[2013] light modulation of volatile organic compounds from petunia flowers and select fruits

8/12/2019 [2013] Light Modulation of Volatile Organic Compounds From Petunia Flowers and Select Fruits

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Postharvest Biology andTechnology 86 (2013) 3744

Contents lists available at SciVerse ScienceDirect

Postharvest Biology and Technology

j ournal homepage: www.elsevier .com/ locate /postharvbio

Light modulation ofvolatile organic compounds from petunia flowers

and select fruits

Thomas A. Colquhouna,c, Michael L. Schwietermanc, Jessica L. Gilbertb,d,Elizabeth A.Jaworskia, Kelly M. Langera, Correy R.Jonesa, Gabrielle V. Rushinga,TiaM. Huntera,James Olmsteadb,d, David G. Clarka,c,d, Kevin M. Foltab,c,d,

a Department of Environmental Horticulture, University of Florida, Gainesville, FL 32611, USAb Horticultural Sciences Department,University of Florida, Gainesville, FL 32611, USAc Graduate Program for PlantMolecular andCellular Biology, Gainesville, FL 32611, USAd Graduate Program for thePlant Molecular Breeding Initiative, Gainesville, FL 32611,USA

a r t i c l e i n f o

Article history:

Received19November2012

Accepted10 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 specific light wavelengths to expression ofspecific genes and the metabolic networks they

govern have been well defined. However, little emphasis has been placed on how discrete wavelengths

oflight, alone or in combination,may be applied to manipulatepostharvestqualities ofhigh-valuehorti-

cultural crops. Using narrow-bandwidthLED lightwe test the hypothesis that discrete lightwavelengths

can affect the accumulation ofvolatile compounds known to affect aroma or taste in select flower and

fruit products. Volatile benzenoid/phenylpropanoidemission frompetuniaflowers could bealteredwith

light application. Levels ofa key floral volatile, 2-phenylethanol, increased with a red and far-red light

treatment. Similar experimentsdemonstrated that fruit volatile profiles oftomato, strawberry, and blue-

berry canbe manipulatedwith specific light treatments. These results suggest that compounds affecting

sensory qualities offlowers and fruits can be modified by adjustment ofambient light conditions. These

findings open new areas of inquiry about how the fragrance and flavor of flowers and fruits may be

improvedwith 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 photosensoryreceptor proteins, each adapted to

sense and relay information about the incident light spectrum. In

thecase of horticultural crops, light quantity, quality, andduration

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 influence 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: [email protected](K.M. Folta).

of various wavelengths of light on plant productivity (Barrero

et al., 2012; Frantzet al., 2004; Li andMa, 2012;Preusset al., 2012;

Reynolds et al., 2012; Singh et al., 2011). Although yield is often

affected,qualitiessuch as ripening, color andnutraceutical 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 artificial 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 andChilders, 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). Specific combinations and concentra-

tions of volatile organic compounds can impart distinct fragrances

and flavors to flowers and fruits (Klee, 2010; Tieman et al., 2012;

Underwoodet al., 2005;Vogel et al., 2010) during gustation(retro-

nasal), and fragrance during inhalation (ortho-nasal), adding value

forproductqualityandultimatelya consumerssensoryexperience

(Goff and Klee, 2006; Small et al., 2004).

0925-5214/$ seefrontmatter 2013 Publishedby Elsevier B.V.

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38 T.A. Colquhoun et al. / Postharvest Biology and Technology 86 (2013) 3744

The literature shows evidence of environmental factors influ-

encing the production of these volatile molecules in planta

(Oyama-Okuboetal., 2005;Watsonet al., 2002). Numerousaspects

ofplantorganic compoundmetabolismareinfluenced by light con-

ditions, e.g. the cellular redox state, cyclic nucleotide metabolism

in bean, phenylpropanoid production inArabidopsis (Brown et al.,

1989;Dietz andPfannschmidt, 2011; Jin et al., 2000). Additionally,

plant volatile production can be influenced by light quantity over

thecourse of fruitdevelopment in strawberry (Watsonetal., 2002).

Terpenoids have been showntobemodulatedby thephytochrome

photosensors (Peer and Langenheim, 1998; Tanaka et al., 1998). It

wasdemonstratedthatwhensweetbasilplantsweregrownon col-

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

LEDbased light platform (Zhang et al., 2011) to exposeflowersand

fruits to specificwavelengthsof light.Five lighting conditionswere

employed: white, blue, red, far-red, and dark (Fig. 1).

Harvested petunia flowers, 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 influenced by light quality. These results

have created a foundation for future identification of light regimes

that mayalter plant product post-harvest quality for consumers.

