植物细胞自噬基因的功能与转录调控机制

doi:10.11983/CBB20159

植物学报 ›› 2021, Vol. 56 ›› Issue (2): 201-217.DOI: 10.11983/CBB20159

植物细胞自噬基因的功能与转录调控机制

李喜豹, 赖敏怡, 梁山, 王小菁, 高彩吉, 杨超()

华南师范大学生命科学学院, 广东省植物发育生物工程重点实验室, 广州 510631

收稿日期:2020-09-29 接受日期:2020-12-25 出版日期:2021-03-01 发布日期:2021-03-17
通讯作者: 杨超
作者简介:^*E-mail: 798916640@qq.com
第一联系人：† 共同第一作者
基金资助:
广东省自然科学基金(2018A030310505);亚热带农业生物资源保护与利用国家重点实验室开放课题(SKLCUSA- b201812)

Function and Transcriptional Regulation of Autophagy-related Genes in Plants

Xibao Li, Minyi Lai, Shan Liang, Xiaojing Wang, Caiji Gao, Chao Yang()

Guangdong Provincial Key Laboratory of Biotechnology for Plant Development, School of Life Sciences, South China Normal University, Guangzhou 510631, China

Received:2020-09-29 Accepted:2020-12-25 Online:2021-03-01 Published:2021-03-17
Contact: Chao Yang
About author:First author contact:† These authors contributed equally to this paper

摘要/Abstract

摘要： 自噬(autophagy)是真核生物长期进化形成的一种高度保守的细胞内物质降解和周转途径, 通过形成双层膜结构的自噬体将包裹其中的待降解大分子物质, 如受损伤的蛋白质、蛋白质复合物和细胞器, 运送至液泡或溶酶体进行降解并产生可循环利用的降解产物。细胞自噬在植物生长发育和环境应答等过程中发挥重要作用。在拟南芥(Arabidopsis thaliana)和水稻(Oryza sativa)等模式植物中已鉴定到40多个自噬基因, 并发现其中多个基因在植物叶片衰老、种子成熟等发育阶段以及营养饥饿、干旱和病原菌侵染等逆境胁迫响应过程中显著上调表达, 但具体的转录激活或抑制机制有待阐明。该文综述了自噬基因在植物生长发育和胁迫应答过程中的功能与转录调控网络。

关键词: 植物细胞自噬, ATG基因, 转录调控, 胁迫应答, 生长发育

Abstract: Macroautophagy (hereafter termed autophagy) is an evolutionarily conserved cellular degradation and recyc- ling pathway in eukaryotes. In this pathway, cellular substances, such as dysfunctional proteins and damaged organelles, are sequestered by a double-membrane structure, autophagosome, and eventually delivered to the lysosomes or vacuoles for degradation and recycling. Autophagy plays essential roles in plant growth and development, as well as in response to environmental stresses. Up to now, more than 40 autophagy-related (ATG) genes have been identified in model plants such as Arabidopsis thaliana and Oryza sativa. It is well established that a large number of ATG genes are up-regulated during specific developmental stages such as leaf senescence and seed maturation, as well as when plants encounter adverse environmental conditions, for example, nutrient starvation, drought or pathogens infection and so on. However, the transcriptional activation or repression mechanisms of ATG genes during these biological processes are largely unknown and need further study. In this review, we summarized the roles and the well-established transcriptional regulation network of ATG genes during plant growth, development and stress responses.

Key words: plant autophagy, ATG gene, transcriptional regulation, stress response, growth and development

李喜豹, 赖敏怡, 梁山, 王小菁, 高彩吉, 杨超. 植物细胞自噬基因的功能与转录调控机制. 植物学报, 2021, 56(2): 201-217.

Xibao Li, Minyi Lai, Shan Liang, Xiaojing Wang, Caiji Gao, Chao Yang. Function and Transcriptional Regulation of Autophagy-related Genes in Plants. Chinese Bulletin of Botany, 2021, 56(2): 201-217.

