依据理化性质分析牡丹花色形成的影响因素
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摘要
对5个不同花色的牡丹品种花瓣解剖结构、色素分布、理化性状及类黄酮合成相关基因的表达进行了观测与分析,结果表明:不同花色牡丹品种的表皮细胞形态存在差异,类黄酮和金属元素(Al和Fe)质量分数与牡丹花色值显著相关。类黄酮为主要影响因素,天竺葵素、矢车菊素和芍药素作为主要色素,槲皮素和木樨草素作为辅助色素,共同参与了红色系的形成;查尔酮、芹菜素、槲皮素和金圣草黄素影响了黄色花的形成。结合8个类黄酮生物合成相关结构基因的表达模式分析显示,CHI、DFR、ANS基因控制花青苷的合成,CHI、F3H、F3’H和FLS基因控制黄酮、黄酮醇以及查尔酮的合成,分别在红色及黄色品种中高表达。因此,牡丹不同花色的形成主要受相关基因表达以及类黄酮质量分数的调控,而金属元素(Al、Fe)和花瓣表皮细胞形态也能在一定程度上影响花色的呈现。
We observed and analyzed the anatomical structure,pigment distribution,physicochemical characteristics and the expression profiles of flavonoid biosynthetic genes in five differentially colored cultivars of tree peony.The results showed that the morphology of epidermal cells in differentially colored cultivars of tree peony is distinct,and the contents of flavonoid and metal elements(Al and Fe) were significantly correlated with the flower color indices.Pelargonidin-cyanidin-and peonidin-based anthocyanins primarily determined the red pigmentation,while quercetin and luteolin acted as copigments to affect the formation of red flower colors; Chalcone,apigenin,quercetin and chrysoeriol determined the formation of yellow flower color.Comprehensive analysis of expression patterns of eight flavonoid biosynthetic structural genes indicated that CHS,CHI,DFR,and ANS genes controlling the synthesis of anthocyanins,and F3 H,F3'H,and FLS genes regulating mainly the synthesis of flavonoids,flavonols,and chalcone,exhibited high expressly levels in red and yellow cultivars,respectively.Therefore,the formation of different flower colors of tree peony is mainly regulated by the expression of related genes and the contents of flavonoids.Furthermore,the accumulation of Al and Fe as well as morphology of petal epidermal cells can also affect the flower color to some extent.
We observed and analyzed the anatomical structure,pigment distribution,physicochemical characteristics and the expression profiles of flavonoid biosynthetic genes in five differentially colored cultivars of tree peony.The results showed that the morphology of epidermal cells in differentially colored cultivars of tree peony is distinct,and the contents of flavonoid and metal elements(Al and Fe) were significantly correlated with the flower color indices.Pelargonidin-cyanidin-and peonidin-based anthocyanins primarily determined the red pigmentation,while quercetin and luteolin acted as copigments to affect the formation of red flower colors; Chalcone,apigenin,quercetin and chrysoeriol determined the formation of yellow flower color.Comprehensive analysis of expression patterns of eight flavonoid biosynthetic structural genes indicated that CHS,CHI,DFR,and ANS genes controlling the synthesis of anthocyanins,and F3 H,F3'H,and FLS genes regulating mainly the synthesis of flavonoids,flavonols,and chalcone,exhibited high expressly levels in red and yellow cultivars,respectively.Therefore,the formation of different flower colors of tree peony is mainly regulated by the expression of related genes and the contents of flavonoids.Furthermore,the accumulation of Al and Fe as well as morphology of petal epidermal cells can also affect the flower color to some extent.
引文
[1] 赵云鹏,陈发棣,郭维明.观赏植物花色基因工程研究进展[J].植物学通报,2003,20(1):51-58.
[2] QI Y Y, LOU Q, LI H B, et al. Anatomical and biochemical studies of bicolored flower development in Muscari latifolium[J]. Protoplasma,2013,250(6):1273-1281.
[3] SHOJI K, MIKI N, NAKAJIMA N, et al. Perianth bottom-specific blue color development in Tulip cv. murasakizuisho requires ferric ions[J]. Plant Cell Physiol,2007,48(2):243-251.
[4] NODA K, GLOVER B J, LINSTEAD P, et al. Flower colour intensity depends on specialized cell shape controlled by a Myb-related transcription factor[J]. Nature,1994,369:661-664.
[5] VIGNOLINI S, MOYROUD E, HINGANT T, et al. The flower of Hibiscus trionumis both visibly and measurably iridescent[J]. New Phytologist,2015,205(1):97-101.
[6] YOSHIDA K, KONDO T, OKAZAKI Y, et al. Cause of blue petal colour[J]. Nature, 1995,373:291-291.
[7] ZHAO D Q, TAO J. Recent advances on the development and regulation of flower color in ornamental plants[J]. Frontiers in Plant Science,2015,6:1-13.
