Regulation of PIN-FORMED-mediated Polar Auxin Transport in Plant Gravitropism

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

Abstract

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

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.

Figures/Tables 3

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

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

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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