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物种	发育时期及组织	检测方法	自噬基因表达	参考文献
拟南芥 (Arabidopsis thaliana)	自然衰老的叶片(10%黄化的叶片)	Microarray	5个AtATG上调	Buchanan-Wollaston et al. 2005
	衰老期叶片(7-39天的叶片)	Microarray	28个AtATG上调	Breeze et al. 2011
	授粉后12-30天的种子	qRT-PCR	31个AtATG上调	Di Berardino et al. 2018
	授粉后的乳突细胞	RNA-Seq	9个AtATG上调	Ye et al. 2020
玉米 (Zea mays)	授粉后2-24天的种子	RNA-Seq	11个ZmATG上调	Li et al. 2015
	授粉后12-24天的胚乳		29个ZmATG上调
	授粉后0-20天的叶片		30个ZmATG上调
烟草 (Nicotiana tabacum)	授粉后4-20天的种子	qRT-PCR	27个NtATG上调	Zhou et al. 2015
烟草 (Nicotiana tabacum)	萌发0.25-4小时的花粉	qRT-PCR	29个NtATG上调	Zhao et al. 2020
蓖麻 (Ricinus communis)	授粉后20-50天的胚乳	RNA-Seq, qRT-PCR	34个RcATG上调	Han et al. 2020
	授粉后20-30天的种皮		31个RcATG上调
	种子萌发后1-10天的胚乳		34个RcATG上调

物种	发育时期及组织	检测方法	自噬基因表达	参考文献
拟南芥 (Arabidopsis thaliana)	自然衰老的叶片(10%黄化的叶片)	Microarray	5个AtATG上调	Buchanan-Wollaston et al. 2005
	衰老期叶片(7-39天的叶片)	Microarray	28个AtATG上调	Breeze et al. 2011
	授粉后12-30天的种子	qRT-PCR	31个AtATG上调	Di Berardino et al. 2018
	授粉后的乳突细胞	RNA-Seq	9个AtATG上调	Ye et al. 2020
玉米 (Zea mays)	授粉后2-24天的种子	RNA-Seq	11个ZmATG上调	Li et al. 2015
	授粉后12-24天的胚乳		29个ZmATG上调
	授粉后0-20天的叶片		30个ZmATG上调
烟草 (Nicotiana tabacum)	授粉后4-20天的种子	qRT-PCR	27个NtATG上调	Zhou et al. 2015
烟草 (Nicotiana tabacum)	萌发0.25-4小时的花粉	qRT-PCR	29个NtATG上调	Zhao et al. 2020
蓖麻 (Ricinus communis)	授粉后20-50天的胚乳	RNA-Seq, qRT-PCR	34个RcATG上调	Han et al. 2020
	授粉后20-30天的种皮		31个RcATG上调
	种子萌发后1-10天的胚乳		34个RcATG上调

