植物线粒体铁硫簇合成系统及其调控的研究进展

doi:10.11983/CBB24103

植物学报 ›› 2025, Vol. 60 ›› Issue (4): 499-514.DOI: 10.11983/CBB24103 cstr: 32102.14.CBB24103

植物线粒体铁硫簇合成系统及其调控的研究进展

谢涛, 章一帆, 刘云辉, 游慧玉, 夏季奔奔, 马蓉, 张春妮, 华学军^*()

浙江理工大学, 生命科学与医药学院, 杭州 310018

收稿日期:2024-07-13 接受日期:2024-10-14 出版日期:2025-07-10 发布日期:2024-10-16
通讯作者: *华学军, 教授, 分别于北京大学、中国农业科学院和比利时根特大学获学士、硕士和博士学位; 经历加拿大多伦多大学博士后, 加拿大卡尔加里大学高级研究助理, 2005年入选中国科学院“百人计划”, 2018年聘为浙江理工大学生医学院教授, 专注于植物发育与逆境应答领域科学研究。主持和参加多项国家自然科学基金面上项目、973项目、863项目、转基因专项、科技部国际合作项目等。在Proc Natl Acad Sci USA、PLoS Genetics、Plant Physiology、Plant Journal、Plant Biotechnology Journal、Plant Physiology & Biochemistry和Plant & Cell Physiology等国际知名学术期刊发表论文80余篇。授权国家发明专利5项。培养硕士和博士30余名。
植物抗逆分子机制研究团队聚焦拟南芥和番茄等逆境生物学, 利用遗传学、表观遗传学和生理生化等手段, 阐明植物响应非生物胁迫(干旱和高盐等)的分子机制与遗传调控网络, 并探索其在提高作物抗逆能力方面的应用。重点挖掘胁迫响应基因(如转录因子、转运蛋白)的功能验证与通路解析; 深入解析ROS和钙信号等介导的胁迫感知与传递机制, 尤其是在基因转录对盐胁迫记忆的表观遗传学机理、线粒体功能稳态及铁硫簇合成代谢对逆境胁迫响应等方面取得重大研究成果。E-mail: xjhua@zstu.edu.cn
基金资助:
国家自然科学基金(32170307)

Research Progress on the Iron-sulfur Cluster Synthesis System and Regulation in Plant Mitochondria

Tao Xie, Yifan Zhang, Yunhui Liu, Huiyu You, Jibenben Xia, Rong Ma, Chunni Zhang, Xuejun Hua^*()

School of Life Sciences and Medicine, Zhejiang Sci-Tech University, Hangzhou 310018, China

Received:2024-07-13 Accepted:2024-10-14 Online:2025-07-10 Published:2024-10-16
Contact: *E-mail: xjhua@zstu.edu.cn

摘要/Abstract

摘要： 铁硫[Fe-S]簇作为铁硫蛋白的辅助因子广泛参与光合作用、呼吸作用和电子传递等多种生物学过程, 并参与合成一些必需的维生素和辅助因子等。其在细胞内的生物合成受到一系列蛋白的催化和调控, 并被区隔在不同的亚细胞结构中。线粒体是细胞能量代谢的主要场所, 其中许多关键代谢酶是铁硫蛋白, 需要线粒体铁硫簇组装系统ISC (iron-sulfur cluster)提供铁硫簇。目前, 得益于细菌和酵母中的相关研究成果, 植物线粒体ISC系统中的重要催化和调控蛋白的鉴定与功能分析, 以及铁硫簇在植物生长发育中的功能研究取得了长足进步。对铁硫簇合成系统中植物特有组分的发掘与鉴定, 以及铁硫簇合成系统响应环境胁迫的机制也日益引起人们的重视。该文对植物铁硫簇合成机制特别是线粒体ISC合成系统的研究进展进行总结, 同时对ISC合成系统的重要基因在植物生长发育和非生物胁迫响应中的作用进行了简要综述。

