Postharvest Biology and Technology 148 (2019) 192–199

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Postharvest Biology and Technology

journal homepage: www.elsevier.com/locate/postharvbio

Postharvest light-emitting diode irradiation of sweet cherries (Prunus avium T

L.) promotes accumulation of anthocyanins
⁎
Doris Kokalj , Emil Zlatić, Blaž Cigić, Rajko Vidrih
Department of Food Science and Technology, Biotechnical Faculty, University of Ljubljana, Jamnikarjeva 101, SI-1000, Ljubljana, Slovenia

A R T I C LE I N FO A B S T R A C T

Keywords: Light influences postharvest formation of bioactive compounds in fruit and vegetables. The objectives of this
Sweet cherry (Prunus avium L.) study were to determine the physicochemical changes and phenylalanine ammonia lyase and flavonoid 3′-hy-
Light emitting diode droxylase activities in sweet cherries (Prunus avium L.) irradiated with light-emitting diodes. Cherries were
UV–B stored under light-emitting diodes for 10 days, exposed to UV-B and blue light, and to the combination of white,
Blue light
blue and green light. Irradiation with blue light significantly increased the anthocyanin content (cyanidin 3-O-
Anthocyanins
Phenylalanine ammonia lyase
glucoside, cyanidin 3-O-rutinoside) and significantly influenced the CIE color parameters hue, C* and ΔE.
Combined white-blue-green light provoked similar but less pronounced effects, while UV-B light was similar to
control (in the dark). Blue and white-blue-green light increased phenylalanine ammonia lyase activity. Light
irradiation had no significant effects on ascorbic acid and the phenolic profile. Highly significant correlations
were found between anthocyanins and phenylalanine ammonia lyase on the one side, and the color parameters
hue, C* and ΔE on the other.

1. Introduction
Considerable efforts are being made to increase the availability of
bioactive compounds with beneficial effects on human health. Sweet
cherries (Prunus avium L.) are a rich source of these compounds, and
especially of phenolics, which have important roles in preventing sev-
eral diseases (Boyer and Liu, 2004; McCune et al., 2010; Yao et al.,
2004). Additionally, cherries are considered a highly nutritious fruit
with significant levels of important nutrients and bioactive compounds,
which include glucose, fructose, vitamin C, anthocyanins, quercetin,
flavan-3-ols, flavonols, and hydroxycinnamates (Chockchaisawasdee
et al., 2016; Martini et al., 2017). Anthocyanins have health promoting
effects, which include anti-cancer activities and the prevention of car-
diovascular diseases (He and Giusti, 2010; Wang and Stoner, 2008).
However, the levels of these compounds in cherries can vary depending
on a range of preharvest and postharvest factors, such as cultivar, de-
gree of ripeness, growing conditions, and preharvest and postharvest
treatments (Gonçalves et al., 2007; Serrano et al., 2005).
The skin color of cherry fruit is one of the main attributes that in-
fluence consumer acceptance (Crisosto et al., 2003), and it is the most
important indicator of quality and maturity of fresh cherries. The skin
color is mainly influenced by the content and distribution of antho-
cyanins in the skin (Esti et al., 2002), the levels and types of colorless
phenolics, and external factors such as light and temperature (Gao and
Mazza, 1995). Five anthocyanins have been described for sweet cher-
ries: cyanidin 3-O-glucoside (Cy-3-glu) and cyanidin 3-O-rutinoside
(Cy-3-r) as the major anthocyanins, and peonidin 3-O-glucoside, pe-
largonidin 3-O-rutinoside, and peonidin 3-O-rutinoside as the minor
anthocyanins. However, the composition of the individual anthocya-
nins depends greatly on the cherry cultivar (Chockchaisawasdee et al.,
2016). In addition to anthocyanins, other phenolic compounds in
cherries include hydroxycinnamic acids, flavan-3-ols, and flavonols,
which have been reported to be associated with color and sensory
qualities (Chockchaisawasdee et al., 2016; Martini et al., 2017).
Phenylalanine ammonia lyase (PAL) is an entry-point enzyme in the
phenylpropanoid pathway. PAL catalyzes non-oxidative deamination of
L-phenylalanine to cinnamic acid, which is subsequently converted to
naringenin, the first of the flavonoid molecules in the flavonoid
pathway, and further to flavones, dihydroflavonols, and anthocyani-
dins. PAL induction has also been linked to stress responses (Flores

Abbreviations: ΔE, total color difference; 3pCoQA, 3-p-coumaroylquinic acid; 4-CQA, cryptochlorogenic acid; C*, chroma; CAT, catechin; CHA, chlorogenic acid;
CIE, Commission Internationale de l'Eclairage; Cy-3-glu, cyanidin 3-O-glucoside; Cy-3-r, cyanidin 3-O-rutinoside; ECAT, epicatechin; F3´H, flavonoid 3′-hydroxylase;
Hue, hue angle; ICHA, isochlorogenic acid; K3R, kaempferol-3-O-rutinoside; LED, light emitting diode; NCHA, neochlorogenic acid; PAL, phenylalanine ammonia-
lyase; procy-B2, procyanidin B2; procy-tri, procyanidin trimer; Q3R, quercetin-3-O-rutinoside; QGLUR, quercetin-3-(2G-glucosyl-rutinoside); WBG, white-blue-green
⁎
Corresponding author.
E-mail address: doris.kokalj@bf.uni-lj.si (D. Kokalj).

