GENETICS OF FLOWER COLOUR
Important pigments responsible for flowers’ colours are flavonoids (in particular anthocyanins,
anthocyanidins, favonols and favones) and carotenoids (carotenes and xanthophylls). Other compounds
involved in pigmentation are betalains (betacyanins and betaxanthins and chlorophylls (α and β).
Pigments have proper subcellular location and are mainly located in the upper epidermal cells of petals.
Flavonoids are accumulated into vacuoles, whereas carotenoids are deposited in plastids. The
morphology of petal epidermal cells contributes to flower colour: when the light penetrate and
adsorbed into conical cells, flowers will show dark colour, whereas fat cells reflect light promoting pale
flowers.
Flavonoids have been extensively studied in ornamental species, petunia, herbaceous peony,
Torenia fournieri, Ipomoea, C. morifolium and rose. Flavonoids containing anthocyanidins provide
colours as orange red (pelargonidin), purplish red (cyanidin), blue (delphinidin), purple violet (malvidin),
rose (peonidin), and purple violet (petunidin), while those containing anthoxanthins give rise to white,
light and dark yellow. The anthocyanin biosynthetic cascade is well known and three steps are
characterized by specifc genes:
1) phenylalanine ammonia lyase (PAL), cinnamate-4-hydroxylase (C4H), and 4-coumarate:CoA ligase
(4CL);
2) chalcone synthase (CHS), chalcone isomerase (CHI), flavanone-3-hydroxylase (F3H), flavonoid 3′-
hydroxylase (F3′H), and favonoid 3′,5′-hydroxylase (F3′5′H);
3) dihydroflavonol 4-reductase (DFR), anthocyanidin synthase (ANS), flavonoid glucosyltransferase
(UFGT), and methyl transferase (MT).
CHS is the first enzyme in the flavonoids metabolic flow and it is ubiquitous in ornamental
plants. Down-regulation with antisense CHS construct caused the complete loss of pigmentation
resulting in white flowers in petunia and chrysanthemum. Petunia transformed with CHS1 of Freesia
resulted in new plants with pink flowers. In rose and carnation, F3′5′H gene is not present; hence they
do not have bluish petals. Since carotenoids are involved in many other important aspects of the plant
(they are involved in photoprotection, nutraceutical value, and are precursors of vitamin A and of
abscisic acid), the key genes have been identified and characterized: phytoene synthase (PSY), phytoene
desaturase (PDS), zeta-carotene desaturase (ZDS), carotenoid isomerase (CRTISO), lycopene epsilon-
cyclase (LCYE) and lycopene beta-cyclase (LCYB).
GENETICS OF FLOWER FRAGRANCE
The fragrance of flowers is specific to each plant and is made up of a mixture of small (100–250 Da)
volatile molecules that mostly belong to three groups of compounds: terpenoids, phenylpropanoids /
benzenoids, and fatty acid derivatives. The VOCs (volatile organic compounds) are synthetized mainly in
flowers (largely in the petals).
Terpenoids, divided into monoterpenoids and sesquiterpenoids, are the largest class of floral volatiles.
They derive from the MVA (mevalonic acid) and MEP (methylerythritol phosphate) pathways in the
cytosol and plastids, respectively. Terpene synthases (TPSs) are the key enzymes in the biosynthetic
pathways of terpenoids. Monoterpenes, which include limonene, (E)-b-ocimene, myrcene, linalool, and
alpha- and beta-pinene, are commonly found in floral volatiles of lilium, rose, orchid, snapdragon,
petunia, Freesia, Alstroemeria, Clarkia, and Iris.
The second major class of plant VOCs are represented by phenylpropanoids and benzenoids and derive
from shikimic acid via phenylanaline, cinnamic acid and further decarboxylation and ring oxidation. They
characterize the floral scent of different species such as snapdragon, petunia and rose.
The third class of flower VOCs are the fatty acid derivatives, which are synthesized via the lipoxygenase
pathway from the unsaturated C18 linoleic and linolenic fatty acids. The products, short chain alcohols
and aldehydes, are important constituents in the floral volatile perfume of several plant species such as
carnation, wild snapdragon and orchids of genus Ophrys.
‘SCENTbase’ and ‘SuperScent’ are two databases in which plant volatile chemicals are collected. Prior to
the development of high-throughput technologies, only classic biochemical approaches could be applied
to isolate floral scent genes. Transcriptome and genome analysis of large numbers of active genes
combined with complex metabolite networks and advances in gas chromatography–mass spectrometry
(GC–MS) analysis allowed the identification of novel scent genes in several ornamental species (ESM_5).
Garden roses are selected primarily for fragrance, whereas marketed roses bred for cut flower
production often lack perfume. In reality, even fragrance-free roses emit small quantities of fragrant
molecules, indicating they have not completely lost their ability to produce them, but according to a
recent hypothesis, it would be a malfunction of the biosynthetic pathway of VOCs. Unfortunately, the
biosynthetic pathways of many rose scent compounds are not completely known. To identify genes
involved in rose scent, high-throughput transcriptomic approaches were applied to petal tissues
(ESM_5).
