麝香草酚调控水稻幼苗根的耐盐作用
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摘要
外源调控作物耐盐是保证盐渍化土壤中农作物安全生产的重要措施.新型植物源天然化合物麝香草酚作为一种医药活性物质已有大量报道,但其对植物的生理调控活性却鲜有报道.本文以水稻幼苗为研究材料,通过植物生理学、组织化学、生物化学等手段,探讨了麝香草酚调控水稻抗盐胁迫的作用方式.结果显示:(1)外源麝香草酚处理能够显著缓解盐胁迫对水稻幼苗根的抑制,并呈现出一定的浓度效应;(2)外源麝香草酚处理显著缓解了盐胁迫诱导的根尖细胞死亡效应;(3)外源麝香草酚处理有效清除了盐胁迫诱导根尖过量累积的过氧化氢和超氧阴离子;(4)盐胁迫显著抑制根尖超氧化合物歧化酶(SOD)和过氧化氢酶(CAT)活性,而添加麝香草酚则显著提升了盐胁迫下这两种抗氧化酶的活力;(5)外源麝香草酚处理显著提高了盐胁迫下根尖抗坏血酸(As A)和谷胱甘肽(GSH)的含量.这些结果表明,麝香草酚能够通过提升水稻幼苗根尖的抗氧化能力,抵御盐胁迫诱导的氧化损伤和细胞死亡效应.本研究对盐渍化土壤中水稻的安全生产提供了重要的科学依据和技术手段.
Exogenous regulation of salt tolerance is an important approach for the safety of crop production in salinity soil.Thymol is a novel plant-derived natural compound,which has been extensively reported to have medicinal activity.However,the regulation of plant physiology by thymol is still elusive.In the present study,physiological,histochemical,and biochemical methods were used to investigate thymol-modulated salt tolerance in rice seedlings.Basically,the results showed that,(1) Exogenous treatment with thymol significantly attenuated salt-induced root inhibition in a dose-dependent manner.(2) Exogenous treatment with thymol remarkably blocked salt-induced cell death in root tip.(3) Exogenous treatment with thymol scavenged the over-generated hydrogen peroxide and superoxide radical in root tip under salt stress.(4)Salt stress significantly inhibited the activity of superoxide dismutase(SOD)and catalase(CAT)in root tip,while thymol treatment pronouncedly enhance the activity of these two anti-oxidative enzymes under salt stress.(5) Exogenous treatment with thymol induced significant increase in the content of ascorbic acid(As A) and glutathione(GSH) in root tip under salt stress.These results suggested that thymol was capable of conferring salt-induced oxidative injury and cell death by enhancing the anti-oxidative capacity in the root tip of rice seedlings.The current study provides important technological and scientific basis to ensure the safety of crop production in salinity soil.
Exogenous regulation of salt tolerance is an important approach for the safety of crop production in salinity soil.Thymol is a novel plant-derived natural compound,which has been extensively reported to have medicinal activity.However,the regulation of plant physiology by thymol is still elusive.In the present study,physiological,histochemical,and biochemical methods were used to investigate thymol-modulated salt tolerance in rice seedlings.Basically,the results showed that,(1) Exogenous treatment with thymol significantly attenuated salt-induced root inhibition in a dose-dependent manner.(2) Exogenous treatment with thymol remarkably blocked salt-induced cell death in root tip.(3) Exogenous treatment with thymol scavenged the over-generated hydrogen peroxide and superoxide radical in root tip under salt stress.(4)Salt stress significantly inhibited the activity of superoxide dismutase(SOD)and catalase(CAT)in root tip,while thymol treatment pronouncedly enhance the activity of these two anti-oxidative enzymes under salt stress.(5) Exogenous treatment with thymol induced significant increase in the content of ascorbic acid(As A) and glutathione(GSH) in root tip under salt stress.These results suggested that thymol was capable of conferring salt-induced oxidative injury and cell death by enhancing the anti-oxidative capacity in the root tip of rice seedlings.The current study provides important technological and scientific basis to ensure the safety of crop production in salinity soil.
引文
[1]VINOCUR B,ALTMAN A.Recent advances in engineering plant tolerance to abiotic stress:achievements and limitations[J].Current opinion in biotechnology,2005,16(2):123-132.
[2]孙伟,郑崇珂,解丽霞,等.水稻对盐胁迫的生理和分子反应研究进展[J].山东农业科学,2016,48(4):148-153.
[3]陈绍荣,邵建华,王喜江,等.我国土壤盐渍化的综合治理[J].化肥工业,2013,40(5):65-69.
[4]HOSSAIN M A,HASANUZZAMAN M,FUJITA M.Coordinate induction of antioxidant defense and glyoxalase system by exogenous proline and glycinebetaine is correlated with salt tolerance in mung bean[J].Frontiers of agriculture in China,2011,5(1):1-14.
