设施番茄盐毒害及缓解机理研究
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摘要
土壤盐渍化是蔬菜栽培的重要限制因子,造成作物生长受阻,产量降低。由于我国设施栽培条件简陋,可控性差,栽培管理技术不足,设施土壤盐渍化常与其他逆境条件共同作用,影响作物生产。番茄(Lycopersicon esculentum Mill.)是我国主要的设施栽培作物之一,喜光耐热,易受低温、弱光和土壤盐渍化影响。因此探讨番茄盐毒害及缓解机理,对于提高设施园艺生产效率具有重要的理论意义和实践价值。本试验采用水培的方法,研究了盐毒害对番茄幼苗的伤害机理,并探讨了嫁接和水杨酸对番茄盐毒害的缓解效应。取得了以下主要研究结果:
1.不同根际温度下盐胁迫对番茄光合特性和营养吸收的影响
盐胁迫和根际低温明显抑制了番茄地上部和地下部的生长,复合处理加重了抑制程度。盐胁迫打破了番茄叶片和根部的营养元素平衡,提高了番茄叶片和根系中Na含量,降低了K、Mg和根系中Ca含量,根际低温降低了番茄对K、Ca、Mg的吸收能力。盐胁迫降低了番茄叶片叶绿素a和总叶绿素含量,根际低温加剧了盐胁迫下叶绿素含量的降低。光化学猝灭系数(qP)、天线色素光能转化效率(Fv'/Fm')、电子传递速率(ETR)和表观量子效率(ΦPSⅡ)在盐胁迫后均明显降低,根际低温处理进一步降低了PSⅡ的实际活性。盐胁迫后番茄净光合速率降低,根际低温处理后,下降幅度更大。
2.根际低温和盐胁迫对番茄氧化胁迫的复合效应
盐胁迫和根际低温提高了番茄叶片O_2·~-产生速率和H_2O_2含量,造成了番茄的氧化伤害,MDA含量上升和电解质渗透率提高,在复合处理中氧化伤害程度最高。盐胁迫诱导了番茄Fe-SOD和Cu,Zn-SOD活性,根际低温诱导了Fe-SOD和Mn-SOD活性。CAT对盐胁迫敏感,盐胁迫和根际低温均降低了其活性,复合处理活性最低。GPX、APX和GR在盐胁迫和根际低温处理后活性上升,在复合处理中上升幅度较大,说明其在清除叶片活性氧方面有重要作用。盐胁迫和根际低温处理后番茄叶片谷胱甘肽和抗坏血酸含量明显降低,复合处理含量最低,说明胁迫后番茄还原态受到破坏。
3.不同光照条件下盐胁迫对番茄光合特性的影响
盐胁迫抑制了番茄生长,弱光下这种抑制程度相对较低。盐胁迫后番茄净光合速率(Pn)、气孔导度(Gs)和胞间CO_2浓度(Ci)均降低,气孔限制值(Ls)上升,表明盐胁迫引起的光合速率下降可能是由气孔限制所致。盐胁迫提高了暗呼吸速率(Rd)和光补偿点(LCP),番茄利用弱光能力降低;盐胁迫降低了光饱和点(LSP),对强光的抵御能力降低。弱光条件下,Rd、LCP和LSP均降低,利用弱光能力加强,这是番茄对弱光环境的适应。100 mmol·L~(-1)的NaCl处理对Fv/Fm没有明显影响,降低了番茄qP、Fv'/Fm'和ΦPSⅡ。弱光提高了PSⅡ的实际运行活性,但由于光照过低,ETR和净光合速率明显低于强光。
4.不同光照条件下盐胁迫对番茄元素分布、膜系统ATP酶和焦磷酸酶活性的影响
盐胁迫提高了番茄根系和叶片中Na含量,降低了K含量。弱光对盐胁迫下Na和K含量没有显著影响,提高了番茄对P、Ca、Mg、Fe和Zn的吸收能力。强光下,NaCl处理显著提高了番茄质膜H~+-ATPase和Ca~(2+)-ATPase的活性,弱光条件下,上升幅度降低。强光下,NaCl处理明显诱导了液泡膜H~+-ATPase、Ca~(2+)-ATPase和H~+-PPase活性,弱光处理降低了液泡膜ATP酶和焦磷酸酶活性。
5.不同盐分形态对番茄幼苗根系呼吸和活性氧代谢的影响
设施土壤次生盐渍化的盐分种类与滨海盐土不同,阴离子以NO_3~-为主,阳离子以Ca~(2+)为主。Ca(NO_3)_2处理和NaCl处理对番茄的毒害方式存在差异。盐胁迫下番茄根系总呼吸速率和细胞色素呼吸速率明显下降,相较Ca(NO3)_2处理,NaCl处理细胞色素呼吸下降幅度更大;在盐胁迫下抗氰呼吸速率上升,NaCl处理上升更明显。盐胁迫明显增加了活性氧含量,NaCl处理上升幅度更大。NaCl胁迫下,根系SOD、CAT和GPX活性显著增加,但在处理后期上升幅度下降,Ca(NO_3)_2胁迫后抗氧化酶活性一直上升,NaCl胁迫根系膜脂过氧化程度显著高于NaCl处理。NaCl处理抗氰呼吸的增强可以弥补细胞色素呼吸引起的ATP不足,并可能参与活性氧的清除。
6.嫁接提高番茄耐盐性机理研究