[8] 吕长平,陈海霞,陈建.外源GA3对非洲菊舌状花着色的影响[J].中国农学通报,2012,28(22):212-215.
[9] HE Q L, SHEN Y, WANG M, et al. Natural variation in petal color in Lycoris longituba revealed by anthocyanin components[J]. PLoS ONE,2011,6(8).doi:10.1371/journal.pone0020098.
[10] 张晶晶,王亮生,刘政安,等.牡丹花色研究进展[J].园艺学报,2006,33(6):1383-1388.
[11] 华梅,原晓龙,杨卫,等.HPLC分析6种不同花色滇牡丹花瓣中花青素和黄酮[J].西部林业科学,2017,46(6):40-45.
[12] SAKATA Y, AOKI N, SADANOBU T, et al. Petal coloration and pigmentation of tree peony bred and selected in Daikon Island (Shimane prefecture)[J]. Journal of the Japanese Society for Horticultural Science,2008,64(2):351-357.
[13] 周琳,王雁,任磊,等.牡丹二氢黄酮醇4-还原酶基因PsDFR1的克隆及表达分析[J].植物生理学报,2011,47(9):885-892.
[14] ZHOU L, WANG Y, PENG Z H. Molecular characterization and expression analysis of chalcone synthase gene during flower development in tree peony (Paeonia suffruticosa)[J]. African Journal of Biotechnology,2011,10(8):1275-1284.
[15] ZHOU L, WANG Y, REN L, et al. Overexpression of Ps-CHI1, a homologue of the chalcone isomerase gene from tree peony (Paeonia suffruticosa Pall.), reduces the intensity of flower pigmentation in transgenic tobacco[J]. Plant Cell, Tissue and Organ Culture,2014,116(3):285-295.
[16] SHI Q Q, ZHOU L, LI K, et al. Isolation and expression of PlbHLH3 transcription factor genes in Paeonia lutea[J]. Forest Research,2015,28(4):488-496.
[17] 樊金玲,朱文学,巩卫东,等.牡丹花色苷的热稳定性和降解动力学[J].应用化学,2010,27(2):231-236.
[18] HANBURY A, SERRA J. Mathematical morphology in the CIELAB space[J]. Image Anal Stereol,2002,21(3):201-206.
[19] ZHANG J, WANG L S, GAO J M, et al. Determination of anthocyanins and exploration of relationship between their composition and petal coloration in crape myrtle (Lagerstroemia hybrid)[J]. Journal of Integrative Plant Biology,2008,50(5):581-588.
[20] 裴仁济,陈小强,孙宁,等.非洲紫罗兰花瓣细胞pH值测定及F3’5’H基因片段克隆[J].天津农学院学报,2012,19(1):11-14.
[21] 赵昶灵,郭维明,陈俊愉.梅花花色色素种类和含量的初步研究[J].北京林业大学学报,2004,26(2):68-73.
[22] 白新祥,胡可,戴思兰,等.不同花色菊花品种花色素成分的初步分析[J].北京林业大学学报,2006,28(5):84-88.
[23] 莫玉楠,张鲁刚,王国芳.橙色大白菜类胡萝卜素提取与测定方法研究[J].西北农林科技大学学报(自然科学版),2014,42(3):206-214.
[24] YANG R Z, WEI X L, GAO F F. Simultaneous analysis of anthocyanins and flavonols in petals of lotus (Nelumbo) cultivars by high-performance liquid chromatography-photodiode array detection/electrospray ionization mass spectrometry[J]. Journal of Chromatography A,2009,1216(1):106-112.
[25] LI C H, DU H, WANG L S, et al. Flavonoid composition and antioxidant activity of tree peony (Paeonia section Moutan) yellow flowers[J]. Journal of Agricultural and Food Chemistry,2009,57(18):8496-8503.
[26] LIVAK K J, SCHMITTGEN T D. Analysis of relative gene expression data using real-time quantitative PCR and the 2(T)(-Delta C) method[J]. Methods,2001,25(4):402-408.
[27] LUO J R, DUAN J J, HUO D, et al. Transcriptomic analysis reveals transcription factors related to leaf anthocyanin biosynthesis[J]. Molecules,2017,22(12):2186.
[28] ZHAO D Q, WEI M R, LIU D, et al. Anatomical and biochemical analysis reveal the role of anthocyanins in flower coloration of herbaceous peony[J]. Plant Physiology and Biochemistry,2016,102:97-106.
[29] YOSHIDA K, TOYAMA-KATO Y, KAMEDA K, et al. Sepal color variation of Hydrangea macrophylla and vacuolar pH measured with a proton-selective microelectrode[J]. Plant and Cell Physiology,2003,44(3):262-268.