2. Materials and methods

2.1. Narrow-bandwidth LED light platform

Thelighttreatmentsweregeneratedusinga lightemittingdiode

(LED) platform (Zhang et al., 2011). A dark treatmentand four light

treatments were tested: white, blue, red, and far-red (Fig. 1). In all

cases, light treatments were 50molm2 s1 in separate illumina-tion chambers within an environmentally controlled and actively

ventilated area (221.5 C). The control treatment (white light)

wasgeneratedbycoolwhitefluorescentbulbs,whilethe darktreat-

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 representthewaveformgeneratedat a fluence rate of 50molm2 s1.

B=blue, R= red, FR= far-red, HBW=half-bandwidth.

Fig. 1. Fruits and flowerswere treated without photoperiod. Spec-

troradiometer readingswere obtainedwith a StellarNet deviceand

visualized on SpectraWiz software (Stellar Net, Tampa, FL).

2.2. Petunia

In all cases of petunia experimentation, Petuniahybrida cv.

Mitchell Diploid (MD) plants (Mitchell et al., 1980) were grown

in a glass greenhouse to reproductive maturity, from seed. Devel-

oping MD flowers 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 flower; tagged flowerswere excised at the petiole,

and placedin a 4mLglass vialwith 2mLof tap water.

In the initial experiment (Fig. 2), thepreparedMDflowers were

placed in each of five light environments: white, blue, red, far-

red, and dark. Six flowers 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

replicatesper experiment.Multiple replications of thisexperiment

were performedwith 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 flowers 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 wasalso included.At 18:00h, allflowers

were removed from their respective light treatments. The recepta-

cle was detached, and two flowerswere each inserted into a single

glass tube for volatile collection, totaling four biological replicates

perexperiment.Multiple replicationsof this experimentwere per-

formedwith similar results observed.

2.3. Tomato

Field-grown tomatoes (Solanum lycopersicum, M82) were har-

vested at breaker stage and allowed to ripen under five different

light conditions:white,blue, red, far-red, anddark. 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 performedwith similar

results observed.

2.4. Strawberry

Field-grown mature Fragariaananassa Strawberry Festival

fruit wereharvested in themorning and chilled at4 C overnight in

dark conditions. Seven berries were selected based on uniformity

of appearance per treatment, and were placed into clear plastic

containers thenext morning, followedby thetreatmentconditions

for 8h. Light environments tested were white, blue, red, far-red,anddark. After 8h, light-treated fruit samples from each 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 replicationsof this experimentwere performedwith

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


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T.A. Colquhoun et al. / Postharvest Biology and Technology 86 (2013) 3744 39

Fig. 2. Histograms showing the detected emission levels of floral volatile benzenoids/phenylpropanoids (FVBPs) from Petuniahybrida cv. Mitchell Diploid after 8h light

treatments (50molm2 s1). The Y-axis is in molkg1 s1 , andthe X-axis is general white light (W), blue light (Bl), red (R), far-red F-R), and dark (D) lighting conditions

(meanse; n=3). Identified FVBPs include: phenylacetaldehyde, 2-phenylethanol, benzyl benzoate, methyl benzoate, methyl salicylate, benzyl alcohol, benzaldehyde,

isoeugenol, and eugenol. Lower case letters above the standarderror bars indicate significant differences at P


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3. Results

3.1. Petunia floral volatile emission after varying spectral light quality treatments

A model plant systemexamined was Petuniahybrida Mitchell Diploid (MD)

for floral volatile benzenoid/phenylpropanoid(FVBP) emission.MD floral fragrance

production isawell characterizedbiologicalprocessconsistingofa relativelysimple,

but concentrated fragrance profile (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 specified lighting conditions. Detected amounts of FVBPsunder lighttreatmentswerecomparable towhathasbeencommonlyreportedin the

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 flowers resulted in the significant reduction of most of the FVBP

compounds like:methyl benzoate, benzaldehyde,phenylacetaldehyde, isoeugenol,

and eugenol (Fig. 2). No light treatments significantly affected the emission of MD

floral volatile phenylpropanoids, isoeugenol and eugenol. The blue light treatment

only resulted in a significant changeof benzaldehyde emission,whichwas slightly

reduced compared to white light treated flowers.

The most obvious changein FVBP emissionwas observedwith thered and far-

red lighttreatments,whichincreasedphenylacetaldehydeemission by2.7-fold(red)

and 2.3-fold (far-red), and increased 2-phenylethanol emission by 9.9-fold (red)

and5.8-fold(far-red). These treatments alsoresulted in an increasedbenzylalcohol

andbenzylbenzoate emission.Methyl salicylate emissionwassignificantlyreduced

after the far-red light treatment(Fig.2).

