Postharvest Biology and Technology 86 (2013) 3744

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Postharvest Biology and Technology
j our nal homepage: www. el sevi er . com/ l ocat e/ post har vbi o
Light modulation of volatile organic compounds from petunia owers
and select fruits
Thomas A. Colquhoun
a,c
, Michael L. Schwieterman
c
, Jessica L. Gilbert
b,d
,
Elizabeth A. Jaworski
a
, Kelly M. Langer
a
, Correy R. Jones
a
, Gabrielle V. Rushing
a
,
Tia M. Hunter
a
, James Olmstead
b,d
, David G. Clark
a,c,d
, Kevin M. Folta
b,c,d,
a
Department of Environmental Horticulture, University of Florida, Gainesville, FL 32611, USA
b
Horticultural Sciences Department, University of Florida, Gainesville, FL 32611, USA
c
Graduate Programfor Plant Molecular and Cellular Biology, Gainesville, FL 32611, USA
d
Graduate Programfor the Plant Molecular Breeding Initiative, Gainesville, FL 32611, USA
a r t i c l e i n f o
Article history:
Received 19 November 2012
Accepted 10 June 2013
Keywords:
Flavor
Flowers
Fruits
Light
Smell
Volatiles
a b s t r a c t
Light intensity, duration, direction, and wavelength are informative to plants. The biochemical circuits
that connect specic light wavelengths to expression of specic genes and the metabolic networks they
govern have been well dened. However, little emphasis has been placed on how discrete wavelengths
of light, alone or in combination, may be applied to manipulate postharvest qualities of high-value horti-
cultural crops. Using narrow-bandwidth LED light we test the hypothesis that discrete light wavelengths
can affect the accumulation of volatile compounds known to affect aroma or taste in select ower and
fruit products. Volatile benzenoid/phenylpropanoid emission from petunia owers could be altered with
light application. Levels of a key oral volatile, 2-phenylethanol, increased with a red and far-red light
treatment. Similar experiments demonstrated that fruit volatile proles of tomato, strawberry, and blue-
berry can be manipulated with specic light treatments. These results suggest that compounds affecting
sensory qualities of owers and fruits can be modied by adjustment of ambient light conditions. These
ndings open new areas of inquiry about how the fragrance and avor of owers and fruits may be
improved with simple changes in postharvest light conditions.
2013 Published by Elsevier B.V.
1. Introduction
Plant growth and development is a product of the genetic
potential of the plant and how it responds to stimuli from the
ambient environment. An element of this interaction is facilitated
by a suite of plant photosensory receptor proteins, each adapted to
sense and relay information about the incident light spectrum. In
the case of horticultural crops, light quantity, quality, and duration
inform the plant of current conditions that ultimately contribute
to plant productivity and product quality.
Light signaling pathways are well understood in the model
system Arabidopsis thaliana. Light signals are transduced through
well-described pathways that inuence many aspects of plant
growth and development (Chen et al., 2004b). These pathways
have been translated to a large number of crop species, where
genetic and photophysiological analyses demonstrate the effects

Corresponding author at: Horticultural Sciences Department, University of

Florida, Gainesville, FL 32611, USA. Tel.: +1 352 273 4812.
E-mail address: kfolta@u.edu (K.M. Folta).
of various wavelengths of light on plant productivity (Barrero
et al., 2012; Frantz et al., 2004; Li and Ma, 2012; Preuss et al., 2012;
Reynolds et al., 2012; Singh et al., 2011). Although yield is often
affected, qualities such as ripening, color and nutraceutical content
are also affected by the light environment. In practice, light is
a passive entity, driving plant processes based on light quantity
and quality from a natural or articial environment. However,
light may also be used to control growth and development by
manipulating the light spectrum itself. A change in the ambient
spectrum can alter plant behavior or potentially affect quality of
plant products (Folta and Childers, 2008).