https://doi.org/10.1016/j.postharvbio.2018.11.011
Received 14 July 2018; Received in revised form 31 October 2018; Accepted 11 November 2018
0925-5214/ © 2018 Elsevier B.V. All rights reserved.
D. Kokalj et al.
et al., 2014; Irisarri et al., 2016). Flavonoid 3′-hydroxylase (F3´H) is a
precursor for cyanidin-based and delphinidin-based anthocyanins, and
is also involved in the biosynthetic pathways of flavonols and flavan-3-
ols (Benmeziane et al., 2016; Jeong et al., 2006).
Light quality and quantity are one of the most important factors that
influence biosynthesis of secondary metabolites, with effects on an-
thocyanin synthesis and antioxidant potential reported for different
fruit and vegetables (Arakawa et al., 2016; Bakhshi and Arakawa, 2006;
Fan et al., 2013). Several studies have reported that using light emitting
diodes (LEDs) as a light source can have valuable effects on vegetable
growth, quality, and/or biochemical responses, such as increased phe-
nolic compounds and improved antioxidant activity (Brazaitytė et al.,
2009; Shin et al., 2012; Wu et al., 2007).
There are scientific evidences that the shelf-life of fruit is correlated
with their content of bioactive compounds. Therefore, increases in
natural antioxidant levels in fruit might represent a valuable strategy to
maintain fruit quality and shelf-life. As well as preharvest application of
LED as a light source, postharvest application has shown significant
effects on fruit antioxidants and might represent a promising tool to
increase fruit shelf-life (Scattino et al., 2014; Sgherri et al., 2015). Ac-
cumulation of many secondary metabolites, including phenols, flavo-
noids, carotenoids, and anthocyanins, depends on the spectral compo-
sition of the light the plants receive (Ouzounis et al., 2015). UV-B
radiation stimulates production of reactive oxygen species, which act as
signaling molecules that trigger plant responses, such as enhanced
production of health-promoting compounds (Jenkins, 2009). Several
studies have shown the potential use of UV-B light irradiation as a
postharvest treatment to promote accumulation of total phenolics,
which results in higher antioxidant capacity during storage (Castagna
et al., 2013; Darré et al., 2017; Formica-Oliveira et al., 2017; Kokalj
et al., 2017; Liu et al., 2011). According to Alothman et al. (2009a;
2009b), UV irradiation of fresh-cut fruit leads to increased antioxidants,
polyphenols, and flavonoids. Furthermore, Hasperué et al. (2016) in-
vestigated the effects of white-blue LED light irradiation on the shelf-
life of broccoli heads, where the treated samples had twice the chlor-
ophyll content and higher levels of carotenoids than the dark-stored
controls. Blue LED irradiation can induce fruit ripening in postharvest
peaches (Gong et al., 2015), while tomatoes treated with blue light
delay softening and ripening (Dhakal and Baek, 2014). Additionally,
Kadomura-Ishikawa et al. (2013) reported that blue light (465 nm) in-
duced anthocyanin accumulation in strawberry fruit. Similarly, sup-
plemental blue light (470 nm) increased the antioxidant capacity, total
phenolic content, ascorbic acid content, and titratable acidity (Xu et al.,
2014b). Shi et al. (2014) reported that blue light (470 nm) enhanced
the content of total anthocyanins and increased the expression of an-
thocyanin biosynthetic and regulatory genes in Chinese bayberry fruit.
There is a certain relationship between light irradiation and fresh-
ness of exposed vegetables and fruit. Implementation of LEDs in storage
chambers, refrigerators, or coolers might enhance the postharvest life
or lengthen the ripening process. Therefore, some existing refrigerators
are already provided with such illumination. Pérez-Ambrocio et al.
(2018) showed that irradiation with blue and UV-C light can increase
the bioactive compounds in habanero pepper over their 30-day storage
under refrigeration.
The objectives of the present study on sweet cherries were to: (1)
investigate the effects of postharvest blue, UV-B, and combined white,
blue, and green (WBG) light irradiation on the development of skin
color; (2) determine the changes in the contents of anthocyanins, as-
corbic acid, and phenolics during LED irradiation; and (3) determine
any changes in PAL and F3´H activities during LED irradiation.
2. Materials and methods
2.1. Chemicals
Ethanol, acetic acid, formic acid, ascorbic acid, ammonium formate,
193
Postharvest Biology and Technology 148 (2019) 192–199
metaphosphoric acid, 2-mercaptoethanol, polyvinylpyrrolidone,
chlorogenic acid, p-coumaric acid, procyanidin B2, quercetin, kaemp-
ferol, FeSO4·7H2O, α-ketoglutaric acid, naringenin, cinnamic acid, and
eriodictyol were from Sigma-Aldrich (Steinheim, Germany). Methanol
and acetonitrile were from Honeywell Riedel-de Haën (Seelze,
Germany). TRIS, ethyl acetate and L-phenylalanine were from Merck
(Darmstadt, Germany). Anthocyanin standards were from
Extrasynthèse S.A. (Genay, France).
2.2. Fruit material and sampling process
‘Van’ sweet cherry (Prunus avium L.) fruit were harvested at com-
mercial maturity stage in Brdo pri Lukovici, Slovenia (46°16′88″ N,
14°68′29″ E; 380 m above sea level) during the 2016 growing season.
The fruit were sorted for fruit size and color, and the most re-
presentative lots were used for these studies. Three hundred and eighty-
