质膜Na+/H+逆向转运蛋白SOS1在植物离子稳态平衡中的作用

doi:10.13560/j.cnki.biotech.bull.1985.2023-0793

摘要/Abstract

摘要：

植物通过一系列复杂转运系统调节离子稳态以适应盐渍环境，SOS（salt overly sensitive）信号通路是植物响应非生物胁迫的主要信号途径，主要由质膜Na⁺/H⁺逆向转运蛋白SOS1、丝氨酸/苏氨酸类蛋白激酶SOS2和钙感应器SOS3组成。SOS1作为SOS信号通路的主要成员之一，广泛存在于高等植物中，由于早期的进化差异，可能导致不同物种SOS1的结构和理化性质存在一定的特异性。SOS1蛋白为一个同型二聚体，每个单体由跨膜和胞内结构域组成，这为整合来自不同途径的信号和调节Na⁺转运提供了稳定的对接平台。SOS1基因转录水平受到不同胁迫条件的调控，通过Ca²⁺信号调控、磷酸化、自抑制和与离子转运体协同调控等机制可抑制或激活SOS1活性。该蛋白具有调控植物昼夜节律和pH以及维持离子稳态等功能，在植物逆境胁迫响应中发挥重要作用。论文对SOS1的结构、功能、调控机制及其维持植物离子稳态平衡作用等方面的研究进展加以综述，并对其未来研究方向进行展望，以期为农作物抗逆性遗传改良提供理论支持和优异基因资源。

关键词: SOS1, Na⁺外排, Na⁺长距离运输, 离子稳态, SOS信号通路, 耐盐性, 氧化应激

Abstract:

Plants regulate ion homeostasis to adapt to saline environment through a series of complex transport systems. SOS（salt over sensitive）signal pathway is the major signal pathway for plants to respond to abiotic stress, which is mainly composed of plasma membrane Na⁺/H⁺ antiporter SOS1, serine/threonine protein kinase SOS2, and calcium sensor SOS3. As one of the main members of SOS signaling pathway, SOS1 widely exists in higher plants. Due to the early evolutionary differences, the structural, physical, and chemical properties of SOS1 from different species had certain specificity. The SOS1 protein is a homodimer, and each monomer is composed of transmembrane and intracellular domains, which provides a stable docking platform for integrating signals from different pathways and regulating Na⁺ transport. Transcription level of the SOS1 gene was regulated by different stress conditions. SOS1 activity was inhibited or activated through Ca²⁺ signal regulation, phosphorylation, self-inhibition, and synergistic regulation with other ion transporters. SOS1 has shown to regulate circadian rhythm and pH, and maintain ion homeostasis in plants, which plays an important role in the response of plant to abiotic stress. In this review, the structure, function, regulation mechanism of SOS1 and its role in maintaining plant ion homeostasis are reviewed, and its future research direction is also prospected. The information in this review provide a theoretical support and excellent genetic resources for the genetic improvement of crops to produce new, stress-resistant varieties.

Key words: SOS1, Na⁺ extrusion, Na⁺ long-distance transport, ion homeostasis, SOS signaling pathway, salt tolerance, oxidative stress

朱业胜, 伍国强, 魏明. 质膜Na⁺/H⁺逆向转运蛋白SOS1在植物离子稳态平衡中的作用[J]. 生物技术通报, 2023, 39(12): 16-32.

ZHU Ye-sheng, WU Guo-qiang, WEI Ming. Roles of Plasma Membrane Na⁺/H⁺ Antiporter SOS1 in Maintaining Ionic Homeostasis of Plants[J]. Biotechnology Bulletin, 2023, 39(12): 16-32.

