MBF1调控植物热应答与生长发育分子机制研究进展

doi:10.11983/CBB21220

植物学报 ›› 2022, Vol. 57 ›› Issue (1): 56-68.DOI: 10.11983/CBB21220

MBF1调控植物热应答与生长发育分子机制研究进展

秦怡¹, 刘艳爽¹^,², 仇柳柳¹, 周敏¹, 杜小杉¹, 戴绍军¹^,^*(), 孙美红¹^,^*()

¹上海师范大学生命科学学院, 上海植物种质资源工程技术研究中心, 上海 200234
²东北林业大学生命科学学院, 东北盐碱植被恢复与重建教育部重点实验室, 哈尔滨 150040

收稿日期:2021-12-16 接受日期:2022-01-18 出版日期:2022-01-01 发布日期:2022-01-19
通讯作者: 戴绍军,孙美红
作者简介:sunmeihong@shnu.edu.cn
* E-mail: daishaojun@shnu.edu.cn;
基金资助:
国家自然科学基金(31902012);国家自然科学基金(32070300);上海市科委科研计划(YDZX20203100003927);上海市科委科研计划(19YF1436500)

Advance in Molecular Mechanism of MBF1 Regulating Plant Heat Response and Development

Yi Qin¹, Yanshuang Liu¹^,², Liuliu Qiu¹, Min Zhou¹, Xiaoshan Du¹, Shaojun Dai¹^,^*(), Meihong Sun¹^,^*()

¹Shanghai Plant Germplasm Resources Engineering Technology Research Center, College of Life Sciences, Shanghai Normal University, Shanghai 200234, China
²Key Laboratory of Saline-alkali Vegetation Ecology Restoration (Northeast Forestry University), Ministry of Education, College of Life Sciences, Northeast Forestry University, Harbin 150040, China

Received:2021-12-16 Accepted:2022-01-18 Online:2022-01-01 Published:2022-01-19
Contact: Shaojun Dai,Meihong Sun

摘要/Abstract

摘要： MBF1是一种进化上高度保守的转录共激活因子, 存在于所有真核生物中, 可通过连接基础转录机器组分与转录因子来激活基因转录。植物MBF1具有多种重要生物学功能, 包括调控植物生长发育和逆境适应等。该文综述了植物MBF1分子结构与调控机制相关研究进展, 重点总结了AtMBF1c参与植物热胁迫应答调控的分子机制。

关键词: 热胁迫, MBF1, 植物, 转录调控

Abstract: MBF1 (multiprotein bridging factor 1) is a transcriptional co-activator, which is highly conserved in evolution. It exists in all eukaryotes and can activate gene transcription by connecting components of the basal transcription machinery and transcription factors. Plant MBF1 plays important roles in a variety of biological processes, including controlling plant growth and development, and adversity adaptation. This review demonstrates the advance in the molecular structure and regulation mechanism of plant MBF1, and highlights the molecular mechanism of AtMBF1c in the regulation of plant heat stress response.

Key words: heat stress, MBF1, plant, transcriptional regulation

秦怡, 刘艳爽, 仇柳柳, 周敏, 杜小杉, 戴绍军, 孙美红. MBF1调控植物热应答与生长发育分子机制研究进展. 植物学报, 2022, 57(1): 56-68.

Yi Qin, Yanshuang Liu, Liuliu Qiu, Min Zhou, Xiaoshan Du, Shaojun Dai, Meihong Sun. Advance in Molecular Mechanism of MBF1 Regulating Plant Heat Response and Development. Chinese Bulletin of Botany, 2022, 57(1): 56-68.

