蛋白质磷酸化修饰与种子休眠及萌发调控

doi:10.11983/CBB21011

植物学报 ›› 2021, Vol. 56 ›› Issue (4): 488-499.DOI: 10.11983/CBB21011

蛋白质磷酸化修饰与种子休眠及萌发调控

赵晓亭¹, 毛凯涛¹, 徐佳慧¹^,², 郑钏¹^,³, 罗晓峰¹, 舒凯¹^,^*()

¹西北工业大学生态环境学院, 西安 710129
²中国农业大学农学院, 北京 100193
³四川农业大学生态农业研究所, 成都 611130

收稿日期:2021-01-15 接受日期:2021-04-19 出版日期:2021-07-01 发布日期:2021-06-30
通讯作者: 舒凯
作者简介:*E-mail: kshu@nwpu.edu.cn
基金资助:
国家自然科学基金(31872804);国家自然科学基金(31701064)

Protein Phosphorylation and Its Regulatory Roles in Seed Dormancy and Germination

Xiaoting Zhao¹, Kaitao Mao¹, Jiahui Xu¹^,², Chuan Zheng¹^,³, Xiaofeng Luo¹, Kai Shu¹^,^*()

¹School of Ecology and Environment, Northwestern Polytechnical University, Xi'an 710129, China
²College of Agronomy and Biotechnology, China Agricultural University, Beijing 100193, China
³Institute of Ecological Agriculture, Sichuan Agricultural University, Chengdu 611130, China

Received:2021-01-15 Accepted:2021-04-19 Online:2021-07-01 Published:2021-06-30
Contact: Kai Shu

摘要/Abstract

摘要： 种子休眠与萌发是截然不同而又紧密联系的两个生理过程, 也是植物生命周期中的关键阶段, 对自然状态下的植物物种繁殖与地理分布以及农业生产均具有重要意义, 且两个过程受不同内源激素和环境信号之间的精确互作调控。大量研究表明, 蛋白质磷酸化修饰作为一种重要的翻译后修饰方式, 参与调控种子休眠与萌发以及植物逆境胁迫响应等过程并发挥重要作用。该文简要介绍了蛋白质磷酸化、去磷酸化修饰过程及其功能, 系统总结了蛋白质磷酸化修饰在种子休眠与萌发过程中的调控作用, 并展望了未来的研究方向。

关键词: 种子, 休眠, 萌发, 蛋白质磷酸化, 植物激素

Abstract: Seed dormancy and germination are two distinct but closely related physiological processes, which are also key stages during plant life-cycle and have great significance to agricultural production, plant species reproduction, and geographical distribution. These processes are precisely regulated by interactions between different endogenous phytohormones and environmental signals. A large number of studies have shown that protein phosphorylation, plays an important role in regulating seed dormancy and germination, as well as plant response to stresses. This review paper briefly introduces the procedures and functions of protein phosphorylation and dephosphorylation modification, and summarizes the regulatory roles of protein phosphorylation modification in seed dormancy and germination. Finally, some future research directions are prospected.

Key words: seed, dormancy, germination, protein phosphorylation, phytohormone

赵晓亭, 毛凯涛, 徐佳慧, 郑钏, 罗晓峰, 舒凯. 蛋白质磷酸化修饰与种子休眠及萌发调控. 植物学报, 2021, 56(4): 488-499.

Xiaoting Zhao, Kaitao Mao, Jiahui Xu, Chuan Zheng, Xiaofeng Luo, Kai Shu. Protein Phosphorylation and Its Regulatory Roles in Seed Dormancy and Germination. Chinese Bulletin of Botany, 2021, 56(4): 488-499.

