Isolation and Expression of PlbHLH3 Transcription Factor Genes in Paeonia lutea

SHI Qian-qian; ZHOU Lin; LI Kui; WANG Yan

Volume 28 Issue 4

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Isolation and Expression of PlbHLH3 Transcription Factor Genes in Paeonia lutea

1.
Laboratory of Tree Breeding and Cultivation, State Forestry Administration, Research Institute of Forestry, Chinese Academy of Forestry, Beijing 100091, China

Received Date: 2014-08-29

Abstract

Twenty-four unigene sequences shared high homology with bHLH transcription factor protein involved in plant anthocyanin biosynthesis were obtained from previous-constructed tree peony (Paeonia lutea) petal transcriptome database and named as PlbHLH1~24. By amino acid sequence comparing and phylogenetic tree analysis, it showed that PlbHLH3 was considered to be related to regulate anthocyanin biosynthesis, contained a 2 040 bp ORF encoding 679 amino acid residues with typical bHLH structural domain; the predicted protein sequence of PlbHLH3 also shared high similarity with other bHLH transcription factor that related to anthocyanin biosynthesis such as VvMYC1 and MdbHLH3. Relative Real-Time PCR analysis indicated that PlbHLH3 expressed in different tissues of P. lutea and P. delavayi, the expression in carpel, anther and petal was significantly higher than that in sepal and leaf; PlbHLH3 reached the highest abundance at early stage of P. lutea,. For P. delavayi, the PlbHLH3 reached the lowest level at early stage, and was significantly or extremely significantly lower than that in other four stages. In conclusion, it is inferred that PlbHLH3 might associate with the regulatory of anthocyanin biosynthesis in P. lutea and this would provide the basis for insight into the molecular mechanisms underlying tree peony yellow flower pigmentation.
- Paeonia lutea,
- bHLH,
- anthocyanin biosynthesis,
- gene cloning,
- expression analysis