[30] YOSHIDA K, MIKI N, MOMONOI K, et al. Synchrony between flower opening and petal-color change from red to blue in morning glory, Ipomoea tricolor cv. Heavenly Blue[J]. Proceedings of the Japan Academy,2009,85(6):187-197.
[31] SHI Q Q, LI L, ZHOU L, et al. Morphological and biochemical studies of the yellow and purple-red petal pigmentation in Paeonia delavayi[J]. Hortscience,2018,53(8):1-7.
[32] WANG L S, SHIRAISHI A, HASHIMOTO F, et al. Analysis of petal anthocyanins to investigate flower coloration of Zhongyuan (Chinese) and Daikon Island (Japanese) tree peony cultivars[J]. Journal of Plant Research,2001,114:33-43.
[33] 周琳,王雁,律春燕,等.云南野生黄牡丹花色素成分的鉴定[J].东北林业大学学报,2011,39(8):52-54.
[34] NAKATSUKA T, NISHIHARA M, MISHIBA K, et al. Temporal expression of flavonoid biosynthesis-related genes regulates flower pigmentation in gentian plants[J]. Plant Science,2005,168(5):1309-1318.
[35] ZHAO D Q, TAO J, HAN C X, et al. Flower color diversity revealed by differential expression of flavonoid biosynthetic genes and flavonoid accumulation in herbaceous peony (Paeonia lactiflora Pall.)[J]. Molecular Biology Reports,2012,39(12):11263-11275.
[36] ZHAO D Q, TANG W H, HAO Z J, et al. Identification of flavonoids and expression of flavonoid biosynthetic genes in two coloured tree peony flowers[J]. Biochemical and Biophysical Research Communications,2015,459(3):450-456.
[37] ZHANG X P, ZHAO M Y, GUO J, et al. Anatomical and biochemical analyses reveal the mechanism of double-color formation in Paeonia suffruticosa ‘Shima Nishiki’[J]. 3 Biotech,2018,8:420.
[2] QI Y Y, LOU Q, LI H B, et al. Anatomical and biochemical studies of bicolored flower development in Muscari latifolium[J]. Protoplasma,2013,250(6):1273-1281.
[3] SHOJI K, MIKI N, NAKAJIMA N, et al. Perianth bottom-specific blue color development in Tulip cv. murasakizuisho requires ferric ions[J]. Plant Cell Physiol,2007,48(2):243-251.
[4] NODA K, GLOVER B J, LINSTEAD P, et al. Flower colour intensity depends on specialized cell shape controlled by a Myb-related transcription factor[J]. Nature,1994,369:661-664.
[5] VIGNOLINI S, MOYROUD E, HINGANT T, et al. The flower of Hibiscus trionumis both visibly and measurably iridescent[J]. New Phytologist,2015,205(1):97-101.
[6] YOSHIDA K, KONDO T, OKAZAKI Y, et al. Cause of blue petal colour[J]. Nature, 1995,373:291-291.
[7] ZHAO D Q, TAO J. Recent advances on the development and regulation of flower color in ornamental plants[J]. Frontiers in Plant Science,2015,6:1-13.
[8] 吕长平,陈海霞,陈建.外源GA3对非洲菊舌状花着色的影响[J].中国农学通报,2012,28(22):212-215.
[9] HE Q L, SHEN Y, WANG M, et al. Natural variation in petal color in Lycoris longituba revealed by anthocyanin components[J]. PLoS ONE,2011,6(8).doi:10.1371/journal.pone0020098.
[10] 张晶晶,王亮生,刘政安,等.牡丹花色研究进展[J].园艺学报,2006,33(6):1383-1388.
[11] 华梅,原晓龙,杨卫,等.HPLC分析6种不同花色滇牡丹花瓣中花青素和黄酮[J].西部林业科学,2017,46(6):40-45.
[12] SAKATA Y, AOKI N, SADANOBU T, et al. Petal coloration and pigmentation of tree peony bred and selected in Daikon Island (Shimane prefecture)[J]. Journal of the Japanese Society for Horticultural Science,2008,64(2):351-357.
[13] 周琳,王雁,任磊,等.牡丹二氢黄酮醇4-还原酶基因PsDFR1的克隆及表达分析[J].植物生理学报,2011,47(9):885-892.
[14] ZHOU L, WANG Y, PENG Z H. Molecular characterization and expression analysis of chalcone synthase gene during flower development in tree peony (Paeonia suffruticosa)[J]. African Journal of Biotechnology,2011,10(8):1275-1284.
[15] ZHOU L, WANG Y, REN L, et al. Overexpression of Ps-CHI1, a homologue of the chalcone isomerase gene from tree peony (Paeonia suffruticosa Pall.), reduces the intensity of flower pigmentation in transgenic tobacco[J]. Plant Cell, Tissue and Organ Culture,2014,116(3):285-295.
