Biosynthesis and metabolic engineering of anthocyanins in Arabidopsis thaliana

Recent Patents on Biotechnology

Volumn 8, Issue 1, 2014, Pages 47-60

  Shi, Ming Zhu ^a   Xie, De Yu ^a  

^a NORTH CAROLINA STATE UNIVERSITY   (United States)

Author keywords

Anthocyanins; Arabidopsis thaliana; Biosynthetic pathway; Structural diversity; Transcriptional regulation

Indexed keywords

BIOCHEMISTRY; BIOSYNTHESIS; METABOLIC ENGINEERING; METABOLISM; PLANTS (BOTANY); TRANSCRIPTION;

'CURRENT; ANTHOCYANIN BIOSYNTHESIS; ARABIDOPSIS; ARABIDOPSIS THALIANA; BIOSYNTHETIC PATHWAY; IDEAL MODEL; PATHWAY GENES; PLANT BIOLOGY; STRUCTURAL DIVERSITY; TRANSCRIPTIONAL REGULATION;

ANTHOCYANINS;

ANTHOCYANIN; FLAVONOID; GLUTATHIONE TRANSFERASE; JASMONIC ACID; NITROGEN; SUCROSE; TRANSCRIPTION FACTOR; GLYCOSYLTRANSFERASE;

ACYLATION; ARABIDOPSIS THALIANA; BIOSYNTHESIS; GENE EXPRESSION; GENE SILENCING; GLYCOSYLATION; LIGHT; METABOLIC ENGINEERING; MICROARRAY ANALYSIS; NONHUMAN; PRIORITY JOURNAL; PROTEIN PROTEIN INTERACTION; REVERSE TRANSCRIPTION POLYMERASE CHAIN REACTION; REVIEW; SIGNAL TRANSDUCTION; STRUCTURAL BIOINFORMATICS; TRANSCRIPTION INITIATION; ARABIDOPSIS; CHEMISTRY; GENE EXPRESSION REGULATION; GENETICS; METABOLISM; PATENT; TRANSGENIC PLANT;

ANTHOCYANINS; ARABIDOPSIS; GENE EXPRESSION REGULATION, PLANT; GLYCOSYLTRANSFERASES; METABOLIC ENGINEERING; PATENTS AS TOPIC; PLANTS, GENETICALLY MODIFIED;

EID: 84924062061     PISSN: 18722083     EISSN: None     Source Type: Journal    
DOI: 10.2174/1872208307666131218123538     Document Type: Review

References (132)

