植物向重力反应中PIN-FORMED介导的生长素极性运输调控

doi:10.13560/j.cnki.biotech.bull.1985.2023-0882

摘要/Abstract

摘要：

植物的向性，即植物对光或重力等环境刺激信号产生的定向生长反应。在向重力性反应中，植物器官将重力感知为定向环境信号，来控制其器官的生长方向以促进生存。植物激素生长素及其极性运输在植物向重力反应中起着决定性的调控作用。质膜定位的生长素输出蛋白PIN-FORMED（PIN）通过动态的亚细胞极性定位，改变生长素运输的方向以响应环境刺激，由此植物器官间建立的生长素浓度梯度是细胞差异化伸长和器官弯曲的基础，来调控植物的形态建成和生长发育过程。本文主要讨论发生在植物重力感受细胞内早期重力感知和信号转导机制的最新研究进展、PIN介导的生长素极性运输、PIN的极性定位以及质膜蛋白丰度的调控机制等。

关键词: 向重力性, PIN, 生长素转运, 极性, 内膜运输

Abstract:

Plant tropisms are the directional growth response of plants to environmental stimuli such as light or gravity. In gravitropism, plant organs sense directional environmental cues to control growth orientation to promote their survival. The phytohormone auxin and its polar transport serve as a major coordinative signal in plant gravitropism. The PIN family auxin exporters, through their dynamic polar subcellular localizations at the plasma membrane（PM）, redirect polar auxin transport in response to environmental stimuli, thus the auxin gradients across tissues or organs underlie differential cell elongation and bending, which regulates the morphological formation and growth process of plants. In this review, we focus on recent advances in the mechanisms underlying the perception and signal transduction of gravity that take place within the sensing cells in the early process of gravitropism, the regulatory mechanisms of PIN-mediated auxin polar transport, PIN polar localization, and PM protein abundance.

Key words: gravitropism, PIN, auxin transport, polarity, endomembrane trafficking

王贤, 彭亚坤, 陈猛, 孔梦娟, 谭树堂. 植物向重力反应中PIN-FORMED介导的生长素极性运输调控[J]. 生物技术通报, 2024, 40(3): 25-40.

WANG Xian, PENG Ya-kun, CHEN Meng, KONG Meng-juan, TAN Shu-tang. Regulation of PIN-FORMED-mediated Polar Auxin Transport in Plant Gravitropism[J]. Biotechnology Bulletin, 2024, 40(3): 25-40.

图/表 3

图1 根和地上部重力感应位置及相关调控基因（A）拟南芥花序茎的结构示意图。（B）拟南芥花序茎纵剖面的结构示意图：表皮，皮层及内皮层。（C）拟南芥内皮层细胞的结构示意图。淀粉体沿重力矢量向细胞底部沉积，内胚层细胞几乎完全被大的中央液泡充满。（D）拟南芥根冠的结构示意图，由四层小柱细胞（蓝色）和周围的侧根冠细胞组成。（E）拟南芥根冠柱细胞（也称作小柱细胞）的结构示意图。细胞核位于顶端，液泡位于中央，淀粉体受重力在细胞底部沉积。在垂直生长下，LZY蛋白通过CCL结构域可以与RLD（RCC1-like domain）蛋白的BRX结构域相互作用。（F）拟南芥根的重定向。（G）重定向的重力刺激激活MKK5-MPK3激酶模块介导的LZY蛋白磷酸化，随后，磷酸化的LZYs与TOC蛋白相互作用促使LZY蛋白由质膜向淀粉体转运。淀粉体沉降介导LZY蛋白转移到根冠柱细胞新的下侧质膜

Fig. 1 Gravity-sensing positions and related regulators in roots and shoots （A）A schematic diagram showing the stem（inflorescence）of Arabidopsis.（B）Schematic structure of the Arabidopsis inflorescent stem longitudinal region: epidermis, cortex, and endodermis.（C）A schematic diagram of the endodermal cell in Arabidopsis. Amyloplasts sediment toward the bottom of the cell according to the gravity vector. The endodermal cell is occupied by a large central vacuole.（D）Schematic structure of the Arabidopsis root cap, which is composed of four layers of columella cells (blue) and the surrounding lateral root cap cells. （E）A schematic diagram of columella cells in Arabidopsis. Nucleus is in the upper part of the cell, and the vacuole located in the center of cell. Amyloplasts sediment toward the bottom of the cell according to the gravity vector. Under vertical growth conditions, the CCL domain of LZYs interacts with the BRX domain of RLD（RCC1-like domain）. （F）Root reorientation of Arabidopsis thaliana.（G）Redirected gravity stimulation activates the LZY protein phosphorylation mediated by the MKK5-MPK3 kinase module, and subsequently, phosphorylated LZYs interacting with the TOC protein prompted the transport of LZY protein from the plasma membrane to statoliths. Amyloplast sedimentation guides the LAZY proteins to distribute onto the new lower side of the plasma membrane in columella cells

图2 翻译后修饰调节PIN蛋白定位 PIN蛋白回收和液泡降解依赖于网格蛋白介导的内吞作用、囊泡运输和内膜融合进行。翻译后修饰决定了PIN蛋白的亚细胞定位。蛋白激酶PID、D6PK、CAMEL、MPKs和CRKs，直接磷酸化PINs并改变PINs的极性和转运活性。蛋白磷酸酶PP2A、PP6、PP1参与PIN蛋白的去磷酸化。PAX募集BRX来调节局部PI（4,5）P2的生物合成，从而调节PIN1的内吞作用。对于PIN2，磷酸化的PIN2蛋白可以募集MAB4/MEL，后者通过内体运输途径进一步调节PIN2的极性定位。在这些调节因子中，PAX/D6PK仅定位于质膜基部

Fig. 2 Post-translational modifications regulate the localization of PIN The recycling and vacuolar degradation of PIN proteins rely on clathrin-mediated endocytosis, vesicle trafficking and endomembrane fusion. Posttranslational modifications determine the subcellular localizations of PIN proteins. Protein kinases, including PID, D6PK, CAMEL, MPKs, and CRKs, directly phosphorylate PINs and determine PIN polarity or activity of transport. Protein phosphatases PP2A, PP6, and PP1 are involved in the dephosphorylation of PINs. PAX recruits BRX to regulate local PI（4,5）P2 biosynthesis, thus modulating the endocytosis of PINs. For PIN2, phosphorylated PIN2 protein recruits MAB4/MEL proteins, which further regulate PIN2 polarity through the endosomal trafficking pathway. Among these regulators, PAX and D6PK are localized to the basal side of plasma membrane

表1 PINs蛋白相关的抑制剂

Table 1 Related inhibitors of PINs

PIN蛋白相关的抑制剂 Inhibitors of PINs	功能 Function	参考文献 References
抑草生NPA	PIN蛋白结合抑制剂	[118⇓-120,123⇓⇓ -126]
萘普生Naproxen	PIN蛋白结合抑制剂	[127]
黄酮醇Flavonols	PIN蛋白结合抑制剂	[116]
褪黑素Melatonin	抑制PIN1, PIN3 和PIN7表达	[128]
S-亚硝基谷胱甘肽GSNO	抑制PIN2内吞	[115]
ES4 Endosidin4	ARF-GEFs抑制剂	[129]
BFA Brefeldin A	ARF-GEFs抑制剂	[67⇓-69]
渥曼青霉素Wortmannin	抑制蛋白靶向液泡分选和内吞	[130-131]
2,3,5-三碘苯甲酸 TIBA	抑制PIN在内膜系统的运输	[67]

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