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红树林湿地一氧化氮释放及其对红树植物耐盐性的调控研究 【作者】肖强 【导师】郑海雷

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&|!g*NE0`@Ft0红树林湿地一氧化氮释放及其对红树植物耐盐性的调控研究水利论文/Y;RXr;m-h"j
Nitric Oxide Emission from Mangrove Wetland and Regulation of NO on Salt Tolerance for Mangrove Plants水利论文%uA sy2CZg.w4BY

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作者肖强导师】郑海雷
A!}6b y0F*U#J0【作者基本信息】厦门大学,植物学,2007年,博士

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【中文摘要】 本文研究了外源NO供体硝普钠(sodium nitroprusside,SNP)释放NO规律以及SNP浓度、光照强度、温度、溶液pH值和振荡频率对其释放过程的影响;采用化学发光法研究了秋茄、白骨壤和木榄叶片在不同氮源、光照强度和氧气供应下NO气体释放速率的变化规律;采用静态箱-气袋采样法研究了红树林湿地NO释放通量的日变化以及环境因素对红树林湿地NO释放通量的影响;在这些研究基础上,详细探讨了NO对桐花树气孔运动的调控以及NO对红树植物白骨壤、桐花树、木榄和秋茄耐盐性的调控。主要结论如下:1.本研究表明,SNP释放NO的速率受SNP浓度、温度、振荡频率、光照强度以及溶液pH等多种因素综合影响,其中SNP浓度以及温度与NO释放速率之间剂量关系可用Logistic曲线方程描述;溶液pH与NO释放速率之间剂量关系符合指数递减模型(ExpDec1);而光照强度以及振荡频率与NO释放速率之间剂量关系则可用指数增长模型(ExpGro1)描述。通过本研究揭示了环境因子对外源NO供体释放NO规律的影响,为合理解释外源NO供体对植物的剂量效应提供了依据。2.首次采用NO和NO2化学发光分析法研究了木榄...更多、秋茄和白骨壤离体叶片在不同氮源、光照强度和氧气供应条件下NO释放速率变化规律。结果表明:在缺乏氮源供应、以铵态氮作为氮源和以精氨酸作为氮源时,木榄叶片在各种情况下均无NO释放;以亚硝酸盐作为氮源时,在黑暗条件下NO释放速率均要高于在光照情况下NO释放速率。在缺氧情况下,NO释放要高于有氧情况;并且,在供氧充足时,NO释放速率要高出以硝酸盐作为底物时7倍以上。在离体秋茄叶片中,当以硝酸盐为氮源情况下,并且在有氧情况下,叶片几乎不释放NO,只是从光-暗转换过程中有NO释放,而缺氧则促进了NO释放,无氧情况下,光暗转换中NO释放速率达到光照下10倍以上;而在亚硝酸盐为氮源情况下,各种条件下均有NO释放,而氧气供应似乎对秋茄叶片NO释放速率影响并不如木榄中显著。在白骨壤离体叶片中,厌氧似乎抑制了黑暗下NO的释放。这些研究表明红树植物释放NO的速率受氮素供应形式、光照强度以及氧气供应等因素制约。3.首次采用静态箱气袋采样法以及NO和NO2化学发光分析法观测研究了福建九龙江红树林自然保护区红树林湿地NO释放通量的日变化规律。分别对裸地光滩的高潮带、中潮带和低潮带,木榄林地的高潮带和中潮带(低潮带无木榄分布)NO释放通量进行了研究。结果表明:在不受潮水淹没情况下,光滩NO释放表现为日间极大值型,且极大值位于上午9时与午后14时之间。进一步,NO释放与光强没有相关性,NO通量变化总体来说受温度影响,高温可以促进NO释放。木榄林下中潮带NO释放通量表现为日间极大值型,且最大值出现在早上7-9时;木榄林下高潮带NO释放通量具有和光滩完全不同的特点,其极大值发生在下午15时和夜间23时,并不表现出明显的的规律性。NO通量日变化极大值与土壤温度、电导率、盐度、样地pH及Eh之间均无单一的相关关系。4.探索一种用材简单、操作方便、真实性强的观察桐花树叶片气孔的制片技术,并利用该技术研究不同浓度、不同时间的SNP处理对桐花树气孔开闭的影响,探讨了NO调控的气孔运动与外源Ca2+的关系以及NO与H2O2在调节气孔运动过程中的效应。结果表明:搅碎法、指甲油印迹法、牛皮胶印迹法三种观察气孔方法中,牛皮胶印迹法是观察气孔开度变化的最佳方法。NO能够诱导桐花树气孔的快速关闭,并且NO的作用表现出明显的时间效应与浓度效应。NO导的气孔关闭与高浓度Ca2+的参与有密切关系,NO与H2O2存在明显的协同效应,可以促进气孔以更快的速度关闭。5.研究了在600 mmol·L-1NaCl处理下,外源一氧化氮供体SNP对白骨壤幼苗叶片中叶绿素和游离脯氨酸含量,O2-·、丙二醛(malondialdehyde,MDA)和过氧化氢(hydrogen peroxide,H2O2)水平、抗氧化酶活性以及抗氧化小分子抗坏血酸(Ascorbate Acid,AsA)、多酚、还原型谷胱甘肽(reduced glutathione,GSH)和类胡萝卜素(carotenoid,Car)等含量的影响。结果表明:100μmol·L-1SNP可以显著缓解盐胁迫引起的白骨壤叶片叶绿素含量下降,促进脯氨酸含量降低。NO也可以增强超氧化物歧化酶(superoxide dismutase,SOD)、愈创木酚过氧化酶(guaiacol peroxidase,GPX)和抗坏血酸过氧化物酶(ascorbate peroxidase,APX)活性,从而显著减少NaCl处理下白骨壤叶片中O2-·和H2O2的积累;NO对脂氧合酶(lipoxygenase,LOX)活性的抑制,减轻了盐胁迫下白骨壤膜脂过氧化程度。SNP虽然促进了AsA消耗,但同时也提高了GSH、多酚和Car等抗氧化小分子含量,从而缓解了NaCl处理下白骨壤叶片的活性氧胁迫。APX和过氧化氢酶(catalase,CAT)等活性氧(reactive oxygen species,ROS)代谢相关酶可能作为NO的直接靶子受到NO调控,从而在白骨壤耐盐中发挥重要作用。6.研究了生长三个月的桐花树幼苗,在含350 mmol·L-1NaCl的溶液中培养下,NO对盐胁迫下抗氧化酶的活性,光合色素、ASA、GSH及叶片中多酚含量以及蛋白表达的影响。结果表明:当加入100μmol·L-1SNP后,经盐处理的植株叶片中,总叶绿素和类胡萝卜素含量增加。NO促进了叶绿体的发育,提高了叶绿体基质片层致密度。NO通过改变小分子抗氧化剂含量和抗氧化酶活性进而减轻桐花树叶片的氧化性损伤。一方面,在盐处理的叶片中ASA和多酚含量都有所增加;另一方面,在NO处理下,SOD和CAT活性的变化不大,而当加入100μmol·L-1SNP后,GPX活性明显增加。研究提示,NO提高桐花树的耐盐能力主要在于其调节小分子抗氧化剂含量和GPX的活性。在NaCl处理下,NO可以诱导桐花树叶片中某些蛋白表达水平上调,这些蛋白可能参与了NO对桐花树耐盐性的调控。7.研究了在15‰和30‰NaCl处理下,SNP对秋茄幼苗叶片中叶绿素含量,渗透调节物质可溶性糖、淀粉、游离氨基酸和脯氨酸含量、MDA和H2O2水平、抗氧化酶活性以及抗氧化小分子多酚和Car等含量的影响。结果表明:100μmol·L-1SNP可以显著提高NaCl处理下秋茄叶片叶绿素含量;促进脯氨酸含量上升。增强秋茄渗透调节能力。NO也可以增强GPX和CAT活性,从而显著减少NaCl处理下白骨壤叶片中H2O2的积累;减轻了盐胁迫下白骨壤膜脂过氧化程度。和桐花树不同的是,除了GPX,CAT活性也受到NO调控,从而在秋茄耐盐中发挥重要作用。8.系统研究了在45天砂培情况下,SNP对500 mmol·L-1盐胁迫下木榄幼苗叶片中叶绿素和脯氨酸含量、MDA和水平、抗氧化酶活性和同功酶表达水平以及抗氧化小分子ASA和Car等含量的影响。结果表明:100μmol·L-1SNP可以维持NaCl处理下秋茄叶片叶绿素含量;促进脯氨酸含量上升,增强木榄渗透调节能力。NO也可以增强GPX、APX和CAT活性,从而显著减少NaCl处理下木榄叶片中H2O2的积累;减轻了盐胁迫下木榄膜脂过氧化程度。但NO并不引起特异定同功酶条带的产生或消失。在木榄耐盐机制中,与H2O2清除相关的三种重要的酶GPX、APX和CAT都作为NO的直接靶子受到NO调控。  还原水利论文_sx!J I%j)A.L"fMR([

