花青素苷合成途径中结构基因的表达对菊花和瓜叶菊花色的影响
|
摘要
菊花(Chrysanthemum x morifolium Ramat.)是中国传统名花,其花色变异丰富,但独缺蓝色系；瓜叶菊(Senecio cruentus Masson ex L'Herit; Pericallis cruenta L'Herit)是菊科千里光属广泛栽培的观赏植物,具有典型的蓝色系。本研究通过对比菊花和瓜叶菊花青素苷生物合成途径上关键结构基因的表达差异,探讨菊花蓝色系缺失的原因,分析花发育过程中蓝色花形成的分子生物学机理,对于开展花色改良的分子育种具有重要的理论意义和实际应用价值。此外,本研究通过探讨结构基因表达和花色表现之间的关系,期望为花色基因工程育种提供更多的基因资源和参考依据。本研究主要取得了如下成果：
1.花色表现是由色素组成决定的。将瓜叶菊品种‘春潮’和菊花品种‘丽金’分为4个色系,初步测定瓜叶菊白色系中仅含有黄酮类化合物；瓜叶菊红色系、紫色系、蓝色系和菊花粉色系中含有黄酮类化合物和花青素苷；菊花黄色系中含有黄酮类化合物、花青素苷和类胡萝卜素；
2.分离得到了与花青素苷合成途径中相关的关键结构基因。利用RT-PCR的方法分离了菊花花青素苷合成途径中关键结构基因的cDNA全长：CHI、F3H, F3'H和ANS,以及瓜叶菊ANS cDNA全长和4个结构基因片段：CHS、CHI、F3H、F3'H。对这些基因进行的生物信息学分析显示：上述基因编码的氨基酸序列与其它物种同源基因相比一致性较高；在氨基酸序列上具有重要的酶活性位点和保守的功能结构域；推测菊花CHI和F3H是亲水性较强的蛋白,而菊花F3'H、ANS和瓜叶菊ANS是跨膜蛋白。
3.采用半定量RT-PCR的方法分析CHS, CHI, F3H, F3'H, DFR、F3’5’H、3MaT这7个结构基因在瓜叶菊不同色系以及舌状花不同发育阶段的表达模式。结果表明：在白色花中,不存在CHS的表达；在红色花中,检测到F3’H基因的高丰度表达,但没有检测到F3’5’H的转录本；在蓝色花中,没有F3’H表达的信号,但在花序开放的第Ⅰ阶段有较强的F3’5’H的表达信号；而在紫色花中,可以同时检测到F3’H和F3’5’H的转录本。此外，瓜叶菊花青素苷合成相关结构基因在花序开放初期高丰度表达,随后逐渐降低,在第Ⅳ阶段表达量再次出现升高现象,而在花序开放末期表达量极低或没有表达信号。
4.对菊花品种‘丽金’不同色系舌状花中关键结构基因的表达模式分析表明,结构基因的表达量伴随着花序的开放呈现先上升后下降的趋势,大部分基因的表达高峰期出现在花序发育的第Ⅱ或第Ⅲ阶段；在舌状花开放末期,几乎检测不到基因表达的信号。F3’H和ANS在紫色花的第Ⅰ阶段表达量最高,而CHS、F3H和DFR在第Ⅱ或第Ⅲ阶段高丰度表达；结构基因在粉色系和红色系舌状花中的表达模式类似,但在粉色花中,F3’H的平均表达量低于红色花和紫色花。
5.关键结构基因在瓜叶菊和菊花不同组织器官中的表达结果表明,根中检测不到任何基因的转录本,除了DFR和ANS是花器官特异表达的基因外,F3’5’H在瓜叶菊舌状花和叶片中表达,而其余基因则在茎、叶中均有表达。
6.黑暗和高温处理菊花红色系、紫色系品种发现,舌状花不能正常呈色或严重褪色。基因表达分析表明,黑暗处理后,调节基因MYB和结构基因CHI、F3H, F3’H、DFR、ANS的表达受到抑制,WD40和CHS的表达丰度减弱,推测MYB是调控结构基因表达的重要转录因子；在菊花花青素苷合成相关结构基因的启动子上,可能存在光响应元件,可以与MYB相结合。高温会抑制菊花花序发育中、后期阶段的结构基因表达,但对发育前期基因的表达影响较弱,推测高温影响了菊花花青素苷合成相关基因的表达及花青素苷的稳定性；但高温处理后,调节基因MYB和WD40的表达量均有所上升,推测转录因子除了与菊花花色形成相关外,还可能与菊花的抗逆性有关。
7.对比瓜叶菊和菊花花青素苷呈色的差异,以及结构基因的表达模式,可得到以下结论：(1)基因表达是花色表现的根本原因；(2)菊花缺少蓝色系的主要原因是其体内没有F3’5’H基因的表达,且与之竞争底物的F3’H基因表达量较强,因此F3’H和F3’5’H是决定蓝色花能否形成的两个关键基因；(3)光照和温度与菊花舌状花呈色密切相关。
Chrysanthemum (Chrysanthemum×morifolium Ramat.) is one of the traditional cut flowers and pot flowers in China. Blue-flowering is the only type absent from colorful chrysanthemum varieties. Cineraria (Senecio cruentus Masson ex L'Herit; Pericallis cruentia L'Herit), which has blue varieties, is a popular potted flower in spring available at local markets in China. In this study, the difference on gene expression patterns involved in anthocyanin biosynthesis between cineraria and chrysanthemum was compared. It is helpful to explain the reasons and the molecular mechanism of the lack of blue-flowering chrysanthemum. These kinds of studies have important academic significances and practical values. There are 7 major results in this study.
(1) According to phenotype of flower coloration, chrysanthemum‘Regan’and cineraria‘Chunchao’were divided into 4 color series. In white ray florets of cineraria, flavonoids were major pigment. There were flavonoids and anthocyanins in cineraria red, purple and blue ray florets, and in chrysanthemum pink ray florets. Carotenoids were detected in yellow ray florets of chrysanthemum.
(2) Using semi-quantitative reverse transcriptase polymerase chain reaction (RT-PCR), this study isolated four internal segments and one full-length cDNA from cineraria flowers: chalcone synthase(CHS), chalcone isomerase (CHI), flavanone 3-hydroxylase (F3H), flavonoid 3'-hydroxylase (F3'H) and anthocyanidin synthase (ANS). In addition, four full-length cDNAs from chrysanthemum flowers were isolated by RT-PCR and PCR-RACE: chalcone isomerase (CHI), flavanone 3-hydroxylase (F3H), flavonoid 3'-hydroxylase (F3'H) and anthocyanidin synthase (ANS). The results of bioinformation suggested that key structual genes shared about 60%-93% homology with the accessed genes from other species in GenBank. And the putative protein contained conserved domains. CHI and F3H from chrysanthemum were considered hydrophilic proteins, F3'H and ANS may be membrane proteins.
(3) The expression patterns of CHS, CHI, F3H, F3'H, DFR, flavonoid 3',5'-hydroxylase (F3'5'H) and anthocyanin 3-O-glucoside-6"-O -malonyltransferase (3MaT) in five developmental stages of cineraria with different flower colors were determined for the first time by RT-PCR. The results indicated that there is no CHS transcript in white flowers, In red flowers, F3'H mRNA was expressed at high level. But no F3'5'H transcript was found. F3'5'H was highly expressed in Stage I but no F3'H mRNA was observed in blue flowers. And the transcripts of F3'H and F3'5'H were found in purple petals. In addition, these genes were expressed at high level in early stage and underwent moderate decreases in expression. Interestingly, genes expression level increased again in Stage IV. However, in Stage V, there is little expression level could be detected in petals.
(4) During seven developmental stages of chrysanthemum, most of genes were highly expressed in Stage II or Stage III and underwent moderate decreases. Little expression level could be detected in ray floret in Stage VII. CmF3'H mRNA were expressed at high level in Stage I in purple flowers. Average levels of CmF3'H expression in pink ray floret are lower than in red and purple ray florets.
(5) Expression of the structral genes in different tissues showed that they were not detected in roots. DFR and ANS were floral organ-specific genes. F3'5'H transcripts was detected in ray florets and leaves. Other genes expressed both in stems and leaves.
(6) After dark and high temperature treatment, chrysanthemum ray florets faded seriously. When inflorescences from Stage I were covered with aluminium foil, the gene expression of MYB and the expression of CHI, F3H, F3'H, DFR, ANS were inhibited. This results implyed that MYB was a key transcriptional factor regulating structural genes expression. However, MYB and WD40 were up-regulated after high temperature treatment, whereas the expression of most structural genes were inhibited in middle and late stages. It can be speculated that high temperature influenced the stability of anthocyanin. Furthermore, MYB and WD40 might be related to stress of chrysanthemum.
(7) According to our results, it could be concluded that gene expression was the underlying causes of flower pigmentation. Besides, there are two key reasons of the lack of blue-flowering chrysanthemum:(1) F3'5'H gene deletion; (2) F3'H strongly expressed. Light and temperature were closely related to chrysanthemum ray florets pigmentation.
1.花色表现是由色素组成决定的。将瓜叶菊品种‘春潮’和菊花品种‘丽金’分为4个色系,初步测定瓜叶菊白色系中仅含有黄酮类化合物；瓜叶菊红色系、紫色系、蓝色系和菊花粉色系中含有黄酮类化合物和花青素苷；菊花黄色系中含有黄酮类化合物、花青素苷和类胡萝卜素；
2.分离得到了与花青素苷合成途径中相关的关键结构基因。利用RT-PCR的方法分离了菊花花青素苷合成途径中关键结构基因的cDNA全长：CHI、F3H, F3'H和ANS,以及瓜叶菊ANS cDNA全长和4个结构基因片段：CHS、CHI、F3H、F3'H。对这些基因进行的生物信息学分析显示：上述基因编码的氨基酸序列与其它物种同源基因相比一致性较高；在氨基酸序列上具有重要的酶活性位点和保守的功能结构域；推测菊花CHI和F3H是亲水性较强的蛋白,而菊花F3'H、ANS和瓜叶菊ANS是跨膜蛋白。
3.采用半定量RT-PCR的方法分析CHS, CHI, F3H, F3'H, DFR、F3’5’H、3MaT这7个结构基因在瓜叶菊不同色系以及舌状花不同发育阶段的表达模式。结果表明：在白色花中,不存在CHS的表达；在红色花中,检测到F3’H基因的高丰度表达,但没有检测到F3’5’H的转录本；在蓝色花中,没有F3’H表达的信号,但在花序开放的第Ⅰ阶段有较强的F3’5’H的表达信号；而在紫色花中,可以同时检测到F3’H和F3’5’H的转录本。此外，瓜叶菊花青素苷合成相关结构基因在花序开放初期高丰度表达,随后逐渐降低,在第Ⅳ阶段表达量再次出现升高现象,而在花序开放末期表达量极低或没有表达信号。
4.对菊花品种‘丽金’不同色系舌状花中关键结构基因的表达模式分析表明,结构基因的表达量伴随着花序的开放呈现先上升后下降的趋势,大部分基因的表达高峰期出现在花序发育的第Ⅱ或第Ⅲ阶段；在舌状花开放末期,几乎检测不到基因表达的信号。F3’H和ANS在紫色花的第Ⅰ阶段表达量最高,而CHS、F3H和DFR在第Ⅱ或第Ⅲ阶段高丰度表达；结构基因在粉色系和红色系舌状花中的表达模式类似,但在粉色花中,F3’H的平均表达量低于红色花和紫色花。
5.关键结构基因在瓜叶菊和菊花不同组织器官中的表达结果表明,根中检测不到任何基因的转录本,除了DFR和ANS是花器官特异表达的基因外,F3’5’H在瓜叶菊舌状花和叶片中表达,而其余基因则在茎、叶中均有表达。
6.黑暗和高温处理菊花红色系、紫色系品种发现,舌状花不能正常呈色或严重褪色。基因表达分析表明,黑暗处理后,调节基因MYB和结构基因CHI、F3H, F3’H、DFR、ANS的表达受到抑制,WD40和CHS的表达丰度减弱,推测MYB是调控结构基因表达的重要转录因子；在菊花花青素苷合成相关结构基因的启动子上,可能存在光响应元件,可以与MYB相结合。高温会抑制菊花花序发育中、后期阶段的结构基因表达,但对发育前期基因的表达影响较弱,推测高温影响了菊花花青素苷合成相关基因的表达及花青素苷的稳定性；但高温处理后,调节基因MYB和WD40的表达量均有所上升,推测转录因子除了与菊花花色形成相关外,还可能与菊花的抗逆性有关。
7.对比瓜叶菊和菊花花青素苷呈色的差异,以及结构基因的表达模式,可得到以下结论：(1)基因表达是花色表现的根本原因；(2)菊花缺少蓝色系的主要原因是其体内没有F3’5’H基因的表达,且与之竞争底物的F3’H基因表达量较强,因此F3’H和F3’5’H是决定蓝色花能否形成的两个关键基因；(3)光照和温度与菊花舌状花呈色密切相关。
Chrysanthemum (Chrysanthemum×morifolium Ramat.) is one of the traditional cut flowers and pot flowers in China. Blue-flowering is the only type absent from colorful chrysanthemum varieties. Cineraria (Senecio cruentus Masson ex L'Herit; Pericallis cruentia L'Herit), which has blue varieties, is a popular potted flower in spring available at local markets in China. In this study, the difference on gene expression patterns involved in anthocyanin biosynthesis between cineraria and chrysanthemum was compared. It is helpful to explain the reasons and the molecular mechanism of the lack of blue-flowering chrysanthemum. These kinds of studies have important academic significances and practical values. There are 7 major results in this study.
(1) According to phenotype of flower coloration, chrysanthemum‘Regan’and cineraria‘Chunchao’were divided into 4 color series. In white ray florets of cineraria, flavonoids were major pigment. There were flavonoids and anthocyanins in cineraria red, purple and blue ray florets, and in chrysanthemum pink ray florets. Carotenoids were detected in yellow ray florets of chrysanthemum.
(2) Using semi-quantitative reverse transcriptase polymerase chain reaction (RT-PCR), this study isolated four internal segments and one full-length cDNA from cineraria flowers: chalcone synthase(CHS), chalcone isomerase (CHI), flavanone 3-hydroxylase (F3H), flavonoid 3'-hydroxylase (F3'H) and anthocyanidin synthase (ANS). In addition, four full-length cDNAs from chrysanthemum flowers were isolated by RT-PCR and PCR-RACE: chalcone isomerase (CHI), flavanone 3-hydroxylase (F3H), flavonoid 3'-hydroxylase (F3'H) and anthocyanidin synthase (ANS). The results of bioinformation suggested that key structual genes shared about 60%-93% homology with the accessed genes from other species in GenBank. And the putative protein contained conserved domains. CHI and F3H from chrysanthemum were considered hydrophilic proteins, F3'H and ANS may be membrane proteins.
(3) The expression patterns of CHS, CHI, F3H, F3'H, DFR, flavonoid 3',5'-hydroxylase (F3'5'H) and anthocyanin 3-O-glucoside-6"-O -malonyltransferase (3MaT) in five developmental stages of cineraria with different flower colors were determined for the first time by RT-PCR. The results indicated that there is no CHS transcript in white flowers, In red flowers, F3'H mRNA was expressed at high level. But no F3'5'H transcript was found. F3'5'H was highly expressed in Stage I but no F3'H mRNA was observed in blue flowers. And the transcripts of F3'H and F3'5'H were found in purple petals. In addition, these genes were expressed at high level in early stage and underwent moderate decreases in expression. Interestingly, genes expression level increased again in Stage IV. However, in Stage V, there is little expression level could be detected in petals.
(4) During seven developmental stages of chrysanthemum, most of genes were highly expressed in Stage II or Stage III and underwent moderate decreases. Little expression level could be detected in ray floret in Stage VII. CmF3'H mRNA were expressed at high level in Stage I in purple flowers. Average levels of CmF3'H expression in pink ray floret are lower than in red and purple ray florets.
(5) Expression of the structral genes in different tissues showed that they were not detected in roots. DFR and ANS were floral organ-specific genes. F3'5'H transcripts was detected in ray florets and leaves. Other genes expressed both in stems and leaves.
(6) After dark and high temperature treatment, chrysanthemum ray florets faded seriously. When inflorescences from Stage I were covered with aluminium foil, the gene expression of MYB and the expression of CHI, F3H, F3'H, DFR, ANS were inhibited. This results implyed that MYB was a key transcriptional factor regulating structural genes expression. However, MYB and WD40 were up-regulated after high temperature treatment, whereas the expression of most structural genes were inhibited in middle and late stages. It can be speculated that high temperature influenced the stability of anthocyanin. Furthermore, MYB and WD40 might be related to stress of chrysanthemum.
(7) According to our results, it could be concluded that gene expression was the underlying causes of flower pigmentation. Besides, there are two key reasons of the lack of blue-flowering chrysanthemum:(1) F3'5'H gene deletion; (2) F3'H strongly expressed. Light and temperature were closely related to chrysanthemum ray florets pigmentation.
引文
1.白新祥.菊花花色形成的表型分析[M].北京林业大学博士论文,2007.
2.白新祥,戴思兰.蓝色花观赏植物及其在园林中的应用.张启翔.中国观赏园艺研究进展2004[M].北京：中国林业出版社,2004：69-75.
3.戴思兰.中国菊花与世界园艺[J].河北科技师范学院学报,2004,18(2)：1-5,9.
4.韩科厅,胡可,戴思兰.观赏植物花色的分子设计[J].分子植物育种,2008,6(1)：16-24.
5.胡可,韩科厅,戴思兰.环境因子调控植物花青素合成及呈色的机理[J].植物学报,2010,45(3)：307-317.
6.胡可,孟丽,韩科厅,孙翊,戴思兰.瓜叶菊花青素合成关键结构基因的分离及表达分析[J].园艺学报,2009,36(7)：1013-1022.
7.黄金霞,王亮生,李晓梅,鲁迎青.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.
8.冷欣夫,邱星辉主编.细胞色素P450酶系的结构,功能与应用前景[M].北京：科学出版社,2001：107-128.
9.李鸿渐,邵建文.中国菊花品种资源的调查收集与分类[J].南京农业大学,1990,13(1)：30-36.
10.刘明,赵琦,王小菁,赵玉锦,童哲.植物的光受体及其调控机制的研究[J].生物学通报,2005,40(5)：10-12.
11.马月萍,戴思兰.高等植物成花分子机理的研究进展[J].分子植物育种,2007,5(z1)：21-28.
12.孟丽.蓝色花形成关键基因的分离及其表达分析[M].北京林业大学博士论文,2006.
13.孟丽,戴思兰.瓜叶菊F3’5’H基因cDNA的克隆、序列分析及其原核表达[J].分子植物育种,2005,3(6)：780-786.
14.孟丽,周琳,张明姝,戴思兰.一种有效的花瓣总RNA的提取方法[J].生物技术,2006,16(1)：38-40.
15.孙明霞,王宝增,范海,赵可夫.叶片中的花色素苷及其对植物适应环境的意义[J].植物生理学通讯,2003,39(6)：688-694.
16.孙卫,李崇晖,王亮生,戴思兰,徐彦军.花青素苷成分对瓜叶菊花色的影响[J].园艺学报,2009,36(12)：1775-1782.
17.孙卫,李崇晖,王亮生,戴思兰.菊花不同花色品种中花青素苷代谢分析[J].植物学报,2010,45(3)：327-336
18.田义,王强,张利义,康国栋,杨玲,郝红梅,杨振英,丛佩华.外源腐胺促进苹果果皮花青素苷积累的效应[J].植物学报,2009,44(3)：310-316.
19.王曼,王小菁.蓝光和蔗糖对拟南芥花色素苷积累和CHS基因表达的影响[J].热带亚热带植物学报,2004,12(3)：252-256.
20.徐清燏,戴思兰.蓝色花卉分子育种[J].分子植物育种,2004,2(1)：93-99.
