硫化氢对镉胁迫下小麦幼苗的生长及生理指标的影响
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摘要
为了解硫化氢(H_2S)对Cd~(2+)胁迫下小麦幼苗生长的缓解机理,以小麦品种苏科麦一号为材料,在水培条件下,测定外源H_2S(供体为NaHS溶液)预处理对Cd~(2+)胁迫下(CdCl_2浓度为50μmol·L~(-1))小麦幼苗生长和生理指标的影响。结果显示,外源H_2S预处理显著提高Cd~(2+)胁迫下小麦幼苗的株高、地上部生物量、非蛋白巯基化合物含量、叶片光合色素含量、净光合速率、叶绿素荧光相关指标(PSⅡ的光合性能指数、单位激发态面积吸收的能量、用于还原Q_A和用于电子传递的能量、非光化学淬灭)和根部Cd含量;显著降低小麦叶片中的过氧化物质(丙二醛、过氧化氢和超氧根阴离子)含量及地上部分Cd含量和Cd转运系数。综上所述,外源H_2S预处理可清除Cd~(2+)胁迫小麦幼苗叶片中的活性氧自由基,减少其对光合色素分子的破坏,降低膜脂过氧化程度,保护光合电子传递系统,增加热耗散,提高光合能力,缓解Cd~(2+)对小麦幼苗生长的抑制,以50μmol·L~(-1)的NaHS效果最好。
To explore the beneficial effect of hydrogen sulfide(H_2S) on the growth of wheat seedlings under Cd~(2+) stress,a wheat cultivar Sukemai 1 was used as the experimental material in this study.The effects of H_2S(different concentrations of NaHS were served as H_2S donor) on the growth and physiological changes of wheat seedlings under Cd~(2+) stress(CdCl_2 concentration of 50 μmol·L~(-1)) were studied through hydroponic method.The results showed that exogenous H_2S significantly improved the plant height,shoot biomass,non-protein thiol content,photosynthetic pigment contents,net photosynthetic rate,chlorophyll fluorescence parameters including photosynthetic performance index of PS II,the absorption of light energy,trapped energy fluxes for reducing Q_A,the electron transport in PS II cross-section and non-photochemical quenching of wheat leaves under Cd~(2+) stress.Cd concentration in roots were also increased.While the content of antioxidants,such as malonaldehyde,hydrogen peroxide and superoxide anion in wheat leaves,Cd concentration in shoots,and Cd transport factor were significantly reduced by pretreatment of H_2S on wheat seedlings under Cd~(2+) stress.In summary,the harmful effects of Cd~(2+) on wheat seedlings were mitigated by exogenous H_2S through scavenging reactive oxygen free radicals in the leaves to decrease the destruction of photosynthetic pigments,to reduce the damage from peroxidation of membrane lipid,to protect the photosynthetic electron transfer system,to increase the heat dissipation,as well as to improve the photosynthetic capacity,and to alleviate the growth inhibition caused by Cd~(2+),and the optimal NaHS concentration was 50 μmol·L~(-1).
To explore the beneficial effect of hydrogen sulfide(H_2S) on the growth of wheat seedlings under Cd~(2+) stress,a wheat cultivar Sukemai 1 was used as the experimental material in this study.The effects of H_2S(different concentrations of NaHS were served as H_2S donor) on the growth and physiological changes of wheat seedlings under Cd~(2+) stress(CdCl_2 concentration of 50 μmol·L~(-1)) were studied through hydroponic method.The results showed that exogenous H_2S significantly improved the plant height,shoot biomass,non-protein thiol content,photosynthetic pigment contents,net photosynthetic rate,chlorophyll fluorescence parameters including photosynthetic performance index of PS II,the absorption of light energy,trapped energy fluxes for reducing Q_A,the electron transport in PS II cross-section and non-photochemical quenching of wheat leaves under Cd~(2+) stress.Cd concentration in roots were also increased.While the content of antioxidants,such as malonaldehyde,hydrogen peroxide and superoxide anion in wheat leaves,Cd concentration in shoots,and Cd transport factor were significantly reduced by pretreatment of H_2S on wheat seedlings under Cd~(2+) stress.In summary,the harmful effects of Cd~(2+) on wheat seedlings were mitigated by exogenous H_2S through scavenging reactive oxygen free radicals in the leaves to decrease the destruction of photosynthetic pigments,to reduce the damage from peroxidation of membrane lipid,to protect the photosynthetic electron transfer system,to increase the heat dissipation,as well as to improve the photosynthetic capacity,and to alleviate the growth inhibition caused by Cd~(2+),and the optimal NaHS concentration was 50 μmol·L~(-1).
