Sử dụng axit biric sau thu hoạch trên nho để cải thiẹn năng suất

1

Postharvest Application of Boric Acid on Grape to Improve Shelf-

2

life and Maintain the Quality

3
4

Hui-Jie Lia,b, Jia-Bing Jiaoa,b, Yi-Man Fanga,b, Yang-Yang Zhanga,b, Da-Long Guoa,b*

5

a

6

Luoyang 471023, P. R. China

7

b

8

Horticultural Plants, Luoyang 471023, P. R. China

9

* Corresponding author: Da-Long Guo, E-mail:

College of Horticulture and Plant Protection, Henan University of Science and Technology,

Henan Engineering Technology Research Center of Quality Regulation and Controlling of

10
11
12
13
14
15
16
17
18
19
20
21
22
1

Electronic copy available at: />

23

ABSTRACT

24

Boric acid (BA) is commercially acceptable and economically feasible material to enhance shelf-

25

life of pear, orange and other horticultural plants. Here, we investigated the effect of BA on the

26

shelf-life and postharvest quality of table grape (cv. ‘Kyoho’). Grape berries were immersed in BA

27

solution with different concentrations (0.00 [as control], 0.01, 0.03, 0.05 M) for 10 min and stored

28

at room temperature for 10 days. Compared with the control, BA treatment groups maintained

29

higher berry firmness by inhibiting the activity of polygalacturonase (PG) and cellulase, although

30

not all indexes and treatments had advantages after BA treatment. At the same time, BA treated

31

grapes maintained higher antioxidant enzyme activities such as catalase (CAT), superoxide

32

dismutase (SOD) and lower metabolic toxic products like superoxide anion (𝑂2_ ) production rate,

33

malondialdehyde (MDA) and hydrogen peroxide (H2O2) content than control. The experiment

34

results showed that postharvest application of BA effectively delay the senescence of grapes

35

compared with the control, and 0.01 M BA treatment had the most obvious effect.

36

Keywords: Kyoho, Boric acid, postharvest, antioxidant enzyme

37
38
39
40
41
42
43
44
2

Electronic copy available at: />

45

1.Introduction

46

Grape (Vitis vinifera L.) is one of the most widely consuming fruits all over the world (Li et al.,

47

2018). At present, China has become one of the world's biggest grape producers (Khan et al., 2020).

48

Over 84% of the total land used for grape production in China is cultivated for the table grapes (Sun

49

et al., 2020). However, table grapes, as non-climacteric fruits with thin pericarp and succulent flesh,

50

are easily infected by plant pathogens and exposed to serious water loss, which could result in a

51

high fungal decay rate (Meng et al., 2010). Postharvest senescence and disease have recently

52

seriously restricted the market development of table grapes. Therefore, it is necessary to explore

53

some strategies for table grapes storage.

54

In this context, different methods such as preharvest and postharvest applications of kombucha

55

(Zhou et al., 2019), short-term high CO2 (Vazquez-Hernandez et al., 2018), Aloe vera gel (Ehtesham

56

Nia et al., 2021), aerosol with calcium-based (Cherviak et al., 2021), edible coatings (1.5% chitosan

57

and 1.0% poly-ε- lysine) (Chen et al., 2019) were used to maintain the firmness and inhibit the decay

58

of fresh table grapes. In addition, chemical fungicides are widely used in vineyards to control

59

postharvest diseases of grapes (Ehtesham Nia et al., 2021). However, a large quantity of chemical

60

spraying will lead to adverse effects on consumer health and the environment. Hence, more

61

environmentally friendly and economical methods need to be explored urgently to solve these

62

problems preferably.

63

Boron was accepted as an essential nutrient for all vascular plants, animals and humans. Boron

64

regulated the metabolic activities by interacting with magnesium, calcium and vitamin D, which are

65

all necessary for bone metabolism (Devirian and Volpe, 2003). Boron has disinfectant and

66

bactericidal properties which inhibits the fruit decay after harvest and plays a crucial role in
3

Electronic copy available at: />

67

maintaining the rigidity of fruit cytoderm and phenolic concentration (Kaur et al., 2019). It was

68

reported that boron has a good effect on the prevention and control of gray mold of table grapes

69

caused by B. cinerea (Qin et al., 2010). Additionally, many studies have revealed that boric acid has

70

chemical properties inhibiting the initial increase of ethylene production (Ahmadnia et al., 2013)

71

and suppressing the activity of ACC synthase and ACC oxidase (Moon et al., 2020). It has also been

72

reported that boric acid enhances the storage life of tomatoes (Wang and Morris, 1992), retains the

73

storability and quality of pear fruits (Kaur et al., 2019), extends shelf life and quality maintenance

74

of Guava (Singh et al., 2017) and improves postharvest quality of Cut Carnation (Ahmadnia et al.,

75

2013), and so on. However, little information is available on the effect of boric acid application on

76

the postharvest quality of table grapes during storage.