21.许志茹,李玉花.‘津田芜菁’花色素苷生物合成相关基因的表达.植物生理与分子生物学学报32,583-586[J].植物生理学通讯,2006,44(5)：931-935.
22.杨致荣,王兴春,李西明,杨长登.高等植物转录因子的研究进展[J].遗传,2004,26(3)：403-408.
23.张剑亮,潘大仁,周以飞,吴明伟,柯祥德,林钿.PCR介导观赏向日葵DFR基因改造及其真核表达载体构建研究初报[J].福建农林大学学报,2008,37(2)：166-169.
24.张明姝.菊花查尔酮合成酶基因的克隆与序列分析[M].北京林业大学博士论文,2006.
25.张树林.菊花品种分类的研究[J].园艺学报,1965,4(1)：35-46.
26. Aharoni A, Vos C H R D, Wein M, Sun Z, Greco R, Kroon A, Mol J N M and O'Connell A P. The strawberry FaMYBl transcription factor suppresses anthocyanin and flavonol accumulation in transgenic tobacco [J]. The Plant Journal.2001,28(3):319-332.
27. Aida R, Kishimoto S, Tanaka Y and Shibata M. Modification of flower color in torenia (Torenia fournieri Lind.) by genetic transformation [J]. Plant Science.2000,153(1):33-42.
28. Akashi T, Ishizaki M, Aoki T and Ayabe S. Isoflavonoid production by adventitious-root cultures of Iris germanica (Iridaceae)[J]. Plant Biotechnology.2005,22(3):207-215.
29. Akira N, Yoko I and Masumi Y. Spatial and temporal expression of chalcon e synthase and dihydroflavonol 4-reductase genes in the Asiatic hybrid lily [J]. Plant science.2003,165(4): 759-767.
30. Albert N W, Lewis D H, Zhang H, Irving L J, Jameson P E and Davies K M. Light-induced vegetative anthocyanin pigmentation in Petunia [J]. Journal of Experimental Botany.2009: erp097.
31. Alfenito M R, Souer E, Goodman C D, Buell R, Mol J, Koes R and Walbot V. Functional complementation of anthocyanin sequestration in the vacuole by widely divergent glutathione S-transferases [J]. Plant Cell.1998,10(7):1135-1150.
32. Alla M M N and Younis M E. Herbicide effects on phenolic metabolism in maize (Zea mays L.)and soybean (Glycine max L.) seedling [J]. Journal of Experimental Botany.1995,46(11): 1731-1736.
33. Almeida J, Carpenter R, Robbins T P, Martin C and Coen E S. Genetic interactions underlying flower color patterns in Antirrhinum majus [J]. Genes & Development.1989,3(11):1758-1767.
34. Alvarez I, Costa A and Feliner G. Selecting Single-Copy Nuclear Genes for Plant Phylogenetics: A Preliminary Analysis for the Senecioneae (Asteraceae)[J]. Journal of Molecular Evolution. 2008,66(3):276-291.
35. Ashihara H, Deng W W, Mullen W and Crozier A. Distribution and biosynthesis of flavan-3-ols in Camellia sinensis seedlings and expression of genes encoding biosynthetic enzymes[J]. Phytochemistry.71(5-6):559-566.
36. Ban Y, Honda C, Hatsuyama Y, Igarashi M, Bessho H and Moriguchi T. Isolation and functional analysis of a MYB transcription factor gene that is a key regulator for the development of red coloration in apple skin [J]. Plant Cell Physiol.2007,48(7):958-970.
37. Banon S, Gonzalez A, Cano E A, Franco J A, Fernandez J A. Growth, development and colour response of potted Dianthus caryophyllus cv. Mondriaan to paclobutrazol treatment[J]. Scienta horticulturae.2002,94(3-4):371-377.
38. Barb J G, Werner D J and Griesbach R J. Genetics and biochemistry of flower color in stokes aster [J]. Journal of the American Society for Horticultural Science.2008,133(4):569-578.
39. Beld M, Martin C, Huits H, Stuitje A R and Gerats A G M. Flavonoid synthesis in Petunia hybrida:Partial characterization of dihydroflavonol-4-reductase genes [J]. Plant Molecular Biology.1989,13(5):491-502.
40. Biolley J P, Jay M. Anthocyanins in modern roses:chemical and colorimetric features in relation to the colour range[J]. Journal of Experimental Botany.1993,44(268):1725-1734.
41. Biran I and Halevy A H. Effects of varying light intensities and temperature treatments applied to whole plants, or locally to leaves or flower buds, on growth and pigmentation of'Baccara'roses [J]. Physiologia Plantarum.1974,31(3):175-179.
42. Boddu J, Svabek C, Sekhon R, Gevens A, Nicholson R L, Jones A D, Pedersen J F, Gustine D L and Chopra S. Expression of a putative flavonoid 3'-hydroxylase in sorghum mesocotyls synthesizing 3-deoxyanthocyanidin phytoalexins [J]. Physiological and Molecular Plant Pathology.2004,65(2):101-113.
43. Bogs J, Jaffe F W, Takos A M, Walker A R and Robinson S P. The grapevine transcription factor VvMYBPA1 regulates proanthocyanidin synthesis during fruit development [J]. Plant Physiology. 2007,143(3):1347-1361.
44. Bolwell G P, Bozak K and Zimmerlin A. Plant cytochrome P450 [J]. Phytochemistry.1994, 37(6):1491-1506.
45. Bongue-Bartelsman M, O'Neill S D, Tong Y and Yoder J I. Characterization of the gene encoding dihydroflavonol 4-reductase in tomato[J]. Gene.1994,138(1-2):153-157.
46. Borevitz J O, Xia Y, Blount J, Dixon R A and Lamb C. Activation tagging identifies a conserved MYB regulator of phenylpropanoid biosynthesis [J]. Plant Cell.2000,12(12):2383-2394.
47. Boss P K, Davies C and Robinson S P. Anthocyanin composition and anthocyanin pathway gene expression in grapevine sports differing in berry skin colour [J]. Australian Journal of Grape and Wine Research.1996,2(3):163-170.
48. Bradley M J, Davies K M, Deroles S C, Bloor S J and Lewis D H. The maize Lc regulatory gene up-regulates the flavonoid biosynthetic pathway of Petunia [J]. The Plant Journal.1998,13(3): 381-392.
49. Britsch L, Ruhnau-Brich B and Forkmann G. Molecular cloning, sequence analysis, and in vitro expression of flavanone 3 beta-hydroxylase from Petunia hybrida [J]. Journal of Biological Chemistry.1992,267(8):5380-5387.
50. Broun P. Transcriptional control of flavonoid biosynthesis: A complex network of conserved regulators involved in multiple aspects of differentiation in Arabidopsis [J]. Current Opinion in Plant Biology.2005,8(3):272-279.
51. Brugliera F, Barri-Rewell G, Holton T A and Mason J G. Isolation and characterization of a flavonoid 3'-hydroxylase cDNA clone corresponding to the htl locus of Petunia hybrida [J]. The Plant Journal.1999,19(4):441-451.
52. Carey C C, Strahle J T, Selinger D A and Chandler V L. Mutations in the pale aleurone color1 regulatory gene of the zea mays anthocyanin pathway have distinct phenotypes relative to the functionally similar transparent testa glabral gene in Arabidopsis thaliana [J]. Plant Cell.2004, 16(2):450-464.
53. Castellarin S, Di Gaspero G, Marconi R, Nonis A, Peterlunger E, Paillard S, Adam-Blondon A and Testolin R. Colour variation in red grapevines(Vitis vinifera L.):genomic organisation, expression of flavonoid 3'-hydroxylase, flavonoid 3',5'-hydroxylase genes and related metabolite profiling of red cyanidin-/blue delphinidin-based anthocyanins in berry skin[J]. BMC Genomics. 2006,7:12.
54. Castellarin S and Di Gaspero G. Transcriptional control of anthocyanin biosynthetic genes in extreme phenotypes for berry pigmentation of naturally occurring grapevines [J]. BMC Plant Biology.2007,7(1):46.
55. Chalker-Scott L. Environmental significance of anthocyanins in plant stress responses [J]. Photochemistry and Photobiology.1999,70(1):1-9.
56. Chandler V L, Radicella J P, Robbins T P, Chen J and Turks D. Two regulatory genes of the maize anthocyanin pathway are homologous:Isolation of B utilizing R genomic sequences [J]. Plant Cell.1989,1(12):1175-1183.
57. Charrier B, Coronado C, Kondorosi A and Ratet P. Molecular characterization and expression of alfalfa (Medicago sativa L.) flavanone-3-hydroxylase and dihydroflavonol-4-reductase encoding genes [J]. Plant Molecular Biology.1995,29(4):773-786.
58. Chatterjee M, Sharma P and Khurana J P. Cryptochrome 1 from Brassica napus is up-regulated by blue light and controls hypocotyl/stem growth and anthocyanin accumulation [J]. Plant Physiology.2006,141(1):61-74.
59. Cheng L, Xu Y, Grotewold E, Jin Z, Wu F, Fu C and Zhao D. Characterization of Anthocyanidin Synthase (ANS) Gene and anthocyanidin in rare medicinal plant-Saussurea medusa[J]. Plant Cell, Tissue and Organ Culture.2007,89(1):63-73.
60. Cho S, Chen W and Muehlbauer F J. Constitutive expression of the Flavanone 3-hydroxylase gene related to pathotype-specific ascochyta blight resistance in Cicer arietinum L.[J]. Physiological and Molecular Plant Pathology.2005,67(2):100-107.
61. Christie J M and Jenkins G I. Distinct UV-B and UV-A/blue light signal transduction pathways induce chalcone synthase gene expression in Arabidopsis cells [J]. Plant Cell.1996,8(9): 1555-1567.
62. Christie P, Alfenito M and Walbot V. Impact of low-temperature stress on general phenylpropaniod and anthocyanin pathways:enhancement of transcript abundance and anthocyanin pigmentation in maize seedlings [J]. Planta.1994,194(4):541-549.
63. Cominelli E, Gusmaroli G, Allegra D, Galbiati M, Wade H K, Jenkins G I and Tonelli C. Expression analysis of anthocyanin regulatory genes in response to different light qualities in Arabidopsis thaliana [J]. Journal of Plant Physiology.2008,165(8):886-894.
64. Cone K C, Burr F A and Burr B. Molecular analysis of the maize anthocyanin regulatory locus C1 [J]. Proceedings of the National Academy of Sciences of the United States of America.1986, 83(24):9631-9635.
65. Cone K C, Cocciolone S M, Burr F A and Burr B. Maize anthocyanin regulatory gene pl is a duplicate of cl that functions in the plant [J]. Plant Cell.1993,5(12):1795-1805.
66. Courtney-Gutterson N, Napoli C, Lemieux C, Morgan A, Firoozabady E and Robinson K E P. Modification of flower color in florist's chrysanthemum:Production of a white-flowering variety through molecular genetics [J]. Nat Biotech.1994,12(3):268-271.
67. David W. Regulation of flower pigmentation and growth: Multiple signaling pathways control anthocyanin synthesis in expanding petals [J]. Physiologia Plantarum.2000,110(2):152-157.
68. Davies K M. A cDNA clone for flavanone 3-hydroxylase from Malus[J]. Plant Physiol.1993, 103(1):291.
69. Davies K M, Schwinn K E, Deroles S C, Manson D G, Lewis D H, Bloor S J and Bradley J M. Enhancing anthocyanin production by altering competition for substrate between flavonol synthase and dihydroflavonol 4-reductase [J]. Euphytica.2003,131(3):259-268.
70. de Vetten N, Quattrocchio F, Mol J and Koes R. The anll locus controlling flower pigmentation in petunia encodes a novel WD-repeat protein conserved in yeast, plants, and animals [J]. Genes & Development.1997,11(11):1422-1434.
71. Deboo G B, Albertsen M C and Taylor L P. Flavanone 3-hydroxylase transcripts and flavonol accumulation are temporally coordinate in maize anthers [J]. The Plant Journal.1995,7(5): 703-713.
72. Deroles S C, Bradley J M, Schwinn K E, Markham K R, Bloor S, Manson D G and Davies K M. An antisense chalcone synthase cDNA leads to novel colour patterns in lisianthus(Eustoma grandiflorum) flowers [J]. Molecular Breeding.1998,4(1):59-66.
73. Dixon R A, Harrison M J and Lamb C J. Early events in the activation of plant defense responses [J]. Annual Review of Phytopathology.1994,32(1):479-501.
74. Dixon R A and Steele C L. Flavonoids and isoflavonoids- a gold mine for metabolic engineering [J]. Trends in Plant Science.1999,4(10):394-400.
75. Dong Y H, Beuning L, Davies K, Mitra D, Morris B and Kootstra A. Expression of pigmentation genes and photo-regulation of anthocyanin biosynthesis in developing royal gala apple flowers [J]. Australian Journal of Plant Physiology.1998,25(2):245-252.
76. Dooner H K. Coordinate genetic regulation of flavonoid biosynthetic enzymes in maize [J]. Molecular and General Genetics MGG.1983,189(1):136-141.
77. Druka A, Kudrna D, Rostoks N, Brueggeman R, von Wettstein D and Kleinhofs A. Chalcone isomerase gene from rice (Oryza sativa) and barley(Hordeum vulgare): physical, genetic and mutation mapping[J]. Gene.2003,302(1-2):171-178.
78. Dubos C, Gourrierec J L, Baudry A, Huep G, Lanet E, Debeaujon I, Routaboul J M, Alboresi A, Weisshaar B and Lepiniec L. MYBL2 is a new regulator of flavonoid biosynthesis in Arabidopsis thaliana [J]. The Plant Journal.2008,55(6):940-953.
79. Durbin M L, McCaig B and Clegg M T. Molecular evolution of the chalcone synthase multigene family in the morning glory genome [J]. Plant Molecular Biology.2000,42(1):79-92.
80. Elomaa P, Honkanen J, Puska R, Seppanen P, Helariutta Y, Mehto M, Kotilainen M, Nevalainen L and Teeri T H. Agrobacterium-mediated transfer of antisense chalcone synthase cDNA to Gerbera hybrida inhibits flower pigmentation [J]. Nature Biotechnology 1993,11:508-511.
81. Elomaa P, Mehto M, Kotilainen M, Yrjo Helariutta, Nevalainen L and Teeri T H. A bHLH transcription factor mediates organ, region and flower type specific signals on dihydroflavonol-4-reductase (dfr) gene expression in the inflorescence of Gerbera hybrida (Asteraceae) [J]. The Plant Journal.1998,16(1):93-99.
82. Epping B, Kittel M, Ruhnau B and Hemleben V. Isolation and sequence analysis of a chalcone synthase cDNA of Matthiola incana R. Br. (Brassicaceae)[J]. Plant Molecular Biology.1990, 14(6):1061-1063.
83. Espley R, Hellens R, Putterill J, Stevenson D, Kutty-Amma S and Allan A. Red colouration in apple fruit is due to the activity of the MYB transcription factor, MdMYB10 [J]. Plant J.2007, 49(3):414-427.
84. Farzad M, Griesbach R, Hammond J, Weiss M R and Elmendorf H G. Differential expression of three key anthocyanin biosynthetic genes in a color-changing flower, Viola cornuta cv. Yesterday, Today and Tomorrow [J]. Plant Science.2003,165(6):1333-1342.
85. Feldbrugge M, Sprenger M, Hahlbrock K and Weisshaar B. PcMYB1, a novel plant protein containing a DNA-binding domain with one MYB repeat, interacts in vivo with a light-regulatory promoter unit [J]. Plant Journal.1997,11(5):1079-1093.
86. Feller A, Hernandez J M and Grotewold E. An ACT-like domain participates in the dimerization of several plant basic-helix-loop-helix transcription factors [J]. Journal of Biological Chemistry. 2006,281(39):28964-28974.
87. Ferrer J L, Jez J M, Bowman M E, Dixon R A and Noel J P. Structure of chalcone synthase and the molecular basis of plant polyketide biosynthesis [J]. Nature Structural Biology.1999,6(8): 775-784.
88. Forkmann G and Martens S. Metabolic engineering and applications of flavonoids [J]. Current Opinion in Biotechnology.2001,12(2):155-160.
89. Forkmann G and Ruhnau B. Distinct substrate specificity of dihydroflavonol 4-reductase from flowers of Petunia hybrida [J]. Zeitschrift fur Naturforschung.1987,42:1146-1148.
90. Fukada-Tanaka S, Hoshino A, Hisatomi Y, Habu Y, Hasebe M and Iida S. Identification of New Chalcone Synthase Genes for Flower Pigmentation in the Japanese and Common Morning Glories[J]. Plant Cell Physiol.1997,38(6):754-758.
91. Fukui Y, Tanaka Y, Kusumi T, Iwashita T and Nomoto K. A rationale for the shift in colour towards blue in transgenic carnation flowers expressing the flavonoid 3',5'-hydroxylase gene [J]. Phytochemistry 2003,63(1):15-23.
92. Fukusaki E-i, Kawasakia K, Kajiyamaa S-i, Ana C I, Suzukib K, Tanakac Y and Kobayashia A. Flower color modulations of Torenia hybrida by downregulation of chalcone synthase genes with ma interference [J]. Journal of Biotechnology 2004,111(3):229-240.
93. Gerats A, Huits H, Vrijlandt E, Marana C, Souer E and Beld M. Molecular characterization of a nonautonomous transposable element (dTph1) of petunia [J]. Plant Cell.1990,2(11):1121-1128.
94. Goff S A, Cone K C and 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 and Development.1992,6(5):864-875.
95. Goff S A, Klein T M, Roth B A, Fromm M E, Cone K C, Radicella J P and Chandler V L. Transactivation of anthocyanin biosynthetic genes following transfer of B regulatory genes into maize tissues [J]. The EMBO Journal.1990,9(8):2517-2522.
96. Goldsbrough A P, Tong Y and Yoder J I. Lc as a non-destructive visual reporter and transposition excision marker gone for tomato [J]. The Plant Journal.1996,9(6):927-933.
97. Gong Z, Yamazaki M, Sugiyama M, Tanaka Y and Saito K. Cloning and molecular analysis of structural genes involved in anthocyanin biosynthesis and expressed in a forma-specific manner in Perilla frutescens [J]. Plant Molecular Biology.1997,35(6):915-927.
98. Gong Z Z, Yamagishi E, Yamazaki M and Saito K. A constitutively expressed Myc-like gene involved in anthocyanin biosynthesis from Perilla frutescens: molecular characterization, heterologous expression in transgenic plants and transactivation in yeast cells [J]. Plant Molecular Biology.1999,41(1):33-44.
99. Gonnet J F. CIElab measurement, a precies communication in flower colour: An example with carnation (Dianthus caryophyllus) cultivars [J]. Journal of horticultural science.1993,68(4): 499-510.
100. Goodrich J, Carpenter R and Coen E S. A common gene regulates pigmentation pattern in diverse plant species [J]. Cell.1992,68(5):955-964.
101. Goto T, Kondo T, Kawai T and Tamura H. Structure of cinerarin, a tetra-acylated anthocyanin isolated from the blue garden cineraria, Senecio cruentus [J]. Tetrahedron letters.1984,25(52): 6021-6024.
102. Griesbach R J. Correlation of pH and light intensity on flower color in potted Eustoma grandiflorum Grise [J]. HortScience.1992,27(7):817-818.
103. Grotewold E. The genetics and biochemistry of floral pigments [J]. Annual Review of Plant Biology.2006,57(1):761-780.
104. Grotewold E and Peterson T. Isolation and characterization of a maize gene encoding chalcone flavonone isomerase[J]. Molecular and General Genetics MGG.1994,242(1):1-8.
105. Grotewold E, Sainz M B, Tagliani L, Hernandez J M, Bowen B and Chandler V L. Identification of the residues in the Myb domain of maize C1 that specify the interaction with the bHLH cofactor R [J]. Proceedings of the National Academy of Sciences of the United States of America. 2000,97(25):13579-13584.