1. Chalker-Scott L. Environmental significance of anthocyanins in plant stress responses. Photochem Photobiol Sci 1999; 70 (1): 1-9.
2. 2in red and green leaves after mechanical injury. Plant Cell Environ 2002; 25: 1261-9.
3. Neill SO, Gould KS. Anthocyanins in leaves: light attenuators or antioxidants. Funct Plant Biol 2003; 30: 865-73.
4. Hatier J-HB, Gould KS. Anthocyanin function in vegetative organs. In: Gould K, Davies K, Winefield C, Eds. Anthocyanins: biosynthesis, functions, and applications. New York: Springer 2009: pp. 1-20.
5. Steyn WJ, Wand SJE, Holcroft DM, Jacobs G. Anthocyanins in vegetative tissues: a proposed unified function in photoprotection. New Phytol 2002; 155: 349-61.
6. Zhang Y, Zheng S, Liu Z, Wang L, Bi Y. Both HY5 and HYH are necessary regulators for low temperature-induced anthocyanin accumulation in Arabidopsis seedlings. J Plant Physiol 2011; 168 (4): 367-74.
7. He J, Giusti MM. Anthocyanins: natural colorants with healthpromoting properties. Annu Rev Food Sci Technol 2010; 1: 163-87.
8. Pascual-Teresa DS, Moreno DA, Garcia-Viguera C. Flavanols and anthocyanins in cardiovascular health: a review of current evidence. Int J Mol Sci 2010; 11 (4): 1679-703.
9. Toufektsian MC, De Lorgeril M, Nagy N, et al. Chronic dietary intake of plant-derived anthocyanins protects the rat heart against ischemia-reperfusion injury. J Nutr 2008; 138 (4): 747-52.
10. Jing P, Bomser JA, Schwartz SJ, He J, Magnuson BA, Giusti MM. Structure-function relationships of anthocyanins from various anthocyanin-rich extracts on the inhibition of colon cancer cell growth. J Agric Food Chem 2008; 56 (20): 9391-8.
11. Speciale A, Canali R, Chirafisi J, Saija A, Virgili F, Cimino F. Cyanidin 3-O-glucoside protection against TNF-alpha-induced endothelial dysfunction: involvement of nuclear factor-kappaB signaling. J Agric Food Chem 2010; 58 (22): 12048-54.
12. Ghosh D, Konishi T. Anthocyanins and anthocyanin-rich extracts: role in diabetes and eye function. Asia Pac J Clin Nutr 2007; 16 (2): 200-8.
13. Butelli E, Titta L, Giorgio M, et al. Enrichment of tomato fruit with health-promoting anthocyanins by expression of select transcription factors. Nat Biotechnol 2008; 26 (11): 1301-8.
14. Bloor SJ, Abrahams S. The structure of the major anthocyanin in Arabidopsis thaliana. Phytochemistry 2002; 59 (3): 343-6.
15. Rowan DD, Cao M, Lin-Wang K, et al. Environmental regulation of leaf colour in red 35S: PAP1 Arabidopsis thaliana. New Phytol 2009; 182 (1): 102-15.
16. Shi MZ, Xie DY. Features of anthocyanin biosynthesis in pap1-D and wild-type Arabidopsis thaliana plants grown in different light intensity and culture media conditions. Planta 2010; 231 (6): 1385-400.
17. Shi MZ, Xie DY. Engineering of red cells of Arabidopsis thaliana and comparative genome-wide gene expression analysis of red cells versus wild-type cells. Planta 2011; 233 (4): 787-805.
18. Tohge T, Nishiyama Y, Hirai MY, et al. Functional genomics by integrated analysis of metabolome and transcriptome of Arabidopsis plants over-expressing an MYB transcription factor. Plant J 2005; 42 (2): 218-35.
19. Borevitz JO, Xia Y, Blount J, Dixon RA, Lamb C. Activation tagging identifies a conserved MYB regulator of phenylpropanoid biosynthesis. Plant Cell 2000; 12 (12): 2383-94.
20. Tatsuzawa F, Saito N, Shinoda K, Shigihara A, Honda T. Acylated cyanidin 3-sambubioside-5-glucosides in three garden plants of the Cruciferae. Phytochem 2006; 67 (12): 1287-95.