[5]MAHAJAN S,TUTEJA N.Cold,salinity and drought stresses:an overview[J].Archives of biochemistry and biophysics,2005,444(2):139-158.
[6]BENSMIRA M,JIANG B,NSABIMANA C,et al.Effect of lavender and thyme incorporation in sunflower seed oil on its resistance to frying temperatures[J].Food research international,2007,40(3):341-346.
[7]AESCHBACH R,LOLIGER J,SCOTT B C,et al.Antioxidant actions of thymol,carvacrol,6-gingerol,zingerone and hydroxytyrosol[J].Food and chemical toxicology,1994,32(1):31-36.
[8]KRUK I,MICHALSKA T,LICHSZTELD K,et al.The effect of thymol and its derivatives on reactions generating reactive oxygen species[J].Chemosphere,2000,41(7):1 059-1 064.
[9]ALAM K,NAGI M N,BADARY O A,et al.The protective action of thymol against carbon tetrachloride hepatotoxicity in mice[J].Pharmacological research,1999,40(2):159-163.
[10]IPEK E,TUYLU B A,ZEYTINOGLU H.Effects of carvacrol on sister chromatid exchanges in human lymphocyte cultures[M]//Animal cell technology:basic and applied aspects.Netherlands:Springer,2003:471-474.
[11]BENARFA A,COMBENS S,PREZIOSI-BELLOY L,et al.Antimicrobial activity of carvacrol related to its chemical structure[J].Letters in applied microbiology,2006,43(2):149-154.
[12]KARKABOUNAS S,KOSTOULA O K,DASKALOU T,et al.Anticarcinogenic and antiplatelet effects of carvacrol[J].Exp oncol,2006,28(2):121-125.
[13]DANDLEN S A,LIMA A S,MENDES M D,et al.Antioxidant activity of six Portuguese thyme species essential oils[J].Flavour and fragrance journal,2010,25(3):150-155.
[14]SAAD A,FADLI M,BOUAZIZ M,et al.Anticandidal activity of the essential oils of Thymus maroccanus and Thymus broussonetii and their synergism with amphotericin B and fluconazol[J].Phytomedicine,2010,17(13):1 057-1 060.
[15]NAVARRO D,DIAZ-MULA H M,GUILLEN F,et al.Reduction of nectarine decay caused by Rhizopus stolonifer,Botrytis cinerea and Penicillium digitatum with Aloe vera gel alone or with the addition of thymol[J].International journal of food microbiology,2011,151(2):241-246.
[16]CASTILLO S,PEREZ A C O,Martínez R D,et al.The essential oils thymol and carvacrol applied in the packing lines avoid lemon spoilage and maintain quality during storage[J].Food control,2014,35(1):132-136.
[17]中国香料香精化妆品协会.欧盟批准百里香酚用作植保产品[J].国内外香化信息,2013(7):21.
[18]张猛,陈健,薛延丰,等.植物源化合物百里香酚对西瓜枯萎病菌的抑菌机制初探[J].江苏农业学报,2013,29(5):1 019-1 024.
[19]GAO T,ZHOU H,ZHOU W,et al.The fungicidal activity of thymol against Fusarium graminearum via inducing lipid peroxidation and disrupting ergosterol biosynthesis[J].Molecules,2016,21(6):770.
[20]DUAN Y,ZHANG W,Li B,et al.An endoplasmic reticulum response pathway mediates programmed cell death of root tip induced by water stress in Arabidopsis[J].New phytologist,2010,186(3):681-695.
[21]OROZCO C M,RYAN C A.Hydrogen peroxide is generated systematically in plant leaves by wounding and systemin via the octadecanoid pathway[J].Proceedings of the national academy of sciences,1999,96(11):6 553-6 557.
[22]YAMAMOTO Y,KOBAYASHI Y,MATSUMOTO H.Lipid peroxidation is an early symptom triggered by aluminum,but not the primary cause of elongation inhibition in pea roots[J].Plant physiology,2001,125(1):199-208.
[23]LISO R,DE TULLIO M C,CIRACI S,et al.Localization of ascorbic acid,ascorbic acid oxidase,and glutathione in roots of Cucurbita maxima L[J].Journal of experimental botany,2004,55(408):2 589-2 597.
[24]XUE Y F,LIU L,LIU Z P,et al.Protective role of Ca against Na Cl toxicity in Jerusalem artichoke by up-regulation of antioxidant enzymes[J].Pedosphere,2008,18(6):766-774.
[25]CHEN J,SHIYAB S,HAN F X,et al.Bioaccumulation and physiological effects of mercury in Pterisvittata and Nephrolepisexaltata[J].Ecotoxicology,2009,18(1):110-121.
[26]BRADFORD M M.A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of protein-dye binding[J].Analytical biochemistry,1976,72(1/2):248-254.