[16] SHI Q Q, ZHOU L, LI K, et al. Isolation and expression of PlbHLH3 transcription factor genes in Paeonia lutea[J]. Forest Research,2015,28(4):488-496.
[17] 樊金玲,朱文学,巩卫东,等.牡丹花色苷的热稳定性和降解动力学[J].应用化学,2010,27(2):231-236.
[18] HANBURY A, SERRA J. Mathematical morphology in the CIELAB space[J]. Image Anal Stereol,2002,21(3):201-206.
[19] ZHANG J, WANG L S, GAO J M, et al. Determination of anthocyanins and exploration of relationship between their composition and petal coloration in crape myrtle (Lagerstroemia hybrid)[J]. Journal of Integrative Plant Biology,2008,50(5):581-588.
[20] 裴仁济,陈小强,孙宁,等.非洲紫罗兰花瓣细胞pH值测定及F3’5’H基因片段克隆[J].天津农学院学报,2012,19(1):11-14.
[21] 赵昶灵,郭维明,陈俊愉.梅花花色色素种类和含量的初步研究[J].北京林业大学学报,2004,26(2):68-73.
[22] 白新祥,胡可,戴思兰,等.不同花色菊花品种花色素成分的初步分析[J].北京林业大学学报,2006,28(5):84-88.
[23] 莫玉楠,张鲁刚,王国芳.橙色大白菜类胡萝卜素提取与测定方法研究[J].西北农林科技大学学报(自然科学版),2014,42(3):206-214.
[24] YANG R Z, WEI X L, GAO F F. Simultaneous analysis of anthocyanins and flavonols in petals of lotus (Nelumbo) cultivars by high-performance liquid chromatography-photodiode array detection/electrospray ionization mass spectrometry[J]. Journal of Chromatography A,2009,1216(1):106-112.
[25] LI C H, DU H, WANG L S, et al. Flavonoid composition and antioxidant activity of tree peony (Paeonia section Moutan) yellow flowers[J]. Journal of Agricultural and Food Chemistry,2009,57(18):8496-8503.
[26] LIVAK K J, SCHMITTGEN T D. Analysis of relative gene expression data using real-time quantitative PCR and the 2(T)(-Delta C) method[J]. Methods,2001,25(4):402-408.
[27] LUO J R, DUAN J J, HUO D, et al. Transcriptomic analysis reveals transcription factors related to leaf anthocyanin biosynthesis[J]. Molecules,2017,22(12):2186.
[28] ZHAO D Q, WEI M R, LIU D, et al. Anatomical and biochemical analysis reveal the role of anthocyanins in flower coloration of herbaceous peony[J]. Plant Physiology and Biochemistry,2016,102:97-106.
[29] YOSHIDA K, TOYAMA-KATO Y, KAMEDA K, et al. Sepal color variation of Hydrangea macrophylla and vacuolar pH measured with a proton-selective microelectrode[J]. Plant and Cell Physiology,2003,44(3):262-268.
[30] YOSHIDA K, MIKI N, MOMONOI K, et al. Synchrony between flower opening and petal-color change from red to blue in morning glory, Ipomoea tricolor cv. Heavenly Blue[J]. Proceedings of the Japan Academy,2009,85(6):187-197.
[31] SHI Q Q, LI L, ZHOU L, et al. Morphological and biochemical studies of the yellow and purple-red petal pigmentation in Paeonia delavayi[J]. Hortscience,2018,53(8):1-7.
[32] WANG L S, SHIRAISHI A, HASHIMOTO F, et al. Analysis of petal anthocyanins to investigate flower coloration of Zhongyuan (Chinese) and Daikon Island (Japanese) tree peony cultivars[J]. Journal of Plant Research,2001,114:33-43.
[33] 周琳,王雁,律春燕,等.云南野生黄牡丹花色素成分的鉴定[J].东北林业大学学报,2011,39(8):52-54.
[34] NAKATSUKA T, NISHIHARA M, MISHIBA K, et al. Temporal expression of flavonoid biosynthesis-related genes regulates flower pigmentation in gentian plants[J]. Plant Science,2005,168(5):1309-1318.
[35] ZHAO D Q, TAO J, HAN C X, et al. Flower color diversity revealed by differential expression of flavonoid biosynthetic genes and flavonoid accumulation in herbaceous peony (Paeonia lactiflora Pall.)[J]. Molecular Biology Reports,2012,39(12):11263-11275.
[36] ZHAO D Q, TANG W H, HAO Z J, et al. Identification of flavonoids and expression of flavonoid biosynthetic genes in two coloured tree peony flowers[J]. Biochemical and Biophysical Research Communications,2015,459(3):450-456.
[37] ZHANG X P, ZHAO M Y, GUO J, et al. Anatomical and biochemical analyses reveal the mechanism of double-color formation in Paeonia suffruticosa ‘Shima Nishiki’[J]. 3 Biotech,2018,8:420.
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