观赏海棠不同叶位色素组分动态研究
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摘要
以57个观赏海棠品种为研究对象,对上位叶(Upper leaf)、中位叶(Middle leaf)、下位叶(Lower leaf)的色素组分进行测定,研究了海棠品种不同叶位间色素组分关系以及动态规律,为海棠优良观叶种质的筛选以及叶色改良提供理论参考。基于紫外—可见分光光度计的色素绝对含量测定和基于Unispec-SC光谱分析仪的色素相对含量测定,分别构建了花青素(Anth)、类胡萝卜素(Car)、叶绿素(Chl)3大色素组分绝对含量与相对含量之间的拟合函数:y_(Anth)=0.522 1x~2-0.111 4x+0.013 3(R~2=0.979 3);y_(Car)=0.456 2x+0.072 6(R~2=0.980 9);y_(Chl)=1.428 7x-0.672 6(R~2=0.901 6)。采用Origin 9.0软件构建了57个海棠品种的3大色素组分含量及权重的三维分布图。结果表明:1)基于紫外—可见分光光度计的色素绝对含量测定和基于Unispec-SC光谱分析仪的色素相对含量测定,分别构建了花青素(Anth)、类胡萝卜素(Car)、叶绿素(Chl)3大色素组分绝对含量与相对含量之间的拟合函数。2)基于上位叶叶绿素含量、花青素含量及其权重的聚类分析表明:57个海棠品种可分为2大色系4个子色系类群,即绿色系(含深绿色系、褐绿色系)、红色系(含紫红色系、红棕色系),各色系间差异显著。3)绿色系之A_1子色系含有较低的花青素含量,且随着叶位的下降,其含量变化平稳,绿色呈现效果较好。4)A_2子色系在上位叶阶段虽含有一定量的花青素,但不足以使叶片呈现出"全红"状态,因此绿色表达效果不如A_1子色系。5)红色系之B_1子色系的花青素含量及稳定性皆高于B_2子色系,在上位叶、中位叶,B_1子色系的花青素含量分别为B_2子色系的2.1倍和2.6倍,显著提高了呈色效果。6)此外,在中位叶,B_1子色系的花青素含量虽较上位叶有所淡化,但仍达到了B_2子色系在上位叶的含量水平。
The pigment components of upper leaf,middle leaf and lower leaf were determined by 57 ornamental crabapple species.The pigment components of different leaf positions of crabapple were studied systematically.The relationship and dynamic laws provide a theoretical reference for screening fine leafy germplasm and improving leaf color of crabapple species.Based on the determination of absolute content of pigments by ultraviolet-visible spectrophotometer and the determination of relative content of pigments by UnispecSC spectrometer,the fitting functions between absolute and relative contents of anthocyanins(Anth),carotenoids(Car)and chlorophyll(Chl)were constructed as follows:y_(Anth)=0.522 1x~2-0.111 4x+0.013(R~2=0.979);y_(Car)=0.456 2x+0.072 6(R~2=0.980 9 9);y_(Chl)=1.428 7x-0.672 6(R~2=0.901 6).Using Origin 9.0 software,the three-dimensional distribution maps of the content and weight of the three major pigments in 57 cultivars of Begonia crabapple were constructed.The results show that 1)Fitting functions between absolute and relative contents of anthocyanins(Anth),carotenoids(Car)and chlorophyll(Chl)were constructed based on the determination of absolute content of pigments by ultraviolet-visible spectrophotometer and the determination of relative content of pigments by Unispec-SC spectrometer;2)Cluster analysis based on chlorophyll content,anthocyanin content and weight of the upper leaves revealed that 57 cultivars could be divided into two big color lines and four sub-color groups,namely green line(dark green line,brown green line),red line(purple red line,red brown line).There were significant differences among different color lines;3)The green A_1 sub-color system contained a lower anthocyanin content,and as the leaf position decreased,its content changed smoothly,and the green color was better;4)Although A_2sub-color system contained a certain amount of anthocyanin in the upper leaf stage,it was not enough to make the leaves show a"red"state,so the green expression effect was inferior to A_1 seed color system;5)The anthocyanin content and stability of B1sub-color system in red line were higher than those of B_2 seed color system,in upper and middle leaves,the anthocyanin content of B_1 seed color system was 2.1 times and 2.6 times of that of B_2 sub-color system respectively,which significantly improved the color rendering effect;6)In addition,in the middle lobe,the anthocyanin content of the B_1 sub-color system was lighter than that of the upper leaf,but it still reached the level of the B_2 sub-color in the upper leaf.
