Scientia Horticulturae 198 (2016) 304–310

Contents lists available at ScienceDirect

Scientia Horticulturae
journal homepage: www.elsevier.com/locate/scihorti

Effect of foliar treatment of sodium selenate on postharvest decay and

quality of tomato fruits
Zhu Zhu a,∗ , Yanli Chen a , Xueji Zhang a , Miao Li b
a
Research Center for Bioengineering and Sensing Technology, University of Science & Technology Beijing, Beijing 100083, China
b
Key Laboratory of Agri-Food Safety of Anhui Province, School of Plant Protection, Anhui Agriculture University, Hefei 230036, China

a r t i c l e i n f o a b s t r a c t

Article history: Postharvest decay and quality deterioration of fruits are responsible for significant economic losses in
Received 24 July 2015 the tomato industry. Selenium (Se) is an important microelement due to its antioxidant properties that
Received in revised form allows detoxifying various reactive oxygen species (ROS). In the present study, the effect of a foliar treat-
24 November 2015
ment of Se prior to fruit development on the subsequent postharvest decay and quality of tomato fruits
Accepted 1 December 2015
was investigated. Foliar treatment of 1 mg L−1 sodium selenate at the time of fruit onset and develop-
ment effectively controlled gray mold rot caused by Botrytis cinerea on both inoculated and naturally
Keywords:
infected tomato fruit. Se treatment reduced lipid peroxidation, enhanced antioxidant enzyme activity
Selenium
Tomato fruit
of glutathione peroxidase and superoxide dismutase, and increased the concentration of non-enzymatic
Postharvest decay antioxidants, such as reduced ascorbate and reduced glutathione in the tomato fruits. Se also contributed
Quality to the maintenance of fruit quality during storage. The results suggested that Se stimulated the antioxi-
dant defense system in tomato plants, thus making the fruits more resistant to postharvest decay caused
by gray mold. Selenium treatment represents a promising strategy for managing postharvest decay and
maintaining the quality of tomato fruits.
© 2015 Elsevier B.V. All rights reserved.

1. Introduction increasing indications that Se may also have beneficial biological

Tomatoes (Solanum lycopersicon L.) are one of the major veg-
etable components of the human diet worldwide. The fleshy fruits,
however, are susceptible to decay and loss of quality once they have
been harvested. Gray mold rot, caused by Botrytis cinerea, is the
major postharvest disease of tomato fruits (Dal Bello et al., 2008;
Fagundes et al., 2014). A number of alternative strategies to the
use of synthetic, chemical fungicides have been evaluated for their
ability to prevent postharvest decay and extend the shelf-life of
tomato fruits, including the application of physical agents (Stevens
et al., 2004; Bailen et al., 2006), biological control agents (Lee et al.,
2006; Wang et al., 2010), and chemical treatments that induce host
resistance (Liu et al., 2007; Lai et al., 2011; Zhu and Tian, 2012).
Selenium (Se) is an essential micronutrient that plays a role
in antioxidant defense systems and maintenance of hormone bal-
ance in human and animal cells. It is an integral part of the
enzyme, glutathione peroxidase (GPX), and a number of other
selenoproteins (Ellis and Salt, 2003). Even though higher plants
may not require Se and are sensitive to Se toxicity, there are
∗ Corresponding author. Fax: +86 10 82376993.
E-mail address: zhuzhu@ustb.edu.cn (Z. Zhu).
effects on higher plants due to its antioxidant properties (Stadtman,
1990; Roman et al., 2014). Se at low concentrations can stimulate
plant growth (Hartikainen et al., 2000), ameliorate the detrimental
effects of diverse environmental stresses including cold (Chu et al.,
2010), high temperature (Djanaguiraman et al., 2010), UV-B (Yao
et al., 2010), drought (Hasanuzzaman and Fujita, 2011), desiccation
(Pukacka et al., 2011), water (Wang, 2011), salt (Hasanuzzaman
et al., 2011), heavy metals (Kumaret al., 2012) and senescence (Xue
et al., 2001; Djanaguiraman et al., 2005). Applications of Se have
been found to prolong the shelf life of peach and pear (Pezzarossa
et al., 2012), and alter the level of antioxidant compounds in tomato
fruits (Schiavon et al., 2013). While the possible mechanisms asso-
ciated with Se-enhanced resistance and tolerance to environmental
stresses in plants have not been fully demonstrated, Feng et al.
(2013) have implicated its potential role in the regulation of reac-
tive oxygen species (ROS), and the maintenance of cellular structure
and function under abiotic stress conditions. In addition to the gen-
eration of ROS by abiotic stresses, invasion of plant tissues by fungal
pathogens also induces the generation of ROS in plants, resulting
in oxidative stress which can affect the integrity of cell membranes
and interfere with key cellular functions (Xu and Tian, 2008). To our
knowledge, however, little information is available about the effect
of Se on pathogen-induced oxidative stress in harvested fruit.

