UV-B辐射胁迫对药用植物次生代谢产物的影响研究进展
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摘要
UV-B辐射胁迫对地球生物的生命活动产生极大影响,其中药用植物次生代谢产物积累是对UV-B辐射胁迫的一种响应,其相关研究对提高药用植物次生代谢产物的产量、品质和合成效率,推动药用植物次生代谢产物的开发和利用,推进增加其药效技术的调控和创新具有参考意义。综述了UV-B辐射胁迫对药用植物次生代谢产物(萜类化合物、酚类化合物和含氮化合物)产量、质量及合成效率的影响,并深入地探讨了UV-B辐射胁迫对部分药用植物次生代谢产物的合成酶、合成途径及主要分子机制的影响。此外,还对UV-B辐射与其他因子复合作用作了概述。
UV-B radiation stress significantly affects all life activities of earth organisms.The accumulation of secondary metabolites in medicinal plants is one of the response mechanism of UV-B radiation stress.The significance of relevant studies provide references for improving the yield,quality,and the synthetic efficiency of secondary metabolites in medicinal plants; and promoting its development and utilization,also enhancing its efficacy by relevant technical regulations and renovations.This study systematically investigated the impacts of the enhanced UV-B radiation on the yield,quality,synthetic efficiency of secondary metabolites produced by medicinal plants including terpene,phenolic,and nitrogen-containingcompound.Furthermore,the study explored a portion of secondary metabolites to discover the influences of enhanced UV-B radiation on their synthase,synthetic pathway,and the primary molecular mechanism.In addition,this study summarized the co-effects of UV-B radiation and other factors.
UV-B radiation stress significantly affects all life activities of earth organisms.The accumulation of secondary metabolites in medicinal plants is one of the response mechanism of UV-B radiation stress.The significance of relevant studies provide references for improving the yield,quality,and the synthetic efficiency of secondary metabolites in medicinal plants; and promoting its development and utilization,also enhancing its efficacy by relevant technical regulations and renovations.This study systematically investigated the impacts of the enhanced UV-B radiation on the yield,quality,synthetic efficiency of secondary metabolites produced by medicinal plants including terpene,phenolic,and nitrogen-containingcompound.Furthermore,the study explored a portion of secondary metabolites to discover the influences of enhanced UV-B radiation on their synthase,synthetic pathway,and the primary molecular mechanism.In addition,this study summarized the co-effects of UV-B radiation and other factors.
引文
[1]徐佳妮,雷梦琦,鲁瑞琪,等.UV-B辐射增强对植物影响的研究进展[J].基因组学与应用生物学,2015,34(6):1347-1352.
[2]利荣千,王建波.植物逆境细胞及生理学[M].湖北:武汉大学出版社,2002:321-327.
[3]吴永波,薛建辉.UV-B辐射增强对植物影响的研究进展[J].世界林业研究,2004,17(3):29-31.
[4]邱德有.试论药用植物有效成分基因调控的研究进展[J].世界科学技术,2000(3):30-34.
[5]陈晓亚,刘培.植物次生代谢的分子生物学及基因工程[J].生命科学,1996(2):8-11.
[6]刘佳钰,王蓉,张弛,等.UV-B辐射与植物次级代谢产物变化的研究进展[J].基因组学与应用生物学,2017,36(8):3157-3166.
[7]Pan W S,Zheng L P,Tian H.Transcriptome responses involved in artemisinin production in Artemisia annua L.under UV-B radiation[J].Journal of Photochemistry and Photobiology B:Biology,2014,140:292-300.
[8]王园,党悦方,张典,等.夏枯草幼苗中有效成分与生理指标对增强UV-B辐射的动态响应[J].西北大学学报(自然科学版),2017,47(3):414-421.
[9]莫运才,曾令杰,黄涵,等.UV-B辐射对铁皮石斛叶片光合色素、类黄酮及PAL酶活性的影响[J].贵州农业科学,2015,43(7):34-37.
[10]Takshak S,Agrawal S B.Secondary metabolites and phenylpropanoid pathway enzymes as influenced under supplemental ultraviolet-B radiation in Withania somnifera Dunal,an indigenous medicinal plant[J].Journal of Photoehemistry&Photobiology,2014,40:332-343.
[11]郭建军,贵梦园,张佩佩,等.景天三七对紫外线B辐射敏感性研究[J].河南农业大学学报,2017,51(1):71-75.
[12]褚润,陈年来,韩国君,等.UV-B辐射增强对芦苇生长及生理特性的影响[J].环境科学学报,2018(5):2074-2081.