随着盐浓度的增加,番茄生长受抑制程度逐渐提高。自根嫁接对番茄生长和耐盐性没有明显影响,砧木嫁接明显缓解了盐胁迫引起的生长受阻,归因于其具有较高的光合活性和抗氧化能力。100和150 mmol·L~(-1)NaCl处理后,自根和自根嫁接番茄qP、ETR、Fv'/Fm'和ΦPSⅡ明显下降,砧木嫁接苗基本不受影响。因此,砧木嫁接苗在盐胁迫后Pn较高。盐胁迫后,砧木嫁接番茄膜脂过氧化程度较低,与抗氧化系统的响应相关。盐胁迫后,自根和自根嫁接番茄叶片CAT和DHAR活性降低,砧木嫁接番茄CAT和DHAR活性上升。盐胁迫提高了番茄叶片SOD和GPX活性,在自根和自根嫁接苗中上升幅度更大。砧木嫁接番茄具有较高的GR活性,在各盐分处理浓度下,其活性均高于自根和自根嫁接苗。盐胁迫提高了番茄叶片和根系Na含量,降低了K含量和K/Na比,砧木嫁接番茄K、Na含量与自根苗相比没有明显差异。
7.水杨酸对盐胁迫下番茄光合作用和活性氧清除系统的影响
1 mmol·L~(-1)的SA叶面喷施明显缓解了盐胁迫对植株生长的抑制作用。这种缓解效应一方面归因于SA叶面喷施缓解了盐胁迫引起的PSⅡ活性降低,电子用于CO_2固定的比例增加,提高了盐胁迫下番茄的净光合速率;另一方面,归因于SA诱导了番茄抗氧化系统活性,提高了SOD、CAT、GPX和DHAR活性,缓解了盐胁迫引起的还原型谷胱甘肽GSH和还原型抗坏血酸ASC含量的降低,降低了O_2~-的产生速率和H_2O_2含量,使盐胁迫条件下MDA含量和电解质渗透率明显降低。
8.水杨酸对盐胁迫下番茄元素分布和根系膜系统的影响
SA处理提高了番茄对K的选择吸收能力,提高了盐胁迫后番茄根系Ca和Mg含量,降低了Na含量,使番茄维持较高的K/Na比。SA处理显著提高了正常生长和胁迫条件下番茄根系质膜H~+-ATPase和Ca~(2+)-ATPase活性。SA处理提高了盐胁迫下番茄液泡膜H~+-ATPase和H~+-PPase活性,对液泡膜Ca~(2+)-ATPase活性没有明显影响。SA处理缓解了盐胁迫条件下番茄根系NADPH氧化速率、NADH氧化速率、Fe(CN)_6~(3+)还原速率和EDTA-Fe~(3+)还原速率的上升,使其接近于正常生长水平。
High concentrations of salts in soils account for large decreases in the yield of a wide variety of crops.Because of the limitation of greenhouse and bad management, crops are routinely subjected to a combination of different abiotic stresses in China. Tomato(Lycopersicon esculentum Mill) is widely planted in greenhouse and easily affected by low temperature,low light intensity and salinity.Therefore,it is very important to study the physiological mechanisms of salt toxicity in tomato and to take measures to decrease the toxicity.In this study,hydroponic culture experiments were conducted to investigate the physiological mechanisms of salt toxicity to tomato and the effects of low temperature,light intensity,grafting and salicylic acid.The main results obtained were presented as follows:
1.Effects of salt stress on plant growth,photosynthesis,nutrient element uptake in tomato plants under different root zone temperature.