four cherries were randomly divided into four treatment units. All of
the cherries were stored in a storage chamber at constant temperature
(1 ± 0.5 °C) under the different light conditions. Samplings were
conducted on days 2, 4, 7, and 10. On each sampling day twenty-four
cherries were randomly removed from each treatment and divided into
four samples, each composed of six cherries. The pitted cherries were
frozen in liquid nitrogen, stored at −80 °C, and ground to a powder
prior to the analyses.
2.3. Light treatments
The light treatments were conducted in storage chambers with
constant temperature (0.5 °C) and 80% to 85% relative humidity. The
cherries were placed on a perforated tray (stems down) and con-
tinuously irradiated with UV-B light, blue light, and WBG light. The UV-
B light LEDs with hemispherical lenses (UVLUX305-HL-3) at
310 ± 5 nm were from Roithner Lasertechnik (Vienna, Austria). These
were installed in a box (0.50 × 0.35 × 0.40 m; L × W × H) and a total
radiant flux 0.046 W m−2 was provided, as emission comparable to the
average solar radiation at the Earth surface at 310 nm. For blue light
treatment, eight LEDs with lambertian high dome lenses (H2A1-H450,
Roithner Lasertechnik) at 444 ± 5 nm were installed in a box
(0.28 × 0.20 × 0.26 m; L × W × H). Total radiant flux at the peak of
emission was adjusted to 1 W m-2, which is comparable to the average
values of solar irradiation at the Earth surface at ∼450 nm. The total
radiant flux at the fruit surface was 23 W m-2. The WBG light LEDs
contained three LED emitters (Kodenshi Corp., Japan): white light
(LW801-1ABPW); blue light (LZ801-006D) at 465 nm to 478 nm, and
peak emission at 470 nm; and green light (LY801-003D) at 518 nm to
532 nm, and peak emission at 520 nm. Six LEDs were installed in a box
(0.32 × 0.24 × 0.17 m; L × W × H) and provided a total radiant flux
of 3.6 W m-2 at the fruit surface, while the individual emitters of blue
and green light provided total radiant flux of 0.05 W m-2 and 0.02 W m-
2
, respectively.
2.4. Color measurements
To investigate the development of the cherry skin color in response
to irradiation, the skin color was measured with a colorimeter (CR-400;
Minolta, Kyoto, Japan) and the CIE parameters (L*, a*, b*) were de-
termined on the same eight fruits per treatment, throughout the 10-day
storage. Each fruit was measured three times. The hue angle (°) was
obtained as arctg(b*/a*) and the Chroma (C*) was calculated as (a*2+
b*2)1/2. The total color difference (ΔE) was obtained as (Δa*2 + Δb*2 +
ΔL*2)1/2 and classified as very distinct when ΔE > 3, distinct when
1.5 < ΔE < 3, and small when ΔE < 1.5.
2.5. Extraction and HPLC analysis of phenolic compounds
One gram of grounded frozen sample was extracted with 3 mL 70%
D. Kokalj et al.
ethanol containing 1% formic acid. These samples were then cen-
trifuged (16,100× g; 3 min) and filtered through 0.20-μm PTFE syringe
filters (Macherey-Nagel, Düren, Germany).
The phenolic compounds were analyzed on an HPLC system
(Agilent 1260 Infinity; Agilent Technologies, Palo Alto, USA) with a
diode array detector, and the wavelength set at 280, 320, 370, and
520 nm. Separation of the phenolic compounds was carried out on a
C18 column (100 × 2.1 mm i.d.; 2.7 μm Poroshell 120 EC; Agilent
Technologies, USA), at a flow rate of 0.2 mL min−1. The mobile phase
consisted of 1% (v/v) aqueous formic acid (eluant A) and methanol
(eluant B), using the following elution gradient: 0–20 min, 2%-23% B;
20–40 min, 23%-35% B; 40–46 min, 35%-38% B; 46–60 min, 60% B;
60–65 min, 95% B. The column temperature was set at 30 °C and the
injection volume of the samples was 1 μL. The contents of the phenolic
compounds were calculated using the external standard calibration
method. For the compounds without standards available, quantification
was estimated using similar compounds: neochlorogenic acid, crypto-
chlorogenic acid, and isochlorogenic acid were calculated as equiva-
lents of chlorogenic acid; 3-p-coumaroylquinic acid as equivalents of p-
coumaric acid; procyanidin trimer as equivalents of procyanidin B2;
quercetin-3-(2G-glucosyl-rutinoside) and quercetin-3-O-rutinoside as
equivalents of quercetin; and kaempferol-3-O-rutinoside as equivalents
of kaempferol. The contents are expressed in mg kg−1 fresh weight.
A triple quadrupole mass spectrometer (Agilent 6460; Agilent
Technologies, Palo Alto, USA) coupled to an electrospray ionization
interface (Agilent Jet Stream) were used for mass analysis of the phe-
nolic compounds. The mass spectra were recorded in positive and ne-
gative modes, from m/z 100 to 1000. The electrospray ionization source
parameters were as follows: gas temperature, 300 °C; drying gas flow,
6 L min−1; capillary voltage, 4000 V; sheath gas temperature, 350 °C;
sheath gas flow, 12 L min−1; and nebulizer pressure, 35 psi. The poly-
phenolic compounds were identified by comparing their retention
times, UV spectra, MS and MS/MS data with those of the reference
standards.
2.6. Enzyme preparations
Frozen cherries were ground in liquid nitrogen, and then the crude
enzymes were extracted in suitable buffers: for PAL (EC 4.3.1.5),
250 mM TRIS buffer (pH 8.5); for F3´H (EC 1.14.13.21), 100 mM TRIS
(pH 7.25). The experimental scheme with detailed procedures is given
in Fig. 1.
2.7. Enzyme assays
Once the enzyme extracts were obtained, the PAL and F3´H activ-