[13]Si C,Yao X Q,He X L,et al.Effects of enhanced UV-B radiation on biochemical traits in post harvest flowers of medicinal chrysanthemum[J].Photochemistry&Photobiology,2015,91(4):845-850.
[14]张小冰.植物次生代谢产物的功能及应用概述[J].生物学教学,2013,8(11):7-10.
[15]杨逢建,庞海河,张学科.光胁迫对南方红豆杉叶片中叶绿体色素和紫杉醇含量的影响[J].植物研究,2007,7(5):556-558.
[16]崔磊.光酶诱导提高南方红豆杉枝叶中紫杉烷的含量及适用于光酶诱导的仪器的设计[D].杭州:浙江大学,2011.
[17]李德文,刘英,李美兰,等.NO对UV-B辐射下南方红豆杉幼苗生理特性的影响[J].东北林业大学学报,2015,3(7):27-30.
[18]郑雯.南方红豆杉和银杏叶响应紫外辐射及土牛膝治疗大鼠关节炎的蛋白质组学研究[D].杭州:浙江大学,2016.
[19]Rai R,Meena R P,Smita S S,et al.UV-B and UV-C pre-treatments induce physiological changes and artemisinin biosynthesis in Artemisia annua L.—An antimalarial plant[J].J Photochem Photobiol B,2011,105(3):216-225.
[20]付金颖,杨超,李德文,等.环境UV-B辐射增强对叶用杜仲林叶片光合作用与活性物质含量的影响[J].安徽农业科学,2017,45(26):6-10.
[21]Gil M,Pontin M,Berli F,et al.Metabolism of terpenes in the response of grape(Vitis vinifera L.)leaf tissues to UV-B radiation[J].Phytochemistry,2012,77:89-98.
[22]马春晖.不同发育阶段祁菊花序对UV-B辐射的响应[D].保定:河北大学,2016.
[23]Gu X D,Sun M Y,Zhang L,et al.UV-B induced changes in the secondary metabolites of Morus alba L.leaves[J].Molecules,2010,15(5):2980-2993.
[24]Zhu W,Yang B,Komatsu S,et al.Binary stress induces an increase in indole alkaloid biosynthesis in Catharanthus roseus[J].Frontiers in Plant Science,2015,6:582.
[25]Ning W,Peng X,Ma L,et al.Enhanced secondary metabolites production and antioxidant activity in post harvest Lonicera japonica(Thunb.)in response to UV radiation[J].Innovative Food Science&Emerging Technologies,2012,3:231-243.
[26]Schenke D,Boettcher C,Scheel D.Crosstalk between abiotic ultraviolet-B stress and biotic(flg 22)stress signaling in Arabidopsis prevents flavonol accumulation in favor of pathogen defense compound production[J].Plant,Cell&Environment,2011,34(11):1849-1864.
[27]Takshak S,Agrawal S B.Effect of ultraviolet-B radiation on biomass production,lipid peroxidation,reactive oxygen species,and antioxidants in Withania somnifera[J].Biologia Plantarum,2014,58(2):328-334.
[28]Tripathi R,Agrawal S B.Evaluation of changes in lipid peroxidation,ROS production,surface structures,secondary metabolites,and yield of linseed(Linum usitatissimum L.)under individual and combined stress of ultraviolet-B and ozone using open top chambers[J].Indian Journal of Biochemistry&Biophysics,2013,50(4):318-325.
[29]Chen H,Feng Y,Wang L,et al.Transcriptome profiling of the UV-B stress response in the desert shrub,Lycium ruthenicum[J].Molecular Biology Reports,2015,42(3):639-649.
[30]杨丙贤.圆锥铁线莲叶片响应紫外诱导机制及香豆素生物合成途径的系统性研究[D].杭州:浙江大学,2016.
[31]Liu M L,Cao B,Zhou S H,et al.Responses of the flavonoid pathway to UV-B radiation stress and the correlation with the lipid antioxidant characteristics in the desert plant Caryopteris mongolica[J].Acta Ecological Sinica,2012,32(3):150-155.
[32]Kumari R,Agrawal S B.Supplemental UV-B induced changes in leaf morphology,physiology,and secondary metabolites of an Indian aromatic plant Cymbopogon citratus(D.C.)Staph under natural field conditions[J].International Journal of Environment Studies,2010,67(5):655-675.
[33]高欣.增补UV-B辐射对黄檗幼苗生理及次生代谢产物的含量的影响[D].哈尔滨:东北林业大学,2015.