Salt stress and low root zone temperature inhibited the growth of shoots and roots, more significantly at combination of salt and low root zone temperature.Salt stress increased Na content in leaves and roots,decreased Ca content in roots,K and Mg content in leaves and roots,especially at low root temperature.Salt stress decreased chlorophyll content,photochemical quenching(qP),the efficiency of excitation energy capture by open PSII reaction center(Fv'/Fm'),electron transport rate(ETR) and the quantum efficiency of non-cyclic electron transport of PSII(ΦPSII),more significantly at low root zone temperature.As a result,the net photosynthesis rate was decreased.The promotion of photochemical quenching(NPQ) due to the increase of de-epoxidation extent of xanthophylls cycle and an elevation of water use efficiency (WUE) were observed in tomato plant under salt stress.
2.Combined effects of salinity and low root zone temperature on oxidative stress in tomato plants.
O_2~(·-) producing rate,H_2O_2 and MDA content were increased in salt-stressed plants, especially under low root zone temperature.Salt stress led to an elevation in activities of Mn-SOD and Cu,Zn-SOD,while low root zone temperature increased activities of Fe-SOD and Mn-SOD,which play its important role in removing O_2·~-.CAT were sensitive to salinity,and its activity significantly decreased under salt stress and low root temperature,while activities of GPX,APX and GR increased under salt stress and low root temperature,which indicated their important roles in scavenging reactive oxygen species in tomato tolerance to low root zone temperature and salinity.
3.Effects of salinity on photosynthesis characteristics in tomato plants under different light intensities.
Salinity inhibited plant growth and the inhibition in plants under low light intensity was less significantly compared to those under high intensity.Salt stress decreased net photosynthetic rate(Pn),stomatal conductance(Gs) and intracellular CO_2(Ci).Therefore,it could be concluded that stomatal limitation might be a dominant factor on Pn decrease under salt stress.Salt stress increased dark respiration rate(Rd) and light consumption point(LCP),decreased light saturated point(LSP). Rd,LCP and LSP decreased in salt-stressed plants under low light intensity compared with salt stressed plants under high light intensity.100 mmol·L~(-1) NaCl had no influence on Fv/Fm,decreased qP,Fv'/Fm' andΦPSII under high light intensity.The PSII activity under low light intensity was higher than that of high light intensity. ETR and Pn were much lower in low light intensity,thus the growth rate of salt stressed plant under low light intensity was much slower.
4.Effects of salinity on nutrient contents,ATPase activity in plasma membrane and ATPase,PPase activity in tonoplast of tomato under different light intensities
Salt stress increased Na content,decreased K content in tomato plants under both light intensities.Low light intensity decreased the inhibitory effects for absorption of P,Ca,Mg,Fe and Zn under salt stress.
Salt stress promoted the activities of H~+-ATPase and Ca~(2+)-ATPase in the plasma membrane of tomato roots,more significantly under high light intensity.Salt stress increased the activities of H~+-ATPase,Ca~(2+)-ATPase and H~+-PPase in the tonoplast of tomato root,more significantly under high light intensity.
5.Effects of iso-osmotic Ca(NO_3)_2 and NaCl stress on respiration and metabolism of reactive oxygen in roots of tomato seedlings
The salt type in protected greenhouse soil is different from that in coast saline soil.The main cation of accumulated salt is Ca~(2+),while the main anion is NO_3~-.Salt stress decreased the total respiration rate due to the decrease of cytochrome respiration,particularly under NaCl stress.Salt stress increased the KCN-resistant respiration rate,however,this effect was more significantly under NaCl stress.The producing rate of O_2·~- and H_2O_2 content were increased by salt stress, especially under NaCl stress.The activities of SOD,CAT and GPX were significantly increased by NaCl treatment,however,the activities decreased after 8 d of treatment.The activities of these enzymes increased all the time under Ca(NO_3)_2 treatment.From the results,we could conclude. that the increase of alternative pathway under NaCl stress might be contributed to the removal of ROS.