21. Nakabayashi R, Kusano M, Kobayashi M, et al. Metabolomicsoriented isolation and structure elucidation of 37 compounds including two anthocyanins from Arabidopsis thaliana. Phytochem 2009; 70 (8): 1017-29.
22. Saito N, Tatsuzawa F, Nishiyama A, Yokoi M, Shigihara A, Honda T. Acylated cyanidin 3-sambubioside-5-glucosides in Matthiola incana. Phytochem 1995; 38 (4): 1027-32.
23. Pourcel L, Irani NG, Lu Y, Riedl K, Schwartz S, Grotewold E. The formation of Anthocyanic Vacuolar Inclusions in Arabidopsis thaliana and implications for the sequestration of anthocyanin pigments. Mol Plant 2010; 3 (1): 78-90.
24. Luo J, Nishiyama Y, Fuell C, et al. Convergent evolution in the BAHD family of acyl transferases: identification and characterization of anthocyanin acyl transferases from Arabidopsis thaliana. Plant J 2007; 50 (4): 678-95.
25. Yonekura-Sakakibara K, Nakayama T, Yamazaki M, Saito K. Modification and stabilization of anthocyanins. In: Gould K, Davies K, Winefield C, Eds. Anthocyanins: Biosynthesis, functions and apllications. New York: Springer 2009: pp. 169-85.
26. Yonekura-Sakakibara K, Fukushima A, Nakabayashi R, et al. Two glycosyltransferases involved in anthocyanin modification delineated by transcriptome independent component analysis in Arabidopsis thaliana. Plant J 2012; 69 (1): 154-67.
27. Fraser CM, Thompson MG, Shirley AM, et al. Related Arabidopsis serine carboxypeptidase-like sinapoylglucose acyltransferases display distinct but overlapping substrate specificities. Plant Physiol 2007; 144 (4): 1986-99.
28. Nakayama T, Suzuki H, Nishino T. Anthocyanin acyltransferases: specificities, mechanism, phylogenetics, and applications. J Mol Catal B: Enzym 2003; 23: 117-32.
29. Joshi CP, Chiang VL. Conserved sequence motifs in plant Sadenosyl-L-methionine-dependent methyltransferases. Plant Mol Biol 1998; 37 (4): 663-74.
30. Huang J, Gu M, Lai Z, et al. Functional analysis of the Arabidopsis PAL gene family in plant growth, development, and response to environmental stress. Plant Physiol 2010; 153 (4): 1526-38.
31. Rohde A, Morreel K, Ralph J, et al. Molecular phenotyping of the pal1 and pal2 mutants of Arabidopsis thaliana reveals far-reaching consequences on phenylpropanoid, amino acid, and carbohydrate metabolism. Plant Cell 2004; 16 (10): 2749-71.
32. Olsen KM, Lea US, Slimestad R, Verheul M, Lillo C. Differential expression of four Arabidopsis PAL genes; PAL1 and PAL2 have functional specialization in abiotic environmental-triggered flavonoid synthesis. J Plant Physiol 2008; 165 (14): 1491-9.
33. Ehlting J, Buttner D, Wang Q, Douglas CJ, Somssich IE, Kombrink E. Three 4-coumarate: coenzyme A ligases in Arabidopsis thaliana represent two evolutionarily divergent classes in angiosperms. Plant J 1999; 19 (1): 9-20.
34. Lepiniec L, Debeaujon I, Routaboul JM, et al. Genetics and biochemistry of seed flavonoids. Annu Rev Plant Biol 2006; 57: 405-30.
35. Shirley BW, Kubasek WL, Storz G, et al. Analysis of Arabidopsis mutants deficient in flavonoid biosynthesis. Plant J 1995; 8 (5): 659-71.
36. Winkel-Shirley B. Flavonoid Biosynthesis. A Colorful Model for Genetics, Biochemistry, Cell Biology, and Biotechnology. Plant Physiology 2001; 126: 485-93.
37. Jorgensen K, Rasmussen AV, Morant M, et al. Metabolon formation and metabolic channeling in the biosynthesis of plant natural products. Curr Opin Plant Biol 2005; 8 (3): 280-91.
38. Winkel BS. Metabolic channeling in plants. Annu Rev Plant Biol 2004; 55: 85-107.
39. Winkel-Shirley B. Evidence for enzyme complexes in the phenylpropanoid and flavonoid pathways. Physiologia Plantarum 1999; 107: 142-9.