ities were measured as previously described by Halbwirth et al. (2009),
with modifications. The following chemicals were used in these assays:
5 mM FeSO4·7H2O (in milliQ water); 100 mM α-ketoglutaric acid (in
milliQ water); 33 mM naringenin (in ethanol); and 20 mM L-phenyla-
lanine (in 250 mM TRIS buffer, pH 8.5). For the PAL assay, L-phenyla-
lanine was used as substrate, while naringenin was the substrate in the
F3´H assay. Evaluation of the enzyme activities was based on the de-
termined content of cinnamic acid produced in the PAL assay, and
eriodictyol in the F3´H assay. The contents of cinnamic acid and erio-
dictyol were analyzed on an HPLC system according to the method of
analysis of phenolic compounds as described in Section 2.5. The data
are calculated as factors in comparison with the content of the product
in the control assay (day 0), which represented factor 1.
2.8. Ascorbic acid content
Cherry extracts were obtained from 1 g frozen sample which was
extracted with 5 mL 2% metaphosphoric acid at room temperature for
3 min. The samples were then centrifuged (16,100× g; 3 min) and fil-
tered through 0.20-μm PTFE syringe filters (Macherey-Nagel, Düren,
194
Postharvest Biology and Technology 148 (2019) 192–199
Germany).
Ascorbic acid was determined on an HPLC system (1260 Infinity;
Agilent Technologies, Santa Clara, CA, USA) equipped with a diode
array detector set at 254 nm. The separations were on a C-18 column
(100 mm × 2 mm; i.d. 3 μm; Scherzo SM-C18; Imtakt, Kyoto, Japan),
with flow rate of 0.3 mL min−1. The mobile phase was aqueous 5 mM
ammonium formate and 0.05% (v/v) formic acid (eluant C), and 0.3%
formic acid in acetonitrile (eluant D). The linear gradient operated
under the following conditions: 0–3 min: 0%-10% D; 3–4 min: 10%-
100% D, and 4–6 min: 100% D. Chromatograms were recorded at 40 °C,
and the temperature of the automatic sample feeder was set to 4 °C. The
content of ascorbic acid was calculated from the calibration curve
plotted for the standard solutions, and is expressed in mg kg-1 fresh
weight.
2.9. Statistical analysis
Statistical analyses were performed using the R commander soft-
ware (version 3.3.3). To ensure appropriateness of ANOVA, the var-
iances between treatments were determined using Levene’s tests
(α > 0.05). Further ANOVA was performed with post-hoc Tukey’s tests.
*, P < 0.05; **, P < 0.01; ***, P < 0.001.
3. Results and discussion
Sweet cherry quality is determined by attributes that are important
for marketability, among which visual appearance is one of the most
important. Therefore, the influence of different light treatments on the
development of cherry skin color was evaluated. During the storage, the
CIE parameters, and consequently hue and C*, decreased under the
control and all of the treatments (Table 1). L* and b* of all of the
samples were lower after 2 days of storage. For the fruit irradiated with
blue and WBG light, a* was also lower after 2 days, while for the control
group and the fruit treated with UV-B, a* started to decrease after 7
days of treatment. Severe changes in visual external fruit skin appeared
after 7 days of storage, with ΔE of all of the samples > 4. The highest
impact on color change was observed for the blue light. In general, UV-
B light irradiation produced no particular changes in the color para-
meters of the cherry skins, as compared to the control group.
After 10 days of storage, there were no significant differences
(P < 0.001) for hue, C*, and ΔE between the control group and the UV-
B light irradiated samples, as well as between the blue and WBG light
irradiated samples (Table 1). A decrease in hue is attributed to con-
version of the color from a slightly orange shade to a more pronounced
red, while a decrease in C* indicates decreased tonality of the fruit
color. The C* of the control cherries and cherries irradiated with UV-B,
blue and WBG light decreased by 3.80, 3.53, 8.54, and 8.14, respec-
tively. Treatment with the blue light resulted in the highest ΔE (9.11),
followed by the WBG light (6.95), while ΔE of the control and the UV-B
light irradiated samples was comparable. We found lower values of L*,
a*, b*, hue and C* for blue and WBG treated fruit than control and UV-B
treated fruit. Similar data were reported by Gonçalves et al. (2007),
who showed that the chromatic functions of C* and hue correlate with
the evolution of color during the storage of sweet cherries.
Two major anthocyanins found in sweet cherries were evaluated:
Cy-3-r as the main anthocyanin in this ‘Van’ cherry cultivar; and Cy-3-
glu, which represented ∼3% of the total anthocyanin content (Table 2).
Pelargonidin 3-O-rutinoside is a minor anthocyanin in sweet cherries;
however, its content varied, and in several samples it was below the
limit of detection (data not shown). Overall, the contents of both, Cy-3-r
and Cy-3-glu increased during storage under all of the treatments. This
accumulation of anthocyanins in sweet cherries during storage is in
agreement with data from Gonçalves et al. (2004). In the present study,
both Cy-3-r and Cy-3-glu increased in the control fruit during storage;
Cy-3-r from an initial 951.8 mg kg−1 to 1281.9 mg kg−1 at the end of
the treatments, and Cy-3-glu from 30.7 mg kg−1 to 37.6 mg kg−1. At
D. Kokalj et al. Postharvest Biology and Technology 148 (2019) 192–199