There is great interest in understanding planthuman interac-
tion with regard to plant produced volatile organic compounds
(Du et al., 2011; Dudareva and Pichersky, 2008; Miyazaki et al.,
2012; Tieman et al., 2012). Specic combinations and concentra-
tions of volatile organic compounds can impart distinct fragrances
and avors to owers and fruits (Klee, 2010; Tieman et al., 2012;
Underwood et al., 2005; Vogel et al., 2010) during gustation (retro-
nasal), and fragrance during inhalation (ortho-nasal), adding value
for product quality andultimately a consumers sensory experience
(Goff and Klee, 2006; Small et al., 2004).
0925-5214/$ see front matter 2013 Published by Elsevier B.V.
http://dx.doi.org/10.1016/j.postharvbio.2013.06.013
38 T.A. Colquhoun et al. / Postharvest Biology and Technology 86 (2013) 3744
The literature shows evidence of environmental factors inu-
encing the production of these volatile molecules in planta
(Oyama-Okubo et al., 2005; Watsonet al., 2002). Numerous aspects
of plant organic compoundmetabolismare inuencedby light con-
ditions, e.g. the cellular redox state, cyclic nucleotide metabolism
in bean, phenylpropanoid production in Arabidopsis (Brown et al.,
1989; Dietz and Pfannschmidt, 2011; Jin et al., 2000). Additionally,
plant volatile production can be inuenced by light quantity over
the course of fruit development instrawberry (Watsonet al., 2002).
Terpenoids have been shown to be modulated by the phytochrome
photosensors (Peer and Langenheim, 1998; Tanaka et al., 1998). It
was demonstratedthat whensweet basil plants were grownoncol-
ored mulches, the volatile compounds emitted from fresh leaves
varied with the color of mulch used (Loughrin and Kasperbauer,
2003).
The central hypothesis of this work is that plant volatile emis-
sion could be reproducibly manipulated by variation in light
quality. To directly test this hypothesis, we exploited the capac-
ity to control discrete spectral quality using a narrow-bandwidth
LED based light platform(Zhang et al., 2011) to expose owers and
fruits to specic wavelengths of light. Five lighting conditions were
employed: white, blue, red, far-red, and dark (Fig. 1).
Harvested petunia owers, tomato, strawberry, and blueberry
fruits were analyzed for the emission of key volatile compounds
subsequent to treatments with different wavelengths of light.
Results show that the emissions of discrete volatiles important to
plant product quality are inuenced by light quality. These results
have created a foundation for future identication of light regimes
that may alter plant product post-harvest quality for consumers.
2. Materials and methods
2.1. Narrow-bandwidth LED light platform
Thelight treatments weregeneratedusinga light emittingdiode
(LED) platform(Zhang et al., 2011). A dark treatment and four light
treatments were tested: white, blue, red, and far-red (Fig. 1). In all
cases, light treatments were 50mol m
2
s
1
in separate illumina-
tion chambers within an environmentally controlled and actively
ventilated area (221.5

C). The control treatment (white light)

was generatedbycool whiteuorescent bulbs, whilethedarktreat-
ments were performed in an identical light-tight enclosure under
the same ambient conditions. The light treatments were gener-
ated using the Flora Lamp LED arrays (Light Emitting Computers,
Victoria, BC). The spectra used in these experiments are shown in
Fig. 1. Spectroradiometer readings of the light qualities used in this study. All
treatments represent the waveformgenerated at a uence rate of 50mol m
2
s
1
.
B =blue, R=red, FR=far-red, HBW=half-bandwidth.
Fig. 1. Fruits and owers were treated without photoperiod. Spec-
troradiometer readings were obtained with a StellarNet device and
visualized on SpectraWiz software (Stellar Net, Tampa, FL).
2.2. Petunia
In all cases of petunia experimentation, Petunia hybrida cv.
Mitchell Diploid (MD) plants (Mitchell et al., 1980) were grown
in a glass greenhouse to reproductive maturity, from seed. Devel-
oping MD owers were tagged at stage 6 and allowed to grow to
stage 8 (Colquhoun et al., 2010). The morning of what would be
a stage 8 open ower; tagged owers were excised at the petiole,
and placed in a 4mL glass vial with 2mL of tap water.