References

[1]	黄金霞,王亮生,李晓梅,等.花色变异的分子基础与进化模式研究进展[J].植物学通报,2006, 23(4): 321-333.
[2]	Holton T A, Cornish E C. Genetics and biochemistry of anthocyanin biosynthesis[J]. Plant Cell, 1995, 7(7): 1071-1083.
[3]	Ramsay N A, Glover B J. MYB-bHLH-WD40 protein complex and the evolution of cellular diversity[J]. Trends Plant Sci, 2005,10(2): 63-70.
[4]	张全琪,朱家红,倪燕妹,等.植物bHLH转录因子的结构特点及其生物学功能[J].热带亚热带植物学报, 2000,19(1):84-90.
[5]	Nesi N, Debeaujon I, Jond C, et al. The TT8 gene encodes a basic helix-loop-helix domain protein required for expression of DFR and BAN genes in Arabidopsis siliques[J]. Plant Cell, 2000, 12 (10): 1863-1878.
[6]	Heisler MG, Atkinson A, Bylstra YH, et al. SPATULA, a gene that controls development of carpel margin tissues in Arabidopsis, encodes a bHLH protein[J]. Development,2001,128:1089-1098.
[7]	Sorensen A M, Kröber S, Unte U S, et al. The Arabidopsis ABORTED MICROSPORES (AMS) gene encodes a MYC class transcription factor[J]. Plant J,2003, 33(2): 413-423.
[8]	Leivar P, Monte E, Oka Y, et al. Multiple phytochrome-interacting bHLH transcription factors repress premature seedling photomorphogenesis in darkness[J]. Curr Biol,2008, 18 (23): 1815-1823.
[9]	Abe H, Urao T, Ito T, et al. Arabidopsis AtMYC2 (bHLH) and AtMYB2 (MYB) function as transcriptional activators in abscisic acid signaling[J]. Plant Cell, 2003,15 (1): 63-78.
[10]	Lorenzo O, Chico JM, Sánchez-Serrano JJ, et al. JASMONATE-INSENSITIVE1 encodes a MYC transcription factor essential to discriminate between different jasmonate-regulated defense responses in Arabidopsis[J]. Plant Cell, 2004,16 (7): 1938-1950.
[11]	Li H, Sun J, Xu Y, et al. The bHLH-type transcription factor AtAIB positively regulates ABA response in Arabidopsis. Plant Mol Biol.2007, 65 (5): 655-665.
[12]	Bou-Torrent J, Roig-Villanova I, Galstyan A, et al. PAR1 and PAR2 integrate shade and hormone transcriptional networks[J]. Plant Signal Behav. 2008, 3 (7): 453-454.
[13]	Rampey R A, Woodward AW, et al. An Arabidopsis basic helix-loop-helix leucine zipper protein modulates metal homeostasis and auxin conjugate responsiveness[J]. Genetics. 2006,174 (4): 1841-1857.
[14]	Long TA, Tsukagoshi H, Busch W, et al. The bHLH transcription factor POPEYE regulates response to iron deficiency in Arabidopsis roots[J]. Plant Cell,2010,22 (7): 2219-2236.
[15]	Ludwig SR, Habera LF, Dellaporta SL, et al. Lc, a member of the maize R gene family responsible for tissuespecific anthocyanin production, encodes a protein similar to transcriptional activators and contains the myc-homology region[J]. Proc Natl Acad Sci USA, 1998,86 (18): 7092-7096.
[16]	ChandlerV L, Radieella J P, Robbins T P, et al. Two regulatory genes of the maize anthocyanin pathway are homologous:Isolation of B utilizing R genomic sequenees[J]. Plant Cell,1989,1(12):1175-1183.
[17]	Goff S A, Cone K C, Chandler V L. Functional analysis of the transcriptional activator encoded by the maize B gene: evidence for a direct functional interaction between two classes of regulatory proteins[J]. Genes Dev., 1992,6 (5): 864-875.
[18]	Goodrich J, Carpenter R and Coen E S. 1992. A common gene regulates pigmentation patten in diverse plant species. Cell, 68(5):955-964.
[19]	Elomaa P, Mehto M, Kotilainen M, et al. A bHLH transcription factor mediates organ, region and flower type specific signals on dihydrofl avonol-4-reductase (dfr) gene expression in the inflorescence of Gerbera hybrida (Asteraceae). Plant J., 1998,16(1):93-99.
[20]	Sakamoto W, Ohmori T, Kageyama K,et al. The purple leaf (Pl) locus of rice:The pl allele has a complex organization and includes two Genes encoding basie helix-loop-helix proteins involved in anthocyanin biosynthesis. Plant Cell Physiol, 2001, 42(9):982-991.
[21]	Park K, Ishikawa N, Morita Y, et al. A bHLH regulatory gene in the common morning glory, Ipomoea Purpurea, controls anthocyanin biosynthesis in flowers, proanthocyanidin and phytolnelanin pigmentation in seeds, and seed trichome formation.The Plant Joumal, 2007b, 49(4):641-654
[22]	Nakatsuka T, Haruta K S, Pitaksutheepong C, et al. Identification and characterization of R2R3-MYB and bHLH transcription factors regulating anthocyanin biosynthesis in gentian flowers[J]. Plant Cell Physiol. 2008, 49: 1818-1829.
[23]	Espley R V, Hellens R P, Putterill J, et al. Red colouration in apple fruit due to the activity of the MYB transcription factor, MdMYB10[J]. Plant J.,2007,49:414-427.
[24]	Franken P., Schrell S., Peterson P A., et al. Molecular analysis of protein domain function encoded by the myb-homologous maize genes CI, Zm1 and Zm 38[J]. The Plant Journal,1994,6 (1):21-30.
