植物类受体蛋白激酶的研究进展

西北植物学报,2009,29(4) :0851-0858　　文章编号:100024025(2009) 0420851208

植物类受体蛋白激酶的研究进展

闫　锋1, 祝传书2, 庞保平13

(1内蒙古农业大学农学院, 呼和浩特010019;2西北农林科技大学, 陕西杨陵712100)

摘　要:植物类受体蛋白激酶(receptor 2like protein kinase ,RL Ks ) , 发生磷酸化、去磷酸化反应而开启或关闭下游靶蛋白, 调节植物固有免疫反应, 诱导抗病防御反应。酶的功能、。酶的结构、, 生理生化功能及应用提供参考。

关键词:植物; 类受体蛋白激酶; 中图分类号:Q942.6

in R esearch of Plant R eceptor 2like Protein Kinases

YAN Feng 1,ZHU Chuan 2shu 2,PAN G Bao 2ping 13

(1College of Agriculture , Inner Mongolia Agricultural University , Huhhot 010019, China ; 2Nort hwest Sci 2Tech University of Agriculture and Forestry , Yangling ,Shaanxi 712100,China )

Abstract :Signaling molecules are recognized t hrough ext racellular ligand 2binding domain of receptors in liv 2ing plant s. Pho sp horylation and dep ho sp horylation of kinases is a common mechanism for positively or neg 2atively regulating kinase 2mediated signaling involved in plant innate immunity ,and t hen stimulates resist 2ance to pat hogens. To date ,t he research of t he f unction of t he plant receptor 2like protein kinases (RL Ks ) , mechanisms of signaling pat hways and ligand 2binding have already been t he importance in t his field. In t his review ,t he f unction and regulation of RL Ks in resistance to pat hogens of plant s was presented in order to offer t he references to t he p hysiological and biochemical f unctio n of t he RL Ks. K ey w ords :plant ;receptor 2like protein kinase ;signal t ransduction ;resistance

　　植物细胞具有感受生物和非生物因子刺激的能力, 并能通过细胞膜内外信号传导, 最终调整其转录组与蛋白质组, 从而对胞外信号做出恰当的反应。类受体蛋白激酶(receptor 2like p rotein kinase , RL Ks ) 属于蛋白激酶的一个亚家族[1], 胞内激酶区通过磷酸化或去磷酸化, 开启或关闭下游靶蛋白, 将胞外信号转换为胞质信号。

1989年Lawton 通过分析动物蛋白的丝氨酸/

苏氨酸激酶保守域, 首次从四季豆(Phaseol us v ul 2

[2]

garis L. ) 中成功克隆到植物蛋白激酶PV P K 21, 随后Ferreira [3]等在植物中也发现了一个相似蛋白P34cdc2。Walker 等[4]第一次成功克隆到玉米的类受体蛋白激酶ZmP K1。这些研究使得植物体内的类受体蛋白激酶开始受到关注。通过预测分析, 拟南芥基因组中包含约600多个类受体蛋白激酶[5], 作为单子叶模式植物的水稻基因组中RL Ks 数量大

3收稿日期:2009201205; 修改稿收到日期:2009203225

基金项目:国家自然科学基金(30460076) 作者简介:闫　锋(1980-) , 男(汉族) , 在读博士研究生, 主要从事化学生态与分子生态学研究。E 2mail :yffwll@163. com

3通讯作者:庞保平, 教授, 博士生导师, 主要从事昆虫生态学及害虫综合治理的教学和研究工作。E 2mail :pangbp@imau. edu. cn

852西　北　植　物　学　报　　　　　　　　　　　　　　　　　　　29卷

约是拟南芥的2倍[6]。

虽然植物拥有如此众多类受体蛋白激酶, 但目前对类受体蛋白激酶的功能、信号传导和配体识别等方面了解甚少。本文就关于植物类受体蛋白激酶的结构、功能及其在抗病防御反应中的作用研究进展做一简要综述。

外信号作用, 是信号识别中心[8]。②胞外受体结构

域(ext racellular ligand 2binding domain , ECLB ) , 一般胞外受体结构域部分差异比较大(如图1中L RR 、PR5K 、EGF 、B 2Lectin 、S 、TN FR 结构域) , 此结构域与信号分子(配体) 结合, 促使受体形成二聚体激活胞内激酶域的级联反应[9], ECLB 主要是开启信号的传递, 这在单子叶植物和双子叶植物中得到证实[6]。因此, 该区域被认为是信号结合中心[8,9]。③跨膜结构域(t ransmembrane domain , TM ) , 一般由22～28, 该结构域将胞外和胞内部分联系到一起[]白酶接触反应域(,P KC ) [10213], 该区域位) , 植物的蛋白激酶区由丝/, 该区域的一个重要特征是带有磷酸化位点。激酶域的可逆磷酸化反应受蛋白激酶和蛋白磷酸酯酶调控, 也是信号级联反应中心, 其中有的在激酶域有一个羧基C 端, 且一般是由0～70个氨基酸构成[8,13]