77

Accordingly, the objective of the present study was to evaluate the potential of postharvest

78

treatment of boric acid to extend the shelf life of table grape during ambient temperature storage. It

79

was hypothesized that different concentrations of boric acid would enhance the storability and

80

quality of table grape (‘Kyoho’).

81

2. Materials and methods

82

2.1. Plant material and experimental treatment

83

Grape berries of ‘Kyoho’ collecting from a grape vineyard in Luoyang, Henan province, China

84

were employed in this study. Grape berries were sampled based on the uniformity in shape and

85

appearance, absence of visible defects. All grape samples were harvested after ripening and

86

analyzed at the Henan University of Science and Technology. BA solutions at different

87

concentrations (0.00, 0.01, 0.03, 0.05 M) were prepared with distilled water. The sampled grape

88

berries were divided into four sets and immersed in four BA solutions for 10 min, then placed at
4

Electronic copy available at: />

89

room temperature for 10 days. Samples were taken every 2 days for a total of 6 times, i.e., sampling

90

at 0, 2, 4, 6, 8, 10 days after the treatment, respectively. A portion of each sample used for the

91

evaluation of weight loss, berry firmness, and total soluble solid (TSS) content. The remain samples

92

were collected and stored at -40 ℃ for subsequent analysis of the physiological indicators.

93

2.2. Determination of the weight loss rate, firmness, TSS content

94
95
96
97

The weight of boric acid treated ‘Kyoho’ berries was measured at 0 storage day and 10 storage
day using an analytical balance. The weight loss rate was calculated by the following formula:

Weight loss (%) =

Initial weight ‒ final weight
× 100
Initial weight

The firmness and TSS content of berries treated with BA were measured using the durometer

98

(FT-327, Wuxi, China) and handheld refractometer (WYT-4, Shanghai, China), respectively.

99

2.3. Determination of content of ascorbic acid (AsA)

100

AsA was measured according to some modified method (Ge et al., 2015). Frozen tissue was

101

homogenized with 4.0 mL of prechilled 5% metaphosphoric acid and centrifuged at 12,000 rpm for

102

10 min at 4 ℃. The supernatant was used to measure the content of AsA. The mixture solution was

103

measured at 525 nm, and expressed as mg AsA/g FW. A standard curve with ascorbic acid was used

104

to calculate the content of AsA.

105

2.4. Determination of the superoxide anion (𝑂𝟐_ ) production rate and content of H2O2

106

The modified method (Ge et al., 2015; Yang et al., 2013) was employed to measure the 𝑂2_

107

production rate. The absorbance of the extracting solution was recorded at 530 nm. A standard curve

108

with sodium nitrite was used to calculate the 𝑂2_ production rate following the reaction equation of

109

𝑂2_ with hydroxylamine. The production rate of 𝑂2_ was expressed as nmol/min/g FW.

110

H2O2 content of the grape berries was measured spectrophotometrically after reaction with
5

Electronic copy available at: />

111

potassium iodide (Chakrabarty and Datta, 2007). The reaction mixture was measured at 390 nm, 10%

112

TCA solution was used as control. The content of H2O2 was calculated using a standard curve with

113

known concentrations of H2O2.

114

2.5. Determination of SOD activity and CAT activity

115

Superoxide dismutase (SOD) and Catalase (CAT) were extracted and assayed according to the

116

modified methods (Sun et al., 2011). Frozen grape berry tissue was extracted for 1 min with 2.0 mL

117

of 0.05 M sodium phosphate buffer (pH 7.8) containing 0.1% (w/v) polyvinyl pyrrolidone. The

118

extract solution was centrifuged for 20 min at 12,000 rpm at 4 ℃. The supernatant was collected

119

for the determination of SOD activity and CAT activity.