[25]	Spelt C, Quattrocchio F, Mol JNM, et al.Anthocyanin1 of petunia encodes a basic helix-loop-helix protein that directly activates transcription of structural anthocyanin genes[J]. Plant Cell, 2000,12 (9): 1619-1631.
[26]	Yamagishi M, Shimoyamada Y, Nakatsuka T, et al. Two R2R3-MYB genes, homologs of petunia AN2, regulate anthocyanin biosyntheses in flower tepals, tepal spots and leaves of Asiatic hybrid lily[J]. Plant Cell Physiol, 2010, 51:463-474.
[27]	Wang Kui-lin, Bolitho, K., Grafton, K., et al. An R2R3 MYB transcription factor associated with regulation of the anthocyanin biosynthetic pathway in Rosaceae[J]. BMC Plant Biology, 2010,10: 50.
[28]	王勇,陈克平,姚勤. bHLH转录因子家族研究进展[J].遗传,2008, 30 (7): 821-830.
[29]	方文培. 中国芍药属植物的研究[J]. 植物分类学报, 1958, 7(4): 297-323.
[30]	王莲英. 中国牡丹品种图志[M]. 北京: 中国林业出版社, 1997, 2-7.
[31]	李嘉珏. 中国牡丹与芍药[M]. 北京: 中国林业出版社, 1999.
[32]	龚询.滇牡丹复合群体分类研究[J]. 7 昆明植物研究所,1990.
[33]	王志芳, 王雁, 岳桦. 珍稀资源: 黄牡丹[J].中国城市林业,2007, 5(2): 59- 60.
[34]	Zhou L, Wang Y, Ren L, et al. Overexpression of Pl-CHI1, a homologue of the chalcone isomerase gene from tree peony (Paeonia suffruticosa), reduces the intensity of flower pigmentation in transgenic tobacco[J]. Plant Cell, Tissue and Organ Culture, 2014, 16 (3): 285-295.
[35]	Zhou L, Wang Y, Peng Z. Molecular characterization and expression analysis of chalcone synthase gene during flower development in tree peony (Paeonia suffruticosa)[J]. Afr J Biotechnol, 2011, 10(8):1275-1284.
[36]	Fan C, Purugganan M D, Thomas D T, et al. Heterogeneous evolution of the Myc-like Anthocyanin regulatory gene and its phylogenetic utility in Cornus L.(Cornaceae)[J]. Mol Phylogenet Evol,2004, 33(3): 580-594.
[37]	Jiang Y, Deyholos M K. Comprehensive transcriptional profiling of NaCl-stressed Arabidopsis roots reveals novel classes of responsive genes[J]. BMC Plant Biol, 2006, 6(1): 25.
[38]	Jiang Y, Yang B, Deyholos M K. Functional characterization of the Arabidopsis bHLH92 transcription factor in abiotic stress[J]. Mol Genet Genomics, 2009, 282(5): 503-516.
[39]	Li XX,Duan XP,Jiang HX, et al. Genome-wide analysis of basic/helix-loop-helix transcription factor family in rice and Arabidopsis[J]. Plant Physiology, 2006, 141(4):1167-1184.
[40]	Hichri I, Heppel S C, Pillet J, et al. The basic helix-loop-helix transcription factor MYC1 is involved in the regulation of the flavonoid biosynthesis pathway in grapevine[J]. Mol Plant, 2010, 3(3): 509-523.
[41]	Xie X B, Li S, Zhang R F, et al. The bHLH transcription factor MdbHLH3 promotes anthocyanin accumulation and fruit colouration in response to low temperature in apples[J]. Plant Cell Environ, 2012, 35(11): 1884-1897.
[42]	Quattrocchio F, Wing J F, Va K, et al. Analysis of bHLH and MYB domain proteins: species-specific regulatory differences are caused by divergent evolution of target anthocyanin genes[J]. Plant J, 1998,13(4): 475-488.
[43]	Martin C, Prescott A, Mackay S, et al. Control of anthocyanin biosynthesis in flowers of Antirrhinum majus. Plant J.1991,1 (1): 37-49.
[44]	Quattrocchio F,Wing J F,Leppen H,et al. Regulatory genes controlling anthocyanin pigmentation are functionally conserved among plant species and have distinct sets of target genes[J]. The Plant Cell, 1993, 5 (11): 1497-1512.
[45]	Quattrocchio F, Baudry A, Lepiniec L, et al. The regulation of flavonoid biosynthesis. In: Grotewold E(ed) The science of flavonoids[J]. New York, Springer, pp.2006: 97-122.
[46]	de Vetten N, Quattrocchio F, Mol J, et al. The an11 locus controlling flower pigmentation in petunia encodes a novel WD-repeat protein conserved in yeast, plants, and animals[J]. Genes & Development, 1997,11:1422-1434.
[47]	Elomaa P, Mehto M, Kotilainen M, et al. A bHLH transcription factor mediates organ, region and flower type specific signals on dihydrofl avonol-4-reductase (dfr) gene expression in the inflorescence of Gerbera hybrida (Asteraceae). Plant J., 1998,16(1):93-99.
[48]	Li C, Du H, Wang L, et al. Flavonoid composition and antioxidant activity of tree peony (Paeonia section Moutan) yellow flowers[J]. J Agric Food Chem, 2009, 57: 8496-8503.
[49]	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.
[50]	Wang LS, Hashimoto F, Shiraishi A, et al. Phenetics in tree peony species from China by flower pigment cluster analysis[J].J Plant Res, 2001a, 114: 213-221.
[51]	Wang LS, 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]. J Plant Res,2001b, 114:33-43.
[52]	韩科厅,赵莉,唐杏姣, 等.菊花花色素苷合成关键基因表达与花色表型的关系[J].园艺学报, 2012,39 (3):516-524.