引文
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[27]TIAN B H,QIAO Z J,ZHANG L P,et al.Hydrogen sulfide and proline cooperate to alleviate cadmium stress in foxtail millet seedlings [J].Plant Physiology & Biochemistry,2016,109:297.
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[35]ZIVCAK M,BRESTIC M,OLSOVSKA K,et al.Performance index as a sensitive indicator of water stress in Triticum aestivum L.[J].Plant Soil & Environment,2008,4:138.
[36]KALAJI H M,GOVINDJEE,BOSA K,et al.Effects of salt stress on photosystem II efficiency and CO2 assimilation in two Syrian barley landraces [J].Environmental & Experimental Botany,2011,73:69.
[37]HU K D,BAI G S,LI W J,et al.Sulfur dioxide promotes germination and plays an antioxidant role in cadmium-stressed wheat seeds [J].Plant Growth Regulation,2015,75:276.
[38]CHEN Z,CHEN M,JIANG M.Hydrogen sulfide alleviates mercury toxicity by sequestering it in roots or regulating reactive oxygen species productions in rice seedlings [J].Plant Physiology & Biochemistry,2017,111:183.
[39]KOTRBA P,NAJMANOVA J,MACEK T,et al.Genetically modified plants in phytoremediation of heavy metal and metalloid soil and sediment pollution [J].Biotechnology Advances,2009,27:802.
[40]KABIL O,MOTL N,BANERJEE R.H2S and its role in redox signaling [J].Biochim Biophys Acta,2014,1844:1357.
[41]ANJUM N A,AHMAD I,MOHMOOD I,et al.Modulation of glutathione and its related enzymes in plants' responses to toxic metals and metalloids—A review [J].Environmental & Experimental Botany,2012,75:307.
[2]尚二萍,许尔琪,张红旗,等.中国粮食主产区耕地土壤重金属时空变化与污染源分析[J].环境科学,2018(10):6. SHANG E P,XU E Q,ZHANG H Q,et al.Spatial-temporal trends and pollution source analysis in heavy metal contamination of cultivated soils in five major grain producing regions of China [J].Environmental Science,2018(10):6.
[3]SHIM D,HWANG J U,LEE J,et al.Orthologs of the class A4 heat shock transcription factor HsfA4a confer cadmium tolerance in wheat and rice [J].Plant Cell,2009,21(12):4031.
[4]LIN Y F,AARTS M G M.The molecular mechanism of zinc and cadmium stress response in plants [J].Cellular & Molecular Life Sciences,2012,69(19):3187.
[5]CLEMENS S,AARTS M G M,THOMINE S,et al.Plant science:The key to preventing slow cadmium poisoning [J].Trends in Plant Science,2013,18(2):92.
[6]DIAS M C,MONTEIRO C,MOUTINHO-PEREIRA J,et al.Cadmium toxicity affects photosynthesis and plant growth at different levels [J].Acta Physiologiae Plantarum,2013,35(4):1281.
[7]孟自力,贾斌,尹海燕,等.镉胁迫对小麦生长发育的影响[J].中国农学通报,2018,34(23):28. MENG Z L,JIA B,YIN H Y,et al.Effects of cadmium stress on the growth and development of wheat [J].Chinese Agricultural Science Bulletin,2018,34(23):28.