In the initial experiment (Fig. 2), the prepared MD owers were
placed in each of ve light environments: white, blue, red, far-
red, and dark. Six owers were exposed to each light condition
for 8h and removed at 18:00h. The corollas were then removed
from the receptacle and two corollas were each inserted into a
single glass tube for volatile collection, totaling three biological
replicates per experiment. Multiple replications of this experiment
were performed with similar results observed.
To determine the length of time required to obtain a light-
induced volatile response, a time course was conducted. Samples
of eight owers were exposed to a far-red light environment for
0, 2, 4, 6, or 8h. Flowers were kept under white light (control)
conditions until entering the far-red light treatment (Fig. S1). A
dark environment control was also included. At 18:00h, all owers
were removed fromtheir respective light treatments. The recepta-
cle was detached, and two owers were each inserted into a single
glass tube for volatile collection, totaling four biological replicates
per experiment. Multiple replications of this experiment were per-
formed with similar results observed.
2.3. Tomato
Field-grown tomatoes (Solanum lycopersicum, M82) were har-
vested at breaker stage and allowed to ripen under ve different
light conditions: white, blue, red, far-red, and dark. After 10d, mul-
tiple fruits per treatment were diced, pooled, and 100g samples
were loaded into glass tubes in triplicate per experiment (n=3) for
volatile collection (Schmelz et al., 2001; Tieman et al., 2006). Mul-
tiple replications of this experiment were performed with similar
results observed.
2.4. Strawberry
Field-grown mature Fragaria ananassa Strawberry Festival
fruit were harvested in the morning and chilled at 4

C overnight in
dark conditions. Seven berries were selected based on uniformity
of appearance per treatment, and were placed into clear plastic
containers the next morning, followed by the treatment conditions
for 8h. Light environments tested were white, blue, red, far-red,
and dark. After 8h, light-treated fruit samples fromeach treatment
were pooled, homogenized in a blender, and 20g of homogenate
was loaded in triplicate (n=3) into glass tubes for volatile collec-
tion. Multiple replications of this experiment were performed with
similar results observed.
2.5. Blueberry
Field-grown Vaccinium corymbosum Scintilla fruit were har-
vested 1d prior to light treatments. Mature blueberry fruits
were harvested in the morning and chilled at 4

C overnight
in dark conditions. The next morning selected uniform fruit
were spread in a single layer and placed in white, blue, red,
far-red, and dark conditions for a 8h treatment. After the
T.A. Colquhoun et al. / Postharvest Biology and Technology 86 (2013) 3744 39
Fig. 2. Histograms showing the detected emission levels of oral volatile benzenoids/phenylpropanoids (FVBPs) fromPetunia hybrida cv. Mitchell Diploid after 8h light
treatments (50mol m
2
s
1
). The Y-axis is in mol kg
1
s
1
, and the X-axis is general white light (W), blue light (Bl), red (R), far-red F-R), and dark (D) lighting conditions
(meanse; n=3). Identied FVBPs include: phenylacetaldehyde, 2-phenylethanol, benzyl benzoate, methyl benzoate, methyl salicylate, benzyl alcohol, benzaldehyde,
isoeugenol, and eugenol. Lower case letters above the standard error bars indicate signicant differences at P<0.05 by a one-way ANOVA (t-distribution).
treatments, the berries were removed, pooled by treatment,
blended 15s, and 30g of homogenate was loaded in triplicate
(n=3) into glass tubes for volatile collection. Multiple replica-
tions of this experiment were performed with similar results
observed.
2.6. Volatile collection
Samples were loaded as described previously in this section
into thin walled glass tubes (2.5cm i.d., 61cm long, and 300mL
volume) attached to a pushpull dynamic headspace volatile
collection system with a column containing approximately 50mg
HaySep Q 80-100 porous polymer adsorbent (Hayes Separations
Inc., Bandera, TX) at the tube outlet to capture volatile organic
compounds over a period of 1h, all at roomtemperature. Volatiles
were eluted from the column with 150L methylene chloride
spiked with 5L of nonyl acetate as an elution standard. Two
L of elution samples were run on an Agilent 7890A series gas
chromatograph ame ionization detector. Peaks from the chro-
matography traces of compounds within a sample elution were
inferred using Chemstation software (Agilent Technologies, Santa
Clara, CA). Peak areas of detected analytes were used to determine
the mass of each compound within a sample via calculations
that adjusted for a nonyl acetate elution standard and original
biological sample mass (Schmelz et al., 2001, 2004). GCMS was
used to conrmthe presence of compounds and identify retention
times in conjunction with chemical standards (SigmaAldrich, St.