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通讯作者: 陈斌, bchen63@163.com

1.
沈阳化工大学材料科学与工程学院沈阳 110142

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Isolation and Expression of PlbHLH3 Transcription Factor Genes in Paeonia lutea

1. Laboratory of Tree Breeding and Cultivation, State Forestry Administration, Research Institute of Forestry, Chinese Academy of Forestry, Beijing 100091, China

Received Date: 2014-08-29

Abstract: Twenty-four unigene sequences shared high homology with bHLH transcription factor protein involved in plant anthocyanin biosynthesis were obtained from previous-constructed tree peony (Paeonia lutea) petal transcriptome database and named as PlbHLH1~24. By amino acid sequence comparing and phylogenetic tree analysis, it showed that PlbHLH3 was considered to be related to regulate anthocyanin biosynthesis, contained a 2 040 bp ORF encoding 679 amino acid residues with typical bHLH structural domain; the predicted protein sequence of PlbHLH3 also shared high similarity with other bHLH transcription factor that related to anthocyanin biosynthesis such as VvMYC1 and MdbHLH3. Relative Real-Time PCR analysis indicated that PlbHLH3 expressed in different tissues of P. lutea and P. delavayi, the expression in carpel, anther and petal was significantly higher than that in sepal and leaf; PlbHLH3 reached the highest abundance at early stage of P. lutea,. For P. delavayi, the PlbHLH3 reached the lowest level at early stage, and was significantly or extremely significantly lower than that in other four stages. In conclusion, it is inferred that PlbHLH3 might associate with the regulatory of anthocyanin biosynthesis in P. lutea and this would provide the basis for insight into the molecular mechanisms underlying tree peony yellow flower pigmentation.

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Reference (52)