106. Guo J, Han W and Wang M H. Ultraviolet and environmental stresses involved in the induction and regulation of anthocyanin biosynthesis: A review [J]. African Journal of Biotechnology.2008, 7(25):4966-4972.
107. Gutterson N C, Napoli C, Lemieux C, Morgan A, Firoozabady E and Robinson K E. Modification of flower color in florist's chrysanthemum: production of a white-genetics [J]. Nature Biotechnology.1994,12:268-271.
108. Halevy A. Manipulation of flowering[M]. London: Butterworth,1987:363-378.
109. Han Y, Ming F, Wang J, Ye M and Shen D. A novel chalcone synthase gene from Phalaenopsis orchid that alters floral morphology in transgenic tobacco plants [J]. Plant Molecular Biology Reporter.2005,23:193a-193m.
110. Harborne J B and Williams C A. Advances in flavonoid research since 1992 [J]. Phytochemistry. 2000,55(6):481-504.
111. Hartmann U, Sagasser M, Mehrtens F, Stracke R and Weisshaar B. Differential combinatorial interactions of cis-acting elements recognized by R2R3-MYB, BZIP, and BHLH factors control light-responsive and tissue-specific activation of phenylpropanoid biosynthesis genes [J]. Plant Molecular Biology.2005,57(2):155-171.
112. Hartmann U, Valentine W J, Christie J M, Hays J, Jenkins G I and Weisshaar B. Identification of UV/blue light-response elements in the Arabidopsis thaliana chalcone synthase promoter using a homologous protoplast transient expression system [J]. Plant Molecular Biology.1998,36(5): 741-754.
113. Helariutta Y, Elomaa P, Kotilainen M, Griesbach R J, Schroder J and Teeri T H. Chalcone synthase-like genes active during corolla development are differentially expressed and encode enzymes with different catalytic properties in Gerbera hybrida (Asteraceae)[J]. Plant Molecular Biology.1995,28(1):47-60.
114. Helariutta Y, Elomaa P, Kotilainen M, Seppanen P and Teeri T H. Cloning of cDNA coding for dihydroflavonol-4-reductase (DFR) and characterization of dfr expression in the corollas of Gerbera hybrida var. Regina (Compositae) [J]. Plant Molecular Biology.1993,22(2):183-193.
115. Heller W, Forkmann G, Britsch L and Grisebach H. Enzymatic reduction of (+)-dihydroflavonols to flavan-3,4-cis-diols with flower extracts from Matthiola incana and its role in anthocyanin biosynthesis [J]. Planta.1985,165(2):284-287.
116. Hirner A A and Seitz H U. Isoforms of chalcone synthase in Daucus carota L. And their differential expression in organs from the european wild carrot and in ultraviolet-A-irradiated cell cultures [J]. Planta.2000,210(6):993-998.
117. Hisatomi Y, Yoneda Y, Kasahara K, Inagaki Y and Iida S. DNA rearrangements at the region of the dihydroflavonol 4-reductase gene for flower pigmentation and incomplete dominance in morning glory carrying the mutable flaked mutation[J]. TAG Theoretical and Applied Genetics. 1997,95(4):509-515.
118. Holton T and Cornish E. Genetics and biochemistry of anthocyanin biosynthesis [J]. Plant Cell. 1995,7(7):1071-1083.
119. Hondo T, Yoshida K, Nakagawa A, Kawai T, Tamura H and Goto T. Structural basis of blue-colour development in flower petals from Commelina communis [J]. Nature.1992,358: 515-518.
120. Hoshino A, Johzuka-Hisatomi Y and Iida S. Gene duplication and mobile genetic elements in the morning glories [J]. Gene.2001,265(1-2):1-10.
121. Hoshino A, Abe Y, Saito N, Inagaki Y and Iida S. The Gene Encoding Flavanone 3-Hydroxylase is Expressed Normally in the Pale Yellow Flowers of the Japanese Morning Glory Carrying the speckled Mutation Which Produce Neither Flavonol nor Anthocyanin but Accumulate Chalcone, Aurone and Flavanone[J]. Plant Cell Physiol.1997,38(8):970-974.
122. Hoshino A, Morita Y, Choi J D, Saito N, Toki K, Tanaka Y and Iida S. Spontaneous mutations of the flavonoid 3'-hydroxylase gene conferring reddish flowers in the three morning glory species[J]. Plant and Cell Physiology.2003,44(10):990-1001.
123. Hrazdina G and Jensen R A. Spatial organization of enzymes in plant metabolic pathways [J]. Annual Review of Plant Physiology and Plant Molecular Biology.1992,43(1):241-267.
124. Hrazdina G, Marx G A and Hoch H C. Distribution of secondary plant metabolites and their biosynthetic enzymes in pea (Pisum sativum L.) leaves:anthocyanins and flavonol glycosides [J]. Plant Physiolgy.1982,70(3):745-748.
125. Huang Z, Liang M, Peng J, Xing T and Wang X. Exogenous ammonium inhibits petal pigmentation and expansion in Gerbera hybrida [J]. Physiologia Plantarum.2008,133(2): 254-265.
126. Huh E, Shin H, Choi S, Kwon O and Lee Y. Thermosusceptible developmental stage in anthocyanin accumulation and color response to high temperature in red chrysanthemum cultivars [J]. Korean Journal of Horticultural Science and Technology.2008,26(4):357-361.
127. Huits H S M, Gerats A G M, Kreike M M, Mol J N M and Koes R E. Genetic control of dihydroflavonol 4-reductase gene expression in Petunia hybrida [J]. The Plant Journal.1994,6(3): 295-310.
128. Inagaki Y, Johzuka-Hisatomi Y, Mori T, Takahashi S, Hayakawa Y, Peyachoknagul S, Ozeki Y and Iida S. Genomic organization of the genes encoding dihydroflavonol 4-reductase for flower pigmentation in the Japanese and common morning glories [J]. Gene.1999,226(2):181-188.
129. Islam M S, Jalaluddin M, Garner J O, Yoshimoto M and Yamakawa O. Artificial shading and temperature influence on anthocyanin compositions in sweetpotato leaves [J]. HortScience.2005, 40:176-180.
130. Jaakola L, Maatta K, Pirttila A M, Torronen R, Karenlampi S and Hohtola A. Expression of genes involved in anthocyanin biosynthesis in relation to anthocyanin, proanthocyanidin, and flavonol levels during bilberry fruit development [J]. Plant Physiology.2002,130(2):729-739.
131. Jackson D, Culianez-Macia F, Prescott A G, Roberts K and Martin C. Expression patterns of myb genes from antirrhinum flowers [J]. Plant Cell.1991,3(2):115-125.
132. Jackson D, Roberts K and Martin C. Temporal and spatial control of expression of anthocyanin biosynthetic genes in developing flowers of Antirrhinum majus [J]. The Plant Journal.1992,2(4): 425-434.
133. Jez J M, Bowman M E, Dixon R A and Noel J P. Structure and mechanism of the evolutionarily unique plant enzyme chalcone isomerase [J]. Nature Structural and Molecular Biology.2000, 7(9):786-791.
134. Jez J M and Noel J P. Mechanism of chalcone synthase. pKa of the catalytic cysteine and the role of the conserved histidine in a plant polyketide synthase [J]. Journal of Biological Chemistry. 2000,275(50):39640-39646.
135. Johzuka-Hisatomi Y, Hoshino A, Mori T, Habu Y and Iida S. Characterization of the chalcone synthase genes expressed in flowers of the common and Japanese morning glories [J]. Genes and Genetic Systems.1999,74(4):141-147.
136. Jonsson L M V, Donker-Koopman W E and Schram A W. Turnover of anthocyanins and tissue compartmentation of anthocyanin biosynthesis in flowers of Petunia hybrida [J]. Multilingue. 1984,115(1):29-37.
137. Kaltenbach M, Schroder G, Schmelzer E, Lutz V and Schroder J. Flavonoid hydroxylase from Catharanthus roseus: cDNA, heterologous expression, enzyme properties and cell-type specific expression in plants[J]. Plant J.1999,19(2):183-193.
138. Katsumoto Y, Fukuchi-Mizutani M, Fukui Y, Brugliera F, Holton T A, et al. Engineering of the rose flavonoid biosynthetic pathway successfully generated blue-hued flowers accumulating delphinidin [J]. Plant Cell Physiol.2007,48(11):1589-1600.
139. Kawabata S, Ohta M, Kusuhara Y and Sakiyama R. Influences of low light intensities on the pigmentation of Eustoma grandiflorum flowers [J]. Acta Horticulturae.1995,405:173-178.
140. Kim S, Jones R, Yoo K-S and Pike L M. The L locus, one of complementary genes required for anthocyanin production in onions (Allium cepa), encodes anthocyanidin synthase [J]. TAG Theoretical and Applied Genetics.2005,111(1):120-127.
141.Kitada C, Gong Z, Tanaka Y, Yamazaki M and Saito K. Differential expression of two cytochrome P450s involved in the biosynthesis of flavones and anthocyanins in chemo-varietal forms of Perillafrutescens[J]. Plant Cell Physiol.2001,42(12):1338-1344.
142. Kobayashi, Kobayashi S, Ishimaru, Ishimaru M, Hiraoka, Hiraoka K, Honda and Honda C. Myb-related genes of the kyoho grape(Vitis labruscana) regulate anthocyanin biosynthesis [J]. Planta.2002,215(6):924-933.
143. Koes R, Verweij W and Quattrocchio F. Flavonoids: A colorful model for the regulation and evolution of biochemical pathways [J]. Trends in Plant Science.2005,10(5):236-242.
144. Koes R E, Spelt C E and Mol J N M. The chalcone synthase multigene family of Petunia hybrida (V30):Differential, light-regulated expression during flower development and UV light induction [J]. Plant Molecular Biology.1989,12(2):213-225.
145. Koes R E, van Blokland R, Quattrocchio F, van Tunen A J and Mol J. Chalcone synthase promoters in petunia are active in pigmented and unpigmented cell types [J]. Plant Cell.1990, 2(5):379-392.
146. Kranz H D, Denekamp M, Greco R, Jin H, Leyva A, et al. Towards functional characterisation of the members of the R2R3-MYB gene family from Arabidopsis thaliana[J]. The Plant Journal. 1998,16(2):263-276.
147. Kreuzaler F, Ragg H, Fautz E, Kuhn D N and Hahlbrock K. UV-induction of chalcone synthase mRNA in cell suspension cultures of Petroselinum hortense [J]. Proceedings of the National Academy of Sciences.1983,80(9):2591-2593.
148. Kusumi T, Tanaka Y, Suzuki K and Katsumoto Y. Development of flower colour modification technology and new floricultural varieties [J]. Nippon Nogeikagaku Kaishi.2002,76(8):700-707.
149. Lalusin A G, Ohta M and Fujimura T. Temporal and spatial expression of genes involved in anthocyanin biosynthesis during sweet potato (Ipomoea batatas [L.] Lam.) root development [J]. International Journal of Plant Sciences.2006,167(2):249-256.
150. Leyva A, Jarillo J A, Salinas J and Martinez-Zapater J M. Low temperature induces the accumulation of phenylalanine ammonia-lyase and chalcone synthase mrnas of Arabidopsis thaliana in a light-dependent manner [J]. Plant Physiology.1995,108(1):39-46.
151. Li F, Jin Z, Qua W, Zhao D and Ma F. Cloning of a cDNA encoding the Saussurea medusa chalcone isomerase and its expression in transgenic tobacco [J]. Plant Physiology and Biochemistry.2006,44(7-9):455-461.
152. Lillo C, Lea U S and Ruoff P. Nutrient depletion as a key factor for manipulating gene expression and product formation in different branches of the flavonoid pathway [J]. Plant, Cell and Environment.2008,31:587-601.
153. Liu C-J, Blount J W, Steele C L and Dixon R A. Bottlenecks for metabolic engineering of isoflavone glycoconjugates in Arabidopsis [J]. Proceedings of the National Academy of Sciences of the United States of America.2002,99(22):14578-14583.
154. Lloyd A, Walbot V and Davis R. Arabidopsis and nicotiana anthocyanin production activated by maize regulators R and C1 [J]. Science.1992,258(5089):1773-1775.
155. Loreti E, Povero G, Novi G, Solfanelli C, Alpi A and Perata P. Gibberellins, jasmonate and abscisic acid modulate the sucrose-induced expression of anthocyanin biosynthetic genes in Arabidopsis [J]. New Phytologist.2008,179(4):1004-1016.
156. Lu Q and Lu Q N. cDNA cloning and expression of anthocyanin biosynthetic genes in wild potato (Solanum pinnatisectum) [J]. African Journal of Biotechnology.2006,5(10):811-818.
157. Ludwig S R, Habera L F, Dellaporta S L and Wessler S R. Lc, a member of the maize R gene family responsible for tissue-specific anthocyanin production, encodes a protein similar to transcriptional activators and contains the myc-homology region [J]. Proceedings of the National Academy of Sciences of the United States of America.1989,86(18):7092-7096.
158. Lukacin R and Britsch L. Identification of strictly conserved histidine and arginine residues as part of the active site in Petunia hybrida flavanone 3beta-hydroxylase[J]. Eur J Biochem.1997, 249(3):748-757.
159. Lulin M, Yi S, Aizhong C, Zengjun Q, Liping X, Peidu C, Dajun L and Xiu-e W. Molecular cloning and characterization of an up-regulated UDP-glucosyltransferase gene induced by DON from Triticum aestivum L. cv. Wangshuibai[J]. Molecular Biology Reports.37(2):785-795.
160. Ma L, Liu B, Gao D, Pang X, Lǖ S, Yu H, Wang H, Yan F, Li Z and Li Y. Molecular cloning and overexpression of a novel UDP-glucosyltransferase elevating salidroside levels in Rhodiola sachalinensis[J]. Plant cell reports.2007,26(7):989-999.
161. McKhann H I and Hirsch A M. Isolation of chalcone synthase and chalcone isomerase cDNAs from alfalfa (Medicago sativa L.):highest transcript levels occur in young roots and root tips[J]. Plant Molecular Biology.1994,24(5):767-777.
162. Mano H, Ogasawara F, Sato K, Higo H and Minobe Y. Isolation of a regulatory gene of anthocyanin biosynthesis in tuberous roots of purple-fleshed sweet potato [J]. Plant Physiology. 2007,143(3):1252-1268.
163. Markham K R. Novel anthocyanins produced in petals of genetically transformed lisianthus [J]. Phytochemistry.1996,42(4):1035-1038.
164. Marrs K A, Alfenito M R, Lloyd A M and Walbot V. A glutathione S-transferase involved in vacuolar transfer encoded by the maize gene Bronze-2 [J]. Nature.1995,375(6530):397-400.
165. Martin C and Paz-Ares J. MYB transcription factors in plants [J]. Trends in Genetics.1997, 13(2):67-73.
166. Martin C, Prescott A, Mackay S, Bartlett J and Vrijlandt E. Control of anthocyanin biosynthesis in flowers of Antirrhinum majus [J]. The Plant Journal.1991,1(1):37-49.
167. Mathews H, Clendennen S K, Caldwell C G, Liu X L, Connors K, Matheis N, Schuster D K, Menasco D J, Wagoner W, Lightner J and Wagner D R. Activation tagging in tomato identifies a transcriptional regulator of anthocyanin biosynthesis, modification, and transport [J]. Plant Cell. 2003,15(8):1689-1703.
168. Mato M, Onozaki T, Ozeki Y, Higeta D, Itoh Y, Yoshimoto Y, Ikeda H, Yoshida H and Shibata M. Flavonoid biosynthesis in white-flowered sim carnations (Dianthus caryophyllus) [J]. Scientia Horticulturae.2000,84(3-4):333-347.
169. Matus J T, Loyola R, Vega A, Pena-Neira A, Bordeu E, Arce-Johnson P and Alcalde J A. Post-veraison sunlight exposure induces myb-mediated transcriptional regulation of anthocyanin and flavonol synthesis in berry skins of Vitis vinifera [J]. Journal of Experimental Botany.2009, 60(3):853-867.
170. Mehrtens F, Kranz H, Bednarek P and Weisshaar B. The Arabidopsis transcription factor MYB 12 is a flavonol-specific regulator of phenylpropanoid biosynthesis [J]. Plant Physiol.2005, 138(2):1083-1096.
171. Meng X, Xing T and Wang X. The role of light in the regulation of anthocyanin accumulation in Gerbera hybrida [J]. Plant Growth Regulation.2004,44(3):243-250.
172. Mol J, Cornish E, Mason J and Koes R. Novel coloured flowers [J]. Current Opinion in Biotechnology.1999,10(2):198-201.
173. Mol J, Grotewold E and Koes R. How genes paint flowers and seeds [J]. Trends Plant Sci.1998, 3(6):212-217.
174. Mol J, Jenkins G, Schafer E and Weiss D. Signal perception, tranduction, and gene expression involved in anthocyanin biosynthesis [J]. Critical reviews in plant sciences.1996,15(5-6) 525-557.
175. Mori K, Goto-Yamamoto N, Kitayama M and Hashizume K. Loss of anthocyanins in red-wine grape under high temperature [J]. Journal of Experimental Botany.2007:on line.
176. Mori K, Sugaya S and Gemma H. Decreased anthocyanin biosynthesis in grape berries grown under elevated night temperature condition [J]. Scientia Horticulturae.2005,105(3):319-330.
177. Morita Y, Saitoh M, Hoshino A, Nitasaka E and Iida S. Isolation of cDNAs for R2R3-MYB, bHLH and WDR transcriptional regulators and identification of c and ca mutations conferring white flowers in the Japanese morning glory [J]. Plant Cell Physiology.2006,47(4):457-470.
178. Moscovici S, Moalem-Beno D and Weiss D. Leaf-mediated light responses in petunia flowers [J]. Plant Physiology.1996,110(4):1275-1282.
179. Muir S R, Collins G J, Robinson S, Hughes S, Bovy A, Ric De Vos C H, van Tunen A J and Verhoeyen M E. Overexpression of petunia chalcone isomerase in tomato results in fruit containing increased levels of flavonols [J]. Nature Biotechnology.2001,19(5):470-474.
180. Nakajima J-i, Tanaka Y, Yamazaki M and Saito K. Cdna clonlng and gene expression of anthocyanidin synthase from Torenia fiournieri [J]. Plant Biotechnology.2000,17(4):331-335.
181. Nakamura N, Fukuchi-Mizutani M, Miyazaki K, Suzuki K and Tanaka Y. RNAi suppression of the anthocyanidin synthase gene in Torenia hybrida yields white flowers with higher frequency and better stability than antisense and sense suppression [J]. Plant Biotechnology.2006,23(1): 13-18.
182. Nakatsuka A, Izumi Y and Yamagishi M. Spatial and temporal expression of chalcone synthase and dihydroflavonol 4-reductase genes in the Asiatic hybrid lily[J]. Plant Science.2003,165: 759-767.
183. Nakatsuka A, Yamagishi M, Nakano M, Tasaki K and Kobayashi N. Light-induced expression of basic helix-loop-helix genes involved in anthocyanin biosynthesis in flowers and leaves of Asiatic hybrid lily [J]. Scientia Horticulturae.2009, in press.
184. Nakatsuka T, Nishihara M, Mishiba K, Hirano H and Yamamura S. Two different transposable elements inserted in flavonoid 3',5'-hydroxylase gene contribute to pink flower coloration in Gentiana scabra [J]. Molecular Genetics and Genomics. 2006,275(3):231-241.
185. Nakatsuka T, Nishihara M, Mishiba K and Yamamura S. Temporal expression of flavonoid biosynthesis-related genes regulates flower pigmentation in gentian plants [J]. Plant Science. 2005a,168(5):1309-1318.
186. Nakatsuka T, Nishihara M, Mishiba K and Yamamura S. Two different mutations are involved in the formation of white-flowered gentian plants [J]. Plant Science.2005b,169(5):949-958.