Louis, Missouri) run on the GC (Table S1).
40 T.A. Colquhoun et al. / Postharvest Biology and Technology 86 (2013) 3744
3. Results
3.1. Petunia oral volatile emission after varying spectral light quality treatments
A model plant system examined was Petunia hybrida Mitchell Diploid (MD)
for oral volatile benzenoid/phenylpropanoid (FVBP) emission. MD oral fragrance
productionis awell characterizedbiological process consistingof arelativelysimple,
but concentrated fragrance prole (Colquhoun and Clark, 2011; Colquhoun et al.,
2010; Schuurink et al., 2006; Verdonk et al., 2003). Flowers from MD plants were
excised at stage 8 (Colquhoun et al., 2010), placed in tap water, and moved from
greenhouse conditions to specied lighting conditions. Detected amounts of FVBPs
under light treatments werecomparabletowhat has beencommonlyreportedinthe
literature (Boatright et al., 2004; Colquhoun et al., 2010; Underwood et al., 2005;
Verdonk et al., 2005), suggesting the treatment conditions were acceptable. Dark
treatment of MD owers resulted in the signicant reduction of most of the FVBP
compounds like: methyl benzoate, benzaldehyde, phenylacetaldehyde, isoeugenol,
and eugenol (Fig. 2). No light treatments signicantly affected the emission of MD
oral volatile phenylpropanoids, isoeugenol and eugenol. The blue light treatment
only resulted in a signicant change of benzaldehyde emission, which was slightly
reduced compared to white light treated owers.
The most obvious change in FVBP emission was observed with the red and far-
redlight treatments, whichincreasedphenylacetaldehydeemissionby2.7-fold(red)
and 2.3-fold (far-red), and increased 2-phenylethanol emission by 9.9-fold (red)
and 5.8-fold (far-red). These treatments also resulted in an increased benzyl alcohol
and benzyl benzoate emission. Methyl salicylate emission was signicantly reduced
after the far-red light treatment (Fig. 2).
3.2. Petunia oral volatile emission after treatment with far-red light over a time
course
Because far-red light treatments produced the highest number of signicantly
affected FVBPs (phenylacetaldehyde, 2-phenylethanol, benzyl benzoate, benzyl
Fig. 3. Histograms showing the detected emission levels of oral volatile benzenoids/phenylpropanoids (FVBPs) from Petunia hybrida cv. Mitchell Diploid after an 8h
treatment of white or varied durations of far-red light (50mol m
2
s
1
). The Y-axis is in mol kg
1
s
1
, and the X-axis is general white light for 8h (0), 6h under white light
and 2h under far-red treatment (2), 4h under white light and 4h under far-red treatment (4), 2h under white light and 6h (6) under far-red treatment, and 8h (8) under
far-red treatment (meanse; n=3). Identied FVBPs include: phenylacetaldehyde, 2-phenylethanol, benzyl benzoate, methyl benzoate, methyl salicylate, benzyl alcohol,
benzaldehyde, isoeugenol, and eugenol. Refer to supplemental Fig. 1 for a detailed description of the experimental design. Lower case letters above the standard error bars
indicate signicant differences at P<0.05 by a one-way ANOVA (t-distribution).
T.A. Colquhoun et al. / Postharvest Biology and Technology 86 (2013) 3744 41
alcohol, and methyl salicylate), we chose to further examine the time course of
their accumulation in response to this treatment. Flowers placed under white light
to far-red light treatment for 8h maintained FVBP levels that were similar to those
shown in Fig. 2 (Fig. 3).
