鸳鸯茉莉花色变化过程中的生理生化特性研究
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摘要
以3年生鸳鸯茉莉(Brunfelsia acuminata Benth.)5个不同花期(花蕾期、初开期、全开期、过渡期、末花期)的花瓣为试材,研究其开花过程中花瓣的色度值、类黄酮、类胡萝卜素、花色苷含量、细胞p H值以及相关酶活性的变化,并分析各指标之间的相关性。结果显示,除全开期外,其它4个时期各色度值变化不明显,初开期、全开期和过渡期,明亮参数L值逐渐增大,红绿参数a值、黄蓝参数b值和彩度C值急剧降低;类黄酮含量在过渡期最低,其它阶段无显著差异;类胡萝卜素含量变化呈上升-下降-稳定的趋势,全开期达到最高;花色苷含量则随开花进程不断下降,花蕾期最高;花蕾期、初开期、全开期的SOD活性变化不显著,过渡期时显著升高,末花期又迅速回落到最低值;POD活性不断上升,末花期为花蕾期的28.73倍;CAT活性前期变化不显著,末花期时有一定幅度的升高;花瓣色度各指标、花色苷含量、上述酶的活性、p H值等几个参数间均存在极显著相关性,但SOD活性、类胡萝卜素、类黄酮与所有参数的相关性均不显著。研究结果表明花色苷是影响鸳鸯茉莉花色变化的决定性代谢产物,POD和CAT活性是调节花色苷降解速率的主要抗氧化酶。
To explain the floral chromogenic mechanism of Brunfelsia acuminata Benth.,3-yearold B. acuminata petals during different flowering stages,including unfold-petal stage(UPS),initial-flowering stage(IFS),full-flowering stage(FFS),transitional-flowering stage(TFS),and flowering-wilting stage(FWS),were used and the floral color index,flavonoid content(FC),anthocyanin content(AC),carotenoid content(CC),p H index,relative enzyme activity and their correlations were measured. Results showed the flowering color change was not obvious during the UPS-IFS and TFS-FWS,the L value(bright parameter) increased gradually,whereas the A value(red/green parameter),B value(blue/yellow parameter),and C value(chroma) reduced sharply. The flavonoid content was lowest during the TFS,and there were no significant differences among the other stages. The carotenoid content trend wasincreasing-decreasing-stable, and the content during the FFS was the highest. The anthocyanin content decreased continuously from the UPS. The SOD activity from the UPS to FFS changed non-significantly but increased during the TFS and rapidly dropped to the lowest level during the FWS. The POD activity increased continuously and was highest during the FWS. The CAT activity changed non-significantly,and only increased in the late flowering stage. The correlations among the floral color index,anthocyanin content,POD activity,CAT activity,and p H value were significant, but were not significant among SOD activity,carotenoid content, flavonoid content, and other indexes. Anthocyanin was the decisive metabolite of B. acuminata flower color,and the activities of POD and CAT were the main antioxidant enzymes,which were involved in the adjustment of the anthocyanin degradation rate.
To explain the floral chromogenic mechanism of Brunfelsia acuminata Benth.,3-yearold B. acuminata petals during different flowering stages,including unfold-petal stage(UPS),initial-flowering stage(IFS),full-flowering stage(FFS),transitional-flowering stage(TFS),and flowering-wilting stage(FWS),were used and the floral color index,flavonoid content(FC),anthocyanin content(AC),carotenoid content(CC),p H index,relative enzyme activity and their correlations were measured. Results showed the flowering color change was not obvious during the UPS-IFS and TFS-FWS,the L value(bright parameter) increased gradually,whereas the A value(red/green parameter),B value(blue/yellow parameter),and C value(chroma) reduced sharply. The flavonoid content was lowest during the TFS,and there were no significant differences among the other stages. The carotenoid content trend wasincreasing-decreasing-stable, and the content during the FFS was the highest. The anthocyanin content decreased continuously from the UPS. The SOD activity from the UPS to FFS changed non-significantly but increased during the TFS and rapidly dropped to the lowest level during the FWS. The POD activity increased continuously and was highest during the FWS. The CAT activity changed non-significantly,and only increased in the late flowering stage. The correlations among the floral color index,anthocyanin content,POD activity,CAT activity,and p H value were significant, but were not significant among SOD activity,carotenoid content, flavonoid content, and other indexes. Anthocyanin was the decisive metabolite of B. acuminata flower color,and the activities of POD and CAT were the main antioxidant enzymes,which were involved in the adjustment of the anthocyanin degradation rate.