187. Nakatsuka T, Sato K, Takahashi H, Yamamura S and Nishihara M. Cloning and characterization of the UDP-glucose:anthocyanin 5-O-glucosyltransferase gene from blue-flowered gentian [J]. Journal of Experimental Botany.2008,59(6):1241-1252.
188. Nakayama M, Koshioka M, Shibata M, Hiradate S, Sugie H and Yamaguchi M A. Identification of cyanidin 3-O-(3,6"-O-dimalonyl-β-glucopyranoside) as a flower pigment of chrysanthemum (Dendranthema grandiflorum) [J]. Bioscience, biotechnology, and biochemistry.1997,61(9): 1607-1608.
189. Napoli C, Lemieux C and Jorgensen R. Introduction of a chimeric chalcone synthase gene into petunia results in reversible co-suppression of homologous genes in trans [J]. Plant Cell.1990, 2(4):279-289.
190. Nesi N, Debeaujon I, Jond C, Pelletier G, Caboche M and Lepiniec L. 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.
191. Nesi N, Debeaujon I, Jond C, Stewart A J, Jenkins G I, Caboche M and Lepiniec L. The TRANSPARENT TESTA16 locus encodes the ARABIDOPSIS BSISTER MADS domain protein and is required for proper development and pigmentation of the seed coat [J]. Plant Cell.2002, 14(10):2463-2479.
192. Nesi N, Jond C, Debeaujon I, Caboche M and Lepiniec L. The Arabidopsis TT2 gene encodes an R2R3 MYB domain protein that acts as a key determinant for proanthocyanidin accumulation in developing seed [J]. Plant Cell.2001,13(9):2099-2114.
193. Nielsen K, Deroles S C, Markham K R, Bradley M J, Podivinsky E and Manson D. Antisense flavonol synthase alters copigmentation and flower color in lisianthus [J]. Molecular Breeding. 2002,9(4):217-229.
194. Nishihara M, Nakatsuka T and Yamamura S. Flavonoid components and flower color change in transgenic tobacco plants by suppression of chalcone isomerase gene [J]. FEBS Letters.2005, 579(27):6074-6078.
195. Noda N, Kanno Y, Kato N, Kazuma K and Suzuki M. Regulation of gene expression involved in flavonol and anthocyanin biosynthesis during petal development in lisianthus (Eustoma grandiflorum) [J]. Physiologia Plantarum.2004,122(3):305-313.
196. Nozaki K, Takamura T and Fukai S. Effects of high temperature on flower colour and anthocyanin content in pink flower genotypes of greenhouse chrysanthemum(Chrysanthemum morifolium Ramat.) [J]. The Journal of Horticultural Science and Biotechnology 2006,81(4): 728-734.
197. Ogata J, Itoh Y, Ishida M, Yoshida H and Ozeki Y. Cloning and heterologous expression of cDNAs encoding flavonoid glucosyltransferases from Dianthus caryophyllus[J]. Plant Biotechnology.2004,21(5):367-375.
198. Okinaka Y, Shimada Y, Nakano-Shimada R, Ohbayashi M, Kiyokawa S and Kikuchi Y. Selective accumulation of delphinidin derivatives in tobacco using a putative flavonoid 3',5'-hydroxylase cDNA from Campanula medium[J]. Biosci Biotechnol Biochem.2003,67(1): 161-165.
199. Olsen K M, Hehn A, Jugde H, Slimestad R, Larbat R, Bourgaud F and Lillo C. Identification and characterisation of CYP75A31, a new flavonoid 3',5'-hydroxylase, isolated from Solanum lycopersicum[J]. BMC Plant Biol.10:21.
200. Oud J S N, Schneiders H, Kool A J and van Grinsven M Q J M. Breeding of transgenic orange Petunia hybrida varieties [J]. Euphytica.1995,84(3):175-181.
201. Pang Y, Wenger J P, Saathoff K, Peel G J, Wen J, et al. A WD40 repeat protein from Eedicago truncatula is necessary for tissue-specific anthocyanin and proanthocyanidin biosynthesis but not for trichome development [J]. Plant Physiolgy.2009,151(3):1114-1129.
202. Park J S, Choung M G, Kim J B, Hahn B S, Kim J B, Bae S C, Roh K H, Kim Y H, Cheon C I, Sung M K and Cho K J. Genes up-regulated during red coloration in UV-B irradiated lettuce leaves [J]. Plant Cell Reports.2007a,26(4):507-516.
203. Park K-I, Ishikawa N, Morita Y, Choi J-D, Hoshino A and Iida S. A bHLH regulatory gene in the common morning glory, Ipomoea purpurea, controls anthocyanin biosynthesis in flowers, proanthocyanidin and phytomelanin pigmentation in seeds, and seed trichome formation [J]. The Plant Journal.2007b,49(4):641-654.
204. Paz-Ares J, Ghosal D, Wienand U, Peterson P A and Saedler H. The regulatory cl locus of Zea mays encodes a protein with homology to myb proto-oncogene products and with structural similarities to transcriptional activators [J]. The EMBO Journal.1987,6(12):3553-3558.
205. Pelletier M K, Murrell J R and Shirley B W. Characterization of flavonol synthase and leucoanthocyanidin dioxygenase genes in arabidopsis (further evidence for differential regulation of "Early" And "Late" Genes) [J]. Plant Physiol.1997,113(4):1437-1445.
206. Pelletier M K and Winkel-Shirley B. Analysis of flavanone 3-hydroxylase in Arabidopsis seedlings (coordinate regulation with chalcone synthase and chalcone isomerase) [J]. Plant Physiol.1996,111(1):339-345.
207. Petroni K, Cominelli E, Consonni G, Gusmaroli G, Gavazzi G and Tonelli C. The developmental expression of the maize regulatory gene hopi determines germination-dependent anthocyanin accumulation [J]. Genetics.2000,155(1):323-336.
208. Piazza P, Procissi A, Jenkins G I and Tonelli C. Members of the cl/pll regulatory gene family mediate the response of maize aleurone and mesocotyl to different light qualities and cytokinins [J]. Plant Physiology.2002,128(3):1077-1086.
209. Polashock J J, Griesbach R J, Sullivan R F and Vorsa N. Cloning of a cDNA encoding the cranberry dihydroflavonol-4-reductase (DFR) and expression in transgenic tobacco [J]. Plant science.2002,163(2):241-251.
210. Poudel P R, Mochioka R, Beppu K and Kataoka I. Influence of temperature on berry composition of interspecific hybrid wine grape'kadainou r-1'(Vitis ficifolia var. Ganebu x V. Vinifera'muscat of alexandria') [J]. Journal of the Japanese Society for Horticultural Science. 2009,78(2):169-174.
211. Prescott A G, Stamford N P J, Wheeler G and Firmin J L. In vitro properties of a recombinant flavonol synthase from Arabidopsis thaliana [J]. Phytochemistry.2002,60(6):589-593.
212. Quattrocchio F, Verweij W, Kroon A, Spelt C, Mol J and Koes R. pH4 of petunia is an R2R3 myb protein that activates vacuolar acidification through interactions with basic-helix-loop-helix transcription factors of the anthocyanin pathway [J]. Plant Cell.2006,18(5):1274-1291.
213. Quattrocchio F, Wing J, Leppen H, Mol J and Koes R. Regulatory genes controlling anthocyanin pigmentation are functionally conserved among plant species and have distinct sets of target genes [J]. Plant Cell.1993,5(11):1497-1512.
214. Quattrocchio F, Wing J, van der Woude K, Mol J and Koes R. Analysis of bHLH and MYB domain proteins: species-specific regulatory differences are caused by divergent evolution of target anthocyanin genes [J]. Plant Journal.1998,13(4):475-488.
215. Rajendran L, Ravishankar G A, Venkataraman L V and Prathiba K R. Anthocyanin production in callus cultures of Daucus carota as influenced by nutrient stress and osmoticum [J]. Biotechnology Letters.1992,14(8):707-712.
216. Ramsay N A and Glover B J. MYB-bHLH-WD40 protein complex and the evolution of cellular diversity [J]. Trends in Plant Science.2005,10(2):63-70.
217. Ramsay N A, Walker A R, Mooney M and Gray J C. Two basic-helix-loop-helix genes (MYC-146 and GL3) from arabidopsis can activate anthocyanin biosynthesis in a white-flowered Matthiola incana mutant [J]. Plant Molecular Biology.2003,52(3):679-688.
218. Reddy A M, Reddy V S, Scheffler B E, Wienand U and Reddy A R. Novel transgenic rice overexpressing anthocyanidin synthase accumulates a mixture of flavonoids leading to an increased antioxidant potential [J]. Metabolic Engineering.2007,9(1):95-111.
219. Reddy A R, Britsch L, Salamini F, Saedler H and Rohde W. The A1 (anthocyanin-1) locus in Zea mays encodes dihydroquercetin reductase [J]. Plant Science.1987,52(1-2):7-13.
220. Reif H J, Niesbach U, Deumling B and Saedler H. Cloning and analysis of two genes for chalcone synthase from Petunia hybrida [J]. Molecular and General Genetics MGG.1985,199(2): 208-215.
221. Riechmann J L and Ratcliffe O J. A genomic perspective on plant transcription factors [J]. Current Opinion in Plant Biology.2000,3(5):423-434.
222. Ronchi A, Farina G, Gozzo F and Tonelli C. Effects of a triazolic fungicide on maize plant metabolism:Modifications of transcript abundance in resistance-related pathways [J]. Plant Science.1997,130(1):51-62.
223. Rosati C, Cadic A, Duron M, Ingouff M and Simoneau P. Molecular characterization of the anthocyanidin synthase gene in Forsythia x intermedia reveals organ-specific expression during flower development [J]. Plant Science.1999,149(1):73-79.
224. Rosso M G, Li Y, Strizhov N, Reiss B, Dekker K and Weisshaar B. An Arabidopsis thaliana T-DNA mutagenized population (GABI-Kat) for flanking sequence tag-based reverse genetics[J]. Plant Molecular Biology.2003,53(1):247-259.
225. Rowan D D, Cao M S, Lin-Wang K, Cooney J M, Jensen D J, Austin P T, Hunt M B, Norling C, Hellens R P, Schaffer R J and Allan A C. Environmental regulation of leaf colour in red 35S:PAP1 Arabidopsis thaliana [J]. New Phytologist.2009,182(1):102-115.
226. Saito K, Kobayashi M, Gong Z, Tanaka Y and Yamazaki M. Direct evidence for anthocyanidin synthase as a 2-oxoglutarate-dependent oxygenase: Molecular cloning and functional expression of cdna from a red forma of Perilla frutescens [J]. The Plant Journal.1999,17(2):181-189.
227. Sakamoto W, Ohmori T, Kageyama K, Miyazaki C, Saito A, Murata M, Noda K and Maekawa M. The purple leaf (Pl)lovud of rice: The Plw allele has a complex organization and includes two genes encoding basic helix-loop-helix proteins involved in anthocyanin biosynthesis [J]. Plant Cell Physiol.2001,42(9):982-991.
228. Shih C H, Chu H, Tang L K, Sakamoto W, Maekawa M, Chu I K, Wang M and Lo C. Functional characterization of key structural genes in rice flavonoid biosynthesis[J]. Planta.2008, 228(6):1043-1054.
229. Schijlen E G W M, Ric de Vos C H, van Tunen A J and Bovy A G. Modification of flavonoid biosynthesis in crop plants [J]. Phytochemistry.2004,65(19):2631-2648.
230. Schmelzer E, Jahnen W and Hahlbrock K. In situ localization of light-induced chalcone synthase mrna, chalcone synthase, and flavonoid end products in epidermal cells of parsley leaves [J]. Proceedings of the National Academy of Sciences.1988,85(9):2989-2993.
231. Schoenbohm C, Martens S, Eder C, Forkmann G and Weisshaar B. Identification of the Arabidopsis thaliana flavonoid 3'-hydroxylase gene and functional expression of the encoded P450 enzyme[J]. Biol Chem.2000,381(8):749-753.
232. Schulze-Lefert P, Becker-Andre M, Schulz W, Hahlbrock K and Dangl J L. Functional architecture of the light-responsive chalcone synthase promoter from parsley [J]. Plant Cell.1989, 1(7):707-714.
233. Schwinn K, Venail J, Shang Y, Mackay S, Alm V, Butelli E, Oyama R, Bailey P, Davies K and Martin C. A small family of MYB-regulatory genes controls floral pigmentation intensity and patterning in the genus antirrhinum [J]. Plant Cell.2006,18(4):831-851.
234. Schwinn K E, Davies K M, Deroles S C, Markham K R, Miller R M, Bradley J M, Manson D G and Given N K. Expression of an antirrhinum majus UDP-glucose:Flavonoid-3-o-glucosyltransferase transgene alters flavonoid glycosylation and acylation in lisianthus(Eustoma grandiflorum Grise.) [J]. Plant science.1997,125(1):53-61.
235. Seitz C, Eder C, Deiml B, Kellner S, Martens S and Forkmann G. Cloning, functional identification and sequence analysis of flavonoid 3'-hydroxylase and flavonoid 3',5'-hydroxylase cdnas reveals independent evolution of flavonoid 3',5'-hydroxylase in the asteraceae family [J]. Plant Molecular Biology.2006,61(3):365-381.
236. Seitz C, Oswald N, Borstling D, Forkmann G and Martens S. Being acyanic: An unavoidable fate for many white flowers? [J]. Acta Horticulturae.2003,612:83-88.
237. Shaked-Sachray L, Weiss D, Reuveni M, Nissim-Levi A and Oren-Shamir M. Increased anthocyanin accumulation in aster flowers at elevated temperatures due to magnesium treatment [J]. Physiologia Plantarum.2002,114(4):559-565.
238. Sheehan M J, Farmer P R and Brutnell T P. Structure and expression of maize phytochrome family homeologs [J]. Genetics.2004,167(3):1395-1405.
239. Shen G, Pang Y, Wu W, Deng Z, Zhao L, Cao Y, Sun X and Tang K. Cloning and characterization of a flavanone 3-hydroxylase gene from Ginkgo biloba [J]. Bioscience Reports. 2006,26(1):19-29.
240. Shimada S, Takahashi K, Sato Y and Sakuta M. Dihydroflavonol 4-reductase cDNA from non-anthocyanin-producing species in the Caryophyllales[J]. Plant and Cell Physiology.2004, 45(9):1290.
241. Shimada Y, Nakano-Shimada R, Ohbayashi M, Okinaka Y, Kiyokawa S and Kikuchi Y. Expression of chimeric P450 genes encoding flavonoid-3',5'-hydroxylase in transgenic tobacco and petunia plants(1)[J]. FEBS Lett.1999,461(3):241-245.
242. Shiono M, Matsugaki N and Takeda K. Structure of the blue cornflower pigment: Packaging red-rose anthocyanin as part of a'superpigment'in another flower turns it brilliant blue [J]. Nature. 2005,436:791.
243. Shirley B, Kubasek W, Storz G, Bruggemann E, Koornneef M, Ausubel F and Goodman H. Analysis of Arabidopsis mutants deficient in flavonoid biosynthesis [J]. The Plant Journal.1995, 8(5):659-671.
244. Shirley B W, Hanley S and Goodman H M. Effects of ionizing radiation on a plant genome: Analysis of two arabidopsis transparent testa mutations [J]. Plant Cell.1992,4(3):333-347.
245. Shvarts M, Borochov A and Weiss D. Low temperature enhances petunia flower pigmentation and induces chalcone synthase gene expression [J]. Physiologia Plantarum.1997,99(1):67-72.
246. Smith T F, Gaitatzes C, Saxena K and Neer E J. The wd repeat: A common architecture for diverse functions [J]. Trends in Biochemical Sciences.1999,24(5):181-185.
247. Solfanelli C, Poggi A, Loreti E, Alpi A and Perata P. Sucrose-specific induction of the anthocyanin biosynthetic pathway in Arabidopsis [J]. Plant Physiology.2006,140(2):637-646.
248. Sommer H and Saedler H. Structure of the chalcone synthase gene of Antirrhinum majus[J], Molecular and General Genetics MGG.1986,202(3):429-434.
249. Sparvoli F, Martin C, Scienza A, Gavazzi G and Tonelli C. Cloning and molecular analysis of structural genes involved in flavonoid and stilbene biosynthesis in grape (Vitis vinifera L.)[J]. Plant Molecular Biology.1994a,24(5):743-755.
250. Sparvoli F, Martin C, Scienza A, Gavazzi G and Tonelli C. Cloning and molecular analysis of structural genes involved in flavonoid and stilbene biosynthesis in grape (Vitis vinifera L.)[J]. Plant Mol Biol.1994b,24(5):743-755.
251. Spelt C, Quattrocchio F, Mol J and Koes R. anthocyaninl of petunia encodes a basic helix-loop-helix protein that directly activates transcription of structural anthocyanin genes [J]. Plant Cell.2000,12(9):1619-1632.
252. Stiles E A, Cech N B, Dee S M and Lacey E P. Temperature-sensitive anthocyanin production in flowers of Plantago lanceolata [J]. Physiologia Plantarum.2007,129(4):756-765.
253. Stracke R, Werber M and Weisshaar B. The R2R3-MYB gene family in Arabidopsis thaliana [J]. Current Opinion in Plant Biology.2001,4(5):447-456.
254. Suzuki H, Nakayama T, Yamaguchi M and Nishino T. cDNA cloning and characterization of two Dendranthema× morifolium anthocyanin malonyltransferases with different functional activities[J]. Plant Science.2004a,166(1):89-96.
255. Suzuki H, Sawada S Y, Yonekura-Sakakibara K, T N, Yamaguchi M A and Nishino T. Identification of a cdna encoding malonyl-coenzyme a:Anthocyanidin 3-O-glucoside 6"-O-malonyltransferase from cineraria (Senecio cruentus) flowers [J]. Plant Biotechnology. 2003,20(3):229-234.
256. Suzuki H, Shin'ya Sawada K, Nagae S, Yamaguchi M, Nakayama T and Nishino T. Identification and characterization of a novel anthocyanin malonyltransferase from scarlet sage (Salvia splendens) flowers:an enzyme that is phylogenetically separated from other anthocyanin acyltransferases[J]. The Plant Journal.2004b,38(6):994-1003.
257. Suzuki K, Xue H, Tanaka Y, Fukui Y, Fukuchi-Mizutani M, Murakami Y, Katsumoto Y, Tsuda S and Kusumi T. Flower color modifications of Torenia hybrida by cosuppression of anthocyanin biosynthesis genes [J]. Molecular breeding:new strategies in plant improvement.2000,6(3): 239-246.
258. Tanaka Y. Flower colour and cytochromes P450 [J]. Phytochemistry Reviews.2006,5(2): 283-291.
259. Tanaka Y, Katsumoto Y, Brugliera F and Mason J. Genetic engineering in floriculture [J]. Plant Cell, Tissue and Organ Culture.2005,80(1):1-24.
260. Tanaka Y and Ohmiya A. Seeing is believing: Engineering anthocyanin and carotenoid biosynthetic pathways [J]. Current Opinion in Biotechnology.2008,19(2):190-197.
261. Tanaka Y, Sasaki N and Ohmiya A. Biosynthesis of plant pigments: anthocyanins, betalains and carotenoids [J]. The Plant Journal.2008,54(4):733-749.
262. Tanaka Y, Tsuda S and Kusumi T. Metabolic engineering to modify flower color [J]. Plant Cell Physiol.1998,39(11):1119-1126.
263. Tatsuzawa F, Yukawa T, Shinoda K, Saito N. Acylated anthocyanins in the flowers of genus Dendrobium section Phalaenanthe (Orchidaceae) [J]. Biochemical Systematics and Ecology. 2005,33(6):625-629.
264. Taylor L P and Briggs W R. Genetic regulation and photocontrol of anthocyanin accumulation in maize seedlings [J]. Plant Cell.1990,2(2):115-127.