关键词: 铁硫簇, 线粒体, ISC系统, 植物, 胁迫响应

Abstract: Iron-sulfur [Fe-S] clusters, which act as cofactors for iron-sulfur proteins, are ubiquitously implicated in a diverse array of biological processes, such as photosynthesis, respiration, electron transport, and the biosynthesis of essential vitamins and cofactors. Their intracellular biogenesis is modulated by a suite of proteins that catalyze and regulate the process, with compartmentalization occurring within discrete subcellular compartments. Mitochondria, as the central organelles for cellular energy metabolism, harbor numerous key metabolic enzymes that are iron-sulfur proteins, necessitating the provision of iron-sulfur clusters by the mitochondrial assembly machinery, the iron-sulfur cluster (ISC). Advancements in research, particularly from bacteria and yeast systems, have facilitated significant strides in the identification and functional characterization of pivotal catalytic and regulatory proteins within the plant mitochondrial ISC system. Additionally, considerable progress has been made in the research of the role in iron-sulfur clusters in the plant growth and development. The elucidation of plant-specific components within the iron-sulfur cluster synthesis machinery and the mechanisms by which this system responds to environmental stress are areas of growing interest. This manuscript provides a comprehensive review of the current state of research on the mechanism of iron-sulfur cluster synthesis in plants, with a particular focus on the mitochondrial ISC assembly system. It also provides a succinct synopsis of the role of key genes within the ISC system in plant growth and development, as well as their involvement in the response to abiotic stressors.

Key words: iron-sulfur clusters, mitochondria, ISC system, plants, stress response

谢涛, 章一帆, 刘云辉, 游慧玉, 夏季奔奔, 马蓉, 张春妮, 华学军. 植物线粒体铁硫簇合成系统及其调控的研究进展. 植物学报, 2025, 60(4): 499-514.

Tao Xie, Yifan Zhang, Yunhui Liu, Huiyu You, Jibenben Xia, Rong Ma, Chunni Zhang, Xuejun Hua. Research Progress on the Iron-sulfur Cluster Synthesis System and Regulation in Plant Mitochondria. Chinese Bulletin of Botany, 2025, 60(4): 499-514.

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链接本文: https://www.chinbullbotany.com/CN/10.11983/CBB24103

https://www.chinbullbotany.com/CN/Y2025/V60/I4/499

图/表 4

图1 铁硫簇的基本结构(参考Lu, 2018) (A) 由4个半胱氨酸连接的[2Fe-2S]型铁硫簇; (B) 由3个半胱氨酸和1个组氨酸连接的NEET类[2Fe-2S]型铁硫簇; (C) 由2个半胱氨酸和2个组氨酸连接的Rieske类[2Fe-2S]簇; (D) 由3个半胱氨酸残基连接的[3Fe-4S]簇; (E) 由4个半胱氨酸残基连接的[4Fe-4S]簇; (F) 由4个半胱氨酸残基配位的[4Fe-4S]簇, 可以硫醇配位键连接血红素。

Figure 1 The basic structure of iron-sulfur clusters (adapted from Lu, 2018) (A) [2Fe-2S] type iron-sulfur clusters linked by 4 cysteines; (B) NEET [2Fe-2S] iron-sulfur clusters connected by three cysteines and one histidine; (C) Rieske [2Fe-2S] connected by two cysteines and two histidine; (D) [3Fe-4S] connected by three cysteine residues; (E) [4Fe-4S] linked by four cysteine residues; (F) [4Fe-4S] coordinated by four cysteine residues, can bind heme with mercaptan coordination bonds.

图2 拟南芥线粒体ISC合成途径模式图(参考Lill, 2009) 模式图参考自酵母线粒体ISC合成途径模型, 红色方框中的组分在拟南芥中尚未见文献报道。第1步中AtNfs1-AtIsd11复合物将半胱氨酸脱硫, Frataxin可能提供铁。未被鉴定的同源铁转运蛋白Mrs3和Mrs4可将胞质中的铁转运至线粒体与Frataxin结合, AtMFDR和AtMFDX可能帮助NADPH转移电子。铁和硫在AtIsu1上合成铁硫簇。第2步, AtHscA2和AtHscB与AtIsu1结合, 并且促进铁硫簇从AtIsu1上释放。第3步是将铁硫簇运送至不同的铁硫蛋白中, 如GrxS15、线粒体铁硫蛋白、顺乌头酸酶、生物合成酶以及电子传递链复合物I、II和III。部分铁硫蛋白转运需要铁硫簇组装因子1、组装因子2 (AtIscA1和AtIscA2)和组装因子AtIBA57参与。