The pigment components of upper leaf,middle leaf and lower leaf were determined by 57 ornamental crabapple species.The pigment components of different leaf positions of crabapple were studied systematically.The relationship and dynamic laws provide a theoretical reference for screening fine leafy germplasm and improving leaf color of crabapple species.Based on the determination of absolute content of pigments by ultraviolet-visible spectrophotometer and the determination of relative content of pigments by UnispecSC spectrometer,the fitting functions between absolute and relative contents of anthocyanins(Anth),carotenoids(Car)and chlorophyll(Chl)were constructed as follows:y_(Anth)=0.522 1x~2-0.111 4x+0.013(R~2=0.979);y_(Car)=0.456 2x+0.072 6(R~2=0.980 9 9);y_(Chl)=1.428 7x-0.672 6(R~2=0.901 6).Using Origin 9.0 software,the three-dimensional distribution maps of the content and weight of the three major pigments in 57 cultivars of Begonia crabapple were constructed.The results show that 1)Fitting functions between absolute and relative contents of anthocyanins(Anth),carotenoids(Car)and chlorophyll(Chl)were constructed based on the determination of absolute content of pigments by ultraviolet-visible spectrophotometer and the determination of relative content of pigments by Unispec-SC spectrometer;2)Cluster analysis based on chlorophyll content,anthocyanin content and weight of the upper leaves revealed that 57 cultivars could be divided into two big color lines and four sub-color groups,namely green line(dark green line,brown green line),red line(purple red line,red brown line).There were significant differences among different color lines;3)The green A_1 sub-color system contained a lower anthocyanin content,and as the leaf position decreased,its content changed smoothly,and the green color was better;4)Although A_2sub-color system contained a certain amount of anthocyanin in the upper leaf stage,it was not enough to make the leaves show a"red"state,so the green expression effect was inferior to A_1 seed color system;5)The anthocyanin content and stability of B1sub-color system in red line were higher than those of B_2 seed color system,in upper and middle leaves,the anthocyanin content of B_1 seed color system was 2.1 times and 2.6 times of that of B_2 sub-color system respectively,which significantly improved the color rendering effect;6)In addition,in the middle lobe,the anthocyanin content of the B_1 sub-color system was lighter than that of the upper leaf,but it still reached the level of the B_2 sub-color in the upper leaf.
引文
[1]姜卫兵,庄猛,韩浩章,等.彩叶植物呈色机理及光合特性研究进展[J].园艺学报,2005,32(2):352-358.
[2]SCHMITZER V,OSTERC G,VEBERIC R,et al.Correlation between chromaticity values and major anthocyanins in seven Acer palmatum Thunb.cultivars.[J].Scientia Horticulturae,2009,119(4):442-446.
[3]王美玲,艾希珍,郑楠,等.紫甘蓝不同叶位叶片色素含量及需光特性的研究[J].西北农业学报,2008,17(1):221-225.
[4]朱书香,杨建民,王中华,等.4种李属彩叶植物色素含量与叶色参数的关系[J].西北植物学报,2009,29(8):1663-1669.
[5]张潇,汪莹.彩叶植物呈色机理与影响花色素苷因素研究进展[J].四川林业科技,2015,36(1):28-30.