。

1　植物体类受体蛋白激酶的结构及分类

1. 1　植物类受体蛋白激酶的结构

植物RL Ks 具有高度保守的氨基酸序列, 普遍具有一种二叶状(bilobal ) 结构[7]。另外, 植物细胞表面的受体都定位在质膜上[8], 由于其受体含有的结构域不同, 不同类型的RL Ks 在细胞信号传导中作用也各异[8]。大多数植物RL Ks 结构域(图1) [8,9]:①N 端信号肽(2peptide ,SP ) , 的RL Ks 激酶除外) , RL , 在蛋白质合成过程中, 可锚定蛋白质于内质网上, 起识别胞

图1　植物类受体蛋白激酶(RL Ks ) 结构示意图

Ser/Thr. 丝氨酸/苏氨酸激酶域;J M. 近膜区;SP. 信号肽;B 2Lectin. 球形甘露糖结合域;S. S 结构域; TN FR. 肿瘤坏死因子受体; PAN. 调节蛋白2蛋白互作或蛋白2醣互作结构域; EGF. 表皮生长类因子; PR5K. 病程相关5蛋白受体激酶;L RR. 富含亮氨酸重复序列;

TM. 跨膜域;Unkown. 未知。FL S2(At5g46330. 1) [14,15]、Xa21(U37133) [16]和Xa26(A Y364476) [17]都属于

富含亮氨酸重复序列型类受体蛋白激酶; Pr5k (U48695) [18]属于PR5型类受体蛋白激酶;Wak1(AJ 009696) [19]属于表皮生长因子型类受体蛋白激酶、Pi 2d2(AP005967) [20]属于外源类凝集素型类受体蛋白激酶、Cr4(U67422) [21]属于类肿瘤坏死因子型受体蛋白激酶;SR K6(M76647) [22]和Zmpk1(X67733) [4]具有S 结构域。比例尺是以氨基酸个数为单位(个)

Fig. 1　Plant receptor 2like protein kinases (RL Ks ) structure

Ser/Thr. Serine/t hreonine kinase domain ;J M. J uxtamembrane ;SP. Signal peptides ;B 2Lectin. Bulb 2type mannose 2specific binding lectin domain ;S. S 2Locus ; TN FR. Tumor necrosis factor receptor 2like ; PAN. A domain for mediating protein 2protein or protein 2carbohydrate

interactions ; EGF. Epidermal growt h factor 2like repeat s ; PR5K. Pat hogenesis 2related protein 52like receptor kinase ;

L RR. Leucine 2rich repeat s ; TM. Transmembrane domain ; FL S2(At5g46330. 1) [14,15],Xa21(U37133) [16]and Xa26(A Y364476) [17]contain L RR domain ; Pr5k (U48695) [18],Wak1(AJ 009696) [19],Pi 2d2(AP005967) [20], Cr4(U67422) [21]encode a PR5K 2RL Ks ,EGF 2RL Ks ,B 2Lectin 2RL Ks , TN FR 2RL Ks ,respectively ;SR K6(M76647) [22]and Zmpk1(X67733) [4]contain S 2Locus domain. A particular scale is determined according to number of specific amino acids

