Research Progress in Alleviating Effects of Exogenous Substances on Salt Stress in Rice

doi:10.16819/j.1001-7216.2023.220404

Abstract

Abstract:

Salt stress is one of the main abiotic factors hindering rice growth, development and grain yield. By the use of exogenous substances, adverse effects can be alleviated and its growth and yield traits can be improved. This work reviews the regulation mechanism and mitigation effect of different exogenous substances (auxin, gibberellin, cytokinin, abscisic acid, ethylene, melatonin, salicylic acid, polyamine, brassinosteroid and jasmonic acid as well as calcium and silicon ionic exogenous substances) on rice salt stress. Finally, the utilization and research work of exogenous substances on salt tolerance of rice is prospected in order to lay a theoretical basis for improving rice growth, yield and quality in saline soil.

Key words: exogenous substance, salt stress, rice

摘要：

盐胁迫是影响水稻生长发育和产量的主要非生物因素之一，外源物质能有效缓解盐对水稻的毒害作用并增加水稻产量。本文主要综述了不同外源物质(生长素、赤霉素、细胞分裂素、脱落酸、乙烯、褪黑素、水杨酸、多胺、油菜素类固醇、茉莉酸类外源生长调节物质以及钙、硅离子类外源物)对盐胁迫下水稻生长的不同调控机理及其缓解效应，并进一步对外源物质在水稻耐盐上的利用与研究工作进行了展望，以期为改善盐渍土环境下水稻的生长发育以及提高产量和品质提供理论依据。

关键词: 外源物质, 盐胁迫, 水稻

LIU Shuli, ZHANG Rui, Shahid HUSSAIN, WANG Yang, CHEN Yinglong, WEI Huanhe, HOU Hongyan, DAI Qigen. Research Progress in Alleviating Effects of Exogenous Substances on Salt Stress in Rice[J]. Chinese Journal OF Rice Science, 2023, 37(1): 1-15.

刘淑丽, 张瑞, 王洋, 陈英龙, 韦还和, 侯红燕, 戴其根. 外源物质对水稻盐胁迫缓解效应研究进展[J]. 中国水稻科学, 2023, 37(1): 1-15.