120

SOD activity was determined by measuring its ability to inhibit the photochemical reduction of

121

nitro blue tetrazolium (NBT). A total of 0.5 mL of enzyme solution was added into 3.0 mL of assay

122

reagent consisted of 130 mM methionine, 30 μM EDTA, 750 μM NBT, 20 mM riboflavin in 0.05

123

M sodium phosphate buffer (pH 7.8). The reaction solutions were incubated for 20 min under 4000

124

lux illumination. The absorbance of sample was spectrophotometrically measured at 560 nm and

125

0.05 M sodium phosphate buffer (pH 7.8) was used as control. The SOD activity was expressed as

126

U/g FW, where 1U is the amount of enzyme that caused 50% inhibition of NBT reduction.

127

The assay mixture for determining CAT activity consisted of 0.3 mL of 0.1 M H2O2 prepared by

128

0.05 M sodium phosphate buffer (pH 7.8) and 0.5 mL of enzyme solution. The decrease in

129

absorbance at 240 nm was recorded for 2 min at 25 ℃. And the CAT activity was expressed as U/g

130

FW/min, where 1U was defined as the amount of enzyme that caused a change of 0.01 in absorbance

131

per minute.

132

2.6. Determination of malondialdehyde (MDA) content

6

Electronic copy available at: />

133

The modified method (Ehtesham Nia et al., 2021) was employed to measure the MDA content.

134

Frozen grape berry tissue was homogenized for 1 min in 5.0 mL of 10% (w/v) trichloroacetic acid.

135

The homogenate was centrifuged for 15 min at 12,000 rpm. Three milliliter of the supernatant was

136

added to 3.0 mL of 0.67% (w/v) trichloroacetic acid. The mixture solution was heated for 20 min at

137

100 ℃, quickly cooled in an ice-bath for 10 min and then centrifuged for 15 min with12,000 rpm

138

at 4 ℃. Absorbances were measured at 532, 450 and 600 nm. MDA concentration was calculated

139

as follows: MDA content (mmol /g FW) = [6.45 (OD532 − OD600) − 0.56OD450] × 5 mL/0.5 g.

140

2.7. Determination of polygalacturonase and cellulase activity

141

PG and cellulase activity were measured using described methods (Abu-Sarra and Abu-Goukh,

142

2015). The reaction mixture containing 0.5ml crude enzyme, 2.0 ml 0.5% pectin was incubated at

143

37 ℃ for 30 min. After the constant temperature reaction, DNS was added, and the mixed solution

144

was boiled for 5 minutes. Absorbance was measured at 540nm. One pectinase activity unit was 1.0

145

mg galacturonic acid produced by pectin decomposition at 37 ℃ per gram of fresh sample per

146

minute.

147

The cellulase activity was determined by the same procedure as PG assay, but the reaction

148

temperature and time was 40 ℃, 60min, and the substrate was 1% carboxymethyl cellulose. The

149

cellulase activity unit was 1.0 mg glucose produced by the decomposition of carboxymethyl

150

cellulose at 40 ℃ per gram of fresh sample per minute.

151

2.8. Statistical analysis

152

The data presented as mean ± standard deviation (SD) from three replicates were tested using the

153

SPSS 21.0 software. Differences at P＜0.05 were considered significant. Figures were produced

154

using GraphPad Prism 9.0.
7

Electronic copy available at: />

155

3. Results

156

3.1. Effects of BA treatment on the weight loss and firmness of ‘Kyoho’ berries

157

In the 0.01 M BA treatment group, the weight loss rate was significantly lower than the control,

158

while there was no significant difference among the other groups (Fig. 1A). These data indicate that

159

the three treatments of boric acid are not all beneficial to the index of water loss.

160

The firmness of grape berries gradually decreased during storage (Fig. 1B), and it was

161

significantly higher in the 0.01 M BA treatment group than that of berries in the other BA treatment

162

and the control groups. This suggests that 0.01 M BA treatment could effectively prevent the

163

reduction in grape quality and firmness during storage (Fig. 1B).

164
165

Fig. 1. Effects of boric acid (BA) treatment on the weight loss rate(A)and firmness(B) of ‘Kyoho’

166

grape berries. The concentrations of boric acid treatment were 0.00, 0.01, 0.03 and 0.05M,

167

respectively. Vertical bars indicate mean ± standard deviation (SD). n = 3 replicates. The bars

168

followed by the same letter are not significantly different at P＜0.05.

