植物乙烯信号转导研究进展

过去10年,对模式植物拟南芥的分子遗传学研究建立了植物乙烯信号转导线性模型.乙烯结合到受体上,经一条MAPK级联反应和转录级联途径将信号转导而产生乙烯反应.拟南芥乙烯受体家族由5个成员构成,ETR1、ERS1、ETR2、ERS2和EIN4.乙烯受体包括三个结构域:乙烯结合结构域、组氨酸激酶结构域和反应调控结构域.乙烯受体定位于内质网,与CTR1协同负调控乙烯反应.ENI2、EIN3/EIL、ERF1依次位于CTR1下游,正调控乙烯反应.EIN3属于转录激活因子调控蛋白家族,受转录后调控.乙烯稳定EIN3结构,EBF1/EBF2促进EIN3分解.ERF1是转录调控因子家族成员之一,是EIN3/EIL的直接作用目标.     

乙烯信号转导的分子机制

气态植物激素乙烯在植物生长发育和应对生物及非生物胁迫过程中起着重要作用。在过去的十几年中, 对模式植物拟南芥的分子遗传研究已建立从信号感知到转录调控的乙烯信号转导线性模型。拟南芥共有5个乙烯受体ETR1、ERS1、ETR2、ERS2和EIN4, 目前已知ETR1定位在内质网上, 与类似于Raf的蛋白激酶CTR1协同负调控乙烯反应。EIN2和EIN3/EILs位于CTR1下游, 正调控乙烯反应。两个F-box蛋白EBF1和EBF2通过泛素/26S蛋白体降解途径调控EIN3的稳定性。5’→3’的外切核酸酶EIN5通过启动EBF1和EBF2 mRNA的降解, 拮抗EBF1和EBF2对EIN3的负反馈调控。目前对于乙烯信号转导途径关键组分的生化功能和乙烯下游反应途径的了解甚少, 乙烯信号转导途径与其它途径之间还存在着广泛的交叉反应, 这些问题的解决将大大增加我们对乙烯信号转导途径的了解。     

植物激素乙烯作用机制的最新进展

气体植物激素乙烯在植物生长发育及应对胁迫的防御反应中起重要调控作用．通过20多年的研究,利用模式植物拟南芥,勾画出一条自内质网膜受体至细胞核内转录因子的线性乙烯信号转导通路．本文概述了研究乙烯信号转导的方法及乙烯信号转导的基本过程;阐述了最新发现的乙烯信号从内质网膜传递到细胞核的分子机制,即原本定位于内质网膜上的EIN2蛋白其C端被剪切之后进入细胞核,然后通过抑制EBF1／2而稳定转录因子EIN3／EIL1;根据最近多个小组报道EIN3／EIL1直接调控除乙烯响应基因之外的其他生物学过程相关基因,提出了EIN3／EIL1可以作为网络节点整合多条信号通路的新观点;通过分析不同信号通路调控EIN3／EIL1的方式,发现不仅EIN3／EIL1的蛋白稳定性受到调控,而且其转录活性还受到诸如JAZ,DELLA等转录调节因子的调控．本文展望了未来乙烯信号转导通路的研究方向与研究热点．     

拟南芥乙烯信号转导机理的遗传学和化学生物学研究

乙烯信号途径的建立得益于一系列的突变体研究,EIN3是乙烯信号转导通路的核心转录因子,EIN3的蛋白质含量严格受F-BOX蛋白EBF1/EBF2的降解调控。为了进一步挖掘乙烯信号途径的新组分和深入研究EIN3及其下游的信号组分,作者筛选了四个不同来源的T-DNA库,并利用转基因植物EIN3ox作为遗传背景,进行了EIN3下游的抑制子筛选工作,还利用化学遗传学的方法筛选了四个小分子库。     

乙烯的生物合成与信号传递

乙烯是气体植物激素, 它在植物的生长发育过程中有很多作用。所以了解乙烯的生物合成及其信号转导是非常重要的。二十年来, 通过筛选有异于正常三重反应的突变体, 人们发现了乙烯信号转导的粗略轮廓。在拟南芥中, 有5个受体蛋白感受乙烯, ETR1、ERS1、ETR2、ERS2、EIN4。它们表现出功能冗余, 是乙烯信号的负调控因子, 在植物体内以二聚体的形式存在。ETR1的N端与乙烯结合时需要 铜离子(Ⅰ)的参与。尽管已经发现ETR1有组氨酸激酶活性, 而其它受体有丝氨酸/苏氨酸激酶活性, 但受体参与乙烯信号转导的机制还不是很清楚。受体与Raf类蛋白激酶CTR1相互作用, CTR1是乙烯反应的负调控因子。CTR1蛋白失活使EIN2蛋白活化。EIN2的N端是跨膜结构域, 与Nramp家族金属离子转运蛋白的跨膜结构域类似。EIN2的C端是一个新的未知结构域, 与乙烯信号途径的下游组分相互作用。EIN3位于EIN2的下游, EIN3和EILs诱导ERF1和其它转录因子的表达, 这些转录因子依次激活乙烯反应目的基因的表达, 表现出乙烯的反应。EIN3受到蛋白酶体介导的蛋白降解途径的调节。由于乙烯是一种多功能的植物激素, 其信号途径与其它信号途径有多重的交叉。     

