高浓度臭氧对美国薄荷(Monarda didyma L.)叶片光合及抗性生理特征的影响
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摘要
以一种常见药食两用植物——美国薄荷(Monarda didyma L.)为实验材料,利用开顶箱(OTC)模拟法,研究了不同臭氧浓度(40 (CK)、80、120和160 nmol·mol-1)下薄荷叶片生理生态指标的变化规律。结果表明:(1) 120、160 nmol·mol-1浓度的O3经过14 d的熏蒸对薄荷造成了明显的伤害,叶片表面出现了大量黄褐色斑块,80 nmol·mol-1处理下则无明显的可见伤害症状;(2)处理14 d时,高浓度O3熏蒸(160 nmol·mol-1)降低了薄荷叶片的净光合速率(Pn)、气孔导度(Gs)、PSⅡ电子传递量子产率(ΦPSⅡ)和PSⅡ最大光化学效率(Fv/Fm),与对照相比分别下降了30.2%、53.1%、25.7%和3.6%;(3)处理14 d时,与对照相比,高浓度O3熏蒸(160 nmol·mol-1)使得薄荷叶片的丙二醛(MDA)含量显著增加(74.6%)、超氧阴离子(O2·-)产生速率显著上升(31.0%)(P <0.05);而超氧化物歧化酶(SOD)活性(19.5%)及可溶性蛋白含量(20.9%)显著下降(P<0.05),表明高浓度O3导致了严重的膜脂过氧化和氧化胁迫,并且破坏了植株抗氧化系统的功能;脱落酸(ABA)含量显著增加(31.0%),表明O3熏蒸显著促进了植物叶片的衰老和脱落(黄叶率增加)。因此,O3污染可能会对薄荷等芳香类植物的生长及其食用和药用价值带来不利的影响。
An experiment was conducted to examine the variations of the physiological characters in leaves of Monarda didyma L. exposed to different ozone concentrations( 40( CK),80,120 and 160 nmol· mol-1) in open top chambers( OTCs). The results were as follows:( 1) An obviously visible leaf injury was observed under 120 and 160 nmol·mol-1 O3 with fumigation of 14 days,and a large number of yellow-brown patches appeared on the leaves,while no visible injury was observed under 80 nmol·mol-1 O3.( 2) After 14 days fumigation,elevated O3( 160 nmol·mol-1) decreased net photosynthetic rate( Pn),stomatal conductance( Gs),PS Ⅱ electron transfer quantum yield( ΦPS Ⅱ) and PS Ⅱ maximum photochemical efficient( Fv/Fm) by30.2%,53.1%,25.7% and 3.6%,respectively.( 3) Elevated O3( 160 nmol·mol-1) increased malondialdehyde( MDA) content( 74.6%) and the production rate of superoxide radical( O2·-)( 31.0%) after fumigation of 14 days,while significantly deceased superoxide dismutase( SOD)activity( 19.5%) and soluble protein content( 20.9%),indicating that elevated O3 led to serious lipid peroxidation and oxidative stress and destroyed the function of antioxidant system in plants.The content of abscisic acid( ABA) significantly increased( 31.0%),suggesting the acceleration of the aging and abscission of plant leaves( the proportion of yellow leaves increased). Consequently,O3 pollution may have adverse effects on the growth of Monarda didyma and its edible and medicinal values.
