滇牡丹类黄酮2'-O-萄糖基转移酶基因的鉴定及表达分析
|
摘要
在被子植物中,2'-O-葡萄糖基转移酶能够影响花色色素分子的生物合成,在类黄酮上添加葡萄糖基配体,是形成黄色色素的关键酶。本研究采用RT-PCR技术首次从滇牡丹中克隆得到一个具完整开放阅读框的类黄酮-2'-O-葡萄糖基转移酶(Pd2'GT)基因,对其进行生物信息学分析和转录模式分析。研究发现该基因全长1 428 bp,可编码的蛋白氨基酸475个,该蛋白无信号肽和跨膜结构,是一种在细胞质基质中发挥功能的不稳定蛋白;核酸序列与葡萄(Vitis vinifera)糖基转移酶的核酸序列具78%的一致性,说明该基因可能编码一个新蛋白,其蛋白序列含糖基转移酶超家族结构域PSPG,其氨基酸残基序列为WAPQVAILSHRATGG FVSHCGWNSILESLWFGVPIAALPMYAEQQ,含典型的糖基转移酶识别区和UDP糖基配体绑定位点;该蛋白与苹果(Malus domestica,NP_001315903)、西洋梨(Pyrus communis,D3UAG1)、仙客来(Cyclamen persicum,BAF75895)和长春花(Catharanthus roseus,BAF75901)的2'GT聚为一类,该组的糖基受体均为查尔酮;转录模式分析表明:Pd2'GT基因在黄色花瓣中表达量最高,在花蕾期该基因的积累水平最高。本研究通过对滇牡丹转录组数据的分析,分离并克隆得到Pd2'GT基因,为通过基因工程手段培育具新性状的观赏牡丹提供研究依据。
In angiosperms, the 2'-O-glucosyltransferases(2'GT) can affect the biosynthesis of color pigment molecules, and adding glucosyl ligand to flavonoids is the key enzyme to form xanthophyl. The present study cloned a flavonoid 2'-O-glycosyltransferase(Pd2'GT) gene with complete ORF from Paeonia delavayi for the first time with the method of RT-PCR, and its bioinformatics and transcriptional pattern analysis were carried out. The results showed that the full length of Pd2'GT gene was 1 428 bp, encoding a protein of 475 amino acids. The protein had no signal peptide and transmembrane structure, which was an unstable protein functioning in cytoplasmic matrix.The identity between the nucleotide sequence of Pd2'GT and the glycosyltransferase gene of Vitis vinifera was78%, indicating that the Pd2'GT gene might encode a new protein. Its protein sequence contained glycosyltransferase superfamily domain PSPG, and its amino acid residue sequence was WAPQVAILSHRATGGFVSHCGWNSILESL WFG VPIAALPMYAEQQ, including the typical identifiable sites of glycosyltransferase and the binding sites ofUDP sugar moiety. This protein clustered together with the 2'GT of Malus domestica(NP_001315903), Pyrus communis(D3 UAG1), Cyclamen persicum(BAF75895), and Catharanthus roseus(BAF75901), and the sugar acceptor of this group was chalcone. The transcriptional pattern analysis clarified that the highest expression of Pd2'GT gene was in the yellow petals of P. delavyi, and the highest accumulation level of this gene was in the budding phase. In this study, the Pd2'GT gene was isolated and cloned through the analysis of the transcriptional data of Paeonia delavayi, which could provide research reference for the breeding of new ornamental peony by genetic engineering.
In angiosperms, the 2'-O-glucosyltransferases(2'GT) can affect the biosynthesis of color pigment molecules, and adding glucosyl ligand to flavonoids is the key enzyme to form xanthophyl. The present study cloned a flavonoid 2'-O-glycosyltransferase(Pd2'GT) gene with complete ORF from Paeonia delavayi for the first time with the method of RT-PCR, and its bioinformatics and transcriptional pattern analysis were carried out. The results showed that the full length of Pd2'GT gene was 1 428 bp, encoding a protein of 475 amino acids. The protein had no signal peptide and transmembrane structure, which was an unstable protein functioning in cytoplasmic matrix.The identity between the nucleotide sequence of Pd2'GT and the glycosyltransferase gene of Vitis vinifera was78%, indicating that the Pd2'GT gene might encode a new protein. Its protein sequence contained glycosyltransferase superfamily domain PSPG, and its amino acid residue sequence was WAPQVAILSHRATGGFVSHCGWNSILESL WFG VPIAALPMYAEQQ, including the typical identifiable sites of glycosyltransferase and the binding sites ofUDP sugar moiety. This protein clustered together with the 2'GT of Malus domestica(NP_001315903), Pyrus communis(D3 UAG1), Cyclamen persicum(BAF75895), and Catharanthus roseus(BAF75901), and the sugar acceptor of this group was chalcone. The transcriptional pattern analysis clarified that the highest expression of Pd2'GT gene was in the yellow petals of P. delavyi, and the highest accumulation level of this gene was in the budding phase. In this study, the Pd2'GT gene was isolated and cloned through the analysis of the transcriptional data of Paeonia delavayi, which could provide research reference for the breeding of new ornamental peony by genetic engineering.