Fig. 1. Detailed procedure for the enzyme preparation and assays.

Table 1
Color measures for the ‘Van’ sweet cherries during storage under the control (in the dark) and irradiation treatments.a.
Day Storage CIE parameters hue C* ΔE

ΔL*
Δa *
Δb*

0 Control 30.26 ± 2.35 50.49 ± 2.21

UV-B 30.15 ± 1.53 50.02 ± 2.85
Blue 29.29 ± 2.25 49.85 ± 2.91
WBG 29.21 ± 2.77 49.55 ± 2.53
2 Control −0.66 ± 0.58 0.20 ± 1.36 bc* −0.98 ± 1.91 a* 29.17 ± 2.36 ab* 50.17 ± 2.35 ab* 2.31 ± 1.62
UV-B −0.73 ± 1.78 0.81 ± 0.69 c* 0.53 ± 0.66 b* 30.21 ± 1.33 a* 50.98 ± 3.19 a* 2.41 ± 0.85
Blue −0.63 ± 0.93 −0.26 ± 0.55 ab* −0.76 ± 0.88 a* 28.70 ± 2.05 ab* 49.25 ± 2.46 ab* 2.11 ± 0.74
WBG −0.40 ± 1.86 −1.07 ± 1.45 a* −1.39 ± 1.53 a* 28.30 ± 3.11 b* 47.95 ± 3.89 b* 3.14 ± 1.25
4 Control −0.67 ± 0.91 0.20 ± 1.82 bc* −0.55 ± 1.82 b* 29.55 ± 2.45 a* 50.39 ± 3.57 a* 2.75 ± 1.11 ab*
UV-B −0.89 ± 1.40 0.91 ± 1.37 c* 0.01 ± 1.04 b* 29.68 ± 1.23 a* 50.81 ± 1.93 a* 2.41 ± 1.29 b*
Blue −0.42 ± 0.63 −0.99 ± 0.74 a* −2.08 ± 1.21 a* 27.74 ± 2.08 b* 47.99 ± 2.48 b* 2.69 ± 1.20 ab*
WBG −1.02 ± 2.37 −0.65 ± 1.24 ab* −1.82 ± 1.37 a* 27.62 ± 2.86 b* 48.11 ± 3.28 b* 3.64 ± 0.91 a*
7 Control −0.60 ± 0.95 b* −2.03 ± 1.80 −3.03 ± 1.21 b* 28.27 ± 2.21 a*** 47.23 ± 3.37 4.11 ± 1.65 b*
UV-B −1.94 ± 1.87 a* −1.42 ± 1.14 −2.49 ± 0.92 b* 28.39 ± 1.51 a*** 47.57 ± 3.63 4.24 ± 0.94 b*
Blue −2.89 ± 0.87 a* −2.74 ± 1.63 −4.91 ± 1.57 a* 25.52 ± 2.30 b*** 45.15 ± 3.55 6.57 ± 1.76 a*
WBG −2.09 ± 1.45 a* −2.75 ± 2.36 −4.64 ± 1.39 a* 25.61 ± 2.74 b*** 44.95 ± 4.40 6.32 ± 1.90 a*
10 Control −2.87 ± 0.87 b* −2.16 ± 2.59 b* −3.93 ± 1.63 b* 27.36 ± 2.09 a*** 46.69 ± 4.13 a*** 5.90 ± 2.00 b***
UV-B −3.19 ± 3.17 ab* −1.86 ± 1.70 b* −3.97 ± 1.89 b* 27.09 ± 1.72 a*** 46.49 ± 1.80 a*** 5.99 ± 3.20 b***
Blue −4.77 ± 1.99 a* −5.54 ± 2.95 a* −8.05 ± 1.91 a* 23.08 ± 3.01 b*** 41.31 ± 5.22 b*** 11.22 ± 2.98 a***
WBG −2.70 ± 1.84 b* −5.34 ± 2.69 a* −7.44 ± 1.22 a* 23.76 ± 2.21 b*** 41.41 ± 3.66 b*** 10.09 ± 1.33 a***

a
hue, hue angle; C*, chroma; ΔE, total color difference; WBG, white, blue and green light.
Data are means ± SD (n = 24). Means with different letters within each row are significantly different.
*
P < 0.05.
**
P < 0.01.
***
P < 0.001.