[27]GILL S S,TUTEJA N.Reactive oxygen species and antioxidant machinery in abiotic stress tolerance in crop plants[J].Plant physiology and biochemistry,2010,48(12):909-930.
[28]ALSCHER R G,ERTURK N,HEATH L S.Role of superoxide dismutases(SODs)in controlling oxidative stress in plants[J].Journal of experimental botany,2002,53(372):1 331-1 341.
[29]VAN B F,DAT J F.Reactive oxygen species in plant cell death[J].Plant physiology,2006,141(2):384-390.
[30]DUPUY L,GREGORY P J,BENGOUGH A G.Root growth models:towards a new generation of continuous approaches[J].Journal of experimental botany,2000,61:2 131-2 143.
[31]KRUK I,MICHALSSKA T,LICHSZTEDLD K,et al.The effect of thymol and its derivatives on reactions generating reactive oxygen species[J].Chemosphere,2000,41(7):1 059-1 064.
[32]AL-MALKI A L.Antioxidant properties of thymol and butylated hydroxytoluene in carbon tetrachloride-induced mice liver injury[J].Journal of king abdulaziz university-science,2010,22:239-248.
[33]ABD-ELHAKIM Y M,MOHAMED W A M.Assessment of the role of thymol in combating chromium(Ⅵ)-induced oxidative stress in isolated rat erythrocytes in vitro[J].Toxicological and environmental chemistry,2016,98(10):1-14.
[34]SHETTIGAR N B,DAS S,RAO N B,et al.Thymol,a monoterpene phenolic derivative of cymene,abrogates mercury-induced oxidative stress resultant cytotoxicity and genotoxicity in hepatocarcinoma cells[J].Environmental toxicology,2015,30(8):968-980.
[35]KIM Y S,HWANG J W,KANG S H,et al.Thymol from Thymus quinquecostatus Celak.protects against tert-butyl hydroperoxideinduced oxidative stress in Chang cells[J].Journal of natural medicines,2014,68(1):154-162.
[36]MITTLER R.Oxidative stress,antioxidants and stress tolerance[J].Trends in plant science,2002,7(9):405-410.
[2]孙伟,郑崇珂,解丽霞,等.水稻对盐胁迫的生理和分子反应研究进展[J].山东农业科学,2016,48(4):148-153.
[3]陈绍荣,邵建华,王喜江,等.我国土壤盐渍化的综合治理[J].化肥工业,2013,40(5):65-69.
[4]HOSSAIN M A,HASANUZZAMAN M,FUJITA M.Coordinate induction of antioxidant defense and glyoxalase system by exogenous proline and glycinebetaine is correlated with salt tolerance in mung bean[J].Frontiers of agriculture in China,2011,5(1):1-14.
[5]MAHAJAN S,TUTEJA N.Cold,salinity and drought stresses:an overview[J].Archives of biochemistry and biophysics,2005,444(2):139-158.
[6]BENSMIRA M,JIANG B,NSABIMANA C,et al.Effect of lavender and thyme incorporation in sunflower seed oil on its resistance to frying temperatures[J].Food research international,2007,40(3):341-346.
[7]AESCHBACH R,LOLIGER J,SCOTT B C,et al.Antioxidant actions of thymol,carvacrol,6-gingerol,zingerone and hydroxytyrosol[J].Food and chemical toxicology,1994,32(1):31-36.
[8]KRUK I,MICHALSKA T,LICHSZTELD K,et al.The effect of thymol and its derivatives on reactions generating reactive oxygen species[J].Chemosphere,2000,41(7):1 059-1 064.
[9]ALAM K,NAGI M N,BADARY O A,et al.The protective action of thymol against carbon tetrachloride hepatotoxicity in mice[J].Pharmacological research,1999,40(2):159-163.
[10]IPEK E,TUYLU B A,ZEYTINOGLU H.Effects of carvacrol on sister chromatid exchanges in human lymphocyte cultures[M]//Animal cell technology:basic and applied aspects.Netherlands:Springer,2003:471-474.
[11]BENARFA A,COMBENS S,PREZIOSI-BELLOY L,et al.Antimicrobial activity of carvacrol related to its chemical structure[J].Letters in applied microbiology,2006,43(2):149-154.
[12]KARKABOUNAS S,KOSTOULA O K,DASKALOU T,et al.Anticarcinogenic and antiplatelet effects of carvacrol[J].Exp oncol,2006,28(2):121-125.
[13]DANDLEN S A,LIMA A S,MENDES M D,et al.Antioxidant activity of six Portuguese thyme species essential oils[J].Flavour and fragrance journal,2010,25(3):150-155.
[14]SAAD A,FADLI M,BOUAZIZ M,et al.Anticandidal activity of the essential oils of Thymus maroccanus and Thymus broussonetii and their synergism with amphotericin B and fluconazol[J].Phytomedicine,2010,17(13):1 057-1 060.
