A review of target gene specificity of flavonoid R2R3-MYB transcription factors and a discussion of factors contributing to the target gene selectivity

doi:10.1007/s11515-013-1281-z

Front Biol

2013, Vol. 8

Issue (6) : 577-598 https://doi.org/10.1007/s11515-013-1281-z

REVIEW

A review of target gene specificity of flavonoid R2R3-MYB transcription factors and a discussion of factors contributing to the target gene selectivity

Yunsong LAI¹, Huanxiu LI¹(

), Masumi YAMAGISHI²

1. Institute of Pomology and Olericulture, Sichuan Agricultural University, Chengdu 611130, China; 2. Research Faculty of Agriculture, Hokkaido University, Kita-Ku, Sapporo 060-8589, Japan

Download: PDF(848 KB) HTML
Export: BibTeX | EndNote | Reference Manager | ProCite | RefWorks

Abstract

Flavonoid biosynthetic genes are often coordinately regulated in a temporal manner during flower or fruit development, resulting in specific accumulation profiles of flavonoid compounds. R2R3-MYB-type transcription factors (TFs) “recruit” a set of biosynthetic genes to produce flavonoids, and, therefore, R2R3-MYBs are responsible for the coordinated expression of structural genes. Although a wealth of information regarding the identified and functionally characterized R2R3-MYBs that are involved in flavonoid accumulation is available to date, this is the first review on the global regulation of MYB factors in the flavonoid pathway. The data presented in this review demonstrate that anthocyanin, flavone/flavonol/3-deoxyflavonoid (FFD), proanthocyanidin (PA), and isoflavonoid are independently regulated by different subgroups of R2R3-MYBs. Furthermore, FFD-specific R2R3-MYBs have a preference for early biosynthetic genes (EBGs) as their target genes; anthocyanin-specific R2R3-MYBs from dicot species essentially regulate late biosynthetic genes (LBGs); the remaining R2R3-MYBs have a wider range of target gene specificity. To elucidate the nature of the differential target gene specificity between R2R3-MYBs, we analyzed the DNA binding domain (also termed the MYB-domain) of R2R3-MYBs and the distribution of the recognition cis-elements. We identified four conserved amino acid residues located in or just before helix-3 of dicot anthocyanin R2R3-MYBs that might account for the different recognition DNA sequence and subsequently the different target gene specificity to the remaining R2R3-MYB TFs.

Keywords MYB cis-element DNA-binding domain flavonoid transcription factor target gene specificity

Corresponding Author(s): LI Huanxiu,Email:hxli62@163.com

Issue Date: 01 December 2013

Cite this article:

Yunsong LAI,Huanxiu LI,Masumi YAMAGISHI. A review of target gene specificity of flavonoid R2R3-MYB transcription factors and a discussion of factors contributing to the target gene selectivity[J]. Front Biol, 2013, 8(6): 577-598.

URL:

https://academic.hep.com.cn/fib/EN/10.1007/s11515-013-1281-z
https://academic.hep.com.cn/fib/EN/Y2013/V8/I6/577

Fig.1 Schematic of the major branch pathways of flavonoid biosynthesis. The major classes of flavonoid products are shown in boxes. CHS, chalcone synthase; CHI, chalcone isomerase; F3H, flavanone 3-hydroxylase; F3′H, flavanone 3′-hydroxylase; F3′5′H, flavanone 3′5′-hydroxylase; DFR, dihydroflavonol 4-reductase; ANS, anthocyanin synthase; GT, glycosyltransferases; AT, acyltransferase; CHR, chalcone reductase; IFS, isoflavone synthase; STS, stilbene synthase; AS, aurone synthase; FLS, flavonol synthase; FSN, flavone synthase; LAR, leucoanthocyanidin reductase; ANR, anthocyanin reductase; F2H, flavanone 2-hydroxylase. 160x139mm (200 × 200 DPI)

Species	MYB 1	Target genes 2, 3	Functional description
Anthocyanin-specific R2R3-MYBs in dicots
Antirrhinum majus	Roseal 1	CHS, CHI, F3H, F3′H, FLS, DFR, ANS, UFGT, AT	Floral pigmentation
Arabidopsis thaliana	Roseal 2	CHS, CHI, F3H, F3′H, FLS, DFR, ANS, UFGT, AT	Floral pigmentation
Brassica oleracea	Venosa	CHS, CHI, F3H, F3′H, FLS, DFR, ANS, UFGT, AT	Floral pigmentation
Capsicum annuum	PAP1	PAL1, CHS, F3′H, DFR, ANS, UFGT, GST, GT	Pigments in vegetative organs
Citrus sinensis	PAP2	PAL1, CHS, F3′H, DFR, ANS, UFGT, GST, GT	Pigments in vegetative organs
Epimedium sagittatum	MYB113	PAL1, CHS, F3′H, DFR, ANS, UFGT, GST, GT	Pigments in vegetative organs
Garcinia mangostana	MYB114	PAL1, CHS, F3′H, DFR, ANS, UFGT, GST, GT	Pigments in vegetative organs
Gentiana triflora	MYB2	CHS, CHI, F3H, F3′H, DFR, ANS, TT8, EGL3	Red cabbage, purple kale,
Gerbera hybrida	A	CHS, DFR, ANS	cauliflower
Ipomoea batatas	MYBA	DFR, ANS	Pigments in fruits
Ipomoea nil	MYBA1	DFR	Red flesh of blood orange
Solanum lycopersicum	MYB10	CHS, F3′5′H, 5AT	Red leaves
Malus × domestica	MYB3	Strongly induces LBGs	Fruit coloration
Medicago truncatula	MYB10	CHS, CHI, F3H, DFR, ANS, GT	Floral pigmentation
Myrica rubra	MYB1	CHS, CHI, F3H, F3′H, DFR, ANS, GT, GST	Floral pigmentation
Nicotiana tabacum	MYB1	Induces both EBGs and LBGs	Purple flesh of tuberous roots
Petunia hybrid	ANT1	DFR, GT	Floral pigmentation
Pyrus communis	MYB1 ⁴	ANS	Anthocyanin accumulation
Pyrus pyrifolia	MYBA ⁴	CHS, CHI, F3H, DFR, ANS, UFGT	Skin coloration
Prunus persica	MYB10	CHS, CHI, F3′H, DFR1, ANS, GT, LAR, ANR	Skin coloration
Solanum lycopersicum	LAP1	CHS, CHI, F3H, F3′H, DFR, ANS, UFGT	Skin and flesh coloration
Solanum tuberosum	MYB1	CHS, CHI, F3H, DFR, ANS	Anthocyanin accumulation
Vitis labruscana	MYBAN2	DFR	Fruit coloration
Vitis vinifera	AN2	Induces flavonoid genes except FLS, 3GT, AT	Floral pigmentation
	DPL	Induces flavonoid genes except F3′5′H	Floral pigmentation
	PHZ	PAL, CHS, CHI, F3H, DFR, ANS, UFGT	Anthocyanin in petal veins
	MYB10	PAL, CHS, CHI, F3H, DFR, ANS	Light-induced bud flush
	MYB10	DFR, UFGT, LAR	Fruit skin pigmentation
	MYB10	Induces both EBGs and LBGs	Fruit skin pigmentation
	ANT1	PAL, CHS, CHI, F3H, DFR, ANS, UFGT	Fruit skin pigmentation
	D	CHI, FLS1, ANS, ANR, UFGT	Anthocyanin accumulation
	MYBA		Tuber skin
	MYBA		Berry skin
			Berry skin
Anthocyanin-specific R2R3-MYBs in monocots
Lilium hybrid	MYB12	CHS, F3H, DFR	Floral pigmentation
Oncidium	MYB1	Transient expression induces CHI, DFR	Floral pigmentation
Zea mays	C1	At least FLS1, DFR, ANS, UFGT, GST	Anthocyanin, flavonol
	PL	At least CHS, DFR, UFGT	Vegetative and floral tissues
Flavone-, flavonol-, and 3-deoxyflavonoid-specific R2R3-MYBs
Gentiana triflora	MYBP3	CHS, F3′H, FNSII, F3′5′H	Flavone in young buds
Solanum lycopersicum	MYB12	PAL, C3H, CHS, CHI, F3H, F3′H, FLS, GT	Red peel
Arabidopsis thaliana	MYB12	CHS, CHI, F3H, FLS1, F3′H, DFR	Flavonol mainly in roots
Vitis vinifera	MYB11	CHS, CHI, F3H, FLS1, F3′H, DFR	Flavonol mainly in cotyledons
Sorghum bicolor	MYB111	CHS, CHI, F3H, FLS1, F3′H, DFR	Flavonol mainly in cotyledons
Zea mays	MYBF1	CHI, FLS1, ANS, ANR, UFGT	Flavonol in flowers, skin
	MYBY1	CHS, CHI, DFR	Phlobaphene
	P	CHS, CHI, F3′H, FLS1, DFR, F2H1	Flavonol, maysin, phlobaphene
Proanthocyanidin-specific R2R3-MYBs
Arabidopsis thaliana	TT2	CHS, CHI, F3H, F3′H, FLS, DFR, ANS, BAN	PA in seeds
Brassica napus	MYBTT2	CHS, F3′H, F3′5′H, DFR, ANR, LAR	PA in seeds
Diospyros kaki	MYB2	ANR, LAR	Wound-induced PA
Fragaria × ananassa	MYB4	F3′H, DFR, ANS, 3GT, ANR, LAR	PA in fruit; wound-induced PA
Lotus japonicas	MYB9	F3′H, DFR, ANS, 3GT, ANR, LAR	PA in young fruits
Medicago truncatula	MYB11	ANR	PA in young fruits
Populus tremuloides	MYBTT2	WD40	PA in roots and stems
Prunus persica	PAR	Strongly induces LBGs	PA in seed coat
Theobroma cacao	MYB134	DFR, LAI, UFGT	Inducible PA in leaves
Trifolium arvense	MYBPA1	CHS, CHI, FLS, F3H, DFR, ANS, BAN, UFGT	PA in fruits
Vaccinium corymbosum	MYBPA	Strongly induces LBGs in T. repens	PA and anthocyanin
Vitis vinifera	MYB14	CHI, FLS1, F3′5′H1, ANS, LAR1, LAR2,	PA in leaves
	MYBPA1	ANR, UFGT	PA in young berries
	MYBPA1	DFR, ANS, LAR1, LAR2, ANR, GT, PA1	PA mainly in seeds
	MYBPA2	CHI, F3′5′H, ANS, ANR, LAR1, UFGT	PA mainly in berry skin
	MYB5a	CHI, F3′5′H, ANS, ANR, LAR1, UFGT	PA in berries; general flavonoid
	MYB5b		PA in berries; general flavonoid
Isoflavonoid and general flavonoids R2R3-MYBs, MYB repressors
Glycine max	MYB176	CHS8	R1-MYB; isoflavonoid
Antirrhinum majus	MYB305	CHS, CHI, F3H, DFR, AS	Flavonoid in flowers
Arabidopsis thaliana	MYB340	CHS, CHI, F3H, DFR, AS	Flavonoid in flowers
Fragaria × ananassa	MYB308	Suppresses4CL, CHSin tobacco	Repressor of lignin
Malus × domestica	MYB330	Suppresses4CL, CHSin tobacco	Repressor of lignin
Perilla frutescens	MYB4	Suppresses CHS, C4H	Phenylpropanoid
Petunia hybrid	MYBL2	Suppresses CHS, CHI, F3H, F3′H, DFR, ANS, TT8	R3-MYB; anthocyanin
Pisum sativum	MYB1	Suppresses CHS, F3H, DFR, ANS, GT, RT,	Repressor of flavonoid
	MYB6	GST in tobacco	Repressor of flavonoid
	PfMYBP1	Suppresses all flavonoid genes in Arabidopsis	R3-MYB; anthocyanin
	MYB3	DFR	Flavonoid in petals
	MYB26	CHSJ	Phenylpropano
			id in flowers

