植物谷氨酸受体的研究进展

doi:10.11983/CBB15212

植物学报 ›› 2016, Vol. 51 ›› Issue (6): 827-840.DOI: 10.11983/CBB15212

植物谷氨酸受体的研究进展

何明洁¹, 孙伊辰², 程晓园², 时冬雪², 李迪秦¹, 陈益银³, 冯永坤², 刘璐², 范腾飞², 杨超³, 曹凤秋^4,,A;^*, 刘来华^1,^2,,A;^*

¹湖南农业大学农学院, 长沙 410128
²中国农业大学资源与环境学院, 北京 100193
³重庆烟草科学研究所, 重庆 400716
⁴中国科学院上海生命科学研究院植物生理生态研究所, 中国科学院上海植物逆境生物学研究中心, 上海 200032

收稿日期:2015-12-03 接受日期:2016-04-01 出版日期:2016-11-01 发布日期:2016-12-02
通讯作者: 曹凤秋,刘来华
作者简介:
# 共同第一作者
基金资助:
国家自然科学基金(No;Y333ZA1D11)、高等学校博士学科点专项科研基金(No.20134320110015)和中国烟草总公司重庆市公司科技计划(No;NY20140401070017)

Current Research Advances on Glutamate Receptors (GLRs) in Plants

Mingjie He^1,2†, Yichen Sun^2†, Xiaoyuan Cheng², Dongxue Shi², Diqin Li¹, Yiyin Chen³, Yongkun Feng², Lu Liu², Tengfei Fan², Chao Yang³, Fengqiu Cao^4*, Laihua Liu^1,2*

¹College of Agriculture Sciences, Hunan Agricultural University, Changsha 410128, China
²College of Resources and Environmental Sciences, China Agricultural University, Beijing 100193, China
³Institute of Tobacco Research of Chongqing, Chongqing 400716, China
⁴Shanghai Center for Plant Stress Biology, Institute of Plant Physiology Ecology, Shanghai Institutes for Biological Sciences, Chinese Academy of Sciences, Shanghai 200032, China

Received:2015-12-03 Accepted:2016-04-01 Online:2016-11-01 Published:2016-12-02
Contact: Cao Fengqiu,Liu Laihua
About author:
# Co-first authors

摘要/Abstract

摘要： 离子型谷氨酸受体(iGLuR)是哺乳动物中一类由L-氨基酸(如谷氨酸、甘氨酸)等配体门控的阳离子通道, 具有调节兴奋性神经信号传递和引导神经元发育等分子功能。1998年研究者在拟南芥(Arabidopsis thaliana)中发现了20个与iGLuRs同源的序列(即AtGLRs), 它们的功能涉及植物光信号传递、根尖分生细胞活性、花粉管生长、胞质Ca²⁺流以及应答多种生物和非生物环境胁迫等。该文从谷氨酸受体(GLR)的结构特征、离子通道激活与配体的关系、表达模式及可能的生物学功能等方面, 综述了近十几年来关于植物GLR和氨基酸信号的研究成果, 旨在为相关领域的同行提供有益的参考。

Abstract: In mammals, ionotropic glutamate receptors (iGLuR) are amino acids (e.g. glutamate and glycine) -gated cation channels, and exhibit molecular functions in the regulation of excitatory neurotransmission as well as in directing neuron growth. Since 1998 twenty genes homologous to iGLuR have been identified in Arabidopsis genome (termed AtGLRs), with reported functions involved in many biological processes including light signaling, root-tip meristematic cell activity, pollen tube growth, cytosolic calcium ion flux and response to varied biotic and abiotic stresses. This paper comprehensively summarizes research achievements or advances in terms of plant glutamate receptors and amino acid (e.g. glutamate) signaling in the past more than ten years, with major issues focusing on e.g. the protein structure of GLRs, a relationship between activation of ion channels and their ligands, their gene expression patterns as well as possible biological roles in plants, thus hopefully providing valuable information for researchers related to this field.

何明洁, 孙伊辰, 程晓园, 时冬雪, 李迪秦, 陈益银, 冯永坤, 刘璐, 范腾飞, 杨超, 曹凤秋, 刘来华. 植物谷氨酸受体的研究进展. 植物学报, 2016, 51(6): 827-840.

