叶绿体TOC-TIC蛋白复合体转运机制研究进展

doi:10.16420/j.issn.0513-353x.2020-0860

摘要/Abstract

摘要：

叶绿体蛋白是植物进行光合作用所必不可缺的物质。参与光合作用的大部分叶绿体蛋白在细胞质中合成,经过TOC-TIC复合体转运至叶绿体中行使其功能。本文中对参与叶绿体蛋白转运的TOC-TIC复合体、转运肽和分子伴侣蛋白的特征、作用及蛋白转运途径进行了总结,并对近年来关于叶绿体蛋白转运研究的新发现进行了介绍和评述。

关键词: 叶绿体, 蛋白转运, TOC-TIC复合体, TOC, TIC

Abstract:

Chloroplast protein is an indispensable substance for plant photosynthesis. Most of proteins involved in photosynthesis are first synthesised in the cytoplasm and subsequently transported into chloroplast by TOC-TIC complexes,then a series of life activities can be carried out. In this review,the characteristics and roles of TOC-TIC complexes,transport peptide and chaperone protein as well as protein transport pathways were summarized,and some new findings on chloroplast protein transport in recent years were introduced and discussed.

Key words: chloroplast, protein transport, TOC-TIC complex, TOC, TIC

中图分类号:

Q732

王琪, 李云洲, 须文, 闫见敏. 叶绿体TOC-TIC蛋白复合体转运机制研究进展[J]. 园艺学报, 2021, 48(4): 689-704.

WANG Qi, LI Yunzhou, XU Wen, YAN Jianmin. Research Advance of Protein Transport Mechanism of TOC-TIC Complexes in Plant Chloroplast[J]. Acta Horticulturae Sinica, 2021, 48(4): 689-704.

图/表 5

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组分 Component	预测功能 Proposed function	拟南芥中同源基因 Homologues in Arabidopsis	突变体 Mutant name	缺失突变体表现型 Knockout mutant phenotype(s)	参考文献 Reference
Toc159	识别前体蛋白 Preprotein recognition	AtToc159	ppi2/toc159	白化 Albino	Bauer et al.2000 Kubis et al.2004
		AtToc132	toc132	无明显表型 None detected	Kubis et al.2004
		AtToc120	toc120	无明显表型 None detected	Ivanova et al.2004 Kubis et al.2004
			toc132toc120 双突变	白化 Albino	Kubis et al.2004
		AtToc90	ppi4/toc90	无明显表型 None detected	Hiltbrunner et al.2004 Kubis et al.2004
			ppi2ppi4 双突变	白化 Albino	Hiltbrunner et al.2004 Kubis et al.2004
Toc34	识别前体蛋白 Preprotein recognition	AtToc33	ppi1/toc33	幼苗苍白转绿 Albino turn green	Jarvis et al.1998 Kubis et al.2003
Toc34	识别前体蛋白 Preprotein recognition	AtToc34	ppi3/toc34	影响根部发育 Stunted roots	Constan et al.2004a
Toc75	形成易位通道 Translocation channel	AtToc75-Ⅲ	toc75-Ⅲ	胚胎致死 Embryo lethal	Baldwin et al.2005
		AtToc75-Ⅳ	toc75-IV	质体结构异常 Etioplast defects	Baldwin et al.2005
		AtToc75-Ⅰ	—	—	Baldwin et al.2005
Toc64	结合转运肽 Binding transit peptide 调控转运效率 Transport efficiency	AtToc64-Ⅲ	toc64-III	—	Qbadou et al.2006 Aronsson et al.2007
		AtToc64-Ⅴ	toc64-V	—	Aronsson et al.2007
		AtToc64-Ⅰ	toc63-I	—	Aronsson et al.2007
Tic22		AtTic22-Ⅲ	—	—	—
		AtTic22-Ⅳ	—	—	—
Tic20	参与TOC-TIC超复合体形成 TOC-TIC Supercomplex	AtTic20-Ⅰ	tic20-I	白化 Albino	Chen et al.2002;Teng et al.2006
		AtTic20-Ⅳ	tic20-IV	无明显表型 None detected	Hirabayashi et al.2011
			tic20-Itic20-IV 双突变	胚胎致死 Embryo lethal	Hirabayashi et al.2011; Kasmati et al.2011
		AtTic20-Ⅱ	—	—	—
		AtTic20-Ⅴ	—	—	—
Tic21	参与易位通道的构成 Translocation channel	AtTic21	pic1	白化 Albino	Teng et al.2006;Duy et al.2007
Tic110	参与易位通道构成 Translocation channel	AtTic110	tic110	胚胎致死 Embryo lethal	Kovacheva et al.2005;Inaba et al.2005
Tic40	促进转运 Transport rate promotion	AtTic40	tic40	浅绿色 Pale-green	Chou et al.2003; Kovacheva et al.2005
Tic55	氧化还原调控 Redox regulation	AtTic55	tic55	无明显表型 None detected	Boij et al.2009
Tic62	氧化还原调控 Redox regulation	AtTic62	tic62	—	Benz et al.2009
Tic32	钙依赖调控 Calcium regulation	AtTic32-Ⅳa	tic32-IVa	胚胎致死 Embryo lethal	H?rmann et al.2004
Tic32	钙依赖调控 Calcium regulation	AtTic32-Ⅳb	—	—	—
Tic236	连接TOC与TIC TOC-TIC connection	AtTic236	tic236	胚胎致死 Embryo lethal	Chen et al.2018