195
D. Kokalj et al. Postharvest Biology and Technology 148 (2019) 192–199

Table 2
Anthocyanin contents of the ‘Van’ sweet cherries during storage under the
control (in the dark) and irradiation treatments.a.
Day Storage Anthocyanin (mg kg−1)

Cyanidin 3-O-
glucoside

0 Control 30.7 ± 5.0 951 Cyanidin

3-O-rutinoside.8 ± 115.2
2 Control 34.0 ± 4.8 1144.4 ± 116.3
UV-B 45.3 ± 10.1 1344.1 ± 202.5
Blue 36.8 ± 4.5 1183.8 ± 164.7
WBG 49.6 ± 6.7 1507.2 ± 159.3
4 Control 33.3 ± 6.9 b 1049.8 ± 83.3 b
UV-B 34.6 ± 4.6 b 1136.8 ± 116.1 b Fig. 3. Flavonoid 3′-hydroxylase relative activities for the ‘Van’ sweet cherries
Blue 51.9 ± 9.4 a 1749.0 ± 358.5 a
during their storage under the control (in the dark) and irradiation treatments
WBG 40.3 ± 5.7 ab 1344.9 ± 160.8 ab
(as indicated). Data are means ± SD (n = 4). Means with different letters are
7 control 35.0 ± 3.1 b 1190.3 ± 83.4 b
UV-B 39.1 ± 2.6 b 1260.4 ± 119.2 b significantly different.
Blue 68.2 ± 14.0 a 1876.3 ± 281.7 a
WBG 46.7 ± 9.1 b 1597.4 ± 265.3 ab
applying blue light with wavelength around 450 nm resulted in high
10 control 37.6 ± 4.2 b 1281.9 ± 105.1 c
UV-B 40.6 ± 5.3 b 1348.2 ± 84.1 c amounts of anthocyanins and developed a dark red color in ‘Jonathan’
Blue 106.3 ± 25.3 a 2252.1 ± 160.4 a apples. However, Sgherri et al. (2015) reported increased anthocyanins
WBG 62.7 ± 6.3 b 1893.8 ± 112.2 b in peaches during UVeB light irradiation, while in the present study
a
there were no significant changes between the control and UV-B light
WBG, white, blue and green light.
treated fruit, although anthocyanin contents in both of these groups
Data are means ± SD (n = 4). Means with different letters within each row are
increased by about 35%–40%. These data are in agreement with those
significantly different (P < 0.05).
of Gonçalves et al. (2007), who studied the evolution of color in sweet
cherries during postharvest storage at different temperatures. Data for
the anthocyanin content were significantly and negatively correlated
with the CIE parameters and hue and C* as was reported by Gonçalves
et al. (2007).
The PAL activity for all of the treatments increased during storage
(Fig. 2). The highest increase was seen for the blue light (almost 5-fold),
followed by WBG light, with an almost 4-fold increase. No significant
differences in PAL activity were found between the UV-B light and
control samples after 10 days. Similar data were also reported by Xu
et al. (2014a), who reported that PAL activities in both control and blue
light treated strawberries increased during the initial 6 days of storage,
and it was significantly higher for the blue light, which correlated with
anthocyanin accumulation. PAL activity, which is linked to stress re-
sponses, was correlated with the anthocyanin content. Indeed, there
Fig. 2. Phenylalanine ammonia lyase relative activities for the ‘Van’ sweet was highly significant correlation between Cy-3-r and PAL activity
cherries during their storage under the control (in the dark) and irradiation (0.910, P < 0.001). On the other hand, there was no correlation be-
treatments (as indicated). Data are means ± SD (n = 4). Means with different
tween any of the treatments and F3´H activity (data not shown). As
letters are significantly different.
seen in Table 3, the data for the PAL activity were highly correlated to
the CIE parameters and to hue, C*, and ΔE.
the end of the treatments, Cy-3-r in the irradiated fruit varied from The data for the F3´H activity did not appear to show any patterns
1348.2 mg kg−1 to 2252.1 mg kg−1, while Cy-3-glu varied from (Fig. 3). On the fourth day, there was a slight increase, which was more
40.6 mg kg−1 to 106.3 mg kg−1. Their highest contents were obtained pronounced in the control fruit. By the end of the treatments, the F3´H
after blue light irradiation, where Cy-3-r and Cy-3-glu increased by activity in the control and the UV-B, blue, and WBG light treated sweet
approximately 137% and 246%, respectively. These were followed by cherries was reduced to 29%, 35%, 48%, and 56%, respectively, com-
WBG light, where Cy-3-r and Cy-3-glu increased by 99% and 104%, pared to their initial activities.
respectively. A recent study by Arakawa et al. (2016) also showed that

Table 3
Correlations for the individual anthocyanin contents and the PAL activities with the changes in the color measures for the ‘Van’ sweet cherries during storage under
the control (in the dark) and irradiation treatments.a.
ΔL* Δa* Δb* hue C* ΔE PAL

Cyanidin 3-O-glucoside −0.710** −0.758*** −0.745*** −0.833*** −0.798*** 0.785*** 0.844***

Cyanidin 3-O-rutinoside −0.656** −0.807*** −0.796*** −0.892*** −0.852*** 0.790*** 0.910***
PAL −0.844*** −0.867*** −0.899*** −0.929*** −0.902*** 0.873*** /

a
hue, hue angle; C*, chroma; ΔE, total color difference; PAL, phenylalanine ammonia-lyase.
Data are means ± SD.
* P < 0.05.
** P < 0.01.
*** P < 0.001.