6.The physiological mechanism of salt toxicity in tomato mitigated by grafting
The inhibition of tomato growth increased with increasing NaCl concentration. Self-grafted plants shared the same responding with non-grafted plants under salt stress,while stock-grafted plants were more resistant to salt stress,which contributed to higher photosynthesis ability and antioxidant capacity.100 mmol·L~(-1) or 150 mmol·L~(-1) NaCl decreased qP,ETR,Fv'/Fm' andΦPSII in non-grafted plants and self-grafted plants,while not in stock-grafted plants.Thus,Pn in stock-grafted plants was higher than that of non-grafted and self-grafted plants.The MDA content increased in salt-stressed plants,more dramatically in non-grafted and self-grafted plants. Salt stress decreased CAT and DHAR activity in non-grafted and self-grafted plants,while increased in stock-grafted plants.Salt stress increased SOD and GPX activity,more significantly in non-grafted and self-grafted plants.GR activity was much higher in stock-grafted plants under different salt concentrations than that of non-grafted and self-grafted plants.Salt stress increased Na content,decreased K content in non-grafted,self-grafted and stock-grafted plants.
7.Effect of Salicylic acid on photosynthesis and the activity of ROS scavenging system under salt stress
1.0 mmol·L~(-1) SA foliar spray mitigated the inhibition of tomato plant growth caused by sat stress.One aspect was attributed to the elevation of PSII activity and enhancement of Pn under salt stress.On the other hand,SA foliar spray increased activities of antioxidant enzymes(SOD,CAT,GPX and DHAR) under salinity and increased ASC and GSH contents in tomato leaves,which could efficiently scavenging reactive oxygen species,and reduced the O_2·~- productivity rate and H_2O_2 content,and this was indicated by lower MDA content and electrolyte leakage.
8.Effects of SA on nutrient contents and plasma membrane activities of tomato roots under salt stress
SA foliar spray increased absorption of K,Ca and Mg under salt stress,decreased Na content,and thus increased K/Na ratio in tomato plants.
The activity of of H~+-ATPase and Ca~(2+)-ATPase in plasma membrane and H~+-ATPase and H~+-PPase in tonoplast were increased by salt stress and promoted further by SA foliar spray.Salt stress increased Ca~(2+)-ATPase activity in tonoplast, while there were no difference between SA foliar and not.SA foliar spray decreased NADH oxidation rate,NADPH oxidation rate,Fe(CN)_6~(3+) reduction rate and EDTA-Fe~(3+) reduction rate under salt stress,which was close to the control.
1.不同根际温度下盐胁迫对番茄光合特性和营养吸收的影响
盐胁迫和根际低温明显抑制了番茄地上部和地下部的生长,复合处理加重了抑制程度。盐胁迫打破了番茄叶片和根部的营养元素平衡,提高了番茄叶片和根系中Na含量,降低了K、Mg和根系中Ca含量,根际低温降低了番茄对K、Ca、Mg的吸收能力。盐胁迫降低了番茄叶片叶绿素a和总叶绿素含量,根际低温加剧了盐胁迫下叶绿素含量的降低。光化学猝灭系数(qP)、天线色素光能转化效率(Fv'/Fm')、电子传递速率(ETR)和表观量子效率(ΦPSⅡ)在盐胁迫后均明显降低,根际低温处理进一步降低了PSⅡ的实际活性。盐胁迫后番茄净光合速率降低,根际低温处理后,下降幅度更大。
2.根际低温和盐胁迫对番茄氧化胁迫的复合效应
