铜胁迫对甘蔗生长及抗氧化酶活性的影响
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摘要
甘蔗是重要的糖能兼用植物,同时也是潜在的重金属土壤修复植物。为了探明甘蔗对铜胁迫的生理响应,该研究采用营养液培养的方法分析了铜胁迫对甘蔗生长、叶片相对电导率、叶绿素含量及根系抗氧化酶活性等的影响。结果表明:甘蔗地上部分生物、叶绿素含量随铜浓度增加而下降,地上部分铜含量和叶片的相对电导率则随铜浓度增加而提高。根系的丙二醛(MDA)含量在胁迫24 h开始发生显著的变化,在400μmol·L~(-1)铜处理下比对照显著增加了25.5%,随着胁迫时间的延长,丙二醛含量增加幅度上升。到胁迫72 h,100μmol·L~(-1)铜处理的MDA含量也显著的提高。根系SOD、POD和CAT酶活性在胁迫24 h开始发生显著的变化,但不同酶活性变化存在差异。SOD酶活性除了在胁迫的72 h随铜浓度增加而提高,其他时间表现为在100μmol·L~(-1)铜处理下活性下降,随着铜浓度继续增加SOD酶活性提高,到400μmol·L~(-1)铜处理SOD酶活性又出现下降。POD酶活性在处理的24、48和72 h表现为随浓度增加先提高后下降的趋势,但在处理7 d时随铜浓度增加而提高。CAT酶活性随着浓度的升高和胁迫时间的延长而下降,到了胁迫的第7天,三个铜处理下CAT酶活性分别下降了89.98%、96.88%和98.50%。由此可见,铜胁迫严重影响甘蔗的生长,较短时间的处理就已经引起了甘蔗根系的氧化胁迫,根系中SOD、POD和CAT酶活性的变化在甘蔗对抗铜引起的氧化胁迫中起到重要作用。
Sugarcane is the most important sugar and energy crop. It is also the potential plant for remediation of high metal-polluted soil. To evaluate the physiological response of sugarcane to copper( Cu) stress,the weight of sugarcane,content of chlorophyll,relative electrical conductivity in leaves and activities of antioxidant enzymes in roots were examined by hydroponics. The results revealed that the growth of sugarcane inhibited by Cu stress. The weight of shoot and content of chlorophyll decreased with the increase of Cu content in nutrient solution,while the copper content in shoot and relative electrical conductivity in leaves increased in response to copper stress. The malonaldehyde( MDA) content was increased significantly by 25.5% under 400 μmol·L~(-1) treatment at 24 h when compared with the control,and then the rate increase raised with the prolonging of treatment time. The MDA content increased notably after 72 h under 100μmol·L~(-1) Cu treatment. The activities of SOD,POD and CAT changed significantly after 24 h treatment,but notable differences were found among different enzymes. The activities of SOD firstly dropped under 100 μmol·L~(-1) treatment,and then increased with the increase of Cu content in solution under 200 μmol·L~(-1) treatment,following with decrease of enzyme activities under 400 μmol·L~(-1) treatment at 24 h,48 h and 7 d. However,the activities of SOD increased with the increase of Cu content in solution at 72 h. The activities of POD increased with the increase of Cu content in solution,following with decline of enzyme activities under high content of Cu treatment at 24,48 and 7 h except at 72 d. The activities of CAT decreased significantly with the increase of Cu content in solution and with the prolonging of stress time. The activities of CAT declined by 89.98%,96.88% and 98.50% under three Cu treatments,respectively on the seventh day. Thus,the growth of sugarcane was greatly inhibited by copper stress and the change of SOD,POD and CAT activities might play an important role in alleviating oxidative stress caused by Cu stress.