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链接本文: https://www.chinbullbotany.com/CN/10.11983/CBB21220

https://www.chinbullbotany.com/CN/Y2022/V57/I1/56

图/表 4

表1 不同植物物种中MBF1蛋白家族成员

Table 1 Members of the MBF1 protein family in different plant species

物种	拉丁名	蛋白名	蛋白编号(JGI数据库)	参考文献
拟南芥	Arabidopsis thaliana	AtMBF1a	AT2G42680.1	Tsuda et al., 2004
		AtMBF1b	AT3G58680.1
		AtMBF1c	AT3G24500.1
番茄	Solanum lycopersicum	SlMBF1a	Solyc10g007350.3.1	Zhang et al., 2019
		SlMBF1b	Solyc12g014290.2.1
		SlMBF1c	Solyc07g062400.3.1
		SlMBF1d	Solyc09g055470.1.1
		SlER24	Solyc01g104740.3.1
水稻	Oryza sativa	OsMBF1a	LOC_Os08g27850.1	Zhang et al., 2019
水稻	Oryza sativa	OsMBF1c	LOC_Os06g39240.1	Zhang et al., 2019
菠菜	Spinacia oleracea	SoMBF1b	Spov3_C0009.00073	Xu et al., 2017
菠菜	Spinacia oleracea	SoMBF1c	Spov3_C0062.00022	Xu et al., 2017
马铃薯	S. tuberosum	StMBF1a	PGSC0003DMP400012592	Yu et al., 2021
		StMBF1b	PGSC0003DMP400026892
		StMBF1c	PGSC0003DMP400051869
		StMBF1d	PGSC0003DMP400051868
小麦	Triticum aestivum	TaMBF1a-1	Traes_2AL_A9D390619.1	Qin et al., 2015; Tian et al., 2021
		TaMBF1a-2	Traes_2BL_4F31B5695.1
		TaMBF1a-3	Traes_2DL_D4AB94C53.1
		TaMBF1a-4	Traes_3AS_719D37CCA.1
		TaMBF1a-5	Traes_3B_EC2B74116.1
		TaMBF1b	Traes_4DS_F1C77E7B0.1
		TaMBF1c-7A	Traes_7AL_FA77CC1F41.1
		TaMBF1c-7B	Traes_7BL_A002364C5.1
		TaMBF1c-7C	Traes_7DL_D5AD8EB4B.1
葡萄	Vitis vinifera	VvMBF1a	VIT_212s0028g02020.1	Yan et al., 2014
		VvMBF1a-like	VIT_219s0014g01260.1
		VvMBF1c	VIT_211s0016g04080.1
大豆	Glycine max	GmMBF1a-1	Glyma.06G276200.1.p	Tsuda et al., 2004
		GmMBF1a-2	Glyma.06G276300.1.p
		GmMBF1a-3	Glyma.12G129100.1.p
玉米	Zea mays	ZmMBF1a	ZmPHB47.01G345100.1.p	Tsuda et al., 2004
		ZmMBF1b	ZmPHB47.04G054300.1.p
		ZmMBF1c	ZmPHB47.09G119100.1.p
蓖麻	Ricinus communis	RcMBF1b	27894.m000799	Tsuda et al., 2004
蓖麻	Ricinus communis	RcMBF1c	29912.m005549	Tsuda et al., 2004
蒺藜苜蓿	Medicago truncatula	MtMBF1b-1	Medtr2g084220.1	Tsuda et al., 2004
		MtMBF1b-2	Medtr4g080090.1
		MtMBF1b-3	Medtr6g018330.1
		MtMBF1c	Medtr6g086280.1

图1 植物MBF1系统发育关系、基因结构和保守基序分析 (A) 植物MBF1的系统发育关系、基因结构和保守基序分布图。11个物种MBF1蛋白的全长蛋白质序列来自JGI数据库(https:// phytozome-next.jgi.doe.gov/) (表1)。用MEGA7.0软件基于邻接法构建同源系统进化树, 绿色背景表示I类MBF1蛋白亚家族, 红色背景表示II类MBF1蛋白亚家族。利用GSDS2.0软件(http://gsds.cbi.pku.edu.cn/)在线绘制基因结构图, 上游5'非编码区/下游3'非编码区用蓝色方框表示, 外显子用黄色方框表示, 内含子用黑色直线显示。使用MEME网站(https://meme-suite.org/meme/ tools/meme)在线预测保守基序。通过NCBI网站CDD数据库(https://www.ncbi.nlm.nih.gov/cdd)鉴定蛋白质序列的保守结构域及功能。利用Tbtools软件绘制保守基序分布图, 5个保守基序用不同颜色方框表示。(B) 11个物种MBF1蛋白中预测到的保守基序序列。