引文
Altschul S.F.,Mad den T.L.,Schffer A.A.,Zhang J.,Zhang Z.Miller W.,and Lipman D.J.,1997,Gapped BLAST and PSI-BLAST:a new generation of protein database search programs,Nucleic Acids Res.,25(17):3389-3402
Asen S.,Stewart R.N.,and Norris K.H.,1972,Co-pigmentation of anthocyanins in plant tissues and its effect on color,Phytochemistry,11(3):1139-1144
Bairoch A.,1991,PROSITE:a dictionary of sites and patterns in proteins,Nucleic Acids Res.,19(S1):2241-2245
Cabrita L.,Fossen T.,and Andersen.M.,2000,Colour and stability of the six common anthocyanidin 3-glucosides in aqueous solutions,Food Chemistry,68(1):101-107
Fukuchi-Mizutani M.,Okuhara H.,Fukui Y.,Nakao M.,Katsumoto Y.,Yonekura-Sakakibara K.,Kusumi T.,Hase T.and Tanaka Y.,2003,Biochemical and molecular characterization of a novel UDP-glucose:anthocyanin 3'-O-glucosyltransferase,a key Enzyme for blue anthocyanin biosynthesis,from gentian,Plant Physiol.,132(3):1652-1663
Gachon C.M.M.,Langlois-Meurinne M.,Saindrenan P.,2005,Plant secondary metabolism glycosyltransferases:the emerging functional analysis,Trends Plant Sci.,10(11):542-549
Gong X.,Pan Y.Z.,and Yang Z.Y.,2003,The diversities and value of present situation of Paeonia delavayi,Xibei Zhiwu Xuebao(Acta Botanica Boreali-Occidentalia Sinica),23(2)218-223(龚洵,潘跃芝,杨志云,2003,滇牡丹的多样性和现状评估,西北植物学报,23(2):218-223)
Gosch C.,Halbwirth H.,Schneider B.,Holscher D.,and Stich K.2010,Cloning and heterologous expression of glycosyltransferases from Malus×domestica and Pyrus communis,which convert phloretin to phloretin 2'-O-glucoside(phloridzin),Plant Sci.,178(3):299-306
Grotewold E.,2006,The genetics and biochemistry of floral pigments,Annu.Rev.Plant Biol.,57(1):761-780
Hall D.,Yuan X.X.,Murata J.,and Luca V.D.,2012,Molecular cloning and biochemical characterization of the UDP-glucose:flavonoid 3-O-glucosyltransferase from concord grape(Vitis labrusca),Phytochemistry,74(3):90-99
Hong D.Y.,Pan K.Y.,and Yu H.,1998,Taxonomy of the Paeonia delavayi complex(Paeoniaceae),Ann.Mo.Bot.Gard.,85(4):554-564
Hughes J.,and Hughes M.A.,1994,Multiple secondary plant product UDP-glucose glucosyltransferase genes expressed in cassava(Manihot esculenta Crantz)cotyledons,DNA Seq.,5(1):41-49
Jia Z.D.,Ma P.Y.,Bian X.F.,Yang Q.,Guo X.D.,and Xie Y.Z.,2014,Biosynthesis metabolic pathway and molecular regulation of plants anthocyanin,Xibei Zhiwu Xuebao(Acta B-otanica Boreali-Occidentalia Sinica),34(7):1496-1506(贾赵东,马佩勇,边小峰,杨清,郭小丁,谢一芝,2014,植物花青素合成代谢途径及其分子调控,西北植物学报,34(7):1496-1506)
Kahn R.A.,Bak S.,Svendsen I.,Halkier B.A.,and Mller B.L.,1997,Isolation and reconstitution of cytochrome P450ox and in vitro reconstitution of the entire biosynthetic pathway of the cyanogenic glucoside dhurrin from Sorghum,Plant Physiol.,115(4):1661-1670
Kobayashi S.,Ishimaru M.,Ding C.K.,Yakushiji H.,and Goto N.,2001,Comparison of UDP-glucose:flavonoid 3-O-glucosyltransferase(UFGT)gene sequences between white grapes(Vitis vinifera)and their sports with red skin,Plant Sci.,160(3):543-550