Sugarcane is the most important sugar and energy crop. It is also the potential plant for remediation of high metal-polluted soil. To evaluate the physiological response of sugarcane to copper( Cu) stress,the weight of sugarcane,content of chlorophyll,relative electrical conductivity in leaves and activities of antioxidant enzymes in roots were examined by hydroponics. The results revealed that the growth of sugarcane inhibited by Cu stress. The weight of shoot and content of chlorophyll decreased with the increase of Cu content in nutrient solution,while the copper content in shoot and relative electrical conductivity in leaves increased in response to copper stress. The malonaldehyde( MDA) content was increased significantly by 25.5% under 400 μmol·L~(-1) treatment at 24 h when compared with the control,and then the rate increase raised with the prolonging of treatment time. The MDA content increased notably after 72 h under 100μmol·L~(-1) Cu treatment. The activities of SOD,POD and CAT changed significantly after 24 h treatment,but notable differences were found among different enzymes. The activities of SOD firstly dropped under 100 μmol·L~(-1) treatment,and then increased with the increase of Cu content in solution under 200 μmol·L~(-1) treatment,following with decrease of enzyme activities under 400 μmol·L~(-1) treatment at 24 h,48 h and 7 d. However,the activities of SOD increased with the increase of Cu content in solution at 72 h. The activities of POD increased with the increase of Cu content in solution,following with decline of enzyme activities under high content of Cu treatment at 24,48 and 7 h except at 72 d. The activities of CAT decreased significantly with the increase of Cu content in solution and with the prolonging of stress time. The activities of CAT declined by 89.98%,96.88% and 98.50% under three Cu treatments,respectively on the seventh day. Thus,the growth of sugarcane was greatly inhibited by copper stress and the change of SOD,POD and CAT activities might play an important role in alleviating oxidative stress caused by Cu stress.
引文
BHADURI AM,FULEKAR MH,2012.Antioxidant enzyme responses of plants to heavy metal stress[J].Rev Environ Sci Biotechnol,11(1):55-69.
CAO YN,MA CX,CHEN GC,et al.,2017.Physiological and biochemical responses of Salix integra Thunb.under copper stress as affected by soil flooding[J].Eniron Poll,225:644-653.
CHEN JX,WAGN XF,2006.Plant physiology experiment instruction[M].2nd ed.Guangzhou:South China University of Technology Press:81-84.[陈建勋,王晓峰,2006.植物生理学实验指导[M].2版.广州:华南理工大学出版社:81-84.]
DING JH,XUE ZL,YANG CY,2013.Effect of salicylic acid on membrane lipid peroxidation in rice seedlings under copper stress[J].Heilongjiang Agric Sci,(1):14-18.[丁佳红,薛正莲,杨超英,2013.水杨酸对铜胁迫下水稻幼苗膜脂过氧化作用的影响[J].黑龙江农业科学,(1):14-18.]
FAMOSO AN,CLARK RT,SHAFF JE,et al.,2010.Development of a novel aluminum tolerance phenotyping platform used for comparisons of cereal aluminum tolerance and investigations into rice aluminum tolerance mechanisms[J].Plant Physiol,153(4):1678-1691.
GE SF,ZHANG Y,WU YH,et al.,2014.Effect of copper pollution on the growth and physiological characters of tobacco(Nicotiana tabacum L.)[J].J Zhejiang Norm Univ(Nat Sci Ed),37(2):219-224.[葛淑芳,章艺,吴玉环,等,2014.铜污染对烟草生长及其生理特性的影响[J].浙江师范大学学报(自然科学版),37(2):219-224.]
GONG Q,KANG Q,WANG L,et al.,2018.Toxicity mechanism of heavy metal copper to plants:A review[J].JSouthern Agric,49(3):469-475.[公勤,康群,王玲,等,2018.重金属铜对植物毒害机理的研究现状及展望[J].南方农业学报,49(3):469-475.]
GUO JW,CUI XX,ZHANG YB,et al.,2010.Cu uptake by sugarcane and its effects on sugarcane yield and quality[J].Soils,42(4):606-610.[郭家文,崔雄维,张跃彬,等,2010.重金属铜在甘蔗体内的吸收及对甘蔗产量和品质的影响[J].土壤,42(4):606-610.]
HU ZB,CHEN YH,WANG GP,et al.,2006.Effects of copper stress on growth,chlorophyll fluorescence parameters and antioxidant enzyme activities of Zea mays seedlings[J].Chin Bull Bot,23(2):129-137.[胡筑兵,陈亚华,王桂萍,等,2006.铜胁迫对玉米幼苗生长、叶绿素荧光参数和抗氧化酶活性的影响[J].植物学通报,23(2):129-137.]
JIN Y,FU QL,ZHENG J,et al.,2012.Research status on phytoremediation of copper contaminated soil with hyperaccumulator[J].J Agric Sci Technol,14(4):93-100.[金勇,付庆灵,郑进,等,2012.超积累植物修复铜污染土壤的研究现状[J].中国农业科技导报,14(4):93-100.]