[34]莫运才.UV-B辐射结合不同氮素营养水平对铁皮石斛生长及主要活性成分的影响[D].广州:广东药科大学,2016.
[35]冯源,李云霞,董晓东.滇黄芩幼苗对增强UV-B辐射与干旱胁迫的生理响应[J].湖北农业科学,2016,55(2):413-417.
[36]Kalidhasan N,Bhagavan N B,Kannan N D.Ultraviolet-B(280-320 nm)enhanced radiation induced changes in secondary metabolites and photosystem-Ⅱof medicinal plant,Withania somnifera Dunal.[J].Journal of Medicinal Plant Research,2013,7(42):3112-3120.
[37]Amako K,Chen G X,Asada K.Separate assays specific for ascorbate peroxidase and guaiacol peroxidase and for the chloroplastic and cytosolic isozymes of ascorbate peroxidase in plants[J].Plant&Cell Physiology,1994,35(3):497-504.
[38]Yamasaki H,Sakihama Y,Ikehara N.Flavonoid-peroxidase reaction as a detoxification mechanism of plant cells against H2O2[J].Plant Physiology,1997,115(4):1405-1412.
[39]Harborne J B,Williams C A.Advances in flavonoid research since 1992[J].Phytochemistry,2000,55(6):481-504.
[40]Suzuki T,Honda Y,Funatsuki W,et al.Purification and characterization of flavonol 3-glycosidase,and its activity during the ripening in tartary buckwheat seeds[J].Plant Science,2002,163(3):417-423.
[41]Suzuki T,Honda Y,Mukasa Y J.Effects of UV-B radiation,cold and desiccation stress on rutin concentration and rutin glucosidase activity in tartary buckwheat(Fagopyrum tataricum)leaves[J].Plant Science,2005,168(5):1303-1307.
[42]党悦方.增强UV-B辐射对夏枯草生长及多种化学指标成分的影响[D].西安:西北大学,2015.
[43]李德文,李美兰,于景华,等.外源NO对UV-B胁迫下红豆杉抗氧化系统的影响[J].生态学杂志,2012,31(9):2203-2208.
[44]范方媛.UV-B辐射对离体茶树新梢物质代谢及其相关基因表达的影响[D].杭州:浙江大学,2014.
[45]孙莹.紫外线-B(UV-B)辐射增强对葡萄果实品质的影响[D].杨凌:西北农林科技大学,2009.
[46]孙莹,张振文,岳泰新,等.不同强度紫外线UV-B辐射对葡萄品质的影响[J].中国酿造,2009(10):15-18.
[47]李亚敏,岳明,林小英.补充UV-B辐射对浙贝母生长、生理及生物碱的影响[J].华西药学杂志,2008(1):77-80.
[48]温泉,张楠,曹瑞霞,等.增强UV-B对黄连代谢及小檗碱含量的影响[J].中国中药杂志,2011,36(22):3063-3069.
[49]周心渝,曹瑞霞,张红敏,等.半夏光合生理特性及总生物碱对增强UV-B的响应[J].西南大学学报(自然科学版),2013,35(6):166-172.
[50]郭玉梅,曾令杰,梁淑颖,等.UV-B辐射对铁皮石斛生长及主要次生代谢产物的影响[J].北方园艺,2016(17):154-156.
[51]王海霞.增强UV-B辐射对喜树幼苗的生理影响[D].西安:西北大学,2008.
[52]王海霞,刘文哲.增强UV-B辐射对喜树幼苗生物量和两种生物碱含量的影响[J].植物科学学报,2011,29(6):712-717.
[53]王玲丽,周晓君,刘文哲.增强UV-B辐射对喜树生理指标及喜树碱含量的影响[J].西北植物学报,2016,36(5):979-986.
[54]卢克欢,苏贝贝,郭双,等.UV-B辐射对颠茄氮代谢及次生代谢产物含量的影响[J].中国生物化学与分子生物学报,2017,33(10):1062-1069.
[55]苏贝贝.UV-B辐射对颠茄生理特性及次生代谢产物含量的影响[D].重庆:西南大学,2016.
[56]Mewis I,Appel H M,Hom A,et al.Major signaling pathways modulate Arabidopsis thaliana(L.)glucosinolate accumulation and response to both phloem feeding and chewing insects[J].Plant Physiology,2005,138(2):1149-1162.