265. Togami J, Tamura M, Ishiguro K, Hirose C, Okuhara H, Ueyama Y, Nakamura N, Yonekura-Sakakibara K, Fukuchi-Mizutani M and Suzuki K. Molecular characterization of the flavonoid biosynthesis of Verbena hybrida and the functional analysis of verbena and Clitoria ternatea F3'5'H genes in transgenic verbena[J]. Plant Biotechnology.2006,23(1):5-11.
266. Toguri T, Umemoto N, Kobayashi O and Ohtani T. Activation of anthocyanin synthesis genes by white light in eggplant hypocotyl tissues, and identification of an inducible P-450 cDNA [J]. Plant Molecular Biology.1993,23(5):933-946.
267. Torskangerpoll K, Nφbaeb R, Nodland E,φvstedal D O, Andersen φ M. Anthocyanin content of Tulipa species and cultivars and its impact on tepal colours[J]. Biochemical Systematics and Ecology,2005,33(5):499-510.
268. Toyama-Kato Y, Yoshida K, Fujimori E, Haraguchi H, Shimizu Y and Kondo T. Analysis of metal elements of hydrangea sepals at various growing stages by ICP-AES [J]. Biochemical Engineering Journal.2003,14(3):237-241.
269. Tunen A J v, Koes R E, Spelt C E, Krol A R v d, Stuitje A R and Mol J N. Cloning of the two chalcone flavanone isomerase genes from Petunia hybrida:coordinate, light-regulated and differential expression of flavonoid genes[J]. The EMBO Journal.1988,7(5):1257-1263.
270. Turnbull J J, Nakajima J-i, Welford R W D, Yamazaki M, Saito K and Schofield C J. Mechanistic studies on three 2-oxoglutarate-dependent oxygenases of flavonoid biosynthesis [J]. Journal of Biological Chemistry.2004,279(2):1206-1216.
271. Ueyama Y, Suzuki K-i, Fukuchi-Mizutani M, Fukui Y, Miyazaki K, Ohkawa H, Kusumi T and Tanaka Y. Molecular and biochemical characterization of torenia flavonoid 3'-hydroxylase and flavone synthase Ⅱ and modification of flower color by modulating the expression of these genes [J]. Plant Science.2002,163(2):253-263.
272. van der Krol A R, Mur L A, Beld M, Mol J N M and Stuitje A R. Flavonoid genes in petunia: addition of a limited number of gene copies may lead to a suppression of gene expression [J]. the Plant Cell.1990,2(4):291-299.
273. van Houwelingen A, Souer E, Spelt K, Kloos D, Mol J and Koes R. Analysis of flower pigmentation mutants generated by random transposon mutagenesis in Petunia hybrida [J]. The Plant Journal.1998,13(1):39-50.
274. van Tunen A J, Hartman S A, Mur L A and Mol J N M. Regulation of chalcone flavanone isomerase (CHI) gene expression in Petunia hybrida:The use of alternative promoters in corolla, anthers and pollen [J]. Plant molecular biology.1989,12(5):539-551.
275. van Tunen A J, Koes R E, Spelt C E, van der Krol A R, Stuitje A R and Mol J N. Cloning of the two chalcone flavanone isomerase genes from Petunia hybrida:Coordinate, light-regulated and differential expression of flavonoid genes [J]. The EMBO Journal.1988,7(5):1257-1263.
276. van Tunen A J, Mur L A, Brouns G S, Rienstra J D, Koes R E and Mol J. Pollen- and anther-specific chi promoters from petunia:Tandem promoter regulation of the chiA gene [J]. Plant Cell.1990,2(5):393-401.
277. Walker A R, Davison P A, Bolognesi-Winfield A C, James C M, Srinivasan N, Blundell T L, Esch J J, Marks M D and Gray J C. The TRANSPARENT TESTA GLABRA1 locus, which regulates trichome differentiation and anthocyanin biosynthesis in Arabidopsis, encodes a wd40 repeat protein [J]. Plant Cell.1999,11(7):1337-1350.
278. Wang H M and To K Y. Agrobacterium-mediated transformation in the high-value medicinal plant Echinacea purpurea [J]. Plant Science.2004,166(4):1087-1096.
279. Wang J, Ming F, Han Y and Shen D. Flavonoid-3',5'-hydroxylase from Phalaenopsis: A novel member of cytochrome P450s, its cDNA cloning, endogenous expression and molecular modeling [J]. Biotechnology Letters.2006,28(5):327-334.
280. Wang L S, Shiraishi A, Hashimoto F, Aoki N, Shimizu K and Sakata Y. Analysis of Petal Anthocyanins to investigate flower coloration of Zhongyuan (Chinese) and Daikon Island (Japanese) tree peony cultivars. Journal of Plant Research [J].2001,114(1):33-43.
281. Weiss D and Halevy A H. The role of light reactions in the regulation of anthocyanin synthesis in Petunia corollas [J]. Physiologia Plantarum.1991,81(1):127-133.
282. Weiss D, van Blokland R, Kooter J M, Mol J N M and van Tunen A J. Gibberellic acid regulates chalcone synthase gene transcription in the corolla of Petunia hybrida [J]. Plant Physiology.1992, 98(1):191-197.
283. Whittall J, Voelckel C, Kliebenstein D and Hodges S. Convergence, constraint and the role of gene expression during adaptive radiation:floral anthocyanins in Aquilegia[J]. Molecular Ecology.2006,15(14):4645-4657.
284. Winkel-Shirley B. Flavonoid biosynthesis. A colorful model for genetics, biochemistry, cell biology, and biotechnology [J]. Plant Physiol.2001,126(2):485-493.
285. Winkel-Shirley B. Biosynthesis of flavonoids and effects of stress [J]. Current Opinion in Plant Biology.2002,5(3):218-223
286. Xu F, Cheng H, Cai R, Li L, Chang J, Zhu J, Zhang F, Chen L, Wang Y, Cheng S and Cheng S Y. Molecular cloning and function analysis of an anthocyanidin synthase gene from Ginkgo biloba, and its expression in abiotic stress responses [J]. Molecules and Cells.2008,26(6): 536-547.
287. Yamagishi M, Shimoyamada Y, Nakatsuka T and Masuda K. 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(3):463-474.
288. Yamazaki M, Makita Y, Springob K and Saito K. Regulatory mechanisms for anthocyanin biosynthesis in chemotypes of Perilla frutescens var. Crispa [J]. Biochemical Engineering Journal. 2003,14(3):191-197.
289. Yang J, Huang J, Gu H, Zhong Y and Yang Z. Duplication and adaptive evolution of the chalcone synthase genes of Dendranthema (Asteraceae)[J]. Molecular biology and evolution. 2002,19(10):1752-1759.
290. Yang S, Sweetman J P, Amirsadeghi S, Barghchi M, Huttly A K, Chung W-I and Twell D. Novel anther-specific myb genes from tobacco as putative regulators of phenylalanine ammonia-lyase expression [J]. Plant Physiol.2001,126(4):1738-1753.
291. Yoshida K, Kitahara S, Ito D and Kondo T. Ferric ions involved in the flower color development of the himalayan blue poppy, meconopsis grandis [J]. Phytochemistry.2006,67(10):992-998.
292. Yoshihara N, Imayama T, Matsuo Y, Fukuchi-Mizutani M, Tanaka Y, Ino I and Yabuya T. Characterization of cDNA clones encoding anthocyanin 3-p-coumaroyltransferase from Iris hollandica[J]. Plant Science.2006,171(5):632-639.
293. Zhou B, Li Y, Xu Z, Yan H, Homma S and Kawabata S. Ultraviolet a-specific induction of anthocyanin biosynthesis in the swollen hypocotyls of turnip(Brassica rapa) [J]. Journal of Experimental Botany.2007,58(7):1771-1781.
294. Zhou W, Huang C, Gong Y, Feng Q and Gao F. Molecular cloning and expression analysis of an ans gene encoding anthocyanidin synthase from purple-fleshed sweet potato [Ipomoea batatas (L.) Lam] [J]. Plant Molecular Biology Reporter.2010,28(1):112-121.
295. Zufall R A and Rausher M D. The Genetic Basis of a Flower Color Polymorphism in the Common Morning Glory (Ipomoea purpurea)[J]. Journal of Heredity.2003,94(6):442-448.
296. Zuker A, Tzfira T, Ben-Meir H, Ovadis M, Shklarman E, Itzhaki H, Forkmann G, Martens S, Neta-Sharir I, Weiss D and Vainstein A. Modification of flower color and fragrance by antisense suppression of the flavanone 3-hydroxylase gene [J]. Molecular Breeding 2002,9(1):33-41.
2.白新祥,戴思兰.蓝色花观赏植物及其在园林中的应用.张启翔.中国观赏园艺研究进展2004[M].北京：中国林业出版社,2004：69-75.
3.戴思兰.中国菊花与世界园艺[J].河北科技师范学院学报,2004,18(2)：1-5,9.
4.韩科厅,胡可,戴思兰.观赏植物花色的分子设计[J].分子植物育种,2008,6(1)：16-24.
5.胡可,韩科厅,戴思兰.环境因子调控植物花青素合成及呈色的机理[J].植物学报,2010,45(3)：307-317.
6.胡可,孟丽,韩科厅,孙翊,戴思兰.瓜叶菊花青素合成关键结构基因的分离及表达分析[J].园艺学报,2009,36(7)：1013-1022.
7.黄金霞,王亮生,李晓梅,鲁迎青.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.
8.冷欣夫,邱星辉主编.细胞色素P450酶系的结构,功能与应用前景[M].北京：科学出版社,2001：107-128.
9.李鸿渐,邵建文.中国菊花品种资源的调查收集与分类[J].南京农业大学,1990,13(1)：30-36.
10.刘明,赵琦,王小菁,赵玉锦,童哲.植物的光受体及其调控机制的研究[J].生物学通报,2005,40(5)：10-12.
11.马月萍,戴思兰.高等植物成花分子机理的研究进展[J].分子植物育种,2007,5(z1)：21-28.
12.孟丽.蓝色花形成关键基因的分离及其表达分析[M].北京林业大学博士论文,2006.
13.孟丽,戴思兰.瓜叶菊F3’5’H基因cDNA的克隆、序列分析及其原核表达[J].分子植物育种,2005,3(6)：780-786.
14.孟丽,周琳,张明姝,戴思兰.一种有效的花瓣总RNA的提取方法[J].生物技术,2006,16(1)：38-40.
15.孙明霞,王宝增,范海,赵可夫.叶片中的花色素苷及其对植物适应环境的意义[J].植物生理学通讯,2003,39(6)：688-694.
16.孙卫,李崇晖,王亮生,戴思兰,徐彦军.花青素苷成分对瓜叶菊花色的影响[J].园艺学报,2009,36(12)：1775-1782.
17.孙卫,李崇晖,王亮生,戴思兰.菊花不同花色品种中花青素苷代谢分析[J].植物学报,2010,45(3)：327-336
18.田义,王强,张利义,康国栋,杨玲,郝红梅,杨振英,丛佩华.外源腐胺促进苹果果皮花青素苷积累的效应[J].植物学报,2009,44(3)：310-316.
19.王曼,王小菁.蓝光和蔗糖对拟南芥花色素苷积累和CHS基因表达的影响[J].热带亚热带植物学报,2004,12(3)：252-256.
20.徐清燏,戴思兰.蓝色花卉分子育种[J].分子植物育种,2004,2(1)：93-99.
21.许志茹,李玉花.‘津田芜菁’花色素苷生物合成相关基因的表达.植物生理与分子生物学学报32,583-586[J].植物生理学通讯,2006,44(5)：931-935.
22.杨致荣,王兴春,李西明,杨长登.高等植物转录因子的研究进展[J].遗传,2004,26(3)：403-408.
23.张剑亮,潘大仁,周以飞,吴明伟,柯祥德,林钿.PCR介导观赏向日葵DFR基因改造及其真核表达载体构建研究初报[J].福建农林大学学报,2008,37(2)：166-169.
24.张明姝.菊花查尔酮合成酶基因的克隆与序列分析[M].北京林业大学博士论文,2006.
25.张树林.菊花品种分类的研究[J].园艺学报,1965,4(1)：35-46.
26. Aharoni A, Vos C H R D, Wein M, Sun Z, Greco R, Kroon A, Mol J N M and O'Connell A P. The strawberry FaMYBl transcription factor suppresses anthocyanin and flavonol accumulation in transgenic tobacco [J]. The Plant Journal.2001,28(3):319-332.
27. Aida R, Kishimoto S, Tanaka Y and Shibata M. Modification of flower color in torenia (Torenia fournieri Lind.) by genetic transformation [J]. Plant Science.2000,153(1):33-42.
28. Akashi T, Ishizaki M, Aoki T and Ayabe S. Isoflavonoid production by adventitious-root cultures of Iris germanica (Iridaceae)[J]. Plant Biotechnology.2005,22(3):207-215.
29. Akira N, Yoko I and Masumi Y. Spatial and temporal expression of chalcon e synthase and dihydroflavonol 4-reductase genes in the Asiatic hybrid lily [J]. Plant science.2003,165(4): 759-767.
30. Albert N W, Lewis D H, Zhang H, Irving L J, Jameson P E and Davies K M. Light-induced vegetative anthocyanin pigmentation in Petunia [J]. Journal of Experimental Botany.2009: erp097.
31. Alfenito M R, Souer E, Goodman C D, Buell R, Mol J, Koes R and Walbot V. Functional complementation of anthocyanin sequestration in the vacuole by widely divergent glutathione S-transferases [J]. Plant Cell.1998,10(7):1135-1150.
32. Alla M M N and Younis M E. Herbicide effects on phenolic metabolism in maize (Zea mays L.)and soybean (Glycine max L.) seedling [J]. Journal of Experimental Botany.1995,46(11): 1731-1736.
33. Almeida J, Carpenter R, Robbins T P, Martin C and Coen E S. Genetic interactions underlying flower color patterns in Antirrhinum majus [J]. Genes & Development.1989,3(11):1758-1767.
34. Alvarez I, Costa A and Feliner G. Selecting Single-Copy Nuclear Genes for Plant Phylogenetics: A Preliminary Analysis for the Senecioneae (Asteraceae)[J]. Journal of Molecular Evolution. 2008,66(3):276-291.
35. Ashihara H, Deng W W, Mullen W and Crozier A. Distribution and biosynthesis of flavan-3-ols in Camellia sinensis seedlings and expression of genes encoding biosynthetic enzymes[J]. Phytochemistry.71(5-6):559-566.
36. Ban Y, Honda C, Hatsuyama Y, Igarashi M, Bessho H and Moriguchi T. Isolation and functional analysis of a MYB transcription factor gene that is a key regulator for the development of red coloration in apple skin [J]. Plant Cell Physiol.2007,48(7):958-970.
37. Banon S, Gonzalez A, Cano E A, Franco J A, Fernandez J A. Growth, development and colour response of potted Dianthus caryophyllus cv. Mondriaan to paclobutrazol treatment[J]. Scienta horticulturae.2002,94(3-4):371-377.
38. Barb J G, Werner D J and Griesbach R J. Genetics and biochemistry of flower color in stokes aster [J]. Journal of the American Society for Horticultural Science.2008,133(4):569-578.
39. Beld M, Martin C, Huits H, Stuitje A R and Gerats A G M. Flavonoid synthesis in Petunia hybrida:Partial characterization of dihydroflavonol-4-reductase genes [J]. Plant Molecular Biology.1989,13(5):491-502.
40. Biolley J P, Jay M. Anthocyanins in modern roses:chemical and colorimetric features in relation to the colour range[J]. Journal of Experimental Botany.1993,44(268):1725-1734.
41. Biran I and Halevy A H. Effects of varying light intensities and temperature treatments applied to whole plants, or locally to leaves or flower buds, on growth and pigmentation of'Baccara'roses [J]. Physiologia Plantarum.1974,31(3):175-179.
42. Boddu J, Svabek C, Sekhon R, Gevens A, Nicholson R L, Jones A D, Pedersen J F, Gustine D L and Chopra S. Expression of a putative flavonoid 3'-hydroxylase in sorghum mesocotyls synthesizing 3-deoxyanthocyanidin phytoalexins [J]. Physiological and Molecular Plant Pathology.2004,65(2):101-113.
43. Bogs J, Jaffe F W, Takos A M, Walker A R and Robinson S P. The grapevine transcription factor VvMYBPA1 regulates proanthocyanidin synthesis during fruit development [J]. Plant Physiology. 2007,143(3):1347-1361.
44. Bolwell G P, Bozak K and Zimmerlin A. Plant cytochrome P450 [J]. Phytochemistry.1994, 37(6):1491-1506.
45. Bongue-Bartelsman M, O'Neill S D, Tong Y and Yoder J I. Characterization of the gene encoding dihydroflavonol 4-reductase in tomato[J]. Gene.1994,138(1-2):153-157.
46. Borevitz J O, Xia Y, Blount J, Dixon R A and Lamb C. Activation tagging identifies a conserved MYB regulator of phenylpropanoid biosynthesis [J]. Plant Cell.2000,12(12):2383-2394.
47. Boss P K, Davies C and Robinson S P. Anthocyanin composition and anthocyanin pathway gene expression in grapevine sports differing in berry skin colour [J]. Australian Journal of Grape and Wine Research.1996,2(3):163-170.
48. Bradley M J, Davies K M, Deroles S C, Bloor S J and Lewis D H. The maize Lc regulatory gene up-regulates the flavonoid biosynthetic pathway of Petunia [J]. The Plant Journal.1998,13(3): 381-392.
49. Britsch L, Ruhnau-Brich B and Forkmann G. Molecular cloning, sequence analysis, and in vitro expression of flavanone 3 beta-hydroxylase from Petunia hybrida [J]. Journal of Biological Chemistry.1992,267(8):5380-5387.
50. Broun P. Transcriptional control of flavonoid biosynthesis: A complex network of conserved regulators involved in multiple aspects of differentiation in Arabidopsis [J]. Current Opinion in Plant Biology.2005,8(3):272-279.
51. Brugliera F, Barri-Rewell G, Holton T A and Mason J G. Isolation and characterization of a flavonoid 3'-hydroxylase cDNA clone corresponding to the htl locus of Petunia hybrida [J]. The Plant Journal.1999,19(4):441-451.
52. Carey C C, Strahle J T, Selinger D A and Chandler V L. Mutations in the pale aleurone color1 regulatory gene of the zea mays anthocyanin pathway have distinct phenotypes relative to the functionally similar transparent testa glabral gene in Arabidopsis thaliana [J]. Plant Cell.2004, 16(2):450-464.
53. Castellarin S, Di Gaspero G, Marconi R, Nonis A, Peterlunger E, Paillard S, Adam-Blondon A and Testolin R. Colour variation in red grapevines(Vitis vinifera L.):genomic organisation, expression of flavonoid 3'-hydroxylase, flavonoid 3',5'-hydroxylase genes and related metabolite profiling of red cyanidin-/blue delphinidin-based anthocyanins in berry skin[J]. BMC Genomics. 2006,7:12.
54. Castellarin S and Di Gaspero G. Transcriptional control of anthocyanin biosynthetic genes in extreme phenotypes for berry pigmentation of naturally occurring grapevines [J]. BMC Plant Biology.2007,7(1):46.
55. Chalker-Scott L. Environmental significance of anthocyanins in plant stress responses [J]. Photochemistry and Photobiology.1999,70(1):1-9.
56. Chandler V L, Radicella J P, Robbins T P, Chen J and Turks D. Two regulatory genes of the maize anthocyanin pathway are homologous:Isolation of B utilizing R genomic sequences [J]. Plant Cell.1989,1(12):1175-1183.
57. Charrier B, Coronado C, Kondorosi A and Ratet P. Molecular characterization and expression of alfalfa (Medicago sativa L.) flavanone-3-hydroxylase and dihydroflavonol-4-reductase encoding genes [J]. Plant Molecular Biology.1995,29(4):773-786.