引文
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[8]付林江,李厚华,李玲,于航,王拉岐.金银花花色变化原因分析[J].林业科学,2013,49(10):155-161.Fu LJ,Li HH,Li L,Yu H,Wang LQ.Reason of flower color change in Lonicera japonica[J].Scientia Silvae Sinicae,2013,49(10):155-161.
[9]邓惠,向甘驹,郭友好,杨春锋.秦岭忍冬属4种植物的繁育系统及花色变化的研究[J].植物科学学报,2017,35(1):1-12.Deng H,Xiang GJ,Guo YH,Yang CF.Study on the breeding system and floral color change of four Lonicera species in the Qinling mountains[J].Plant Science Journal,2017,35(1):1-12.
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[14]李玲,杜绍华,张超,高金锋,李彦慧.4种花色仙客来色素及相关生理生化研究[J].河北农业大学学报,2012,35(2):25-29.Li L,Du SH,Zhang C,Gao JF,Li YH.A study on the pigment content and other related physiological and biochemical indices in four color Cylamen cultivars[J].Journal of Agricultural University of Hebei,2012,35(2):25-29.
[15]朱满兰,王亮生,张会金,徐彦军,郑绪辰,王丽金.耐寒睡莲花瓣中花青素苷组成及其与花色的关系[J].植物学报,2012,47(5):437-453.Zhu ML,Wang LS,Zhang HJ,Xu YJ,Zheng XC,Wang LJ.Relationship between the composition of anthocyanins and flower color variation in hardy water lily(Nymphaea spp.)cultivars[J].Chinese Bulletin of Botany,2012,47(5):437-453.
[16]程怡,张云婷,王清明,马建伟,张勇.月季花发育过程中花色变化的生理生化研究[J].西北植物学报,2014,34(4):733-739.Cheng Y,Zhang YT,Wang QM,Ma JW,Zhang Y.Correlation analysis of flower color during flower development of Rosa hybrida[J].Acta Botanica Boreali-Occidentalia Sinica,2014,34(4):733-739.
[17]Yan J,Wang ML,Zhang L.Light induces petal color change in Quisqualis indica[J].Plant Divers,2018,40:28-34.
[18]于晓南,张启翔.观赏植物的花色素苷与花色[J].林业科学,2002,38(3):147-153.Yu XN,Zhang QX.Anthocyanin in ornamental plant and color express[J].Scientia Silvae Sinicae,2002,38(3):147-153.
[19]戴思兰,洪艳.基于花青素苷合成和呈色机理的观赏植物花色改良分子育种[J].中国农业科学,2016,49(3):529-542.Dai SL,Hong Y.Molecular breeding for flower colors modification on ornamental plants based on the mechanism of anthocyanins biosynthesis and coloration[J].Scientia Agricultural Sinica,2016,49(3):529-542.
[20]安田齐著.花色的生理生物化学[M].傅玉兰译.北京:中国林业出版社,1989.
[21]于国瑞.色彩构成[M].修订版.北京:清华大学出版社,2012.
[22]Vaknin H,Bar-Akiva A,Ovadia R,Nissim-Levi A,Forer I,et al.Active anthocyanin degradation in Brunfelsia calycina flowers[J].Planta,2005,222(1):19-26.
[23]陈小琴,郭志雄,林琳,王蓓,柯毅湧.鸳鸯茉莉不同发育时期POD同工酶体系优化[J].热带作物学报,2014,35(8):1510-1516.Chen XQ,Guo ZX,Lin L,Wang B,Ke YY.POD isozyme system optimization of Brunfelsia acuminata(Solanaceae)in different development stages[J].Chinese Journal of Tropical Crops,2014,35(8):1510-1516.
[24]Qiu J,Xue XF,Chen FD,Li CH,Bolat N,et al.Quality evaluation of snow lotus(Saussurea):Quantitative chemical analysis and antioxidant activity assessment[J].Plant Cell Rep,2010,29:1325-1337.