LI X,RAO HH,YANG CL,et al.,2016.Study on physiological mechanism of exogenous abscisic acid alleviating copper stress on wheat seedling[J].J Tritic Crops,36(6):759-764.[李雪,饶红红,杨彩玲,等,2016.外源脱落酸缓解小麦幼苗铜胁迫伤害的生理机制初探[J].麦类作物学报,36(6):759-764.]
LI ZG,YUAN LX,WANG QL,et al.,2013.Combined action of antioxidant defense system and osmolytes in chilling shockinduced chilling tolerance in Jatropha curcas seedlings[J].Acta Physiol Plant,35(7):2127-2136.
LIU P,ZHANG YY,WU YH,et al.,2009.Effects of copper stress on nutrient uptake and physiological characteristics of tobacco roots[J].J Zhejiang Norm Univ(Nat Sci Ed),32(4):442-447.[刘鹏,张艳英,吴玉环,等,2009.铜胁迫对烟草养分吸收和根系生理的影响[J].浙江师范大学学报(自然科学版),32(4):442-447.]
LOU LQ,SHEN ZG,LI XD,2004.The copper tolerance mechanisms of Elsholtzia haichowensis,a plant from copperenriched soils[J].Environ Exp Bot,51(2):111-120.
MOSTOFA MG,HOSSAIN MA,FUJITA M,et al.,2015.Physiological and biochemical mechanisms associated with trehalose-induced copper-stress tolerance in rice[J].Sci Rep,5:11433.
PENG XY,SONG M,2011.Effects of exogenous cysteine on growth,copper accumulation and antioxidative systems in whteat seedlings under Cu stress[J].Acta Ecol Sin,31(12):3504-3511.[彭向永,宋敏,2011.外源半胱氨酸对铜胁迫下小麦幼苗生长、铜积累量及抗氧化系统的影响[J].生态学报,31(12):3504-3511.]
RAVET K,PILON M,2013.Copper and iron homeostasis in plants:The challenges of oxidative stress[J].Antioxid Redox Sign,19(9):919-932.
SERENO ML,ALMEIDA RS,NISHIMURA DS,et al.,2007.Response of sugarcane to increasing concentrations of copper and cadmium and expression of metallothionein genes[J].J Plant Physiol,164(11):1499-1515.
TANG ZC,1999.Modern plant physiology experiment[M].Beijing:Science Press:302-305.[汤章城,1999.现代植物生理学实验指南[M].北京:科学出版社:302-305.]
WANG SZ,JIN ZX,LI YL,et al.,2015.Effects of arbuscular mycorrhizal fungi inoculation on the photosynthetic pigment contents,anti-oxidation capacity and membrane lipid peroxidation of Elsholtzia splendens leaves under copper stress[J].Acta Ecol Sin,35(23):7699-7708.[王穗子,金则新,李月灵,等,2015.铜胁迫条件下AMF对海州香薷光合色素含量、抗氧化能力和膜脂过氧化的影响[J].生态学报,35(23):7699-7708.]
WANG XW,HUANG GQ,XU JC,et al.,2014.Effects of copper stresses and intercropping on antioxidant enzyme activities and malondialdehyde contents in maize[J].J AgroEnviron Sci,33(10):1890-1896.[王晓维,黄国勤,徐健程,等,2014.铜胁迫和间作对玉米抗氧化酶活性及丙二醛含量的影响[J].农业环境科学学报,33(10):1890-1896.]
WANG XW,XU JC,SUN DP,et al.,2016.Effects of biochar on chlorophyll and protective enzyme activity of rape seedlings in red soil under copper stress[J].J Agro-Environ Sci,35(4):640-646.[王晓维,徐健程,孙丹平,等,2016.生物炭对铜胁迫下红壤地油菜苗期叶绿素和保护性酶活性的影响[J].农业环境科学学报,35(4):640-646.]
XUE YW,WANG YF,ZHAO CJ,et al.,2016.Effect of copper stress on germination and antioxidant system in wheat seedlings[J].Acta Agric Univ Jiangxi,38(1):54-59.[薛盈文,王玉凤,赵长江,等,2016.铜胁迫对小麦种子萌发及幼苗抗氧化系统的影响[J].江西农业大学学报,38(1):54-59.]