Tab.1 Cloned and identified genes that regulate the flavonoid biosynthetic pathway.

Fig.2 Phylogenetic tree based on DNA binding domains showing that flavonoid-regulating R2R3-MYBs form nine clades. The tree was constructed using MEGA 4.0 (neighbor joining) and displays only topology. The amino acid sequences were retrieved from the DDBJ/EMBL/GenBank databases with the accession numbers shown in brackets. LhMYB12 and LhMYB6 (red dot) are from the monocot lily but cluster with dicot species; gentian GtMYB3 (red dot) clusters with VvMYB5a and VvMYB5b that regulate PA synthesis. Flavonoid branches that each clade is related are given in the outer ring. FFD, flavone/flavonol/3-deoxyflavonoid; GF, general flavonoids; PA, proanthocyanidin; AN, anthocyanin. 166 × 160mm (200 × 200 DPI)

Fig.3 Alignment of the MYB-domain of known flavonoid R2R3-MYB TFs using the ClustalW2 program. The boxed residues are those that are conserved within clade 9 but different from the remaining R2R3-MYBs. 185 × 172mm (136 × 136 DPI)

Tab.2 -elements interacting with flavonoid R2R3-MYBs.

1	Aharoni A, De Vos C H, Wein M, Sun Z, Greco R, Kroon A, Mol J N, O’Connell A P (2001). The strawberry FaMYB1 transcription factor suppresses anthocyanin and flavonol accumulation in transgenic tobacco. Plant J , 28(3): 319–332 doi: 10.1046/j.1365-313X.2001.01154.x pmid:11722774
2	Akagi T, Ikegami A, Tsujimoto T, Kobayashi S, Sato A, Kono A, Yonemori K (2009). DkMyb4 is a Myb transcription factor involved in proanthocyanidin biosynthesis in persimmon fruit. Plant Physiol , 151(4): 2028–2045 doi: 10.1104/pp.109.146985 pmid:19783643
3	Akagi T, Ikegami A, Yonemori K (2010). DkMyb2 wound-induced transcription factor of persimmon (Diospyros kaki Thunb.), contributes to proanthocyanidin regulation. Planta , 232(5): 1045–1059 doi: 10.1007/s00425-010-1241-7 pmid:20690029
4	Albert N W, Lewis D H, Zhang H, Schwinn K E, Jameson P E, Davies K M (2011). Members of an R2R3-MYB transcription factor family in Petunia are developmentally and environmentally regulated to control complex floral and vegetative pigmentation patterning. Plant J , 65(5): 771–784 doi: 10.1111/j.1365-313X.2010.04465.x pmid:21235651
5	Allan A C, Hellens R P, Laing W A (2008). MYB transcription factors that colour our fruit. Trends Plant Sci , 13(3): 99–102 doi: 10.1016/j.tplants.2007.11.012 pmid:18280199
6	Ang L H, Chattopadhyay S, Wei N, Oyama T, Okada K, Batschauer A, Deng X W (1998). Molecular interaction between COP1 and HY5 defines a regulatory switch for light control of Arabidopsis development. Mol Cell , 1(2): 213–222 doi: 10.1016/S1097-2765(00)80022-2 pmid:9659918
7	Ballester A R, Molthoff J, de Vos R, Hekkert Bt, Orzaez D, Fernández-Moreno J P, Tripodi P, Grandillo S, Martin C, Heldens J, Ykema M, Granell A, Bovy A (2010). Biochemical and molecular analysis of pink tomatoes: deregulated expression of the gene encoding transcription factor SlMYB12 leads to pink tomato fruit color. Plant Physiol , 152(1): 71–84 doi: 10.1104/pp.109.147322 pmid:19906891
8	Ban Y, Honda C, Hatsuyama Y, Igarashi M, Bessho H, Moriguchi T (2007). Isolation and functional analysis of a MYB transcription factor gene that is a key regulator for the development of red coloration in apple skin. Plant Cell Physiol , 48(7): 958–970 doi: 10.1093/pcp/pcm066 pmid:17526919
9	Baudry A, Heim M A, Dubreucq B, Caboche M, Weisshaar B, Lepiniec L (2004). TT2, TT8, and TTG1 synergistically specify the expression of BANYULS and proanthocyanidin biosynthesis in Arabidopsis thaliana. Plant J , 39(3): 366–380 doi: 10.1111/j.1365-313X.2004.02138.x pmid:15255866
10	Boddu J, Jiang C, Sangar V, Olson T, Peterson T, Chopra S (2006). Comparative structural and functional characterization of sorghum and maize duplications containing orthologous myb transcription regulators of 3-deoxyflavonoid biosynthesis. Plant Mol Biol , 60(2): 185–199 doi: 10.1007/s11103-005-3568-1 pmid:16429259
11	Boddu J, Svabek C, Ibraheem F, Jones A D, Chopra S (2005). Characterization of a deletion allele of a sorghum Myb gene yellow seed1 showing loss of 3-deoxyflavonoids. Plant Sci , 169(3): 542–552 doi: 10.1016/j.plantsci.2005.05.007
12	Bogs J, Downey M O, Harvey J S, Ashton A R, Tanner G J, Robinson S P (2005). Proanthocyanidin synthesis and expression of genes encoding leucoanthocyanidin reductase and anthocyanidin reductase in developing grape berries and grapevine leaves. Plant Physiol , 139(2): 652–663 doi: 10.1104/pp.105.064238 pmid:16169968
13	Bogs J, Jaffé F W, Takos A M, Walker A R, Robinson S P (2007). The grapevine transcription factor VvMYBPA1 regulates proanthocyanidin synthesis during fruit development. Plant Physiol , 143(3): 1347–1361 doi: 10.1104/pp.106.093203 pmid:17208963
14	Borevitz J O, Xia Y, Blount J, Dixon R A, Lamb C (2000). Activation tagging identifies a conserved MYB regulator of phenylpropanoid biosynthesis. Plant Cell , 12(12): 2383–2394 pmid:11148285
15	Borovsky Y, Oren-Shamir M, Ovadia R, De Jong W, Paran I (2004). The A locus that controls anthocyanin accumulation in pepper encodes a MYB transcription factor homologous to Anthocyanin2 of Petunia. Theor Appl Genet , 109(1): 23–29 doi: 10.1007/s00122-004-1625-9 pmid:14997303
16	Boss P K, Davies C, Robinson S P (1996). Analysis of the expression of anthocyanin pathway genes in developing Vitis vinifera L. cv Shiraz grape berries and the implications for pathway regulation. Plant Physiol , 111(4): 1059–1066 pmid:12226348
17	Bovy A, de Vos R, Kemper M, Schijlen E, Almenar Pertejo M, Muir S, Collins G, Robinson S, Verhoeyen M, Hughes S, Santos-Buelga C, van Tunen A (2002). High-flavonol tomatoes resulting from the heterologous expression of the maize transcription factor genes LC and C1. Plant Cell , 14(10): 2509–2526 doi: 10.1105/tpc.004218 pmid:12368501
18	Bushman B S, Snook M E, Gerke J P, Szalma S J, Berhow M A, Houchins K E, McMullen M D (2002). Two loci exert major effects on chlorogenic acid synthesis in maize silks. Crop Sci , 42(5): 1669–1678 doi: 10.2135/cropsci2002.1669
19	Chiou C Y, Yeh K W (2008). Differential expression of MYB gene (OgMYB1) determines color patterning in floral tissue of Oncidium Gower Ramsey. Plant Mol Biol , 66(4): 379–388 doi: 10.1007/s11103-007-9275-3 pmid:18161007
20	Chiu L W, Li L (2012). Characterization of the regulatory network of BoMYB2 in controlling anthocyanin biosynthesis in purple cauliflower. Planta , 236(4): 1153–1164 doi: 10.1007/s00425-012-1665-3 pmid:22644767
21	Chiu L W, Zhou X, Burke S, Wu X, Prior R L, Li L (2010). The purple cauliflower arises from activation of a MYB transcription factor. Plant Physiol , 154(3): 1470–1480 doi: 10.1104/pp.110.164160 pmid:20855520
22	Cocciolone S M, Nettleton D, Snook M E, Peterson T (2005). Transformation of maize with the p1 transcription factor directs production of silk maysin, a corn earworm resistance factor, in concordance with a hierarchy of floral organ pigmentation. Plant Biotechnol J , 3(2): 225–235 doi: 10.1111/j.1467-7652.2005.00120.x pmid:17173622