x`(f;I[sPo0【英文摘要】 The aim of the presented studies was to investigate the rate of NO release of the most frequently applied NO donors sodium nitroprusside(SNP).The influence of SNP concentration,light,temperature,pH and surge frequency on the rate of NO release was investigated.The rate of NO release from leaves of mangrove,including Bruguiera gymnorrhiza,Kandelia candel and Avicennia marina under the influence of different nitrogen sources,light intensity,oxygen was observed.Using static chamber-gas bag techniques and a chemiluminescence NOx analyzer manufactured by Teledyne Inc.USA(model 200A),NO emission from mangrove wetland and diurnal variation pattern under the influence of environmental factor were carefully monitored. Based on above results,effect of NO on the leaf stomatal movement of Aegiceras corniculatum and the order and mechanism of NO on salt resistance in A.marina,A. corniculatum,K.candel and B.gymnorrhiza were investigated.1.The rate of NO release was affected by SNP concentration,lig...更多ht intensity, temperature and pH of solution and shake frequency.Dose-dependent correlation between the rate of NO release and SNP concentration,temperature were fitted to Logistic equation while dose-dependent correlation between the rate of NO release and pH of solution were fitted to exponential decrease equation.However,light intensity,shake frequency were fitted to exponential increase equation.Based on above results,it is likely to provide clue to interpret dose-dependent correlation between NO donor and physiological effect.2.Using chemiluminescence NOx analyzer,the rate of NO release from leaves of mangrove,including B.gymnorrhiza,K.candel and A.marina under the influence of different nitrogen source,light,oxygen was investigated.Leaves of B.gymnorrhiza grown on nitrogen absence or ammonia,arginine as the N source did not emit NO, either in air or under anoxia,which was interpreted to indicate that nitrate reducase(NR)was the only NO source.However,when supplied with nitrite,NO emission in dark were higher than in light.Furthermore,NO emission increased strongly in anoxia.Under air condition,NO emission rate in leaves of B.gymnorrhiza grown on nitrite as N source were more than 7 times higher than those grown on nitrate as N source.Detached leaves of K.candel grown on nitrate as N source, emitted only small amount of NO in air whereas NO emission increased strongly under anoxia.NO emission rate in the light-dark was morn than 10 times higher than in the light under air condition.NO emission occurred in different treatment in K. candel leaves from nitrite-fed plants.Furthermore,NO emission rate increased in the light under air condition.However,leaves of A.marina from nitrite-fed plants emitted a little NO in the dark under anoxia condition.Our results suggested that different species of mangrove showed diverse NO emission