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陈悦 孙明哲 贾博为 冷月 孙晓丽*
黑龙江八一农垦大学 / 作物逆境分子生物学实验室
摘要 Abstract
AP2/ERF (APETALA2/ethylene responsive factor)是植物特有的转录因子家族,广泛参与生长发育和胁迫应答等多种生物学进程。水稻是我国重要的粮食作物,但在其生长过程中受到多种逆境的影响。目前研究发现AP2/ERF转录因子在水稻逆境应答中扮演十分重要的角色。本文结合国内外研究进展,综述了水稻AP2/ERF转录因子的分类和结构,并梳理了不同亚家族的AP2/ERF转录因子响应病害、干旱、高盐和低温胁迫的功能和机制研究进展,为深入阐释水稻AP2/ERF转录因子调控逆境应答的分子网络及品种改良提供参考。
水稻(Oryza sativa L.)是全球最重要的粮食作物之一,我国2/3的人口以水稻为食。水稻生产对保障国家粮食安全至关重要,但温度、水分、病虫害等胁迫严重威胁水稻高产稳产。近年来,在水稻逆境应答分子机制方面,国内外科学家进行了大量工作,并取得了一系列重要成果[1-3],初步建立了水稻参与逆境胁迫的分子调控网络,其中转录因子(transcription factors)处于调控网络的核心位置。
01
水稻AP2/ERF转录因子家族的分类与结构特征
1.1 水稻AP2/ERF转录因子家族的分类
图1 植物AP2/ERF转录因子的结构域[9]
Fig. 1 Conserved domains of AP2/ERF transcription factors in plants[9]
02
水稻AP2/ERF转录因子参与逆境应答的分子机制研究进展
水稻生长过程中会受到多种生物和非生物胁迫,导致大面积减产,严重威胁我国水稻高产稳产和粮食安全。近年来,植物学家对水稻响应外界逆境的分子机制和信号网络开展了大量研究工作。AP2/ERF转录因子,尤其是ERF和DREB亚家族,在水稻逆境应答信号传导通路中占据重要的位置。
正常生长条件下,AP2/ERF基因通常表达水平较低,当遭遇逆境胁迫后表达显著升高。AP2/ERF可结合GCC-box、DRE/CRT或GT-1 (GAAAAA)等元件,调节下游靶基因表达,最终引起生理生化变化,帮助抵抗外界逆境胁迫。接下来本文将综述水稻AP2/ERF家族参与不同逆境应答的研究进展(表3)。
DREB亚家族正调控水稻干旱胁迫应答过程。OsDREB1A、OsDREB1E和OsDREB1F受干旱胁迫诱导,与DRE/CRT元件结合[19,57-58]。在拟南芥中过表达OsDREB1A可诱导AtDREB1A靶基因的表达,增强了转基因株系耐旱性[59]。OsDREB1E、OsDREB1F、OsDREB1G过表达能增强转基因水稻耐旱性[57-58]。OsDREB2A、OsDREB2B和OsDREB2C基因表达均受干旱胁迫诱导[59]。Cui等[66]利用4ABRA (four copies 49-bp responsible sequence of ABA)启动子诱导OsDREB2A基因表达,增强了转基因水稻株系的耐旱性。在拟南芥中过表达OsDREB2B诱导AtDREB2A靶基因的表达,增强了转基因拟南芥株系耐旱性[64],暗示OsDREB1A/2B与AtDREB1A/2A可能具有相似作用。此外,Zhao等[67]发现干旱和ABA能诱导水稻DREB基因ARAG1 (ABA responsive AP2-like gene)的转录,敲除ARAG使水稻种子萌发和幼苗生长对ABA高度敏感。过表达OsAP21提高转基因拟南芥株系中RD29B 表达水平,暗示OsAP21可能在干旱胁迫应答中发挥作用[68]。综上,DREB亚家族通过与DRE/CRT元件结合,激活干旱响应基因表达,调控水稻对干旱胁迫的应答。
03
总结与展望
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