23	Cone K C, Burr F A, Burr B (1986). Molecular analysis of the maize anthocyanin regulatory locus C1. Proc Natl Acad Sci U S A , 83(24): 9631–9635 doi: 10.1073/pnas.83.24.9631 pmid:3025847
24	Cone K C, Cocciolone S M, Burr F A, Burr B (1993). Maize anthocyanin regulatory gene pl is a duplicate of c1 that functions in the plant. Plant Cell , 5(12): 1795–1805 pmid:8305872
25	Czemmel S, Heppel S C, Bogs J (2012). R2R3 MYB transcription factors: key regulators of the flavonoid biosynthetic pathway in grapevine. Protoplasma , 249(S2 Suppl 2): S109–S118 doi: 10.1007/s00709-012-0380-z pmid:22307206
26	Czemmel S, Stracke R, Weisshaar B, Cordon N, Harris N N, Walker A R, Robinson S P, Bogs J (2009). The grapevine R2R3-MYB transcription factor VvMYBF1 regulates flavonol synthesis in developing grape berries. Plant Physiol , 151(3): 1513–1530 doi: 10.1104/pp.109.142059 pmid:19741049
27	Dare A P, Schaffer R J, Lin-Wang K, Allan A C, Hellens R P (2008). Identification of a cis-regulatory element by transient analysis of co-ordinately regulated genes. Plant Methods , 4(1): 17 doi: 10.1186/1746-4811-4-17 pmid:18601751
28	Deluc L, Barrieu F, Marchive C, Lauvergeat V, Decendit A, Richard T, Carde J P, Mérillon J M, Hamdi S (2006). Characterization of a grapevine R2R3-MYB transcription factor that regulates the phenylpropanoid pathway. Plant Physiol , 140(2): 499–511 doi: 10.1104/pp.105.067231 pmid:16384897
29	Deluc L, Bogs J, Walker A R, Ferrier T, Decendit A, Merillon J M, Robinson S P, Barrieu F (2008). The transcription factor VvMYB5b contributes to the regulation of anthocyanin and proanthocyanidin biosynthesis in developing grape berries. Plant Physiol , 147(4): 2041–2053 doi: 10.1104/pp.108.118919 pmid:18539781
30	Devic M, Guilleminot J, Debeaujon I, Bechtold N, Bensaude E, Koornneef M, Pelletier G, Delseny M (1999). The BANYULS gene encodes a DFR-like protein and is a marker of early seed coat development. Plant J , 19(4): 387–398 doi: 10.1046/j.1365-313X.1999.00529.x pmid:10504561
31	Dhaubhadel S, Li X (2010). A new client for 14-3-3 proteins: GmMYB176, an R1 MYB transcription factor. Plant Signal Behav , 5(7): 921–923 doi: 10.4161/psb.5.7.12133 pmid:20622511
32	Dhaubhadel S, McGarvey B D, Williams R, Gijzen M (2003). Isoflavonoid biosynthesis and accumulation in developing soybean seeds. Plant Mol Biol , 53(6): 733–743 doi: 10.1023/B:PLAN.0000023666.30358.ae pmid:15082922
33	Dixon R A, Xie D Y, Sharma S B (2005). Proanthocyanidins—a final frontier in flavonoid research? New Phytol , 165(1): 9–28 doi: 10.1111/j.1469-8137.2004.01217.x pmid:15720617
34	Downey M O, Harvey J S, Robinson S P (2003a). Analysis of tannins in seeds and skins of Shiraz grapes throughout berry development. Aust J Grape Wine Res , 9(1): 15–27 doi: 10.1111/j.1755-0238.2003.tb00228.x
35	Downey M O, Harvey J S, Robinson S P (2003b). Synthesis of flavonols and expression of flavonol synthase genes in the developing grape berries of Shiraz and Chardonnay (Vitis vinifera L.). Aust J Grape Wine Res , 9(2): 110–121 doi: 10.1111/j.1755-0238.2003.tb00261.x
36	Du H, Feng B R, Yang S S, Huang Y B, Tang Y X (2012a). The R2R3-MYB transcription factor gene family in maize. PLoS One , 7(6): e37463 doi: 10.1371/journal.pone.0037463 pmid:22719841
37	Du H, Huang Y, Tang Y (2010). Genetic and metabolic engineering of isoflavonoid biosynthesis. Appl Microbiol Biotechnol , 86(5): 1293–1312 doi: 10.1007/s00253-010-2512-8 pmid:20309543
38	Du H, Yang S S, Liang Z, Feng B R, Liu L, Huang Y B, Tang Y X (2012b). Genome-wide analysis of the MYB transcription factor superfamily in soybean. BMC Plant Biol , 12(1): 106 doi: 10.1186/1471-2229-12-106 pmid:22776508
39	Dubos C, Le Gourrierec J, Baudry A, Huep G, Lanet E, Debeaujon I, Routaboul J M, Alboresi A, Weisshaar B, Lepiniec L (2008). MYBL2 is a new regulator of flavonoid biosynthesis in Arabidopsis thaliana. Plant J , 55(6): 940–953 doi: 10.1111/j.1365-313X.2008.03564.x pmid:18532978
40	Dubos C, Stracke R, Grotewold E, Weisshaar B, Martin C, Lepiniec L (2010). MYB transcription factors in Arabidopsis. Trends Plant Sci , 15(10): 573–581 doi: 10.1016/j.tplants.2010.06.005 pmid:20674465
41	Elomaa P, Uimari A, Mehto M, Albert V A, Laitinen R A, Teeri T H (2003). Activation of anthocyanin biosynthesis in Gerbera hybrida (Asteraceae) suggests conserved protein-protein and protein-promoter interactions between the anciently diverged monocots and eudicots. Plant Physiol , 133(4): 1831–1842 doi: 10.1104/pp.103.026039 pmid:14605235
42	Espley R V, Hellens R P, Putterill J, Stevenson D E, Kutty-Amma S, Allan A C (2007). Red colouration in apple fruit is due to the activity of the MYB transcription factor, MdMYB10. Plant J , 49(3): 414–427 doi: 10.1111/j.1365-313X.2006.02964.x pmid:17181777
43	Falcone Ferreyra M L, Rius S, Emiliani J, Pourcel L, Feller A, Morohashi K, Casati P, Grotewold E (2010). Cloning and characterization of a UV-B-inducible maize flavonol synthase. Plant J , 62(1): 77–91 doi: 10.1111/j.1365-313X.2010.04133.x pmid:20059741
44	Falcone Ferreyra M L, Rius S P, Casati P (2012) Flavonoids: biosynthesis, biological functions, and biotechnological applications. Front Plant Sci 3: Article 222
45	Feldbrügge M, Sprenger M, Hahlbrock K, Weisshaar B (1997). PcMYB1, a novel plant protein containing a DNA-binding domain with one MYB repeat, interacts in vivo with a light-regulatory promoter unit. Plant J , 11(5): 1079–1093 doi: 10.1046/j.1365-313X.1997.11051079.x pmid:9193077
46	Feng S, Wang Y, Yang S, Xu Y, Chen X (2010). Anthocyanin biosynthesis in pears is regulated by a R2R3-MYB transcription factor PyMYB10. Planta , 232(1): 245–255 doi: 10.1007/s00425-010-1170-5 pmid:20422209
47	Ferrer J L, Austin M B, Stewart C Jr, Noel J P (2008). Structure and function of enzymes involved in the biosynthesis of phenylpropanoids. Plant Physiol Biochem , 46(3): 356–370 doi: 10.1016/j.plaphy.2007.12.009 pmid:18272377
48	Frampton J, Gibson T J, Ness S A, D?derlein G, Graf T (1991). Proposed structure for the DNA-binding domain of the Myb oncoprotein based on model building and mutational analysis. Protein Eng , 4(8): 891–901 doi: 10.1093/protein/4.8.891 pmid:1817253
49	Fujiwara H, Tanaka Y, Yonekura-Sakakibara K, Fukuchi-Mizutani M, Nakao M, Fukui Y, Yamaguchi M, Ashikari T, Kusumi T (1998). cDNA cloning, gene expression and subcellular localization of anthocyanin 5-aromatic acyltransferase from Gentiana triflora. Plant J , 16(4): 421–431 doi: 10.1046/j.1365-313x.1998.00312.x pmid:9881162
50	Gabrielsen O S, Sentenac A, Fromageot P (1991). Specific DNA binding by c-Myb: evidence for a double helix-turn-helix-related motif. Science , 253(5024): 1140–1143 doi: 1887237" target="_blank">10.1126/science. pmid:1887237 pmid:1887237