40. Achnine L, Blancaflor EB, Rasmussen S, Dixon RA. Colocalization of L-phenylalanine ammonia-lyase and cinnamate 4-hydroxylase for metabolic channeling in phenylpropanoid biosynthesis. Plant Cell 2004; 16 (11): 3098-109.
41. Gomez C, Conejero G, Torregrosa L, Cheynier V, Terrier N, Ageorges A. In vivo grapevine anthocyanin transport involves vesicle-mediated trafficking and the contribution of anthoMATE transporters and GST. Plant J 2011; 67 (6): 960-70.
42. Zhao J, Dixon RA. The 'ins' and 'outs' of flavonoid transport. Trends Plant Sci 2009; 15 (2): 72-80.
43. +-antiporter active in proanthocyanidin-accumulating cells of the seed coat. Plant Cell 2007; 19 (6): 2023-38.
44. Goodman CD, Casati P, Walbot V. A multidrug resistanceassociated protein involved in anthocyanin transport in Zea mays. Plant Cell 2004; 16 (7): 1812-26.
45. +-dependent acylated anthocyanin transporters. Plant Physiol 2009; 150 (1): 402-15.
46. Frank S, Keck M, Sagasser M, Niehaus K, Weisshaar B, Stracke R. Two differentially expressed MATE factor genes from apple complement the Arabidopsis transparent testa12 mutant. Plant Biol (Stuttg) 2011; 13 (1): 42-50.
47. Debeaujon I, Peeters AJ, Leon-Kloosterziel KM, Koornneef M. The TRANSPARENT TESTA12 gene of Arabidopsis encodes a multidrug secondary transporter-like protein required for flavonoid sequestration in vacuoles of the seed coat endothelium. Plant Cell 2001; 13 (4): 853-71.
48. +-ATPase is required for the formation of proanthocyanidins in the seed coat endothelium of Arabidopsis thaliana. Proc Natl Acad Sci U S A 2005; 102 (7): 2649-54.
49. Kitamura S, Shikazono N, Tanaka A. TRANSPARENT TESTA 19 is involved in the accumulation of both anthocyanins and proanthocyanidins in Arabidopsis. Plant J 2004; 37 (1): 104-14.
50. Poustka F, Irani NG, Feller A, et al. A trafficking pathway for anthocyanins overlaps with the endoplasmic reticulum-to-vacuole protein-sorting route in Arabidopsis and contributes to the formation of vacuolar inclusions. Plant Physiol 2007; 145 (4): 1323-35.
51. Wangwattana B, Koyama Y, Nishiyama Y, Kitayama M, Yamazaki M, Saito K. Characterization of PAP1-upregulated glutathione Stransferease genes in Arabidopsis thaliana. Plant Biotech J 2008; 25: 191-6.
52. Li X, Gao P, Cui D, et al. The Arabidopsis tt19-4 mutant differentially accumulates proanthocyanidin and anthocyanin through a 3' amino acid substitution in glutathione S-transferase. Plant Cell Environ 2011; 34 (3): 374-88.
53. Sun Y, Li H, Huang JR. Arabidopsis TT19 functions as a carrier to transport anthocyanin from the cytosol to tonoplasts. Mol Plant 2012; 5 (2): 387-400.
54. Xie DY, Shi MZ. Differentiation of programmed Arabidopsis cells. Bioeng Bugs 2012; 3 (1): 54-9.
55. Petroni K, Tonelli C. Recent advances on the regulation of anthocyanin synthesis in reproductive organs. Plant Sci 2011; 181 (3): 219-29.
56. Hichri I, Barrieu F, Bogs J, Kappel C, Delrot S, Lauvergeat V. Recent advances in the transcriptional regulation of the flavonoid biosynthetic pathway. J Exp Bot 2011; 62 (8): 2465-83.
57. Stracke R, Jahns O, Keck M, et al. Analysis of Production of flavonol glycosides-dependent flavonol glycoside accumulation in Arabidopsis thaliana plants reveals MYB11-, MYB12-and MYB111-independent flavonol glycoside accumulation. New Phytol 2010; 188 (4): 985-1000.
58. Stracke R, Werber M, Weisshaar B. The R2R3-MYB gene family in Arabidopsis thaliana. Curr Opin Plant Biol 2001; 4 (5): 447-56.