Far-red light treatment elevated phenylacetaldehyde emission after 2h, and
4 h, then remained constant for the duration of the experiment, while emission of
2-phenylethanol increased over time with a similar prole as emission of pheny-
lacetaldehyde (Fig. 3). Benzyl alcohol was elevated by 2h of far-red light treatment
and reached a maximum level after 8h of treatment. Benzyl benzoate and benzal-
dehyde were elevated by the far-red light treatment and both reached a maximum
level after 4h of treatment (Fig. 3). Methyl salicylate emission was reduced after 2h
of far-red light treatment, and remained consistent afterward. No signicant change
was detected for methyl benzoate or isoeugenol emission through any time-point
of far-red treatment compared the white light treatment.
The time course of the far-red light affect was tested by monitoring phenylac-
etaldehyde and 2-phenylethanol emission. MD owers were exposed to the far-red
light conditions for 8h, removed fromthe far-red light conditions, and placed into
white light conditions. Volatiles were collected every 4h for a total of ve volatile
collections (Fig. S2). Emission of phenylacetaldehyde and 2-phenylethanol from
MD owers was similar to previous results after the 8h far-red light treatment
(Figs. 2 and 3). MD volatile emission increased after subjective nightfall, and the
far-red light effect diminished compared to the controls (Fig. S2).
3.3. Tomato fruit volatile emission after varying spectral light quality treatments
Fruits from tomato (Solanum lycopersicum) cv. M82 were tested for volatile
emission. Tomato fruit volatile compound production is well characterized, but
unlike petunia fragrance, tomatofruit volatile proles canconsist of a large andcom-
plexnumber of chemical species (Butteryet al., 1987; Tiemanet al., 2012). Therefore,
we focused on a simple set of volatile compounds that have demonstrated impor-
tance intomatoavor. Cis-3-hexen-1-ol, 2-methyl butanal, and3-methyl-1-butanol
have been shown to contribute to overall tomato avor intensity (Tieman et al.,
2012). Tomato fruits were harvested at breaker stage and allowed to ripen under
white light, blue, red, far-red, and dark conditions. Compared to white light treat-
ments, the blue light treatment resulted in no signicant differences in tomato fruit
volatile emission of these four compounds (Fig. 4). In contrast, the dark treatment
resulted in a 5-fold increase of 3-methyl-1-butanol, 2.6-fold increase in 2-methyl
butanal, and a signicant increase of cis-3-hexen-1-ol compared to white light con-
ditions. Redlight treatment resultedinsignicant increases of 2-methyl butanal and
3-methyl-1-butanol, along with a reduction of cis-3-hexenal (Fig. 4). Far-red light
treatments resulted in signicant increases of all four volatile compounds with a
notable 2.2-fold increase of cis-3-hexenal.
3.4. Strawberry fruit volatile emission after varying spectral light quality
treatments
Volatile emissions from strawberry (Fragaria ananassa) fruits contain a large
array of compounds (Du et al., 2011; Maarse, 1991). We focused on a subset of
volatile compounds that likely contribute to strawberry sensory perception and
could be analyzed consistently. These include cis-3-hexen-1-ol, which was present
in large amounts as well as ethyl caproate and methyl butyrate, two compounds
well represented in the strawberry literature (Hakala et al., 2002; Jetti et al., 2007;
Olbricht et al., 2008), along with hexyl butyrate, a ve carbon extension of methyl
butyrate, known to be a volatile with fruity aroma. Mature strawberry fruit was har-
vested in the morning and chilled at 4

C overnight in dark conditions. Compared

to white light treatments strawberry cis-3-hexen-1-ol emission was not signi-
cantly altered in any of the other light treatments (Fig. 5). In contrast, ethyl caproate
emission was dramatically reduced in all light treatments compared to white light.
Methyl butyrate emission was signicantly increased after far-red light treatments,
while hexyl butyrate emission was detected at signicantly reduced amounts under
blue light treatments (Fig. 5).