51	Gao J, Shen X, Zhang Z, Peng R, Xiong A, Xu J, Zhu B, Zheng J, Yao Q (2011). The myb transcription factor MdMYB6 suppresses anthocyanin biosynthesis in transgenic Arabidopsis. Plant Cell Tissue Organ Cult , 106(2): 235–242 doi: 10.1007/s11240-010-9912-4
52	Goff S A, Cone K C, Chandler V L (1992). Functional analysis of the transcriptional activator encoded by the maize B gene: evidence for a direct functional interaction between two classes of regulatory proteins. Genes Dev , 6(5): 864–875 doi: 10.1101/gad.6.5.864 pmid:1577278
53	Goff S A, Cone K C, Fromm M E (1991). Identification of functional domains in the maize transcriptional activator C1: comparison of wild-type and dominant inhibitor proteins. Genes Dev , 5(2): 298–309 doi: 10.1101/gad.5.2.298 pmid:1995419
54	Gong Z Z, Yamazaki M, Saito K (1999). A light-inducible Myb-like gene that is specifically expressed in red Perilla frutescens and presumably acts as a determining factor of the anthocyanin forma. Mol Gen Genet , 262(1): 65–72 doi: 10.1007/PL00008639 pmid:10503537
55	Gonzalez A, Zhao M, Leavitt J M, Lloyd A M (2008). Regulation of the anthocyanin biosynthetic pathway by the TTG1/bHLH/Myb transcriptional complex in Arabidopsis seedlings. Plant J , 53(5): 814–827 doi: 10.1111/j.1365-313X.2007.03373.x pmid:18036197
56	Grotewold E (2006). The genetics and biochemistry of floral pigments. Annu Rev Plant Biol , 57(1): 761–780 doi: 10.1146/annurev.arplant.57.032905.105248 pmid:16669781
57	Grotewold E, Athma P, Peterson T (1991). Alternatively spliced products of the maize P gene encode proteins with homology to the DNA-binding domain of myb-like transcription factors. Proc Natl Acad Sci U S A , 88(11): 4587–4591 doi: 10.1073/pnas.88.11.4587 pmid:2052542
58	Grotewold E, Chamberlin M, Snook M, Siame B, Butler L, Swenson J, Maddock S, St Clair G, Bowen B (1998). Engineering secondary metabolism in maize cells by ectopic expression of transcription factors. Plant Cell , 10(5): 721–740 pmid:9596632
59	Grotewold E, Drummond B J, Bowen B, Peterson T (1994). The myb-homologous P gene controls phlobaphene pigmentation in maize floral organs by directly activating a flavonoid biosynthetic gene subset. Cell , 76(3): 543–553 doi: 10.1016/0092-8674(94)90117-1 pmid:8313474
60	Hancock K R, Collette V, Fraser K, Greig M, Xue H, Richardson K, Jones C, Rasmussen S (2012). Expression of the R2R3-MYB transcription factor TaMYB14 from Trifolium arvense activates proanthocyanidin biosynthesis in the legumes Trifolium repens and Medicago sativa. Plant Physiol , 159(3): 1204–1220 doi: 10.1104/pp.112.195420 pmid:22566493
61	Hartmann U, Sagasser M, Mehrtens F, Stracke R, Weisshaar B (2005). 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. Plant Mol Biol , 57(2): 155–171 doi: 10.1007/s11103-004-6910-0 pmid:15821875
62	Heppel S C, Jaffé F W, Takos A M, Schellmann S, Rausch T, Walker A R, Bogs J (2013). Identification of key amino acids for the evolution of promoter target specificity of anthocyanin and proanthocyanidin regulating MYB factors. Plant Mol Biol , 82(4-5): 457–471 doi: 10.1007/s11103-013-0074-8 pmid:23689818
63	Hichri I, Barrieu F, Bogs J, Kappel C, Delrot S, Lauvergeat V (2011a). Recent advances in the transcriptional regulation of the flavonoid biosynthetic pathway. J Exp Bot , 62(8): 2465–2483 doi: 10.1093/jxb/erq442 pmid:21278228
64	Hichri I, Deluc L, Barrieu F, Bogs J, Mahjoub A, Regad F, Gallois B, Granier T, Trossat-Magnin C, Gomès E, Lauvergeat V (2011b). A single amino acid change within the R2 domain of the VvMYB5b transcription factor modulates affinity for protein partners and target promoters selectivity. BMC Plant Biol , 11(1): 117 doi: 10.1186/1471-2229-11-117 pmid:21861899
65	Huang W, Sun W, Lv H, Luo M, Zeng S, Pattanaik S, Yuan L, Wang Y (2013). A R2R3-MYB transcription factor from Epimedium sagittatum regulates the flavonoid biosynthetic pathway. PLoS One , 8(8): e70778 doi: 10.1371/journal.pone.0070778 pmid:23936468
66	Ikegami A, Eguchi S, Kitajima A, Inoue K, Yonemori K (2007). Identification of genes involved in proanthocyanidin biosynthesis of persimmon (Diospyros kaki) fruit. Plant Sci , 172(5): 1037–1047 doi: 10.1016/j.plantsci.2007.02.010
67	Jackson D, Culianez-Macia F, Prescott A G, Roberts K, Martin C (1991). Expression patterns of myb genes from Antirrhinum flowers. Plant Cell , 3(2): 115–125 pmid:1840903
68	Jia L, Clegg M T, Jiang T (2004). Evolutionary dynamics of the DNA-binding domains in putative R2R3-MYB genes identified from rice subspecies indica and japonica genomes. Plant Physiol , 134(2): 575–585 doi: 10.1104/pp.103.027201 pmid:14966247
69	Jin H, Cominelli E, Bailey P, Parr A, Mehrtens F, Jones J, Tonelli C, Weisshaar B, Martin C (2000). Transcriptional repression by AtMYB4 controls production of UV-protecting sunscreens in Arabidopsis. EMBO J , 19(22): 6150–6161 doi: 10.1093/emboj/19.22.6150 pmid:11080161
70	Jung C S, Griffiths H M, De Jong D M, Cheng S, Bodis M, Kim T S, De Jong W S (2009). The potato developer (D) locus encodes an R2R3 MYB transcription factor that regulates expression of multiple anthocyanin structural genes in tuber skin. Theor Appl Genet , 120(1): 45–57 doi: 10.1007/s00122-009-1158-3 pmid:19779693
71	Kobayashi S, Ishimaru M, Hiraoka K, Honda C (2002). Myb-related genes of the Kyoho grape ( Vitis labruscana) regulate anthocyanin biosynthesis. Planta , 215(6): 924–933 doi: 10.1007/s00425-002-0830-5 pmid:12355152
72	Koes R, Verweij W, Quattrocchio F (2005). Flavonoids: a colorful model for the regulation and evolution of biochemical pathways. Trends Plant Sci , 10(5): 236–242 doi: 10.1016/j.tplants.2005.03.002 pmid:15882656
73	Kong Q, Pattanaik S, Feller A, Werkman J R, Chai C, Wang Y, Grotewold E, Yuan L (2012). Regulatory switch enforced by basic helix-loop-helix and ACT-domain mediated dimerizations of the maize transcription factor R. Proc Natl Acad Sci U S A , 109(30): E2091–E2097 doi: 10.1073/pnas.1205513109 pmid:22778424
74	Lai Y S, Shimoyamada Y, Nakayama M, Yamagishi M (2012). Pigment accumulation and transcription of LhMYB12 and anthocyanin biosynthesis genes during flower development in the Asiatic hybrid lily (Lilium spp.). Plant Sci , 193-194: 136–147 doi: 10.1016/j.plantsci.2012.05.013 pmid:22794927
75	Laitinen R A, Ainasoja M, Broholm S K, Teeri T H, Elomaa P (2008). Identification of target genes for a MYB-type anthocyanin regulator in Gerbera hybrida. J Exp Bot , 59(13): 3691–3703 doi: 10.1093/jxb/ern216 pmid:18725377
76	Lee J, He K, Stolc V, Lee H, Figueroa P, Gao Y, Tongprasit W, Zhao H, Lee I, Deng X W (2007). Analysis of transcription factor HY5 genomic binding sites revealed its hierarchical role in light regulation of development. Plant Cell , 19(3): 731–749 doi: 10.1105/tpc.106.047688 pmid:17337630
77	Lesnick M L, Chandler V L (1998). Activation of the maize anthocyanin gene a2 is mediated by an element conserved in many anthocyanin promoters. Plant Physiol , 117(2): 437–445 doi: 10.1104/pp.117.2.437 pmid:9625696