model in response to nitrogen supply,light intensity and oxygen.Whether these variations were correlated to different regulate mechanism in response to environmental stress among different species were remained to be studied.3.Using static chamber-gas bag techniques and a chemiluminescence NOx analyzer,NO emission from mangrove wetland in Mangrove Nature Reserves Jiulongjiang estuarine,Fugong Town,Longhai County,Fujian Province(24°29'N, 17°55'E)and diurnal variation pattern under the influence of environmental factor were carefully monitored.Our results suggest that during the period of non-waterlogging,the diurnal variation pattern in nude beach,with the maximum emission at 9:00-14:00,was determined by temperature.There were no significant correlations between light intensity and NO emission in nude beach.The diurnal variation pattern in mid tidal zones of B.gymnorrhiza region,with the maximum emission at 7:00-9:00,was not correlated with temperature variation.However,the diurnal variation pattern in high tidal zones of B.gymnorrhiza region,with the maximum emission at 15:00-23:00,was different from mid tidal zones and nude beach.Furthermore,there were no significant correlations between soil temperature, conductance,salinity,pH,Eh and NO emission from soil.4.The experiment is mainly to find out a best way to observe the stomatal movement of A.corniculatum,based on it to study the effects of NO on the leaf stomatal movement under different concentrations and different treated time,to study the effects of Ca2+in the regulation of stomatal movement induced by exogenous nitric oxide,to study the interactional effects of NO and H2O2 in the regulation of stomatal movement.Our results suggest that the oxhide pastern is the best way to observe stomatal movement of A.corniculatum compared with the puddle method and the nail poish pastern.The inductions of stomatal closure by NO were enhanced by higher concerntration of Ca2+,and the role of NO appears to demonstrate to be obvious time and concentration effects.We also prove that H2O2 could probably act as upstream component of NO signaling and NO regulate H2O2 generation in stomatal closure in guard cells.5.The contents of photosynthetic pigment and proline,producing rate of superoxide radicals,contents of H2O2,MDA,AsA,GSH,polyphenol and Car, activities of the antioxidant enzymes involved in ROS scavenging invoked by SNP, were surveyed in A.marina seedling leaves under 600 mmol·L-1NaCl stress.The results showed that the decrease in content of total chlorophyll in NaCl-treated seedlings was alleviated when 100μmol·L-1SNP was supplied.NO promoted decrease in proline content.The activities of SOD,GPX and APX were promoted by NO,thus reduced the producing rate of superoxide radicals and content of H2O2 in A. marina leaves under NaCl stress.Meanwhile,Lipid peroxidation was alleviated by inhibited LOX activity.In addition,the oxidation of AsA was enhanced as well as contents of GSH,polyphenol and Car was increased by SNP treatment.It was also shown that APX and CAT are likely to be the major NO targets,which activities were regulated by NO,thus play important role in A.marina salt-tolerance.6.Three-month-old healthy seedlings of A.corniculatum a true mangrove,raised