Figure 1 The analysis of the phylogenetic relationships, gene structures and conserved motifs of plant MBF1 (A) Phylogenetic relationship, gene structure and conserved motif distribution map of plant MBF1. The full-length protein sequences of MBF1 proteins from 11 species are downloaded from the JGI database (https://phytozome-next.jgi.doe.gov/) (table 1). MEGA7.0 software is used to construct a Neighbor-Joining homologous phylogenetic tree. The green background indicates the type I subfamily, and the red background indicates the type II subfamily. GSDS2.0 software (http://gsds.cbi.pku.edu.cn/) is used to draw a gene structure diagram online. 5'UTR/3'UTR are represented by blue boxes, exons are represented by yellow boxes, and introns are represented by black straight lines. The MEME website (https://meme-suite.org/meme/tools/meme) is used to predict conserved motifs. Conserved domains and functions are identified through the NCBI website CDD database (https://www.ncbi.nlm.nih.gov/cdd). Tbtools software is used to draw a map of the conserved motifs, and the 5 conserved motifs are indicated by boxes with different colors. (B) The predicted conserved motif sequences in MBF1 proteins of 11 species.

图2 不同物种MBF1s同源序列比对分析家蚕BmMBF1和酿酒酵母yMBF1蛋白质序列从Ensembl数据库(https://ensemblgenomes.org/)下载, 蛋白编号分别为BGIBMGA 007702和YOR298C-A。利用DNAMAN软件对蛋白序列进行比对。紫色框表示N端MBF1结构域, 蓝色框表示C端HTH_3结构域。拟南芥和水稻MBF1蛋白质序列及蛋白编号见表1。

Figure 2 Homologous sequence alignment analysis of MBF1s of different species The protein sequences of Bombyx mori BmMBF1 and Saccharomyces cerevisiae yMBF1 are downloaded from Ensembl database (https://ensemblgenomes.org/), the corresponding protein accession number is BGIBMGA007702 and YOR298C-A. DNAMAN software is used for sequence alignment. The purple box indicates the N-terminal MBF1 domain, and the blue box indicates the C-terminal HTH_3 domain. The protein sequences and protein accession number of Arabidopsis and rice MBF1 are listed in Table 1.

图3 MBF1调控热胁迫应答信号通路热胁迫通过激活质膜CNGC2引起Ca2+内流, 通过激活质膜结合的RBOHD导致ROS积累。而Ca2+信号和ROS信号通过未知途径激活AtMBF1c及其下游靶基因调节的热胁迫应答。热激转录因子HSFA1与HSFA2相互作用, 直接调控AtMBF1c、HSFA2、HSFBs和DREB2A的表达。同时, AtMBF1c通过与DREB2A、HSFB2A和HSFB2B的启动子HSE元件结合, 调控其基因表达, 提高热胁迫耐受性。DREB2A与DPB3-1、NF-Y A2和NF-Y B3形成的三聚体共激活复合物互作, 增强其对下游靶基因HSFA3的转录激活, 提高植物的耐热性。DREB2A也促进HSP70、HSP18.2和At1g52560的表达, 增强植株耐热性。AtSAP5作为上游调控因子与AtMBF1c发生互作并激活AtMBF1c, 调节细胞核中HSP18.2的表达, 提高植物耐热性。热胁迫诱导TPS5的表达, AtMBF1c与TPS5互作, 通过促进海藻糖的合成和积累提高耐热性。热胁迫诱导AtWRKY39、AtWRKY25、AtWRKY26和AtWRKY33的表达。AtWRKY39通过AtMBF1c调节水杨酸(SA)信号通路下游PR1基因表达, 提高耐热性。AtWRKY25、AtWRKY26和AtWRKY33通过AtMBF1c调节乙烯信号通路下游基因的表达提高耐热性, 同时通过促进HSP70、HSP101、HSFA2和HSFB1的表达提高耐热性。CNGC2: 环核苷酸门控离子通道2; DPB3-1 (NF-YC10): DNA聚合酶II亚基B3-1; DRE: 脱水应答元件; DREB2A: 干旱应答元件结合蛋白2A; HSE: 热休克响应元件; HSFA1/2/3: 热激转录因子A1/2/3; HSFB2A/B: 热激转录因子B2A/B; HSP18.2/70/101: 热激蛋白18.2/70/101; MBF1c: 多蛋白桥梁因子1c; NF-Y A2/B3: 核因子-Y A2/B3; PR1: 病程相关因子1; RBOHD: 呼吸爆发氧化同源蛋白D; ROS: 活性氧; SAP5: 胁迫相关蛋白5; TPS5: 海藻糖磷酸合成酶5。蓝色实线箭头表示蛋白互作; 绿色实线箭头表示基因编码蛋白; 黑色实线箭头表示直接转录激活; 黑色虚线箭头表示间接转录激活。