[15]NAVARRO D,DIAZ-MULA H M,GUILLEN F,et al.Reduction of nectarine decay caused by Rhizopus stolonifer,Botrytis cinerea and Penicillium digitatum with Aloe vera gel alone or with the addition of thymol[J].International journal of food microbiology,2011,151(2):241-246.
[16]CASTILLO S,PEREZ A C O,Martínez R D,et al.The essential oils thymol and carvacrol applied in the packing lines avoid lemon spoilage and maintain quality during storage[J].Food control,2014,35(1):132-136.
[17]中国香料香精化妆品协会.欧盟批准百里香酚用作植保产品[J].国内外香化信息,2013(7):21.
[18]张猛,陈健,薛延丰,等.植物源化合物百里香酚对西瓜枯萎病菌的抑菌机制初探[J].江苏农业学报,2013,29(5):1 019-1 024.
[19]GAO T,ZHOU H,ZHOU W,et al.The fungicidal activity of thymol against Fusarium graminearum via inducing lipid peroxidation and disrupting ergosterol biosynthesis[J].Molecules,2016,21(6):770.
[20]DUAN Y,ZHANG W,Li B,et al.An endoplasmic reticulum response pathway mediates programmed cell death of root tip induced by water stress in Arabidopsis[J].New phytologist,2010,186(3):681-695.
[21]OROZCO C M,RYAN C A.Hydrogen peroxide is generated systematically in plant leaves by wounding and systemin via the octadecanoid pathway[J].Proceedings of the national academy of sciences,1999,96(11):6 553-6 557.
[22]YAMAMOTO Y,KOBAYASHI Y,MATSUMOTO H.Lipid peroxidation is an early symptom triggered by aluminum,but not the primary cause of elongation inhibition in pea roots[J].Plant physiology,2001,125(1):199-208.
[23]LISO R,DE TULLIO M C,CIRACI S,et al.Localization of ascorbic acid,ascorbic acid oxidase,and glutathione in roots of Cucurbita maxima L[J].Journal of experimental botany,2004,55(408):2 589-2 597.
[24]XUE Y F,LIU L,LIU Z P,et al.Protective role of Ca against Na Cl toxicity in Jerusalem artichoke by up-regulation of antioxidant enzymes[J].Pedosphere,2008,18(6):766-774.
[25]CHEN J,SHIYAB S,HAN F X,et al.Bioaccumulation and physiological effects of mercury in Pterisvittata and Nephrolepisexaltata[J].Ecotoxicology,2009,18(1):110-121.
[26]BRADFORD M M.A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of protein-dye binding[J].Analytical biochemistry,1976,72(1/2):248-254.
[27]GILL S S,TUTEJA N.Reactive oxygen species and antioxidant machinery in abiotic stress tolerance in crop plants[J].Plant physiology and biochemistry,2010,48(12):909-930.
[28]ALSCHER R G,ERTURK N,HEATH L S.Role of superoxide dismutases(SODs)in controlling oxidative stress in plants[J].Journal of experimental botany,2002,53(372):1 331-1 341.
[29]VAN B F,DAT J F.Reactive oxygen species in plant cell death[J].Plant physiology,2006,141(2):384-390.
[30]DUPUY L,GREGORY P J,BENGOUGH A G.Root growth models:towards a new generation of continuous approaches[J].Journal of experimental botany,2000,61:2 131-2 143.
[31]KRUK I,MICHALSSKA T,LICHSZTEDLD K,et al.The effect of thymol and its derivatives on reactions generating reactive oxygen species[J].Chemosphere,2000,41(7):1 059-1 064.
[32]AL-MALKI A L.Antioxidant properties of thymol and butylated hydroxytoluene in carbon tetrachloride-induced mice liver injury[J].Journal of king abdulaziz university-science,2010,22:239-248.
[33]ABD-ELHAKIM Y M,MOHAMED W A M.Assessment of the role of thymol in combating chromium(Ⅵ)-induced oxidative stress in isolated rat erythrocytes in vitro[J].Toxicological and environmental chemistry,2016,98(10):1-14.
[34]SHETTIGAR N B,DAS S,RAO N B,et al.Thymol,a monoterpene phenolic derivative of cymene,abrogates mercury-induced oxidative stress resultant cytotoxicity and genotoxicity in hepatocarcinoma cells[J].Environmental toxicology,2015,30(8):968-980.
[35]KIM Y S,HWANG J W,KANG S H,et al.Thymol from Thymus quinquecostatus Celak.protects against tert-butyl hydroperoxideinduced oxidative stress in Chang cells[J].Journal of natural medicines,2014,68(1):154-162.
[36]MITTLER R.Oxidative stress,antioxidants and stress tolerance[J].Trends in plant science,2002,7(9):405-410.