59. Dubos C, Stracke R, Grotewold E, Weisshaar B, Martin C, Lepiniec L. MYB transcription factors in Arabidopsis. Trends Plant Sci 2010; 15 (10): 573-81.
60. Gonzalez A, Zhao M, Leavitt JM, Lloyd AM. Regulation of the anthocyanin biosynthetic pathway by the TTG1/bHLH/Myb transcriptional complex in Arabidopsis seedlings. Plant J 2008; 53 (5): 814-27.
61. Zhou LL, Shi MZ, Xie DY. Regulation of anthocyanin biosynthesis by nitrogen in TTG1-GL3/TT8-PAP1-programmed red cells of Arabidopsis thaliana. Planta 2012; 236 (3): 825-37.
62. Li X, Gao MJ, Pan HY, Cui DJ, Gruber MY. Purple canola: Arabidopsis PAP1 increases antioxidants and phenolics in Brassica napus leaves. J Agric Food Chem 2010; 58 (3): 1639-45.
63. Zhang Y, Yan YP, Wang ZZ. The Arabidopsis PAP1 transcription factor plays an important role in the enrichment of phenolic acids in Salvia miltiorrhiza. J Agric Food Chem 2010.
64. Zvi MM, Shklarman E, Masci T, et al. PAP1 transcription factor enhances production of phenylpropanoid and terpenoid scent compounds in rose flowers. New Phytol 2012; 195 (2): 335-45.
65. Gatica-Arias A, Farag MA, Stanke M, Matousek J, Wessjohann L, Weber G. Flavonoid production in transgenic hop (Humulus lupulus L.) altered by PAP1/MYB75 from Arabidopsis thaliana L. Plant Cell Rep 2012; 31 (1): 111-9.
66. Zuluaga DL, Gonzali S, Loreti E, et al. Arabidopsis thaliana MYB75/PAP1 transcription factor induces anthocyanin production in transgenic tomato plants. Funct Plant Biol 2008 (35): 606-18.
67. Bailey PC, Martin C, Toledo-Ortiz G, et al. Update on the basic helix-loop-helix transcription factor gene family in Arabidopsis thaliana. Plant Cell 2003; 15 (11): 2497-502.
68. Heim MA, Jakoby M, Werber M, Martin C, Weisshaar B, Bailey PC. The basic helix-loop-helix transcription factor family in plants: a genome-wide study of protein structure and functional diversity. Mol Biol Evol 2003; 20 (5): 735-47.
69. Li X, Duan X, Jiang H, et al. Genome-wide analysis of basic/helixloop-helix transcription factor family in rice and Arabidopsis. Plant Physiol 2006; 141 (4): 1167-84.
70. Toledo-Ortiz G, Huq E, Quail PH. The Arabidopsis basic/helixloop-helix transcription factor family. Plant Cell 2003; 15 (8): 1749-70.
71. Baudry A, Heim MA, Dubreucq B, Caboche M, Weisshaar B, Lepiniec L. TT2, TT8, and TTG1 synergistically specify the expression of Banyuls and proanthocyanidin biosynthesis in Arabidopsis thaliana. Plant J 2004; 39 (3): 366-80.
72. Payne CT, Zhang F, Lloyd AM. GL3 encodes a bHLH protein that regulates trichome development in arabidopsis through interaction with GL1 and TTG1. Genetics 2000; 156 (3): 1349-62.
73. Nesi N, Debeaujon I, Jond C, Pelletier G, Caboche M, Lepiniec L. The TT8 gene encodes a basic helix-loop-helix domain protein required for expression of DFR and BAN genes in Arabidopsis siliques. Plant Cell 2000; 12 (10): 1863-78.
74. Zhao M, Morohashi K, Hatlestad G, Grotewold E, Lloyd A. The TTG1-bHLH-MYB complex controls trichome cell fate and patterning through direct targeting of regulatory loci. Development 2008; 135 (11): 1991-9.
75. Zhang F, Gonzalez A, Zhao M, Payne CT, Lloyd A. A network of redundant bHLH proteins functions in all TTG1-dependent pathways of Arabidopsis. Development 2003; 130 (20): 4859-69.
76. Ramsay NA, Walker AR, Mooney M, Gray JC. Two basic-helixloop-helix genes (MYC-146 and GL3) from Arabidopsis can activate anthocyanin biosynthesis in a white-flowered Matthiola incana mutant. Plant Mol Biol 2003; 52 (3): 679-88.