Figure 3 Signaling pathway of MBF1 regulating heat stress response Heat stress causes Ca2+ influx by activating the plasma membrane-localized protein CNGC2, and leads to the accumulation of ROS by activating the plasma membrane-bound RBOHD. The Ca2+ signal and ROS signal activate the heat stress response by regulating AtMBF1c and its downstream target genes through unknown pathways. The heat stress transcription factor HSFA1 interacts with HSFA2 and directly regulates the expression of AtMBF1c, HSFA2, HSFBs and DREB2A. At the same time, AtMBF1c binds to the HSE elements of DREB2A, HSFB2A and HSFB2B promoters to regulate their gene expression and improve heat stress tolerance. DREB2A interacts with the trimeric co-activation complex formed by DPB3-1, NF-Y A2 and NF-Y B3 to enhance the transcriptional activation of the downstream target gene HSFA3 and improve plant heat tolerance. DREB2A also promotes the expression of HSP70, HSP18.2 and At1g52560 to enhance plant heat tolerance. As the upstream regulator, AtSAP5 interacts with and activates AtMBF1c in the nucleus, regulating the expression of HSP18.2 and improving plant heat tolerance. Heat stress induces the expression of TPS5. AtMBF1c interacts with TPS5 to improve heat tolerance by promoting the synthesis and accumulation of trehalose. Heat stress induces the expression of AtWRKY39, AtWRKY25, AtWRKY26 and AtWRKY33. AtWRKY39 regulates the expression of the downstream gene of salicylic acid (SA) signaling pathway PR1 through AtMBF1c to improve heat tolerance. AtWRKY25, AtWRKY26 and AtWRKY33 regulate the expression of downstream genes in the ethylene (ET) signaling pathway through AtMBF1c to improve heat resistance, and at the same time promote the expression of HSP70, HSP101, HSFA2, and HSFB1. CNGC2: Cyclic nucleotide-gated channel 2; DPB3-1 (NF-YC10): DNA polymerase II subunit B3-1; DRE: Dehydration-responsive element; DREB2A: Dehydration responsive element-binding protein 2A; HSE: Heat shock elements; HSFA1/2/3: Heat stress transcription factor A1/2/3; HSFB2A/B: Heat stress transcription factor B2A/B; HSP18.2/ 70/101: Heat shock protein 18.2/70/101; MBF1c: Multiprotein bridging factor 1c; NF-Y A2/B3: Nuclear factor Y A2/B3; PR1: Pathogenesis-related factor 1; RBOHD: Respiratory burst oxidase homologue D; ROS: Reactive oxygen species; SAP5: Stress-associated protein 5; TPS5: Trehalose phosphate synthetase 5. The blue solid arrow indicates protein interaction; the green solid arrow indicates gene encoding protein; the black solid arrow indicates direct transcription activation; the black dashed arrow indicates indirect transcription activation.

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MBF1调控植物热应答与生长发育分子机制研究进展

Advance in Molecular Mechanism of MBF1 Regulating Plant Heat Response and Development

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