组分 Component	预测功能 Proposed function	拟南芥中同源基因 Homologues in Arabidopsis	突变体 Mutant name	缺失突变体表现型 Knockout mutant phenotype(s)	参考文献 Reference
Toc159	识别前体蛋白 Preprotein recognition	AtToc159	ppi2/toc159	白化 Albino	Bauer et al.2000 Kubis et al.2004
		AtToc132	toc132	无明显表型 None detected	Kubis et al.2004
		AtToc120	toc120	无明显表型 None detected	Ivanova et al.2004 Kubis et al.2004
			toc132toc120 双突变	白化 Albino	Kubis et al.2004
		AtToc90	ppi4/toc90	无明显表型 None detected	Hiltbrunner et al.2004 Kubis et al.2004
			ppi2ppi4 双突变	白化 Albino	Hiltbrunner et al.2004 Kubis et al.2004
Toc34	识别前体蛋白 Preprotein recognition	AtToc33	ppi1/toc33	幼苗苍白转绿 Albino turn green	Jarvis et al.1998 Kubis et al.2003
Toc34	识别前体蛋白 Preprotein recognition	AtToc34	ppi3/toc34	影响根部发育 Stunted roots	Constan et al.2004a
Toc75	形成易位通道 Translocation channel	AtToc75-Ⅲ	toc75-Ⅲ	胚胎致死 Embryo lethal	Baldwin et al.2005
		AtToc75-Ⅳ	toc75-IV	质体结构异常 Etioplast defects	Baldwin et al.2005
		AtToc75-Ⅰ	—	—	Baldwin et al.2005
Toc64	结合转运肽 Binding transit peptide 调控转运效率 Transport efficiency	AtToc64-Ⅲ	toc64-III	—	Qbadou et al.2006 Aronsson et al.2007
		AtToc64-Ⅴ	toc64-V	—	Aronsson et al.2007
		AtToc64-Ⅰ	toc63-I	—	Aronsson et al.2007
Tic22		AtTic22-Ⅲ	—	—	—
		AtTic22-Ⅳ	—	—	—
Tic20	参与TOC-TIC超复合体形成 TOC-TIC Supercomplex	AtTic20-Ⅰ	tic20-I	白化 Albino	Chen et al.2002;Teng et al.2006
		AtTic20-Ⅳ	tic20-IV	无明显表型 None detected	Hirabayashi et al.2011
			tic20-Itic20-IV 双突变	胚胎致死 Embryo lethal	Hirabayashi et al.2011; Kasmati et al.2011
		AtTic20-Ⅱ	—	—	—
		AtTic20-Ⅴ	—	—	—
Tic21	参与易位通道的构成 Translocation channel	AtTic21	pic1	白化 Albino	Teng et al.2006;Duy et al.2007
Tic110	参与易位通道构成 Translocation channel	AtTic110	tic110	胚胎致死 Embryo lethal	Kovacheva et al.2005;Inaba et al.2005
Tic40	促进转运 Transport rate promotion	AtTic40	tic40	浅绿色 Pale-green	Chou et al.2003; Kovacheva et al.2005
Tic55	氧化还原调控 Redox regulation	AtTic55	tic55	无明显表型 None detected	Boij et al.2009
Tic62	氧化还原调控 Redox regulation	AtTic62	tic62	—	Benz et al.2009
Tic32	钙依赖调控 Calcium regulation	AtTic32-Ⅳa	tic32-IVa	胚胎致死 Embryo lethal	H?rmann et al.2004
Tic32	钙依赖调控 Calcium regulation	AtTic32-Ⅳb	—	—	—
Tic236	连接TOC与TIC TOC-TIC connection	AtTic236	tic236	胚胎致死 Embryo lethal	Chen et al.2018