http://dx.doi.org/10.1016/j.scienta.2015.12.002
0304-4238/© 2015 Elsevier B.V. All rights reserved.
 Z. Zhu et al. / Scientia Horticulturae 198 (2016) 304–310
The objective of the present study was to investigate the effect
of Se on (i) postharvest decay in tomato fruits caused by gray mold
(B. cinerea); (ii) antioxidant compounds in fruits inoculated with
B. cinerea, with a special focus on glutathione peroxidase (GPX),
superoxide dismutase (SOD), reduced ascorbate (AsA), and reduced
glutathione (GSH); (iii) lipid peroxidation; and (iv) fruit quality
parameters during storage.
2. Materials and methods
2.1. Fruit and Se treatment
Experiments were conducted on tomato plants (S. lycopersi-
con, cv. Provence) grown in a greenhouse located in the Shandong
province of China. The physical composition and pH of the soil
were as follows: 49% sand, 29% silt, and 22% clay, organic carbon
1.61%, and pH 6.5. Plants were grown using standard cultural prac-
tices, with regular supply of fertilizer and supplementary lighting
when required. Flowers were tagged on the day of anthesis, and
fruits were harvested at the mature green stage, which occurred at
approximately six weeks post anthesis. A series of sodium selenate
concentrations (0.1, 1, 10 mg Se L−1 ) were evaluated in a prelim-
inary experiment, and 1 mg Se L−1 was selected as the optimal
concentration for further study.
Foliar spray of 1 mg Se L−1 as sodium selenate (Na2 SeO4 ) was
applied to tomato plants at approximately four weeks after trans-
planting, when the onset of flowering had commenced. The sodium
selenate was obtained from Sigma–Aldrich (Shanghai) Trading Co.,
Ltd., as a colorless, crystalline powder. Twenty eight plants were
used in Se treatment and each plant was sprayed with 1 L of the
Se solution. Twenty eight additional plants were sprayed with an
equal volume of water and served as the control. During foliar Se
treatment, the soil was covered to avoid Se contamination. The
experiments were conducted once in 2013 and twice in 2014. In
each experiment, fruits were harvested three times at the mature
green stage (between 40 and 55 days after Se treatment). At the
time of each harvest, 150 fruits were obtained from each of the
control and Se-treated plants, respectively. Half of the fruits from
Se-treated plants and non-treated control were subjected to inoc-
ulation with spores of B. cinerea, while the other half were used to
monitor natural infections and measure quality parameters.
2.2. Determination of Se content
Se content was determined as described by Pedrero et al. (2006).
Briefly, dried fruit tissue (100 mg) from 10 fruits per treatment was
digested with 1.5 mL of concentrated HNO3 (65%) and 1.5 mL of
H2 O2 (30%) in an analytical microwave oven. The resulting solution
was diluted to 35 mL with deionized water and analyzed by induc-
tively coupled plasma mass spectroscopy (ICP-MS). Each treatment
consisted of three replicates, and the experiment was repeated
three times.
2.3. Fungal culture
B. cinerea was isolated from naturally infected tomato fruit.
The culture was purified by single spore isolation and cultured on
potato dextrose agar (PDA) at 25 ◦ C for 14 days. Conidia were col-
lected by flooding the culture surface with sterile distilled water
containing 0.05% (v/v) Tween-80. The suspension was filtered
through four layers of sterile cheesecloth and adjusted to the con-
centration of 1 × 104 spores mL−1 using a hemocytometer.
305
2.4. Effect of Se on postharvest decay in tomato fruit