from propagules in normal nursery conditions were subjected to 350 mmol·L-1NaCl under hydroponic culture.SNP was applied to investigate the effect of NO on activation potentials for antioxidative enzymes against salt stress.Changes in the contents of photosynthetic pigments,ASA,GSH and polyphenol in leaves,activities of the antioxidative enzymes including CAT,GPX and SOD in leaves were assayed to evaluate the effects of NO.Also,expression of proteins in leaves of A.corniculatum induced by NO were analyzed by two-dimensional gel electrophoresis(2-DE).The results showed that the content of total chlorophyll and carotenoid increased in NaCl-treated seedlings when 100μmol·L-1SNP was supplied.NO treatment increased amount of thylakoid staking in the plastids.NO relieved the oxidative damage in leaves of A.corniculatum by changing the content of small molecule antioxidant and the activity of antioxidant enzyme.Firstly,NO treatment increased ASA and polyphenol content in NaCl-treated leaves,secondly,activities of SOD and CAT did not show change under NO treatment whereas GPX activity increased significantly when 100μmol·L-1SNP was supplied.In our present study,NO enhanced salt tolerance in A.corniculatum mainly due to its regulating ability on the content of small molecule antioxidant and GPX activity.Furthermore,five up-regulated protein spots was observed in NO treated plants which suggested these protein may involved in salt resistance regulated by NO.7.Changes in the contents of photosynthetic pigments,soluble sugar,starch,total free amino acid,proline and polyphenol,levels of MDA and H2O2 in K.candel leaves, activities of the antioxidative enzymes including CAT,GPX and SOD in K.candel leaves were assayed to evaluate the effects of NO under 15‰and 30‰NaCl treatment.The results showed that the content of total chlorophyll and proline increased in NaCl-treated seedlings when 100μmol·L-1SNP was supplied.NO relieved the oxidative damage in leaves of K.candel by enhancing activities of GPX and CAT,thus reduced H2O2 content,alleviate lipid peroxidation.GPX and CAT are likely to be the major NO targets,which play important role in K.candel under salt stress.8.Three-month-old healthy seedlings of B.gymnorrhiza a true mangrove,raised from propagules in normal nursery conditions were subjected to 500 mmol·L-1NaCl under sand culture during 45 days.SNP was applied to investigate the effect of NO on activation potentials for antioxidative enzymes against salt stress.Changes in the contents of photosynthetic pigments,proline,ASA and Car,level of MDA and H2O2 in leaves,activities of CAT,GPX,APX and SOD in leaves were assayed to evaluate the effects of NO.The results showed that the content of proline increased.NO reduced H2O2 content by enhancing activities of GPX,APX and CAT.Furthermore, NO regulate antioxidative enzymes mainly by regulating antioxdative enzymes activites instead of altering the express of specific isoform bands.  还原

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HA*T#B;a jV*E0【中文关键词】 红树林; 一氧化氮; 耐盐水利论文7j FASa Jt~
【英文关键词】 mangrove; nitric oxide; salt-tolerance
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TAG: 红树林 湿地 植物 肖强
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