ZHAO QF,YIN JM,GAO XX,et al.,2017.Effects of copper or zinc stress on antioxidant system in wheat roots[J].J NWNorm Univ(Nat Sci Ed),53(4):92-97.[赵庆芳,殷金梅,高晓霞,等,2017.铜或锌胁迫对小麦根中抗氧化系统的影响[J].西北师范大学学报(自然科学版),53(4):92-97.]
ZHAO XQ,ZHANG HY,LIU WZ,2012.Research advances in transport-related proteins of copper in plant[J].Guihaia,32(2):280-284.[赵雪芹,张海燕,刘维仲,2012.植物铜转运相关蛋白研究进展[J].广西植物,32(2):280-284.]
CAO YN,MA CX,CHEN GC,et al.,2017.Physiological and biochemical responses of Salix integra Thunb.under copper stress as affected by soil flooding[J].Eniron Poll,225:644-653.
CHEN JX,WAGN XF,2006.Plant physiology experiment instruction[M].2nd ed.Guangzhou:South China University of Technology Press:81-84.[陈建勋,王晓峰,2006.植物生理学实验指导[M].2版.广州:华南理工大学出版社:81-84.]
DING JH,XUE ZL,YANG CY,2013.Effect of salicylic acid on membrane lipid peroxidation in rice seedlings under copper stress[J].Heilongjiang Agric Sci,(1):14-18.[丁佳红,薛正莲,杨超英,2013.水杨酸对铜胁迫下水稻幼苗膜脂过氧化作用的影响[J].黑龙江农业科学,(1):14-18.]
FAMOSO AN,CLARK RT,SHAFF JE,et al.,2010.Development of a novel aluminum tolerance phenotyping platform used for comparisons of cereal aluminum tolerance and investigations into rice aluminum tolerance mechanisms[J].Plant Physiol,153(4):1678-1691.
GE SF,ZHANG Y,WU YH,et al.,2014.Effect of copper pollution on the growth and physiological characters of tobacco(Nicotiana tabacum L.)[J].J Zhejiang Norm Univ(Nat Sci Ed),37(2):219-224.[葛淑芳,章艺,吴玉环,等,2014.铜污染对烟草生长及其生理特性的影响[J].浙江师范大学学报(自然科学版),37(2):219-224.]
GONG Q,KANG Q,WANG L,et al.,2018.Toxicity mechanism of heavy metal copper to plants:A review[J].JSouthern Agric,49(3):469-475.[公勤,康群,王玲,等,2018.重金属铜对植物毒害机理的研究现状及展望[J].南方农业学报,49(3):469-475.]
GUO JW,CUI XX,ZHANG YB,et al.,2010.Cu uptake by sugarcane and its effects on sugarcane yield and quality[J].Soils,42(4):606-610.[郭家文,崔雄维,张跃彬,等,2010.重金属铜在甘蔗体内的吸收及对甘蔗产量和品质的影响[J].土壤,42(4):606-610.]
HU ZB,CHEN YH,WANG GP,et al.,2006.Effects of copper stress on growth,chlorophyll fluorescence parameters and antioxidant enzyme activities of Zea mays seedlings[J].Chin Bull Bot,23(2):129-137.[胡筑兵,陈亚华,王桂萍,等,2006.铜胁迫对玉米幼苗生长、叶绿素荧光参数和抗氧化酶活性的影响[J].植物学通报,23(2):129-137.]
JIN Y,FU QL,ZHENG J,et al.,2012.Research status on phytoremediation of copper contaminated soil with hyperaccumulator[J].J Agric Sci Technol,14(4):93-100.[金勇,付庆灵,郑进,等,2012.超积累植物修复铜污染土壤的研究现状[J].中国农业科技导报,14(4):93-100.]
LI X,RAO HH,YANG CL,et al.,2016.Study on physiological mechanism of exogenous abscisic acid alleviating copper stress on wheat seedling[J].J Tritic Crops,36(6):759-764.[李雪,饶红红,杨彩玲,等,2016.外源脱落酸缓解小麦幼苗铜胁迫伤害的生理机制初探[J].麦类作物学报,36(6):759-764.]
LI ZG,YUAN LX,WANG QL,et al.,2013.Combined action of antioxidant defense system and osmolytes in chilling shockinduced chilling tolerance in Jatropha curcas seedlings[J].Acta Physiol Plant,35(7):2127-2136.