[1]	黄金霞,王亮生,李晓梅,等.花色变异的分子基础与进化模式研究进展[J].植物学通报,2006, 23(4): 321-333.
[2]	Holton T A, Cornish E C. Genetics and biochemistry of anthocyanin biosynthesis[J]. Plant Cell, 1995, 7(7): 1071-1083.
[3]	Ramsay N A, Glover B J. MYB-bHLH-WD40 protein complex and the evolution of cellular diversity[J]. Trends Plant Sci, 2005,10(2): 63-70.
[4]	张全琪,朱家红,倪燕妹,等.植物bHLH转录因子的结构特点及其生物学功能[J].热带亚热带植物学报, 2000,19(1):84-90.
[5]	Nesi N, Debeaujon I, Jond C, et al. The TT8 gene encodes a basic helix-loop-helix domain protein required for expression of DFR and BAN genes in Arabidopsis siliques[J]. Plant Cell, 2000, 12 (10): 1863-1878.
[6]	Heisler MG, Atkinson A, Bylstra YH, et al. SPATULA, a gene that controls development of carpel margin tissues in Arabidopsis, encodes a bHLH protein[J]. Development,2001,128:1089-1098.
[7]	Sorensen A M, Kröber S, Unte U S, et al. The Arabidopsis ABORTED MICROSPORES (AMS) gene encodes a MYC class transcription factor[J]. Plant J,2003, 33(2): 413-423.
[8]	Leivar P, Monte E, Oka Y, et al. Multiple phytochrome-interacting bHLH transcription factors repress premature seedling photomorphogenesis in darkness[J]. Curr Biol,2008, 18 (23): 1815-1823.
[9]	Abe H, Urao T, Ito T, et al. Arabidopsis AtMYC2 (bHLH) and AtMYB2 (MYB) function as transcriptional activators in abscisic acid signaling[J]. Plant Cell, 2003,15 (1): 63-78.
[10]	Lorenzo O, Chico JM, Sánchez-Serrano JJ, et al. JASMONATE-INSENSITIVE1 encodes a MYC transcription factor essential to discriminate between different jasmonate-regulated defense responses in Arabidopsis[J]. Plant Cell, 2004,16 (7): 1938-1950.
[11]	Li H, Sun J, Xu Y, et al. The bHLH-type transcription factor AtAIB positively regulates ABA response in Arabidopsis. Plant Mol Biol.2007, 65 (5): 655-665.
[12]	Bou-Torrent J, Roig-Villanova I, Galstyan A, et al. PAR1 and PAR2 integrate shade and hormone transcriptional networks[J]. Plant Signal Behav. 2008, 3 (7): 453-454.
[13]	Rampey R A, Woodward AW, et al. An Arabidopsis basic helix-loop-helix leucine zipper protein modulates metal homeostasis and auxin conjugate responsiveness[J]. Genetics. 2006,174 (4): 1841-1857.
[14]	Long TA, Tsukagoshi H, Busch W, et al. The bHLH transcription factor POPEYE regulates response to iron deficiency in Arabidopsis roots[J]. Plant Cell,2010,22 (7): 2219-2236.
[15]	Ludwig SR, Habera LF, Dellaporta SL, et al. Lc, a member of the maize R gene family responsible for tissuespecific anthocyanin production, encodes a protein similar to transcriptional activators and contains the myc-homology region[J]. Proc Natl Acad Sci USA, 1998,86 (18): 7092-7096.
[16]	ChandlerV L, Radieella J P, Robbins T P, et al. Two regulatory genes of the maize anthocyanin pathway are homologous:Isolation of B utilizing R genomic sequenees[J]. Plant Cell,1989,1(12):1175-1183.
[17]	Goff S A, Cone K C, Chandler V L. Functional analysis of the transcriptional activator encoded by the maize B gene: evidence for a direct functional interaction between two classes of regulatory proteins[J]. Genes Dev., 1992,6 (5): 864-875.
[18]	Goodrich J, Carpenter R and Coen E S. 1992. A common gene regulates pigmentation patten in diverse plant species. Cell, 68(5):955-964.
[19]	Elomaa P, Mehto M, Kotilainen M, et al. A bHLH transcription factor mediates organ, region and flower type specific signals on dihydrofl avonol-4-reductase (dfr) gene expression in the inflorescence of Gerbera hybrida (Asteraceae). Plant J., 1998,16(1):93-99.
[20]	Sakamoto W, Ohmori T, Kageyama K,et al. The purple leaf (Pl) locus of rice:The pl allele has a complex organization and includes two Genes encoding basie helix-loop-helix proteins involved in anthocyanin biosynthesis. Plant Cell Physiol, 2001, 42(9):982-991.
[21]	Park K, Ishikawa N, Morita Y, et al. A bHLH regulatory gene in the common morning glory, Ipomoea Purpurea, controls anthocyanin biosynthesis in flowers, proanthocyanidin and phytolnelanin pigmentation in seeds, and seed trichome formation.The Plant Joumal, 2007b, 49(4):641-654
[22]	Nakatsuka T, Haruta K S, Pitaksutheepong C, et al. Identification and characterization of R2R3-MYB and bHLH transcription factors regulating anthocyanin biosynthesis in gentian flowers[J]. Plant Cell Physiol. 2008, 49: 1818-1829.
[23]	Espley R V, Hellens R P, Putterill J, et al. Red colouration in apple fruit due to the activity of the MYB transcription factor, MdMYB10[J]. Plant J.,2007,49:414-427.
[24]	Franken P., Schrell S., Peterson P A., et al. Molecular analysis of protein domain function encoded by the myb-homologous maize genes CI, Zm1 and Zm 38[J]. The Plant Journal,1994,6 (1):21-30.
[25]	Spelt C, Quattrocchio F, Mol JNM, et al.Anthocyanin1 of petunia encodes a basic helix-loop-helix protein that directly activates transcription of structural anthocyanin genes[J]. Plant Cell, 2000,12 (9): 1619-1631.