Li C.,Du H.,Wang L.,Shu Q.,Zheng Y.,Xu Y.,Zhang J.,Zhang J.,Yang R.,and Ge Y.,2009,Flavonoid composition and antioxidant activity of tree peony(Paeonia section Moutan)yellow flowers,J.Agric.Food Chem.,57(18):8496-8503
Li X.,Liu Y.,Li H.Z.,Wang Z.Y.,and Chen H.Y.,2011,Cloning and expression characterization of the anthocyanin 5-O-glucosyltransferase in eggplant,Shanghai Jiaotong Daxue Xuebao(Journal of Shanghai Jiao Tong University),29(6):1-5(李翔,刘杨,李怀志,王中彦,陈火英,2011,茄花青素5-O糖基转移酶基因克隆与表达特征分析,上海交通大学学报,29(6):1-5)
Mol J.,Cornish E.,Mason J.,and Koes R.,1999,Novel coloured flowers,Curr.Opin.Biotechnol.,10(2):198-201
Ogata J.,Itoh Y.,Ishida M.,Yoshida H.,and Ozeki Y.,2004,Cloning and heterologous expression of c DNAs encoding flavonoid glucosyltransferases from Dianthus caryophyllus,Plant Biotechnology,21(5):367-375
Ren X.X.,Zhang Y.,Xue J.Q.,Zhu F.Y.,Shi F.R.,Wang S.L.,and Zhang X.X.,2015,Genetic diversity analysis of natural populations in Paeonia delavayi,Zhiwu Yichuan Ziyuan X-uebao(Journal of Plant Genetic Resources),16(4):772-780(任秀霞,张盈,薛璟祺,朱富勇,石丰瑞,王顺利,张秀新2015,滇牡丹天然居群的遗传多样性分析,植物遗传资源学报,16(4):772-780)
Shi Q.Q.,Zhou L.,Wang Y.,Li K.,Zheng B.Q.,and Miao K.2015,Transcriptomic analysis of Paeonia delavayi wild population flowers to identify differentially expressed genes involved in purple-red and yellow petal pigmentation,PLo SOne,10(8):e0135038
Tanaka Y.,Brugliera F.,and Chandler S.,2009,Recent progress of flower colour modification by biotechnology,Int.J.Mol Sci.,10(12):5350-5369
Tanaka Y.,Sasaki N.,and Ohmiya A.,2008,Biosynthesis of plant pigments:anthocyanins,betalains and carotenoids,Plant J.,54(4):733-749
Togami J.,Okuhara H.,Nakamura N.,Ishiguro K.,Hirose C.Ochiai M.,Fukui Y.,Yamaguchi M.A.,and Tanaka Y.2011,Isolation of c DNAs encoding tetrahydroxychalcone2'-glucosyltransferase activity from carnation,cyclamen and catharanthus,Plant Biotechnology,28(2):231-238
Vogt T.,and Jones P.,2000,Glycosyltransferases in plant natura product synthesis characterization of a supergene family,T-rends Plant Sci.,5(9):380-386
Wang W.Y.,Li H.M.,Dai Y.M.,Zou H.,Wu S.J.,Li Y.S.,and Lin J.B.,2015,Cloning and expression analysis of flavonoid 3-O-glucosyltransferase gene from Narcissus tazetta var.Chinensis,Fujian Nongye Xuebao(Fujian Journal of Agricultural Sciences),30(6):577-581(王伟英,李海明戴艺民,邹晖,吴水金,李跃森,林江波,2015,中国水仙类黄酮3-氧-葡糖基转移酶基因的克隆与序列分析福建农业学报,30(6):577-581)
Wang Y.,Wang C.C.,Zhou X.,Bi W.,Yang Y.M.,and Wang J.2015,Cloning and functional analysis of a flavonoid-3-O-glucosyltransferase gene from Indosasa hispida,Guangxi Zhiwu(Guihaia),35(2):244-249(王毅,王晨晨,周旭,毕玮,杨宇明,王娟,2015,七彩红竹中类黄酮-3-O-葡萄糖基转移酶基因的克隆及功能分析,广西植物,35(2)244-249)