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[99]	Zhang XJ, Yang GY, Shi R, Han XM, Qi LW, Wang RG, Xiong LM, Li GJ (2013). Arabidopsis cysteine-rich receptor-like kinase 45 functions in the responses to abscisic acid and abiotic stresses. Plant Physiol Biochem 67, 189-198. DOI URL
[100]	Zhao R, Sun HL, Mei C, Wang XJ, Yan L, Liu R, Zhang XF, Wang XF, Zhang DP (2011a). The Arabidopsis Ca²⁺-dependent protein kinase CPK12 negatively regulates abscisic acid signaling in seed germination and post-germination growth. New Phytol 192, 61-73. DOI PMID
[101]	Zhao R, Wang XF, Zhang DP (2011b). CPK12: a Ca²⁺- dependent protein kinase balancer in abscisic acid signaling. Plant Signal Behav 6, 1687-1690. DOI PMID
[102]	Zhu SY, Yu XC, Wang XJ, Zhao R, Li Y, Fan RC, Shang Y, Du SY, Wang XF, Wu FQ, Xu YH, Zhang XY, Zhang DP (2007). Two calcium-dependent protein kinases, CPK4 and CPK11, regulate abscisic acid signal transduction in Arabidopsis. Plant Cell 19, 3019-3036. DOI URL

酶	大类	关键基因	分子功能	参考文献
蛋白激酶	MAPKs	MPK8	通过与GA反应的转录因子TCP14相互作用磷酸化TCP14, 增强其转录活性, 调控从休眠到萌发的转换。	Zhang et al., 2019
		MKK1、 MPK6	参与ABA和糖调节的种子萌发, 通过增强ABA的合成与信号强度, 维持种子休眠, 抑制种子萌发。	Xing et al., 2009
		Raf10、 Raf11	过表达Raf10和Raf11, 使种子对ABA敏感性增强, ABI3和ABI5表达上调, 从而延缓种子萌发。	Lee et al., 2015
		MKK3	MKK3-A高表达促进小麦种子休眠释放, 且MKK3可以与Qsd2-AK相互作用, 调控大麦种子休眠。	Nakamura et al., 2016; Torada et al., 2016
	CPKs	CPK4、 CPK11	过表达CPK4和CPK11, 种子表现出对ABA敏感表型, 萌发受到明显抑制。	Zhu et al., 2007
		CPK12	CPK12通过与ABI2相互作用磷酸化ABI2, 并下调ABF1和ABF4表达, 负调控ABA信号, 促进种子萌发。	Zhao et al., 2011a, 2011b
		CPK32	CPK32通过与ABF4相互作用磷酸化ABF4, 增强其转录活性, 促进ABA信号转导, 抑制种子萌发。	Choi et al., 2005
	SnRKs	SnRK2.2、 SnRK2.3	ABA信号通路复合物PYLs-ABA-PP2C通过激活SnRK2.2和SnRK2.3激酶活性, 激活下游转录因子, 诱导种子对ABA的响应。	Finkelstein et al., 2008; Fujii and Zhu, 2009