Mingjie He, Yichen Sun, Xiaoyuan Cheng, Dongxue Shi, Diqin Li, Yiyin Chen, Yongkun Feng, Lu Liu, Tengfei Fan, Chao Yang, Fengqiu Cao, Laihua Liu. Current Research Advances on Glutamate Receptors (GLRs) in Plants. Chinese Bulletin of Botany, 2016, 51(6): 827-840.

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链接本文: https://www.chinbullbotany.com/CN/10.11983/CBB15212

https://www.chinbullbotany.com/CN/Y2016/V51/I6/827

图/表 3

图1 来自细菌、拟南芥和人类的谷氨酸受体同源蛋白的系统进化树氨基酸序列的对比分析采用Lasergene DNA*软件的Clustal W方法, 并使用PAUP* 4.10b软件分析生成进化树。At: Arabidopsis thaliana (拟南芥); Hs: Homo sapiens (人类); Ssp: Synechocystis sp. (单细胞蓝藻菌属, 菌株PCC6803)。AtGLR蛋白序列取自Tair, HsGLuR1-7及SspGLuR0取自NCBI数据库。基因库登录编号或基因模式所编码的可能性完整蛋白质序列名称如下: AtGLR1.1 (At3g04110)、AtGLR1.2 (At5g48400)、AtGLR1.3 (At5g48410)、AtGLR1.4 (At3g07520)、AtGLR2.1 (At5g27100)、AtGLR2.2 (At2g24720)、AtGLR2.3 (At2g24710)、AtGLR2.4 (At4g31710)、AtGLR2.5 (At5g11210)、AtGLR2.6 (At5g11180)、AtGLR2.7 (At2g29120)、AtGLR2.8 (At2g29110)、AtGLR2.9 (At2g29100)、AtGLR3.1 (At2g17260)、AtGLR3.2 (At4g35290)、AtGLR3.3 (At1g42540)、AtGLR3.4 (At1g05200)、AtGLR3.5 (At2g32390)、AtGLR3.6 (At3g51480)和AtGLR3.7 (At2g32400), HsGLuR1 (NP_000818.1)、HsGLuR2 (NP_786944.1)、HsGLuR3 (NP_015564.1)、HsGLuR4 (NP_000820.1)、HsGLuR5 (NP_000821.1)、 HsGLuR6 (CAC67487.1)和HsGLuR7 (NP_000822)以及SspGLuR0 (BAA17851.1)。

Figure 1 Phylogenetic tree of glutamate receptor homologs from Arabidopsis, human beings and bacteriumThe amino acid sequence alignment was performed using Clustal W (from Lasergene DNA*) and the phylogenetic tree was generated by parsimony analysis using PAUP* 4.10b software. At: Arabidopsis thaliana; Hs: Homo sapiens; Ssp, Synechocystis sp. (strain) PCC6803. AtGLR protein sequences were extracted from Tair, HsGLuR1-7 and SspGLuR protein sequences were obtained from NCBI database. GenBank accession number or gene models for obtaining putative full-length protein sequences are described as below: AtGLR1.1 (At3g04110), AtGLR1.2 (At5g48400), AtGLR1.3 (At5g48410), AtGLR1.4 (At3g07520), AtGLR2.1 (At5g27100), AtGLR2.2 (At2g24720), AtGLR2.3 (At2g24710), AtGLR2.4 (At4g31710), AtGLR2.5 (At5g11210), AtGLR2.6 (At5g11180), AtGLR2.7 (At2g29120), AtGLR2.8 (At2g29110), AtGLR2.9 (At2g29100), AtGLR3.1 (At2g17260), AtGLR3.2 (At4g35290), AtGLR3.3 (At1g42540), AtGLR3.4 (At1g05200), AtGLR3.5 (At2g32390), AtGLR3.6 (At3g51480), AtGLR3.7 (At2g32400); HsGLuR1 (NP_000818.1), HsGLuR2 (NP_786944.1), HsGLuR3 (NP_015564.1), HsGLuR4 (NP_000820.1), HsGLuR5 (NP_000821.1), HsGLuR6 (CAC67487.1), HsGLuR7 (NP_000822); SspGLuR0 (BAA17851.1).