Wang Y.,Zhao N.,Yuan X.L.,Chen Z.H.,Yang Y.M.,and Wang J.,2017,Identification and expression analysis of a flavonoid 3-O-glucosyltransferase gene in Paeonia delavayi,Jiyinzuxue Yu Yingyong Shengwuxue(Genomics and Applied Biology),36(4):1556-1562(王毅,赵能,原晓龙,陈中华,杨宇明,王娟,2017,滇牡丹类黄酮3-O-葡萄糖基转移酶基因的鉴定及表达分析,基因组学与应用生物学,36(4):1556-1562)
Yonekura-Sakakibara K.,2009,Functional genomics of family 1glycosyltransferases in Arabidopsis,Plant Biotechnology,26(3):267-274
Yoshida H.,Itoh Y.,Ozeki Y.,Iwashina T.,and Yamaguchi M.A.,2004,Variation in chalcononaringenin 2'-O-glucoside content in the petals of carnations(Dianthus caryophyllus)bearing yellow flowers,Sci.Hortic.,99(2):175-186
Zhang H.W.,Liang Y.,Liu X.Y.,and Tan W.P.,2015,Cloning and expression analysis of UFGT in onion,Henongxue Bao(Journal of Nuclear Agricultural Science),29(9):1677-1686(张洪伟,梁毅,刘小义,谭武平,2015,洋葱UFGT基因的克隆和表达分析,核农学报,29(9):1677-1686)
Zhang Y.L.,Li Z.H.,Ma H.,Wang Y.,Li W.J.,Liu X.X.,and Wan Y.M.,2011,A study on characters variation of different flower color groups of Paeonia delavayi(Paeoniaceae),Zhiwu Fenlei Yu Ziyuan Xuebao(Plant Diversity and Resources),33(2):229-236(张艳丽,李正红,马宏,王雁,李文娟,刘秀贤,万友名,2011,滇牡丹花色类群性状变异分析,植物分类与资源学报,33(2):229-236)
Zhao Y.P.,Chen F.D.,and Guo W.M.,2003 Advances in genetic engineering of flower color of ornamental plants,Zhiwuxue Tongbao(Chinese Bullein of Botany),20(1):51-58(赵云鹏,陈发棣,郭维明,2003,观赏植物花色基因工程研究进展,植物学通报,20(1):51-58)
Zhou L.,Wang Y.,Lv C.Y.,and Peng Z.H.,2011,Identification of components of flower pigments in petals of Paeonia lutea wild population in Yunnan,Dongbei Linye Daxue Xuebao(Journal of Northeast Forestry University),39(8):52-54(周琳,王雁,律春燕,彭镇华,2011,云南野生黄牡丹花色素成分的鉴定,东北林业大学学报,39(8):52-54)
Asen S.,Stewart R.N.,and Norris K.H.,1972,Co-pigmentation of anthocyanins in plant tissues and its effect on color,Phytochemistry,11(3):1139-1144
Bairoch A.,1991,PROSITE:a dictionary of sites and patterns in proteins,Nucleic Acids Res.,19(S1):2241-2245
Cabrita L.,Fossen T.,and Andersen.M.,2000,Colour and stability of the six common anthocyanidin 3-glucosides in aqueous solutions,Food Chemistry,68(1):101-107
Fukuchi-Mizutani M.,Okuhara H.,Fukui Y.,Nakao M.,Katsumoto Y.,Yonekura-Sakakibara K.,Kusumi T.,Hase T.and Tanaka Y.,2003,Biochemical and molecular characterization of a novel UDP-glucose:anthocyanin 3'-O-glucosyltransferase,a key Enzyme for blue anthocyanin biosynthesis,from gentian,Plant Physiol.,132(3):1652-1663
Gachon C.M.M.,Langlois-Meurinne M.,Saindrenan P.,2005,Plant secondary metabolism glycosyltransferases:the emerging functional analysis,Trends Plant Sci.,10(11):542-549
Gong X.,Pan Y.Z.,and Yang Z.Y.,2003,The diversities and value of present situation of Paeonia delavayi,Xibei Zhiwu Xuebao(Acta Botanica Boreali-Occidentalia Sinica),23(2)218-223(龚洵,潘跃芝,杨志云,2003,滇牡丹的多样性和现状评估,西北植物学报,23(2):218-223)
Gosch C.,Halbwirth H.,Schneider B.,Holscher D.,and Stich K.2010,Cloning and heterologous expression of glycosyltransferases from Malus×domestica and Pyrus communis,which convert phloretin to phloretin 2'-O-glucoside(phloridzin),Plant Sci.,178(3):299-306