58. Chatterjee M, Sharma P and Khurana J P. Cryptochrome 1 from Brassica napus is up-regulated by blue light and controls hypocotyl/stem growth and anthocyanin accumulation [J]. Plant Physiology.2006,141(1):61-74.
59. Cheng L, Xu Y, Grotewold E, Jin Z, Wu F, Fu C and Zhao D. Characterization of Anthocyanidin Synthase (ANS) Gene and anthocyanidin in rare medicinal plant-Saussurea medusa[J]. Plant Cell, Tissue and Organ Culture.2007,89(1):63-73.
60. Cho S, Chen W and Muehlbauer F J. Constitutive expression of the Flavanone 3-hydroxylase gene related to pathotype-specific ascochyta blight resistance in Cicer arietinum L.[J]. Physiological and Molecular Plant Pathology.2005,67(2):100-107.
61. Christie J M and Jenkins G I. Distinct UV-B and UV-A/blue light signal transduction pathways induce chalcone synthase gene expression in Arabidopsis cells [J]. Plant Cell.1996,8(9): 1555-1567.
62. Christie P, Alfenito M and Walbot V. Impact of low-temperature stress on general phenylpropaniod and anthocyanin pathways:enhancement of transcript abundance and anthocyanin pigmentation in maize seedlings [J]. Planta.1994,194(4):541-549.
63. Cominelli E, Gusmaroli G, Allegra D, Galbiati M, Wade H K, Jenkins G I and Tonelli C. Expression analysis of anthocyanin regulatory genes in response to different light qualities in Arabidopsis thaliana [J]. Journal of Plant Physiology.2008,165(8):886-894.
64. Cone K C, Burr F A and Burr B. Molecular analysis of the maize anthocyanin regulatory locus C1 [J]. Proceedings of the National Academy of Sciences of the United States of America.1986, 83(24):9631-9635.
65. Cone K C, Cocciolone S M, Burr F A and Burr B. Maize anthocyanin regulatory gene pl is a duplicate of cl that functions in the plant [J]. Plant Cell.1993,5(12):1795-1805.
66. Courtney-Gutterson N, Napoli C, Lemieux C, Morgan A, Firoozabady E and Robinson K E P. Modification of flower color in florist's chrysanthemum:Production of a white-flowering variety through molecular genetics [J]. Nat Biotech.1994,12(3):268-271.
67. David W. Regulation of flower pigmentation and growth: Multiple signaling pathways control anthocyanin synthesis in expanding petals [J]. Physiologia Plantarum.2000,110(2):152-157.
68. Davies K M. A cDNA clone for flavanone 3-hydroxylase from Malus[J]. Plant Physiol.1993, 103(1):291.
69. Davies K M, Schwinn K E, Deroles S C, Manson D G, Lewis D H, Bloor S J and Bradley J M. Enhancing anthocyanin production by altering competition for substrate between flavonol synthase and dihydroflavonol 4-reductase [J]. Euphytica.2003,131(3):259-268.
70. de Vetten N, Quattrocchio F, Mol J and Koes R. The anll locus controlling flower pigmentation in petunia encodes a novel WD-repeat protein conserved in yeast, plants, and animals [J]. Genes & Development.1997,11(11):1422-1434.
71. Deboo G B, Albertsen M C and Taylor L P. Flavanone 3-hydroxylase transcripts and flavonol accumulation are temporally coordinate in maize anthers [J]. The Plant Journal.1995,7(5): 703-713.
72. Deroles S C, Bradley J M, Schwinn K E, Markham K R, Bloor S, Manson D G and Davies K M. An antisense chalcone synthase cDNA leads to novel colour patterns in lisianthus(Eustoma grandiflorum) flowers [J]. Molecular Breeding.1998,4(1):59-66.
73. Dixon R A, Harrison M J and Lamb C J. Early events in the activation of plant defense responses [J]. Annual Review of Phytopathology.1994,32(1):479-501.
74. Dixon R A and Steele C L. Flavonoids and isoflavonoids- a gold mine for metabolic engineering [J]. Trends in Plant Science.1999,4(10):394-400.
75. Dong Y H, Beuning L, Davies K, Mitra D, Morris B and Kootstra A. Expression of pigmentation genes and photo-regulation of anthocyanin biosynthesis in developing royal gala apple flowers [J]. Australian Journal of Plant Physiology.1998,25(2):245-252.
76. Dooner H K. Coordinate genetic regulation of flavonoid biosynthetic enzymes in maize [J]. Molecular and General Genetics MGG.1983,189(1):136-141.
77. Druka A, Kudrna D, Rostoks N, Brueggeman R, von Wettstein D and Kleinhofs A. Chalcone isomerase gene from rice (Oryza sativa) and barley(Hordeum vulgare): physical, genetic and mutation mapping[J]. Gene.2003,302(1-2):171-178.
78. Dubos C, Gourrierec J L, Baudry A, Huep G, Lanet E, Debeaujon I, Routaboul J M, Alboresi A, Weisshaar B and Lepiniec L. MYBL2 is a new regulator of flavonoid biosynthesis in Arabidopsis thaliana [J]. The Plant Journal.2008,55(6):940-953.
79. Durbin M L, McCaig B and Clegg M T. Molecular evolution of the chalcone synthase multigene family in the morning glory genome [J]. Plant Molecular Biology.2000,42(1):79-92.
80. Elomaa P, Honkanen J, Puska R, Seppanen P, Helariutta Y, Mehto M, Kotilainen M, Nevalainen L and Teeri T H. Agrobacterium-mediated transfer of antisense chalcone synthase cDNA to Gerbera hybrida inhibits flower pigmentation [J]. Nature Biotechnology 1993,11:508-511.
81. Elomaa P, Mehto M, Kotilainen M, Yrjo Helariutta, Nevalainen L and Teeri T H. A bHLH transcription factor mediates organ, region and flower type specific signals on dihydroflavonol-4-reductase (dfr) gene expression in the inflorescence of Gerbera hybrida (Asteraceae) [J]. The Plant Journal.1998,16(1):93-99.
82. Epping B, Kittel M, Ruhnau B and Hemleben V. Isolation and sequence analysis of a chalcone synthase cDNA of Matthiola incana R. Br. (Brassicaceae)[J]. Plant Molecular Biology.1990, 14(6):1061-1063.
83. Espley R, Hellens R, Putterill J, Stevenson D, Kutty-Amma S and Allan A. Red colouration in apple fruit is due to the activity of the MYB transcription factor, MdMYB10 [J]. Plant J.2007, 49(3):414-427.
84. Farzad M, Griesbach R, Hammond J, Weiss M R and Elmendorf H G. Differential expression of three key anthocyanin biosynthetic genes in a color-changing flower, Viola cornuta cv. Yesterday, Today and Tomorrow [J]. Plant Science.2003,165(6):1333-1342.
85. Feldbrugge M, Sprenger M, Hahlbrock K and Weisshaar B. PcMYB1, a novel plant protein containing a DNA-binding domain with one MYB repeat, interacts in vivo with a light-regulatory promoter unit [J]. Plant Journal.1997,11(5):1079-1093.
86. Feller A, Hernandez J M and Grotewold E. An ACT-like domain participates in the dimerization of several plant basic-helix-loop-helix transcription factors [J]. Journal of Biological Chemistry. 2006,281(39):28964-28974.
87. Ferrer J L, Jez J M, Bowman M E, Dixon R A and Noel J P. Structure of chalcone synthase and the molecular basis of plant polyketide biosynthesis [J]. Nature Structural Biology.1999,6(8): 775-784.
88. Forkmann G and Martens S. Metabolic engineering and applications of flavonoids [J]. Current Opinion in Biotechnology.2001,12(2):155-160.
89. Forkmann G and Ruhnau B. Distinct substrate specificity of dihydroflavonol 4-reductase from flowers of Petunia hybrida [J]. Zeitschrift fur Naturforschung.1987,42:1146-1148.
90. Fukada-Tanaka S, Hoshino A, Hisatomi Y, Habu Y, Hasebe M and Iida S. Identification of New Chalcone Synthase Genes for Flower Pigmentation in the Japanese and Common Morning Glories[J]. Plant Cell Physiol.1997,38(6):754-758.
91. Fukui Y, Tanaka Y, Kusumi T, Iwashita T and Nomoto K. A rationale for the shift in colour towards blue in transgenic carnation flowers expressing the flavonoid 3',5'-hydroxylase gene [J]. Phytochemistry 2003,63(1):15-23.
92. Fukusaki E-i, Kawasakia K, Kajiyamaa S-i, Ana C I, Suzukib K, Tanakac Y and Kobayashia A. Flower color modulations of Torenia hybrida by downregulation of chalcone synthase genes with ma interference [J]. Journal of Biotechnology 2004,111(3):229-240.
93. Gerats A, Huits H, Vrijlandt E, Marana C, Souer E and Beld M. Molecular characterization of a nonautonomous transposable element (dTph1) of petunia [J]. Plant Cell.1990,2(11):1121-1128.
94. Goff S A, Cone K C and 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 and Development.1992,6(5):864-875.
95. Goff S A, Klein T M, Roth B A, Fromm M E, Cone K C, Radicella J P and Chandler V L. Transactivation of anthocyanin biosynthetic genes following transfer of B regulatory genes into maize tissues [J]. The EMBO Journal.1990,9(8):2517-2522.
96. Goldsbrough A P, Tong Y and Yoder J I. Lc as a non-destructive visual reporter and transposition excision marker gone for tomato [J]. The Plant Journal.1996,9(6):927-933.
97. Gong Z, Yamazaki M, Sugiyama M, Tanaka Y and Saito K. Cloning and molecular analysis of structural genes involved in anthocyanin biosynthesis and expressed in a forma-specific manner in Perilla frutescens [J]. Plant Molecular Biology.1997,35(6):915-927.
98. Gong Z Z, Yamagishi E, Yamazaki M and Saito K. A constitutively expressed Myc-like gene involved in anthocyanin biosynthesis from Perilla frutescens: molecular characterization, heterologous expression in transgenic plants and transactivation in yeast cells [J]. Plant Molecular Biology.1999,41(1):33-44.
99. Gonnet J F. CIElab measurement, a precies communication in flower colour: An example with carnation (Dianthus caryophyllus) cultivars [J]. Journal of horticultural science.1993,68(4): 499-510.
100. Goodrich J, Carpenter R and Coen E S. A common gene regulates pigmentation pattern in diverse plant species [J]. Cell.1992,68(5):955-964.
101. Goto T, Kondo T, Kawai T and Tamura H. Structure of cinerarin, a tetra-acylated anthocyanin isolated from the blue garden cineraria, Senecio cruentus [J]. Tetrahedron letters.1984,25(52): 6021-6024.
102. Griesbach R J. Correlation of pH and light intensity on flower color in potted Eustoma grandiflorum Grise [J]. HortScience.1992,27(7):817-818.
103. Grotewold E. The genetics and biochemistry of floral pigments [J]. Annual Review of Plant Biology.2006,57(1):761-780.
104. Grotewold E and Peterson T. Isolation and characterization of a maize gene encoding chalcone flavonone isomerase[J]. Molecular and General Genetics MGG.1994,242(1):1-8.
105. Grotewold E, Sainz M B, Tagliani L, Hernandez J M, Bowen B and Chandler V L. Identification of the residues in the Myb domain of maize C1 that specify the interaction with the bHLH cofactor R [J]. Proceedings of the National Academy of Sciences of the United States of America. 2000,97(25):13579-13584.
106. Guo J, Han W and Wang M H. Ultraviolet and environmental stresses involved in the induction and regulation of anthocyanin biosynthesis: A review [J]. African Journal of Biotechnology.2008, 7(25):4966-4972.
107. Gutterson N C, Napoli C, Lemieux C, Morgan A, Firoozabady E and Robinson K E. Modification of flower color in florist's chrysanthemum: production of a white-genetics [J]. Nature Biotechnology.1994,12:268-271.
108. Halevy A. Manipulation of flowering[M]. London: Butterworth,1987:363-378.
109. Han Y, Ming F, Wang J, Ye M and Shen D. A novel chalcone synthase gene from Phalaenopsis orchid that alters floral morphology in transgenic tobacco plants [J]. Plant Molecular Biology Reporter.2005,23:193a-193m.
110. Harborne J B and Williams C A. Advances in flavonoid research since 1992 [J]. Phytochemistry. 2000,55(6):481-504.
111. Hartmann U, Sagasser M, Mehrtens F, Stracke R and Weisshaar B. Differential combinatorial interactions of cis-acting elements recognized by R2R3-MYB, BZIP, and BHLH factors control light-responsive and tissue-specific activation of phenylpropanoid biosynthesis genes [J]. Plant Molecular Biology.2005,57(2):155-171.
112. Hartmann U, Valentine W J, Christie J M, Hays J, Jenkins G I and Weisshaar B. Identification of UV/blue light-response elements in the Arabidopsis thaliana chalcone synthase promoter using a homologous protoplast transient expression system [J]. Plant Molecular Biology.1998,36(5): 741-754.
113. Helariutta Y, Elomaa P, Kotilainen M, Griesbach R J, Schroder J and Teeri T H. Chalcone synthase-like genes active during corolla development are differentially expressed and encode enzymes with different catalytic properties in Gerbera hybrida (Asteraceae)[J]. Plant Molecular Biology.1995,28(1):47-60.
114. Helariutta Y, Elomaa P, Kotilainen M, Seppanen P and Teeri T H. Cloning of cDNA coding for dihydroflavonol-4-reductase (DFR) and characterization of dfr expression in the corollas of Gerbera hybrida var. Regina (Compositae) [J]. Plant Molecular Biology.1993,22(2):183-193.
115. Heller W, Forkmann G, Britsch L and Grisebach H. Enzymatic reduction of (+)-dihydroflavonols to flavan-3,4-cis-diols with flower extracts from Matthiola incana and its role in anthocyanin biosynthesis [J]. Planta.1985,165(2):284-287.
116. Hirner A A and Seitz H U. Isoforms of chalcone synthase in Daucus carota L. And their differential expression in organs from the european wild carrot and in ultraviolet-A-irradiated cell cultures [J]. Planta.2000,210(6):993-998.
117. Hisatomi Y, Yoneda Y, Kasahara K, Inagaki Y and Iida S. DNA rearrangements at the region of the dihydroflavonol 4-reductase gene for flower pigmentation and incomplete dominance in morning glory carrying the mutable flaked mutation[J]. TAG Theoretical and Applied Genetics. 1997,95(4):509-515.
118. Holton T and Cornish E. Genetics and biochemistry of anthocyanin biosynthesis [J]. Plant Cell. 1995,7(7):1071-1083.
119. Hondo T, Yoshida K, Nakagawa A, Kawai T, Tamura H and Goto T. Structural basis of blue-colour development in flower petals from Commelina communis [J]. Nature.1992,358: 515-518.
120. Hoshino A, Johzuka-Hisatomi Y and Iida S. Gene duplication and mobile genetic elements in the morning glories [J]. Gene.2001,265(1-2):1-10.
121. Hoshino A, Abe Y, Saito N, Inagaki Y and Iida S. The Gene Encoding Flavanone 3-Hydroxylase is Expressed Normally in the Pale Yellow Flowers of the Japanese Morning Glory Carrying the speckled Mutation Which Produce Neither Flavonol nor Anthocyanin but Accumulate Chalcone, Aurone and Flavanone[J]. Plant Cell Physiol.1997,38(8):970-974.
122. Hoshino A, Morita Y, Choi J D, Saito N, Toki K, Tanaka Y and Iida S. Spontaneous mutations of the flavonoid 3'-hydroxylase gene conferring reddish flowers in the three morning glory species[J]. Plant and Cell Physiology.2003,44(10):990-1001.
123. Hrazdina G and Jensen R A. Spatial organization of enzymes in plant metabolic pathways [J]. Annual Review of Plant Physiology and Plant Molecular Biology.1992,43(1):241-267.
124. Hrazdina G, Marx G A and Hoch H C. Distribution of secondary plant metabolites and their biosynthetic enzymes in pea (Pisum sativum L.) leaves:anthocyanins and flavonol glycosides [J]. Plant Physiolgy.1982,70(3):745-748.
125. Huang Z, Liang M, Peng J, Xing T and Wang X. Exogenous ammonium inhibits petal pigmentation and expansion in Gerbera hybrida [J]. Physiologia Plantarum.2008,133(2): 254-265.
126. Huh E, Shin H, Choi S, Kwon O and Lee Y. Thermosusceptible developmental stage in anthocyanin accumulation and color response to high temperature in red chrysanthemum cultivars [J]. Korean Journal of Horticultural Science and Technology.2008,26(4):357-361.
127. Huits H S M, Gerats A G M, Kreike M M, Mol J N M and Koes R E. Genetic control of dihydroflavonol 4-reductase gene expression in Petunia hybrida [J]. The Plant Journal.1994,6(3): 295-310.
128. Inagaki Y, Johzuka-Hisatomi Y, Mori T, Takahashi S, Hayakawa Y, Peyachoknagul S, Ozeki Y and Iida S. Genomic organization of the genes encoding dihydroflavonol 4-reductase for flower pigmentation in the Japanese and common morning glories [J]. Gene.1999,226(2):181-188.
129. Islam M S, Jalaluddin M, Garner J O, Yoshimoto M and Yamakawa O. Artificial shading and temperature influence on anthocyanin compositions in sweetpotato leaves [J]. HortScience.2005, 40:176-180.
130. Jaakola L, Maatta K, Pirttila A M, Torronen R, Karenlampi S and Hohtola A. Expression of genes involved in anthocyanin biosynthesis in relation to anthocyanin, proanthocyanidin, and flavonol levels during bilberry fruit development [J]. Plant Physiology.2002,130(2):729-739.
131. Jackson D, Culianez-Macia F, Prescott A G, Roberts K and Martin C. Expression patterns of myb genes from antirrhinum flowers [J]. Plant Cell.1991,3(2):115-125.
132. Jackson D, Roberts K and Martin C. Temporal and spatial control of expression of anthocyanin biosynthetic genes in developing flowers of Antirrhinum majus [J]. The Plant Journal.1992,2(4): 425-434.
133. Jez J M, Bowman M E, Dixon R A and Noel J P. Structure and mechanism of the evolutionarily unique plant enzyme chalcone isomerase [J]. Nature Structural and Molecular Biology.2000, 7(9):786-791.
134. Jez J M and Noel J P. Mechanism of chalcone synthase. pKa of the catalytic cysteine and the role of the conserved histidine in a plant polyketide synthase [J]. Journal of Biological Chemistry. 2000,275(50):39640-39646.
135. Johzuka-Hisatomi Y, Hoshino A, Mori T, Habu Y and Iida S. Characterization of the chalcone synthase genes expressed in flowers of the common and Japanese morning glories [J]. Genes and Genetic Systems.1999,74(4):141-147.
136. Jonsson L M V, Donker-Koopman W E and Schram A W. Turnover of anthocyanins and tissue compartmentation of anthocyanin biosynthesis in flowers of Petunia hybrida [J]. Multilingue. 1984,115(1):29-37.
137. Kaltenbach M, Schroder G, Schmelzer E, Lutz V and Schroder J. Flavonoid hydroxylase from Catharanthus roseus: cDNA, heterologous expression, enzyme properties and cell-type specific expression in plants[J]. Plant J.1999,19(2):183-193.
138. Katsumoto Y, Fukuchi-Mizutani M, Fukui Y, Brugliera F, Holton T A, et al. Engineering of the rose flavonoid biosynthetic pathway successfully generated blue-hued flowers accumulating delphinidin [J]. Plant Cell Physiol.2007,48(11):1589-1600.
139. Kawabata S, Ohta M, Kusuhara Y and Sakiyama R. Influences of low light intensities on the pigmentation of Eustoma grandiflorum flowers [J]. Acta Horticulturae.1995,405:173-178.