An experiment was conducted to examine the variations of the physiological characters in leaves of Monarda didyma L. exposed to different ozone concentrations( 40( CK),80,120 and 160 nmol· mol-1) in open top chambers( OTCs). The results were as follows:( 1) An obviously visible leaf injury was observed under 120 and 160 nmol·mol-1 O3 with fumigation of 14 days,and a large number of yellow-brown patches appeared on the leaves,while no visible injury was observed under 80 nmol·mol-1 O3.( 2) After 14 days fumigation,elevated O3( 160 nmol·mol-1) decreased net photosynthetic rate( Pn),stomatal conductance( Gs),PS Ⅱ electron transfer quantum yield( ΦPS Ⅱ) and PS Ⅱ maximum photochemical efficient( Fv/Fm) by30.2%,53.1%,25.7% and 3.6%,respectively.( 3) Elevated O3( 160 nmol·mol-1) increased malondialdehyde( MDA) content( 74.6%) and the production rate of superoxide radical( O2·-)( 31.0%) after fumigation of 14 days,while significantly deceased superoxide dismutase( SOD)activity( 19.5%) and soluble protein content( 20.9%),indicating that elevated O3 led to serious lipid peroxidation and oxidative stress and destroyed the function of antioxidant system in plants.The content of abscisic acid( ABA) significantly increased( 31.0%),suggesting the acceleration of the aging and abscission of plant leaves( the proportion of yellow leaves increased). Consequently,O3 pollution may have adverse effects on the growth of Monarda didyma and its edible and medicinal values.
引文
付伟,高江艳,徐胜,等.2014.高浓度臭氧对城市白桦和银中杨光合作用的影响.生态学杂志,33(12):3184-3190.
高峰,李品,冯兆忠.2017.臭氧与干旱对植物复合影响的研究进展.植物生态学报,41(2):252-268.
高俊凤.2006.植物生理学实验指导.北京:高等教育出版社.
郭雄飞,黎华寿,杨宝仪,等.2015.秋枫和木棉对大气臭氧浓度升高的生理响应.中南林业科技大学学报,35(2):49-53.
黄玉源,黄益宗,李秋霞,等.2006.臭氧对南方3种木本植物的急性伤害症状及其生理指标变化.生态环境,15(4):674-681.
李莉,田士林.2007.酶联免疫(ELISA)分析晋麦叶片中脱落酸的含量.安徽农业科学,35(23):7098-7099.
李梅,徐胜,张恒庆,等.2014.高浓度O3对水蜡叶表皮气孔及其叶组织结构特征的影响.生态学杂志,33(1):53-58.
李苗苗.2014.几种典型亚热带森林植物对臭氧胁迫的光合生理响应(硕士学位论文).北京:中国林业科学研究院.
列淦文,叶龙华,薛立.2014.臭氧胁迫对植物主要生理功能的影响.生态学报,34(2):294-306.
欧英娟.2014.O3/CO2浓度升高对两种城市绿化植物光合特性及生长发育的影响(硕士学位论文).长沙:湖南农业大学.
阮亚男,徐胜,郭龙,等.2017.大气臭氧浓度升高对银杏叶片活性氧代谢及相关基因表达的影响.应用生态学报,28(11):3479-3486.
王闯,王帅,杨碧波,等.2015.气象条件对沈阳市环境空气臭氧浓度影响研究.中国环境监测,31(3):32-37.
吴丹.2016.地面臭氧污染对树木叶片组织结构的影响(硕士学位论文).北京:北京林业大学.
徐胜,何兴元,陈玮,等.2009.高浓度臭氧对树木生理生态的影响.生态学报,29(1):368-377.
尹海龙,田长彦.2013.氮调控对盐环境下甜菜功能叶光系统Ⅱ荧光特性的影响.植物生态学报,37(2):122-131.
赵诣,徐胜,何兴元,等.2014.三种冷季型草坪草对高浓度O3的生理响应.生态学杂志,12(3):203-208.
郑有飞,石茗化,吴荣军,等.2013.遮阴和臭氧浓度增加对冬小麦叶片光合作用的影响.农业环境科学,32(10):1925-1933.
庄明浩,李迎春,陈双林.2011a.毛竹和四季竹对臭氧胁迫的耐受力差异.生态学杂志,30(10):2191-2196.
庄明浩,李迎春,郭子武,等.2011b.臭氧浓度升高对毛竹叶片光合色素和抗性生理的影响.生态科学,30(6):607-612.
Anav A,De Marco A,Proietti C.2016.Comparing concentration-based(AOT40)and stomatal uptake(PODY)metrics for ozone risk assessment to European forests.Global Change Biology,22:1608-1627.