3.5. Blueberry fruit volatile emission after varying spectral light quality
treatments
Blueberry (V. corymbosum) fruit was tested for volatile emission under various
light conditions. Blueberry fruits produce a relatively large array of volatile com-
pounds much like tomato and strawberry. Therefore, we analyzed a simple set
of volatile compounds with putative importance in blueberry, i.e. hexenal, trans-
2-hexenal, 1-hexanol, trans-2-hexen-1-ol (Parliment and Kolor, 1975; Horvat and
Senter, 1985; Hirvi and Honkanen, 1983). Compared to white light treatments, blue-
berries exposed to far-red light conditions emitted higher levels of hexenal, while
1-hexanol and trans-2-hexen-1-ol emissions were considerably reduced (Fig. 6).
The blue light treatment also resulted in signicant reduction of 1-hexanol and
trans-2-hexen-1-ol emissions compared to white light controls.
4. Discussion
The experiments in this report test the hypothesis that narrow-
bandwidth light treatments can affect the metabolite qualities of
common owers and fruits that produce diverse aromatic prod-
ucts. This hypothesis is based on signicant data that indicate
light-driven gene expression of enzymes inuence steps in these
pathways, likely affecting metabolite levels. The results of these
experiments indicate that in a post-harvest setting, it is possible
to affect the ux of substrates through discrete biochemical nodes
that ultimatelyaffects volatilecompoundproductioninowers and
fruits, simply by applying specic light treatments. It is exciting to
speculate that it may be possible to use light treatments to shape
the avor or aroma qualities of a given genotype of plant, creat-
ing a range of phenotypes from a common genetic background.
More likely, these concepts may be employed to stabilize or induce
desirable avors and aromas of fruits and owers post-harvest.
4.1. Petunia
Petunia hybrida Mitchell Diploid (MD) is an established oral
volatile model systemwith a relatively lowmetabolic background
Fig. 4. Histograms showing the detected emission levels of specic volatile organic compounds from Solanum lycopersicum cv. M82

fruit after 8h light treatments

(50 mol m
2
s
1
). The Y-axis is in mol kg
1
s
1
, and the X-axis is general white light (W), blue (Bl), red (R), far-red (F-R), and dark (D) lighting conditions (meanse;
n =3). Identied tomato fruit volatile compounds include: cis-3-hexenal, 3-methyl-1-butanol, and cis-3-hexen-1-ol, and 2-methyl butanal. Lower case letters above the
standard error bars indicate signicant differences at P<0.05 by a one-way ANOVA (t-distribution).
42 T.A. Colquhoun et al. / Postharvest Biology and Technology 86 (2013) 3744
Fig. 5. Histograms showingthedetectedemissionlevels of specic volatileorganic compounds fromFragaria ananassacv. StrawberryFestival fruit after 8hlight treatments
(50 mol m
2
s
1
). The Y-axis is in mol kg
1
s
1
, and the X-axis is general white light (W), blue (Bl), red (R), far-red (F-R), and dark (D) lighting conditions (meanse; n=3).
Identied strawberry fruit volatile compounds include: cis-3-hexen-1-ol, ethyl caproate, methyl butyrate, and hexyl butyrate. Lower case letters above the standard error
bars indicate signicant differences at P<0.05 by a one-way ANOVA (t-distribution).
Fig. 6. Histograms showing the detected emission levels of specic volatile organic compounds from V. corymbosum cv. Scintilla fruit after 8h light treatments
(50 mol m
2
s
1
). The Y-axis is in mol kg
1
s
1
, and the X-axis is general white light (W), blue (Bl), red (R), far-red (F-R), and dark (D) lighting conditions (meanse;
n =3). Identied strawberry fruit volatile compounds include: hexanal, 1-hexanol, trans-2-hexenal, and trans-2-hexen-1-ol. Lower case letters above the standard error bars
indicate signicant differences at P<0.05 by a one-way ANOVA (t-distribution).
and a comparatively small number of emitted volatiles (Colquhoun
and Clark, 2011; Colquhoun et al., 2010; Schuurink et al., 2006;
Verdonk et al., 2003). The major volatile compounds that are
produced are generated from the petal limb tissue (Underwood
et al., 2005). The genetic regulation and biochemical formation of
MD oral volatile benzenoids/phenylpropanoids have been well
described. Therefore, this system served as a starting point to
understand light-quality-mediated regulation of specic control
points in regard to plant secondary metabolism. MD also has the
advantage of fewother screening pigments in the petals, since the
owers are white. Compared to the other plant products examined
here, the petunia corolla limb tissue is very thin, which allows for
full illumination with the photon energies applied.