3.2. Petunia floral volatile emission after treatment with far-red light over a time

course

Because far-red light treatments produced thehighest number of significantly

affected FVBPs (phenylacetaldehyde, 2-phenylethanol, benzyl benzoate, benzyl

Fig. 3. Histograms showing the detected emission levels of floral volatile benzenoids/phenylpropanoids (FVBPs) from Petuniahybrida cv. Mitchell Diploid after an 8h

treatmentof white or varied durationsof far-red light (50molm2 s1). The Y-axis is inmolkg1 s1, and theX-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 (mean se; n=3). Identified 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 significant differences at P


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alcohol, and methyl salicylate), we chose to further examine the time course of

their accumulation in response to this treatment. Flowers placedunderwhite light

to far-red light treatment for8 h maintained FVBP levelsthatwere similar to those

shownin Fig.2 (Fig. 3).

Far-red light treatment elevated phenylacetaldehyde emission after 2h, and

4h, then remained constant for the duration of the experiment, while emission of

2-phenylethanol increased over time with a similar profile as emission of pheny-

lacetaldehyde (Fig.3). Benzyl alcoholwaselevated by 2h of far-red light treatment

and reached a maximum level after 8h of treatment. Benzyl benzoate andbenzal-

dehydewere elevated by thefar-red light treatmentand both reached a maximum

level after4 h of treatment(Fig.3). Methyl salicylate emissionwasreducedafter 2hoffar-redlight treatment, andremained consistent afterward.Nosignificant change

was detected formethyl benzoate or isoeugenol emission through any time-point

offar-red treatment compared thewhite light treatment.

The time course of the far-red light affectwas tested bymonitoring phenylac-

etaldehyde and2-phenylethanol emission. MDflowerswereexposedto the far-red

light conditions for 8h, removed from the far-red light conditions, andplaced into

white light conditions. Volatileswere collected every 4h for a total of five volatile

collections (Fig. S2). Emission of phenylacetaldehyde and 2-phenylethanol from

MD flowers 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 comparedto thecontrols (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

unlikepetuniafragrance,tomatofruitvolatileprofilescan consistof a largeandcom-

plexnumberofchemicalspecies(Butteryet al.,1987;Tiemanetal., 2012). Therefore,

we focused on a simple set of volatile compounds that have demonstrated impor-

tancein tomatoflavor.Cis-3-hexen-1-ol,2-methyl butanal,and 3-methyl-1-butanol

have been shown to contribute to overall tomato flavor intensity (Tieman et al.,

2012). Tomato fruits were harvested at breaker stage and allowed to ripen under

white light, blue, red, far-red, anddark conditions. Compared to white light treat-

ments, thebluelight treatment resulted in no significant 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 significant increaseof cis-3-hexen-1-ol comparedto white lightcon-

ditions.Red lighttreatmentresulted in significant increasesof 2-methylbutanal and

3-methyl-1-butanol, along with a reduction of cis-3-hexenal (Fig. 4). Far-red light

treatments resulted in significant 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 (Fragariaananassa) 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 analyzedconsistently. These include cis-3-hexen-1-ol,whichwas 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 five carbon extension of methyl

butyrate, known tobea volatilewith fruity aroma.Maturestrawberry fruitwashar-

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 signifi-

cantlyalteredin any ofthe otherlighttreatments(Fig. 5). In contrast,ethyl caproate

emissionwasdramatically reduced in all light treatments comparedto white light.

Methyl butyrateemissionwas significantly increasedafter far-red lighttreatments,

whilehexylbutyrateemissionwasdetectedat significantly reducedamountsunder

blue light treatments (Fig. 5).

3.5. Blueberry fruit volatile emission after varying spectral light qualitytreatments

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 andHonkanen,1983). Comparedtowhitelighttreatments,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 significant reduction of 1-hexanol and

trans-2-hexen-1-ol emissionscomparedto white light controls.

4. Discussion

Theexperiments in this report test thehypothesis that narrow-bandwidth light treatments can affect the metabolite qualities of

common flowers and fruits that produce diverse aromatic prod-

ucts. This hypothesis is based on significant data that indicate

light-driven gene expression of enzymes influence 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 theflux of substrates through discrete biochemical nodes

thatultimatelyaffectsvolatilecompoundproduction inflowersand

fruits, simplyby applying specific light treatments. It is exciting to

speculate that it may be possible to use light treatments to shape

the flavor or aroma qualities of a given genotype of plant, creat-

ing a range of phenotypes from a common genetic background.

More likely, these conceptsmay beemployed to stabilizeor induce

desirable flavors and aromasof fruits andflowers post-harvest.