4期　　　　　　　　　　　　　　　　闫　锋, 等:植物类受体蛋白激酶的研究进展853

1. 2　植物类受体蛋白激酶的分类

植物RL Ks 的研究是建立在与哺乳动物受体蛋白激酶比较的基础上获得发展的。人们将植物RL K 胞外域及其基因表达模式较相似的归为一类[8]。因此, 前人根据植物RL Ks 胞外结构域不同[7,23225], 可将RL Ks 分为以下6个亚家族(图1) :①富含亮氨酸重复序列(leucine 2rich repeat s , L RRs ) 型[26230],L RR 家族占居整个家族成员一半以上, 它的胞内激酶结构域相对保守, 氨基酸同源性达45%以上, 通过序列比较分析, 胞外受体蛋白序列都含有22～24个富含亮氨酸的氨基酸序列, 其保守核心含有7个氨基酸残基:XXL XL XX (其中X 代表任意氨基酸残基) [26,31]。②PR5型(pat hogenesis 2re 2lated p rotein 52like receptor kinase ,PR5K ) , 胞外域氨基酸序列与pat hogenesis 2相似, 因此称为PR5K [18]]③(epidermal growt h factor 2like repeat s , EGF ) 型[19,33,34], 其内含12个保守的半胱氨酸残基, 与其它类型的RL Ks 不同的是, 其胞外的N 2端不含信号肽序列(见图1Wak1) [19]。它的胞外域与细胞壁成分直接相连, 能介导细胞壁与质膜互作, 该类基因与植物抗病防御反应等有关[33,35]。④外源类凝集素结合域(Lectin 2binding domain ,LB ) 型[20,25,36,37], 目前已知其胞外由2个区域组成(图1Pi 2d2) , 一个区域具有球形结构, 它是蛋白与甘露糖互作或与胞外配体结合的区域[20,38]; 另一个已知区域是PAN 结构域, 它是与蛋白或碳水化合物结合的区域[20]。该基因被推测可能与果胶结合, 并发现其激酶活性能被果胶寡聚物激活[39]。拟南芥中至少有30多个基因属于此类型。⑤类肿瘤坏死因子受体(t umor necrosis factor receptor 2like , TN FR ) 型[21,40], 胞外由一个富含半胱氨酸序列组成, 该类基因与生长发育有关[21,40]。⑥S 2结构域(S 2do main ,S ) 型[4,22], 具有12个保守的富含半胱氨酸残基, 这类基因多数与生长发育、自交不亲和等有关[41,42]。

活RL Ks 胞内激酶域发生磷酸化反应, 将胞外信号转导到胞内, 诱导防御反应, 保护植物免受病原菌的进一步侵害[43]。发生磷酸化的RL Ks 为了保持植物体内一个正常的磷酸化水平还要发生去磷酸化反应[44]。细胞中蛋白质可逆磷酸化是调节酶活性的两个主要翻译后反应, 是细胞内普遍存在的一种调节机制, 可逆磷酸化反应几乎与所有的生理和病理过程息息相关[44246]。RL Ks 可逆磷酸化在植物的防御机制、信号转导中起着极其重要的作用。

2. 1　类受体蛋白激酶的磷酸化反应和抗病防御反应

的λ位磷酸基, 它是植物[47]。研究表明, 拟南芥BRI1(L RR 2RL K ) 突变体能识别云苔素类固醇(Brassinosteroid ,BR ) 并与之结合, 促使胞内BRI1与其特异性受体激酶BA K1(Associated Receptor K inase 1) 结合形成二聚体, 最后导致激酶域的磷酸

化残基(丝氨酸/苏氨酸) 发生自我磷酸化反应[48,49]。Horn 等研究发现, 拟南芥RL K5(L RR 2RL K ) 的自我磷酸化可对自身活性的催化以及与其它蛋白质的互作起重要调控作用[50]。

尽管不同RL Ks 编码的蛋白各不相同, 而且抗性也不同, 但它们在抗病防御反应中都会经历以下3个过程:首先, 病原信号分子(配体) 与RL Ks 胞外结构域识别并结合; 然后, 胞内激酶域发生磷酸化等反应将信号传递到下游靶基因; 最后激活下游靶蛋白表达使植株产生抗性[8,51]。研究表明, 以单体形式存在的RL Ks 不具活性, 被磷酸化的RL Ks 形成复合体后才能调控下游靶标[52,53]。

Song 等[16]研究表明, Xa21在抗水稻白叶枯病(X ant homonas ory z ae pv ory z ae ) 中起重要作用。Xa21胞外域含有21个L RR 基序(图1) , 它参与在细胞表面识别病原物激发子AvrXA21(无毒基因AvrXA21编码) , 二者结合后能形成一个稳定的二聚体结构, 促使胞内Ser/Thr 激酶区部分相互靠拢并激活胞内激酶域发生磷酸化反应, 引起RL Ks 构象发生变化[16,54,55]。被活化的二聚体在XB3(XA21Binding Protein 3) 和WR KYs 转录因子的作用下开启下游防御基因, 使植株对病原物表现出抗性[51,52,56,57]。据研究报道, 拟南芥FL S2在抗细菌性叶斑病中起重要作用, 而且FL S2的胞外结构域与水稻Xa21基因具有相似的胞外域, 胞外也具有

2　植物类受体蛋白激酶在抗病防御反

应中的作用

　　植物体在生长发育过程中能不断感受生物和非生物因子的刺激, 引发一系列信号转导途径, 并做出相应的反应。植物在进行抗病反应时, 当病原物侵染植物后, 植物RL Ks 能识别和接受病原信号物, 激

854西　北　植　物　学　报　　　　　　　　　　　　　　　　　　　29卷

L RR 结构域(图1) 的模式识别受体(Pattern 2recog 2

nitio n receptors ,PRRs ) [58,59]。FL S2能调控拟南芥

产生免疫反应, 诱导抗病防御反应[60]。当病原信号分子flg22(激发子) 与FL S2的胞外域结合后, 引发激酶域发生磷酸化反应, 将信号传递到胞内, 经MA P K 级联反应将信号进一步放大, 在WR KY22

