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Award Review
Institute for Plant Science, Suntory Holdings Ltd., 1-1-1 Wakayamadai, Shimamoto, Osaka 618-8503, Japan
Florigene Pty Ltd., 1 Park Drive, Bundoora, Victoria 3083, Australia
This review was written in response to the corresponding authors receipt of The Japan Prize of Agricultural Science in 2009.
To whom correspondence should be addressed. Tel: +81-75-962-8807; Fax: +81-75-962-3791; E-mail:Yoshikazu Tanaka@suntory.co.jp
Abbreviations: THC, 20 ,40 ,60 ,4-tetrahydroxychalcone; ODG, 2-oxoglutarate dependent dioxygenase; DHK, dihydrokaempferol; FNS, avone
synthase; P450, cytochrome P450; F2H, avanone 2-hydroxylase; F30 H, avonoid 30 -hydroxylase; F30 50 H, avonoid 30 ,50 -hydroxylase; FLS, avonol
synthase; DFR, dihydroavonol 4-reductase; DHM, dihydromyricetin; ANS, anthocyanidin synthase; F3GT, UDP-glucose: avonoid
3-glucosyltransferae; AT, acyltransferase; GST, glutathione S-transferase; MRP, multidrug resistance-associated protein; ABC, ATP binding
cassette; MATE, multidrug and toxic extrusion; bHLH, basic helix loop helix; LAR, leucoanthocyanidin reductase; ANR, anthocyanidin reductase
y
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Fig. 1. Schematic of the General Flavonoid Biosynthetic Pathway Relevant to Flower Color.
Anthocyanidin is further modied with glycosyl, acyl, or methyl groups catalyzed by glycosyltransferase (GT), acyltransferase (AT), and
methyltransferase (MT). Classication of avonoids is shown in parentheses. Abbreviations: CHS, chalcone synthase; F3H, avanone
3-hydroxylase; F30 H, avonoid 30 -hydroxylase; F30 50 H, avonoid 30 ,50 -hydroxylase; DFR, dihydroavonol 4-reductase; ANS, anthocyanidin
synthase; THC40 GT, tetrahydroxychalcone 40 -glucosyltransferase; AS, aurusidin synthase; FLS, avonol synthasae; FNS, avone synthase;
LAR, leucoanthocyanidin reductase; ANR, anthocyanidin reductase.
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Y. TANAKA et al.
protein, R3-MYB (MYBL2), suppresses avonoid biosynthesis by interacting with a MBW complex. The loss
of MYBL2 resulted in a dramatic increase in anthocyanins in Arabidopsis seedlings.58,59) Overexpression of
MYBL2 inhibited the biosynthesis of proanthocyanidins
in seeds.58) Plant-specic NAC transcriptional factor has
been found to regulate anthocyanin biosynthesis. An
Arabidopsis ANAC078 induced anthocyanin accumulation by activating anthocyanin biosynthetic genes via
transcriptional factors regulating biosynthesis under
strong-light conditions.60)
5. Unsolved problems in avonoid biosynthesis and
metabolism
The avonoid biosynthetic pathway is probably the
most thoroughly studied plant secondary metabolism
pathway. Except for the glucosyltransferases and ATs
that are necessary for polyacyl anthocyanins, most of the
enzymatic genes have been characterized. Isolation of
the genes of these enzymes should provide more
molecular tools to engineer anthocyanin structure and
ower color. It is not clear whether these modications
occur in the cytosol or in the vacuole or on membranes.
The enzymes involved in some plant metabolic
pathways, including avonoid biosynthesis, perhaps
form a macromolecular complex (metabolon) for metabolic channeling and ecient biosynthesis.61) Metabolons are thought to be anchored onto the ER membrane
utilizing cytochromes P450 such as F30 H.61) More
experimental data are necessary to conrm this. If a
metabolon is necessary for ecient synthesis of anthocyanins, an enzyme from an exogenous gene must be
compatible with the endogenous metabolon.
The petals of some plants fade or lose color during
development. Anthocyanin degradation and disappearance have been reviewed,62) and also is important in
terms of engineering ower color. Further understanding
of these problems should contribute to engineering of
the pathway.
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Rose
The cultivated rose (Rosa hybrida) was bred from
about eight wild species, including R. chinensis,
R. galica, R. gigantia, R. moschata, R. multiora,
R. wichuraiana, and R. foetida after extensive interspecic hybridization (http://www.gifu-u.ac.jp/fukui/
02-1-2-2.htm). Pink and red colors are derived from
pelargonidin or cyanidin-based anthocyanins. The expression of a pansy (Viola spp) F30 50 H gene resulted in
signicant amounts of delphinidin-derived anthocyanins accumulating in the petals of the transgenic plants.
Expression of the pansy F30 50 H gene in rose cultivars
that have higher vacuolar pH, relatively large amounts
of avonols (co-pigments), and weak or no F30 H
activity resulted in transgenic lines in which 95% of
the anthocyanidins was delphinidin. The colors of the
owers in these lines were of a signicantly bluer hue
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Y. TANAKA et al.
63)
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Line numbers
Color stability
SWB/308-25
SWB/308-35
SWB/308-36
SWB/805-27
SWB/805-36
SWB/1322-45
SWB/1322-90
SWV/1341-17
SWV/1341-85
Unstable
Unstable
Unstable
Stable
Unstable
Stable
Stable
Stable
Unstable
Growth compared
to the host
Poor
Poor
Poor
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Y. TANAKA et al.
Acknowledgments
We wish to thank the current and former members of
Suntory and Florigene for their contributions, over many
years. In order to focus on recent progress, only a
limited number of original papers are cited here.
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