77. Feyissa DN, Lovdal T, Olsen KM, Slimestad R, Lillo C. The endogenous GL3, but not EGL3, gene is necessary for anthocyanin accumulation as induced by nitrogen depletion in Arabidopsis rosette stage leaves. Planta 2009; 230 (4): 747-54.
78. Baudry A, Caboche M, Lepiniec L. TT8 controls its own expression in a feedback regulation involving TTG1 and homologous MYB and bHLH factors, allowing a strong and cell-specific accumulation of flavonoids in Arabidopsis thaliana. Plant J 2006; 46 (5): 768-79.
79. Matsui K, Umemura Y, Ohme-Takagi M. AtMYBL2, a protein with a single MYB domain, acts as a negative regulator of anthocyanin biosynthesis in Arabidopsis. Plant J 2008; 55 (6): 954-67.
80. Maes L, Inze D, Goossens A. Functional specialization of the TRANSPARENT TESTA GLABRA1 network allows differential hormonal control of laminal and marginal trichome initiation in Arabidopsis rosette leaves. Plant Physiol 2008; 148 (3): 1453-64.
81. Koornneef M. The complex syndrome of ttg mutants. Arabidopsis Inform Serv 1981; 18: 45-51.
82. Walker AR, Davison PA, Bolognesi-Winfield AC, et al. The transparent testa glabra1 locus, which regulates trichome differentiation and anthocyanin biosynthesis in Arabidopsis, encodes a WD40 repeat protein. Plant Cell 1999; 11 (7): 1337-50.
83. Cominelli E, Gusmaroli G, Allegra D, et al. Expression analysis of anthocyanin regulatory genes in response to different light qualities in Arabidopsis thaliana. J Plant Physiol 2008; 165 (8): 886-94.
84. Lea US, Slimestad R, Smedvig P, Lillo C. Nitrogen deficiency enhances expression of specific MYB and bHLH transcription factors and accumulation of end products in the flavonoid pathway. Planta 2007; 225 (5): 1245-53.
85. Olsen KM, Slimestad R, Lea US, et al. Temperature and nitrogen effects on regulators and products of the flavonoid pathway: experimental and kinetic model studies. Plant Cell Environ 2009; 32 (3): 286-99.
86. Van Nocker S, Ludwig P. The WD-repeat protein superfamily in Arabidopsis: conservation and divergence in structure and function. BMC Genomics 2003; 4 (1): 50.
87. Balkunde R, Bouyer D, Hulskamp M. Nuclear trapping by GL3 controls intercellular transport and redistribution of TTG1 protein in Arabidopsis. Development 2011; 138 (22): 5039-48.
88. Zimmermann IM, Heim MA, Weisshaar B, Uhrig JF. Comprehensive identification of Arabidopsis thaliana MYB transcription factors interacting with R/B-like BHLH proteins. Plant J 2004; 40 (1): 22-34.
89. Zhu HF, Fitzsimmons K, Khandelwal A, Kranz RG. CPC, a singlerepeat R3 MYB, is a negative regulator of anthocyanin biosynthesis in Arabidopsis. Mol Plant 2009; 2 (4): 790-802.
90. Feller A, Machemer K, Braun EL, Grotewold E. Evolutionary and comparative analysis of MYB and bHLH plant transcription factors. Plant J 2011; 66 (1): 94-116.
91. Tominaga-Wada R, Iwata M, Nukumizu Y, Sano R, Wada T. A full-length R-like basic-helix-loop-helix transcription factor is required for anthocyanin upregulation whereas the N-terminal region regulates epidermal hair formation. Plant Sci 2012; 183: 115-22.
92. Hernandez JM, Feller A, Morohashi K, Frame K, Grotewold E. The basic helix loop helix domain of maize R links transcriptional regulation and histone modifications by recruitment of an EMSYrelated factor. Proc Natl Acad Sci USA 2007; 104 (43): 17222-7.
93. Wada T, Kurata T, Tominaga R, et al. Role of a positive regulator of root hair development, CAPRICE, in Arabidopsis root epidermal cell differentiation. Development 2002; 129 (23): 5409-19.