接杀死作用[57,70272]。同时凋亡的细胞会释放出一种新的信号分子快速传遍整个植株, 使未感病邻近细胞内的水杨酸(SA ) 大量积累[67,74]。研究表明, 利用外源激素SA 处理烟草可诱导其产生SAR [75]。Verberne 等[76]通过提高植物体内SA 的方法, 增强植物的SAR 能力, 使植物的整体抗病水平提高。可见SA 是导致植株产生SA R 的主要因素[77]。目前关于A v r 与RL Ks 的对应关系还不清楚, 有待进一步研究。2. 2　, 对其生的活性是保持正常磷酸的磷酸酯酶(p rotein ho sp ) [51,78,79]。根据PPs 作用的底物不同可将其分为4类[44]:第1类, 酪氨酸磷酸酯酶(Tyr 2PPs ) ; 第2类, 丝氨酸/苏氨酸磷酸酯酶(Ser/Thr 2PPs ) ; 第3类, 丝氨酸/苏氨酸2酪氨酸磷酸酯酶(Ser/Thr 2Tyr 2PPs ) ; 第4类, 依赖金属离子蛋白磷酸酯酶(metalion 2dependent protein p hosp hatase , PPM ) 。

最近研究表明, 水稻在调节自身固有免疫反应中,XB15(XA21Binding p rotein 15) 在保持正常磷酸化水平中起了极其重要的作用[51]。XB15

编码的

和WR KY29转录因子作用下激活防卫基因诱导植

株产生抗病防御反应[14,51,61](图2) 。

植物之所以表现出抗病防御反应特性, 是由于植物能通过胞外的受体识别病原信号分子(配体) , 诱导植物产生过敏反应(hypersensitive response , HR ) 和系统获得抗性(systemic acquired resist 2ance ,SAR ) [62](图2) 。RL Ks 在抗病防御反应途径中引发的一系列反应可通过配体2述[63]。无毒基因(avirulent v , HR , 使(p rogrammed cell deat h , PCD ) , 这是植物普遍具有的一种抗病反应[64269]。植物发生HR 的速度很快, 可有效地抑制病原菌不断蔓延, 在此过程中氧爆(oxygen burst ) 导致活性氧中间体(ractive oxygen species ,ROS ) 产

-生, 促使H 2O 2、超氧阴离子自由基(O 2・) 、羟基自由) 和NO 的浓度增加, 对入侵病原物有直基(HO ・

图2　RL Ks 抗病防御反应示意图

Fig. 2　The schematic representation of RL Ks resistance to pathogens and defense responses

4期　　　　　　　　　　　　　　　　闫　锋, 等:植物类受体蛋白激酶的研究进展855

是第4类PPs , 即PPM 2PP2c [51], 在Xa21胞内近膜区拥有一个能与XB15结合的区域(图2) , 当这2种蛋白发生互作时, 促使磷酸化的RL Ks 失去活性或活性降低, 导致下游靶蛋白不能表达或表达量下降, 从而有效地维持植株体内的磷酸化水平, 也就使植物的抗病防御反应受到抑制, 表明XB15在水稻固有免疫反应中起负调控作用[51]。

在FL S2的近膜区也有一个PPs 的结合区, 能被蛋白磷酸酯酶(kinase 2associated p rotein p ho s 2p hatase , KA PP ) 所识别[80](图2) 。KA PP 在拟南芥防御反应中起负调控作用, 与XB15在水稻防御反应中具有相类似的功能[80,81]。拟南芥FL S2具有识别病原激发子鞭毛蛋白(flagellin ) 的作用[6], 能启动拟南芥的抗病防御反应, KA PP 能有效阻碍FL 的信号转导, 使FL S2化, [79之, 植物RL Ks , 但其参, 仍需要继

续进行深入研究。

3　结语与展望

植物RL Ks 具有不同的胞外域结构, 其功能也各有不同之处。如含有L RR 、PR5K 、EGF 、TN FR 和Lectin 结构域的RL Ks , 它们在抗性防御反应中起重要作用, 尤其对L RR 2RL Ks 研究较多也较为深入, 这些功能已经在拟南芥、水稻、番茄等植物中得到证实。病原信号分子与植物RL Ks 的胞外结构域, 它是抗性防御反应中必不可少的环节, , (配体) 是如何? 植物RL Ks 在发生磷酸化反? 植物病原信号分子的受体结构如何? RL Ks 是否参与抗虫防御反应? RL Ks 是否调控植物挥发物的释放? 这些都是今后研究中急需解决的问题。

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