		SAPK2	ABA受体PYL/RCAR5作用于SAPK2上游, 激活SAPK2激酶活性, 通过OREB1介导ABA信号转导, 激活ABRE启动子活性, 负调控种子萌发。	Kim et al., 2012
类受体激酶	RLKs	GRACE	编码膜蛋白, 在干种子中高表达, 其表达水平受ABA诱导, 进而延缓种子萌发, 维持种子休眠。	Wu et al., 2017
		RPK1	RPK1基因表达受ABA诱导, 通过正调控ABA信号传导, 抑制种子萌发。	Hong et al., 1997; Osakabe et al., 2005
		CRK28	CRK28能够上调ABI3和ABI5的表达, 使ABA反应增强, 削弱种子萌发。	Pelagio-Flores et al., 2019
		CRK45	过表达CRK45能够上调ABA合成及反应基因表达水平, 正调控ABA信号转导, 延缓种子萌发。	Zhang et al., 2013
		CARK1、 CARK6	CARK1和CARK6通过与ABA受体RCAR11-14相互作用, 使其磷酸化, 促进ABA信号转导, 最终抑制种子萌发。	Zhang et al., 2018; Wang et al., 2019
		OsLecRK	OsLecRK表达受萌发信号刺激而上调, 进而与OsADF相互作用, 并进一步转导萌发信号, 上调α-淀粉酶基因表达, 增强种子活力, 促进种子萌发。	Cheng et al., 2013
磷酸酶	PP2Cs	AHG1、 AHG3	DOG1与AHG1/AHG3相互作用, 形成PP2C磷酸酶复合物, 抑制其磷酸酶活性, 进而增强ABA信号, 促使种子休眠。	Née et al., 2017; Nishimura et al., 2018
		PP2C-a10	PP2C-a10与TaDOG1L1和TaDOG1L4互作, 促进小麦种子萌发; 也可与ABA反应基因(ABI3、ABI4、ABI5、EM1和EM6)互作, 降低其表达水平, 促进拟南芥种子萌发。	Yu et al., 2020
		ABI1、 ABI2	在ABA存在条件下, ABA与PYR1/PYL/RCAR受体结合, 抑制ABI1和ABI2活性, 激活SnRK2s的激酶活性, 并磷酸化下游转录因子, 使ABA 信号向下传递, 抑制种子萌发。	Merlot et al., 2001; Raghavendra et al., 2010
		RDO5	RDO5是种子特异性表达的休眠积极调控因子, 独立于ABA对种子休眠的调控, 通过抑制APUM9和APUM11的转录水平调节种子休眠。	Xiang et al., 2014
		FsPP2C1	过表达PP2C1种子萌发率高, 对ABA敏感性低, 且能够在不利的条件 (如甘露醇和盐)下萌发, 通过负调控ABA信号转导, 促进种子萌发。	González-García et al., 2003; Saavedra et al., 2010
		HON	HON通过上调GA合成和响应基因的表达, 下调ABA响应基因和GA分解基因表达, 激活GA信号, 抑制ABA信号, 使种子解除休眠, 向萌发过渡。	Kim et al., 2013
	脂质磷酸酶	LPP2	后熟可激活LPP2的转录活性, 使其表达上调, 抑制种子对ABA的敏感性, 使种子能够完成萌发。	Carrera et al., 2008; Barrero et al., 2009
	线粒体蛋白磷酸酶	SLP2	SLP2与AtMIA40互作, 形成AtSLP2-AtMIA40蛋白质复合体, 通过抑制GA相关过程负调控种子萌发。	Uhrig et al., 2017
	肌醇多聚磷酸1-磷酸酶	FRY1	FRY1突变使IP3大量积累, 导致ABA的诱导和内源RD29A及其它胁迫响应基因的表达显著增强, 负调控ABA和逆境信号, 促进种子萌发。	Xiong et al., 2001