物种	发育时期/组织	胁迫类型/时间	检测方法	自噬基因表达	参考文献
拟南芥 (Arabidopsis thaliana)	4周/叶	淹水胁迫/24小时	qRT-PCR	6个AtATG上调	Chen et al. 2015
	2周/幼苗	缺锌胁迫/14天		13个AtATG上调	Eguchi et al. 2017
	5天/幼苗	42°C热激后/24小时		33个AtATG上调	Sedaghatmehr et al. 2019
	5天/幼苗	光照到黑暗/24小时		11个AtATG上调	Yang et al. 2020a
	30天/茎叶	黑暗胁迫/3天	RNA-Seq	23个AtATG上调	Yan et al. 2019
	4周/叶	黑暗胁迫/6天	Microarray	14个AtATG上调	van der Graaff et al. 2006
水稻 (Oryza sativa)	3周/整株苗	250 mmol·L^-1 NaCl/4小时	qRT-PCR	13个OsATG上调	Xia et al. 2011
		干旱胁迫/4小时		13个OsATG上调
		4°C冷胁迫/4小时		21个OsATG上调
		黑暗胁迫/48小时		19个OsATG上调
		碳饥饿/48小时		17个OsATG上调
	10天/整株苗	氮饥饿/7天		18个OsATG上调
物种	发育时期/组织	胁迫类型/时间		检测方法	自噬基因表达	参考文献
烟草 (Nicotiana tabacum)	2周/整株苗	碳饥饿/48小时		qRT-PCR	17个NtATG上调	Zhou et al. 2015
		氮饥饿/48小时			16个NtATG上调
		黑暗胁迫/48小时			16个NtATG上调
		250 mmol·L^-1 NaCl/4小时			14个NtATG上调
		4°C冷胁迫/4小时			14个NtATG上调
		干旱胁迫/4小时			7个NtATG上调
		1 μmol ·L^-1 NAA/24小时			6个NtATG上调
		5 μmol ·L^-1 2,4-D/24小时			5个NtATG上调
		25 μmol ·L^-1 ABA/24小时			4个NtATG上调
		5 μmol ·L^-1 GA₃/24小时			7个NtATG下调
		500 μmol ·L^-1 SA/24小时			4个NtATG上调
		500 μmol ·L^-1 JA/24小时			11个NtATG上调
		20 μmol ·L^-1 Cu²⁺/24小时			12个NtATG上调
		20 μmol ·L^-1 Ni²⁺/24小时			15个NtATG上调
		40 μmol ·L^-1 Zn²⁺/24小时			13个NtATG上调
		40 μmol ·L^-1 Cd²⁺/24小时			11个NtATG上调
		100 μmol ·L^-1 Mn²⁺/24小时			14个NtATG上调
辣椒 (Capsicum annuum)	5-6叶期/根	200 mmol·L^-1 NaCl/3小时		RNA-Seq	9个CaATG上调	Zhai et al. 2016
	5-6叶期/茎	200 mmol·L^-1 NaCl/3小时			15个CaATG上调
	5-6叶期/根	干旱胁迫/3小时			7个CaATG上调
	5-6叶期/茎	干旱胁迫/3小时			7个CaATG上调
	5-6叶期/整株苗	40°C热胁迫/12小时 (耐热品种)			26个CaATG上调
		40°C热胁迫/12小时 (热敏感品种)			17个CaATG上调
		4°C冷胁迫/3小时			14个CaATG上调
		碳饥饿/2天			10个CaATG上调
小麦 (Triticum aestivum)	10天/叶	150 mmol·L^-1 NaCl/48小时		RNA-Seq	75个TaATG上调	Yue et al. 2018
		4°C冷胁迫/24小时			23个TaATG上调
		42°C热胁迫/6小时			36个TaATG上调
		干旱胁迫/24小时			23个TaATG上调
	2叶期/根-叶	氮饥饿/4天		qRT-PCR	9个TaATG8上调	Zhang et al. 2020
		白粉病真菌侵染/36小时(敏感品种)			9个TaATG8上调
		白粉病真菌侵染/36小时(抗性品种)			3个TaATG8上调
番茄 (Solanum lycopersicum)	6周/叶	干旱胁迫/6天		qRT-PCR	20个SlATG上调	Wang et al. 2015
	6周/叶	500 nmol·L^-1 BL/12小时			8个SlATG上调	Wang et al. 2019
	5周/叶	4°C冷胁迫/24小时			11个SlATG上调	Chi et al. 2020
香蕉 (Musa nana)	5叶期/根	镰刀菌侵染/51小时		qRT-PCR	8个MaATG上调	Wei et al. 2017
木薯 (Manihot esculenta)	30天/叶	单黄胞菌侵染/6小时		qRT-PCR	25个MeATG上调	Yan et al. 2017
葡萄 (Vitis vinifera)	2年/叶	400 μmol ·L^-1 Cu²⁺/36小时		qRT-PCR	24个VvATG上调	Shangguan et al. 2018
线形草沙蚕 (Tripogon loliiformis)	2月/根-茎	脱水处理		RNA-Seq	10个TlATG上调	Williams et al. 2015

