生长素输出载体PIN蛋白的质膜定位机制

doi:10.11983/CBB15017

摘要/Abstract

摘要： 生长素浓度梯度影响植物个体及其器官的形态建成, 而PIN (PIN-FORMED)蛋白决定组织中的生长素流向。细胞质膜的脂筏特性是PIN蛋白在质膜上不均匀分布的基础。与此同时, 网格蛋白介导的胞吞、蛋白质的磷酸化/去磷酸化甚至基因的转录调控影响PIN蛋白的这种极性定位。另外, 在多细胞植物起源之时, PIN蛋白可能经历了从内质网膜定位到质膜定位的转变。

关键词: 生长素, PIN, 胞吞, 磷酸化, 植物进化

Abstract: Auxin concentration affects the morphological establishment of plant body and its organs, while PIN (PIN- FORMED) proteins determine the direction of auxin stream within tissues. The nature of lipid rafts of cellular plasma membrane underlies the uneven distribution of PINs in the plasma membrane. Meanwhile, clathrin-mediated endocytyosis, proteins phosphorylation/dephosphorylation and even genes transcriptional regulation affect this polar location of PINs. Additionally, at the time when multicellular plants emerge, PINs may undergo a transition from endoplasmic reticulum membrane to plasma membrane positioning.

Key words: auxin, PIN, endocytosis, phosphorylation, plant evolution

曹文杰, 李贵生. 生长素输出载体PIN蛋白的质膜定位机制. 植物学报, 2016, 51(2): 265-273.

Wenjie Cao, Guisheng Li. Plasma Membrane Positioning Mechanism of Auxin Efflux Carrier PIN Proteins. Chinese Bulletin of Botany, 2016, 51(2): 265-273.

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链接本文: https://www.chinbullbotany.com/CN/10.11983/CBB15017

https://www.chinbullbotany.com/CN/Y2016/V51/I2/265

图/表 2

图1 PIN蛋白质膜极性定位的细胞学机制质膜上与细胞壁成分相连的部分(●)可以阻止PIN蛋白的扩散, 位于质膜的PID和PP2A分别对PIN蛋白进行磷酸化和去磷酸化。网格蛋白介导PIN蛋白的胞吞, 但是膜上的激酶TMK与ABP1-生长素复合体的结合抑制该过程。进入胞质的PIN蛋白最后到达TGN/EE, 从这里开始它们会通过再循环调控到达质膜的特定部位或者通过降解途径进入液泡。肌动蛋白纤丝介导PIN蛋白的向顶移动, 而微管介导其反方向的运动, 在向基运输中, GNOM是重要成分; 磷酸化的PIN蛋白将到达质膜顶部, 而去磷酸化的PIN蛋白则移向基部。单泛素化促进PIN蛋白的胞吞, 而多泛素化使它进入液泡降解途径。P: 磷酸化PIN蛋白; U: 泛素化PIN蛋白。

Figure 1 The cellular mechanism of the plasma membrane positioning of PINs The parts of plasma membrane connecting to cell wall (●) prevent the diffusion of PINs, and PID and PP2A located in plama membrane respectively phosphorylates and dephosphory- lates PINs. Clathrin mediates the endocytosis of PINs, however this process is inhibited by the binding of kinase TMK to ABP1-auxin complex. PINs already entering into cytoplasm ultimately arrive at TGN/EE, whereby they undergo regulation in recycling and reach certain parts of plasma membrane or enter into vacuole via degradation pathway. Actin mediates the apical movement of PINs while microtubuli mediates the reverse migration, and in the basal transportation GNOM is a key component; phosphorylated PINs will reach the apical membrane while dephosphorylated ones move to the basal. Mono-ubiquitylation promotes endocytosis of PINs while poly-ubiquitylation gets them into vacuole degeradation pathway. P: Phosphorylated PINs; U: Ubiquitylated PINs.

图2 代表性PIN蛋白的系统发育关系 (A) 分支上的靴带分析值分别来自密码子1和2位的最大简约树/ 氨基酸序列的最大似然树/密码子1和2位的最大似然树; (B) 靴带分析值分别来自氨基酸序列的最大简约树/密码子1和2位的邻接树; (C) 靴带分析值分别来自密码子2和3位的最大似然树/氨基酸序列的邻接树。-表示靴带分析值小于50; ?表示不同方法得到不一致的拓扑结构; 分支上竖条的长短表示亲水环的长度, 灰色竖条表示PIN6的长亲水环的定性有争议。

Figure 2 Representative phylogenetic relationships of PINs (A) Bootstrap values on branches are respectively from most parsimony tree of the first and second position of codons/ most likelihood tree of amino acid sequence/most likelihood tree of the first and second position of codons; (B) Bootstrap values are respectively from most parsimony tree of amino acid sequence/neighbour joining tree of the first and second position of codons; (C) Bootstrap values are from most likelihood tree of the second and third position of codons/ neighbour joining tree of amino acid sequence. - indicates values lower than 50; ? indicate incongruent topology between different methods; The length of the vertical bar on branches suggests the length of hydrophilic loop, and the grey colour means that the determination of PIN6 possessing a long hydrophilic loop is equivocal.

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