[26]	Yamagishi M, Shimoyamada Y, Nakatsuka T, et al. Two R2R3-MYB genes, homologs of petunia AN2, regulate anthocyanin biosyntheses in flower tepals, tepal spots and leaves of Asiatic hybrid lily[J]. Plant Cell Physiol, 2010, 51:463-474.
[27]	Wang Kui-lin, Bolitho, K., Grafton, K., et al. An R2R3 MYB transcription factor associated with regulation of the anthocyanin biosynthetic pathway in Rosaceae[J]. BMC Plant Biology, 2010,10: 50.
[28]	王勇,陈克平,姚勤. bHLH转录因子家族研究进展[J].遗传,2008, 30 (7): 821-830.
[29]	方文培. 中国芍药属植物的研究[J]. 植物分类学报, 1958, 7(4): 297-323.
[30]	王莲英. 中国牡丹品种图志[M]. 北京: 中国林业出版社, 1997, 2-7.
[31]	李嘉珏. 中国牡丹与芍药[M]. 北京: 中国林业出版社, 1999.
[32]	龚询.滇牡丹复合群体分类研究[J]. 7 昆明植物研究所,1990.
[33]	王志芳, 王雁, 岳桦. 珍稀资源: 黄牡丹[J].中国城市林业,2007, 5(2): 59- 60.
[34]	Zhou L, Wang Y, Ren L, et al. Overexpression of Pl-CHI1, a homologue of the chalcone isomerase gene from tree peony (Paeonia suffruticosa), reduces the intensity of flower pigmentation in transgenic tobacco[J]. Plant Cell, Tissue and Organ Culture, 2014, 16 (3): 285-295.
[35]	Zhou L, Wang Y, Peng Z. Molecular characterization and expression analysis of chalcone synthase gene during flower development in tree peony (Paeonia suffruticosa)[J]. Afr J Biotechnol, 2011, 10(8):1275-1284.
[36]	Fan C, Purugganan M D, Thomas D T, et al. Heterogeneous evolution of the Myc-like Anthocyanin regulatory gene and its phylogenetic utility in Cornus L.(Cornaceae)[J]. Mol Phylogenet Evol,2004, 33(3): 580-594.
[37]	Jiang Y, Deyholos M K. Comprehensive transcriptional profiling of NaCl-stressed Arabidopsis roots reveals novel classes of responsive genes[J]. BMC Plant Biol, 2006, 6(1): 25.
[38]	Jiang Y, Yang B, Deyholos M K. Functional characterization of the Arabidopsis bHLH92 transcription factor in abiotic stress[J]. Mol Genet Genomics, 2009, 282(5): 503-516.
[39]	Li XX,Duan XP,Jiang HX, et al. Genome-wide analysis of basic/helix-loop-helix transcription factor family in rice and Arabidopsis[J]. Plant Physiology, 2006, 141(4):1167-1184.
[40]	Hichri I, Heppel S C, Pillet J, et al. The basic helix-loop-helix transcription factor MYC1 is involved in the regulation of the flavonoid biosynthesis pathway in grapevine[J]. Mol Plant, 2010, 3(3): 509-523.
[41]	Xie X B, Li S, Zhang R F, et al. The bHLH transcription factor MdbHLH3 promotes anthocyanin accumulation and fruit colouration in response to low temperature in apples[J]. Plant Cell Environ, 2012, 35(11): 1884-1897.
[42]	Quattrocchio F, Wing J F, Va K, et al. Analysis of bHLH and MYB domain proteins: species-specific regulatory differences are caused by divergent evolution of target anthocyanin genes[J]. Plant J, 1998,13(4): 475-488.
[43]	Martin C, Prescott A, Mackay S, et al. Control of anthocyanin biosynthesis in flowers of Antirrhinum majus. Plant J.1991,1 (1): 37-49.
[44]	Quattrocchio F,Wing J F,Leppen H,et al. Regulatory genes controlling anthocyanin pigmentation are functionally conserved among plant species and have distinct sets of target genes[J]. The Plant Cell, 1993, 5 (11): 1497-1512.
[45]	Quattrocchio F, Baudry A, Lepiniec L, et al. The regulation of flavonoid biosynthesis. In: Grotewold E(ed) The science of flavonoids[J]. New York, Springer, pp.2006: 97-122.
[46]	de Vetten N, Quattrocchio F, Mol J, et al. The an11 locus controlling flower pigmentation in petunia encodes a novel WD-repeat protein conserved in yeast, plants, and animals[J]. Genes & Development, 1997,11:1422-1434.
[47]	Elomaa P, Mehto M, Kotilainen M, et al. A bHLH transcription factor mediates organ, region and flower type specific signals on dihydrofl avonol-4-reductase (dfr) gene expression in the inflorescence of Gerbera hybrida (Asteraceae). Plant J., 1998,16(1):93-99.
[48]	Li C, Du H, Wang L, et al. Flavonoid composition and antioxidant activity of tree peony (Paeonia section Moutan) yellow flowers[J]. J Agric Food Chem, 2009, 57: 8496-8503.
[49]	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.
[50]	Wang LS, Hashimoto F, Shiraishi A, et al. Phenetics in tree peony species from China by flower pigment cluster analysis[J].J Plant Res, 2001a, 114: 213-221.
[51]	Wang LS, 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]. J Plant Res,2001b, 114:33-43.
[52]	韩科厅,赵莉,唐杏姣, 等.菊花花色素苷合成关键基因表达与花色表型的关系[J].园艺学报, 2012,39 (3):516-524.

Isolation and Expression of PlbHLH3 Transcription Factor Genes in Paeonia lutea

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通讯作者: 陈斌, bchen63@163.com

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Isolation and Expression of PlbHLH3 Transcription Factor Genes in Paeonia lutea

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