Figure 2 Hypothetical model of the mitochondrial ISC biosynthetic pathway in Arabidopsis thaliana (adapted from Lill, 2009) The schematic diagram refers to the yeast mitochondrial ISC synthesis pathway model, and the components in the red box have not yet been reported in Arabidopsis. In the first step, the AtNfs1-AtIsd11 complex desulfurates cysteine, and Frataxin may provide iron. The unidentified homologous iron transport proteins Mrs3 and Mrs4 can transport iron from the cytoplasm to the mitochondria where it binds with Frataxin, whereas AtMFDR and AtMFDX may assist in electron transfer by NADPH. Iron and sulfur are synthesized into iron-sulfur clusters on AtIsu1. In the second step, AtHscA2 and AtHscB bind to AtIsu1, promoting the release of iron-sulfur clusters from AtIsu1. The third step involves transporting the iron-sulfur clusters to various iron-sulfur proteins, such as GrxS15, mitochondrial iron-sulfur proteins, fumarase, biosynthetic enzymes, and electron transport chain complexes I, II, and III. The transport of some iron-sulfur proteins requires the participation of iron-sulfur cluster assembly factors 1 and 2 (AtIscA1 and AtIscA2), and the assembly factor AtIBA57.

表1 拟南芥中参与线粒体[Fe-S]簇组装基因的突变等位基因的表型

Table 1 Phenotypes of mutant alleles involved in mitochondrial [Fe-S] cluster assembly genes in Arabidopsis thaliana

基因	突变类型	生长表型	参考文献
Frataxin	SALK_021263	生长迟缓、果实鲜重降低且种子数减少	Busi et al., 2006
	SALK_094203	部分种子不育	Busi et al., 2004
	SALK_122008	部分种子不育, 毛绒根	Martin et al., 2009
ISU1	过表达	提升铁的富集能力	Song et al., 2022
	RNAi	茎细, 植株矮小	Frazzon et al., 2007
	SALK_006332	植株矮小	Frazzon et al., 2007
NFS1	过表达	新叶扇形边缘、杂乱花序、莲座叶和尾状叶腋下的腋芽增多, 对病菌的抗性增强	Frazzon et al., 2007 Fonseca et al., 2020
HscB	SALK_099684	根的铁吸收水平降低, 铁在枝叶中积累	Leaden et al., 2016
HscB	SALK_085159	茎部显示出无蜡状表型, 具有光滑的外表	Xu et al., 2009
HscA1	SALK_081383、SALK_081385、 SALK_128982和SALK_140494	茎短, 莲座叶变小	Wei et al., 2019
HscA2	SAIL_354_E09、SAIL_302_G07和 HscA2m点突变	无明显表型, 对外源脯氨酸具有抗性	Wei et al., 2019; Zhang et al., 2025
GrxS15	SALK_112767、GrxS15^amiR、 SALK_112767C、GK-837C05 和SAIL_431_H03	根短, 莲座叶变小, 根尖呼吸速率降低, 对砷胁迫的抗性增强, 早期胚胎致死	Moseler et al., 2015; Ströher et al., 2016
SSR1	FLAG_356A08和FLAG_571A02	根生长受到抑制	Feng et al., 2025

图3 拟南芥ISC合成途径相关基因在不同非生物胁迫下的表达热图不同非生物胁迫下线粒体铁硫蛋白基因的表达热图, 数据来源于Arabidopsis eFP Browser (https://bar.utoronto.ca/efp/cgi-bin/ efpWeb.cgi)。颜色标记表示基因相对对照组的倍数关系。图中展示胁迫处理时间均为24小时。

Figure 3 Heatmap of gene expression related to the ISC synthesis pathway in Arabidopsis thaliana under various abiotic stresses Heatmap of mitochondrial iron-sulfur protein gene expression under various abiotic stresses, with data sourced from the Arabidopsis eFP Browser (https://bar.utoronto.ca/efp/cgi-bin/efpWeb.cgi). The color markings indicate the fold change in gene expression relative to that in the control group. The time of stress treatment shown in the graph is 24 h for all the samples.

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植物线粒体铁硫簇合成系统及其调控的研究进展

Research Progress on the Iron-sulfur Cluster Synthesis System and Regulation in Plant Mitochondria

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