196
 D. Kokalj et al.

Table 4
Phenolic compounds for the ‘Van’ sweet cherries during storage under the control (in the dark) and irradiation treatments.
Day Storage Phenolic compound (mg kg−1)a

NCHA procy-B2 procy-tri 3pCoQA CAT CHA 4-CQA ECAT QGLUR ICHA Q3R K3R

0 Control 577.94 ± 25.88 108.88 ± 7.48 6.47 ± 1.67 55.30 ± 2.51 26.45 ± 5.72 54.02 ± 3.71 13.45 ± 1.83 77.72 ± 8.81 1.84 ± 0.28 17.91 ± 0.98 3.83 ± 0.73 0.45 ± 0.12
2 Control 602.78 ± 28.75 107.09 ± 9.00 5.00 ± 1.15 53.46 ± 5.52 24.23 ± 2.21 53.13 ± 4.46 12.29 ± 1.98 72.13 ± 11.52 2.00 ± 0.07 18.85 ± 0.55 4.24 ± 0.41 0.48 ± 0.13
UV-B 617.41 ± 31.72 111.88 ± 9.06 5.63 ± 1.51 52.07 ± 1.46 24.86 ± 5.29 55.50 ± 4.49 13.50 ± 2.28 77.64 ± 12.08 1.78 ± 0.16 17.64 ± 0.86 5.27 ± 1.20 0.77 ± 0.25
Blue 584.34 ± 32.86 105.32 ± 6.15 5.93 ± 0.36 50.51 ± 3.09 23.44 ± 3.03 52.25 ± 3.05 12.89 ± 1.08 73.61 ± 3.56 1.52 ± 0.20 18.08 ± 0.44 3.94 ± 1.16 0.43 ± 0.21
WBG 636.41 ± 19.55 106.99 ± 2.55 5.65 ± 0.32 53.23 ± 3.27 26.24 ± 4.70 53.08 ± 1.27 13.01 ± 2.09 71.92 ± 7.30 1.71 ± 0.05 18.41 ± 0.68 4.56 ± 0.39 0.54 ± 0.05
4 Control 629.62 ± 47.83 111.21 ± 8.71 6.38 ± 0.67 54.04 ± 6.40 25.94 ± 3.50 55.17 ± 4.32 14.36 ± 1.04 83.24 ± 5.01 1.63 ± 0.35 19.54 ± 1.10 3.44 ± 0.76 0.36 ± 0.10
UV-B 626.04 ± 23.85 108.52 ± 2.72 5.72 ± 0.57 51.56 ± 2.80 23.75 ± 5.27 53.84 ± 1.35 13.90 ± 1.22 79.31 ± 3.16 1.65 ± 0.15 19.67 ± 0.20 3.47 ± 0.27 0.36 ± 0.07
Blue 621.08 ± 60.15 106.10 ± 10.74 5.23 ± 2.49 50.49 ± 7.93 22.65 ± 5.88 52.64 ± 5.33 13.11 ± 2.98 75.32 ± 13.73 1.99 ± 0.55 22.24 ± 4.80 4.76 ± 1.26 0.36 ± 0.09

197
WBG 640.96 ± 26.34 114.93 ± 2.08 6.23 ± 0.27 51.15 ± 3.48 28.04 ± 2.69 57.02 ± 1.03 12.87 ± 1.22 72.74 ± 6.23 1.99 ± 0.12 19.49 ± 0.63 5.91 ± 1.42 0.92 ± 0.41
7 Control 636.48 ± 48.86 112.90 ± 6.63 6.51 ± 0.53 57.30 ± 5.20 26.92 ± 2.31 56.01 ± 3.29 15.37 ± 1.18 87.63 ± 7.62 1.68 ± 0.15 19.05 ± 1.09 4.16 ± 0.79 0.48 ± 0.19
UV-B 566.32 ± 30.89 96.47 ± 5.47 4.89 ± 0.81 51.26 ± 3.29 21.36 ± 3.32 47.86 ± 2.71 11.86 ± 1.20 69.68 ± 6.87 1.42 ± 0.27 17.24 ± 0.70 3.08 ± 0.31 0.24 ± 0.02
Blue 624.80 ± 30.63 107.28 ± 4.17 5.50 ± 0.65 55.25 ± 2.77 24.11 ± 3.23 53.22 ± 2.07 13.88 ± 1.30 78.77 ± 8.22 1.48 ± 0.17 19.06 ± 1.46 4.73 ± 1.17 0.42 ± 0.21
WBG 597.16 ± 57.08 107.31 ± 13.49 5.29 ± 1.06 51.96 ± 5.27 25.37 ± 5.22 53.24 ± 6.69 13.11 ± 1.59 73.24 ± 6.64 1.56 ± 0.30 18.55 ± 1.95 4.29 ± 0.95 0.45 ± 0.21
10 Control 629.68 ± 59.91 109.47 ± 13.20 6.61 ± 2.11 56.04 ± 8.29 26.22 ± 6.46 54.31 ± 6.55 14.23 ± 2.85 81.0 ± 11.35 1.65 ± 0.21 18.17 ± 2.00 4.39 ± 0.89 0.54 ± 0.29
UV-B 572.81 ± 29.32 100.94 ± 5.53 5.81 ± 1.16 52.23 ± 2.84 23.95 ± 2.55 50.08 ± 2.75 13.93 ± 1.97 78.65 ± 9.87 1.26 ± 0.06 16.74 ± 1.48 3.99 ± 0.85 0.45 ± 0.28
Blue 589.65 ± 56.89 104.89 ± 12.34 4.97 ± 1.09 49.30 ± 8.15 23.90 ± 3.89 52.04 ± 6.12 12.19 ± 2.96 67.05 ± 13.22 1.75 ± 0.12 20.52 ± 1.53 6.27 ± 1.17 0.50 ± 0.24
WBG 660.98 ± 23.45 110.51 ± 6.35 7.69 ± 0.77 57.20 ± 1.68 28.04 ± 4.31 54.82 ± 3.15 15.90 ± 1.13 87.03 ± 3.23 1.73 ± 0.34 19.63 ± 1.20 4.94 ± 0.60 0.41 ± 0.12
Mean 607.41 ± 43.06 107.87 ± 8.09 5.95 ± 1.30 53.41 ± 4.63 25.24 ± 4.32 53.51 ± 4.02 13.51 ± 1.89 76.99 ± 9.25 1.71 ± 0.29 18.73 ± 1.82 4.34 ± 1.10 0.48 ± 0.22