Effect of Se on the postharvest decay of tomato fruits caused by
gray mold (B. cinerea) was evaluated by inoculated tomato fruits.
Two wounds (3 mm deep × 3 mm wide) were made at the equator
of each fruit with a sterile nail. Then, 10 ␮L of the spore suspension
of B. cinerea described above was pipetted onto each wound. The
inoculated fruits were put into plastic trays, covered with a plastic
film to maintain a high relative humidity (95%), and stored at 25 ◦ C.
Disease incidence and lesion diameter were determined daily for
3 days. Each treatment consisted of 3 replications, each of which
contains 15 fruits. To assess the effect of Se on natural decay of
tomato fruit, control and Se-treated fruits were stored at 25 ◦ C for
20 days, and naturally-occurring disease incidence was recorded at
5-day intervals. Each treatment contained 15 fruits and consisted
of 3 replications with triplicates.
2.5. Measurement of lipid peroxidation
Fruit tissue from each treatment was sampled daily for 3 days,
beginning one hour after inoculation as 0 day. Ten grams of frozen,
healthy flesh tissue in proximity to the infected, macerated tissue
was obtained from 10 fruits and combined into a single replicate.
Three replicates were obtained from each treatment, and the exper-
iment was repeated three times. Levels of malondialdehyde (MDA)
were measured using the thiobarbituric acid method described
by Wang et al. (2005). Absorbance at 532 nm was recorded and
corrected for non-specific absorbance at 600 nm. The concentra-
tion of MDA was calculated from the extinction coefficient of
155 mM−1 cm−1 and expressed as nmol mg−1 protein.
Lipoxygenase (LOX) activity was measured as described by
Wang et al. (2005). LOX activity was calculated following the
rise in the extinction at A234 using an extinction coefficient of
25,000 M−1 cm−1 and expressed as U mg−1 protein, where one unit
represents 1 ␮mol hydroperoxide produced per minute at 30 ◦ C.
Protein content was determined using BCA Protein Assay Kit
(Pierce, Thermo Scientific, USA) with bovine serum albumin as stan-
dard.
2.6. Antioxidant enzyme activity
Ten grams of healthy fruit tissue was collected from ten fruits
on a daily basis for 3 days as described above. Three replicates were
analyzed from each treatment, and the experiment was repeated
three times. Glutathione peroxidase (GPX) assay was conducted as
described by Pukacka et al. (2011). Briefly, GPX activity was deter-
mined by monitoring the disappearance of NADPH at 340 nm for
15 min, and expressed as U mg−1 protein, where one unit repre-
sents the nmol of NADPH produced per minute at 25 ◦ C.
Superoxide dismutase (SOD) activity was measured at 560 nm
according to the method described by Wang et al. (2005). SOD activ-
ity was also expressed as U mg−1 protein. One unit was defined as
the amount of enzyme that caused 50% decrease in NBT reduction.
2.7. Measurement of reduced ascorbate and reduced glutathione
content
Ten grams of frozen, healthy flesh tissue was collected from
10 fruits as described above. Three replicates were analyzed from
each treatment and the experiment was repeated three times. AsA
content was measured as described by Ding et al. (2007). Deter-
mination was based on the reduction of Fe3+ to Fe2+ by AsA. The
complex of Fe2+ and 2,2-dipyridyl was detected at 525 nm. AsA
content was expressed as ␮mol g−1 .
GSH level was determined using the DTNB-GSSG reductase recy-
cling assay (Loggini et al., 1999). Neutralized samples and samples
306 Z. Zhu et al. / Scientia Horticulturae 198 (2016) 304–310