78	Li L, Ban Z J, Li X H, Wu M Y, Wang A L, Jiang Y Q, Jiang Y H (2012a). Differential expression of anthocyanin biosynthetic genes and transcription factor PcMYB10 in pears (Pyrus communis L.). PLoS One , 7(9): e46070 doi: 10.1371/journal.pone.0046070 pmid:23029391
79	Li Z T, Gmitter F G, Grosser J W, Chen C, Gray D J (2012b). Isolation and characterization of a novel anthocyanin-promoting MYBA gene family in Citrus. Tree Genet Genomes , 8(4): 675–685 doi: 10.1007/s11295-011-0455-6
80	Lin-Wang K, Bolitho K, Grafton K, Kortstee A, Karunairetnam S, McGhie T K, Espley R V, Hellens R P, Allan A C (2010). An R2R3 MYB transcription factor associated with regulation of the anthocyanin biosynthetic pathway in Rosaceae. BMC Plant Biol , 10(1): 50 doi: 10.1186/1471-2229-10-50 pmid:20302676
81	Liu Y (2010) Molecular analysis of genes involved in the synthesis of proanthocyanidins in Theobroma cacao. PhD Dissertation (USA: The Pennsylvania State University).
82	Lüscher B, Eisenman R N (1990). New light on Myc and Myb. Part II. Myb . Genes Dev , 4(12B 12b): 2235–2241 doi: 10.1101/gad.4.12b.2235 pmid:2279697
83	Mackova Z, Koblovska R, Lapcik O (2006). Distribution of isoflavonoids in non-leguminous taxa- an update. Phytochemistry , 67(9): 849–855 doi: 10.1016/j.phytochem.2006.01.020 pmid:16499937
84	Mano H, Ogasawara F, Sato K, Higo H, Minobe Y (2007). Isolation of a regulatory gene of anthocyanin biosynthesis in tuberous roots of purple-fleshed sweet potato. Plant Physiol , 143(3): 1252–1268 doi: 10.1104/pp.106.094425 pmid:17208956
85	Martens S, Mith?fer A (2005). Flavones and flavone synthases. Phytochemistry , 66(20): 2399–2407 doi: 10.1016/j.phytochem.2005.07.013 pmid:16137727
86	Martens S, Preuss A, Matern U (2010). Multifunctional flavonoid dioxygenases: flavonol and anthocyanin biosynthesis in Arabidopsis thaliana L. Phytochemistry , 71(10): 1040–1049 doi: 10.1016/j.phytochem.2010.04.016 pmid:20457455
87	Martin C, Gerats T (1993). Control of pigment biosynthesis genes during petal development. Plant Cell , 5(10): 1253–1264 pmid:12271025
88	Martin C, Prescott A, Mackay S, Bartlett J, Vrijlandt E (1991). Control of anthocyanin biosynthesis in flowers of Antirrhinum majus. Plant J , 1(1): 37–49 doi: 10.1111/j.1365-313X.1991.00037.x pmid:1844879
89	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, Wagner D R (2003). Activation tagging in tomato identifies a transcriptional regulator of anthocyanin biosynthesis, modification, and transport. Plant Cell , 15(8): 1689–1703 doi: 10.1105/tpc.012963 pmid:12897245
90	Matsui K, Umemura Y, Ohme-Takagi M (2008). AtMYBL2, a protein with a single MYB domain, acts as a negative regulator of anthocyanin biosynthesis in Arabidopsis. Plant J , 55(6): 954–967 doi: 10.1111/j.1365-313X.2008.03565.x pmid:18532977
91	Matus J T, Loyola R, Vega A, Pe?a-Neira A, Bordeu E, Arce-Johnson P, Alcalde J A (2009). Post-veraison sunlight exposure induces MYB-mediated transcriptional regulation of anthocyanin and flavonol synthesis in berry skins of Vitis vinifera. J Exp Bot , 60(3): 853–867 doi: 10.1093/jxb/ern336 pmid:19129169
92	Matus J T, Poupin M J, Ca?ón P, Bordeu E, Alcalde J A, Arce-Johnson P (2010). Isolation of WDR and bHLH genes related to flavonoid synthesis in grapevine (Vitis vinifera L.). Plant Mol Biol , 72(6): 607–620 doi: 10.1007/s11103-010-9597-4 pmid:20112051
93	Mehrtens F, Kranz H, Bednarek P, Weisshaar B (2005). The Arabidopsis transcription factor MYB12 is a flavonol-specific regulator of phenylpropanoid biosynthesis. Plant Physiol , 138(2): 1083–1096 doi: 10.1104/pp.104.058032 pmid:15923334
94	Mellway R D, Tran L T, Prouse M B, Campbell M M, Constabel C P (2009). The wound-, pathogen-, and ultraviolet B-responsive MYB134 gene encodes an R2R3 MYB transcription factor that regulates proanthocyanidin synthesis in poplar. Plant Physiol , 150(2): 924–941 doi: 10.1104/pp.109.139071 pmid:19395405
95	Miranda M, Ralph S G, Mellway R, White R, Heath M C, Bohlmann J, Constabel C P (2007). The transcriptional response of hybrid poplar (Populus trichocarpa x P. deltoides) to infection by Melampsora medusae leaf rust involves induction of flavonoid pathway genes leading to the accumulation of proanthocyanidins. Mol Plant Microbe Interact , 20(7): 816–831 doi: 10.1094/MPMI-20-7-0816 pmid:17601169
96	Mol J, Grotewold E, Koes R (1998). How genes paint flowers and seeds. Trends Plant Sci , 3(6): 212–217 doi: 10.1016/S1360-1385(98)01242-4
97	Morita Y, Saitoh M, Hoshino A, Nitasaka E, Iida S (2006). 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. Plant Cell Physiol , 47(4): 457–470 doi: 10.1093/pcp/pcj012 pmid:16446312
98	Morohashi K, Casas M I, Falcone Ferreyra M L, Mejía-Guerra M K, Pourcel L, Yilmaz A, Feller A, Carvalho B, Emiliani J, Rodriguez E, Pellegrinet S, McMullen M, Casati P, Grotewold E (2012). A genome-wide regulatory framework identifies maize pericarp color1 controlled genes. Plant Cell , 24(7): 2745–2764 doi: 10.1105/tpc.112.098004 pmid:22822204
99	Moyano E, Martínez-Garcia J F, Martin C (1996). Apparent redundancy in myb gene function provides gearing for the control of flavonoid biosynthesis in antirrhinum flowers. Plant Cell , 8(9): 1519–1532 pmid:8837506
100	Nakatsuka T, Haruta K S, Pitaksutheepong C, Abe Y, Kakizaki Y, Yamamoto K, Shimada N, Yamamura S, Nishihara M (2008). Identification and characterization of R2R3-MYB and bHLH transcription factors regulating anthocyanin biosynthesis in gentian flowers. Plant Cell Physiol , 49(12): 1818–1829 doi: 10.1093/pcp/pcn163 pmid:18974195
101	Nakatsuka T, Nishihara M, Mishiba K, Yamamura S (2005). Temporal expression of flavonoid biosynthesis-related genes regulates flower pigmentation in gentian plants. Plant Sci , 168(5): 1309–1318 doi: 10.1016/j.plantsci.2005.01.009
102	Nakatsuka T, Saito M, Yamada E, Fujita K, Kakizaki Y, Nishihara M (2012). Isolation and characterization of GtMYBP3 and GtMYBP4, orthologues of R2R3-MYB transcription factors that regulate early flavonoid biosynthesis, in gentian flowers. J Exp Bot , 63(18): 6505–6517 doi: 10.1093/jxb/ers306 pmid:23125348
103	Nesi N, Debeaujon I, Jond C, Pelletier G, Caboche M, Lepiniec L (2000). The TT8 gene encodes a basic helix-loop-helix domain protein required for expression of DFR and BAN genes in Arabidopsis siliques. Plant Cell , 12(10): 1863–1878 pmid:11041882
104	Nesi N, Jond C, Debeaujon I, Caboche M, Lepiniec L (2001). The Arabidopsis TT2 gene encodes an R2R3 MYB domain protein that acts as a key determinant for proanthocyanidin accumulation in developing seed. Plant Cell , 13(9): 2099–2114 pmid:11549766
105	Nishihara M, Nakatsuka T (2011). Genetic engineering of flavonoid pigments to modify flower color in floricultural plants. Biotechnol Lett , 33(3): 433–441 doi: 10.1007/s10529-010-0461-z pmid:21053046