Brbara BM,Edenise SA,Mauro AM,et al.2018.Ozone affects leaf physiology and causes injury to foliage of native tree species from the tropical Atlantic Forest of southern Brazil.Science of the Total Environment,610-611:912-925.
Bortolin RC,Caregnato FF,Armando MD,et al.2014.Effects of chronic elevated ozone concentration on the redox state and fruit yield of red pepper plant Capsicum baccatum.Ecotoxicology and Environmental Safety,100:114-121.
Brando SE,Bulbovas P,Lima MEL,et al.2017.Biochemical leaf traits as indicators of tolerance potential in tree species from the Brazilian Atlantic forest against oxidative environmental stressors.Science of the Total Environment,575:406-417.
Feng Z,Sun J,Wan W,et al.2014.Evidence of widespread ozone induced visible injury on plants in Beijing.China.Environmental Pollution,193:296-301.
Feng ZZ,Hu EZ,Wang XK,et al.2015.Ground-level O3pollution and its impacts on food crops in China:A review.Environmental Pollution,199:42-48.
Feng ZZ,Kobayashi K,Ainsworth E.2008.Impact of elevated ozone concentration on growth,physiology,and yield of wheat(Triticum aestivum L.).Global Change Biology,14:2696-2708.
Ghanbari F,Sayyari M.2018.Controlled drought stress affects the chilling-hardening capacity of tomato seedlings as indicated by changes in phenol metabolisms,antioxidant enzymes activity,osmolytes concentration and abscisic acid accumulation.Scientia Horticulturae,229:167-174.
Ghimire RP,KivimenpM,Kasurinen A,et al.2017.Herbivore-induced BVOC emissions of Scots pine under warming,elevated ozone and increased nitrogen availability in an open-field exposure.Agricultural and Forest Meteorology,242:21-32.
Gong XJ,Chen XS,Feng SQ,et al.2017.Effects of ozone stress on physiological and biochemical characteristics dwarfing rootstock M9T337.Agricultural Science&Technology,18:579-582+606.
Hodges DM,Delong JM,Forney CF,et al.1999.Improving the thiobarbituric acid-substances assay for estimating lipid peroxidation in plant tissues containing anthocyanin and other interfering compounds.Planta,207:604-611.
Huttunen S,Manninen S.2013.A review of ozone responses in Scots pine(Pinus sylvestris).Environmental and Experimental Botany,90:17-31.
Karnosky DF,Skelly JM,Percy KE,et al.2007.Perspectives regarding 50 years of research on effects of tropospheric ozone air pollution on US forests.Environmental Pollution,147:489-506.
KivimenpM,Sutinen S,Valolahti H,et al.2017.Warming and elevated ozone differently modify needle anatomy of Norway spruce(Picea abies)and Scots pine(Pinus sylvestris).Canadian Journal of Forest Research,47:488-499.
Li CH,Meng J,Guo LY,et al.2016.Effects of ozone pollution on yield and quality of winter wheat under flixweed competition.Environmental and Experimental Botany,129:77-84.
Pang J,Kobayashi K,Zhu JG.2009.Yield and photosynthetic characteristics of flag leaves in Chinese rice(Oryza sativa L.)varieties subjected to free-air release of ozone.Agriculture,Ecosystems and Environment,132:203-211.
Pleijel H,Broberg MC,Uddling J,et al.2018.Current surface ozone concentrations significantly decrease wheat growth,yield and quality.Science of the Total Environment,613-614:687-692.
Rao S,Klimont Z,Smith SJ,et al.2017.Future air pollution in the shared socio-economic pathways.Global Environmental Change,42:346-358.
Sui JX,Wen MX,Jia LL,et al.2017.Effects of elevated ozone on Polka Dot Plant(Hypoestes phyllostachya)with variegated leaves.Bulletin of Environmental Contamination and Toxicology,99:445-451.