Table 1
Effect of foliar treatment of 1 mg L−1 of sodium selenate to tomato plants prior to
fruit development on the Se content of the fruit at the mature green stage.

Treatment Selenium content (␮g kg−1 DW)

H2 O 30.2 ± 4.3
Se 500.0 ± 10.6a
a
Indicate significant differences between control and Se-treated fruit according
to the Student’s t-test (P < 0.05). Data represent the mean ± standard error of nine
replicates pooled from three experiments.

pretreated with 2-vinyl pyridine were used for determination of

total glutathione and oxidized glutathione (GSSG), respectively. A
standard curve was used for calculating the amount of total glu-
tathione and GSSG. GSH was determined by subtraction of GSSG
from the total glutathione content. GSH content was expressed as
␮mol g−1 .

2.8. Measurements of quality parameters

Soluble solids content (SSC) was determined using a hand-held

refractometer (Atago, Tokyo, Japan). Titratable acidity (TA) was
determined by titration with 0.01 M NaOH. Flesh firmness was
assessed using a TA-XT2 Texture analyser (Stable Micro Systems,
Surrey, UK) by penetrating the fruit pericarp with a cylindrical
probe (2 mm diameter) at a constant speed (2 mm/s). Fruit color
was measured using a Minolta CR-300 chromameter (Minolta,
Osaka, Japan) as L* , a* , b* values and converted to hue angle (color
wheel, with red-purple at an angle of 0◦ , yellow at 90◦ , bluish-green
at 180◦ ). The color reading was taken twice at the equatorial region Fig. 1. Effect of foliar treatment of 1 mg L−1 of sodium selenate to tomato plants
prior to fruit development on disease incidence (A) and lesion diameter (B) in the
of each fruit and averaged to give a value for each fruit. Each assay of
fruits inoculated with Botrytis cinerea. Water used as the control. Data represent the
quality index contained three replicates with 10 fruits per replicate, mean ± standard error of nine replicates pooled from three experiments. Asterisks
and the experiment was repeated three times. indicate significant differences between control and Se-treated fruit according to
the Student’s t-test (P < 0.05).

2.9. Statistical analysis

Statistical analysis was performed with SPSS 13.0. Data were

analyzed using the Shapiro-Wilk test for normality, and then com-
pared in a Student’s t-test. Differences at P < 0.05 were considered
as significant.

3. Results

3.1. Se accumulation in tomato fruit

As indicated in Table 1, the Se level in mature green fruits

obtained from tomato plants treated with 1 mg Se L−1 as sodium
selenate was 16-fold higher than those sprayed with water. Pre-
liminary tests also indicated that treatment with 1 mg Se L−1 also
had the greatest impact on control of postharvest gray mold fruit
decay.
3.2. Effect of Se on postharvest gray mold fruit decay
In the preliminary study, application of 10 mg Se L−1 as sodium
selenate to tomato plants resulted in a reduction in the yield of
tomato plants by 16%, while 0.1 mg Se L−1 application had no obvi-
ous inhibitory effect on the incidence of postharvest decay (data
not shown). In contrast, 1 mg Se L−1 had a positive effect on inhibi-
tion of postharvest decay, reducing the disease incidence by 25%.
Based on these preliminary results, application of 1 mg Se L−1 was
selected for use in the further studies.
These results indicated that only a single application of
1 mg Se L−1 to tomato plants before fruit development significantly
reduced the decay incidence in the fruits that harvested and stored
long after Se application (Fig. 1A). The lesion diameter of gray mold
Fig. 2. Effect of foliar treatment of 1 mg L−1 of sodium selenate to tomato plants
prior to fruit development on the subsequent natural decay of tomato fruit up to 20
days after harvest. Water used as the control. Data represent the mean ± standard
error of nine replicates pooled from three experiments. Asterisks indicate significant
differences between control and Se-treated fruit according to the Student’s t-test
(P < 0.05).
infections was also significantly smaller in tomato fruits from Se-
treated plants than those sprayed with water (Fig. 1B). Notably,
tomato fruits from Se-treated plants had lower incidence (11.1%)
of natural decay than those from Se-non-treated, water-sprayed
plants (28.9%) after 20 days of storage at 25 ◦ C (Fig. 2). In gen-
eral, 1 mg Se L−1 treatment had significant suppressive effects on
the fruit decay regardless of gray mold infection.