Figures/Tables 1

References 144

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外源物质 Exogenous substances	表型影响 Phenotypic effects	生理生化影响 Physiological and biochemical effects	分子水平影响 Molecular level effects
脱落酸 Abscisic acid	提高秧苗存活率^[38]Increase seedling survival rate	缓解类胡萝卜素减少引起的强光损伤^[51]；增强可溶性蛋白及抗氧化酶活性^[52] Alleviate the damage caused by reduced carotenoid; enhance the content of soluble protein and antioxidant enzymes activities	转录因子OsSAE1可作水稻种子萌发和耐盐性的正调控因子^[53]OsSAE1 could be a positive regulator of seed germination and salt tolerance in rice
乙烯 Ethylene	萌发阶段克服种子休眠^[56]；孕穗成熟阶段抑制生长^[55,57]；促进幼苗根系生长^[59]Overcome seed dormancy during germination; inhibit growth from booting to maturity; promote seedling root growth	增加根系抗氧化活性,提高渗透调节物质含量^[59]Increase root antioxidant activity, increase the content of osmotic regulating substances	通过SNL1(SIN3-like 1)和SNL2(SIN3-like 2)负调控ABA信号^[59]ABA signaling is negatively regulated by SNL1(SIN3-like 1) and SNL2(SIN3-like 2)
茉莉酸类物质 Jasmonates	低浓度促进、高浓度抑制的“双重效应”^[122]" Double effects", promoting at low concentration and inhibiting at high concentration	维持叶绿素和K⁺含量,增加抗氧化酶活性^[124]；缓解光系统II(PSII)活性的降低和离子胁迫与渗透胁迫^[125]；调控植株内源激素水平^[128]Maintain the contents of chlorophyll and K⁺, increase the activitises of antioxidant enzymes; alleviate the reduction of photosystem II(PSII) activity, alleviate ionic and osmotic stress; regulate endogenous hormone levels in plants	TaFBA-2A负调控JA的生物合成^[120]；OSPP65可以通过调节JA和ABA信号通路以及棉子糖代谢途径负调控水稻的耐盐性^[121]；茉莉酸生物合成基因OsOPR7缓解盐胁迫^[127] TaFBA-2A negatively regulates JA biosynthesis; OSPP65 can negatively regulate salt tolerance of rice by regulating JA and ABA signaling pathways and raffinose metabolism pathway; Jasmonate biosynthetic gene OsOPR7 alleviates salt stress
生长素 Indole acetic acid	促进种子萌发^{[12,41,100,109]}；改善幼苗个体形态^{[35⇓⇓-38,99]}；促进植株生长^{[35⇓⇓⇓⇓-40]}；提高产量^[39-40,102] Promote seed germination; improve seedling morphology; boost plant growth; increase grain yield	促进细胞分裂、生长与分化,提高代谢产物含量^[39,40]Promote cell division, growth and differentiation, increase metabolite contents
赤霉素 Gibberellin		维持光合系统的稳定^[42]；促进淀粉合成^[43]Maintain the stability of the photosynthetic system; promote starch synthesis	作为GA的负调控因子，盐胁迫下DELLA蛋白的积累可以诱导ABA合成基因的表达，抑制植物生长的同时提高抗逆性^{[45⇓⇓-48]}As a negative regulator of GA, DELLA protein accumulation under salt stress can induce the expression of ABA synthesis genes, inhibit plant growth, and improve stress tolerance
细胞分裂素 Cytokinin		增强叶片光合速率^[40]Enhance leaf photosynthetic rate