Sumita K,Madhoolika A,Aradhana S.2015.Effects of ambient and elevated CO2and ozone on physiological characteristics,antioxidative defense system and metabolites of potato in relation to ozone flux.Environmental and Experimental Botany,109:276-287.
Sun J,Feng Z,Ort D.2014.Impacts of rising tropospheric ozone on photosynthesis and metabolite levels on field grown soybean.Plant Science,226:147-161.
Tao FL,Feng ZZ,Tang HY,et al.2017.Effects of climate change,CO2and O3on wheat productivity in Eastern China,singly and in combination.Atmospheric Environment,153:182-193.
Tong L,Zhang HL,Yu J,et al.2017.Characteristics of surface ozone and nitrogen oxides at urban,suburban and rural sites in Ningbo,China.Atmospheric Research,187:57-68.
Vu NT,Kang HM,Kim YS,et al.2015.Growth,physiology,and abiotic stress response to abscisic acid in tomato seedlings.Horticulture Environment and Biotechnology,56:294-304.
Wang T,Xue L,Brimblecombe P,et al.2016.Ozone pollution in China:A review of concentrations,meteorological influences,chemical precursors,and effects.Science of the Total Environment,575:1582.
Wu FF,Zheng YF,Wu RJ,et al.2011.Concentration of O3at the atmospheric surface affects the changes characters of antioxidant enzyme activities in Triticum aestivum.Acta Ecologica Sinica,31:4019-4026.
Xin Y,Yuan X,Shang B,et al.2016.Moderate drought did not affect the effectiveness of ethylenediurea(EDU)in protecting Populus cathayana from ambient ozone.Science of the Total Environment,569:1536-1544.
Xu S,Chen W,He X.2012.Responses of growth,photosynthesis and VOC emissions of Pinus tabulaeformis Carr.exposure to elevated CO2and/or elevated O3in an urban area.Bulletin of Environmental Contamination and Toxicology,88:443-448.
Xu S,He X,Chen W,et al.2014.Elevated CO2ameliorated the adverse effect of elevated O3in previous-year and current-year needles of Pinus tabulaeformis in Urban area.Bulletin of Environmental Contamination and Toxicology,92:733-737.
Xu S,He X,Chen W,et al.2015.Differential sensitivity of four urban tree species to elevated O3.Urban Forestry&Urban Greening,14:1166-1173.
Yu H,Cao JX,Chen Z,et al.2018.Effects of elevated O3on physiological and biochemical responses in three kinds of trees native to subtropical forest in China during non-growing period.Environmental Pollution,34:716-725.
高峰,李品,冯兆忠.2017.臭氧与干旱对植物复合影响的研究进展.植物生态学报,41(2):252-268.
高俊凤.2006.植物生理学实验指导.北京:高等教育出版社.
郭雄飞,黎华寿,杨宝仪,等.2015.秋枫和木棉对大气臭氧浓度升高的生理响应.中南林业科技大学学报,35(2):49-53.
黄玉源,黄益宗,李秋霞,等.2006.臭氧对南方3种木本植物的急性伤害症状及其生理指标变化.生态环境,15(4):674-681.
李莉,田士林.2007.酶联免疫(ELISA)分析晋麦叶片中脱落酸的含量.安徽农业科学,35(23):7098-7099.
李梅,徐胜,张恒庆,等.2014.高浓度O3对水蜡叶表皮气孔及其叶组织结构特征的影响.生态学杂志,33(1):53-58.
李苗苗.2014.几种典型亚热带森林植物对臭氧胁迫的光合生理响应(硕士学位论文).北京:中国林业科学研究院.
列淦文,叶龙华,薛立.2014.臭氧胁迫对植物主要生理功能的影响.生态学报,34(2):294-306.
欧英娟.2014.O3/CO2浓度升高对两种城市绿化植物光合特性及生长发育的影响(硕士学位论文).长沙:湖南农业大学.
阮亚男,徐胜,郭龙,等.2017.大气臭氧浓度升高对银杏叶片活性氧代谢及相关基因表达的影响.应用生态学报,28(11):3479-3486.