LIU P,ZHANG YY,WU YH,et al.,2009.Effects of copper stress on nutrient uptake and physiological characteristics of tobacco roots[J].J Zhejiang Norm Univ(Nat Sci Ed),32(4):442-447.[刘鹏,张艳英,吴玉环,等,2009.铜胁迫对烟草养分吸收和根系生理的影响[J].浙江师范大学学报(自然科学版),32(4):442-447.]
LOU LQ,SHEN ZG,LI XD,2004.The copper tolerance mechanisms of Elsholtzia haichowensis,a plant from copperenriched soils[J].Environ Exp Bot,51(2):111-120.
MOSTOFA MG,HOSSAIN MA,FUJITA M,et al.,2015.Physiological and biochemical mechanisms associated with trehalose-induced copper-stress tolerance in rice[J].Sci Rep,5:11433.
PENG XY,SONG M,2011.Effects of exogenous cysteine on growth,copper accumulation and antioxidative systems in whteat seedlings under Cu stress[J].Acta Ecol Sin,31(12):3504-3511.[彭向永,宋敏,2011.外源半胱氨酸对铜胁迫下小麦幼苗生长、铜积累量及抗氧化系统的影响[J].生态学报,31(12):3504-3511.]
RAVET K,PILON M,2013.Copper and iron homeostasis in plants:The challenges of oxidative stress[J].Antioxid Redox Sign,19(9):919-932.
SERENO ML,ALMEIDA RS,NISHIMURA DS,et al.,2007.Response of sugarcane to increasing concentrations of copper and cadmium and expression of metallothionein genes[J].J Plant Physiol,164(11):1499-1515.
TANG ZC,1999.Modern plant physiology experiment[M].Beijing:Science Press:302-305.[汤章城,1999.现代植物生理学实验指南[M].北京:科学出版社:302-305.]
WANG SZ,JIN ZX,LI YL,et al.,2015.Effects of arbuscular mycorrhizal fungi inoculation on the photosynthetic pigment contents,anti-oxidation capacity and membrane lipid peroxidation of Elsholtzia splendens leaves under copper stress[J].Acta Ecol Sin,35(23):7699-7708.[王穗子,金则新,李月灵,等,2015.铜胁迫条件下AMF对海州香薷光合色素含量、抗氧化能力和膜脂过氧化的影响[J].生态学报,35(23):7699-7708.]
WANG XW,HUANG GQ,XU JC,et al.,2014.Effects of copper stresses and intercropping on antioxidant enzyme activities and malondialdehyde contents in maize[J].J AgroEnviron Sci,33(10):1890-1896.[王晓维,黄国勤,徐健程,等,2014.铜胁迫和间作对玉米抗氧化酶活性及丙二醛含量的影响[J].农业环境科学学报,33(10):1890-1896.]
WANG XW,XU JC,SUN DP,et al.,2016.Effects of biochar on chlorophyll and protective enzyme activity of rape seedlings in red soil under copper stress[J].J Agro-Environ Sci,35(4):640-646.[王晓维,徐健程,孙丹平,等,2016.生物炭对铜胁迫下红壤地油菜苗期叶绿素和保护性酶活性的影响[J].农业环境科学学报,35(4):640-646.]
XUE YW,WANG YF,ZHAO CJ,et al.,2016.Effect of copper stress on germination and antioxidant system in wheat seedlings[J].Acta Agric Univ Jiangxi,38(1):54-59.[薛盈文,王玉凤,赵长江,等,2016.铜胁迫对小麦种子萌发及幼苗抗氧化系统的影响[J].江西农业大学学报,38(1):54-59.]
ZHAO QF,YIN JM,GAO XX,et al.,2017.Effects of copper or zinc stress on antioxidant system in wheat roots[J].J NWNorm Univ(Nat Sci Ed),53(4):92-97.[赵庆芳,殷金梅,高晓霞,等,2017.铜或锌胁迫对小麦根中抗氧化系统的影响[J].西北师范大学学报(自然科学版),53(4):92-97.]
ZHAO XQ,ZHANG HY,LIU WZ,2012.Research advances in transport-related proteins of copper in plant[J].Guihaia,32(2):280-284.[赵雪芹,张海燕,刘维仲,2012.植物铜转运相关蛋白研究进展[J].广西植物,32(2):280-284.]