a
WBG, white, blue and green light; NCHA, neochlorogenic acid; procy-B2, procyanidin B2; procy-tri, procyanidin trimer; 3pCoQA, 3-p-coumaroylquinic acid; CAT, catechin; CHA, chlorogenic acid; 4-CQA, crypto-
chlorogenic acid; ECAT, epicatechin; QGLUR, quercetin-3-(2G-glucosyl-rutinoside); ICHA, isochlorogenic acid; Q3R, quercetin-3-O-rutinoside; K3R, kaempferol-3-O-rutinoside.
Data are means ± SD (n = 4).
Postharvest Biology and Technology 148 (2019) 192–199
D. Kokalj et al.
Fig. 4. Ascorbic acid content of the ‘Van’ sweet cherries during their storage
under the control (in the dark) and irradiation treatments (as indicated). Data
are means ± SD (n = 4). Means with different letters are significantly dif-
ferent.
Twelve phenolic compounds were evaluated in these ‘Van’ sweet
cherries during storage under these three different irradiation treat-
ments: neochlorogenic acid, catechin, chlorogenic acid, kaempferol-3-
O-rutinoside, quercetin-3-O-rutinoside, 3-p-coumaroylquinic acid,
cryptochlorogenic acid, epicatechin, isochlorogenic acid, procyanidin
B2, procyanidin trimer, and quercetin-3-(2G-glucosyl-rutinoside). The
predominant hydroxycinnamic acid was neochlorogenic acid, followed
by 3-p-coumaroylquinic acid, which is in agreement with previous re-
ports (Chockchaisawasdee et al., 2016). However, neither monochro-
matic light irradiation nor the combinations of light spectra stimulated
the accumulation of these phenolic compounds (Table 4). These data
are similar to Hagen et al. (2007), where their contents of epicatechin
and procyanidins in postharvest irradiated apples were minimally in-
fluenced by the treatments, while the chlorogenic acid content in the
fruit increased after irradiation with a combination of visible and UV-B
light.
Ascorbic acid varied greatly between the batches of fruit in the
present study, from ∼10 mg kg−1 to 70 mg kg−1, with an overall mean
just below 30 mg kg−1 (data not shown). While ascorbic acid in the
control fruit decreased, there was a small increase in ascorbic acid in
the irradiated fruit after two days (Fig. 4). Interestingly blue and WBG
light, which had the most pronounced effects on the anthocyanins and
PAL activity, negatively influenced the ascorbic acid content only at the
end of storage, i.e. after 10 days of treatment. At the end of the treat-
ments, the ascorbic acid content of the blue in WBG light irradiated
fruit was lower than the initial content, as ∼15 mg kg−1 and 11 mg
kg−1, respectively. On the other hand, the control and UV-B irradiated
fruit had higher ascorbic acid, at nearly 44 mg kg−1 and 43 mg kg−1,
respectively. Changes in the content of ascorbic acid in fruit during
storage depend on the fruit species, maturity, storage conditions, and
other factors. Kalt et al. (1999) reported that there were no losses in
ascorbate in strawberries and highbush blueberries during storage at
various temperatures, while there were losses in lowbush blueberries
and raspberries. On the other hand, Xu et al. (2014b) reported de-
creases in ascorbic acid in strawberries during storage. The literature
data thus show mixed results for ascorbic acid content of fruit during
storage. However, blue light (470 nm) irradiated strawberries showed
higher ascorbic acid than the control fruit (Xu et al., 2014b). Similar
positive effects of blue light irradiation were reported by Kim et al.
(2011) for immature strawberries. Although, the vitamin C content of
all fruit increased during their storage, there was relatively high content
in the fruit treated with blue light (470 nm) (Kim et al., 2011).
4. Conclusions
Synthesis of many bioactive compounds is light dependent. The data
in the present study showed that postharvest irradiation with blue light
might have an impact on the regulation of anthocyanin synthesis and
198
Postharvest Biology and Technology 148 (2019) 192–199
the improved quality of sweet cherries. Blue light provoked the highest
increase in PAL activity, followed by WBG light. Blue light proved to be
the most efficient in accumulation of Cy-3-glu and Cy-3-r. Highly sig-
nificant correlations were found for the individual anthocyanin con-
tents and PAL activities with the changes in the color measures.
Interestingly, WBG light highly positively influenced anthocyanin ac-
cumulation, although its intensity was relatively low. Regarding ben-
eficial eff ;ects, LEDs with lower light intensity might be suitable for
broader commercial use, such as installation in refrigerators, not only to
illuminate the compartments, but also to potentially maintain or even
improve the food quality. It is evident from the present study that
postharvest irradiation can have an impact on the complex regulation
of anthocyanin synthesis, and consequently on sweet cherry color de-
velopment. However, further studies are needed to evaluate the impact
of cultivar, maturity stage, treatment duration and storage temperature
under LEDs irradiation.
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