植物ESCRT复合体的功能研究进展

doi:10.11983/CBB22038

摘要/Abstract

摘要： 真核细胞中, 功能高度保守的内体蛋白分选转运装置ESCRT在胞吞途径和蛋白分泌途径中均扮演重要角色。植物细胞中, 该装置包含ESCRT-I、ESCRT-II、ESCRT-III和VPS4/SKD1复合体4个亚基, 但缺乏ESCRT-0亚基。ESCRT的每个亚基均由多个蛋白构成。目前, 针对ESCRT的研究已经证实, 其在泛素化的膜蛋白进入多囊泡体/液泡前体(MVB/PVC)内腔过程中发挥重要调控作用; 同时在自噬途径以及应对环境胁迫等方面也具有重要的调节功能。该文首先介绍了植物中ESCRT复合体的组成及生物学功能, 然后总结了植物中特有ESCRT复合体组分蛋白的最新研究进展, 最后探讨了有关ESCRT复合体研究中尚未解决的重要科学问题。

关键词: ESCRT复合体, MVB/PVC, 胞吞

Abstract: In eucaryotic cells, the endosomal sorting complex required for transport (ESCRT) machinery, consisting of the conserved multiple sub-complexes, plays important roles in both endocytic and secretory pathways. In plant cells, the ESCRT machinery consists of four ESCRT complexes (ESCRT-I, -II, -III, and the VPS4/SKD1 complex) and several ESCRT accessory proteins, but lacks of the ESCRT-0. Every ESCRT complex contains several subunit proteins. Recent researches have demonstrated that the ESCRT machinery plays an essential role in the vacuolar sorting of ubiquitinated membrane proteins via regulation the biogenesis of multivesicular body (MVB)/PVC (prevacuolar compartment) and autophagic pathway. Recent researches have demonstrated that the ESCRT machinery plays an essential role in the vacuolar sorting of ubiquitinated membrane proteins via regulation the biogenesis of multivesicular body (MVB)/PVC (prevacuolar compartment) and autophagosome as well as response to environmental stresses. In this review, we firstly introduce the subunits and biological functions of the ESCRT machinery in plants, and then summarize the latest research advances on the plant-specific ESCRT complex components. Finally, we discuss future study on the unsolved scientific problems of the ESCRT complex in plants.

Key words: ESCRT complex, MVB/PVC, endocytosis

吴凡, 沈锦波, 胡帅. 植物ESCRT复合体的功能研究进展. 植物学报, 2022, 57(5): 697-712.

Wu Fan, Shen Jinbo, Hu Shuai. Research Advances of the Plant ESCRT Machinery. Chinese Bulletin of Botany, 2022, 57(5): 697-712.

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链接本文: https://www.chinbullbotany.com/CN/10.11983/CBB22038

https://www.chinbullbotany.com/CN/Y2022/V57/I5/697

图/表 5

表1 内体蛋白分选转运装置(ESCRT)组分蛋白及其功能

Table 1 Endosomal sorting complex required for transport (ESCRT) components and their functions in plants