169

3.2. Effects of BA treatment on TSS content and AsA content

170

The variation trend of TSS after BA treatment with different concentrations was consistent with

171

that of the control (Fig. 2A). Among the BA treatment groups, the 0.03 and 0.05 M BA treatment
8

Electronic copy available at: />

172

group showed the highest TSS contents (Fig. 2A).

173

The AsA content of 0.01 M BA treatment grape berries were significantly higher than the control

174

during all storage days (Fig. 2B). In addition, the content of AsA in 0.03 M BA treatment was also

175

significantly higher than the control except at the 8th storage day. (Fig. 2B).

176

177

Fig. 2. Effects of boric acid (BA) treatment on the TSS (A)and AsA content (B) of ‘Kyoho’ grape

178

berries. The concentrations of boric acid treatment were 0.00, 0.01, 0.03 and 0.05M, respectively.

179

Vertical bars indicate mean ± standard deviation (SD). n = 3 replicates. The bars followed by the

180

same letter are not significantly different at P＜0.05.

181

3.3. Effects of BA treatment on MDA content, superoxide anion (𝑂𝟐_ ) production rate and H2O2

182

content

183

Changes in the MDA content of grapes were shown in Fig. 3A. The MDA content of berries

184

increased during the first 4 days of storage, reaching the peak on day 4, and then declined from day

185

4 to day10. At the peak levels, the MDA content was the highest in the 0.03 M BA treatment group

186

and the lowest in the 0.01 M BA treatment group (Fig. 3A). During all storage times other than at

187

the 10th storage day, the MDA content in 0.01 M BA treatment was significantly lower than the

188

control. These data showed that BA inhibited the production of MDA, and it slowed the senescence

9

Electronic copy available at: />

189

rate of grape berries effectively at a concentration of 0.01 M (Fig. 3A).

190

The superoxide anion (𝑂2_ ) production rate of grape berries showed the same increase or decrease

191

trend in all the treatment groups, and the BA treatment groups was lower than the control group on

192

the whole especially in 0.01 M BA treatment group (Fig. 3B). Additionally, the 0.01 BA treatment

193

had the lowest hydrogen peroxide content in grape berries among all the treatments (Fig. 3C).

194
195

Fig. 3. Effects of boric acid (BA) treatment on MDA content (A), superoxide anion (𝑂2_ ) production

196

rate(B) and H2O2 content(C) of ‘Kyoho’ grape berries. The concentrations of boric acid treatment

197

were 0.00, 0.01, 0.03 and 0.05M, respectively. Vertical bars indicate mean ± standard deviation

198

(SD). n = 3 replicates. The bars followed by the same letter are not significantly different at P＜

199

0.05.

200

3.4. Effects of BA treatment on SOD and CAT activities

10

Electronic copy available at: />

201

SOD activity showed similar profiles among the control and BA treatment groups (Fig. 4A). The

202

activity of SOD went up and down three times but the overall activity was increasing gradually

203

during storage (Fig. 4A). The SOD activity of 0.01M BA group was higher than that of the control

204

group on the whole (Fig. 4A).

205

The CAT activity profile was similar among the different BA treatments and the control, with a

206

gradual increase from day 0 to day 2 and from day 4 to day 6, followed by a decrease from day 2 to

207

day 4 and from day6 to day 10 (Fig. 4B). The activity of CAT was significantly higher in the 0.01

208

M BA treatment group than in the control from day 0 to day 8. During the storage, the highest CAT

209

activity was detected on day6 in 0.01 M BA treated berries (Fig. 4B).

210
211

Fig. 4. Effects of boric acid (BA) treatment on SOD(A) and CAT(B) activities in ‘Kyoho’ grape

212

berries. The concentrations of boric acid treatment were 0.00, 0.01, 0.03 and 0.05M, respectively.

213

Vertical bars indicate mean ± standard deviation (SD). n = 3 replicates. The bars followed by the

214

same letter are not significantly different at P＜0.05.

215

3.5. Effects of BA treatment on PG and cellulase activities

216

The activities of PG and cellulase in 0.01 M treatment group were lower than that in control group

217

as a whole (Fig. 5). The PG activity increased gradually and then stabilized in a certain range (Fig.

11

Electronic copy available at: />

218

5A), moreover, the cellulase activity in grape berries increased firstly, then decreased slightly, and

219

significant differences were observed among various BA treatment groups (Fig. 5B).