物种	发育时期/组织	胁迫类型/时间	检测方法	自噬基因表达	参考文献
拟南芥 (Arabidopsis thaliana)	4周/叶	淹水胁迫/24小时	qRT-PCR	6个AtATG上调	Chen et al. 2015
	2周/幼苗	缺锌胁迫/14天		13个AtATG上调	Eguchi et al. 2017
	5天/幼苗	42°C热激后/24小时		33个AtATG上调	Sedaghatmehr et al. 2019
	5天/幼苗	光照到黑暗/24小时		11个AtATG上调	Yang et al. 2020a
	30天/茎叶	黑暗胁迫/3天	RNA-Seq	23个AtATG上调	Yan et al. 2019
	4周/叶	黑暗胁迫/6天	Microarray	14个AtATG上调	van der Graaff et al. 2006
水稻 (Oryza sativa)	3周/整株苗	250 mmol·L^-1 NaCl/4小时	qRT-PCR	13个OsATG上调	Xia et al. 2011
		干旱胁迫/4小时		13个OsATG上调
		4°C冷胁迫/4小时		21个OsATG上调
		黑暗胁迫/48小时		19个OsATG上调
		碳饥饿/48小时		17个OsATG上调
	10天/整株苗	氮饥饿/7天		18个OsATG上调
物种	发育时期/组织	胁迫类型/时间		检测方法	自噬基因表达	参考文献
烟草 (Nicotiana tabacum)	2周/整株苗	碳饥饿/48小时		qRT-PCR	17个NtATG上调	Zhou et al. 2015
		氮饥饿/48小时			16个NtATG上调
		黑暗胁迫/48小时			16个NtATG上调
		250 mmol·L^-1 NaCl/4小时			14个NtATG上调
		4°C冷胁迫/4小时			14个NtATG上调
		干旱胁迫/4小时			7个NtATG上调
		1 μmol ·L^-1 NAA/24小时			6个NtATG上调
		5 μmol ·L^-1 2,4-D/24小时			5个NtATG上调
		25 μmol ·L^-1 ABA/24小时			4个NtATG上调
		5 μmol ·L^-1 GA₃/24小时			7个NtATG下调
		500 μmol ·L^-1 SA/24小时			4个NtATG上调
		500 μmol ·L^-1 JA/24小时			11个NtATG上调
		20 μmol ·L^-1 Cu²⁺/24小时			12个NtATG上调
		20 μmol ·L^-1 Ni²⁺/24小时			15个NtATG上调
		40 μmol ·L^-1 Zn²⁺/24小时			13个NtATG上调
		40 μmol ·L^-1 Cd²⁺/24小时			11个NtATG上调
		100 μmol ·L^-1 Mn²⁺/24小时			14个NtATG上调
辣椒 (Capsicum annuum)	5-6叶期/根	200 mmol·L^-1 NaCl/3小时		RNA-Seq	9个CaATG上调	Zhai et al. 2016
	5-6叶期/茎	200 mmol·L^-1 NaCl/3小时			15个CaATG上调
	5-6叶期/根	干旱胁迫/3小时			7个CaATG上调
	5-6叶期/茎	干旱胁迫/3小时			7个CaATG上调
	5-6叶期/整株苗	40°C热胁迫/12小时 (耐热品种)			26个CaATG上调
		40°C热胁迫/12小时 (热敏感品种)			17个CaATG上调
		4°C冷胁迫/3小时			14个CaATG上调
		碳饥饿/2天			10个CaATG上调
小麦 (Triticum aestivum)	10天/叶	150 mmol·L^-1 NaCl/48小时		RNA-Seq	75个TaATG上调	Yue et al. 2018
		4°C冷胁迫/24小时			23个TaATG上调
		42°C热胁迫/6小时			36个TaATG上调
		干旱胁迫/24小时			23个TaATG上调
	2叶期/根-叶	氮饥饿/4天		qRT-PCR	9个TaATG8上调	Zhang et al. 2020
		白粉病真菌侵染/36小时(敏感品种)			9个TaATG8上调
		白粉病真菌侵染/36小时(抗性品种)			3个TaATG8上调
番茄 (Solanum lycopersicum)	6周/叶	干旱胁迫/6天		qRT-PCR	20个SlATG上调	Wang et al. 2015
	6周/叶	500 nmol·L^-1 BL/12小时			8个SlATG上调	Wang et al. 2019
	5周/叶	4°C冷胁迫/24小时			11个SlATG上调	Chi et al. 2020
香蕉 (Musa nana)	5叶期/根	镰刀菌侵染/51小时		qRT-PCR	8个MaATG上调	Wei et al. 2017
木薯 (Manihot esculenta)	30天/叶	单黄胞菌侵染/6小时		qRT-PCR	25个MeATG上调	Yan et al. 2017
葡萄 (Vitis vinifera)	2年/叶	400 μmol ·L^-1 Cu²⁺/36小时		qRT-PCR	24个VvATG上调	Shangguan et al. 2018
线形草沙蚕 (Tripogon loliiformis)	2月/根-茎	脱水处理		RNA-Seq	10个TlATG上调	Williams et al. 2015

植物细胞自噬基因的功能与转录调控机制

Function and Transcriptional Regulation of Autophagy-related Genes in Plants

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