盐胁迫和根际低温提高了番茄叶片O_2·~-产生速率和H_2O_2含量,造成了番茄的氧化伤害,MDA含量上升和电解质渗透率提高,在复合处理中氧化伤害程度最高。盐胁迫诱导了番茄Fe-SOD和Cu,Zn-SOD活性,根际低温诱导了Fe-SOD和Mn-SOD活性。CAT对盐胁迫敏感,盐胁迫和根际低温均降低了其活性,复合处理活性最低。GPX、APX和GR在盐胁迫和根际低温处理后活性上升,在复合处理中上升幅度较大,说明其在清除叶片活性氧方面有重要作用。盐胁迫和根际低温处理后番茄叶片谷胱甘肽和抗坏血酸含量明显降低,复合处理含量最低,说明胁迫后番茄还原态受到破坏。
3.不同光照条件下盐胁迫对番茄光合特性的影响
盐胁迫抑制了番茄生长,弱光下这种抑制程度相对较低。盐胁迫后番茄净光合速率(Pn)、气孔导度(Gs)和胞间CO_2浓度(Ci)均降低,气孔限制值(Ls)上升,表明盐胁迫引起的光合速率下降可能是由气孔限制所致。盐胁迫提高了暗呼吸速率(Rd)和光补偿点(LCP),番茄利用弱光能力降低;盐胁迫降低了光饱和点(LSP),对强光的抵御能力降低。弱光条件下,Rd、LCP和LSP均降低,利用弱光能力加强,这是番茄对弱光环境的适应。100 mmol·L~(-1)的NaCl处理对Fv/Fm没有明显影响,降低了番茄qP、Fv'/Fm'和ΦPSⅡ。弱光提高了PSⅡ的实际运行活性,但由于光照过低,ETR和净光合速率明显低于强光。
4.不同光照条件下盐胁迫对番茄元素分布、膜系统ATP酶和焦磷酸酶活性的影响
盐胁迫提高了番茄根系和叶片中Na含量,降低了K含量。弱光对盐胁迫下Na和K含量没有显著影响,提高了番茄对P、Ca、Mg、Fe和Zn的吸收能力。强光下,NaCl处理显著提高了番茄质膜H~+-ATPase和Ca~(2+)-ATPase的活性,弱光条件下,上升幅度降低。强光下,NaCl处理明显诱导了液泡膜H~+-ATPase、Ca~(2+)-ATPase和H~+-PPase活性,弱光处理降低了液泡膜ATP酶和焦磷酸酶活性。
5.不同盐分形态对番茄幼苗根系呼吸和活性氧代谢的影响
设施土壤次生盐渍化的盐分种类与滨海盐土不同,阴离子以NO_3~-为主,阳离子以Ca~(2+)为主。Ca(NO_3)_2处理和NaCl处理对番茄的毒害方式存在差异。盐胁迫下番茄根系总呼吸速率和细胞色素呼吸速率明显下降,相较Ca(NO3)_2处理,NaCl处理细胞色素呼吸下降幅度更大;在盐胁迫下抗氰呼吸速率上升,NaCl处理上升更明显。盐胁迫明显增加了活性氧含量,NaCl处理上升幅度更大。NaCl胁迫下,根系SOD、CAT和GPX活性显著增加,但在处理后期上升幅度下降,Ca(NO_3)_2胁迫后抗氧化酶活性一直上升,NaCl胁迫根系膜脂过氧化程度显著高于NaCl处理。NaCl处理抗氰呼吸的增强可以弥补细胞色素呼吸引起的ATP不足,并可能参与活性氧的清除。
6.嫁接提高番茄耐盐性机理研究
随着盐浓度的增加,番茄生长受抑制程度逐渐提高。自根嫁接对番茄生长和耐盐性没有明显影响,砧木嫁接明显缓解了盐胁迫引起的生长受阻,归因于其具有较高的光合活性和抗氧化能力。100和150 mmol·L~(-1)NaCl处理后,自根和自根嫁接番茄qP、ETR、Fv'/Fm'和ΦPSⅡ明显下降,砧木嫁接苗基本不受影响。因此,砧木嫁接苗在盐胁迫后Pn较高。盐胁迫后,砧木嫁接番茄膜脂过氧化程度较低,与抗氧化系统的响应相关。盐胁迫后,自根和自根嫁接番茄叶片CAT和DHAR活性降低,砧木嫁接番茄CAT和DHAR活性上升。盐胁迫提高了番茄叶片SOD和GPX活性,在自根和自根嫁接苗中上升幅度更大。砧木嫁接番茄具有较高的GR活性,在各盐分处理浓度下,其活性均高于自根和自根嫁接苗。盐胁迫提高了番茄叶片和根系Na含量,降低了K含量和K/Na比,砧木嫁接番茄K、Na含量与自根苗相比没有明显差异。
7.水杨酸对盐胁迫下番茄光合作用和活性氧清除系统的影响
1 mmol·L~(-1)的SA叶面喷施明显缓解了盐胁迫对植株生长的抑制作用。这种缓解效应一方面归因于SA叶面喷施缓解了盐胁迫引起的PSⅡ活性降低,电子用于CO_2固定的比例增加,提高了盐胁迫下番茄的净光合速率;另一方面,归因于SA诱导了番茄抗氧化系统活性,提高了SOD、CAT、GPX和DHAR活性,缓解了盐胁迫引起的还原型谷胱甘肽GSH和还原型抗坏血酸ASC含量的降低,降低了O_2~-的产生速率和H_2O_2含量,使盐胁迫条件下MDA含量和电解质渗透率明显降低。
8.水杨酸对盐胁迫下番茄元素分布和根系膜系统的影响
SA处理提高了番茄对K的选择吸收能力,提高了盐胁迫后番茄根系Ca和Mg含量,降低了Na含量,使番茄维持较高的K/Na比。SA处理显著提高了正常生长和胁迫条件下番茄根系质膜H~+-ATPase和Ca~(2+)-ATPase活性。SA处理提高了盐胁迫下番茄液泡膜H~+-ATPase和H~+-PPase活性,对液泡膜Ca~(2+)-ATPase活性没有明显影响。SA处理缓解了盐胁迫条件下番茄根系NADPH氧化速率、NADH氧化速率、Fe(CN)_6~(3+)还原速率和EDTA-Fe~(3+)还原速率的上升,使其接近于正常生长水平。
High concentrations of salts in soils account for large decreases in the yield of a wide variety of crops.Because of the limitation of greenhouse and bad management, crops are routinely subjected to a combination of different abiotic stresses in China. Tomato(Lycopersicon esculentum Mill) is widely planted in greenhouse and easily affected by low temperature,low light intensity and salinity.Therefore,it is very important to study the physiological mechanisms of salt toxicity in tomato and to take measures to decrease the toxicity.In this study,hydroponic culture experiments were conducted to investigate the physiological mechanisms of salt toxicity to tomato and the effects of low temperature,light intensity,grafting and salicylic acid.The main results obtained were presented as follows:
