不同类型盐生植物适应盐胁迫的生理生长机制及Na+逆向转运研究进展

doi:10.13560/j.cnki.biotech.bull.1985.2022-0342

摘要/Abstract

摘要：

盐生植物是指能在离子浓度至少200 mmol/L以上的生境中生长并完成生活史的植物。盐生植物可分为稀盐盐生植物、泌盐盐生植物、拒盐盐生植物三类。本文从生长形态、生理和分子3个方面总结三类盐生植物响应盐胁迫的不同策略及研究进展，发现盐生植物在分子水平上主要通过Na⁺转运蛋白和为其提供能量的两类基因应对体内过高Na⁺，这可能是引起盐生植物生理和生长形态异于非盐生植物的重要因素。其中稀盐盐生植物主要通过液泡离子区隔化应对盐胁迫，并表现出肉质化生长形态；泌盐盐生植物通过将体内盐分排出体外应对盐胁迫，并进化出特有的生理结构——盐腺或盐囊泡；拒盐盐生植物通过将盐离子积累在皮层细胞液泡和根部木质部薄壁细胞中减少向上运输Na⁺，同时根部多栓质化减少Na⁺吸收。本综述旨在为今后研究盐生植物及其耐盐机制提供相关依据，为植物耐盐分子育种奠定基础。

关键词: 盐生植物, Na⁺逆向转运, 耐盐生理机制, 耐盐生长机制

Abstract:

Halophytes refer to plants that can grow and complete their life cycle in habitats with an ion concentration of at least 200 mmol/L. Halophytes can be classified into three types：euhalophytes, recretohalophytes and pseudohalophytes. This review summarizes the different strategies and research progress of the three types of halophytes in response to salt stress from the three aspects of growth morphology, physiology and molecule. It is found that halophytes mainly use Na⁺ transporters genes and energy-providing genes to respond to excessive Na⁺ at the molecular level, which may be an important factor that causes halophytes to be different from non-halophytes in physiology and growth morphology. Euhalophytes respond to salt stress mainly through vacuolar ion compartmentalization and have succulent growth morphology. Recretohalophytes respond to salt through by expelling salt from their bodies and evolve unique physiological structures-salt glands or salt vesicles. Pseudohalophytes reduce the upward transport of Na⁺ by accumulating salt ions in the vacuoles of cortical cells and parenchyma cells of root xylem; meanwhile, root is having suberization to reduce the absorption of Na⁺. The purpose of the review is to provide relevant basis for the study of halophytes and their salt tolerance mechanism and lay a foundation for plant salt tolerance molecular breeding in the future.

Key words: halophytes, Na⁺antiport, physiological mechanism of salt tolerance, growth mechanism of salt tolerance

刘佳欣, 张会龙, 邹荣松, 杨秀艳, 朱建峰, 张华新. 不同类型盐生植物适应盐胁迫的生理生长机制及Na⁺逆向转运研究进展[J]. 生物技术通报, 2023, 39(1): 59-72.

LIU Jia-xin, ZHANG Hui-long, ZOU Rong-song, YANG Xiu-yan, ZHU Jian-feng, ZHANG Hua-xin. Research Progress in Na⁺Antiport and Physiological Growth Mechanisms of Differernt Halophytes Adapted to Salt Stress[J]. Biotechnology Bulletin, 2023, 39(1): 59-72.

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基因 Gene	基因类型 Gene type	基因功能 Gene function	基因来源 Gene resource	参考文献 Reference
NsNHX1	液泡膜Na⁺/H⁺逆向转运蛋白基因	调控Na⁺液泡区隔化	西伯利亚白刺Nitraria sibirica	[20]
SsNHX1	液泡膜Na⁺/H⁺逆向转运蛋白基因	调控Na⁺液泡区隔化	盐地碱蓬Suaeda salsa	[30-31]
SbNHX1	液泡膜Na⁺/H⁺逆向转运蛋白基因	调控Na⁺液泡区隔化和维持离子稳态的过程	海蓬子Sal Icornia brachiate	[32-33]
SeNHX1	液泡膜Na⁺/H⁺逆向转运蛋白基因	调控Na⁺液泡区隔化	盐角草Sal Icornia europaea	[34]
KfNHX1	液泡膜Na⁺/H⁺逆向转运蛋白基因	可能增强将Na⁺区隔化至液泡的能力	盐爪爪Kalidium foliatum	[35]
HcNHX1	液泡膜Na⁺/H⁺逆向转运蛋白基因	离子区隔化	盐穗木Halostachys caspica	[36-37]
NsSOS1	质膜Na⁺/H⁺逆向转运蛋白基因	将胞质中过量的Na⁺排出到胞外	西伯利亚白刺Nitraria sibirica	[22,38]
SsHKT1	高亲和性K⁺转运蛋白基因	减少根木质部Na⁺回收，协调SsSOS1和SsNHX1维持Na⁺积累	盐地碱蓬Suaeda salsa	[30]
SsHAK2	高亲和性K⁺转运蛋白基因	参与K⁺吸收及转运过程	盐地碱蓬Suaeda salsa	[39]
AQP	编码水通道蛋白基因	离子区隔化	盐地碱蓬Suaeda salsa	[40]
HcPIP1	水通道蛋白家族亚类质膜内嵌蛋白基因	在胁迫时能够通过增加根的生长以抵御胁迫的影响	盐穗木Halostachys caspica	[41]
NsVP1	液泡膜H⁺-PPase基因	调控Na⁺液泡区隔化	西伯利亚白刺Nitraria sibirica	[20]
KfVP1	液泡膜H⁺-PPase基因	离子的调控运输中发挥作用	盐爪爪Kalidium foliatum	[42]
HcVP1	液泡膜H⁺-PPase基因	增加液泡中Na⁺的积累来增强转基因拟南芥的耐盐性	盐穗木Halostachys caspica	[43]
HcVHA-B	液泡膜H⁺-ATPase亚基B基因	增加液泡中Na⁺的积累来增强转基因拟南芥的耐盐性	盐穗木Halostachys caspica	[43]