140. Kim S, Jones R, Yoo K-S and Pike L M. The L locus, one of complementary genes required for anthocyanin production in onions (Allium cepa), encodes anthocyanidin synthase [J]. TAG Theoretical and Applied Genetics.2005,111(1):120-127.
141.Kitada C, Gong Z, Tanaka Y, Yamazaki M and Saito K. Differential expression of two cytochrome P450s involved in the biosynthesis of flavones and anthocyanins in chemo-varietal forms of Perillafrutescens[J]. Plant Cell Physiol.2001,42(12):1338-1344.
142. Kobayashi, Kobayashi S, Ishimaru, Ishimaru M, Hiraoka, Hiraoka K, Honda and Honda C. Myb-related genes of the kyoho grape(Vitis labruscana) regulate anthocyanin biosynthesis [J]. Planta.2002,215(6):924-933.
143. Koes R, Verweij W and Quattrocchio F. Flavonoids: A colorful model for the regulation and evolution of biochemical pathways [J]. Trends in Plant Science.2005,10(5):236-242.
144. Koes R E, Spelt C E and Mol J N M. The chalcone synthase multigene family of Petunia hybrida (V30):Differential, light-regulated expression during flower development and UV light induction [J]. Plant Molecular Biology.1989,12(2):213-225.
145. Koes R E, van Blokland R, Quattrocchio F, van Tunen A J and Mol J. Chalcone synthase promoters in petunia are active in pigmented and unpigmented cell types [J]. Plant Cell.1990, 2(5):379-392.
146. Kranz H D, Denekamp M, Greco R, Jin H, Leyva A, et al. Towards functional characterisation of the members of the R2R3-MYB gene family from Arabidopsis thaliana[J]. The Plant Journal. 1998,16(2):263-276.
147. Kreuzaler F, Ragg H, Fautz E, Kuhn D N and Hahlbrock K. UV-induction of chalcone synthase mRNA in cell suspension cultures of Petroselinum hortense [J]. Proceedings of the National Academy of Sciences.1983,80(9):2591-2593.
148. Kusumi T, Tanaka Y, Suzuki K and Katsumoto Y. Development of flower colour modification technology and new floricultural varieties [J]. Nippon Nogeikagaku Kaishi.2002,76(8):700-707.
149. Lalusin A G, Ohta M and Fujimura T. Temporal and spatial expression of genes involved in anthocyanin biosynthesis during sweet potato (Ipomoea batatas [L.] Lam.) root development [J]. International Journal of Plant Sciences.2006,167(2):249-256.
150. Leyva A, Jarillo J A, Salinas J and Martinez-Zapater J M. Low temperature induces the accumulation of phenylalanine ammonia-lyase and chalcone synthase mrnas of Arabidopsis thaliana in a light-dependent manner [J]. Plant Physiology.1995,108(1):39-46.
151. Li F, Jin Z, Qua W, Zhao D and Ma F. Cloning of a cDNA encoding the Saussurea medusa chalcone isomerase and its expression in transgenic tobacco [J]. Plant Physiology and Biochemistry.2006,44(7-9):455-461.
152. Lillo C, Lea U S and Ruoff P. Nutrient depletion as a key factor for manipulating gene expression and product formation in different branches of the flavonoid pathway [J]. Plant, Cell and Environment.2008,31:587-601.
153. Liu C-J, Blount J W, Steele C L and Dixon R A. Bottlenecks for metabolic engineering of isoflavone glycoconjugates in Arabidopsis [J]. Proceedings of the National Academy of Sciences of the United States of America.2002,99(22):14578-14583.
154. Lloyd A, Walbot V and Davis R. Arabidopsis and nicotiana anthocyanin production activated by maize regulators R and C1 [J]. Science.1992,258(5089):1773-1775.
155. Loreti E, Povero G, Novi G, Solfanelli C, Alpi A and Perata P. Gibberellins, jasmonate and abscisic acid modulate the sucrose-induced expression of anthocyanin biosynthetic genes in Arabidopsis [J]. New Phytologist.2008,179(4):1004-1016.
156. Lu Q and Lu Q N. cDNA cloning and expression of anthocyanin biosynthetic genes in wild potato (Solanum pinnatisectum) [J]. African Journal of Biotechnology.2006,5(10):811-818.
157. Ludwig S R, Habera L F, Dellaporta S L and Wessler S R. Lc, a member of the maize R gene family responsible for tissue-specific anthocyanin production, encodes a protein similar to transcriptional activators and contains the myc-homology region [J]. Proceedings of the National Academy of Sciences of the United States of America.1989,86(18):7092-7096.
158. Lukacin R and Britsch L. Identification of strictly conserved histidine and arginine residues as part of the active site in Petunia hybrida flavanone 3beta-hydroxylase[J]. Eur J Biochem.1997, 249(3):748-757.
159. Lulin M, Yi S, Aizhong C, Zengjun Q, Liping X, Peidu C, Dajun L and Xiu-e W. Molecular cloning and characterization of an up-regulated UDP-glucosyltransferase gene induced by DON from Triticum aestivum L. cv. Wangshuibai[J]. Molecular Biology Reports.37(2):785-795.
160. Ma L, Liu B, Gao D, Pang X, Lǖ S, Yu H, Wang H, Yan F, Li Z and Li Y. Molecular cloning and overexpression of a novel UDP-glucosyltransferase elevating salidroside levels in Rhodiola sachalinensis[J]. Plant cell reports.2007,26(7):989-999.
161. McKhann H I and Hirsch A M. Isolation of chalcone synthase and chalcone isomerase cDNAs from alfalfa (Medicago sativa L.):highest transcript levels occur in young roots and root tips[J]. Plant Molecular Biology.1994,24(5):767-777.
162. Mano H, Ogasawara F, Sato K, Higo H and Minobe Y. Isolation of a regulatory gene of anthocyanin biosynthesis in tuberous roots of purple-fleshed sweet potato [J]. Plant Physiology. 2007,143(3):1252-1268.
163. Markham K R. Novel anthocyanins produced in petals of genetically transformed lisianthus [J]. Phytochemistry.1996,42(4):1035-1038.
164. Marrs K A, Alfenito M R, Lloyd A M and Walbot V. A glutathione S-transferase involved in vacuolar transfer encoded by the maize gene Bronze-2 [J]. Nature.1995,375(6530):397-400.
165. Martin C and Paz-Ares J. MYB transcription factors in plants [J]. Trends in Genetics.1997, 13(2):67-73.
166. Martin C, Prescott A, Mackay S, Bartlett J and Vrijlandt E. Control of anthocyanin biosynthesis in flowers of Antirrhinum majus [J]. The Plant Journal.1991,1(1):37-49.
167. Mathews H, Clendennen S K, Caldwell C G, Liu X L, Connors K, Matheis N, Schuster D K, Menasco D J, Wagoner W, Lightner J and Wagner D R. Activation tagging in tomato identifies a transcriptional regulator of anthocyanin biosynthesis, modification, and transport [J]. Plant Cell. 2003,15(8):1689-1703.
168. Mato M, Onozaki T, Ozeki Y, Higeta D, Itoh Y, Yoshimoto Y, Ikeda H, Yoshida H and Shibata M. Flavonoid biosynthesis in white-flowered sim carnations (Dianthus caryophyllus) [J]. Scientia Horticulturae.2000,84(3-4):333-347.
169. Matus J T, Loyola R, Vega A, Pena-Neira A, Bordeu E, Arce-Johnson P and Alcalde J A. Post-veraison sunlight exposure induces myb-mediated transcriptional regulation of anthocyanin and flavonol synthesis in berry skins of Vitis vinifera [J]. Journal of Experimental Botany.2009, 60(3):853-867.
170. Mehrtens F, Kranz H, Bednarek P and Weisshaar B. The Arabidopsis transcription factor MYB 12 is a flavonol-specific regulator of phenylpropanoid biosynthesis [J]. Plant Physiol.2005, 138(2):1083-1096.
171. Meng X, Xing T and Wang X. The role of light in the regulation of anthocyanin accumulation in Gerbera hybrida [J]. Plant Growth Regulation.2004,44(3):243-250.
172. Mol J, Cornish E, Mason J and Koes R. Novel coloured flowers [J]. Current Opinion in Biotechnology.1999,10(2):198-201.
173. Mol J, Grotewold E and Koes R. How genes paint flowers and seeds [J]. Trends Plant Sci.1998, 3(6):212-217.
174. Mol J, Jenkins G, Schafer E and Weiss D. Signal perception, tranduction, and gene expression involved in anthocyanin biosynthesis [J]. Critical reviews in plant sciences.1996,15(5-6) 525-557.
175. Mori K, Goto-Yamamoto N, Kitayama M and Hashizume K. Loss of anthocyanins in red-wine grape under high temperature [J]. Journal of Experimental Botany.2007:on line.
176. Mori K, Sugaya S and Gemma H. Decreased anthocyanin biosynthesis in grape berries grown under elevated night temperature condition [J]. Scientia Horticulturae.2005,105(3):319-330.
177. Morita Y, Saitoh M, Hoshino A, Nitasaka E and Iida S. Isolation of cDNAs for R2R3-MYB, bHLH and WDR transcriptional regulators and identification of c and ca mutations conferring white flowers in the Japanese morning glory [J]. Plant Cell Physiology.2006,47(4):457-470.
178. Moscovici S, Moalem-Beno D and Weiss D. Leaf-mediated light responses in petunia flowers [J]. Plant Physiology.1996,110(4):1275-1282.
179. Muir S R, Collins G J, Robinson S, Hughes S, Bovy A, Ric De Vos C H, van Tunen A J and Verhoeyen M E. Overexpression of petunia chalcone isomerase in tomato results in fruit containing increased levels of flavonols [J]. Nature Biotechnology.2001,19(5):470-474.
180. Nakajima J-i, Tanaka Y, Yamazaki M and Saito K. Cdna clonlng and gene expression of anthocyanidin synthase from Torenia fiournieri [J]. Plant Biotechnology.2000,17(4):331-335.
181. Nakamura N, Fukuchi-Mizutani M, Miyazaki K, Suzuki K and Tanaka Y. RNAi suppression of the anthocyanidin synthase gene in Torenia hybrida yields white flowers with higher frequency and better stability than antisense and sense suppression [J]. Plant Biotechnology.2006,23(1): 13-18.
182. Nakatsuka A, Izumi Y and Yamagishi M. Spatial and temporal expression of chalcone synthase and dihydroflavonol 4-reductase genes in the Asiatic hybrid lily[J]. Plant Science.2003,165: 759-767.
183. Nakatsuka A, Yamagishi M, Nakano M, Tasaki K and Kobayashi N. Light-induced expression of basic helix-loop-helix genes involved in anthocyanin biosynthesis in flowers and leaves of Asiatic hybrid lily [J]. Scientia Horticulturae.2009, in press.
184. Nakatsuka T, Nishihara M, Mishiba K, Hirano H and Yamamura S. Two different transposable elements inserted in flavonoid 3',5'-hydroxylase gene contribute to pink flower coloration in Gentiana scabra [J]. Molecular Genetics and Genomics. 2006,275(3):231-241.
185. Nakatsuka T, Nishihara M, Mishiba K and Yamamura S. Temporal expression of flavonoid biosynthesis-related genes regulates flower pigmentation in gentian plants [J]. Plant Science. 2005a,168(5):1309-1318.
186. Nakatsuka T, Nishihara M, Mishiba K and Yamamura S. Two different mutations are involved in the formation of white-flowered gentian plants [J]. Plant Science.2005b,169(5):949-958.
187. Nakatsuka T, Sato K, Takahashi H, Yamamura S and Nishihara M. Cloning and characterization of the UDP-glucose:anthocyanin 5-O-glucosyltransferase gene from blue-flowered gentian [J]. Journal of Experimental Botany.2008,59(6):1241-1252.
188. Nakayama M, Koshioka M, Shibata M, Hiradate S, Sugie H and Yamaguchi M A. Identification of cyanidin 3-O-(3,6"-O-dimalonyl-β-glucopyranoside) as a flower pigment of chrysanthemum (Dendranthema grandiflorum) [J]. Bioscience, biotechnology, and biochemistry.1997,61(9): 1607-1608.
189. Napoli C, Lemieux C and Jorgensen R. Introduction of a chimeric chalcone synthase gene into petunia results in reversible co-suppression of homologous genes in trans [J]. Plant Cell.1990, 2(4):279-289.
190. Nesi N, Debeaujon I, Jond C, Pelletier G, Caboche M and Lepiniec L. 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.
191. Nesi N, Debeaujon I, Jond C, Stewart A J, Jenkins G I, Caboche M and Lepiniec L. The TRANSPARENT TESTA16 locus encodes the ARABIDOPSIS BSISTER MADS domain protein and is required for proper development and pigmentation of the seed coat [J]. Plant Cell.2002, 14(10):2463-2479.
192. Nesi N, Jond C, Debeaujon I, Caboche M and Lepiniec L. The Arabidopsis TT2 gene encodes an R2R3 MYB domain protein that acts as a key determinant for proanthocyanidin accumulation in developing seed [J]. Plant Cell.2001,13(9):2099-2114.
193. Nielsen K, Deroles S C, Markham K R, Bradley M J, Podivinsky E and Manson D. Antisense flavonol synthase alters copigmentation and flower color in lisianthus [J]. Molecular Breeding. 2002,9(4):217-229.
194. Nishihara M, Nakatsuka T and Yamamura S. Flavonoid components and flower color change in transgenic tobacco plants by suppression of chalcone isomerase gene [J]. FEBS Letters.2005, 579(27):6074-6078.
195. Noda N, Kanno Y, Kato N, Kazuma K and Suzuki M. Regulation of gene expression involved in flavonol and anthocyanin biosynthesis during petal development in lisianthus (Eustoma grandiflorum) [J]. Physiologia Plantarum.2004,122(3):305-313.
196. Nozaki K, Takamura T and Fukai S. Effects of high temperature on flower colour and anthocyanin content in pink flower genotypes of greenhouse chrysanthemum(Chrysanthemum morifolium Ramat.) [J]. The Journal of Horticultural Science and Biotechnology 2006,81(4): 728-734.
197. Ogata J, Itoh Y, Ishida M, Yoshida H and Ozeki Y. Cloning and heterologous expression of cDNAs encoding flavonoid glucosyltransferases from Dianthus caryophyllus[J]. Plant Biotechnology.2004,21(5):367-375.
198. Okinaka Y, Shimada Y, Nakano-Shimada R, Ohbayashi M, Kiyokawa S and Kikuchi Y. Selective accumulation of delphinidin derivatives in tobacco using a putative flavonoid 3',5'-hydroxylase cDNA from Campanula medium[J]. Biosci Biotechnol Biochem.2003,67(1): 161-165.
199. Olsen K M, Hehn A, Jugde H, Slimestad R, Larbat R, Bourgaud F and Lillo C. Identification and characterisation of CYP75A31, a new flavonoid 3',5'-hydroxylase, isolated from Solanum lycopersicum[J]. BMC Plant Biol.10:21.
200. Oud J S N, Schneiders H, Kool A J and van Grinsven M Q J M. Breeding of transgenic orange Petunia hybrida varieties [J]. Euphytica.1995,84(3):175-181.
201. Pang Y, Wenger J P, Saathoff K, Peel G J, Wen J, et al. A WD40 repeat protein from Eedicago truncatula is necessary for tissue-specific anthocyanin and proanthocyanidin biosynthesis but not for trichome development [J]. Plant Physiolgy.2009,151(3):1114-1129.
202. Park J S, Choung M G, Kim J B, Hahn B S, Kim J B, Bae S C, Roh K H, Kim Y H, Cheon C I, Sung M K and Cho K J. Genes up-regulated during red coloration in UV-B irradiated lettuce leaves [J]. Plant Cell Reports.2007a,26(4):507-516.
203. Park K-I, Ishikawa N, Morita Y, Choi J-D, Hoshino A and Iida S. A bHLH regulatory gene in the common morning glory, Ipomoea purpurea, controls anthocyanin biosynthesis in flowers, proanthocyanidin and phytomelanin pigmentation in seeds, and seed trichome formation [J]. The Plant Journal.2007b,49(4):641-654.
204. Paz-Ares J, Ghosal D, Wienand U, Peterson P A and Saedler H. The regulatory cl locus of Zea mays encodes a protein with homology to myb proto-oncogene products and with structural similarities to transcriptional activators [J]. The EMBO Journal.1987,6(12):3553-3558.
205. Pelletier M K, Murrell J R and Shirley B W. Characterization of flavonol synthase and leucoanthocyanidin dioxygenase genes in arabidopsis (further evidence for differential regulation of "Early" And "Late" Genes) [J]. Plant Physiol.1997,113(4):1437-1445.
206. Pelletier M K and Winkel-Shirley B. Analysis of flavanone 3-hydroxylase in Arabidopsis seedlings (coordinate regulation with chalcone synthase and chalcone isomerase) [J]. Plant Physiol.1996,111(1):339-345.
207. Petroni K, Cominelli E, Consonni G, Gusmaroli G, Gavazzi G and Tonelli C. The developmental expression of the maize regulatory gene hopi determines germination-dependent anthocyanin accumulation [J]. Genetics.2000,155(1):323-336.
208. Piazza P, Procissi A, Jenkins G I and Tonelli C. Members of the cl/pll regulatory gene family mediate the response of maize aleurone and mesocotyl to different light qualities and cytokinins [J]. Plant Physiology.2002,128(3):1077-1086.
209. Polashock J J, Griesbach R J, Sullivan R F and Vorsa N. Cloning of a cDNA encoding the cranberry dihydroflavonol-4-reductase (DFR) and expression in transgenic tobacco [J]. Plant science.2002,163(2):241-251.
210. Poudel P R, Mochioka R, Beppu K and Kataoka I. Influence of temperature on berry composition of interspecific hybrid wine grape'kadainou r-1'(Vitis ficifolia var. Ganebu x V. Vinifera'muscat of alexandria') [J]. Journal of the Japanese Society for Horticultural Science. 2009,78(2):169-174.
211. Prescott A G, Stamford N P J, Wheeler G and Firmin J L. In vitro properties of a recombinant flavonol synthase from Arabidopsis thaliana [J]. Phytochemistry.2002,60(6):589-593.
212. Quattrocchio F, Verweij W, Kroon A, Spelt C, Mol J and Koes R. pH4 of petunia is an R2R3 myb protein that activates vacuolar acidification through interactions with basic-helix-loop-helix transcription factors of the anthocyanin pathway [J]. Plant Cell.2006,18(5):1274-1291.
213. Quattrocchio F, Wing J, Leppen H, Mol J and Koes R. Regulatory genes controlling anthocyanin pigmentation are functionally conserved among plant species and have distinct sets of target genes [J]. Plant Cell.1993,5(11):1497-1512.
214. Quattrocchio F, Wing J, van der Woude K, Mol J and Koes R. Analysis of bHLH and MYB domain proteins: species-specific regulatory differences are caused by divergent evolution of target anthocyanin genes [J]. Plant Journal.1998,13(4):475-488.
215. Rajendran L, Ravishankar G A, Venkataraman L V and Prathiba K R. Anthocyanin production in callus cultures of Daucus carota as influenced by nutrient stress and osmoticum [J]. Biotechnology Letters.1992,14(8):707-712.
216. Ramsay N A and Glover B J. MYB-bHLH-WD40 protein complex and the evolution of cellular diversity [J]. Trends in Plant Science.2005,10(2):63-70.
217. Ramsay N A, Walker A R, Mooney M and Gray J C. Two basic-helix-loop-helix genes (MYC-146 and GL3) from arabidopsis can activate anthocyanin biosynthesis in a white-flowered Matthiola incana mutant [J]. Plant Molecular Biology.2003,52(3):679-688.
218. Reddy A M, Reddy V S, Scheffler B E, Wienand U and Reddy A R. Novel transgenic rice overexpressing anthocyanidin synthase accumulates a mixture of flavonoids leading to an increased antioxidant potential [J]. Metabolic Engineering.2007,9(1):95-111.
