脱落酸调控油菜素甾醇初级信号通路的研究
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摘要
植物激素在共同调节植物生长发育和生理生化过程中发挥重要的作用,但目前对激素互作的分子机制了解甚少。
油菜素甾醇(brassinosteroids,BRs)是上世纪70年代发现的第六大植物激素,其在植物的很多发育过程中起着重要作用,包括茎和根中的细胞分裂和伸长、光形态建成、生殖发育、叶片衰老、以及对胁迫的反应等。脱落酸(abscisic acid,ABA)作为另一类作用广泛的植物激素,也调控着植物生长发育的诸多方面,包括参与植物的胚胎发育、种子休眠与萌发、气孔关闭、开花时间和果实成熟等生理过程以及植物对胁迫的适应性反应。已有研究表明,BRs和ABA共同调控数百个基因的表达,从而共同调节很多发育和生理过程。同时,已经发现BR的一些突变体具有对ABA反应改变的表型,两者在发芽率上表现出相互拮抗的生理效应。然而,对于两者互作是否通过修饰或介入到初级信号途径中,抑或通过平行或独立的途径起作用,却仍然未知。
本研究工作主要包括:首先利用BR相关的多种遗传材料,系统研究了BRs和ABA两类激素在调节种子发芽率上具有相互拮抗的作用,为研究信号互作奠定生理基础。接着利用分子(DWF4和CPD转录水平)和生化标记(BES1的磷酸化状态)并结合ABA合成缺失突变体,证明外源的ABA能快速抑制BR的信号输出。再者,研究发现BR受体BRI1的强突变体bri1-116中,ABA仍能调节BR的信号输出;且在BKI1-YFP过表达系中,BKI1-YFP的亚细胞定位也不受ABA的影响,已知油菜素内酯(BRs中最具活性的一种形式)能促使BKI1-YFP从膜上解离;而用LiCl处理使下游组分BIN2失活却抑制了ABA对BR信号输出的调节。最后,利用ABA早期信号突变体abi1和abi2,得出ABA调控BR的初级信号转导途径主要依赖于ABA信号组分ABI2,部分依赖于ABI1。因此我们提出了BRs与ABA信号互作的模型,ABA对BR初级信号转导途径的调节界定在受体BRI复合体的下游,BIN2(含BIN2)的上游。这一模型有力的解释了为什么受BR调节的基因有一大部分也受到了ABA的调节,同时为BRs和ABA信号的互作提供了新的分子机制。
本研究通过遗传、细胞和分子水平的系统深入研究,确定了ABA影响BR的初级信号输出,且界定了信号互作的区域,同时形成了ABA对BR信号输出影响的初步模型。本研究从分子层面有力的支持了BRs和ABA在生理实验中的拮抗效应,具有重要的理论意义。同时为研究激素互作提供了很好的范例,也为植物更好的适应环境提供理论依据和目标基因。
Phytohormones play essential roles in coordinately regulating a large array of developmental processes.Studies have revealed that brassinosteroids and abscisic acid interact to regulate hundreds of genes'expression,governing many biological processes.However,whether their interaction is through modification or intersection of their primary signaling cascades or by independent or parallel pathways remains a big mystery.
Using biochemical and molecular markers of brassinosteroid signaling and abscisic acid biosynthetic mutants,we demonstrated that exogenous abscisic acid rapidly inhibits brassinosteroid signaling outputs as indicated by the phosphorylation status of BES1 and brassinosteroid-responsive gene expression.Experiments using a bri1 null-allele,bri1-116,and analysis of subcellular localization of BKI1-YFP further revealed that the brassinosteroid receptor complex is not required for abscisic acid to act on brassinosteroid signaling outputs.However,when the BR downstream signaling component BIN2 is inhibited by LiCl,abscisic acid failed to inhibit brassinosteroid signaling outputs.Furthermore,using a set of abscisic acid insensitive mutants,we found that regulation of abscisic acid on brassinosteroid primary signaling pathway is dependent on the abscisic acid early signaling components ABI1 and ABI2.We propose that the signaling cascades of abscisic acid and brassinosteroid primarily crosstalk after brassinosteroid perception,but before their transcriptional activation.
This model provides a reasonable explanation for why a large proportion of brassinosteroid-responsive genes are also regulated by abscisic acid,and provides novel insight into the molecular mechanisms by which brassinosteroids could interact with abscisic acid.