复合体	基因	功能	参考文献
ESCRT-0	N.I.
	VPS23A (AT3G12400)	膜蛋白内吞分选; 非生物逆境胁迫应答调控; 细胞分裂; 病毒复制	2020
	VPS23B (AT5G13860)	病毒复制	2009
ESCRT-I	VPS28A (AT4G21560)	胚胎发育; 生长素响应; 膜蛋白内吞分选; 液泡发育	2020
	VPS28B (AT4G05000)	胚胎发育; 生长素响应; 膜蛋白内吞分选; 液泡发育; 植物免疫	2020
	VPS37-1 (AT3G53120)	植物免疫	2013
	VPS37-2 (AT2G36680)
	VPS22 (AT4G27040)	水稻主根生长; 胚乳灌浆	2013
ESCRT-II	VPS25 (AT4G19003)	生长素响应; 主根生长	2014
ESCRT-II	VPS36 (AT5G04920)	胚胎发育; 主根生长; MVB/PVC发育; 液泡发育; 膜蛋白内吞分选	2017
	VPS2.1 (AT2G06530)	胚胎发育; MVB/PVC发育; 膜蛋白内吞分选; 自噬途径	2020
	VPS2.2 (AT5G44560)
	VPS2.3 (AT1G03950)
	VPS20.1 (AT5G63880)	MVB/PVC发育; 膜蛋白内吞分选; 病毒复制	2014
ESCRT-III	VPS20.2 (AT5G09260)
	SNF7A (AT2G19830)	MVB/PVC发育; 膜蛋白内吞分选; 病毒复制	2014
	SNF7B (AT4G29160)	MVB/PVC发育; 植物发育; 病毒复制	2021
	VPS24-1 (AT5G22950)	MVB/PVC发育; 膜蛋白内吞分选; 病毒复制	2019
	VPS24-2 (AT3G45000)
	CHMP1A (AT1G73030)	胚胎发育; MVB/PVC发育; 膜蛋白内吞分选, 自噬途径	2015
ESCRT-III辅助蛋白	CHMP1B (AT1G17730)	胚胎发育; MVB/PVC发育; 膜蛋白内吞分选; 自噬途径	2015
	VPS60-1 (AT3G10640)
	VPS60-2 (AT5G04850)
	CHMP7 (AT3G62080)	衰老; 非生物逆境胁迫应答调控	2016
	ISTL (AT1G34220)	MVB/PVC发育; 植物发育; 植物免疫	2016
VPS4/SKD1复合体	VPS4/SKD1 (AT2G27600)	MVB/PVC发育; 膜蛋白内吞分选; 病毒复制; 植物免疫	2016
VPS4/SKD1复合体	LIP5 (AT4G26750)	MVB/PVC发育; 植物发育; 膜蛋白内吞分选; 植物免疫; 非生物逆境胁迫应答调控	2016
	FREE1 (AT1G20110)	MVB/PVC发育; 液泡发育; 膜蛋白内吞分选; 自噬途径; 非生物逆境胁迫应答调控	2020
植物特有蛋白	FYVE4 (AT1G61690)	MVB/PVC发育; 膜蛋白内吞分选	2021
	PROS (AT4G24370)	VPS4/SKD1复合体活性调控; 细胞延伸	2014
	BRAF (AT5G14020)	MVB/PVC发育; 与FREE1竞争性结合VPS23A	2018
	RST1 (AT3G27670)	胚胎发育; 茎角质层蜡的分布; 植物免疫; MVB/ PVC发育	2019

表1 内体蛋白分选转运装置(ESCRT)组分蛋白及其功能

Table 1 Endosomal sorting complex required for transport (ESCRT) components and their functions in plants

复合体	基因	功能	参考文献
ESCRT-0	N.I.
	VPS23A (AT3G12400)	膜蛋白内吞分选; 非生物逆境胁迫应答调控; 细胞分裂; 病毒复制	2020
	VPS23B (AT5G13860)	病毒复制	2009
ESCRT-I	VPS28A (AT4G21560)	胚胎发育; 生长素响应; 膜蛋白内吞分选; 液泡发育	2020
	VPS28B (AT4G05000)	胚胎发育; 生长素响应; 膜蛋白内吞分选; 液泡发育; 植物免疫	2020
	VPS37-1 (AT3G53120)	植物免疫	2013
	VPS37-2 (AT2G36680)
	VPS22 (AT4G27040)	水稻主根生长; 胚乳灌浆	2013
ESCRT-II	VPS25 (AT4G19003)	生长素响应; 主根生长	2014
ESCRT-II	VPS36 (AT5G04920)	胚胎发育; 主根生长; MVB/PVC发育; 液泡发育; 膜蛋白内吞分选	2017
	VPS2.1 (AT2G06530)	胚胎发育; MVB/PVC发育; 膜蛋白内吞分选; 自噬途径	2020
	VPS2.2 (AT5G44560)
	VPS2.3 (AT1G03950)
	VPS20.1 (AT5G63880)	MVB/PVC发育; 膜蛋白内吞分选; 病毒复制	2014
ESCRT-III	VPS20.2 (AT5G09260)
	SNF7A (AT2G19830)	MVB/PVC发育; 膜蛋白内吞分选; 病毒复制	2014
	SNF7B (AT4G29160)	MVB/PVC发育; 植物发育; 病毒复制	2021
	VPS24-1 (AT5G22950)	MVB/PVC发育; 膜蛋白内吞分选; 病毒复制	2019
	VPS24-2 (AT3G45000)
	CHMP1A (AT1G73030)	胚胎发育; MVB/PVC发育; 膜蛋白内吞分选, 自噬途径	2015
ESCRT-III辅助蛋白	CHMP1B (AT1G17730)	胚胎发育; MVB/PVC发育; 膜蛋白内吞分选; 自噬途径	2015
	VPS60-1 (AT3G10640)
	VPS60-2 (AT5G04850)
	CHMP7 (AT3G62080)	衰老; 非生物逆境胁迫应答调控	2016
	ISTL (AT1G34220)	MVB/PVC发育; 植物发育; 植物免疫	2016
VPS4/SKD1复合体	VPS4/SKD1 (AT2G27600)	MVB/PVC发育; 膜蛋白内吞分选; 病毒复制; 植物免疫	2016
VPS4/SKD1复合体	LIP5 (AT4G26750)	MVB/PVC发育; 植物发育; 膜蛋白内吞分选; 植物免疫; 非生物逆境胁迫应答调控	2016
	FREE1 (AT1G20110)	MVB/PVC发育; 液泡发育; 膜蛋白内吞分选; 自噬途径; 非生物逆境胁迫应答调控	2020
植物特有蛋白	FYVE4 (AT1G61690)	MVB/PVC发育; 膜蛋白内吞分选	2021
	PROS (AT4G24370)	VPS4/SKD1复合体活性调控; 细胞延伸	2014
	BRAF (AT5G14020)	MVB/PVC发育; 与FREE1竞争性结合VPS23A	2018
	RST1 (AT3G27670)	胚胎发育; 茎角质层蜡的分布; 植物免疫; MVB/ PVC发育	2019