[25]李合生.植物生理生化试验原理与技术[M].北京:高等教育出版社,2003.
[26]Pua EC,Davey MR.Plant Developmental Biology-Biotechnological Perspectives[M].Berlin:Springer Heidelberg,2010.
[27]Murray RB,David HL,Kevin MD,Gayle BM,Deepa P,et al.Isolation and antisense suppression of flavonoid 3',5'-hydroxylase modifies flower pigments and color in cyclamen[J].BMC Plant Biol,2010,10:107-118.
[28]Ferrante A,Vernieri P,Tognoni F,Serra G.Changes in abscisic acid and flower pigments during floral senescence of Petunia[J].Biol Plant,2006,50:581-585.
[2]赵昶灵,郭维明,陈俊愉.植物花色形成及其调控机理[J].植物学通报,2005,22(1):70-81.Zhao CL,Guo WM,Chen JY.Formation and regulation of flower color in higher plants[J].Chinese Bulletin of Botany,2005,22(1):70-81.
[3]Graeme DR,Schaefer HM.Floral colour change as a potential signal to pollinators[J].Curr Opin Plant Biol,2016,32:96-100.
[4]张超,陈光,李严慧.二乔玉兰开花过程中花色变化的生理生化机制[J].西北植物学报,2012,32(4):716-721.Zhang C,Chen G,Li YH.Physiological and biochemical mechanism of flower color variation during flowering season in Yulania×soulangeana[J].Acta Botanica BorealiOccidentalia Sinica,2012,32(4):716-721.
[5]贺窑青,马履一,桑子阳.红花玉兰花色形成的初步研究[J].西北植物学报,2010,30(11):2252-2257.He YQ,Ma LY,Sang ZY.Preliminary study on formation of flower color in Magnolia wufengensis[J].Acta Botanica Boreali-Occidentalia Sinica,2010,30(11):2252-2257.
[6]李崇晖,王亮生,舒庆艳,徐彦军,张洁.迎红杜鹃花色素组成及花色在开花过程中的变化[J].园艺学报,2008,35(7):1023-1030.Li CH,Wang LS,Shu QY,Xu YJ,Zhang J.Pigments composition of petals and floral color change during the blooming period in Rhododendron mucronulatum[J].Acta Horticulture Sinica,2008,35(7):1023-1030.
[7]葛雨萱,王亮生,徐彦军,刘政安,李崇晖,贾妮.蜡梅的花色和花色素组成及其在开花过程中的变化[J].园艺学报,2008,35(9):1331-1338.Ge YX,Wang LS,Xu YJ,Liu ZA,Li CH,Jia N.Flower color,pigment composition and their changes during flowering in Chimonanthus praecox[J].Acta Horticulture Sinica,2008,35(9):1331-1338.
[8]付林江,李厚华,李玲,于航,王拉岐.金银花花色变化原因分析[J].林业科学,2013,49(10):155-161.Fu LJ,Li HH,Li L,Yu H,Wang LQ.Reason of flower color change in Lonicera japonica[J].Scientia Silvae Sinicae,2013,49(10):155-161.
[9]邓惠,向甘驹,郭友好,杨春锋.秦岭忍冬属4种植物的繁育系统及花色变化的研究[J].植物科学学报,2017,35(1):1-12.Deng H,Xiang GJ,Guo YH,Yang CF.Study on the breeding system and floral color change of four Lonicera species in the Qinling mountains[J].Plant Science Journal,2017,35(1):1-12.
[10]Farzad M,Griesbach R,Weiss M.Floral color change in Viola cornuta:a model system to study regulation of anthocyanin production[J].Plant Sci,2002,162:225-231.
[11]杨琴,袁涛,孙湘滨.两个牡丹品种开花过程中花色变化的研究[J].园艺学报,2015,42(5):930-938.Yang Q,Yuan T,Sun XB.Preliminary studies on the changes of flower color during the flowering period in two tree peony cultivars[J].Acta Horticulture Sinica,2015,42(5):930-938.
[12]张宝智.江南牡丹花色素组成与开花过程生理特征研究[D].上海:上海交通大学,2013.