图/表 4

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物种名 Species name	基因名称 Gene name	基因登录号 GenBank accession No.	氨基酸 Amino acid	亚细胞定位 Sub-cellular localization	跨膜结构域TMD	亲水性平均值 GRAVY	等电点 pI	分子量 Mw/kD	参考文献 Reference
甜菜 Beta vulgaris	BvSOS1	/	1 162	PM	12	0.085	6.34	128.46	Unpublished
小盐芥thellungiella halophila	ThSOS1	EF207775	1 146	PM	11	0.090	6.40	126.35	[36]
小盐芥thellungiella halophila	ThSOS1S	AB562331	473	PM	11	0.722	6.15	51.19	[37]
冰叶日中花 Mesembryanthemum crystallinum	McSOS1	EF207776	1 115	PM	12	0.110	5.97	127.36	[38]
花花柴 Karelinia caspia	KcSOS1	MN481368	1 136	PM	12	0.070	5.97	126.02	[39]
水稻Oryza sativa	OsSOS1-1	AY785147	1 148	PM	12	0.046	6.77	127.92	[11,21 -22]
水稻Oryza sativa	OsSOS1-2	MG602252	1 148	PM	13	0.047	6.77	127.90	[11,21 -22]
小麦 Triticum aestivum	TaSOS1-1	KJ563230	1 142	PM	12	0.110	7.00	126.22	[24]
小麦 Triticum aestivum	TaSOS1-2	AY326952	1 142	PM	13	0.106	7.40	126.18	[24]
大豆 Glycine max	GmSOS1	JQ287499	1 143	PM	12	0.089	6.30	126.43	[32]
文冠果Xanthoceras sorbifolium	XsSOS1	/	915	PM	6	/	6.19	102.50	[33]
萝卜 Raphanus sativus	RsSOS1	MZ484950	1 137	PM	12	0.093	6.71	125.62	[26]
海马齿Sesuvium portulacastrum	SpSOS1	JX674067	1 155	PM	12	0.063	6.08	127.95	[27]
印度南瓜 Cucurbita pepo	CmaSOS1	NW_019272028	1 142	PM	12	0.079	5.92	126.70	[20]
陆地棉Gossypium hirsutum	GhSOS1	KU994886	1 152	PM	12	0.061	6.32	128.18	[40]
拟南芥Arabidopsis thaliana	AtSOS1	AT2G01980	1 146	PM	13	0.098	7.62	127.19	[16]
荞麦Fagopyrum esculentum	FeSOS1	MH064255	1 132	PM	12	0.087	6.36	126.20	[31]
白刺 Nitraria tangutorum	NtSOS1	KC292267	1 163	PM	11	0.107	6.43	127.59	[41]
黄花草木樨Melilotus officinalis	MoSOS1	MG680455	957	PM	8	0.000	6.10	107.21	[42]
黄花红砂Reaumuria trigyna	RtSOS1	KC292265	1 145	PM	11	0.114	6.10	126.79	[43]
木榄Bruguiera gymnorhiza	BgSOS1	HM054521	1 153	PM	12	0.136	6.19	126.91	[44]
盐生草Halogeton glomeratus	HgSOS1	KT759142	1 165	PM	11	0.086	6.21	128.97	/
牡蒿 Artemisia japonica	AjSOS1	KP896475	1 147	PM	12	0.061	6.08	127.04	[45]