[57]Huyskens-Keil S,Eichholz I,Kroh L W,et al.UV-B induced changes of phenol composition and antioxidant activity in black currant fruit(Ribes nigrum L.)[J].Journal of Applied Botany&Food Quality,2007,81(2):140-144.
[58]Pérez C P,Ulrichs C,Huyskens-Keil S,et al.Composition of carotenoids in tomato fruits as affected by moderate UV-B radiation before harvest[J].Acta Horticulture,2009,821(1):217-222.
[59]Wang Y,Xu W J,Yan X F,et al.Glucosinolate content and related gene expression in response to enhanced UVB radiation in Arabidopsis[J].African Journal of Biotechnology,2011,10(34):6481-6491.
[60]Schreiner M,Krumbein A,Mewis I,et al.Short-term and moderate UV-B radiation effects on secondary plant metabolism in different organs of nasturtium(Tropaeolum majus L.)[J].Innovative Food Science&Emerging Technologies,2009,10(1):93-96.
[61]徐文佳.UV-B辐射对拟南芥芥子油苷代谢的影响[D].哈尔滨:东北林业大学,2010.
[62]牟晓飞,于涛.UV-B辐射对植物物质代谢的影响[J].现代农业科技,2012(8):46,50.
[63]牟晓飞.拟南芥芥子油苷对UV-B辐射的响应[D].哈尔滨:东北林业大学,2012.
[2]利荣千,王建波.植物逆境细胞及生理学[M].湖北:武汉大学出版社,2002:321-327.
[3]吴永波,薛建辉.UV-B辐射增强对植物影响的研究进展[J].世界林业研究,2004,17(3):29-31.
[4]邱德有.试论药用植物有效成分基因调控的研究进展[J].世界科学技术,2000(3):30-34.
[5]陈晓亚,刘培.植物次生代谢的分子生物学及基因工程[J].生命科学,1996(2):8-11.
[6]刘佳钰,王蓉,张弛,等.UV-B辐射与植物次级代谢产物变化的研究进展[J].基因组学与应用生物学,2017,36(8):3157-3166.
[7]Pan W S,Zheng L P,Tian H.Transcriptome responses involved in artemisinin production in Artemisia annua L.under UV-B radiation[J].Journal of Photochemistry and Photobiology B:Biology,2014,140:292-300.
[8]王园,党悦方,张典,等.夏枯草幼苗中有效成分与生理指标对增强UV-B辐射的动态响应[J].西北大学学报(自然科学版),2017,47(3):414-421.
[9]莫运才,曾令杰,黄涵,等.UV-B辐射对铁皮石斛叶片光合色素、类黄酮及PAL酶活性的影响[J].贵州农业科学,2015,43(7):34-37.
[10]Takshak S,Agrawal S B.Secondary metabolites and phenylpropanoid pathway enzymes as influenced under supplemental ultraviolet-B radiation in Withania somnifera Dunal,an indigenous medicinal plant[J].Journal of Photoehemistry&Photobiology,2014,40:332-343.
[11]郭建军,贵梦园,张佩佩,等.景天三七对紫外线B辐射敏感性研究[J].河南农业大学学报,2017,51(1):71-75.
[12]褚润,陈年来,韩国君,等.UV-B辐射增强对芦苇生长及生理特性的影响[J].环境科学学报,2018(5):2074-2081.
[13]Si C,Yao X Q,He X L,et al.Effects of enhanced UV-B radiation on biochemical traits in post harvest flowers of medicinal chrysanthemum[J].Photochemistry&Photobiology,2015,91(4):845-850.
[14]张小冰.植物次生代谢产物的功能及应用概述[J].生物学教学,2013,8(11):7-10.
[15]杨逢建,庞海河,张学科.光胁迫对南方红豆杉叶片中叶绿体色素和紫杉醇含量的影响[J].植物研究,2007,7(5):556-558.
[16]崔磊.光酶诱导提高南方红豆杉枝叶中紫杉烷的含量及适用于光酶诱导的仪器的设计[D].杭州:浙江大学,2011.
[17]李德文,刘英,李美兰,等.NO对UV-B辐射下南方红豆杉幼苗生理特性的影响[J].东北林业大学学报,2015,3(7):27-30.
[18]郑雯.南方红豆杉和银杏叶响应紫外辐射及土牛膝治疗大鼠关节炎的蛋白质组学研究[D].杭州:浙江大学,2016.
[19]Rai R,Meena R P,Smita S S,et al.UV-B and UV-C pre-treatments induce physiological changes and artemisinin biosynthesis in Artemisia annua L.—An antimalarial plant[J].J Photochem Photobiol B,2011,105(3):216-225.