 Z. Zhu et al. / Scientia Horticulturae 198 (2016) 304–310
Fig. 3. Effect of foliar treatment of 1 mg L−1 of sodium selenate to tomato plants prior to fruit development on lipid peroxidation as indicated by malondialdehyde content,
lipoxygenase, glutathione peroxidase, and superoxide dismutase activities in tomato fruits inoculated with Botrytis cinerea, the causal agent of gray mold. Water used as the
control. Data represent the mean ± standard error of nine replicates pooled from three experiments. Asterisks indicate significant differences between control and Se-treated
fruit according to the Student’s t-test (P < 0.05).
3.3. Effect of Se on lipid peroxidation and antioxidant enzyme
activity in tomato fruit
MDA content in inoculated, control fruits increased sharply
and reached a peak on the second day after inoculation, while
MDA formation was significantly less in inoculated fruits from Se-
treated plants (Fig. 3A). LOX activity increased regardless of the
Se-treatment, however, the increase levels were again less in the
fruits from Se-treated plants 2 days after inoculation (Fig. 3B). These
results indicated that Se-treatment on the tomato plants at the time
of fruit onset can ameliorate lipid peroxidation in tomato fruits
infected with gray mold.
Regarding antioxidant enzymes, in both inoculated fruits from
control and Se-treated plants had an initial increase in GPX activ-
ity 2 days after inoculation, followed by a gradual decline (Fig. 3C).
The level of GPX activity, however, was three times higher in the
Se-treated than that in the control. The differences in SOD activity
between the fruits from control and Se-treated plants were similar
to the results obtained on GPX. In the case of SOD activity, how-
ever, the SOD activity levels of both treatment were initially high
and gradually declined over 3-day period but the decline in the
activity was less in the Se-treated compared to that in the control
(Fig. 3D).
3.4. Effect of Se on reduced ascorbate and reduced glutathione
content in tomato fruit
As shown in Fig. 4A, AsA content in inoculated control fruits
increased slightly and then decreased, while AsA level was
markedly higher in inoculated fruits from Se-treated plants. GSH
contents in both inoculated fruits from control and Se-treated
plants were relatively stable, while the overall level of GSH in Se-
treated was at least twice higher than that in the control (Fig. 4B).
307
3.5. Effect of Se on fruit quality
SSC content in all fruits increased during the first 15 days after
harvest and then decreased (Fig. 5A). The SSC level in fruits from
Se-treated plants tended to be slightly higher, especially at harvest
(Day 0) and at day 20, when differences between the treatments
were statistically significant. Titratable acidity (TA) and firmness
declined gradually in both treatments, however, the decline level
in both of these parameters was significantly less in fruits from Se-
treated plants (Fig. 5B and C). Thus, both the decline in TA and the
loss of firmness were delayed in fruits from Se-treated plants. The
hue angle decreased over the 20-day period after harvest, indicat-
ing a change in fruit color from green to red. No differences were
observed, however, in hue angle between the treatments (Fig. 5D).
4. Discussion
This study presents the first report of a positive effect of sele-
nium on postharvest decay in tomato fruit and suggests that
1 mg Se L−1 could significantly reduce the incidence and severity
of postharvest decay in tomato fruit caused by gray mold.
A previous report indicated that Se could protect Brassica juncea
plants, which hyper-accumulated up to 300 mg kg−1 DW of Se,
from herbivory by invertebrates and fungal infections (Hanson
et al., 2003). The basis postulated for the protection was that the
hyper-accumulated levels of Se were toxic to both the herbivores
and fungi. For adults, the dietary intake for Se recommended by
the World Health Organization is 40 ␮g day−1 , whereas a tolera-
ble dose is described as 400 ␮g day−1 (Combs, 2001). The average
fresh weight of a treated tomato fruit is approximately 160 g,
which corresponds to 10 g DW, and the average Se concentration is
500 ␮g kg−1 DW, or, about 5 ␮g Se in each treated fruit. Therefore,
the concentration of selenate used in the current study resulted in
Se-biofortified tomato fruit in which Se level was low enough not
308 Z. Zhu et al. / Scientia Horticulturae 198 (2016) 304–310