[6]姜文龙,范俊俊,张丹丹,等.观赏海棠不同叶位色彩特征及特异种质挖掘[J].园艺学报,2017,44(6):1135-1144.
[7]DANIEL A SIMS,JOHN A GAMON.Relationships between leaf pigment content and spectral reflectance across a wide range of species,leaf structures and developmental stages[J].Remote Sensing of Environment,2002,81(2):337-354.
[8]张往祥,裘靓,江志华,等.观赏海棠品种群开花进程中色素组分动态特征研究[J].园艺学报,2014,41(6):1145-1156.
[9]彭涛,李鹏民,贾裕娇,等.介绍两种无损伤测定植物活体叶片色素含量的方法[J].植物生理学报,2006,42(1):83-86.
[10]LETTS M G,JOHNSON D R E,COBURN C A.Drought stress ecophysiology of shrub and grass functional groups on opposing slope aspects of a temperate grassland valley[J].Botany-botanique,2010,88(9):850-866.
[11]宋艳冬,金爱武,金晓春,等.毛竹叶片光合色素的光谱估算模型[J].北京林业大学学报,2010,32(1):31-34.
[12]申晓慧,姜成,张敬涛,等.不同氮肥水平下大豆叶片光谱反射率与叶绿素含量的相关性研究[J].大豆科学,2012,31(1):73-75.
[13]DANIEL A SIMS,JOHN A GAMON.Relationships between leaf pigment content and spectral reflectance across a wide range of species,leaf structures and developmental stages[J].Remote Sensing of Environment,2002,81(2):337-354.
[14]MERZLYAK M N,SOLOVCHENKO A E,GITELSON A A.Reflectance spectral features and non-destructive estimation of chlorophyll,carotenoid and anthocyanin content in apple fruit[J].Postharvest Biology&Technology,2003,27(2):197-211.
[15]GOULD K S,MARKHAM K R,SMITH R H,et al.Functional role of anthocyanins in the leaves of Quintinia serrata A.Cunn[J].Journal of Experimental Botany,2000,51(347):1107-1115.
[16]莫巍.彩叶树种叶片中花青素含量的测定及动态分析[J].新疆农业科学,2007,44(B06):138-140.
[17]张宪政.植物叶绿素含量测定-丙酮乙醇混合液法[J].辽宁农业科学,1986(3):28-30.
[18]刘晓芳,李萍,黄闽敏,等.红宝石海棠叶片花青素及叶绿素含量的影响因子[J].经济林研究,2007,25(1):38-41.
[19]袁明,万兴智,杜蕾,等.红花檵木叶色变化机理的初步研究[J].园艺学报,2010,37(6):949-956.
[20]薛晓丽.紫叶李叶中花青素含量的测定[J].湖北农业科学,2010,49(4):956-958.
[21]王亚芸,王立英,任建武,等.金叶榆不同叶位叶片呈色生理机制研究[J].中国农学通报,2014,30(16):22-29.
[22]陈延惠,李跃霞,郭晓丽,等.园艺植物叶色变化机制研究进展[J].河南农业科学,2011,40(12):30-34.
[23]张丹丹,范俊俊,王欢,等.不同观赏海棠种质果色动态变化规律研究[J].经济林研究,2017,35(3):161-167.
[24]MOL J,GROTEWOLD E,KOES R.How gens paint flowers and seeds[J].Trends in Plant Science,1998,6(6):212-217.
[25]JOSEPHMOL,GARETHJENKIN S,E B E R H A R DSCH?¤FER,et al.Signal perception,transduction,and gene expression involved in anthocyanin biosynthesis[J].Critical Reviews in Plant Sciences,1996,15(5-6):525-557.
[26]NICOLA A.RAMSAY,BEVERLEY J.Glover.MYB-bHLH-WD40 protein complex and the evolution of cellular diversity.[J].Trends in Plant Science,2005,10(2):63-70.