菠菜 Spinacia oleracea	SoSOS1	HG799055	1 102	/	12	0.164	6.38	121.97	[28]
葡萄 Vitis vinifera	VvSOS1	NM_001281211	1 141	/	12	0.122	6.32	126.30	[46]
黄瓜 Cucumis sativus	CsSOS1	JQ655747	1 144	PM	11	0.078	6.30	127.27	[47]
黑果枸杞Lycium ruthenicum	LrSOS1	MH454108	1 160	/	12	0.062	5.89	128.61	[48]
欧洲油菜 Brassica napus	BnSOS1	EU487184	1 142	PM	11	0.109	6.45	126.01	[26]
胡杨 Populus euphratica	PeSOS1	KX132789	1 145	PM	12	0.141	6.43	126.84	[49]
篦麻 Ricinus communis	RcSOS1	KX943304	1 143	PM	12	0.069	6.62	126.49	[50]
菊芋 Helianthus tuberosus	HtSOS1	KC410809	1 130	PM	12	0.091	6.00	125.00	[51]
盐地碱蓬 Suaeda salsa	SsSOS1	KF914414.1	1 169	PM	12	0.023	6.52	129.20	[52-53]
绿豆 Vigna radiata	VrSOS1	KC855193	1 046	/	11	0.119	6.20	116.10	/
节节麦 Aegilops tauschii	AtASOS1	FN356231	1 142	/	12	0.10	6.85	126.18	/
短柄草Brachypodium sylvaticum	BsSOS1	KF671967	1 140	/	12	0.078	7.63	126.77	/
海滨锦葵Kosteletzkya virginica	KvSOS1	KJ577576	1 147	PM	12	0.073	6.18	127.56	[54]
山萮菜Eutrema salsugineum	EsSOS1	KF671959	1 127	/	12	0.064	6.47	124.51	[55]
一粒小麦Triticum monococcum	TmSOS1	FN356229	1 142	/	12	0.108	6.87	126.33	/
大叶补血草Limonium gmelinii	LgSOS1	EU780458	1 151	PM	13	0.163	6.15	126.52	[56]
藜麦Chenopodium quinoa	CqSOS1	EU024570	1 158	PM	11	0.094	6.27	127.90	[57]
霸王Zygophyllum xanthoxylon	ZxSOS1	GU177864	1 153	PM	12	0.083	6.20	127.88	[58]
番茄Solanum lycopersicum	SlSOS1-1	AB675690	1 151	PM	12	0.069	5.85	127.54	[59]
番茄Solanum lycopersicum	SlSOS1-2	NM_001247769	1 151	PM	13	0.083	5.89	127.50	[59]
苦荞麦Fagopyrum tataricum	FtSOS1	KY659584	1 131	PM	12	0.090	6.13	125.88	[60]
盐角草Salicornia brachiata	SbSOS1	EU879059	1 159	/	11	0.037	5.92	128.55	[61]
山丹 Lilium pumilum	LpSOS1	OM650676	1 161	PM	12	0.107	6.59	129.14	[62]
甘蔗Saccharum officinarum	ScSOS1	KT003285	423	Nucleus	/	-0.396	9.12	47.60	[63]
小花碱茅Puccinellia tenuiflora	PtSOS1-1	EF440291	1 137	PM	11	0.107	6.48	125.50	[64]
小花碱茅Puccinellia tenuiflora	PtSOS1-2	GQ452778	1 143	PM	13	0.103	6.82	126.16	[64]
海岛棉Gossypium barbadense	GbSOS1	/	1 152	PM	/	/	6.42	128.04	[35]