Fig. 4. Effect of foliar treatment of 1 mg L−1 of sodium selenate to tomato plants prior to fruit development on reduced ascorbate and reduced glutathione contents in tomato
fruits inoculated with Botrytis cinerea, the causal agent of gray mold. Water used as the control. Data represent the mean ± standard error of nine replicates pooled from three
experiments. Asterisks indicate significant differences between control and Se-treated fruit according to the Student’s t-test (P < 0.05).

Fig. 5. Effect of foliar treatment of 1 mg L−1 of sodium selenate to tomato plants prior to fruit development on soluble solids content, titratable acidity, firmness, and hue
angle of tomato fruits stored at 25 ◦ C for 20 days. Water used as the control. Data represent the mean ± standard error of nine replicates pooled from three experiments.
Asterisks indicate significant differences between control and Se-treated fruit according to the Student’s t-test (P < 0.05).

to pose a health risk to humans, but rather could even be regarded
as a human dietary supplement.
Plants under pathogens’ attack are subjected to oxidative stress,
and one of the most commonly used indicators of oxidative stress
is the level of lipid peroxidation, expressed as MDA concentration
and LOX activity (Ríos et al., 2008). Therefore, one way of evalu-
ating the protective effect of Se was to evaluate these indicators
of oxidative stress in fruits from control and Se-treated plants that
had been inoculated with gray mold. Results demonstrated that
foliar application of Se prior to fruit development reduced sub-
sequent MDA production and LOX activity in the harvested fruit
when inoculated with B. cinerea, indicating that lipid peroxidation
in fruit tissue was significantly lower in the tomato from Se-treated
plants (Fig. 3A and B). The MDA results were consistent with pre-
vious studies which reported that Se at low concentrations acts as
 Z. Zhu et al. / Scientia Horticulturae 198 (2016) 304–310
an antioxidant and reduces lipid peroxidation (Hartikainen et al.,
2000; Xue et al., 2001; Djanaguiraman et al., 2005).
Plants contain antioxidant enzymes and a wide array of non-
enzymatic antioxidants to deal with injurious levels of ROS. SOD
eliminates O2 − through the formation of H2 O2 , which is a ROS com-
pound. GPX, together with GSH, is a powerful scavenger of H2 O2
and organic peroxides (Ríos et al., 2008). Our data demonstrated
that the levels of GPX and SOD activity in fruits form Se-treated
plants were higher than their corresponding levels in untreated,
control fruit when inoculated with B. cinerea (Fig. 3C and D). Sig-
nificant increases in the GPX and SOD activity have also been
observed in other studies of Se-treated plants subjected to a variety
of stresses, including senescing lettuce (Xue et al., 2001), soy-
bean (Djanaguiraman et al., 2005), and cadmium-stressed marine
red alga (Kumar et al., 2012). Collectively, the data suggest that
increased induction of GPX and SOD in Se-treated plants plays an
important role in ameliorating oxidative stress before ROS levels
can cause injury in cells. AsA and GSH, non-enzymatic antioxidants,
also possess an ability to scavenge ROS and assist the detoxifica-
tion of H2 O2 through the AsA-GSH cycle (Mittler, 2002). In the
present study, the concentrations of AsA and GSH were signifi-
cantly higher in tomato fruits obtained from plants that had been
pre-treated with Se (Fig. 4). These results confirm previous reports
of elevated levels of AsA and GSH in rapeseed (Brassica napus)
seedlings pre-treated with Se and then subjected to drought and
salt stress (Hasanuzzaman and Fujita, 2011; Hasanuzzaman et al.,
2011). It has been suggested that treatment of plants with Se upreg-
ulates the synthesis of enzymes needed to regenerate AsA, and also
enhances the uptake of sulfur, making it more readily available for
the synthesis of GSH (Hasanuzzaman et al., 2011; Feng et al., 2013).
Quality parameters, including SSC, TA, firmness, and color
development were evaluated, in order to determine the effect of Se-
treatments to the tomato plants at the time of fruits onset on tomato
fruit quality during storage. Results showed that Se-treatments
was effective in delaying softening, and maintaining SSC and TA
in tomato fruits regardless of gray mold infection. A similar result