[20]付金颖,杨超,李德文,等.环境UV-B辐射增强对叶用杜仲林叶片光合作用与活性物质含量的影响[J].安徽农业科学,2017,45(26):6-10.
[21]Gil M,Pontin M,Berli F,et al.Metabolism of terpenes in the response of grape(Vitis vinifera L.)leaf tissues to UV-B radiation[J].Phytochemistry,2012,77:89-98.
[22]马春晖.不同发育阶段祁菊花序对UV-B辐射的响应[D].保定:河北大学,2016.
[23]Gu X D,Sun M Y,Zhang L,et al.UV-B induced changes in the secondary metabolites of Morus alba L.leaves[J].Molecules,2010,15(5):2980-2993.
[24]Zhu W,Yang B,Komatsu S,et al.Binary stress induces an increase in indole alkaloid biosynthesis in Catharanthus roseus[J].Frontiers in Plant Science,2015,6:582.
[25]Ning W,Peng X,Ma L,et al.Enhanced secondary metabolites production and antioxidant activity in post harvest Lonicera japonica(Thunb.)in response to UV radiation[J].Innovative Food Science&Emerging Technologies,2012,3:231-243.
[26]Schenke D,Boettcher C,Scheel D.Crosstalk between abiotic ultraviolet-B stress and biotic(flg 22)stress signaling in Arabidopsis prevents flavonol accumulation in favor of pathogen defense compound production[J].Plant,Cell&Environment,2011,34(11):1849-1864.
[27]Takshak S,Agrawal S B.Effect of ultraviolet-B radiation on biomass production,lipid peroxidation,reactive oxygen species,and antioxidants in Withania somnifera[J].Biologia Plantarum,2014,58(2):328-334.
[28]Tripathi R,Agrawal S B.Evaluation of changes in lipid peroxidation,ROS production,surface structures,secondary metabolites,and yield of linseed(Linum usitatissimum L.)under individual and combined stress of ultraviolet-B and ozone using open top chambers[J].Indian Journal of Biochemistry&Biophysics,2013,50(4):318-325.
[29]Chen H,Feng Y,Wang L,et al.Transcriptome profiling of the UV-B stress response in the desert shrub,Lycium ruthenicum[J].Molecular Biology Reports,2015,42(3):639-649.
[30]杨丙贤.圆锥铁线莲叶片响应紫外诱导机制及香豆素生物合成途径的系统性研究[D].杭州:浙江大学,2016.
[31]Liu M L,Cao B,Zhou S H,et al.Responses of the flavonoid pathway to UV-B radiation stress and the correlation with the lipid antioxidant characteristics in the desert plant Caryopteris mongolica[J].Acta Ecological Sinica,2012,32(3):150-155.
[32]Kumari R,Agrawal S B.Supplemental UV-B induced changes in leaf morphology,physiology,and secondary metabolites of an Indian aromatic plant Cymbopogon citratus(D.C.)Staph under natural field conditions[J].International Journal of Environment Studies,2010,67(5):655-675.
[33]高欣.增补UV-B辐射对黄檗幼苗生理及次生代谢产物的含量的影响[D].哈尔滨:东北林业大学,2015.
[34]莫运才.UV-B辐射结合不同氮素营养水平对铁皮石斛生长及主要活性成分的影响[D].广州:广东药科大学,2016.
[35]冯源,李云霞,董晓东.滇黄芩幼苗对增强UV-B辐射与干旱胁迫的生理响应[J].湖北农业科学,2016,55(2):413-417.
[36]Kalidhasan N,Bhagavan N B,Kannan N D.Ultraviolet-B(280-320 nm)enhanced radiation induced changes in secondary metabolites and photosystem-Ⅱof medicinal plant,Withania somnifera Dunal.[J].Journal of Medicinal Plant Research,2013,7(42):3112-3120.
[37]Amako K,Chen G X,Asada K.Separate assays specific for ascorbate peroxidase and guaiacol peroxidase and for the chloroplastic and cytosolic isozymes of ascorbate peroxidase in plants[J].Plant&Cell Physiology,1994,35(3):497-504.
[38]Yamasaki H,Sakihama Y,Ikehara N.Flavonoid-peroxidase reaction as a detoxification mechanism of plant cells against H2O2[J].Plant Physiology,1997,115(4):1405-1412.
[39]Harborne J B,Williams C A.Advances in flavonoid research since 1992[J].Phytochemistry,2000,55(6):481-504.