基因 Gene	基因类型 Gene type	基因功能 Gene function	基因来源 Gene resource	参考文献 Reference
NsNHX1	液泡膜Na⁺/H⁺逆向转运蛋白基因	调控Na⁺液泡区隔化	西伯利亚白刺Nitraria sibirica	[20]
SsNHX1	液泡膜Na⁺/H⁺逆向转运蛋白基因	调控Na⁺液泡区隔化	盐地碱蓬Suaeda salsa	[30-31]
SbNHX1	液泡膜Na⁺/H⁺逆向转运蛋白基因	调控Na⁺液泡区隔化和维持离子稳态的过程	海蓬子Sal Icornia brachiate	[32-33]
SeNHX1	液泡膜Na⁺/H⁺逆向转运蛋白基因	调控Na⁺液泡区隔化	盐角草Sal Icornia europaea	[34]
KfNHX1	液泡膜Na⁺/H⁺逆向转运蛋白基因	可能增强将Na⁺区隔化至液泡的能力	盐爪爪Kalidium foliatum	[35]
HcNHX1	液泡膜Na⁺/H⁺逆向转运蛋白基因	离子区隔化	盐穗木Halostachys caspica	[36-37]
NsSOS1	质膜Na⁺/H⁺逆向转运蛋白基因	将胞质中过量的Na⁺排出到胞外	西伯利亚白刺Nitraria sibirica	[22,38]
SsHKT1	高亲和性K⁺转运蛋白基因	减少根木质部Na⁺回收，协调SsSOS1和SsNHX1维持Na⁺积累	盐地碱蓬Suaeda salsa	[30]
SsHAK2	高亲和性K⁺转运蛋白基因	参与K⁺吸收及转运过程	盐地碱蓬Suaeda salsa	[39]
AQP	编码水通道蛋白基因	离子区隔化	盐地碱蓬Suaeda salsa	[40]
HcPIP1	水通道蛋白家族亚类质膜内嵌蛋白基因	在胁迫时能够通过增加根的生长以抵御胁迫的影响	盐穗木Halostachys caspica	[41]
NsVP1	液泡膜H⁺-PPase基因	调控Na⁺液泡区隔化	西伯利亚白刺Nitraria sibirica	[20]
KfVP1	液泡膜H⁺-PPase基因	离子的调控运输中发挥作用	盐爪爪Kalidium foliatum	[42]
HcVP1	液泡膜H⁺-PPase基因	增加液泡中Na⁺的积累来增强转基因拟南芥的耐盐性	盐穗木Halostachys caspica	[43]
HcVHA-B	液泡膜H⁺-ATPase亚基B基因	增加液泡中Na⁺的积累来增强转基因拟南芥的耐盐性	盐穗木Halostachys caspica	[43]