[8]MUSTAFA A K,GADALLA M M,SNYDER S H.Signaling by gasotransmitters [J].Science Signaling,2009,2(68):re2. Doi:10.1126/scisignal.268re2.
[9]WAGNER C A.Hydrogen sulfide:A new gaseous signal molecule and blood pressure regulator [J].Journal of Nephrology,2009,22(2):173.
[10]TAN B H,WONG P T,BIAN J S.Hydrogen sulfide:A novel signaling molecule in the central nervous system [J].Neurochemistry International,2010,56(1):3.
[11]WILSON L G,BRESSAN R A,FILNER P.Light-dependent emission of hydrogen sulfide from plants [J].Plant Physiology,1978,61(2):186.
[12]DAWOOD M,CAO F B,JAHANGIR M M,et al.Alleviation of aluminum toxicity by hydrogen sulfide is related to elevated ATPase,and suppressed aluminum uptake and oxidative stress in barley [J].Journal of Hazardous Materials,2012,209-210(1):124.
[13]SINGH V P,SINGH S,KUMAR J,et al.Hydrogen sulfide alleviates toxic effects of arsenate in pea seedlings through up-regulation of the ascorbate-glutathione cycle:Possible involvement of nitric oxide [J].Journal of Plant Physiology,2015,181(51):25.
[14]FANG H H,LIU Z Q,JIN Z P,et al.An emphasis of hydrogen sulfide-cysteine cycle on enhancing the tolerance to chromium stress in Arabidopsis [J].Environmental Pollution,2016,213:873.
[15]MOSTOFA M G,RAHMAN A,ANSARY M M U,et al.Hydrogen sulfide modulates cadmium-induced physiological and biochemical responses to alleviate cadmium toxicity in rice [J].Scientific Reports,2014,5(14078):4.
[16]SUN J,WANG R,ZHANG X,et al.Hydrogen sulfide alleviates cadmium toxicity through regulations of cadmium transport across the plasma and vacuolar membranes in Populus euphratica cells [J].Plant Physiol Biochem,2013,65(6):68.
[17]ZHAO F J,JIANG R F,DUNHAM S J,et al.Cadmium uptake,translocation and tolerance in the hyperaccumulator Arabidopsis helleri [J].New Phytologist,2006,172(4):647.
[18]LICHTENTHALER H K.Chlorophylls and carotenoids:Pigments of photosynthetic biomembranes [J].Methods in Enzymology,1987,148C(1):362.
[19]JIN Q J,ZHU K K,CUI W T,et al.Hydrogen gas acts as a novel bioactive molecule in enhancing plant tolerance to paraquat‐induced oxidative stress via the modulation of heme oxygenase-1 signalling system [J].Plant Cell & Environment,2013,36(5):958.
[20]VELIKOVA V,YORDANOV I,EDREVA A.Oxidative stress and some antioxidant systems in acid rain-treated bean plants:Protective role of exogenous polyamines [J].Plant Science,2000,151(1):60.
[21]JIANG M Y,ZHANG J H.Effect of abscisic acid on active oxygen species,antioxidative defence system and oxidative damage in leaves of maize seedlings [J].Plant & Cell Physiology,2001,42(11):1272.
[22]XIA X J,WANG Y J,ZHOU Y H,et al.Reactive oxygen species are involved in brassinosteroid-induced stress tolerance in cucumber [J].Plant Physiology,2009,150(2):812.
[23]RICDEVOS C H,VONK M J,VOOIJS R,et al.Glutathione depletion due to copper-induced phytochelatin synthesis causes oxidative stress in Silene cucubalus [J].Plant Physiology,1992,98(3):854.
[24]WANG Y,WANG C,LIU Y,et al.GmHMA3 sequesters Cd to the root endoplasmic reticulum to limit translocation to the stems in soybean [J].Plant Science,2018,270:23.
