细胞核中的光敏素信号传递.ppt

光敏素通过与PIF1、PIF3、PIF4、PIF5结合,通过抑制负调控幼苗发育中光反应的PIFs促进形态建成。红光激活反应中，光敏素迅速使PIF5和PIF1磷酸化并使之降解 。 PIF1直接与叶绿素合成途径中的protochlorophyllide oxidoreductase (POR) 基因启动子中G-box (CACGTG)结合，调控其表达，同时间接调控PORA、 PORB、 HO3、FeChII的表达 。 PIF3调控叶绿素合成途径关键基因HEMA1（编码glutamyl tRNA reductase）的表达 。 PIF4负调控NIA2 (nitrate reductase 2)的表达，在PHYB信号传导途径中也起负调控作用。 HFR1也是一种bHLH转录因子，与PIF3形成二聚体，在远红光下其mRNA的量是红光下的30倍，表明它是PHYA信号途径中的因子 。 PIF3暗中在核中积累，在光下经PHYA、PHYB、PHYD的介导迅速降解。黑暗中PIF3 的积累需要COP1 （Constitutive photomorphogenesis1）的作用，在光下的降解与COP1无关 。 A model of phytochrome signal transduction. Activated phytochrome (Pfr) is proposed to regulate transcription through several parallel pathways. A rapid response involves Pfr translocation to the nucleus, where it binds transcription factors of the bHLH family (in particular PIF3). Key regulatory transcription factors (RTFs) that are responsible for inducing a range of light regulated genes are subsequently activated. In a second nuclear-localized pathway, phytochromes are proposed to bind response regulators (RR), which stabilize them in the activated form and can induce light-regulated gene expression by inhibiting COP1-, COP10- and CSN-dependent proteolysis of the HY5 transcription factor and by binding to activated cryptochromes (cry). In all cases, regulation of the genes responsible for photomorphogenesis is predicted to require chromatin remodelling mediated by the DET1/DDB1 nucleosome-binding complex. In the cytoplasm, phytochrome may activate gene expression through G-proteins (G), calcium and cGMP-dependent pathways, which are regulated by SUB1. In addition, phytochromes may be sequestered away from the signalling-competent pool by PKS1. Elements involved in signalling from specific photoreceptors or controlling specific responses have not been included. COP1是光调控信号传导途径中的一个成分，是E3泛素连接酶，含有几个可识别区域，包括RING-锌指结合区域、线圈区域、核心区和WD-40 重复区域，是蛋白酶降解复合物的组成成分，分别在细胞质、细胞核定位以及与其他蛋白如SPA1、CIPs(COP1 interacting proteins)共同作用中起作用，COP1在核中的定位受光的调控, 在核中与目标蛋白结合，将目标蛋白送到蛋白酶降解复合体降解。HY5、LAF1、CIP4、