220
221

Fig. 5. Effects of boric acid (BA) treatment on PG(A) and cellulase(B) activities in ‘Kyoho’ grape

222

berries. The concentrations of boric acid treatment were 0.00, 0.01, 0.03 and 0.05M, respectively.

223

Vertical bars indicate mean ± standard deviation (SD). n = 3 replicates. The bars followed by the

224

same letter are not significantly different at P＜0.05.

225

4. Discussion

226

It is essential to maintain the postharvest quality of grape during storage days which is mostly

227

consumed in fresh state (Jung et al., 2018). To improve the shelf life of table grapes, delaying the

228

progression of grape decay because of senescence is very important. Senescence is a complex

229

genetic programming process that is used to describe a series of events that culminate in cell death

230

at the last of a development period, including structural deterioration and macromolecule

231

degradation (Noodén et al., 1997). In table grape, senescence is closely related to reactive oxygen

232

species (ROS) accumulation (Zhang et al., 2019). The excessive production of ROS damages the

233

cells and accelerates the senescence of grapes. Plant cells have developed two main scavenging

234

mechanisms of ROS under oxidative stress which can be categorized as enzymatic system and non-

12

Electronic copy available at: />

235

enzymatic system (Shao et al., 2008). In this study, boric acid treated grapes had lower H2O2 content

236

and superoxide anion (𝑂2_ ) production rate than of the control except for individual storage days in

237

the 0.03 M treatment group (Fig. 3), indicating that boric acid treatment may control the excessive

238

production of ROS to a certain extent during storage of table grape. Meanwhile, boric acid treated

239

grape maintained significantly slightly higher CAT activity and SOD activity than control and the

240

highest CAT activity was detected on the 6th day of storage in 0.01 M BA treated berries (Fig. 4).

241

Membrane deterioration and degradation is an early and essential characteristic of signal

242

transduction pathway that occur in plant senescence (Bhattacharjee, 2005; Fan et al., 1997). In the

243

meantime, as the major cause of plant cell senescence, peroxidation of membrane lipids leads to the

244

loss of membrane integrity, physical structure, and fluidity, which further affects protein function

245

(Shewfelt and Del Rosario, 2000). Loss of membrane integrity is associated with the senescence of

246

grapes, which accompanied by the disorder of ROS, especially the high levels of H2O2 and MDA.

247

Malondialdehyde, as a toxic by-product of ROS metabolism and the end product of lipid

248

peroxidation, has been used to reflect the degree of cell membrane lipid peroxidation (Hodges et al.,

249

1999). The MDA content of 0.01and 0.05 M BA treatment groups were lower than that of the control,

250

and the MDA content of 0.01 M BA treatment group was the lowest. However, compared with the

251

control group, the 0.03 M treatment group had no significant effect on reducing the MDA content

252

(Fig. 3A). The measurement results of hydrogen peroxide content (Fig. 3C) in the samples were

253

similar to MDA, described as above-mentioned. The results showed that BA treatment significantly

254

reduced the over-production of MDA and H2O2 during storage period and inhibited lipid

255

peroxidation. This possibly is related to the previous discovery that boron helps maintain plasma

256

membrane integrity by stimulating the activity of ATPase (Ferreira et al., 2021; Ganie et al., 2013).
13

Electronic copy available at: />

257

In the previous study, some reports demonstrated that boron could control disease in grapevine

258

caused by fungus and gray mold on table grapes caused by B. cinerea (Estevez-Fregoso et al., 2021;

259

Qin et al., 2010). Additionally, Boric acid is commercially acceptable and economically feasible

260

and environmentally safe management strategy to enhance shelf life of a lot of horticultural plants.

261

In the present experiment, there was a reduction in fruit firmness with a storage period in boric acid

262

treated as well as control fruit, and the 0.01 M BA treatment group maintained the highest firmness

263

of grape berries. (Fig. 1B). This probability due to the function of boron in the synthesis of cell wall

264

composition (Liu et al., 2014; Matoh, 1997). The role of borates as antifungal complex in the control

265

of postharvest diseases in various fruits has also been demonstrated (Shi et al., 2011). In addition,

266

this may be concerned with the revelation that boric acid keeps the cell wall rigid by forming links

267

to the carboxyl groups of pectin compounds in the cell wall (O'Neill et al., 2004). This binding

268

resists cell wall deterioration enzymes, including polygalacturonase, cellulase, and inhibits the rate

269

of softening during storage. Meanwhile, BA treated grapes maintained significantly low PG activity

270

and cellulase activity (Fig. 5), which often caused fruit softening and degradation of cell wall

271

components due to depolymerization of celluloses, hemicelluloses and pectin substances which

272

decreased the thickness and rigidity following the degradation of cellulose fiber (Chen et al., 2017;

273

Ge et al., 2019). It is presumably because this element improves carbohydrate metabolism and

274

translocation, whose effect is to provide a substrate for cell respiration and cell wall synthesis. It is

275

also reported to play a role in processes such as cell capture and transport, cell wall formation, cell

276

membrane function and antioxidant defense system (Riaz et al., 2021).