The phenolic methyl ethers were found responsible for the characteristic sweet “tea scent” smell of
some chinese roses. This unique scent is rich in 3,5-dimethoxytoluene (DMT), that is synthesized by two
specific enzymes, orcinol-O-methyl transferases 1 and 2 (OOMT1 and OOMT2). The key enzymes
phenylacetaldehyde synthase and phenylacetaldehyde reductase were identified based on a detailed
study of the main 2-phenylethanol (2PE) pathway. Eugenol synthase (RcEGS1) gene cloned from the
petals of R. chinensis ‘Old Blush’ is involved in the biosynthesis of eugenol in rose. Terpenoids, especially
monoterpene alcohols such as geraniol, are central molecules for the ‘typical rose scent’. Nevertheless,
only few genes of the terpenoid pathway have been characterized: a sesquiterpene synthase, for the
biosynthesis of germacrene D, and two monoterpene synthases for linalool and nerolidol, although
these are not the main terpenoids in rose. It has been recently discovered that rose uses a terpene
synthase independent pathway to produce geraniol.
In lilium “Siberia”, a commercially important cv appreciated by the consumer for its fragrance and large
and snowy white flowers, two monoterpene synthase genes (LoTPS1 and LoTPS3), responsible for
biosynthesis of β-ocimene and linalool, were isolated and functionally characterized. These
monoterpenes compounds were released following a diurnal circadian rhythm. The expression of both
genes occurs in sepals and petals in response to mechanical injury.
In orchids, the monoterpenoid biosynthesis pathway and responsible genes in Phalaenopsis bellina and
P. equestris have been reported; in particular, the tissue and temporal expression of glyceraldehyde-3-
phosphate (GDPS), precursor of geraniol and linalool has been elucidated. In the Vanda orchid, a new 1-
deoxy-D-xylulose 5-phosphate reductoisomerase (DXR) transcript was reported and two fragrance-
related transcripts, VMPAAT encoding a putative alcohol acyltransferase and VMPSTS encoding a
sesquiterpene synthase, were cloned and characterized
GENETICS OF FLOWER SHELF LIFE
Increasing vase life is a key factor to convince consumers to re-purchase cut flowers. To extend the life
of the flower after harvesting, it is necessary to act on several factors before harvesting, including
genotype choice. In many flowers, post-harvest life depends on the appearance of the first symptoms of
petal senescence which show the characteristic "in-rolling" behavior, but also other symptoms may also
appear such as wilting, colour change, petal and bud abscission These symptoms can be delayed by
treatments that block the biosynthesis or action of ethylene. Quite often, the quality and vase life of
flowers is related to the amount of exogenous and endogenous ethylene, a phytohormone that plays a
central role in various metabolic plant processes, such as seed germination, but also flower and leaf
senescence. The most important cut flower species, such as rose, lilium, lisianthus, chrysanthemum or
carnation show different levels of sensitivity to ethylene and even different flower longevity. In many of
these species, breeding approaches, based on cross-pollination coupled with ethylene screening, have
been applied with little success. An alternative solution to improve ornamental’s vase life can be the
exploitation of molecular approaches (Netam 2018). In carnation, transcripts for the various steps of the
ethylene biosynthetic pathway have been identified by transcriptomic analysis. In Rosa hybrida,
ethylene-responsive transcript, potential key regulators of ethylene-influenced cell expansion, were also
identified using a floral transcriptome. In the last years, however, many studies focused on the
knowledge of ethylene biosynthesis. Onozaki et al. (2011), by repeated selection for longer vase life,
produced two carnation cultivars improved for that trait. Their flowers showed a reduced expression of
the three ethylene biosynthesis genes: 1-aminocyclopropane-1-carboxylate synthase (DcACS1 and
DcACS2) and 1-aminocyclopropane-1-carboxylate oxidase (DcACO1). Further, transgenic plants with
longer vase life were obtained by introducing genes encoding 1-aminocyclopropane-1-carboxylate (ACC)
degrading enzymes. In D caryophyllus, Torenia, Petunia, and Begonia, the antisense technology has been
successfully attempted for ACS (ACC synthase) and ACO (ACC oxidase), key enzymes of ethylene
biosynthesis. In carnation, by silencing ACO, vase life was almost doubled compared to the original
variety, increasing from 5 to 9 days (Chandler 2007; Savin et al. 2019). The ethylene receptor 1 (etr1-1)
appeared to be a key gene for further manipulation. In comparison with nontransformed plants,
petunias expressing constitutively a mutated etr1-1 transgene from Arabidopsis were insensitive to
ethylene and showed larger flowers with a longer vase life (Clevenger et al. 2004). They, however,
showed a negative efect in several morpho-physiological traits. Normal plants with flowers showing
higher longevity could be obtained in kalanchoë, Campanula, and D. caryophyllus using a flowerspecifc
fructose-bisphosphatase 1 (FBP1) promoter driving the expression of the same transgene (Olsen et al.
2015). Other genes, tested in Nemesia strumosa and T. fournieri, were less successful than etr1-1 of
Arabidopsis. In Japanese morning glory (I. nil), functional studies highlighted the role of a NAC
(NAM/ATAF1,2/CUC2) type transcription factor, EPHEMERAL 1 (EPH1), in the control of ethylene-
independent flower senescence (Shibuya et al. 2014).