was obtained by Pezzarossa et al. (2012), who reported that firm-
ness and SSC content were higher in peach fruits from Se-treated
plants than those from Se-untreated plants. These results suggest
that the ability of Se to maintain fruit quality may be linked to Se-
inhibition of ethylene production (Pezzarossa et al., 1999; Malorgio
et al., 2009).
In conclusion, foliar pre-treatment of tomato plants with
1 mg Se L−1 prior to fruit development was found to have a posi-
tive impact on reducing fruit decay incidence in harvested tomato
fruits and help maintaining fruit quality during storage. Se con-
tent in fruits from Se-treated tomato plants were sufficiently low
enough to the safety level for consumers. The possible mecha-
nism of enhanced resistance to postharvest fungal pathogens by
Se-treatment to tomato plants appears to be related to the abil-
ity of Se to stimulate the activity of antioxidant enzymes and
non-enzymatic antioxidants, as well as reduce the level of lipid per-
oxidation in tomato fruit during fungal invasion. It is concluded that
Se treatment before fruit development would be a promising alter-
native strategy for managing postharvest decay and maintaining
quality during the storage of tomato fruit.
Acknowledgments
The study was supported by the National Natural Science
Foundation of China (31401545), the China Postdoctoral Science
Foundation (2014M560893), the Fundamental Research Funds for
the Central Universities (FRF-TP-14-016A1), and the Natural Sci-
ence Foundation of Anhui Province (1408085MC68).
309
References
Bailen, G., Guillen, F., Castillo, S., Serrano, M., Valero, D., Martinez-Romero, D., 2006.
Use of activated carbon inside modified atmosphere packages to maintain
tomato fruit quality during cold storage. J. Agric. Food Chem. 54, 2229–2235.
Chu, J.Z., Yao, X.Q., Zhang, Z.N., 2010. Responses of wheat seedlings to exogenous
selenium supply under cold stress. Biol. Trace Elem. Res. 136, 355–363.
Combs, G.F., 2001. Selenium in global food systems. Br. J. Nutr. 85, 517–547.
Dal Bello, G., Rollan, M.C., Lampugnani, G., Arteta, N., Abramoff, C., Ronco, L.,
Stocco, M., 2008. Biocontrol of postharvest grey mould on tomato by yeasts. J.
Phytopathol. 156, 257–263.
Ding, Z.S., Tian, S.P., Zheng, X.L., Zhou, Z.W., Xu, Y., 2007. Responses of reactive
oxygen metabolism and quality in mango fruit to exogenous oxalic acid or
salicylic acid under chilling temperature stress. Physiol. Plant. 130, 112–121.
Djanaguiraman, M., Durga Devi, D., Shanker, A.K., Annie Sheeba, J., Bangarusamy,
U., 2005. Selenium-an antioxidative protectant in soybean during senescence.
Plant Soil 272, 77–86.
Djanaguiraman, M., Prasad, P.V.V., Seppanen, M., 2010. Selenium protects sorghum
leaves from oxidative damage under high temperature stress by enhancing
antioxidant defense system. Plant Physiol. Biochem. 48, 999–1007.
Ellis, D.R., Salt, D.E., 2003. Plants, selenium and human health. Curr. Opin. Plant
Biol. 6, 273–279.
Fagundes, C., Palou, L., Monteiro, A.R., Pérez-Gago, M.B., 2014. Effect of antifungal
hydroxypropyl methylcellulose-beeswax edible coatings on gray mold
development and quality attributes of cold-stored cherry tomato fruit.
Postharvest Biol. Technol. 92, 1–8.
Feng, R.W., Wei, C.Y., Tu, S.X., 2013. The roles of selenium in protecting plants
against abiotic stresses. Environ. Exp. Bot. 87, 58–68.
Hanson, B., Garifullina, G.F., Lindblom, S.D., Wangeline, A., Ackley, A., Kramer, K.,
Norton, A.P., Lawrence, C.B., Pilon-Smits, E.A.H., 2003. Selenium accumulation
protects Brassica juncea from invertebrate herbivory and fungal infection. New
Phytol. 159, 461–469.
Hartikainen, H., Xue, T., Piironen, V., 2000. Selenium as an anti-oxidant and
pro-oxidant in ryegrass. Plant Soil 225, 193–200.
Hasanuzzaman, M., Fujita, M., 2011. Selenium pretreatment upregulates the
antioxidant defense and methylglyoxal detoxification system and confers
enhanced tolerance to drought stress in rapeseed seedlings. Biol. Trace Elem.
Res. 143, 1758–1776.
Hasanuzzaman, M., Hossain, M.A., Fujita, M., 2011. Selenium-induced
up-regulation of the antioxidant defense and methylglyoxal detoxification
system reduces salinity-induced damage in rapeseed seedlings. Biol. Trace
Elem. Res. 143, 1704–1721.
Kumar, M., Bijo, A.J., Baghel, R.S., Reddy, C.R.K., Jha, B., 2012. Selenium and