219. Reddy A R, Britsch L, Salamini F, Saedler H and Rohde W. The A1 (anthocyanin-1) locus in Zea mays encodes dihydroquercetin reductase [J]. Plant Science.1987,52(1-2):7-13.
220. Reif H J, Niesbach U, Deumling B and Saedler H. Cloning and analysis of two genes for chalcone synthase from Petunia hybrida [J]. Molecular and General Genetics MGG.1985,199(2): 208-215.
221. Riechmann J L and Ratcliffe O J. A genomic perspective on plant transcription factors [J]. Current Opinion in Plant Biology.2000,3(5):423-434.
222. Ronchi A, Farina G, Gozzo F and Tonelli C. Effects of a triazolic fungicide on maize plant metabolism:Modifications of transcript abundance in resistance-related pathways [J]. Plant Science.1997,130(1):51-62.
223. Rosati C, Cadic A, Duron M, Ingouff M and Simoneau P. Molecular characterization of the anthocyanidin synthase gene in Forsythia x intermedia reveals organ-specific expression during flower development [J]. Plant Science.1999,149(1):73-79.
224. Rosso M G, Li Y, Strizhov N, Reiss B, Dekker K and Weisshaar B. An Arabidopsis thaliana T-DNA mutagenized population (GABI-Kat) for flanking sequence tag-based reverse genetics[J]. Plant Molecular Biology.2003,53(1):247-259.
225. Rowan D D, Cao M S, Lin-Wang K, Cooney J M, Jensen D J, Austin P T, Hunt M B, Norling C, Hellens R P, Schaffer R J and Allan A C. Environmental regulation of leaf colour in red 35S:PAP1 Arabidopsis thaliana [J]. New Phytologist.2009,182(1):102-115.
226. Saito K, Kobayashi M, Gong Z, Tanaka Y and Yamazaki M. Direct evidence for anthocyanidin synthase as a 2-oxoglutarate-dependent oxygenase: Molecular cloning and functional expression of cdna from a red forma of Perilla frutescens [J]. The Plant Journal.1999,17(2):181-189.
227. Sakamoto W, Ohmori T, Kageyama K, Miyazaki C, Saito A, Murata M, Noda K and Maekawa M. The purple leaf (Pl)lovud of rice: The Plw allele has a complex organization and includes two genes encoding basic helix-loop-helix proteins involved in anthocyanin biosynthesis [J]. Plant Cell Physiol.2001,42(9):982-991.
228. Shih C H, Chu H, Tang L K, Sakamoto W, Maekawa M, Chu I K, Wang M and Lo C. Functional characterization of key structural genes in rice flavonoid biosynthesis[J]. Planta.2008, 228(6):1043-1054.
229. Schijlen E G W M, Ric de Vos C H, van Tunen A J and Bovy A G. Modification of flavonoid biosynthesis in crop plants [J]. Phytochemistry.2004,65(19):2631-2648.
230. Schmelzer E, Jahnen W and Hahlbrock K. In situ localization of light-induced chalcone synthase mrna, chalcone synthase, and flavonoid end products in epidermal cells of parsley leaves [J]. Proceedings of the National Academy of Sciences.1988,85(9):2989-2993.
231. Schoenbohm C, Martens S, Eder C, Forkmann G and Weisshaar B. Identification of the Arabidopsis thaliana flavonoid 3'-hydroxylase gene and functional expression of the encoded P450 enzyme[J]. Biol Chem.2000,381(8):749-753.
232. Schulze-Lefert P, Becker-Andre M, Schulz W, Hahlbrock K and Dangl J L. Functional architecture of the light-responsive chalcone synthase promoter from parsley [J]. Plant Cell.1989, 1(7):707-714.
233. Schwinn K, Venail J, Shang Y, Mackay S, Alm V, Butelli E, Oyama R, Bailey P, Davies K and Martin C. A small family of MYB-regulatory genes controls floral pigmentation intensity and patterning in the genus antirrhinum [J]. Plant Cell.2006,18(4):831-851.
234. Schwinn K E, Davies K M, Deroles S C, Markham K R, Miller R M, Bradley J M, Manson D G and Given N K. Expression of an antirrhinum majus UDP-glucose:Flavonoid-3-o-glucosyltransferase transgene alters flavonoid glycosylation and acylation in lisianthus(Eustoma grandiflorum Grise.) [J]. Plant science.1997,125(1):53-61.
235. Seitz C, Eder C, Deiml B, Kellner S, Martens S and Forkmann G. Cloning, functional identification and sequence analysis of flavonoid 3'-hydroxylase and flavonoid 3',5'-hydroxylase cdnas reveals independent evolution of flavonoid 3',5'-hydroxylase in the asteraceae family [J]. Plant Molecular Biology.2006,61(3):365-381.
236. Seitz C, Oswald N, Borstling D, Forkmann G and Martens S. Being acyanic: An unavoidable fate for many white flowers? [J]. Acta Horticulturae.2003,612:83-88.
237. Shaked-Sachray L, Weiss D, Reuveni M, Nissim-Levi A and Oren-Shamir M. Increased anthocyanin accumulation in aster flowers at elevated temperatures due to magnesium treatment [J]. Physiologia Plantarum.2002,114(4):559-565.
238. Sheehan M J, Farmer P R and Brutnell T P. Structure and expression of maize phytochrome family homeologs [J]. Genetics.2004,167(3):1395-1405.
239. Shen G, Pang Y, Wu W, Deng Z, Zhao L, Cao Y, Sun X and Tang K. Cloning and characterization of a flavanone 3-hydroxylase gene from Ginkgo biloba [J]. Bioscience Reports. 2006,26(1):19-29.
240. Shimada S, Takahashi K, Sato Y and Sakuta M. Dihydroflavonol 4-reductase cDNA from non-anthocyanin-producing species in the Caryophyllales[J]. Plant and Cell Physiology.2004, 45(9):1290.
241. Shimada Y, Nakano-Shimada R, Ohbayashi M, Okinaka Y, Kiyokawa S and Kikuchi Y. Expression of chimeric P450 genes encoding flavonoid-3',5'-hydroxylase in transgenic tobacco and petunia plants(1)[J]. FEBS Lett.1999,461(3):241-245.
242. Shiono M, Matsugaki N and Takeda K. Structure of the blue cornflower pigment: Packaging red-rose anthocyanin as part of a'superpigment'in another flower turns it brilliant blue [J]. Nature. 2005,436:791.
243. Shirley B, Kubasek W, Storz G, Bruggemann E, Koornneef M, Ausubel F and Goodman H. Analysis of Arabidopsis mutants deficient in flavonoid biosynthesis [J]. The Plant Journal.1995, 8(5):659-671.
244. Shirley B W, Hanley S and Goodman H M. Effects of ionizing radiation on a plant genome: Analysis of two arabidopsis transparent testa mutations [J]. Plant Cell.1992,4(3):333-347.
245. Shvarts M, Borochov A and Weiss D. Low temperature enhances petunia flower pigmentation and induces chalcone synthase gene expression [J]. Physiologia Plantarum.1997,99(1):67-72.
246. Smith T F, Gaitatzes C, Saxena K and Neer E J. The wd repeat: A common architecture for diverse functions [J]. Trends in Biochemical Sciences.1999,24(5):181-185.
247. Solfanelli C, Poggi A, Loreti E, Alpi A and Perata P. Sucrose-specific induction of the anthocyanin biosynthetic pathway in Arabidopsis [J]. Plant Physiology.2006,140(2):637-646.
248. Sommer H and Saedler H. Structure of the chalcone synthase gene of Antirrhinum majus[J], Molecular and General Genetics MGG.1986,202(3):429-434.
249. Sparvoli F, Martin C, Scienza A, Gavazzi G and Tonelli C. Cloning and molecular analysis of structural genes involved in flavonoid and stilbene biosynthesis in grape (Vitis vinifera L.)[J]. Plant Molecular Biology.1994a,24(5):743-755.
250. Sparvoli F, Martin C, Scienza A, Gavazzi G and Tonelli C. Cloning and molecular analysis of structural genes involved in flavonoid and stilbene biosynthesis in grape (Vitis vinifera L.)[J]. Plant Mol Biol.1994b,24(5):743-755.
251. Spelt C, Quattrocchio F, Mol J and Koes R. anthocyaninl of petunia encodes a basic helix-loop-helix protein that directly activates transcription of structural anthocyanin genes [J]. Plant Cell.2000,12(9):1619-1632.
252. Stiles E A, Cech N B, Dee S M and Lacey E P. Temperature-sensitive anthocyanin production in flowers of Plantago lanceolata [J]. Physiologia Plantarum.2007,129(4):756-765.
253. Stracke R, Werber M and Weisshaar B. The R2R3-MYB gene family in Arabidopsis thaliana [J]. Current Opinion in Plant Biology.2001,4(5):447-456.
254. Suzuki H, Nakayama T, Yamaguchi M and Nishino T. cDNA cloning and characterization of two Dendranthema× morifolium anthocyanin malonyltransferases with different functional activities[J]. Plant Science.2004a,166(1):89-96.
255. Suzuki H, Sawada S Y, Yonekura-Sakakibara K, T N, Yamaguchi M A and Nishino T. Identification of a cdna encoding malonyl-coenzyme a:Anthocyanidin 3-O-glucoside 6"-O-malonyltransferase from cineraria (Senecio cruentus) flowers [J]. Plant Biotechnology. 2003,20(3):229-234.
256. Suzuki H, Shin'ya Sawada K, Nagae S, Yamaguchi M, Nakayama T and Nishino T. Identification and characterization of a novel anthocyanin malonyltransferase from scarlet sage (Salvia splendens) flowers:an enzyme that is phylogenetically separated from other anthocyanin acyltransferases[J]. The Plant Journal.2004b,38(6):994-1003.
257. Suzuki K, Xue H, Tanaka Y, Fukui Y, Fukuchi-Mizutani M, Murakami Y, Katsumoto Y, Tsuda S and Kusumi T. Flower color modifications of Torenia hybrida by cosuppression of anthocyanin biosynthesis genes [J]. Molecular breeding:new strategies in plant improvement.2000,6(3): 239-246.
258. Tanaka Y. Flower colour and cytochromes P450 [J]. Phytochemistry Reviews.2006,5(2): 283-291.
259. Tanaka Y, Katsumoto Y, Brugliera F and Mason J. Genetic engineering in floriculture [J]. Plant Cell, Tissue and Organ Culture.2005,80(1):1-24.
260. Tanaka Y and Ohmiya A. Seeing is believing: Engineering anthocyanin and carotenoid biosynthetic pathways [J]. Current Opinion in Biotechnology.2008,19(2):190-197.
261. Tanaka Y, Sasaki N and Ohmiya A. Biosynthesis of plant pigments: anthocyanins, betalains and carotenoids [J]. The Plant Journal.2008,54(4):733-749.
262. Tanaka Y, Tsuda S and Kusumi T. Metabolic engineering to modify flower color [J]. Plant Cell Physiol.1998,39(11):1119-1126.
263. Tatsuzawa F, Yukawa T, Shinoda K, Saito N. Acylated anthocyanins in the flowers of genus Dendrobium section Phalaenanthe (Orchidaceae) [J]. Biochemical Systematics and Ecology. 2005,33(6):625-629.
264. Taylor L P and Briggs W R. Genetic regulation and photocontrol of anthocyanin accumulation in maize seedlings [J]. Plant Cell.1990,2(2):115-127.
265. Togami J, Tamura M, Ishiguro K, Hirose C, Okuhara H, Ueyama Y, Nakamura N, Yonekura-Sakakibara K, Fukuchi-Mizutani M and Suzuki K. Molecular characterization of the flavonoid biosynthesis of Verbena hybrida and the functional analysis of verbena and Clitoria ternatea F3'5'H genes in transgenic verbena[J]. Plant Biotechnology.2006,23(1):5-11.
266. Toguri T, Umemoto N, Kobayashi O and Ohtani T. Activation of anthocyanin synthesis genes by white light in eggplant hypocotyl tissues, and identification of an inducible P-450 cDNA [J]. Plant Molecular Biology.1993,23(5):933-946.
267. Torskangerpoll K, Nφbaeb R, Nodland E,φvstedal D O, Andersen φ M. Anthocyanin content of Tulipa species and cultivars and its impact on tepal colours[J]. Biochemical Systematics and Ecology,2005,33(5):499-510.
268. Toyama-Kato Y, Yoshida K, Fujimori E, Haraguchi H, Shimizu Y and Kondo T. Analysis of metal elements of hydrangea sepals at various growing stages by ICP-AES [J]. Biochemical Engineering Journal.2003,14(3):237-241.
269. Tunen A J v, Koes R E, Spelt C E, Krol A R v d, Stuitje A R and Mol J N. Cloning of the two chalcone flavanone isomerase genes from Petunia hybrida:coordinate, light-regulated and differential expression of flavonoid genes[J]. The EMBO Journal.1988,7(5):1257-1263.
270. Turnbull J J, Nakajima J-i, Welford R W D, Yamazaki M, Saito K and Schofield C J. Mechanistic studies on three 2-oxoglutarate-dependent oxygenases of flavonoid biosynthesis [J]. Journal of Biological Chemistry.2004,279(2):1206-1216.
271. Ueyama Y, Suzuki K-i, Fukuchi-Mizutani M, Fukui Y, Miyazaki K, Ohkawa H, Kusumi T and Tanaka Y. Molecular and biochemical characterization of torenia flavonoid 3'-hydroxylase and flavone synthase Ⅱ and modification of flower color by modulating the expression of these genes [J]. Plant Science.2002,163(2):253-263.
272. van der Krol A R, Mur L A, Beld M, Mol J N M and Stuitje A R. Flavonoid genes in petunia: addition of a limited number of gene copies may lead to a suppression of gene expression [J]. the Plant Cell.1990,2(4):291-299.
273. van Houwelingen A, Souer E, Spelt K, Kloos D, Mol J and Koes R. Analysis of flower pigmentation mutants generated by random transposon mutagenesis in Petunia hybrida [J]. The Plant Journal.1998,13(1):39-50.
274. van Tunen A J, Hartman S A, Mur L A and Mol J N M. Regulation of chalcone flavanone isomerase (CHI) gene expression in Petunia hybrida:The use of alternative promoters in corolla, anthers and pollen [J]. Plant molecular biology.1989,12(5):539-551.
275. van Tunen A J, Koes R E, Spelt C E, van der Krol A R, Stuitje A R and Mol J N. Cloning of the two chalcone flavanone isomerase genes from Petunia hybrida:Coordinate, light-regulated and differential expression of flavonoid genes [J]. The EMBO Journal.1988,7(5):1257-1263.
276. van Tunen A J, Mur L A, Brouns G S, Rienstra J D, Koes R E and Mol J. Pollen- and anther-specific chi promoters from petunia:Tandem promoter regulation of the chiA gene [J]. Plant Cell.1990,2(5):393-401.
277. Walker A R, Davison P A, Bolognesi-Winfield A C, James C M, Srinivasan N, Blundell T L, Esch J J, Marks M D and Gray J C. The TRANSPARENT TESTA GLABRA1 locus, which regulates trichome differentiation and anthocyanin biosynthesis in Arabidopsis, encodes a wd40 repeat protein [J]. Plant Cell.1999,11(7):1337-1350.
278. Wang H M and To K Y. Agrobacterium-mediated transformation in the high-value medicinal plant Echinacea purpurea [J]. Plant Science.2004,166(4):1087-1096.
279. Wang J, Ming F, Han Y and Shen D. Flavonoid-3',5'-hydroxylase from Phalaenopsis: A novel member of cytochrome P450s, its cDNA cloning, endogenous expression and molecular modeling [J]. Biotechnology Letters.2006,28(5):327-334.
280. Wang L S, Shiraishi A, Hashimoto F, Aoki N, Shimizu K and Sakata Y. Analysis of Petal Anthocyanins to investigate flower coloration of Zhongyuan (Chinese) and Daikon Island (Japanese) tree peony cultivars. Journal of Plant Research [J].2001,114(1):33-43.
281. Weiss D and Halevy A H. The role of light reactions in the regulation of anthocyanin synthesis in Petunia corollas [J]. Physiologia Plantarum.1991,81(1):127-133.
282. Weiss D, van Blokland R, Kooter J M, Mol J N M and van Tunen A J. Gibberellic acid regulates chalcone synthase gene transcription in the corolla of Petunia hybrida [J]. Plant Physiology.1992, 98(1):191-197.
283. Whittall J, Voelckel C, Kliebenstein D and Hodges S. Convergence, constraint and the role of gene expression during adaptive radiation:floral anthocyanins in Aquilegia[J]. Molecular Ecology.2006,15(14):4645-4657.
284. Winkel-Shirley B. Flavonoid biosynthesis. A colorful model for genetics, biochemistry, cell biology, and biotechnology [J]. Plant Physiol.2001,126(2):485-493.
285. Winkel-Shirley B. Biosynthesis of flavonoids and effects of stress [J]. Current Opinion in Plant Biology.2002,5(3):218-223
286. Xu F, Cheng H, Cai R, Li L, Chang J, Zhu J, Zhang F, Chen L, Wang Y, Cheng S and Cheng S Y. Molecular cloning and function analysis of an anthocyanidin synthase gene from Ginkgo biloba, and its expression in abiotic stress responses [J]. Molecules and Cells.2008,26(6): 536-547.
287. Yamagishi M, Shimoyamada Y, Nakatsuka T and Masuda K. 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(3):463-474.
288. Yamazaki M, Makita Y, Springob K and Saito K. Regulatory mechanisms for anthocyanin biosynthesis in chemotypes of Perilla frutescens var. Crispa [J]. Biochemical Engineering Journal. 2003,14(3):191-197.
289. Yang J, Huang J, Gu H, Zhong Y and Yang Z. Duplication and adaptive evolution of the chalcone synthase genes of Dendranthema (Asteraceae)[J]. Molecular biology and evolution. 2002,19(10):1752-1759.
290. Yang S, Sweetman J P, Amirsadeghi S, Barghchi M, Huttly A K, Chung W-I and Twell D. Novel anther-specific myb genes from tobacco as putative regulators of phenylalanine ammonia-lyase expression [J]. Plant Physiol.2001,126(4):1738-1753.
291. Yoshida K, Kitahara S, Ito D and Kondo T. Ferric ions involved in the flower color development of the himalayan blue poppy, meconopsis grandis [J]. Phytochemistry.2006,67(10):992-998.
292. Yoshihara N, Imayama T, Matsuo Y, Fukuchi-Mizutani M, Tanaka Y, Ino I and Yabuya T. Characterization of cDNA clones encoding anthocyanin 3-p-coumaroyltransferase from Iris hollandica[J]. Plant Science.2006,171(5):632-639.
293. Zhou B, Li Y, Xu Z, Yan H, Homma S and Kawabata S. Ultraviolet a-specific induction of anthocyanin biosynthesis in the swollen hypocotyls of turnip(Brassica rapa) [J]. Journal of Experimental Botany.2007,58(7):1771-1781.
294. Zhou W, Huang C, Gong Y, Feng Q and Gao F. Molecular cloning and expression analysis of an ans gene encoding anthocyanidin synthase from purple-fleshed sweet potato [Ipomoea batatas (L.) Lam] [J]. Plant Molecular Biology Reporter.2010,28(1):112-121.
295. Zufall R A and Rausher M D. The Genetic Basis of a Flower Color Polymorphism in the Common Morning Glory (Ipomoea purpurea)[J]. Journal of Heredity.2003,94(6):442-448.
296. Zuker A, Tzfira T, Ben-Meir H, Ovadis M, Shklarman E, Itzhaki H, Forkmann G, Martens S, Neta-Sharir I, Weiss D and Vainstein A. Modification of flower color and fragrance by antisense suppression of the flavanone 3-hydroxylase gene [J]. Molecular Breeding 2002,9(1):33-41.