277

Our results indicated that postharvest application of BA delayed the senescence process of grapes.

278

In general, 0.01 M BA treatment had the best fresh-keeping effect. First, antioxidant enzymes
14

Electronic copy available at: />

279

maintained high activity. The other is the toxic by-product of aerobic metabolism, which is kept

280

low. However, the 0.03 and 0.05 M treatment groups did not have significant advantages in all

281

indicators, especially in water loss, which may be caused by high or inappropriate concentration.

282

Consequently, senescence of grape berries during storage is moderated by BA treatment.

283

5. Conclusions

284

The present study explored the effect of boric acid on the storage performance of grape berries at

285

room temperature, and further explored the appropriate concentration of BA for postharvest

286

preservation of grape. BA alleviated postharvest senescence of grape to some extent and 0.01 M

287

BA

288

was found most effective in improving the quality of grape after harvest by maintaining higher berry

289

firmness, AsA content and moisture content. Moreover, it also resulted in lower MDA content,

290

superoxide anion (O2_ ) production rate and H2O2 content during storage. Compared with the control,

291

the activities of PG and cellulase enzymes were also lower in 0.01 M BA treated berries. Above all,

292

0.01 M BA postharvest treatment is an effective means to prolong the storage life of grape.

293

Author contributions

294

Manuscript writing: H.J.L. Experiments performance: H.J.L, J.B.J, Y.M.F, Y.Y.Z. Manuscript

295

revision and confirmation: D.L.G. Data analysis: H.J.L. All authors read and approved the final

296

manuscript.

297

Declaration of Competing Interest

298
299
300

The authors declare that there is no conflict of interest in this study.
Acknowledgments
This work was financially supported by Natural Science Foundation of China (NSFC: U1904113),
15

Electronic copy available at: />

301

National Key Research and Development Program of China (2018YFD1000105), and Program for

302

Innovative Research Team (in Science and Technology) in University of Henan Province

303

(21IRTSTHN021).

304
305

References

306

Abu-Sarra, A.F., Abu-Goukh, A.A., 2015. Changes in pectinesterase, polygalacturonase and cellulase

307

activity

308

http://10.1080/00221589.1992.11516284.

during

mango

fruit

ripening.

J.

Hortic.

Sci.

67,

561-568.

309

Ahmadnia, S., Hashemabadi, D., Sedaghathoor, S., 2013. Effects of boric acid on postharvest

310

characteristics of cut Carnation (Dianthus caryophyllus L. cv.‘Nelson’). Ann. Biol. Res. 4, 242-245.

311

Bhattacharjee, S., 2005. Reactive oxygen species and oxidative burst: Roles in stress, senescence and

312

signal transducation in plants. Curr. Sci. 1113-1121.

313

Chakrabarty, D., Datta, S.K., 2007. Micropropagation of gerbera: lipid peroxidation and antioxidant

314

enzyme activities during acclimatization process. Acta Physiol. Plant. 30, 325-331.

315

http://10.1007/s11738-007-0125-3.

316

Chen, R., Wu, P., Cao, D., Tian, H., Chen, C., Zhu, B., 2019. Edible coatings inhibit the postharvest

317

berry abscission of table grapes caused by sulfur dioxide during storage. Postharvest Biol. Technol.

318

152, 1-8. />
319

Chen, Y., Hung, Y.-C., Chen, M., Lin, H., 2017. Effects of acidic electrolyzed oxidizing water on

320

retarding cell wall degradation and delaying softening of blueberries during postharvest storage.

321

LWT 84, 650-657.

322

Devirian, T.A., Volpe, S.L., 2003. The physiological effects of dietary boron.

323

Ehtesham Nia, A., Taghipour, S., Siahmansour, S., 2021. Pre-harvest application of chitosan and

324

postharvest Aloe vera gel coating enhances quality of table grape (Vitis vinifera L. cv. 'Yaghouti')

325

during postharvest period. Food Chem. 347, 129012. http://10.1016/j.foodchem.2021.129012.