Ingeneral, the blue light treatment of MD owers hadlittle to no
effect on volatile emission, but a slight decrease of benzaldehyde
emission relative to white light conditions was observed (Fig. 2).
Benzaldehyde production in MD oral limb cells is putatively cat-
alyzed by a BALDH enzyme, which is localized to the mitochondria
(Long et al., 2009). Currently, benzaldehyde is the only MD volatile
compound thought to originate fromthe mitochondrial subcellular
compartment. Little is known about how blue light signals
affect mitochondrial processes other than their movement (Islam
et al., 2009), and a link between this large-scale movement and
metabolism is difcult to devise. We will now test how blue light
mechanistically affects metabolic processes in this compartment.
In contrast to blue treatments, red and far-red light affected the
emission of phenylacetaldehyde, 2-phenylethanol, benzyl alcohol,
and benzyl benzoate. In these cases both red and far-red had com-
parable effects, suggesting that the response was not phytochrome
reversible. Fig. 2 also shows that methyl salicylate is signicantly
reduced only by far-red light treatment, indicating that methyl sal-
icylate production may be repressed by active phytochrome. In
short, multiple mechanisms may be responsible for the effect that
red and far-red light treatments have on MD oral fragrance.
4.2. Tomato
Tomato fruit volatile proles are muchmore complex compared
to petunia, yet tomato is also a well-established system for char-
acterization of plant volatiles (Buttery et al., 1987; Goff and Klee,
T.A. Colquhoun et al. / Postharvest Biology and Technology 86 (2013) 3744 43
2006; Tieman et al., 2012). There are a number of reports regarding
the role of light in fruit phytonutrient content (Azari et al., 2010).
For example, some of these reports test howlight affects the storage
of juices (Hashizume et al., 2007), or the effect of light treatments
on lycopene accumulation during fruit ripening (Alba et al., 2000).
The analyses herein focused on a subset of volatiles that contribute
signicantly to tomato avor (Tieman et al., 2012). The compounds
cis-3-hexenal (grassy), cis-3-hexen-1-ol (green, leafy), 2-methyl
butanal (cocoa, coffee-like), and 3-methyl-1-butanol (amyl alco-
hol) have important roles in shaping the avor of tomato, with
the latter two providing a perception of sweetness (Tieman et al.,
2012).
Some of the most signicant increases involatile emission, com-
pared to white light treatments, are when fruits ripen in far-red
light or darkness. This result suggests that light is driving a process,
either dependent on photosynthesis, metabolism, or development,
which actually decreases the accumulation of these compounds.
This nding is consistent with reports that showhowgenes known
to participate in light-driven responses affect tomato pigmentation
and nutritional quality. Liu et al. (2004) identied high pigmenta-
tion mutants as HY5 and a COP-like genes. HY5 is known for its
role in phytochrome and cryptochrome signaling. COP1 and COP-
like genes have dened roles in light signaling as well. Although
avors were not assessed in the hp1 and hp2 mutants, it may be
proposed that the increased carotenoids may give rise to changes
in key volatiles (like beta-ionone) that can affect avor (Chen et al.,
2004a; Tieman et al., 2012; Vogel et al., 2010).
Red light treatment of the tomatoes led to a signicant increase
in 2-methyl butanal and 3-methyl-1-butanol, while levels of cis-
3-hexenal were reduced, all compared to white light treatments
(Fig. 4). The experiments performed here illustrate that signicant
change in tomato avor volatiles can be attained by altering the
ripening light environment. These ndings are important because
two volatiles that contribute to sweet avor perception (without
the contribution of sugars) are present at higher levels after red
light treatment than in white light, suggesting that the likability of
tomato may be increased due to a perceived sweeter avor. Such
outcomes present an opportunity to manipulate avors of these
fruits without changing the genetics, adding transgenes, treating
with hormones or affecting plant nutrition. By changing red and
far red mixtures or the daily duration of red and far-red treatments
post-harvest, it may be possible to signicantly remodel the avor
of tomato fruits.