106	Niu S S, Xu C J, Zhang W S, Zhang B, Li X, Lin-Wang K, Ferguson I B, Allan A C, Chen K S (2010). Coordinated regulation of anthocyanin biosynthesis in Chinese bayberry (Myrica rubra) fruit by a R2R3 MYB transcription factor. Planta , 231(4): 887–899 doi: 10.1007/s00425-009-1095-z pmid:20183921
107	Noda N, Kanno Y, Kato N, Kazuma K, Suzuki M (2004). Regulation of gene expression involved in flavonol and anthocyanin biosynthesis during petal development in lisianthus (Eustoma grandiflorum). Physiol Plant , 122(3): 305–313 doi: 10.1111/j.1399-3054.2004.00407.x
108	Ogata K, Hojo H, Aimoto S, Nakai T, Nakamura H, Sarai A, Ishii S, Nishimura Y (1992). Solution structure of a DNA-binding unit of Myb: a helix-turn-helix-related motif with conserved tryptophans forming a hydrophobic core. Proc Natl Acad Sci U S A , 89(14): 6428–6432 doi: 10.1073/pnas.89.14.6428 pmid:1631139
109	Ogata K, Morikawa S, Nakamura H, Sekikawa A, Inoue T, Kanai H, Sarai A, Ishii S, Nishimura Y (1994). Solution structure of a specific DNA complex of the Myb DNA-binding domain with cooperative recognition helices. Cell , 79(4): 639–648 doi: 10.1016/0092-8674(94)90549-5 pmid:7954830
110	Osier T L, Lindroth R L (2001). Effects of genotype, nutrient availability, and defoliation on aspen phytochemistry and insect performance. J Chem Ecol , 27(7): 1289–1313 doi: 10.1023/A:1010352307301 pmid:11504029
111	Palapol Y, Ketsa S, Lin-Wang K, Ferguson I B, Allan A C (2009). A MYB transcription factor regulates anthocyanin biosynthesis in mangosteen (Garcinia mangostana L.) fruit during ripening. Planta , 229(6): 1323–1334 doi: 10.1007/s00425-009-0917-3 pmid:19306102
112	Pattanaik S, Kong Q, Zaitlin D, Werkman J R, Xie C H, Patra B, Yuan L (2010). Isolation and functional characterization of a floral tissue-specific R2R3 MYB regulator from tobacco. Planta , 231(5): 1061–1076 doi: 10.1007/s00425-010-1108-y pmid:20157728
113	Paz-Ares J, Ghosal D, Wienand U, Peterson P A, Saedler H (1987). The regulatory c1 locus of Zea mays encodes a protein with homology to myb proto-oncogene products and with structural similarities to transcriptional activators. EMBO J , 6(12): 3553–3558 pmid:3428265
114	Peel G J, Pang Y, Modolo L V, Dixon R A (2009). The LAP1 MYB transcription factor orchestrates anthocyanidin biosynthesis and glycosylation in Medicago. Plant J , 59(1): 136–149 doi: 10.1111/j.1365-313X.2009.03885.x pmid:19368693
115	Peters D J, Constabel C P (2002). Molecular analysis of herbivore-induced condensed tannin synthesis: cloning and expression of dihydroflavonol reductase from trembling aspen (Populus tremuloides). Plant J , 32(5): 701–712 doi: 10.1046/j.1365-313X.2002.01458.x pmid:12472686
116	Quattrocchio F, Verweij W, Kroon A, Spelt C, Mol J, Koes R (2006). 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. Plant Cell , 18(5): 1274–1291 doi: 10.1105/tpc.105.034041 pmid:16603655
117	Quattrocchio F, Wing J, van der Woude K, Souer E, de Vetten N, Mol J, Koes R (1999). Molecular analysis of the anthocyanin2 gene of petunia and its role in the evolution of flower color. Plant Cell , 11(8): 1433–1444 pmid:10449578
118	Quattrocchio F, Wing J F, van der Woude K, Mol J N, Koes R (1998). Analysis of bHLH and MYB domain proteins: species-specific regulatory differences are caused by divergent evolution of target anthocyanin genes. Plant J , 13(4): 475–488 doi: 10.1046/j.1365-313X.1998.00046.x pmid:9680994
119	Rabinowicz P D, Braun E L, Wolfe A D, Bowen B, Grotewold E (1999). Maize R2R3 Myb genes: Sequence analysis reveals amplification in the higher plants. Genetics , 153(1): 427–444 pmid:10471724
120	Ravaglia D, Espley R V, Henry-Kirk R A, Andreotti C, Ziosi V, Hellens R P, Costa G, Allan A C (2013). Transcriptional regulation of flavonoid biosynthesis in nectarine (Prunus persica) by a set of R2R3 MYB transcription factors. BMC Plant Biol , 13(1): 68 doi: 10.1186/1471-2229-13-68 pmid:23617716
121	Roth B A, Goff S A, Klein T M, Fromm M E (1991). C1- and R-dependent expression of the maize Bz1 gene requires sequences with homology to mammalian myb and myc binding sites. Plant Cell , 3(3): 317–325 pmid:1840914
122	Routaboul J M, Kerhoas L, Debeaujon I, Pourcel L, Caboche M, Einhorn J, Lepiniec L (2006). Flavonoid diversity and biosynthesis in seed of Arabidopsis thaliana. Planta , 224(1): 96–107 doi: 10.1007/s00425-005-0197-5 pmid:16395586
123	Sablowski R W, Moyano E, Culianez-Macia F A, Schuch W, Martin C, Bevan M (1994). A flower-specific Myb protein activates transcription of phenylpropanoid biosynthetic genes. EMBO J , 13(1): 128–137 pmid:8306956
124	Saikumar P, Murali R, Reddy E P (1990). Role of tryptophan repeats and flanking amino acids in Myb-DNA interactions. Proc Natl Acad Sci U S A , 87(21): 8452–8456 doi: 10.1073/pnas.87.21.8452 pmid:2236054
125	Sainz M B, Grotewold E, Chandler V L (1997). Evidence for direct activation of an anthocyanin promoter by the maize C1 protein and comparison of DNA binding by related Myb domain proteins. Plant Cell , 9(4): 611–625 pmid:9144964
126	Saito R, Fukuta N, Ohmiya A, Itoh Y, Ozeki Y, Kuchitsu K, Nakayama M (2006). Regulation of anthocyanin biosynthesis involved in the formation of marginal picotee petals in Petunia. Plant Sci , 170(4): 828–834 doi: 10.1016/j.plantsci.2005.12.003
127	Sasaki M, Ogata K, Hatanaka H, Nishimura Y (2000). Backbone dynamics of the c-Myb DNA-binding domain complexed with a specific DNA. J Biochem , 127(6): 945–953 doi: 10.1093/oxfordjournals.jbchem.a022710 pmid:10833261
128	Schaart J G, Dubos C, Romero De La Fuente I, van Houwelingen A M, de Vos R C, Jonker H H, Xu W, Routaboul J M, Lepiniec L, Bovy A G (2013). Identification and characterization of MYB-bHLH-WD40 regulatory complexes controlling proanthocyanidin biosynthesis in strawberry (Fragariaβ×βananassa) fruits. New Phytol , 197(2): 454–467 doi: 10.1111/nph.12017 pmid:23157553
129	Schr?der G, Brown J W, Schr?der J (1988). Molecular analysis of resveratrol synthase. cDNA, genomic clones and relationship with chalcone synthase. Eur J Biochem , 172(1): 161–169 doi: 10.1111/j.1432-1033.1988.tb13868.x pmid:2450022
130	Schwinn K, Venail J, Shang Y, Mackay S, Alm V, Butelli E, Oyama R, Bailey P, Davies K, Martin C (2006). A small family of MYB-regulatory genes controls floral pigmentation intensity and patterning in the genus Antirrhinum. Plant Cell , 18(4): 831–851 doi: 10.1105/tpc.105.039255 pmid:16531495
131	Sharma M, Chai C, Morohashi K, Grotewold E, Snook M E, Chopra S (2012). Expression of flavonoid 3′-hydroxylase is controlled by P1, the regulator of 3-deoxyflavonoid biosynthesis in maize. BMC Plant Biol , 12(1): 196 doi: 10.1186/1471-2229-12-196 pmid:23113982
132	Solano R, Fuertes A, Sánchez-Pulido L, Valencia A, Paz-Ares J (1997). A single residue substitution causes a switch from the dual DNA binding specificity of plant transcription factor MYB.Ph3 to the animal c-MYB specificity. J Biol Chem , 272(5): 2889–2895 doi: 10.1074/jbc.272.5.2889 pmid:9006933