[25]XIE Y,LI X,LIU X,et al.Application of Aspergillus aculeatus,to rice roots reduces Cd concentration in grain [J].Plant & Soil,2018(422):410.
[26]ZHOU J,HAO M,LIU Y,et al.Effects of exogenous sulfur on growth and Cd uptake in Chinese cabbage(Brassica campestris spp.pekinensis) in Cd-contaminated soil [J].Environmental Science & Pollution Research,2018(25):15823.
[27]TIAN B H,QIAO Z J,ZHANG L P,et al.Hydrogen sulfide and proline cooperate to alleviate cadmium stress in foxtail millet seedlings [J].Plant Physiology & Biochemistry,2016,109:297.
[28]BHARWANA S A,ALI S,FAROOQ M A,et al.Hydrogen sulfide ameliorates lead-induced morphological,photosynthetic,oxidative damages and biochemical changes in cotton [J].Environmental Science & Pollution Research,2014,21(1):723.
[29]CHEN J,WU F H,WANG W H,et al.Hydrogen sulphide enhances photosynthesis through promoting chloroplast biogenesis,photosynthetic enzyme expression,and thiol redox modification in Spinacia oleracea seedlings [J].Journal of Experimental Botany,2011,62(13):4485.
[30]KHAN M I R,NAZIR F,ASGHER M,et al.Selenium and sulfur influence ethylene formation and alleviate cadmium-induced oxidative stress by improving proline and glutathione production in wheat [J].Journal of Plant Physiology,2015,173:13.
[31]KHAN N,ASGHER M,PER T,et al.Ethylene potentiates sulfur-mediated reversal of cadmium inhibited photosynthetic responses in mustard [J].Frontiers in Plant Science,2016,7:12.
[32]LIU L,SUN H,CHEN J,et al.Effects of cadmium(Cd) on seedling growth traits and photosynthesis parameters in cotton(Gossypium hirsutum L.) [J].Plant Omics,2014,7(4):284.
[33]YANG C W,PENG C L,DUAN J,et al.Responses of chlorophyll fluorescence and carotenoids biosynthesis to high light stress in rice seedling leaves at different leaf position [J].Acta Botanica Sinica,2002,44(11):1303.
[34]KALAJI H M,JAJOO A,OUKARROUM A,et al.Chlorophyll a,fluorescence as a tool to monitor physiological status of plants under abiotic stress conditions [J].Acta Physiologiae Plantarum,2016,38(4):102.
[35]ZIVCAK M,BRESTIC M,OLSOVSKA K,et al.Performance index as a sensitive indicator of water stress in Triticum aestivum L.[J].Plant Soil & Environment,2008,4:138.
[36]KALAJI H M,GOVINDJEE,BOSA K,et al.Effects of salt stress on photosystem II efficiency and CO2 assimilation in two Syrian barley landraces [J].Environmental & Experimental Botany,2011,73:69.
[37]HU K D,BAI G S,LI W J,et al.Sulfur dioxide promotes germination and plays an antioxidant role in cadmium-stressed wheat seeds [J].Plant Growth Regulation,2015,75:276.
[38]CHEN Z,CHEN M,JIANG M.Hydrogen sulfide alleviates mercury toxicity by sequestering it in roots or regulating reactive oxygen species productions in rice seedlings [J].Plant Physiology & Biochemistry,2017,111:183.
[39]KOTRBA P,NAJMANOVA J,MACEK T,et al.Genetically modified plants in phytoremediation of heavy metal and metalloid soil and sediment pollution [J].Biotechnology Advances,2009,27:802.
[40]KABIL O,MOTL N,BANERJEE R.H2S and its role in redox signaling [J].Biochim Biophys Acta,2014,1844:1357.
[41]ANJUM N A,AHMAD I,MOHMOOD I,et al.Modulation of glutathione and its related enzymes in plants' responses to toxic metals and metalloids—A review [J].Environmental & Experimental Botany,2012,75:307.