4.3. Strawberry
The various light conditions did not signicantly affect the
accumulation of cis-3-hexen-1-ol in strawberry fruits used here.
However, the fruits exhibited a range of responses to the other
treatments. Methyl butyrate increased signicantly in response to
far-red light, and to a lesser extent to red light and dark treat-
ment (Fig. 5). Ethyl caproate was most abundant under white light
conditions. Exposure to narrow-bandwidth light drove levels of
this volatile down 450-fold, a magnitude of change even greater
than moving the fruits to complete darkness. These data suggest
that the pathways leading to production of ethyl caproate contain
regulatory nodes that require active participation of blue and red
(and possibly far-red) signaling systems. Conversely the indepen-
dent steps may depend on blue, then red light cues (or vice versa)
to produce the enzymes necessary for synthesis of this volatile.
The nal compound assayed, hexyl butyrate, was unaffected in all
light conditions except for blue light. Narrow-bandwidth blue light
decreased hexyl butyrate accumulation approximately 5-fold, sug-
gesting that activation of cryptochromes (or possibly other blue
light sensors) regulates expression, stability, or activity of enzymes
required for its synthesis.
When strawberry is compared to tomato for light control of cis-
3-hexen-1-ol accumulationthe data indicate that light has aneffect
intomato, but not instrawberry fruit tissue. Intomatothe levels are
clearly affected by far-red light (Fig. 4), in a manner suggestive of
phytochrome control. This same pattern is not observed in straw-
berry where the compound is present in similar quantities across
treatments (Fig. 5). In these cases the enzymes that underlie pro-
duction are not regulated in the same way or that substrates are
not comparably available. One simple explanation is that straw-
berry and tomato are remarkably different botanically, and reside
in distant plant families. It may be expected that the response to
the light environment would be different.
4.4. Blueberry
Like strawberry and tomato fruits, blueberry fruits present a
complex prole of avor volatiles. However, unlike strawberry and
tomato, blueberry fruit volatiles have not been as extensively char-
acterized. In response to light, the levels of hexenal emission are
highest after far-red treatment but are not signicantly affected
by other treatments or darkness. This result indicates that rever-
sion of phytochrome to an inactive state may increase the levels
of this compound. Trans-2-hexanal is not signicantly changed
by light treatment, while 1-hexanol and trans-2-hexen-1-ol lev-
els are decreased by blue and far-red treatment. When considered
against petunia, tomato, and strawberry; blueberry exhibited the
least change in response to narrow-bandwidth light environments.
Of course, different compounds were assayed, yet the sets tested
were those most likely to contribute to favorable consumer avor
qualities. The results suggest that blueberry may be the least likely
to be easily altered by application of narrow-bandwidth light.
5. Conclusion
It has been demonstrated for several decades that discrete
qualities of light can generate profound changes in plant gene
transcription, growth, and development. It is therefore not surpris-
ing that the abundance of secondary metabolites would also be
affected by light cues. Here we show that a group of compounds
central to avors and aromas of fruits and owers are affected by
monochromatic light in a post-harvest system. Looking forward,
such technologies could have applications in post-harvest treat-
ments or retail-level storage of fresh fruits and vegetables. It may
be possible to enhance avor in plant produce by designing simple,
safe, andinexpensivelight treatment programs that manipulatethe
quality of compounds that affect consumer liking.
Acknowledgments
This workwas supportedbyfundingfromthe ofce of the Senior
Vice President for Agriculture and Natural Resources and the Insti-
tute for Plant Innovation; all at the University of Florida. Additional
support was provided by the USDA Floral and Nursery Research
Initiative and the American Floral Endowment.
Appendix A. Supplementary data
Supplementary data associated with this article can be
found, in the online version, at http://dx.doi.org/10.1016/j.
postharvbio.2013.06.013.
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