133	Solano R, Nieto C, Avila J, Ca?as L, Diaz I, Paz-Ares J (1995). Dual DNA binding specificity of a petal epidermis-specific MYB transcription factor (MYB.Ph3) from Petunia hybrida. EMBO J , 14(8): 1773–1784 pmid:7737128
134	Spelt C, Quattrocchio F, Mol J N, Koes R (2000). anthocyanin1 of petunia encodes a basic helix-loop-helix protein that directly activates transcription of structural anthocyanin genes. Plant Cell , 12(9): 1619–1632 pmid:11006336
135	Springob K, Nakajima J, Yamazaki M, Saito K (2003). Recent advances in the biosynthesis and accumulation of anthocyanins. Nat Prod Rep , 20(3): 288–303 doi: 10.1039/b109542k pmid:12828368
136	Stracke R, Ishihara H, Huep G, Barsch A, Mehrtens F, Niehaus K, Weisshaar B (2007). Differential regulation of closely related R2R3-MYB transcription factors controls flavonol accumulation in different parts of the Arabidopsis thaliana seedling. Plant J , 50(4): 660–677 doi: 10.1111/j.1365-313X.2007.03078.x pmid:17419845
137	Stracke R, Jahns O, Keck M, Tohge T, Niehaus K, Fernie A R, Weisshaar B (2010). 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 , 188(4): 985–1000 doi: 10.1111/j.1469-8137.2010.03421.x pmid:20731781
138	Stracke R, Werber M, Weisshaar B (2001). The R2R3-MYB gene family in Arabidopsis thaliana. Curr Opin Plant Biol , 4(5): 447–456 doi: 10.1016/S1369-5266(00)00199-0 pmid:11597504
139	Tahirov T H, Sato K, Ichikawa-Iwata E, Sasaki M, Inoue-Bungo T, Shiina M, Kimura K, Takata S, Fujikawa A, Morii H, Kumasaka T, Yamamoto M, Ishii S, Ogata K (2002). Mechanism of c-Myb-C/EBP beta cooperation from separated sites on a promoter. Cell , 108(1): 57–70 doi: 10.1016/S0092-8674(01)00636-5 pmid:11792321
140	Takos A M, Jaffé F W, Jacob S R, Bogs J, Robinson S P, Walker A R (2006). Light-induced expression of a MYB gene regulates anthocyanin biosynthesis in red apples. Plant Physiol , 142(3): 1216–1232 doi: 10.1104/pp.106.088104 pmid:17012405
141	Tamagnone L, Merida A, Parr A, Mackay S, Culianez-Macia F A, Roberts K, Martin C (1998). The AmMYB308 and AmMYB330 transcription factors from antirrhinum regulate phenylpropanoid and lignin biosynthesis in transgenic tobacco. Plant Cell , 10(2): 135–154 pmid:9490739
142	Tanaka Y, Brugliera F, Chandler S (2009). Recent progress of flower colour modification by biotechnology. Int J Mol Sci , 10(12): 5350–5369 doi: 10.3390/ijms10125350 pmid:20054474
143	Terrier N, Torregrosa L, Ageorges A, Vialet S, Verriès C, Cheynier V, Romieu C (2009). Ectopic expression of VvMybPA2 promotes proanthocyanidin biosynthesis in grapevine and suggests additional targets in the pathway. Plant Physiol , 149(2): 1028–1041 doi: 10.1104/pp.108.131862 pmid:19098092
144	Tuerck J A, Fromm M E (1994). Elements of the maize A1 promoter required for transactivation by the anthocyanin B/C1 or phlobaphene P regulatory genes. Plant Cell , 6(11): 1655–1663 pmid:7827497
145	Uimari A, Strommer J (1997). Myb26: a MYB-like protein of pea flowers with affinity for promoters of phenylpropanoid genes. Plant J , 12(6): 1273–1284 doi: 10.1046/j.1365-313x.1997.12061273.x pmid:9450341
146	Verdier J, Zhao J, Torres-Jerez I, Ge S, Liu C, He X, Mysore K S, Dixon R A, Udvardi M K (2012). MtPAR MYB transcription factor acts as an on switch for proanthocyanidin biosynthesis in Medicago truncatula. Proc Natl Acad Sci U S A , 109(5): 1766–1771 doi: 10.1073/pnas.1120916109 pmid:22307644
147	Waiss A C, Chan B G, Elliger C A, Wiseman B R, McMillian W W, Widstrom N W, Zuber M S, Keaster A J (1979). Maysin, a flavone glycoside from corn silks with antibiotic activity toward corn earworm. J Econ Entomol , 72: 256–258
148	Wang X (2011). Structure, function, and engineering of enzymes in isoflavonoid biosynthesis. Funct Integr Genomics , 11(1): 13–22 doi: 10.1007/s10142-010-0197-9 pmid:21052759
149	Wei Y L, Li J N, Lu J, Tang Z L, Pu D C, Chai Y R (2007). Molecular cloning of Brassica napus TRANSPARENT TESTA 2 gene family encoding potential MYB regulatory proteins of proanthocyanidin biosynthesis. Mol Biol Rep , 34(2): 105–120 doi: 10.1007/s11033-006-9024-8 pmid:17115250
150	Winkel-Shirley B (2001). Flavonoid biosynthesis. A colorful model for genetics, biochemistry, cell biology, and biotechnology. Plant Physiol , 126(2): 485–493 doi: 10.1104/pp.126.2.485 pmid:11402179
151	Winkel-Shirley B (2002). Biosynthesis of flavonoids and effects of stress. Curr Opin Plant Biol , 5(3): 218–223 doi: 10.1016/S1369-5266(02)00256-X pmid:11960739
152	Xie D Y, Dixon R A (2005). Proanthocyanidin biosynthesis—still more questions than answers? Phytochemistry , 66(18): 2127–2144 doi: 10.1016/j.phytochem.2005.01.008 pmid:16153412
153	Yamagishi M, Shimoyamada Y, Nakatsuka T, Masuda K (2010). Two R2R3-MYB genes, homologs of Petunia AN2, regulate anthocyanin biosyntheses in flower Tepals, tepal spots and leaves of asiatic hybrid lily. Plant Cell Physiol , 51(3): 463–474 doi: 10.1093/pcp/pcq011 pmid:20118109
154	Yi J, Derynck M R, Li X, Telmer P, Marsolais F, Dhaubhadel S (2010). A single-repeat MYB transcription factor, GmMYB176, regulates CHS8 gene expression and affects isoflavonoid biosynthesis in soybean. Plant J , 62(6): 1019–1034 pmid:20345602
155	Yoshida K, Iwasaka R, Kaneko T, Sato S, Tabata S, Sakuta M (2008). Functional differentiation of Lotus japonicus TT2s, R2R3-MYB transcription factors comprising a multigene family. Plant Cell Physiol , 49(2): 157–169 doi: 10.1093/pcp/pcn009 pmid:18202001
156	Zhang B, Hu Z, Zhang Y, Li Y, Zhou S, Chen G (2012). A putative functional MYB transcription factor induced by low temperature regulates anthocyanin biosynthesis in purple kale (Brassica Oleracea var. acephala f. tricolor). Plant Cell Rep , 31(2): 281–289 doi: 10.1007/s00299-011-1162-3 pmid:21987119
157	Zhang P, Chopra S, Peterson T (2000). A segmental gene duplication generated differentially expressed myb-homologous genes in maize. Plant Cell , 12(12): 2311–2322 pmid:11148280
158	Zhang P, Wang Y, Zhang J, Maddock S, Snook M, Peterson T (2003). A maize QTL for silk maysin levels contains duplicated Myb-homologous genes which jointly regulate flavone biosynthesis. Plant Mol Biol , 52(1): 1–15 doi: 10.1023/A:1023942819106 pmid:12825685
159	Zhao L, Gao L, Wang H, Chen X, Wang Y, Yang H, Wei C, Wan X, Xia T (2013). The R2R3-MYB, bHLH, WD40, and related transcription factors in flavonoid biosynthesis. Funct Integr Genomics , 13(1): 75–98 doi: 10.1007/s10142-012-0301-4 pmid:23184474
160	Zifkin M, Jin A, Ozga J A, Zaharia L I, Schernthaner J P, Gesell A, Abrams S R, Kennedy J A, Constabel C P (2012). Gene expression and metabolite profiling of developing highbush blueberry fruit indicates transcriptional regulation of flavonoid metabolism and activation of abscisic acid metabolism. Plant Physiol , 158(1): 200–224 doi: 10.1104/pp.111.180950 pmid:22086422