326

Estevez-Fregoso, E., Farfán-García, E.D., García-Coronel, I.H., Martínez-Herrera, E., Alatorre, A.,
16

Electronic copy available at: />

327

Scorei, R.I., Soriano-Ursúa, M.A., 2021. Effects of boron-containing compounds in the fungal

328

kingdom. J. Trace Elem. Med. Biol. 65, 126714.

329

Fan, L., Zheng, S., Wang, X., 1997. Antisense suppression of phospholipase D alpha retards abscisic

330

acid-and ethylene-promoted senescence of postharvest Arabidopsis leaves. Plant Cell 9, 2183-2196.

331

Ferreira, G.A., Hippler, F.W., Prado, L.A.d.S., Rima, J.A., Boaretto, R.M., Quaggio, J.A., Faỗanha, A.R.,

332

MattosJr, D., 2021. Boron modulates the plasma membrane H+‐ATPase activity affecting nutrient

333

uptake of Citrus trees. Ann. Appl. Biol. 178, 293-303.

334

Ganie, M.A., Akhter, F., Bhat, M., Malik, A., Junaid, J.M., Shah, M.A., Bhat, A.H., Bhat, T.A., 2013.

335

Boron—a critical nutrient element for plant growth and productivity with reference to temperate

336

fruits. Curr. Sci. 76-85.

337

Ge, Y., Deng, H., Bi, Y., Li, C., Liu, Y., Dong, B., 2015. Postharvest ASM dipping and DPI pre-treatment

338

regulated reactive oxygen species metabolism in muskmelon (Cucumis melo L.) fruit. Postharvest

339

Biol. Technol. 99, 160-167. http://10.1016/j.postharvbio.2014.09.001.

340

Ge, Y., Duan, B., Li, C., Wei, M., Chen, Y., Li, X., Tang, Q., 2019. Application of sodium silicate retards

341

apple softening by suppressing the activity of enzymes related to cell wall degradation. J. Sci. Food

342

Agric. 99, 1828-1833.

343

Hodges, D.M., DeLong, J.M., Forney, C.F., Prange, R.K., 1999. Improving the thiobarbituric acid-

344

reactive-substances assay for estimating lipid peroxidation in plant tissues containing anthocyanin

345

and other interfering compounds. Planta 207, 604-611.

346
347
348

Jung, H.M., Lee, S., Lee, W.-H., Cho, B.-K., Lee, S.H., 2018. Effect of vibration stress on quality of
packaged grapes during transportation. Eng. Agric. Environ. Food 11, 79-83.
Kaur, A., Gill, P.P.S., Jawandha, S.K., Singh, M., 2019. Pre-storage exogenous application of boric acid

349

extends

350

http://10.1016/j.scienta.2019.108616.

351
352
353

storability

and

maintains

quality

of

pear

fruits.

Sci.

Hortic.

256.

Khan, N., Fahad, S., Naushad, M., Faisal, S., 2020. Grape Production Critical Review in the World.
SSRN Electronic Journal. http://10.2139/ssrn.3595842.
Li, X., Dong, S., Su, X., 2018. Copper and other heavy metals in grapes: a pilot study tracing influential
17

Electronic copy available at: />

354

factors and evaluating potential risks in China. Sci. Rep. 8. http://10.1038/s41598-018-34767-z.

355

Liu, G., Dong, X., Liu, L., Wu, L., Jiang, C., 2014. Boron deficiency is correlated with changes in cell

356

wall structure that lead to growth defects in the leaves of navel orange plants. Sci. Hortic. 176, 54-

357

62.

358

Matoh, T., 1997. Boron in plant cell walls. Plant Soil 193, 59-70.

359

Meng, X.-H., Qin, G.-Z., Tian, S.-P., 2010. Influences of preharvest spraying Cryptococcus laurentii

360

combined with postharvest chitosan coating on postharvest diseases and quality of table grapes in

361

storage. LWT 43, 596-601. http://10.1016/j.lwt.2009.10.007.

362

Moon, J., Park, Y.J., Son, S.-H., Roh, J., Youn, J.H., Kim, S.Y., Kim, S.-K., 2020. Brassinosteroids

363

signaling via BZR1 down-regulates expression of ACC oxidase 4 to control growth of Arabidopsis

364

thaliana seedlings. Plant Signal. Behav. 15, 1734333.