图2 拟南芥谷氨酸受体的拓扑学结构(以GLR3.5为例)AtGLR3.5序列取自Aramemnon (http://aramemnon.uni-koeln.de/index.ep)。蛋白拓扑学结构(或疏水性)的预测使用http://www.cbs. dtu.dk/services/TMHMM/。AtGLR3.5包含3个跨膜域(M1、M3和M4)、1个嵌入脂质膜内的结构域M2及2个预测的位于胞质外侧的配体结合区S1和S2, 其碳端位于胞质内。图中的数字表示蛋白质肽链的氨基酸残基位置数。

Figure 2 Putative topology of a representative glutamate receptor (GLR3.5) from ArabidopsisThe protein sequence of AtGLR3.5 was extracted from Aramemnon (http://aramemnon.uni-koeln.de/index.ep). Topological (or hydrophobicity) prediction of protein was performed using a program from a web server http://www.cbs.dtu.dk/services/TMHMM/. AtGLR3.5 is predicted to exhibit 3 trans-membrane domains (M1, M3, and M4), 1 domain (M2) imbedded in a lipid-layer, 2 domains S1 and S2 locating outside cytoplasm for ligand binding, and a C-terminal region towards the cytosol. Numbers indicate positions of amino acid residues in a peptide sequence.

表1 拟南芥谷氨酸受体AtGLRs表达的时空特异性

Table 1 Overview of glutamate receptors-like receptors (AtGLRs) expression pattern in Arabidopsis thaliana

基因名称及登录号	家族	苗龄	表达位点			参考文献
基因名称及登录号	家族	苗龄	器官水平	组织水平	亚细胞水平	参考文献
AtGLR1.1 At3g04110	I	1周	根、茎、叶	根部皮层, 托叶着生部	NR	Chiu et al., 2002; Roy et al., 2008
		4周	根、茎、叶	全展叶叶沿; 根系除根尖外各组织	NR
		8周	主要是根、叶; 花、角果中很微弱	花、角果中检测不到GUS信号	NR
AtGLR1.2 At5g48400		4周后	根、茎、叶、花、角果; 主要是根与角果	花粉管	NR
AtGLR1.3 At5g48410		4周	根、茎、叶, 叶中很微弱	NR	NR
AtGLR1.3 At5g48410		8周	叶和花中微弱; 主要是根与角果	NR	NR
AtGLR1.4 At3g07520		4周后	根、茎、叶、花、角果	NR	NR
AtGLR2.1 At5g27100	II	1周后	根、茎、叶、叶柄	根系除根尖外各组织; 托叶着生部位	NR
AtGLR2.1 At5g27100		4周后	根	根系所有组织; 花药和胚珠检测到微弱且转瞬即逝的GUS信号	NR
AtGLR2.2 At2g24720		4周后	根	NR	NR
AtGLR2.3 At2g24710		4周后	根	NR	NR
AtGLR2.4 At4g31710		4周后	根、角果	NR	NR
AtGLR2.5 At5g11210		4周后	根、茎、叶、花、角果	NR	NR
AtGLR2.6 At5g11180		4周后	根	NR	NR
AtGLR2.7 At2g29120		4周后	叶、角果, 根中微弱	NR	NR
AtGLR2.8 At2g29110		4周后	根、叶柄、叶、角果	NR	NR	Chiu et al., 2002 Roy et al., 2008
AtGLR2.9 At2g29100		4周	根、叶、茎、叶柄	NR	NR
AtGLR2.9 At2g29100		8周	根	NR	NR
AtGLR3.1 At2g17260	III	5天后	根、茎、叶、叶柄	所有维管组织; 保卫细胞	NR	Cho et al., 2009 Kim et al., 2001; Kong et al., 2015; Meyerhoff et al., 2005 Turano et al., 2002 Teardo et al., 2011 Teardo et al., 2015
AtGLR3.1 At2g17260		8周	根、茎、叶、花、角果	花丝和花药连接部位强烈表达	NR
AtGLR3.2 At4g35290		7天后	根、茎、叶柄、叶	所有维管组织及其邻近的导管, 随着植物生长表达更强烈; 根系韧皮部细胞中表达量远高于邻近细胞, 原生韧皮部更为强烈	NR
AtGLR3.2 At4g35290		8周	根、茎、叶、花、角果, 主要是叶与角果	花、茎维管组织; 花芽; 胚珠, 尤其是外皮层和生长中心	NR
AtGLR3.3 At1g42540		4周后	根、茎、叶柄、叶、角果		NR
AtGLR3.4 At1g05200		4周后	根、茎、叶、花、角果, 莲座叶中最强	叶肉组织; 维管束; 保卫细胞, 排水孔; 根内外皮层, 根毛	细胞质膜, 叶绿体内膜
AtGLR3.5 At2g32390		0	种子发芽时强烈, 随发芽完成降低	胚胎与子叶中强烈表达	线粒体内膜; 叶绿体膜
AtGLR3.5 At2g32390		4周后	根、茎、叶、花、角果	NR	线粒体内膜; 叶绿体膜
AtGLR3.7 At2g32400		整个生长阶段	所有器官	所有组织	细胞质膜