基因 Gene	基因类型 Gene type	基因功能 Gene function	基因来源 Gene resource	参考文献 Reference
SaNHX2	液泡膜Na⁺/H⁺逆向转运蛋白基因	控制液泡膜中活性K⁺的摄取，同时调节气孔的关闭	互花米草Spartina alterniflora	[73]
AmNHX1	液泡膜Na⁺/H⁺逆向转运蛋白基因	将Na⁺转运到液泡中	海榄雌Avicennia maritima	[74]
AgNHX1	液泡膜Na⁺/H⁺逆向转运蛋白基因	保持离子平衡	北滨黎Atriplex gmelini	[68]
McNHX1	液泡膜Na⁺/H⁺逆向转运蛋白基因	离子区隔	冰叶日中花Mesembryanthemum crystallinum	[75]
LsNHX2	液泡膜Na⁺/H⁺逆向转运蛋白基因	Na⁺的囊泡化转运活动	中华补血草Limonium sinense	[76]
BgNHX1	液泡膜Na⁺/H⁺逆向转运蛋白基因	Na⁺区隔化作用	木榄Bruguiera gymnorrhiz	[77]
AmSOS1	质膜Na⁺/H⁺逆向转运蛋白基因	将多余的Na⁺泵出细胞	海榄雌Avicennia maritima	[74]
AcSOS1	质膜Na⁺/H⁺逆向转运蛋白基因	盐囊泡Na⁺积累	四翅滨藜Atriplex canescens	[63]
McSOS1	质膜Na⁺/H⁺逆向转运蛋白基因	离子区隔	冰叶日中花Mesembryanthemum crystallinum	[78]
AcHKT1	高亲和K⁺转运蛋白基因	盐囊泡Na⁺积累	四翅滨藜Atriplex canescens	[63]
McHKT1	高亲和K⁺转运蛋白基因	将Na⁺转运到液泡中	冰叶日中花Mesembryanthemum crystallinum	[79]
ThVHAc1	液泡膜H⁺-ATPase亚基c1基因	具有抗逆性作用	刚毛柽柳Tamarix hispida	[80]
SaVHAc1	液泡膜H⁺-ATPase亚基c1基因	参与能量供应	互花米草Spartina alterniflora	[81]
AhVP	液泡膜H⁺-PPase基因	对渗透和/或离子压力的耐受性	滨藜Atriplex halimus	[82-83]
ThVP1	液泡膜H⁺-PPase基因	参与Na⁺的隔离	刚毛柽柳Tamarix hispida	[66]
AmHA1	质膜H⁺-ATPase基因	参与能量供应	海榄雌Avicennia maritima	[74]
PIP and TIP	水通道蛋白基因	参与盐腺脱盐过程中水的再吸收	海茄冬Avicennia officinalis	[84]
CLC	质膜Cl^-通道基因	将Cl^-区域化为囊泡维持电荷平衡	二色补血草Limonium bicolor	[72]

基因 Gene	基因类型 Gene type	基因功能 Gene function	基因来源 Gene resource	参考文献 Reference
SaNHX2	液泡膜Na⁺/H⁺逆向转运蛋白基因	控制液泡膜中活性K⁺的摄取，同时调节气孔的关闭	互花米草Spartina alterniflora	[73]
AmNHX1	液泡膜Na⁺/H⁺逆向转运蛋白基因	将Na⁺转运到液泡中	海榄雌Avicennia maritima	[74]
AgNHX1	液泡膜Na⁺/H⁺逆向转运蛋白基因	保持离子平衡	北滨黎Atriplex gmelini	[68]
McNHX1	液泡膜Na⁺/H⁺逆向转运蛋白基因	离子区隔	冰叶日中花Mesembryanthemum crystallinum	[75]
LsNHX2	液泡膜Na⁺/H⁺逆向转运蛋白基因	Na⁺的囊泡化转运活动	中华补血草Limonium sinense	[76]
BgNHX1	液泡膜Na⁺/H⁺逆向转运蛋白基因	Na⁺区隔化作用	木榄Bruguiera gymnorrhiz	[77]
AmSOS1	质膜Na⁺/H⁺逆向转运蛋白基因	将多余的Na⁺泵出细胞	海榄雌Avicennia maritima	[74]
AcSOS1	质膜Na⁺/H⁺逆向转运蛋白基因	盐囊泡Na⁺积累	四翅滨藜Atriplex canescens	[63]
McSOS1	质膜Na⁺/H⁺逆向转运蛋白基因	离子区隔	冰叶日中花Mesembryanthemum crystallinum	[78]
AcHKT1	高亲和K⁺转运蛋白基因	盐囊泡Na⁺积累	四翅滨藜Atriplex canescens	[63]
McHKT1	高亲和K⁺转运蛋白基因	将Na⁺转运到液泡中	冰叶日中花Mesembryanthemum crystallinum	[79]
ThVHAc1	液泡膜H⁺-ATPase亚基c1基因	具有抗逆性作用	刚毛柽柳Tamarix hispida	[80]
SaVHAc1	液泡膜H⁺-ATPase亚基c1基因	参与能量供应	互花米草Spartina alterniflora	[81]
AhVP	液泡膜H⁺-PPase基因	对渗透和/或离子压力的耐受性	滨藜Atriplex halimus	[82-83]
ThVP1	液泡膜H⁺-PPase基因	参与Na⁺的隔离	刚毛柽柳Tamarix hispida	[66]
AmHA1	质膜H⁺-ATPase基因	参与能量供应	海榄雌Avicennia maritima	[74]
PIP and TIP	水通道蛋白基因	参与盐腺脱盐过程中水的再吸收	海茄冬Avicennia officinalis	[84]
CLC	质膜Cl^-通道基因	将Cl^-区域化为囊泡维持电荷平衡	二色补血草Limonium bicolor	[72]

基因 Gene	基因类型 Gene type	基因功能 Gene function	基因来源Gene resource	参考文献 Reference
cNHX1	液泡膜Na⁺/H⁺逆向转运蛋白基因		柑橘Citrus×paradisi	[101]
PtNHX1	液泡膜Na⁺/H⁺逆向转运蛋白基因	将Na⁺从体内排出	小花碱茅Puccinellia tenuiflora	[98]
Pt SOS1	质膜Na⁺/H⁺逆向转运蛋白基因	将Na⁺从体内排出	小花碱茅P. tenuiflora	[98]
Pt HKT1；5	高亲和K⁺转运蛋白基因	将Na⁺从木质部排到木质部薄壁细胞	小花碱茅P. tenuiflora	[98]