94. Wada T, Tachibana T, Shimura Y, Okada K. Epidermal cell differentiation in Arabidopsis determined by a Myb homolog, CPC. Sci 1997; 277 (5329): 1113-6.
95. Sawa S. Overexpression of the AtmybL2 gene represses trichome development in Arabidopsis. DNA Res 2002; 9 (2): 31-4.
96. Song SK, Ryu KH, Kang YH, et al. Cell fate in the Arabidopsis root epidermis is determined by competition between WEREWOLF and CAPRICE. Plant Physiol 2011; 157 (3): 1196-208.
97. Hsieh LC, Lin SI, Shih AC, et al. Uncovering small RNA-mediated responses to phosphate deficiency in Arabidopsis by deep sequencing. Plant Physiol 2009; 151 (4): 2120-32.
98. Rajagopalan R, Vaucheret H, Trejo J, Bartel DP. A diverse and evolutionarily fluid set of microRNAs in Arabidopsis thaliana. Genes Dev 2006; 20 (24): 3407-25.
99. Velten J, Cakir C, Youn E, Chen J, Cazzonelli CI. Transgene silencing and transgene-derived siRNA production in tobacco plants homozygous for an introduced AtMYB90 construct. PLoS One 2012; 7 (2): e30141.
100. Gou JY, Felippes FF, Liu CJ, Weigel D, Wang JW. Negative regulation of anthocyanin biosynthesis in Arabidopsis by a miR156-targeted SPL transcription factor. Plant Cell 2011; 23 (4): 1512-22.
101. Christie PJ, Alfenito MR, Walbot V. Impact of low-temperature stress on general phenylpropanoid and anthocyanin pathways: Enhancement of transcript abundance and anthocyanin pigmentation in maize seedlings. Planta 1994 (194): 541-9.
102. Leyva A, Jarillo JA, Salinas J, Martinez-Zapater JM. Low temperature induces the accumulation of phenylalanine ammonia-lyase and chalcone synthase mRNAs of Arabidopsis thaliana in a lightdependent manner. Plant Physiol 1995; 108 (1): 39-46.
103. Solfanelli C, Poggi A, Loreti E, Alpi A, Perata P. Sucrose-specific induction of the anthocyanin biosynthetic pathway in Arabidopsis. Plant Physiol 2006; 140 (2): 637-46.
104. Teng S, Keurentjes J, Bentsink L, Koornneef M, Smeekens S. Sucrose-specific induction of anthocyanin biosynthesis in Arabidopsis requires the MYB75/PAP1 gene. Plant Physiol 2005; 139 (4): 1840-52.
105. Shan X, Zhang Y, Peng W, Wang Z, Xie D. Molecular mechanism for jasmonate-induction of anthocyanin accumulation in Arabidopsis. J Exp Bot 2009; 60 (13): 3849-60.
106. Ang LH, Chattopadhyay S, Wei N, et al. Molecular interaction between COP1 and HY5 defines a regulatory switch for light control of Arabidopsis development. Mol Cell 1998; 1 (2): 213-22.
107. Shin J, Park E, Choi G. PIF3 regulates anthocyanin biosynthesis in an HY5-dependent manner with both factors directly binding anthocyanin biosynthetic gene promoters in Arabidopsis. Plant J 2007; 49 (6): 981-94.
108. Brown BA, Headland LR, Jenkins GI. UV-B action spectrum for UVR8-mediated HY5 transcript accumulation in Arabidopsis. Photochem Photobiol 2009; 85 (5): 1147-55.
109. Vandenbussche F, Habricot Y, Condiff AS, Maldiney R, Van der Straeten D, Ahmad M. HY5 is a point of convergence between cryptochrome and cytokinin signalling pathways in Arabidopsis thaliana. Plant J 2007; 49 (3): 428-41.
110. Chang CS, Li YH, Chen LT, et al. LZF1, a HY5-regulated transcriptional factor, functions in Arabidopsis de-etiolation. Plant J 2008; 54 (2): 205-19.
111. Hartmann U, Sagasser M, Mehrtens F, Stracke R, Weisshaar B. Differential combinatorial interactions of cis-acting elements recognized by R2R3-MYB, BZIP, and BHLH factors control lightresponsive and tissue-specific activation of phenylpropanoid biosynthesis genes. Plant Mol Biol 2005; 57 (2): 155-71.