[13]钟培星,王亮生,李珊珊,徐彦军,朱满兰.芍药开花过程中花色和色素的变化[J].园艺学报,2012,39(11):2271-2282.Zhong PX,Wang LS,Li SS,Xu YJ,Zhu ML.The changes of floral color and pigments composition during the flowering period in Paeonia lactiflora petals[J].Acta Horticulture Sinica,2012,39(11):2271-2282.
[14]李玲,杜绍华,张超,高金锋,李彦慧.4种花色仙客来色素及相关生理生化研究[J].河北农业大学学报,2012,35(2):25-29.Li L,Du SH,Zhang C,Gao JF,Li YH.A study on the pigment content and other related physiological and biochemical indices in four color Cylamen cultivars[J].Journal of Agricultural University of Hebei,2012,35(2):25-29.
[15]朱满兰,王亮生,张会金,徐彦军,郑绪辰,王丽金.耐寒睡莲花瓣中花青素苷组成及其与花色的关系[J].植物学报,2012,47(5):437-453.Zhu ML,Wang LS,Zhang HJ,Xu YJ,Zheng XC,Wang LJ.Relationship between the composition of anthocyanins and flower color variation in hardy water lily(Nymphaea spp.)cultivars[J].Chinese Bulletin of Botany,2012,47(5):437-453.
[16]程怡,张云婷,王清明,马建伟,张勇.月季花发育过程中花色变化的生理生化研究[J].西北植物学报,2014,34(4):733-739.Cheng Y,Zhang YT,Wang QM,Ma JW,Zhang Y.Correlation analysis of flower color during flower development of Rosa hybrida[J].Acta Botanica Boreali-Occidentalia Sinica,2014,34(4):733-739.
[17]Yan J,Wang ML,Zhang L.Light induces petal color change in Quisqualis indica[J].Plant Divers,2018,40:28-34.
[18]于晓南,张启翔.观赏植物的花色素苷与花色[J].林业科学,2002,38(3):147-153.Yu XN,Zhang QX.Anthocyanin in ornamental plant and color express[J].Scientia Silvae Sinicae,2002,38(3):147-153.
[19]戴思兰,洪艳.基于花青素苷合成和呈色机理的观赏植物花色改良分子育种[J].中国农业科学,2016,49(3):529-542.Dai SL,Hong Y.Molecular breeding for flower colors modification on ornamental plants based on the mechanism of anthocyanins biosynthesis and coloration[J].Scientia Agricultural Sinica,2016,49(3):529-542.
[20]安田齐著.花色的生理生物化学[M].傅玉兰译.北京:中国林业出版社,1989.
[21]于国瑞.色彩构成[M].修订版.北京:清华大学出版社,2012.
[22]Vaknin H,Bar-Akiva A,Ovadia R,Nissim-Levi A,Forer I,et al.Active anthocyanin degradation in Brunfelsia calycina flowers[J].Planta,2005,222(1):19-26.
[23]陈小琴,郭志雄,林琳,王蓓,柯毅湧.鸳鸯茉莉不同发育时期POD同工酶体系优化[J].热带作物学报,2014,35(8):1510-1516.Chen XQ,Guo ZX,Lin L,Wang B,Ke YY.POD isozyme system optimization of Brunfelsia acuminata(Solanaceae)in different development stages[J].Chinese Journal of Tropical Crops,2014,35(8):1510-1516.
[24]Qiu J,Xue XF,Chen FD,Li CH,Bolat N,et al.Quality evaluation of snow lotus(Saussurea):Quantitative chemical analysis and antioxidant activity assessment[J].Plant Cell Rep,2010,29:1325-1337.
[25]李合生.植物生理生化试验原理与技术[M].北京:高等教育出版社,2003.
[26]Pua EC,Davey MR.Plant Developmental Biology-Biotechnological Perspectives[M].Berlin:Springer Heidelberg,2010.
[27]Murray RB,David HL,Kevin MD,Gayle BM,Deepa P,et al.Isolation and antisense suppression of flavonoid 3',5'-hydroxylase modifies flower pigments and color in cyclamen[J].BMC Plant Biol,2010,10:107-118.
[28]Ferrante A,Vernieri P,Tognoni F,Serra G.Changes in abscisic acid and flower pigments during floral senescence of Petunia[J].Biol Plant,2006,50:581-585.