王闯,王帅,杨碧波,等.2015.气象条件对沈阳市环境空气臭氧浓度影响研究.中国环境监测,31(3):32-37.
吴丹.2016.地面臭氧污染对树木叶片组织结构的影响(硕士学位论文).北京:北京林业大学.
徐胜,何兴元,陈玮,等.2009.高浓度臭氧对树木生理生态的影响.生态学报,29(1):368-377.
尹海龙,田长彦.2013.氮调控对盐环境下甜菜功能叶光系统Ⅱ荧光特性的影响.植物生态学报,37(2):122-131.
赵诣,徐胜,何兴元,等.2014.三种冷季型草坪草对高浓度O3的生理响应.生态学杂志,12(3):203-208.
郑有飞,石茗化,吴荣军,等.2013.遮阴和臭氧浓度增加对冬小麦叶片光合作用的影响.农业环境科学,32(10):1925-1933.
庄明浩,李迎春,陈双林.2011a.毛竹和四季竹对臭氧胁迫的耐受力差异.生态学杂志,30(10):2191-2196.
庄明浩,李迎春,郭子武,等.2011b.臭氧浓度升高对毛竹叶片光合色素和抗性生理的影响.生态科学,30(6):607-612.
Anav A,De Marco A,Proietti C.2016.Comparing concentration-based(AOT40)and stomatal uptake(PODY)metrics for ozone risk assessment to European forests.Global Change Biology,22:1608-1627.
Brbara BM,Edenise SA,Mauro AM,et al.2018.Ozone affects leaf physiology and causes injury to foliage of native tree species from the tropical Atlantic Forest of southern Brazil.Science of the Total Environment,610-611:912-925.
Bortolin RC,Caregnato FF,Armando MD,et al.2014.Effects of chronic elevated ozone concentration on the redox state and fruit yield of red pepper plant Capsicum baccatum.Ecotoxicology and Environmental Safety,100:114-121.
Brando SE,Bulbovas P,Lima MEL,et al.2017.Biochemical leaf traits as indicators of tolerance potential in tree species from the Brazilian Atlantic forest against oxidative environmental stressors.Science of the Total Environment,575:406-417.
Feng Z,Sun J,Wan W,et al.2014.Evidence of widespread ozone induced visible injury on plants in Beijing.China.Environmental Pollution,193:296-301.
Feng ZZ,Hu EZ,Wang XK,et al.2015.Ground-level O3pollution and its impacts on food crops in China:A review.Environmental Pollution,199:42-48.
Feng ZZ,Kobayashi K,Ainsworth E.2008.Impact of elevated ozone concentration on growth,physiology,and yield of wheat(Triticum aestivum L.).Global Change Biology,14:2696-2708.
Ghanbari F,Sayyari M.2018.Controlled drought stress affects the chilling-hardening capacity of tomato seedlings as indicated by changes in phenol metabolisms,antioxidant enzymes activity,osmolytes concentration and abscisic acid accumulation.Scientia Horticulturae,229:167-174.
Ghimire RP,KivimenpM,Kasurinen A,et al.2017.Herbivore-induced BVOC emissions of Scots pine under warming,elevated ozone and increased nitrogen availability in an open-field exposure.Agricultural and Forest Meteorology,242:21-32.
Gong XJ,Chen XS,Feng SQ,et al.2017.Effects of ozone stress on physiological and biochemical characteristics dwarfing rootstock M9T337.Agricultural Science&Technology,18:579-582+606.
Hodges DM,Delong JM,Forney CF,et al.1999.Improving the thiobarbituric acid-substances assay for estimating lipid peroxidation in plant tissues containing anthocyanin and other interfering compounds.Planta,207:604-611.
Huttunen S,Manninen S.2013.A review of ozone responses in Scots pine(Pinus sylvestris).Environmental and Experimental Botany,90:17-31.