1.Effects of salt stress on plant growth,photosynthesis,nutrient element uptake in tomato plants under different root zone temperature.
Salt stress and low root zone temperature inhibited the growth of shoots and roots, more significantly at combination of salt and low root zone temperature.Salt stress increased Na content in leaves and roots,decreased Ca content in roots,K and Mg content in leaves and roots,especially at low root temperature.Salt stress decreased chlorophyll content,photochemical quenching(qP),the efficiency of excitation energy capture by open PSII reaction center(Fv'/Fm'),electron transport rate(ETR) and the quantum efficiency of non-cyclic electron transport of PSII(ΦPSII),more significantly at low root zone temperature.As a result,the net photosynthesis rate was decreased.The promotion of photochemical quenching(NPQ) due to the increase of de-epoxidation extent of xanthophylls cycle and an elevation of water use efficiency (WUE) were observed in tomato plant under salt stress.
2.Combined effects of salinity and low root zone temperature on oxidative stress in tomato plants.
O_2~(·-) producing rate,H_2O_2 and MDA content were increased in salt-stressed plants, especially under low root zone temperature.Salt stress led to an elevation in activities of Mn-SOD and Cu,Zn-SOD,while low root zone temperature increased activities of Fe-SOD and Mn-SOD,which play its important role in removing O_2·~-.CAT were sensitive to salinity,and its activity significantly decreased under salt stress and low root temperature,while activities of GPX,APX and GR increased under salt stress and low root temperature,which indicated their important roles in scavenging reactive oxygen species in tomato tolerance to low root zone temperature and salinity.
3.Effects of salinity on photosynthesis characteristics in tomato plants under different light intensities.
Salinity inhibited plant growth and the inhibition in plants under low light intensity was less significantly compared to those under high intensity.Salt stress decreased net photosynthetic rate(Pn),stomatal conductance(Gs) and intracellular CO_2(Ci).Therefore,it could be concluded that stomatal limitation might be a dominant factor on Pn decrease under salt stress.Salt stress increased dark respiration rate(Rd) and light consumption point(LCP),decreased light saturated point(LSP). Rd,LCP and LSP decreased in salt-stressed plants under low light intensity compared with salt stressed plants under high light intensity.100 mmol·L~(-1) NaCl had no influence on Fv/Fm,decreased qP,Fv'/Fm' andΦPSII under high light intensity.The PSII activity under low light intensity was higher than that of high light intensity. ETR and Pn were much lower in low light intensity,thus the growth rate of salt stressed plant under low light intensity was much slower.