4.1. Petunia

Petuniahybrida MitchellDiploid (MD) is an established floral

volatilemodel systemwith a relatively lowmetabolic background

Fig. 4. Histograms showing the detected emission levels of specific volatile organic compounds from Solanum lycopersicum cv. M82 fruit after 8h light treatments

(50molm2 s1). The Y-axis is in molkg1 s1, 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). Identified 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

standarderror bars indicate significant differences at P


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Fig.5. Histogramsshowingthedetectedemissionlevelsof specificvolatileorganiccompoundsfrom Fragariaananassacv.StrawberryFestivalfruitafter8 h light treatments

(50molm2 s1). The Y-axis is inmolkg1 s1 , and theX-axis is general white light (W), blue (Bl), red (R), far-red (F-R), and dark (D)lighting conditions (mean se; n=3).

Identified 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 significant differences at P


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2006;Tiemanet al., 2012). There area numberof reports regarding

the role of light in fruit phytonutrient content (Azari et al., 2010).

Forexample, someofthesereports testhowlightaffectsthestorage

of juices (Hashizume et al., 2007), or the effect of light treatments

on lycopene accumulationduring fruit ripening (Alba et al., 2000).

Theanalyses herein focused on a subsetof volatiles that contribute

significantlyto tomatoflavor (Tiemanet 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 flavor of tomato, with

the latter two providing a perception of sweetness (Tieman et al.,

2012).

Someofthemostsignificant increasesin volatileemission,com-

pared to white light treatments, are when fruits ripen in far-red

light ordarkness.This resultsuggests that light is driving a process,

either dependentonphotosynthesis,metabolism, or development,

which actually decreases the accumulation of these compounds.

This findingis consistentwith reports that show howgenes known

toparticipate in light-driven responsesaffect tomatopigmentation

and nutritional quality. Liu et al. (2004) identified 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 defined roles in light signaling as well. Although

flavors were not assessed in the hp1 and hp2 mutants, it may be

proposed that the increased carotenoids may give rise to changes

in keyvolatiles (like beta-ionone) that canaffectflavor (Chen et al.,

2004a; Tieman et al., 2012; Vogel et al., 2010).

Red light treatmentof the tomatoes led to a significant 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 performedhere illustrate that significant

change in tomato flavor volatiles can be attained by altering the

ripening light environment. These findings are important because

two volatiles that contribute to sweet flavor perception (without

the contribution of sugars) are present at higher levels after red

light treatment than inwhite light, suggesting that the likability of

tomato may be increased due to a perceived sweeter flavor. Such

outcomes present an opportunity to manipulate flavors of thesefruits without changing the genetics, adding transgenes, treating

with hormones or affecting plant nutrition. By changing red and

farredmixtures or thedailyduration of redandfar-red treatments

post-harvest, it may be possible to significantly remodel theflavor

of tomato fruits.

4.3. Strawberry

The various light conditions did not significantly 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 significantly in response to

far-red light, and to a lesser extent to red light and dark treat-

ment (Fig. 5). Ethyl caproatewasmost abundant underwhite lightconditions. 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 final compound assayed, hexyl butyrate, was unaffected in all

lightconditionsexcept for bluelight.Narrow-bandwidth bluelight

decreasedhexylbutyrateaccumulation approximately 5-fold, sug-

gesting that activation of cryptochromes (or possibly other blue

lightsensors) regulatesexpression,stability,or activityof enzymes

required for its synthesis.

When strawberry is compared to tomatofor light control of cis-

3-hexen-1-ol accumulation thedataindicatethatlight hasaneffect

in tomato, butnot instrawberry fruit tissue. Intomatothe levelsare

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 profileofflavor volatiles. However, unlikestrawberry and

tomato, blueberryfruit volatileshave notbeen as extensively char-

acterized. In response to light, the levels of hexenal emission are

highest after far-red treatment but are not significantly 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 significantly 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

leastchangein response tonarrow-bandwidth lightenvironments.

Of course, different compounds were assayed, yet the sets tested

were those most likely to contribute to favorable consumer flavor

qualities. The results suggest that blueberrymay 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, anddevelopment. It is thereforenot 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 flavors and aromas of fruits and flowers 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

bepossible to enhance flavor in plant produce bydesigningsimple,

safe,and inexpensivelight treatmentprograms thatmanipulatethe

quality of compounds that affect consumer liking.

Acknowledgments

Thisworkwassupportedby funding fromtheofficeof theSenior

Vice President forAgriculture andNatural Resources and the Insti-

tute forPlant Innovation; allat theUniversity of Florida. Additionalsupport was provided by the USDA Floral and Nursery Research

Initiative and theAmerican 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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[2013] light modulation of volatile organic compounds from petunia flowers and select fruits

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