油菜素甾醇(brassinosteroids,BRs)是上世纪70年代发现的第六大植物激素,其在植物的很多发育过程中起着重要作用,包括茎和根中的细胞分裂和伸长、光形态建成、生殖发育、叶片衰老、以及对胁迫的反应等。脱落酸(abscisic acid,ABA)作为另一类作用广泛的植物激素,也调控着植物生长发育的诸多方面,包括参与植物的胚胎发育、种子休眠与萌发、气孔关闭、开花时间和果实成熟等生理过程以及植物对胁迫的适应性反应。已有研究表明,BRs和ABA共同调控数百个基因的表达,从而共同调节很多发育和生理过程。同时,已经发现BR的一些突变体具有对ABA反应改变的表型,两者在发芽率上表现出相互拮抗的生理效应。然而,对于两者互作是否通过修饰或介入到初级信号途径中,抑或通过平行或独立的途径起作用,却仍然未知。
本研究工作主要包括:首先利用BR相关的多种遗传材料,系统研究了BRs和ABA两类激素在调节种子发芽率上具有相互拮抗的作用,为研究信号互作奠定生理基础。接着利用分子(DWF4和CPD转录水平)和生化标记(BES1的磷酸化状态)并结合ABA合成缺失突变体,证明外源的ABA能快速抑制BR的信号输出。再者,研究发现BR受体BRI1的强突变体bri1-116中,ABA仍能调节BR的信号输出;且在BKI1-YFP过表达系中,BKI1-YFP的亚细胞定位也不受ABA的影响,已知油菜素内酯(BRs中最具活性的一种形式)能促使BKI1-YFP从膜上解离;而用LiCl处理使下游组分BIN2失活却抑制了ABA对BR信号输出的调节。最后,利用ABA早期信号突变体abi1和abi2,得出ABA调控BR的初级信号转导途径主要依赖于ABA信号组分ABI2,部分依赖于ABI1。因此我们提出了BRs与ABA信号互作的模型,ABA对BR初级信号转导途径的调节界定在受体BRI复合体的下游,BIN2(含BIN2)的上游。这一模型有力的解释了为什么受BR调节的基因有一大部分也受到了ABA的调节,同时为BRs和ABA信号的互作提供了新的分子机制。
本研究通过遗传、细胞和分子水平的系统深入研究,确定了ABA影响BR的初级信号输出,且界定了信号互作的区域,同时形成了ABA对BR信号输出影响的初步模型。本研究从分子层面有力的支持了BRs和ABA在生理实验中的拮抗效应,具有重要的理论意义。同时为研究激素互作提供了很好的范例,也为植物更好的适应环境提供理论依据和目标基因。
Phytohormones play essential roles in coordinately regulating a large array of developmental processes.Studies have revealed that brassinosteroids and abscisic acid interact to regulate hundreds of genes'expression,governing many biological processes.However,whether their interaction is through modification or intersection of their primary signaling cascades or by independent or parallel pathways remains a big mystery.
Using biochemical and molecular markers of brassinosteroid signaling and abscisic acid biosynthetic mutants,we demonstrated that exogenous abscisic acid rapidly inhibits brassinosteroid signaling outputs as indicated by the phosphorylation status of BES1 and brassinosteroid-responsive gene expression.Experiments using a bri1 null-allele,bri1-116,and analysis of subcellular localization of BKI1-YFP further revealed that the brassinosteroid receptor complex is not required for abscisic acid to act on brassinosteroid signaling outputs.However,when the BR downstream signaling component BIN2 is inhibited by LiCl,abscisic acid failed to inhibit brassinosteroid signaling outputs.Furthermore,using a set of abscisic acid insensitive mutants,we found that regulation of abscisic acid on brassinosteroid primary signaling pathway is dependent on the abscisic acid early signaling components ABI1 and ABI2.We propose that the signaling cascades of abscisic acid and brassinosteroid primarily crosstalk after brassinosteroid perception,but before their transcriptional activation.
This model provides a reasonable explanation for why a large proportion of brassinosteroid-responsive genes are also regulated by abscisic acid,and provides novel insight into the molecular mechanisms by which brassinosteroids could interact with abscisic acid.
引文
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