[27]ALBERT N W,LEWIS D H,ZHANG H,et al.Light-induced vegetative anthocyanin pigmentation in Petunia[J].Journal of Experimental Botany,2009,60(7):2191-2202.
[28]STEYN W J,WAND S J E,JACOBS G,et al.Evidence for a photoprotective function of low-temperature-induced anthocyanin accumulation in apple and pear peel[J].Physiologia Plantarum,2010,136(4):461-472.
[29]WANG H,CHENG L.Differential Effects of Nitrogen Supply On Skin Pigmentation and Flesh Starch Breakdown of Gala’Apple[C]//Ashs Conference.2011.
[30]CASTELLARIN S D,MATTHEWS M A,GASPERO G D,et al.Water deficits accelerate ripening and induce changes in gene expression regulating flavonoid biosynthesis in grape berries[J].Planta,2007,227(1):101-112.
[31]陆秀君,葛根塔娜,梅梅,等.N、P、K配比施肥对美国红枫幼苗生长及叶色变化的影响[J].中南林业科技大学学报,2015,35(5):9-15.
[32]高艳.低氮胁迫诱导海棠花花青素积累及其基因与代谢分析[D].咸阳:西北农林科技大学,2014.
[33]Shvarts M,Borochov A,Weiss D.Low temperature enhances petunia flower pigmentation and induces chalcone synthase gene expression[J].Physiologia Plantarum,2010,99(1):67-72.
[34]安欣,王亚芸,任建武,等.环境因素对彩叶植物色素含量的相关影响[J].中国园艺文摘,2013(11):19-21.
[2]SCHMITZER V,OSTERC G,VEBERIC R,et al.Correlation between chromaticity values and major anthocyanins in seven Acer palmatum Thunb.cultivars.[J].Scientia Horticulturae,2009,119(4):442-446.
[3]王美玲,艾希珍,郑楠,等.紫甘蓝不同叶位叶片色素含量及需光特性的研究[J].西北农业学报,2008,17(1):221-225.
[4]朱书香,杨建民,王中华,等.4种李属彩叶植物色素含量与叶色参数的关系[J].西北植物学报,2009,29(8):1663-1669.
[5]张潇,汪莹.彩叶植物呈色机理与影响花色素苷因素研究进展[J].四川林业科技,2015,36(1):28-30.
[6]姜文龙,范俊俊,张丹丹,等.观赏海棠不同叶位色彩特征及特异种质挖掘[J].园艺学报,2017,44(6):1135-1144.
[7]DANIEL A SIMS,JOHN A GAMON.Relationships between leaf pigment content and spectral reflectance across a wide range of species,leaf structures and developmental stages[J].Remote Sensing of Environment,2002,81(2):337-354.
[8]张往祥,裘靓,江志华,等.观赏海棠品种群开花进程中色素组分动态特征研究[J].园艺学报,2014,41(6):1145-1156.
[9]彭涛,李鹏民,贾裕娇,等.介绍两种无损伤测定植物活体叶片色素含量的方法[J].植物生理学报,2006,42(1):83-86.
[10]LETTS M G,JOHNSON D R E,COBURN C A.Drought stress ecophysiology of shrub and grass functional groups on opposing slope aspects of a temperate grassland valley[J].Botany-botanique,2010,88(9):850-866.
[11]宋艳冬,金爱武,金晓春,等.毛竹叶片光合色素的光谱估算模型[J].北京林业大学学报,2010,32(1):31-34.
[12]申晓慧,姜成,张敬涛,等.不同氮肥水平下大豆叶片光谱反射率与叶绿素含量的相关性研究[J].大豆科学,2012,31(1):73-75.
[13]DANIEL A SIMS,JOHN A GAMON.Relationships between leaf pigment content and spectral reflectance across a wide range of species,leaf structures and developmental stages[J].Remote Sensing of Environment,2002,81(2):337-354.