酶	大类	关键基因	分子功能	参考文献
蛋白激酶	MAPKs	MPK8	通过与GA反应的转录因子TCP14相互作用磷酸化TCP14, 增强其转录活性, 调控从休眠到萌发的转换。	Zhang et al., 2019
		MKK1、 MPK6	参与ABA和糖调节的种子萌发, 通过增强ABA的合成与信号强度, 维持种子休眠, 抑制种子萌发。	Xing et al., 2009
		Raf10、 Raf11	过表达Raf10和Raf11, 使种子对ABA敏感性增强, ABI3和ABI5表达上调, 从而延缓种子萌发。	Lee et al., 2015
		MKK3	MKK3-A高表达促进小麦种子休眠释放, 且MKK3可以与Qsd2-AK相互作用, 调控大麦种子休眠。	Nakamura et al., 2016; Torada et al., 2016
	CPKs	CPK4、 CPK11	过表达CPK4和CPK11, 种子表现出对ABA敏感表型, 萌发受到明显抑制。	Zhu et al., 2007
		CPK12	CPK12通过与ABI2相互作用磷酸化ABI2, 并下调ABF1和ABF4表达, 负调控ABA信号, 促进种子萌发。	Zhao et al., 2011a, 2011b
		CPK32	CPK32通过与ABF4相互作用磷酸化ABF4, 增强其转录活性, 促进ABA信号转导, 抑制种子萌发。	Choi et al., 2005
	SnRKs	SnRK2.2、 SnRK2.3	ABA信号通路复合物PYLs-ABA-PP2C通过激活SnRK2.2和SnRK2.3激酶活性, 激活下游转录因子, 诱导种子对ABA的响应。	Finkelstein et al., 2008; Fujii and Zhu, 2009
		SAPK2	ABA受体PYL/RCAR5作用于SAPK2上游, 激活SAPK2激酶活性, 通过OREB1介导ABA信号转导, 激活ABRE启动子活性, 负调控种子萌发。	Kim et al., 2012
类受体激酶	RLKs	GRACE	编码膜蛋白, 在干种子中高表达, 其表达水平受ABA诱导, 进而延缓种子萌发, 维持种子休眠。	Wu et al., 2017
		RPK1	RPK1基因表达受ABA诱导, 通过正调控ABA信号传导, 抑制种子萌发。	Hong et al., 1997; Osakabe et al., 2005
		CRK28	CRK28能够上调ABI3和ABI5的表达, 使ABA反应增强, 削弱种子萌发。	Pelagio-Flores et al., 2019
		CRK45	过表达CRK45能够上调ABA合成及反应基因表达水平, 正调控ABA信号转导, 延缓种子萌发。	Zhang et al., 2013
		CARK1、 CARK6	CARK1和CARK6通过与ABA受体RCAR11-14相互作用, 使其磷酸化, 促进ABA信号转导, 最终抑制种子萌发。	Zhang et al., 2018; Wang et al., 2019
		OsLecRK	OsLecRK表达受萌发信号刺激而上调, 进而与OsADF相互作用, 并进一步转导萌发信号, 上调α-淀粉酶基因表达, 增强种子活力, 促进种子萌发。	Cheng et al., 2013
磷酸酶	PP2Cs	AHG1、 AHG3	DOG1与AHG1/AHG3相互作用, 形成PP2C磷酸酶复合物, 抑制其磷酸酶活性, 进而增强ABA信号, 促使种子休眠。	Née et al., 2017; Nishimura et al., 2018
		PP2C-a10	PP2C-a10与TaDOG1L1和TaDOG1L4互作, 促进小麦种子萌发; 也可与ABA反应基因(ABI3、ABI4、ABI5、EM1和EM6)互作, 降低其表达水平, 促进拟南芥种子萌发。	Yu et al., 2020
		ABI1、 ABI2	在ABA存在条件下, ABA与PYR1/PYL/RCAR受体结合, 抑制ABI1和ABI2活性, 激活SnRK2s的激酶活性, 并磷酸化下游转录因子, 使ABA 信号向下传递, 抑制种子萌发。	Merlot et al., 2001; Raghavendra et al., 2010
		RDO5	RDO5是种子特异性表达的休眠积极调控因子, 独立于ABA对种子休眠的调控, 通过抑制APUM9和APUM11的转录水平调节种子休眠。	Xiang et al., 2014
		FsPP2C1	过表达PP2C1种子萌发率高, 对ABA敏感性低, 且能够在不利的条件 (如甘露醇和盐)下萌发, 通过负调控ABA信号转导, 促进种子萌发。	González-García et al., 2003; Saavedra et al., 2010
		HON	HON通过上调GA合成和响应基因的表达, 下调ABA响应基因和GA分解基因表达, 激活GA信号, 抑制ABA信号, 使种子解除休眠, 向萌发过渡。	Kim et al., 2013
	脂质磷酸酶	LPP2	后熟可激活LPP2的转录活性, 使其表达上调, 抑制种子对ABA的敏感性, 使种子能够完成萌发。	Carrera et al., 2008; Barrero et al., 2009
	线粒体蛋白磷酸酶	SLP2	SLP2与AtMIA40互作, 形成AtSLP2-AtMIA40蛋白质复合体, 通过抑制GA相关过程负调控种子萌发。	Uhrig et al., 2017
	肌醇多聚磷酸1-磷酸酶	FRY1	FRY1突变使IP3大量积累, 导致ABA的诱导和内源RD29A及其它胁迫响应基因的表达显著增强, 负调控ABA和逆境信号, 促进种子萌发。	Xiong et al., 2001

蛋白质磷酸化修饰与种子休眠及萌发调控

Protein Phosphorylation and Its Regulatory Roles in Seed Dormancy and Germination

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