112. Sivitz AB, Reinders A, Ward JM. Arabidopsis sucrose transporter AtSUC1 is important for pollen germination and sucrose-induced anthocyanin accumulation. Plant Physiol 2008; 147 (1): 92-100.
113. Lei M, Liu Y, Zhang B, et al. Genetic and genomic evidence that sucrose is a global regulator of plant responses to phosphate starvation in Arabidopsis. Plant Physiol 2011; 156 (3): 1116-30.
114. Loreti E, Povero G, Novi G, Solfanelli C, Alpi A, Perata P. Gibberellins, jasmonate and abscisic acid modulate the sucroseinduced expression of anthocyanin biosynthetic genes in Arabidopsis. New Phytol 2008; 179 (4): 1004-16.
115. Jeong SW, Das PK, Jeoung SC, et al. Ethylene suppression of sugar-induced anthocyanin pigmentation in Arabidopsis. Plant Physiol 2010; 154 (3): 1514-31.
116. Kwon Y, Oh JE, Noh H, Hong SW, Bhoo SH, Lee H. The ethylene signaling pathway has a negative impact on sucrose-induced anthocyanin accumulation in Arabidopsis. J Plant Res 2011; 124 (1): 193-200.
117. Lillo C, Lea US, Ruoff P. Nutrient depletion as a key factor for manipulating gene expression and product formation in different branches of the flavonoid pathway. Plant Cell Environ 2008; 31 (5): 587-601.
118. Pourtau N, Jennings R, Pelzer E, Pallas J, Wingler A. Effect of sugar-induced senescence on gene expression and implications for the regulation of senescence in Arabidopsis. Planta 2006; 224 (3): 556-68.
119. Gu Q, Ferrandiz C, Yanofsky MF, Martienssen R. The FRUITFULL MADS-box gene mediates cell differentiation during Arabidopsis fruit development. Development 1998; 125 (8): 1509-17.
120. Jaakola L, Poole M, Jones MO, et al. A SQUAMOSA MADS box gene involved in the regulation of anthocyanin accumulation in bilberry fruits. Plant Physiol 2010; 153 (4): 1619-29.
121. Rubin G, Tohge T, Matsuda F, Saito K, Scheible WR. Members of the LBD family of transcription factors repress anthocyanin synthesis and affect additional nitrogen responses in Arabidopsis. Plant Cell 2009; 21 (11): 3567-84.
122. Kazan K, Manners JM. The interplay between light and jasmonate signalling during defence and development. J Exp Bot 2011; 62 (12): 4087-100.
123. Xu L, Liu F, Lechner E, et al. The SCF(COI1) ubiquitin-ligase complexes are required for jasmonate response in Arabidopsis. Plant Cell 2002; 14 (8): 1919-35.
124. Pauwels L, Goossens A. The JAZ proteins: a crucial interface in the jasmonate signaling cascade. Plant Cell 2011; 23 (9): 3089-100.
125. Qi T, Song S, Ren Q, et al. The Jasmonate-ZIM-domain proteins interact with the WD-Repeat/bHLH/MYB complexes to regulate Jasmonate-mediated anthocyanin accumulation and trichome initiation in Arabidopsis thaliana. Plant Cell 2011; 23 (5): 1795-814.
126. Borevitz J, Xia YJ, Dixon RA, Lamb CJ. Regulation of anthocyanin pigment production. US 6573432 B1, 2003.
127. Espley R, Hellens R, Allan AC. Compositions and methods for modulating pigment production in plants. US 7973216-B2, 2011.
128. Vainstein A, Moyal-Ben-Zvi M, Rimon-Spitzer B. Methods of modulating production of phenylpropanoid compounds in plants. US 20100319091-A1, 2010.
129. Van Breusegem F, Vanderauwera S. Means and methods to modulate flavonoid biosynthesis in plants and plant cells. US 20090100545-A1, 2009.
130. Mouradov A, Spangenberg G. Modification of flavonoid biosynthesis in plants by PAP1. US 8008543-B2, 2011.
131. Mouradov A, Spangenberg G. Modification of flavonoid biosynthesis in plants. US 7960608-B2, 2011.
132. Spangenberg G, Mouradov A. Modification of plant flavonoid metabolism. US 20100186114-A1, 2010.