褪黑素 Melatonin		改善光合作用和离子稳态^[61,62,67]；增强抗氧化与渗透调节能力^[64,66]；增加初级代谢产物的积累^[68]；激活TFs级联反应和植物激素信号等^[70]Improve photosynthesis and ion homeostasis; enhance antioxidant and osmotic adjustment ability; increase the accumulation of primary metabolites; activate TFs cascade and plant hormone signaling	调节水稻钾离子转运蛋白OsHAK基因的表达^[63]；抑制衰老相关基因的表达^[69]；调节ABA生物合成及分解基因的表达^[72] Regulate the expression of K⁺ transporter OsHAK gene in rice; inhibit the expression of genes related to senescence; regulate the expression of ABA biosynthesis and decomposition genes
水杨酸 Salicylic acid		诱导抗氧化防御系统，调控离子平衡、激素稳态和代谢物质^[74]；维持细胞膜稳定^[79]Induce antioxidant defense system, regulate ion balance, hormone homeostasis, and metabolic substances; maintain cell membrane stability	诱导抗盐基因表达^[80]；调控GA与ABA生物合成基因的表达^[81]Induce the expression of salt resistance genes; regulate the expression of GA and ABA biosynthetic genes
多胺 Polyamines		调节植物内源激素水平^[85]；稳定质膜结构^[93]；缓解氧化胁迫和渗透胁迫^[94,95]；提高植株光合色素含量^[97]Regulate endogenous hormone levels in plants; Stabilize plasma membrane structure; alleviate oxidative stress and osmotic stress; increase photosynthetic pigment contents in plants	调控ABA合成途径中关键基因NCED的表达^[88]；抑制光化学反应及叶绿体编码基因的下调^[98] Regulate the expression of NCED, a key gene in ABA synthesis pathway; inhibite photochemical reactions and down-regulate chloroplast coding genes
油菜素类固醇 Brassinosteroids		提高核酸和可溶性蛋白水平^[110]；维持植物细胞的渗透平衡、增加抗氧化酶活性^[112]；激活盐胁迫下的次生代谢功能^[115]；调控IAA、GA水平^[117,118]Increase the levels of nucleic acid and soluble protein; maintain the osmotic balance of plant cells, increase the activities of antioxidant enzymes; activate secondary metabolic functions under salt stress; regulate IAA and GA levels	上调OsBRI1基因的表达,下调OsDWF4基因的表达^[113]；上调类胡萝卜素和类黄酮通路基因^[115]Up-regulate the expression of OsBRI1 gene and down-regulate OsDWF4 gene; up-regulate carotenoid and flavonoid pathway genes
钙 Calcium		改善水稻幼苗的离子稳态、抗氧化防御及乙二醛酶系统^[135]；改善植物光合活性^[136]Improve ion homeostasis, antioxidant defense, and glyoxalase system in rice seedlings; maintain plant photosynthetic activity
硅 Silicon		维持光合系统稳定及离子平衡^[142]；提高抗氧化酶活性^[144]Maintain the stability of photosynthetic system and ion balance; increase the activity of antioxidant enzymes	上调植物根中质膜水通道蛋白基因的表达^[141]；上调OsHAK家族钾吸收基因及水稻钠外排(OsSOS1)、钠区隔化(OsNHX1、OsNHX3 与OsNHX5)基因的表达^[143]Up-regulate expression of plasma membrane aquaporin gene in plant roots; up-regulate OsHAK family potassium uptake genes, rice sodium efflux (OsSOS1), sodium compartments (OsNHX1, OsNHX3 and OsNHX5) gene expression