表1 拟南芥谷氨酸受体AtGLRs表达的时空特异性

Table 1 Overview of glutamate receptors-like receptors (AtGLRs) expression pattern in Arabidopsis thaliana

基因名称及登录号	家族	苗龄	表达位点			参考文献
基因名称及登录号	家族	苗龄	器官水平	组织水平	亚细胞水平	参考文献
AtGLR1.1 At3g04110	I	1周	根、茎、叶	根部皮层, 托叶着生部	NR	Chiu et al., 2002; Roy et al., 2008
		4周	根、茎、叶	全展叶叶沿; 根系除根尖外各组织	NR
		8周	主要是根、叶; 花、角果中很微弱	花、角果中检测不到GUS信号	NR
AtGLR1.2 At5g48400		4周后	根、茎、叶、花、角果; 主要是根与角果	花粉管	NR
AtGLR1.3 At5g48410		4周	根、茎、叶, 叶中很微弱	NR	NR
AtGLR1.3 At5g48410		8周	叶和花中微弱; 主要是根与角果	NR	NR
AtGLR1.4 At3g07520		4周后	根、茎、叶、花、角果	NR	NR
AtGLR2.1 At5g27100	II	1周后	根、茎、叶、叶柄	根系除根尖外各组织; 托叶着生部位	NR
AtGLR2.1 At5g27100		4周后	根	根系所有组织; 花药和胚珠检测到微弱且转瞬即逝的GUS信号	NR
AtGLR2.2 At2g24720		4周后	根	NR	NR
AtGLR2.3 At2g24710		4周后	根	NR	NR
AtGLR2.4 At4g31710		4周后	根、角果	NR	NR
AtGLR2.5 At5g11210		4周后	根、茎、叶、花、角果	NR	NR
AtGLR2.6 At5g11180		4周后	根	NR	NR
AtGLR2.7 At2g29120		4周后	叶、角果, 根中微弱	NR	NR
AtGLR2.8 At2g29110		4周后	根、叶柄、叶、角果	NR	NR	Chiu et al., 2002 Roy et al., 2008
AtGLR2.9 At2g29100		4周	根、叶、茎、叶柄	NR	NR
AtGLR2.9 At2g29100		8周	根	NR	NR
AtGLR3.1 At2g17260	III	5天后	根、茎、叶、叶柄	所有维管组织; 保卫细胞	NR	Cho et al., 2009 Kim et al., 2001; Kong et al., 2015; Meyerhoff et al., 2005 Turano et al., 2002 Teardo et al., 2011 Teardo et al., 2015
AtGLR3.1 At2g17260		8周	根、茎、叶、花、角果	花丝和花药连接部位强烈表达	NR
AtGLR3.2 At4g35290		7天后	根、茎、叶柄、叶	所有维管组织及其邻近的导管, 随着植物生长表达更强烈; 根系韧皮部细胞中表达量远高于邻近细胞, 原生韧皮部更为强烈	NR
AtGLR3.2 At4g35290		8周	根、茎、叶、花、角果, 主要是叶与角果	花、茎维管组织; 花芽; 胚珠, 尤其是外皮层和生长中心	NR
AtGLR3.3 At1g42540		4周后	根、茎、叶柄、叶、角果		NR
AtGLR3.4 At1g05200		4周后	根、茎、叶、花、角果, 莲座叶中最强	叶肉组织; 维管束; 保卫细胞, 排水孔; 根内外皮层, 根毛	细胞质膜, 叶绿体内膜
AtGLR3.5 At2g32390		0	种子发芽时强烈, 随发芽完成降低	胚胎与子叶中强烈表达	线粒体内膜; 叶绿体膜
AtGLR3.5 At2g32390		4周后	根、茎、叶、花、角果	NR	线粒体内膜; 叶绿体膜
AtGLR3.7 At2g32400		整个生长阶段	所有器官	所有组织	细胞质膜

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植物谷氨酸受体的研究进展

Current Research Advances on Glutamate Receptors (GLRs) in Plants

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