receptor on the plasma membrane; (2) Pathogen or PAMP/elicitor recognition by this receptor activates CNGC2 (either by an increase in cytosolic level of cNMP ligand through the up-regulation of a nucleotide triphosphate cyclase or by an unknown mechanism); (3) Activation of inward CNGC2 current results in a (transient) increase in cytosolic Ca2+; (4) Cytosolic Ca2+/CaM level increases due to influx of Ca2+ into the cell; (5) Elevated level of cytosol Ca2+/CaM inhibits CNGC2, ending the transient cytosolic Ca2+ burst.

图4 35S-AtCNGC2-GFP融合蛋白的亚细胞定位(A, A1, A2) 叶表皮细胞; (B, B1, B2) 气孔保卫细胞; (C, C1, C2) 胚轴细胞; (D, D1, D2) 根细胞。其中图A、B、C和D是激光扫描共聚焦显微镜图片; 图A1、B1、C1和D1分别是对应的明场图片; 图A2、B2、C2和D2分别是对应的叠加图片。Bars=20 μm

Figure 4 Subcellular localization of AtCNGC2-GFP in trans- genic Arabidopsis seedlings (A, A1, A2) Leave pavement cells; (B, B1, B2) Stomatal guard cells; (C, C1, C2) Hypocotyl cells; (D, D1, D2) Root cells. Cells were imaged by laser scanning confocal microscopy (Figures A, B, C, D), differential interference contrast microscopy (Figures A1, B1, C1, D1) and merged images (Figures A2, B2, C2, D2). Bars=20 μm

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植物环核苷酸门控离子通道及其功能研究进展

Recent Advances in Cyclic Nucleotide-gated Ion Channels with their Functions in Plants

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