外源物质 Exogenous substances	表型影响 Phenotypic effects	生理生化影响 Physiological and biochemical effects	分子水平影响 Molecular level effects
脱落酸 Abscisic acid	提高秧苗存活率^[38]Increase seedling survival rate	缓解类胡萝卜素减少引起的强光损伤^[51]；增强可溶性蛋白及抗氧化酶活性^[52] Alleviate the damage caused by reduced carotenoid; enhance the content of soluble protein and antioxidant enzymes activities	转录因子OsSAE1可作水稻种子萌发和耐盐性的正调控因子^[53]OsSAE1 could be a positive regulator of seed germination and salt tolerance in rice
乙烯 Ethylene	萌发阶段克服种子休眠^[56]；孕穗成熟阶段抑制生长^[55,57]；促进幼苗根系生长^[59]Overcome seed dormancy during germination; inhibit growth from booting to maturity; promote seedling root growth	增加根系抗氧化活性,提高渗透调节物质含量^[59]Increase root antioxidant activity, increase the content of osmotic regulating substances	通过SNL1(SIN3-like 1)和SNL2(SIN3-like 2)负调控ABA信号^[59]ABA signaling is negatively regulated by SNL1(SIN3-like 1) and SNL2(SIN3-like 2)
茉莉酸类物质 Jasmonates	低浓度促进、高浓度抑制的“双重效应”^[122]" Double effects", promoting at low concentration and inhibiting at high concentration	维持叶绿素和K⁺含量,增加抗氧化酶活性^[124]；缓解光系统II(PSII)活性的降低和离子胁迫与渗透胁迫^[125]；调控植株内源激素水平^[128]Maintain the contents of chlorophyll and K⁺, increase the activitises of antioxidant enzymes; alleviate the reduction of photosystem II(PSII) activity, alleviate ionic and osmotic stress; regulate endogenous hormone levels in plants	TaFBA-2A负调控JA的生物合成^[120]；OSPP65可以通过调节JA和ABA信号通路以及棉子糖代谢途径负调控水稻的耐盐性^[121]；茉莉酸生物合成基因OsOPR7缓解盐胁迫^[127] TaFBA-2A negatively regulates JA biosynthesis; OSPP65 can negatively regulate salt tolerance of rice by regulating JA and ABA signaling pathways and raffinose metabolism pathway; Jasmonate biosynthetic gene OsOPR7 alleviates salt stress
生长素 Indole acetic acid	促进种子萌发^{[12,41,100,109]}；改善幼苗个体形态^{[35⇓⇓-38,99]}；促进植株生长^{[35⇓⇓⇓⇓-40]}；提高产量^[39-40,102] Promote seed germination; improve seedling morphology; boost plant growth; increase grain yield	促进细胞分裂、生长与分化,提高代谢产物含量^[39,40]Promote cell division, growth and differentiation, increase metabolite contents
赤霉素 Gibberellin		维持光合系统的稳定^[42]；促进淀粉合成^[43]Maintain the stability of the photosynthetic system; promote starch synthesis	作为GA的负调控因子，盐胁迫下DELLA蛋白的积累可以诱导ABA合成基因的表达，抑制植物生长的同时提高抗逆性^{[45⇓⇓-48]}As a negative regulator of GA, DELLA protein accumulation under salt stress can induce the expression of ABA synthesis genes, inhibit plant growth, and improve stress tolerance
细胞分裂素 Cytokinin		增强叶片光合速率^[40]Enhance leaf photosynthetic rate
褪黑素 Melatonin		改善光合作用和离子稳态^[61,62,67]；增强抗氧化与渗透调节能力^[64,66]；增加初级代谢产物的积累^[68]；激活TFs级联反应和植物激素信号等^[70]Improve photosynthesis and ion homeostasis; enhance antioxidant and osmotic adjustment ability; increase the accumulation of primary metabolites; activate TFs cascade and plant hormone signaling	调节水稻钾离子转运蛋白OsHAK基因的表达^[63]；抑制衰老相关基因的表达^[69]；调节ABA生物合成及分解基因的表达^[72] Regulate the expression of K⁺ transporter OsHAK gene in rice; inhibit the expression of genes related to senescence; regulate the expression of ABA biosynthesis and decomposition genes
水杨酸 Salicylic acid		诱导抗氧化防御系统，调控离子平衡、激素稳态和代谢物质^[74]；维持细胞膜稳定^[79]Induce antioxidant defense system, regulate ion balance, hormone homeostasis, and metabolic substances; maintain cell membrane stability	诱导抗盐基因表达^[80]；调控GA与ABA生物合成基因的表达^[81]Induce the expression of salt resistance genes; regulate the expression of GA and ABA biosynthetic genes
多胺 Polyamines		调节植物内源激素水平^[85]；稳定质膜结构^[93]；缓解氧化胁迫和渗透胁迫^[94,95]；提高植株光合色素含量^[97]Regulate endogenous hormone levels in plants; Stabilize plasma membrane structure; alleviate oxidative stress and osmotic stress; increase photosynthetic pigment contents in plants	调控ABA合成途径中关键基因NCED的表达^[88]；抑制光化学反应及叶绿体编码基因的下调^[98] Regulate the expression of NCED, a key gene in ABA synthesis pathway; inhibite photochemical reactions and down-regulate chloroplast coding genes
油菜素类固醇 Brassinosteroids		提高核酸和可溶性蛋白水平^[110]；维持植物细胞的渗透平衡、增加抗氧化酶活性^[112]；激活盐胁迫下的次生代谢功能^[115]；调控IAA、GA水平^[117,118]Increase the levels of nucleic acid and soluble protein; maintain the osmotic balance of plant cells, increase the activities of antioxidant enzymes; activate secondary metabolic functions under salt stress; regulate IAA and GA levels	上调OsBRI1基因的表达,下调OsDWF4基因的表达^[113]；上调类胡萝卜素和类黄酮通路基因^[115]Up-regulate the expression of OsBRI1 gene and down-regulate OsDWF4 gene; up-regulate carotenoid and flavonoid pathway genes
钙 Calcium		改善水稻幼苗的离子稳态、抗氧化防御及乙二醛酶系统^[135]；改善植物光合活性^[136]Improve ion homeostasis, antioxidant defense, and glyoxalase system in rice seedlings; maintain plant photosynthetic activity
硅 Silicon		维持光合系统稳定及离子平衡^[142]；提高抗氧化酶活性^[144]Maintain the stability of photosynthetic system and ion balance; increase the activity of antioxidant enzymes	上调植物根中质膜水通道蛋白基因的表达^[141]；上调OsHAK家族钾吸收基因及水稻钠外排(OsSOS1)、钠区隔化(OsNHX1、OsNHX3 与OsNHX5)基因的表达^[143]Up-regulate expression of plasma membrane aquaporin gene in plant roots; up-regulate OsHAK family potassium uptake genes, rice sodium efflux (OsSOS1), sodium compartments (OsNHX1, OsNHX3 and OsNHX5) gene expression