[1]	Yujie Deng, Caixia Lin, Huanjiao Jenny Zhou, Wang Min. Smooth muscle cell differentiation: Mechanisms and models for vascular diseases[J]. Front. Biol., 2017, 12(6): 392-405.
[2]	Madhuchhanda Das, Harischandra Sripathy Prakash, Monnanda Somaiah Nalini. *Antioxidative properties of phenolic compounds isolated from the fungal endophytes of Zingiber nimmonii* (J. Graham) Dalzell.**[J]. Front. Biol., 2017, 12(2): 151-162.
[3]	Pathomwat Wongrattanakamon,Vannajan Sanghiran Lee,Piyarat Nimmanpipug,Supat Jiranusornkul. Nucleotide binding domain 1 pharmacophore modeling for visualization and analysis of P-glycoprotein–flavonoid molecular interactions[J]. Front. Biol., 2016, 11(5): 391-395.
[4]	Rebecca M. FOX,Deborah J. ANDREW. Transcriptional regulation of secretory capacity by bZip transcription factors[J]. Front. Biol., 2015, 10(1): 28-51.
[5]	Yaojuan LU,Longwei QIAO,Guanghua LEI,Ranim R. MIRA,Junxia GU,Qiping ZHENG. Col10a1 gene expression and chondrocyte hypertrophy during skeletal development and disease[J]. Front. Biol., 2014, 9(3): 195-204.
[6]	Francesca FANINI, Ivan VANNINI, Muller FABBRI. MicroRNAs and drug modulation in cancer: an intertwined new story[J]. Front Biol, 2011, 6(5): 351-356.
[7]	Afsar U. AHMED. An overview of inflammation: mechanism and consequences[J]. Front Biol, 2011, 6(4): 274-281.
[8]	Peng ZHAO, Dong-Qiao SHI, Wei-Cai YANG. Patterning the embryo in higher plants: Emerging pathways and challenges[J]. Front Biol, 2011, 06(01): 3-11.
[9]	YUAN Wuzhou, HUANG Xinqiong, ZHU Chuanbing, WANG Yuequn, LI Yongqing, WU Xiushan. Role of the domains of human gene ZNF569 in MAPK pathway[J]. Front. Biol., 2006, 1(4): 357-361.

Viewed

Full text

Abstract

Cited

Shared

Discussed