物种名 Species name	基因名称 Gene name	基因登录号 GenBank accession No.	氨基酸 Amino acid	亚细胞定位 Sub-cellular localization	跨膜结构域TMD	亲水性平均值 GRAVY	等电点 pI	分子量 Mw/kD	参考文献 Reference
甜菜 Beta vulgaris	BvSOS1	/	1 162	PM	12	0.085	6.34	128.46	Unpublished
小盐芥thellungiella halophila	ThSOS1	EF207775	1 146	PM	11	0.090	6.40	126.35	[36]
小盐芥thellungiella halophila	ThSOS1S	AB562331	473	PM	11	0.722	6.15	51.19	[37]
冰叶日中花 Mesembryanthemum crystallinum	McSOS1	EF207776	1 115	PM	12	0.110	5.97	127.36	[38]
花花柴 Karelinia caspia	KcSOS1	MN481368	1 136	PM	12	0.070	5.97	126.02	[39]
水稻Oryza sativa	OsSOS1-1	AY785147	1 148	PM	12	0.046	6.77	127.92	[11,21 -22]
水稻Oryza sativa	OsSOS1-2	MG602252	1 148	PM	13	0.047	6.77	127.90	[11,21 -22]
小麦 Triticum aestivum	TaSOS1-1	KJ563230	1 142	PM	12	0.110	7.00	126.22	[24]
小麦 Triticum aestivum	TaSOS1-2	AY326952	1 142	PM	13	0.106	7.40	126.18	[24]
大豆 Glycine max	GmSOS1	JQ287499	1 143	PM	12	0.089	6.30	126.43	[32]
文冠果Xanthoceras sorbifolium	XsSOS1	/	915	PM	6	/	6.19	102.50	[33]
萝卜 Raphanus sativus	RsSOS1	MZ484950	1 137	PM	12	0.093	6.71	125.62	[26]
海马齿Sesuvium portulacastrum	SpSOS1	JX674067	1 155	PM	12	0.063	6.08	127.95	[27]
印度南瓜 Cucurbita pepo	CmaSOS1	NW_019272028	1 142	PM	12	0.079	5.92	126.70	[20]
陆地棉Gossypium hirsutum	GhSOS1	KU994886	1 152	PM	12	0.061	6.32	128.18	[40]
拟南芥Arabidopsis thaliana	AtSOS1	AT2G01980	1 146	PM	13	0.098	7.62	127.19	[16]
荞麦Fagopyrum esculentum	FeSOS1	MH064255	1 132	PM	12	0.087	6.36	126.20	[31]
白刺 Nitraria tangutorum	NtSOS1	KC292267	1 163	PM	11	0.107	6.43	127.59	[41]
黄花草木樨Melilotus officinalis	MoSOS1	MG680455	957	PM	8	0.000	6.10	107.21	[42]
黄花红砂Reaumuria trigyna	RtSOS1	KC292265	1 145	PM	11	0.114	6.10	126.79	[43]
木榄Bruguiera gymnorhiza	BgSOS1	HM054521	1 153	PM	12	0.136	6.19	126.91	[44]
盐生草Halogeton glomeratus	HgSOS1	KT759142	1 165	PM	11	0.086	6.21	128.97	/
牡蒿 Artemisia japonica	AjSOS1	KP896475	1 147	PM	12	0.061	6.08	127.04	[45]
菠菜 Spinacia oleracea	SoSOS1	HG799055	1 102	/	12	0.164	6.38	121.97	[28]
葡萄 Vitis vinifera	VvSOS1	NM_001281211	1 141	/	12	0.122	6.32	126.30	[46]
黄瓜 Cucumis sativus	CsSOS1	JQ655747	1 144	PM	11	0.078	6.30	127.27	[47]
黑果枸杞Lycium ruthenicum	LrSOS1	MH454108	1 160	/	12	0.062	5.89	128.61	[48]
欧洲油菜 Brassica napus	BnSOS1	EU487184	1 142	PM	11	0.109	6.45	126.01	[26]
胡杨 Populus euphratica	PeSOS1	KX132789	1 145	PM	12	0.141	6.43	126.84	[49]
篦麻 Ricinus communis	RcSOS1	KX943304	1 143	PM	12	0.069	6.62	126.49	[50]
菊芋 Helianthus tuberosus	HtSOS1	KC410809	1 130	PM	12	0.091	6.00	125.00	[51]
盐地碱蓬 Suaeda salsa	SsSOS1	KF914414.1	1 169	PM	12	0.023	6.52	129.20	[52-53]
绿豆 Vigna radiata	VrSOS1	KC855193	1 046	/	11	0.119	6.20	116.10	/
节节麦 Aegilops tauschii	AtASOS1	FN356231	1 142	/	12	0.10	6.85	126.18	/
短柄草Brachypodium sylvaticum	BsSOS1	KF671967	1 140	/	12	0.078	7.63	126.77	/
海滨锦葵Kosteletzkya virginica	KvSOS1	KJ577576	1 147	PM	12	0.073	6.18	127.56	[54]
山萮菜Eutrema salsugineum	EsSOS1	KF671959	1 127	/	12	0.064	6.47	124.51	[55]
一粒小麦Triticum monococcum	TmSOS1	FN356229	1 142	/	12	0.108	6.87	126.33	/
大叶补血草Limonium gmelinii	LgSOS1	EU780458	1 151	PM	13	0.163	6.15	126.52	[56]
藜麦Chenopodium quinoa	CqSOS1	EU024570	1 158	PM	11	0.094	6.27	127.90	[57]
霸王Zygophyllum xanthoxylon	ZxSOS1	GU177864	1 153	PM	12	0.083	6.20	127.88	[58]
番茄Solanum lycopersicum	SlSOS1-1	AB675690	1 151	PM	12	0.069	5.85	127.54	[59]
番茄Solanum lycopersicum	SlSOS1-2	NM_001247769	1 151	PM	13	0.083	5.89	127.50	[59]
苦荞麦Fagopyrum tataricum	FtSOS1	KY659584	1 131	PM	12	0.090	6.13	125.88	[60]
盐角草Salicornia brachiata	SbSOS1	EU879059	1 159	/	11	0.037	5.92	128.55	[61]
山丹 Lilium pumilum	LpSOS1	OM650676	1 161	PM	12	0.107	6.59	129.14	[62]
甘蔗Saccharum officinarum	ScSOS1	KT003285	423	Nucleus	/	-0.396	9.12	47.60	[63]
小花碱茅Puccinellia tenuiflora	PtSOS1-1	EF440291	1 137	PM	11	0.107	6.48	125.50	[64]
小花碱茅Puccinellia tenuiflora	PtSOS1-2	GQ452778	1 143	PM	13	0.103	6.82	126.16	[64]
海岛棉Gossypium barbadense	GbSOS1	/	1 152	PM	/	/	6.42	128.04	[35]