图1 拟南芥中内体蛋白分选转运装置(ESCRT)的作用机制及泛素化的膜蛋白内吞过程胞外蛋白和营养物质及新合成的蛋白分别通过蛋白内吞和分泌途径进入反面高尔基体(TGN)。经泛素化修饰的膜蛋白被ESCRT复合物识别, 并以依赖ESCRT的作用机制通过内腔泡(ILVs)进入MVB/PVC内腔, MVB/PVC与液泡融合, 最终使货物蛋白进入液泡中降解, 与此同时ESCRT-III复合体在VPS4/SKD1 ATP水解酶的作用下解离, 进入下一个循环。PM: 质膜; MVB/PVC: 多囊泡体/液泡前体

Figure 1 The endosomal sorting complex required for transport (ESCRT) mechanism and the endocytosis of ubiquitinated membrane proteins in Arabidopsis Extracellular proteins, materials and newly synthesized proteins are transported to trans Golgi network (TGN) via the endocytic or the secretory pathway, respectively. The ubiquitinated membrane proteins are then recognized by ESCRT machinery and be transported into the lumen of MVB/PVC via intraluminal vesicles (ILVs). After membrane fusion of MVB/PVC and vacuole, the cargo proteins are then transported to vacuole for degradation.The ESCRT-III complex is finally released from MVB/PVC membrane and the components are disassembled from each other by the activation of VPS4/SKD1 ATPase and recycled. PM: Plasma membrane; MVB/PVC: Multivesicular body/prevacuolar compartment

图2 类TOM1 (TOL)家族蛋白与动物及酵母ESCRT-0组分蛋白的氨基酸序列比对横线标注区域分别表示VHS和GAT结构域。

Figure 2 Amino acid sequences alignment of TOM1-like (TOL) family proteins in plant and ESCRT-0 components in animal and yeast Transverse line region indicate the VHS domain and GAT domain, respectively.

图3 不同物种中ESCRT-I组分蛋白(VPS23 (A)、VPS28 (B)和VPS37 (C))间的氨基酸序列比对横线区域表示UEV结构域, 虚线框中表示氨基酸保守区域。

Figure 3 Sequences alignment of ESCRT-I components (VPS23 (A), VPS28 (B) and VPS37 (C)) from different organisms Transverse line and dashed box regions indicate UEV domain and conserved amino acids, respectively.

图4 不同物种中ESCRT-III组分蛋白(VPS2 (A)、VPS20 (B)、VPS24 (C)和SNF7 (D))间的氨基酸序列比对横线区域分别表示保守的CC结构域和SNF7结构域。

Figure 4 Sequences alignment of ESCRT-III components (VPS2 (A), VPS20 (B), VPS24 (C) and SNF7 (D)) from different organisms Transverse lines indicate conserved CC domain and SNF7 domain, respectively.

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