Grotewold E.,2006,The genetics and biochemistry of floral pigments,Annu.Rev.Plant Biol.,57(1):761-780
Hall D.,Yuan X.X.,Murata J.,and Luca V.D.,2012,Molecular cloning and biochemical characterization of the UDP-glucose:flavonoid 3-O-glucosyltransferase from concord grape(Vitis labrusca),Phytochemistry,74(3):90-99
Hong D.Y.,Pan K.Y.,and Yu H.,1998,Taxonomy of the Paeonia delavayi complex(Paeoniaceae),Ann.Mo.Bot.Gard.,85(4):554-564
Hughes J.,and Hughes M.A.,1994,Multiple secondary plant product UDP-glucose glucosyltransferase genes expressed in cassava(Manihot esculenta Crantz)cotyledons,DNA Seq.,5(1):41-49
Jia Z.D.,Ma P.Y.,Bian X.F.,Yang Q.,Guo X.D.,and Xie Y.Z.,2014,Biosynthesis metabolic pathway and molecular regulation of plants anthocyanin,Xibei Zhiwu Xuebao(Acta B-otanica Boreali-Occidentalia Sinica),34(7):1496-1506(贾赵东,马佩勇,边小峰,杨清,郭小丁,谢一芝,2014,植物花青素合成代谢途径及其分子调控,西北植物学报,34(7):1496-1506)
Kahn R.A.,Bak S.,Svendsen I.,Halkier B.A.,and Mller B.L.,1997,Isolation and reconstitution of cytochrome P450ox and in vitro reconstitution of the entire biosynthetic pathway of the cyanogenic glucoside dhurrin from Sorghum,Plant Physiol.,115(4):1661-1670
Kobayashi S.,Ishimaru M.,Ding C.K.,Yakushiji H.,and Goto N.,2001,Comparison of UDP-glucose:flavonoid 3-O-glucosyltransferase(UFGT)gene sequences between white grapes(Vitis vinifera)and their sports with red skin,Plant Sci.,160(3):543-550
Li C.,Du H.,Wang L.,Shu Q.,Zheng Y.,Xu Y.,Zhang J.,Zhang J.,Yang R.,and Ge Y.,2009,Flavonoid composition and antioxidant activity of tree peony(Paeonia section Moutan)yellow flowers,J.Agric.Food Chem.,57(18):8496-8503
Li X.,Liu Y.,Li H.Z.,Wang Z.Y.,and Chen H.Y.,2011,Cloning and expression characterization of the anthocyanin 5-O-glucosyltransferase in eggplant,Shanghai Jiaotong Daxue Xuebao(Journal of Shanghai Jiao Tong University),29(6):1-5(李翔,刘杨,李怀志,王中彦,陈火英,2011,茄花青素5-O糖基转移酶基因克隆与表达特征分析,上海交通大学学报,29(6):1-5)
Mol J.,Cornish E.,Mason J.,and Koes R.,1999,Novel coloured flowers,Curr.Opin.Biotechnol.,10(2):198-201
Ogata J.,Itoh Y.,Ishida M.,Yoshida H.,and Ozeki Y.,2004,Cloning and heterologous expression of c DNAs encoding flavonoid glucosyltransferases from Dianthus caryophyllus,Plant Biotechnology,21(5):367-375
Ren X.X.,Zhang Y.,Xue J.Q.,Zhu F.Y.,Shi F.R.,Wang S.L.,and Zhang X.X.,2015,Genetic diversity analysis of natural populations in Paeonia delavayi,Zhiwu Yichuan Ziyuan X-uebao(Journal of Plant Genetic Resources),16(4):772-780(任秀霞,张盈,薛璟祺,朱富勇,石丰瑞,王顺利,张秀新2015,滇牡丹天然居群的遗传多样性分析,植物遗传资源学报,16(4):772-780)
Shi Q.Q.,Zhou L.,Wang Y.,Li K.,Zheng B.Q.,and Miao K.2015,Transcriptomic analysis of Paeonia delavayi wild population flowers to identify differentially expressed genes involved in purple-red and yellow petal pigmentation,PLo SOne,10(8):e0135038
Tanaka Y.,Brugliera F.,and Chandler S.,2009,Recent progress of flower colour modification by biotechnology,Int.J.Mol Sci.,10(12):5350-5369