365

Noodén, L.D., Guiamét, J.J., John, I., 1997. Senescence mechanisms. Physiol Plant 101, 746-753.

366

O'Neill, M.A., Ishii, T., Albersheim, P., Darvill, A.G., 2004. Rhamnogalacturonan II: structure and

367

function of a borate cross-linked cell wall pectic polysaccharide. Annu. Rev. Plant Biol. 55, 109-

368

139.

369
370

Qin, G., Zong, Y., Chen, Q., Hua, D., Tian, S., 2010. Inhibitory effect of boron against Botrytis cinerea
on table grapes and its possible mechanisms of action. Int. J. Food Microbiol. 138, 145-150.

371

Riaz, M., Kamran, M., El-Esawi, M.A., Hussain, S., Wang, X., 2021. Boron-toxicity induced changes in

372

cell wall components, boron forms, and antioxidant defense system in rice seedlings. Ecotoxicol.

373

Environ. Saf. 216, 112192.

374
375
376
377
378
379
380

Shao, H.-B., Chu, L.-Y., Lu, Z.-H., Kang, C.-M., 2008. Primary antioxidant free radical scavenging and
redox signaling pathways in higher plant cells. Int. J. Biol. Sci. 4, 8.

Shewfelt, R., Del Rosario, B., 2000. The role of lipid peroxidation in storage disorders of fresh fruits and
vegetables. Hortic. Sci. 35, 575-579.
Shi, X.-Q., Li, B.-Q., Qin, G.-Z., Tian, S.-P., 2011. Antifungal activity and possible mode of action of
borate against Colletotrichum gloeosporioides on mango. Plant Dis. 95, 63-69.
Singh, J., Prasad, N., Singh, S.K., 2017. Postharvest Treatment of Guava (Psidium guajava L.) Fruits
18

Electronic copy available at: />

381

with Boric Acid and NAA for Quality Regulation during Ambient Storage. Int. j. bio-resour. stress

382

manag. 8, 201-206. http://10.23910/IJBSM/2017.8.2.1633a.

383

Sun, J., You, X., Li, L., Peng, H., Su, W., Li, C., He, Q., Liao, F., 2011. Effects of a phospholipase D

384

inhibitor on postharvest enzymatic browning and oxidative stress of litchi fruit. Postharvest Biol.

385

Technol. 62, 288-294. http://10.1016/j.postharvbio.2011.07.001.

386

Sun, Y.-D., Guo, D.-L., Yang, S.-D., Zhang, H.-C., Wang, L.-L., Min, L., Yu, Y.-H., 2020. Melatonin

387

treatment improves the shelf-life and postharvest quality of table grape (Vitis labrusca L. cv.

388

‘Fengzao’). J. Berry Res. 10, 665-676. http://10.3233/jbr-200569.

389

Vazquez-Hernandez, M., Navarro, S., Sanchez-Ballesta, M.T., Merodio, C., Escribano, M.I., 2018.

390

Short-term high CO2 treatment reduces water loss and decay by modulating defense proteins and

391

organic osmolytes in Cardinal table grape after cold storage and shelf-life. Sci. Hortic. 234, 27-35.

392

/>
393
394

Wang, S., Morris, S., 1992. Effects of borax and guazatine on the ripening and postharvest diseases of

tomato (cv. Flora-Dade). Acta Hortic. 343, 331-333.

395

Yang, Q., Rao, J., Yi, S., Meng, K., Wu, J., Hou, Y., 2013. Antioxidant enzyme activity and chilling

396

injury during low-temperature storage of Kiwifruit cv. Hongyang exposed to gradual postharvest

397

cooling. Hortic Environ Biotechnol 53, 505-512. http://10.1007/s13580-012-0101-8.

398
399

Zhang, Z., Xu, J., Chen, Y., Wei, J., Wu, B., 2019. Nitric oxide treatment maintains postharvest quality
of table grapes by mitigation of oxidative damage. Postharvest Biol. Technol. 152, 9-18.

400

Zhou, X., Tan, J., Gou, Y., Liao, Y., Xu, F., Li, G., Cao, J., Yao, J., Ye, J., Tang, N., Chen, Z., 2019. The

401

biocontrol of postharvest decay of table grape by the application of kombucha during cold storage.

402

Sci. Hortic. 253, 134-139. />
403

19

Electronic copy available at: />