Karnosky DF,Skelly JM,Percy KE,et al.2007.Perspectives regarding 50 years of research on effects of tropospheric ozone air pollution on US forests.Environmental Pollution,147:489-506.
KivimenpM,Sutinen S,Valolahti H,et al.2017.Warming and elevated ozone differently modify needle anatomy of Norway spruce(Picea abies)and Scots pine(Pinus sylvestris).Canadian Journal of Forest Research,47:488-499.
Li CH,Meng J,Guo LY,et al.2016.Effects of ozone pollution on yield and quality of winter wheat under flixweed competition.Environmental and Experimental Botany,129:77-84.
Pang J,Kobayashi K,Zhu JG.2009.Yield and photosynthetic characteristics of flag leaves in Chinese rice(Oryza sativa L.)varieties subjected to free-air release of ozone.Agriculture,Ecosystems and Environment,132:203-211.
Pleijel H,Broberg MC,Uddling J,et al.2018.Current surface ozone concentrations significantly decrease wheat growth,yield and quality.Science of the Total Environment,613-614:687-692.
Rao S,Klimont Z,Smith SJ,et al.2017.Future air pollution in the shared socio-economic pathways.Global Environmental Change,42:346-358.
Sui JX,Wen MX,Jia LL,et al.2017.Effects of elevated ozone on Polka Dot Plant(Hypoestes phyllostachya)with variegated leaves.Bulletin of Environmental Contamination and Toxicology,99:445-451.
Sumita K,Madhoolika A,Aradhana S.2015.Effects of ambient and elevated CO2and ozone on physiological characteristics,antioxidative defense system and metabolites of potato in relation to ozone flux.Environmental and Experimental Botany,109:276-287.
Sun J,Feng Z,Ort D.2014.Impacts of rising tropospheric ozone on photosynthesis and metabolite levels on field grown soybean.Plant Science,226:147-161.
Tao FL,Feng ZZ,Tang HY,et al.2017.Effects of climate change,CO2and O3on wheat productivity in Eastern China,singly and in combination.Atmospheric Environment,153:182-193.
Tong L,Zhang HL,Yu J,et al.2017.Characteristics of surface ozone and nitrogen oxides at urban,suburban and rural sites in Ningbo,China.Atmospheric Research,187:57-68.
Vu NT,Kang HM,Kim YS,et al.2015.Growth,physiology,and abiotic stress response to abscisic acid in tomato seedlings.Horticulture Environment and Biotechnology,56:294-304.
Wang T,Xue L,Brimblecombe P,et al.2016.Ozone pollution in China:A review of concentrations,meteorological influences,chemical precursors,and effects.Science of the Total Environment,575:1582.
Wu FF,Zheng YF,Wu RJ,et al.2011.Concentration of O3at the atmospheric surface affects the changes characters of antioxidant enzyme activities in Triticum aestivum.Acta Ecologica Sinica,31:4019-4026.
Xin Y,Yuan X,Shang B,et al.2016.Moderate drought did not affect the effectiveness of ethylenediurea(EDU)in protecting Populus cathayana from ambient ozone.Science of the Total Environment,569:1536-1544.
Xu S,Chen W,He X.2012.Responses of growth,photosynthesis and VOC emissions of Pinus tabulaeformis Carr.exposure to elevated CO2and/or elevated O3in an urban area.Bulletin of Environmental Contamination and Toxicology,88:443-448.
Xu S,He X,Chen W,et al.2014.Elevated CO2ameliorated the adverse effect of elevated O3in previous-year and current-year needles of Pinus tabulaeformis in Urban area.Bulletin of Environmental Contamination and Toxicology,92:733-737.
Xu S,He X,Chen W,et al.2015.Differential sensitivity of four urban tree species to elevated O3.Urban Forestry&Urban Greening,14:1166-1173.
Yu H,Cao JX,Chen Z,et al.2018.Effects of elevated O3on physiological and biochemical responses in three kinds of trees native to subtropical forest in China during non-growing period.Environmental Pollution,34:716-725.