Ethylene signaling is critical for synergid cell functional specification and pollen tube attraction

ETHYLENE INSENSITIVE 3 (EIN3) is a key regulator of ethylene signaling, and EIN3‐BINDING F‐BOX1 (EBF1) and EBF2 are responsible for EIN3 degradation. Previous reports have shown that the ebf1 ebf2 double homozygous mutant cannot be identified. In this study, the genetic analysis revealed that the ebf1 ebf2 female gametophyte is defective. The pollination experiment showed that ebf1 ebf2 ovules failed to attract pollen tubes. In female gametophyte/ovule, the synergid cell is responsible for pollen tube attraction. Observation of the pEIN3::EIN3‐GFP transgenic lines showed that EIN3 signal was over‐accumulated at the micropylar end of ebf1 ebf2 female gametophyte. The overexpression of stabilized EIN3 in synergid cell led to the defect of pollen tube guidance. These results suggested that the over‐accumulated EIN3 in ebf1 ebf2 synergid cell blocks its pollen tube attraction which leads to the failure of ebf1 ebf2 homozygous plant. We identified that EIN3 directly activated the expression of a sugar transporter, SENESCENCE‐ASSOCIATED GENE29 (SAG29/SWEET15). Overexpression of SAG29 in synergid cells blocked pollen tube attraction, suggesting that SAG29 might play a role in ethylene signaling to repel pollen tube entry. Taken together, our study reveals that strict control of ethylene signaling is critical for the synergid cell function during plant reproduction.     

EIN2 regulates salt stress response and interacts with a MA3 domain-containing protein ECIP1 in Arabidopsis

Ethylene signalling regulates plant growth and development. However, its roles in salt stress response are less known. Here we studied functions of EIN2, a central membrane protein of ethylene signalling, and its interacting protein ECIP1 in salt stress responses. Mutation of EIN2 led to extreme salt sensitivity as revealed by phenotypic and physiological changes, and overexpression of C-terminus of EIN2 suppressed salt sensitivity in ein2-5, indicating that EIN2 is required for salt tolerance. Downstream components EIN3 and EIL1 are also essential for salt tolerance because ein3-1eil1-1 double mutant showed extreme salt-sensitive phenotype. A MA3 domain-containing protein ECIP1 was further identified to interact with EIN2 in yeast two-hybrid assay and GST pull-down assay. Loss-of-function of ECIP1 resulted in enhanced ethylene response but altered salt response during seed germination and plant growth. Double mutant analysis revealed that ein2-1 was epistatic to ecip1, and ecip1 mutation partially suppressed ethylene-insensitivity of etr2-1 and ein4-1. These studies strengthen that interactions between ECIP1 and EIN2 or ethylene receptors regulate ethylene response and stress response.     

水稻体内的乙烯信号传导途径(综述)

迄今为止,水稻中已经鉴定的有关乙烯信号传导途径中的组分包括乙烯受体、EIN2和EIN3的同系物,CTR1、RTE1、EBF1/2和EIN5的同系物,这些组分在双子叶植物拟南芥和单子叶植物水稻中相对保守。然而,对水稻ein2和eil1突变体的研究发现,两突变体与野生型相比并没有明显的表型差异。由此可以推断,水稻中的乙烯信号可能比拟南芥中的更加复杂。水稻依靠乙烯调节生长发育的许多方面（如对低氧环境的适应）,这些在拟南芥中是不存在的,这表明水稻可能在乙烯信号传导途径中存在新的组分或新的机制。文章就水稻体内乙烯信号传导途径、乙烯信号调节以及乙烯在水稻中的应答进行综述。     

A novel tomato F‐box protein,SlEBF3, is involved in tuning ethylene signaling during plant development and climacteric fruit ripening

Ethylene is instrumental to climacteric fruit ripening and EIN3 BINDING F‐BOX (EBF) proteins have been assigned a central role in mediating ethylene responses by regulating EIN3/EIL degradation in Arabidopsis. However, the role and mode of action of tomato EBFs in ethylene‐dependent processes like fruit ripening remains unclear. Two novel EBF genes, SlEBF3 and SlEBF4, were identified in the tomato genome, and SlEBF3 displayed a ripening‐associated expression pattern suggesting its potential involvement in controlling ethylene response during fruit ripening. SlEBF3 downregulated tomato lines failed to show obvious ripening‐related phenotypes likely due to functional redundancy among SlEBF family members. By contrast, SlEBF3 overexpression lines exhibited pleiotropic ethylene‐related alterations, including inhibition of fruit ripening, attenuated triple‐response and delayed petal abscission. Yeast‐two‐hybrid system and bimolecular fluorescence complementation approaches indicated that SlEBF3 interacts with all known tomato SlEIL proteins and, consistently, total SlEIL protein levels were decreased in SlEBF3 overexpression fruits, supporting the idea that the reduced ethylene sensitivity and defects in fruit ripening are due to the SlEBF3‐mediated degradation of EIL proteins. Moreover, SlEBF3 expression is regulated by EIL1 via a feedback loop, which supposes its role in tuning ethylene signaling and responses. Overall, the study reveals the role of a novel EBF tomato gene in climacteric ripening, thus providing a new target for modulating fleshy fruit ripening.