Spermine alleviates cadmium induced toxicity in the red seaweed Gracilaria
dura by regulating antioxidant system and DNA methylation. Plant Physiol.
Biochem. 51, 129–138.
Lai, T.F., Wang, Y.Y., Li, B.Q., Qin, G.Z., Tian, S.P., 2011. Defense responses of tomato
fruit to exogenous nitric oxide during postharvest storage. Postharvest Biol.
Technol. 62, 127–132.
Lee, J.P., Lee, S.W., Kim, C.S., Son, J.H., Song, J.H., Lee, K.Y., Kim, H.J., Jung, S.J., Moon,
B.J., 2006. Evaluation of formulations of Bacillus licheniformis for the biological
control of tomato gray mold caused by Botrytis cinerea. Biol. Control 37,
329–337.
Liu, J., Tian, S.P., Meng, X.H., Xu, Y., 2007. Effects of chitosan on control of
postharvest diseases and physiological responses of tomato fruit. Postharvest
Biol. Technol. 44, 300–306.
Loggini, B., Scartazza, A., Brugnoli, E., Navari-Izzo, F., 1999. Antioxidative defense
system, pigment composition, and photosynthetic efficiency in two wheat
cultivars subjected to drought. Plant Physiol. 119, 1091–1100.
Malorgio, F., Diaz, K., Ferrante, A., Mensuali, A., Pezzarossa, B., 2009. Effects of
selenium addition on minimally processed leafy vegetables grown in floating
system. J. Sci. Food Agric. 89, 2243–2251.
Mittler, R., 2002. Oxidative stress, antioxidants and stress tolerance. Trends Plant
Sci. 7, 405–410.
Pedrero, Z., Madrid, Y., Cámara, C., 2006. Selenium species bioaccessibility in
enriched radish (Raphanus sativus): a potential dietary source of selenium. J.
Agric. Food Chem. 54, 2412–2417.
Pezzarossa, B., Malorgio, F., Tonutti, P., 1999. Effects of selenium uptake by tomato
plants on senescence, fruit ripening and ethylene evolution. In: Kanellis, A.K.,
Chang, C., Klee, H., Bleecker, A.B., Pech, J.C., Grierson, D. (Eds.), Biology and
Biotechnology of the Plant Hormone Ethylen. , II ed. Kluwer, Dordrecht, pp.
275–276.
Pezzarossa, B., Remorini, D., Gentile, M.L., Massai, R., 2012. Effects of foliar and fruit
addition of sodium selenate on selenium accumulation and fruit quality. J. Sci.
Food Agric. 92, 781–786.
Pukacka, S., Ratajczak, E., Kalemba, E., 2011. The protective role of selenium in
recalcitrant Acer saccharium L. seeds subjected to desiccation. J. Plant Physiol.
168, 220–225.
Ríos, J.J., Blasco, B., Cervilla, L.M., Rosales, M.A., Sanchez-Rodriguez, E., Romero, L.,
Ruiz, J.M., 2008. Production and detoxification of H2 O2 in lettuce plants
exposed to selenium. Ann. Appl. Biol. 154, 107–116.
Roman, M., Jitaru, P., Barbante, C., 2014. Selenium biochemistry and its role for
human health. Metallomics 6, 25–54.
Schiavon, M., dall’Acqua, S., Mietto, A., Pilon-Smits, E.A.H., Sambo, P., Masi, A.,
Malagoli, M., 2013. Selenium fertilization alters the chemical composition and
310 Z. Zhu et al. / Scientia Horticulturae 198 (2016) 304–310
antioxidant constituents of tomato (Solanum lycopersicon L.). J. Agric. Food
Chem. 61, 10542–10554.
Stadtman, T.C., 1990. Selenium biochemistry. Annu. Rev. Biochem. 59, 111–127.
Stevens, C., Liu, J., Khan, V.A., Lu, J.Y., Kabwe, M.K., Wilson, C.L., Igwegbe, E.C.K.,
Chalutz, E., Droby, S., 2004. The effects of low-dose ultraviolet light-C
treatment on polygalacturonase activity, delay ripening and rhizopus soft rot
development of tomatoes. Crop Prot. 23, 551–554.
Wang, C.Q., 2011. Water-stress mitigation by selenium in Trifolium repens L. J. Plant
Nutr. 174, 276–282.
Wang, Y.S., Tian, S.P., Xu, Y., 2005. Effects of high oxygen concentration on pro- and
anti-oxidant enzymes in peach fruit during postharvest periods. Food Chem.
91, 99–104.
Wang, Y.F., Yu, T., Xia, J.D., Yu, D.S., Wang, J., Zheng, X.D., 2010. Biocontrol of
postharvest gray mold of cherry tomatoes with the marine yeast
Rhodosporidium paludigenum. Biol. Control 53, 178–182.
Xu, X.B., Tian, S.P., 2008. Salicylic acid alleviated pathogen-induced oxidative stress
in harvested sweet cherry fruit. Postharvest Biol. Technol. 49, 379–385.
Xue, T.L., Hartikainen, H., Piironen, V., 2001. Antioxidative and growth-promoting
effect of selenium on senescing lettuce. Plant Soil 237, 55–61.
Yao, X.Q., Chu, J.Z., Ba, C.J., 2010. Antioxidant responses of wheat seedlings to
exogenous selenium supply under enhanced ultraviolet-B. Biol. Trace Elem.
Res. 136, 96–105.
Zhu, Z., Tian, S.P., 2012. Resistant responses of tomato fruit treated with exogenous
methyl jasmonate to Botrytis cinerea infection. Sci. Hortic. 142, 38–43.