[40]Suzuki T,Honda Y,Funatsuki W,et al.Purification and characterization of flavonol 3-glycosidase,and its activity during the ripening in tartary buckwheat seeds[J].Plant Science,2002,163(3):417-423.
[41]Suzuki T,Honda Y,Mukasa Y J.Effects of UV-B radiation,cold and desiccation stress on rutin concentration and rutin glucosidase activity in tartary buckwheat(Fagopyrum tataricum)leaves[J].Plant Science,2005,168(5):1303-1307.
[42]党悦方.增强UV-B辐射对夏枯草生长及多种化学指标成分的影响[D].西安:西北大学,2015.
[43]李德文,李美兰,于景华,等.外源NO对UV-B胁迫下红豆杉抗氧化系统的影响[J].生态学杂志,2012,31(9):2203-2208.
[44]范方媛.UV-B辐射对离体茶树新梢物质代谢及其相关基因表达的影响[D].杭州:浙江大学,2014.
[45]孙莹.紫外线-B(UV-B)辐射增强对葡萄果实品质的影响[D].杨凌:西北农林科技大学,2009.
[46]孙莹,张振文,岳泰新,等.不同强度紫外线UV-B辐射对葡萄品质的影响[J].中国酿造,2009(10):15-18.
[47]李亚敏,岳明,林小英.补充UV-B辐射对浙贝母生长、生理及生物碱的影响[J].华西药学杂志,2008(1):77-80.
[48]温泉,张楠,曹瑞霞,等.增强UV-B对黄连代谢及小檗碱含量的影响[J].中国中药杂志,2011,36(22):3063-3069.
[49]周心渝,曹瑞霞,张红敏,等.半夏光合生理特性及总生物碱对增强UV-B的响应[J].西南大学学报(自然科学版),2013,35(6):166-172.
[50]郭玉梅,曾令杰,梁淑颖,等.UV-B辐射对铁皮石斛生长及主要次生代谢产物的影响[J].北方园艺,2016(17):154-156.
[51]王海霞.增强UV-B辐射对喜树幼苗的生理影响[D].西安:西北大学,2008.
[52]王海霞,刘文哲.增强UV-B辐射对喜树幼苗生物量和两种生物碱含量的影响[J].植物科学学报,2011,29(6):712-717.
[53]王玲丽,周晓君,刘文哲.增强UV-B辐射对喜树生理指标及喜树碱含量的影响[J].西北植物学报,2016,36(5):979-986.
[54]卢克欢,苏贝贝,郭双,等.UV-B辐射对颠茄氮代谢及次生代谢产物含量的影响[J].中国生物化学与分子生物学报,2017,33(10):1062-1069.
[55]苏贝贝.UV-B辐射对颠茄生理特性及次生代谢产物含量的影响[D].重庆:西南大学,2016.
[56]Mewis I,Appel H M,Hom A,et al.Major signaling pathways modulate Arabidopsis thaliana(L.)glucosinolate accumulation and response to both phloem feeding and chewing insects[J].Plant Physiology,2005,138(2):1149-1162.
[57]Huyskens-Keil S,Eichholz I,Kroh L W,et al.UV-B induced changes of phenol composition and antioxidant activity in black currant fruit(Ribes nigrum L.)[J].Journal of Applied Botany&Food Quality,2007,81(2):140-144.
[58]Pérez C P,Ulrichs C,Huyskens-Keil S,et al.Composition of carotenoids in tomato fruits as affected by moderate UV-B radiation before harvest[J].Acta Horticulture,2009,821(1):217-222.
[59]Wang Y,Xu W J,Yan X F,et al.Glucosinolate content and related gene expression in response to enhanced UVB radiation in Arabidopsis[J].African Journal of Biotechnology,2011,10(34):6481-6491.
[60]Schreiner M,Krumbein A,Mewis I,et al.Short-term and moderate UV-B radiation effects on secondary plant metabolism in different organs of nasturtium(Tropaeolum majus L.)[J].Innovative Food Science&Emerging Technologies,2009,10(1):93-96.
[61]徐文佳.UV-B辐射对拟南芥芥子油苷代谢的影响[D].哈尔滨:东北林业大学,2010.
[62]牟晓飞,于涛.UV-B辐射对植物物质代谢的影响[J].现代农业科技,2012(8):46,50.
[63]牟晓飞.拟南芥芥子油苷对UV-B辐射的响应[D].哈尔滨:东北林业大学,2012.