Tanaka Y.,Sasaki N.,and Ohmiya A.,2008,Biosynthesis of plant pigments:anthocyanins,betalains and carotenoids,Plant J.,54(4):733-749
Togami J.,Okuhara H.,Nakamura N.,Ishiguro K.,Hirose C.Ochiai M.,Fukui Y.,Yamaguchi M.A.,and Tanaka Y.2011,Isolation of c DNAs encoding tetrahydroxychalcone2'-glucosyltransferase activity from carnation,cyclamen and catharanthus,Plant Biotechnology,28(2):231-238
Vogt T.,and Jones P.,2000,Glycosyltransferases in plant natura product synthesis characterization of a supergene family,T-rends Plant Sci.,5(9):380-386
Wang W.Y.,Li H.M.,Dai Y.M.,Zou H.,Wu S.J.,Li Y.S.,and Lin J.B.,2015,Cloning and expression analysis of flavonoid 3-O-glucosyltransferase gene from Narcissus tazetta var.Chinensis,Fujian Nongye Xuebao(Fujian Journal of Agricultural Sciences),30(6):577-581(王伟英,李海明戴艺民,邹晖,吴水金,李跃森,林江波,2015,中国水仙类黄酮3-氧-葡糖基转移酶基因的克隆与序列分析福建农业学报,30(6):577-581)
Wang Y.,Wang C.C.,Zhou X.,Bi W.,Yang Y.M.,and Wang J.2015,Cloning and functional analysis of a flavonoid-3-O-glucosyltransferase gene from Indosasa hispida,Guangxi Zhiwu(Guihaia),35(2):244-249(王毅,王晨晨,周旭,毕玮,杨宇明,王娟,2015,七彩红竹中类黄酮-3-O-葡萄糖基转移酶基因的克隆及功能分析,广西植物,35(2)244-249)
Wang Y.,Zhao N.,Yuan X.L.,Chen Z.H.,Yang Y.M.,and Wang J.,2017,Identification and expression analysis of a flavonoid 3-O-glucosyltransferase gene in Paeonia delavayi,Jiyinzuxue Yu Yingyong Shengwuxue(Genomics and Applied Biology),36(4):1556-1562(王毅,赵能,原晓龙,陈中华,杨宇明,王娟,2017,滇牡丹类黄酮3-O-葡萄糖基转移酶基因的鉴定及表达分析,基因组学与应用生物学,36(4):1556-1562)
Yonekura-Sakakibara K.,2009,Functional genomics of family 1glycosyltransferases in Arabidopsis,Plant Biotechnology,26(3):267-274
Yoshida H.,Itoh Y.,Ozeki Y.,Iwashina T.,and Yamaguchi M.A.,2004,Variation in chalcononaringenin 2'-O-glucoside content in the petals of carnations(Dianthus caryophyllus)bearing yellow flowers,Sci.Hortic.,99(2):175-186
Zhang H.W.,Liang Y.,Liu X.Y.,and Tan W.P.,2015,Cloning and expression analysis of UFGT in onion,Henongxue Bao(Journal of Nuclear Agricultural Science),29(9):1677-1686(张洪伟,梁毅,刘小义,谭武平,2015,洋葱UFGT基因的克隆和表达分析,核农学报,29(9):1677-1686)
Zhang Y.L.,Li Z.H.,Ma H.,Wang Y.,Li W.J.,Liu X.X.,and Wan Y.M.,2011,A study on characters variation of different flower color groups of Paeonia delavayi(Paeoniaceae),Zhiwu Fenlei Yu Ziyuan Xuebao(Plant Diversity and Resources),33(2):229-236(张艳丽,李正红,马宏,王雁,李文娟,刘秀贤,万友名,2011,滇牡丹花色类群性状变异分析,植物分类与资源学报,33(2):229-236)
Zhao Y.P.,Chen F.D.,and Guo W.M.,2003 Advances in genetic engineering of flower color of ornamental plants,Zhiwuxue Tongbao(Chinese Bullein of Botany),20(1):51-58(赵云鹏,陈发棣,郭维明,2003,观赏植物花色基因工程研究进展,植物学通报,20(1):51-58)
Zhou L.,Wang Y.,Lv C.Y.,and Peng Z.H.,2011,Identification of components of flower pigments in petals of Paeonia lutea wild population in Yunnan,Dongbei Linye Daxue Xuebao(Journal of Northeast Forestry University),39(8):52-54(周琳,王雁,律春燕,彭镇华,2011,云南野生黄牡丹花色素成分的鉴定,东北林业大学学报,39(8):52-54)