4.Effects of salinity on nutrient contents,ATPase activity in plasma membrane and ATPase,PPase activity in tonoplast of tomato under different light intensities
Salt stress increased Na content,decreased K content in tomato plants under both light intensities.Low light intensity decreased the inhibitory effects for absorption of P,Ca,Mg,Fe and Zn under salt stress.
Salt stress promoted the activities of H~+-ATPase and Ca~(2+)-ATPase in the plasma membrane of tomato roots,more significantly under high light intensity.Salt stress increased the activities of H~+-ATPase,Ca~(2+)-ATPase and H~+-PPase in the tonoplast of tomato root,more significantly under high light intensity.
5.Effects of iso-osmotic Ca(NO_3)_2 and NaCl stress on respiration and metabolism of reactive oxygen in roots of tomato seedlings
The salt type in protected greenhouse soil is different from that in coast saline soil.The main cation of accumulated salt is Ca~(2+),while the main anion is NO_3~-.Salt stress decreased the total respiration rate due to the decrease of cytochrome respiration,particularly under NaCl stress.Salt stress increased the KCN-resistant respiration rate,however,this effect was more significantly under NaCl stress.The producing rate of O_2·~- and H_2O_2 content were increased by salt stress, especially under NaCl stress.The activities of SOD,CAT and GPX were significantly increased by NaCl treatment,however,the activities decreased after 8 d of treatment.The activities of these enzymes increased all the time under Ca(NO_3)_2 treatment.From the results,we could conclude. that the increase of alternative pathway under NaCl stress might be contributed to the removal of ROS.
6.The physiological mechanism of salt toxicity in tomato mitigated by grafting
The inhibition of tomato growth increased with increasing NaCl concentration. Self-grafted plants shared the same responding with non-grafted plants under salt stress,while stock-grafted plants were more resistant to salt stress,which contributed to higher photosynthesis ability and antioxidant capacity.100 mmol·L~(-1) or 150 mmol·L~(-1) NaCl decreased qP,ETR,Fv'/Fm' andΦPSII in non-grafted plants and self-grafted plants,while not in stock-grafted plants.Thus,Pn in stock-grafted plants was higher than that of non-grafted and self-grafted plants.The MDA content increased in salt-stressed plants,more dramatically in non-grafted and self-grafted plants. Salt stress decreased CAT and DHAR activity in non-grafted and self-grafted plants,while increased in stock-grafted plants.Salt stress increased SOD and GPX activity,more significantly in non-grafted and self-grafted plants.GR activity was much higher in stock-grafted plants under different salt concentrations than that of non-grafted and self-grafted plants.Salt stress increased Na content,decreased K content in non-grafted,self-grafted and stock-grafted plants.
7.Effect of Salicylic acid on photosynthesis and the activity of ROS scavenging system under salt stress
1.0 mmol·L~(-1) SA foliar spray mitigated the inhibition of tomato plant growth caused by sat stress.One aspect was attributed to the elevation of PSII activity and enhancement of Pn under salt stress.On the other hand,SA foliar spray increased activities of antioxidant enzymes(SOD,CAT,GPX and DHAR) under salinity and increased ASC and GSH contents in tomato leaves,which could efficiently scavenging reactive oxygen species,and reduced the O_2·~- productivity rate and H_2O_2 content,and this was indicated by lower MDA content and electrolyte leakage.
8.Effects of SA on nutrient contents and plasma membrane activities of tomato roots under salt stress
SA foliar spray increased absorption of K,Ca and Mg under salt stress,decreased Na content,and thus increased K/Na ratio in tomato plants.
The activity of of H~+-ATPase and Ca~(2+)-ATPase in plasma membrane and H~+-ATPase and H~+-PPase in tonoplast were increased by salt stress and promoted further by SA foliar spray.Salt stress increased Ca~(2+)-ATPase activity in tonoplast, while there were no difference between SA foliar and not.SA foliar spray decreased NADH oxidation rate,NADPH oxidation rate,Fe(CN)_6~(3+) reduction rate and EDTA-Fe~(3+) reduction rate under salt stress,which was close to the control.
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