[14]MERZLYAK M N,SOLOVCHENKO A E,GITELSON A A.Reflectance spectral features and non-destructive estimation of chlorophyll,carotenoid and anthocyanin content in apple fruit[J].Postharvest Biology&Technology,2003,27(2):197-211.
[15]GOULD K S,MARKHAM K R,SMITH R H,et al.Functional role of anthocyanins in the leaves of Quintinia serrata A.Cunn[J].Journal of Experimental Botany,2000,51(347):1107-1115.
[16]莫巍.彩叶树种叶片中花青素含量的测定及动态分析[J].新疆农业科学,2007,44(B06):138-140.
[17]张宪政.植物叶绿素含量测定-丙酮乙醇混合液法[J].辽宁农业科学,1986(3):28-30.
[18]刘晓芳,李萍,黄闽敏,等.红宝石海棠叶片花青素及叶绿素含量的影响因子[J].经济林研究,2007,25(1):38-41.
[19]袁明,万兴智,杜蕾,等.红花檵木叶色变化机理的初步研究[J].园艺学报,2010,37(6):949-956.
[20]薛晓丽.紫叶李叶中花青素含量的测定[J].湖北农业科学,2010,49(4):956-958.
[21]王亚芸,王立英,任建武,等.金叶榆不同叶位叶片呈色生理机制研究[J].中国农学通报,2014,30(16):22-29.
[22]陈延惠,李跃霞,郭晓丽,等.园艺植物叶色变化机制研究进展[J].河南农业科学,2011,40(12):30-34.
[23]张丹丹,范俊俊,王欢,等.不同观赏海棠种质果色动态变化规律研究[J].经济林研究,2017,35(3):161-167.
[24]MOL J,GROTEWOLD E,KOES R.How gens paint flowers and seeds[J].Trends in Plant Science,1998,6(6):212-217.
[25]JOSEPHMOL,GARETHJENKIN S,E B E R H A R DSCH?¤FER,et al.Signal perception,transduction,and gene expression involved in anthocyanin biosynthesis[J].Critical Reviews in Plant Sciences,1996,15(5-6):525-557.
[26]NICOLA A.RAMSAY,BEVERLEY J.Glover.MYB-bHLH-WD40 protein complex and the evolution of cellular diversity.[J].Trends in Plant Science,2005,10(2):63-70.
[27]ALBERT N W,LEWIS D H,ZHANG H,et al.Light-induced vegetative anthocyanin pigmentation in Petunia[J].Journal of Experimental Botany,2009,60(7):2191-2202.
[28]STEYN W J,WAND S J E,JACOBS G,et al.Evidence for a photoprotective function of low-temperature-induced anthocyanin accumulation in apple and pear peel[J].Physiologia Plantarum,2010,136(4):461-472.
[29]WANG H,CHENG L.Differential Effects of Nitrogen Supply On Skin Pigmentation and Flesh Starch Breakdown of Gala’Apple[C]//Ashs Conference.2011.
[30]CASTELLARIN S D,MATTHEWS M A,GASPERO G D,et al.Water deficits accelerate ripening and induce changes in gene expression regulating flavonoid biosynthesis in grape berries[J].Planta,2007,227(1):101-112.
[31]陆秀君,葛根塔娜,梅梅,等.N、P、K配比施肥对美国红枫幼苗生长及叶色变化的影响[J].中南林业科技大学学报,2015,35(5):9-15.
[32]高艳.低氮胁迫诱导海棠花花青素积累及其基因与代谢分析[D].咸阳:西北农林科技大学,2014.
[33]Shvarts M,Borochov A,Weiss D.Low temperature enhances petunia flower pigmentation and induces chalcone synthase gene expression[J].Physiologia Plantarum,2010,99(1):67-72.
[34]安欣,王亚芸,任建武,等.环境因素对彩叶植物色素含量的相关影响[J].中国园艺文摘,2013(11):19-21.