horticulturae

Review
Advances in the Strategic Approaches of Pre- and Post-Harvest
Treatment Technologies for Peach Fruits (Prunus persica)
Jin Song Shin 1 , Han Sol Park 1 , Ki Won Lee 2 , Ji Seop Song 2 , Hea Yeon Han 2 , Hye Won Kim 3
and Tae Jin Cho 1,2, *

1 Department of Food Regulatory Science, College of Science and Technology, Korea University, 2511, Sejong-ro,
Sejong 30019, Republic of Korea
2 Department of Food and Biotechnology, College of Science and Technology, Korea University, 2511, Sejong-ro,
Sejong 30019, Republic of Korea
3 Department of Animal Sciences, University of Illinois at Urbana-Champaign, Urbana, IL 61801, USA
* Correspondence: microcho@korea.ac.kr; Tel.: +82-44-860-1433

Abstract: Peach (Prunus persica) is one of the representative climacteric fruits susceptible to envi-
ronmental stresses, including microbial contamination. This article analyzed major findings from
the literature on pre- and post-harvest technologies for maintaining the quality of peach fruit to
figure out the strengths and limitations of each treatment strategy. The key implication from studies
of pre-harvest agents directly applied to the fruit surface or supplemented as fertilizer was the
application of a mixture regarding substances with diverse working mechanisms to prevent excessive
use of the agent. The common objectives of previous research on pre-harvest treatments were not
only the improvement in the quality of harvested fruit but also the storability during long-term refrig-
eration due to the short lifespan of peaches. In the case of post-harvest treatments, the efficacy was
considerably affected by various determinant factors (e.g., a cultivar of fruit, the sort of technologies,
and storage environments), and thus operating conditions optimized for peach fruit were described
in this article. Whereas, although the combined treatment of technologies categorized into principles
(physical, chemical, and biological approaches) has been adopted to achieve the synergistic effect,
Citation: Shin, J.S.; Park, H.S.; Lee,
undesirable antagonistic effects (i.e., the inhibition of efficacies expectable from singular treatments)
K.W.; Song, J.S.; Han, H.Y.; Kim,
were also reported to highlight the importance for exploring adequate treatment conditions.
H.W.; Cho, T.J. Advances in the
Strategic Approaches of Pre- and
Post-Harvest Treatment Technologies
Keywords: fruit quality; productivity; fungal infection; long-term fruit storage; fruit ripening;
for Peach Fruits (Prunus persica). microbial safety; climacteric fruit; stone fruit; combined treatment
Horticulturae 2023, 9, 315.
https://doi.org/10.3390/
horticulturae9030315
1. Introduction
Academic Editors: Brian Farneti and
Custódia Maria Luís Gago Peach (Prunus persica) is a globally consumed fruit preferred by consumers due to its
exotic taste and rich nutritional substances (e.g., minerals, sugars, and amino acids) [1].
Received: 12 January 2023 Peaches have several antioxidant compounds, including vitamins, phenolic compounds,
Revised: 23 February 2023
volatiles, carotenoids, and organic acids [2]. However, as a climacteric fruit, peaches are
Accepted: 24 February 2023
susceptible to rot and have a short shelf life due to ethylene emission accompanied by
Published: 28 February 2023
a rising respiration rate during storage [3]. Within the Prunus genus, peach is a stone
fruit that has a thin exocarp or skin with the characteristics of a lignified endocarp and
a fleshy mesocarp [4]. Although the production of peaches and nectarines is steadily
Copyright: © 2023 by the authors.
increasing (Figure 1), the problems in storage and transportation due to their rapid rotting
Licensee MDPI, Basel, Switzerland. and softening at ambient temperature make their exports difficult [5]. Previous studies
This article is an open access article regarding the quality of peaches have been conducted with similar objectives to those
distributed under the terms and of other stone fruits (e.g., cherry, plum, and apricot) [6–8]. However, research focused
conditions of the Creative Commons specifically on peaches is needed because peaches age more rapidly and are more vulnerable
Attribution (CC BY) license (https:// to disease caused by pathogens than other stone fruits [9].
creativecommons.org/licenses/by/ To enhance the quality and extend the storage period of fruits, both pre-harvest
4.0/). and post-harvest treatment technologies have been consistently developed and applied

Horticulturae 2023, 9, 315. https://doi.org/10.3390/horticulturae9030315 https://www.mdpi.com/journal/horticulturae

 peaches is needed because peaches age more rapidly and are more vulnerable to disease
caused by pathogens than other stone fruits [9].
Horticulturae 2023, 9, 315 To enhance the quality and extend the storage period of fruits, both pre-harvest and 2 of 41
post-harvest treatment technologies have been consistently developed and applied from
production to storage [10,11]. Pre-harvest treatment is a method performed before har-
vesting to improvetothe
from production quality
storage of harvested
[10,11]. Pre-harvest fruitstreatment
and/or extend their shelf
is a method life during
performed before
post-harvest storage [12,13]. Post-harvest treatment prevents
harvesting to improve the quality of harvested fruits and/or extend their shelf life the deterioration of during
fruit
quality until consumption
post-harvest storage [12,13]. by consumers
Post-harvest [14,15]. Various
treatment types ofthe
prevents pre- and post-harvest
deterioration of fruit
technologies
quality until have been reported
consumption with differing
by consumers effects,
[14,15]. and even
Various types the
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Previous
relevant review
research articles regarding
conducted to determinepeaches havetreatment
desirable focused on determinant factors of
conditions.
Previous review
fruit characteristics andarticles
specificregarding peaches have
pre- or post-harvest focused on
technology as determinant factors of
follows: pre-harvest
fruit characteristics
conditions linked to the and specific
quality of pre- or post-harvest
peaches [16], the treatmenttechnology as follows:
of salicylic acid topre-harvest
enhance
conditions linked
post-harvest to the
storability of quality
peachesof[17],
peaches [16], the
the effects treatmentapplication
of nitrogen of salicylicon acid
thetointernal
enhance
quality of peaches [18], and the use of oligosaccharides to improve fruit preservationinternal
post-harvest storability of peaches [17], the effects of nitrogen application on the after
quality[19].
harvest of peaches
However, [18],a and the use of oligosaccharides
comprehensive to improve
analysis of the literature onfruit preservation
experimental after
condi-
harvest
tions and [19]. However,
the results a comprehensive
of various analysis of the
pre- and post-harvest literature
treatments on experimental
optimally designedcondi- for
tionsfruits
peach and the results to
according of the
various pre- and post-harvest
categorization of the principles treatments optimally
of technologies designed
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peach fruits
chemical, and according
biological to the categorization
approaches) and theof the principles
application of technologies
strategies (i.e.,or
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been reported. application
This strategies
review article (i.e., singular
suggests a strategic or ap-
com-
bined treatments) has rarely been reported. This review article
proach for the pre- and post-treatment of peaches based on relevant research with the suggests a strategic approach
for the pre-
following and(1)
topics: post-treatment
the summaryofofpeaches based on
major quality relevant
factors research
selected with the following
for evaluating values
of treatment technologies; (2) the distinct implications of studies on physical,ofchemical,
topics: (1) the summary of major quality factors selected for evaluating values treatment
technologies;
and biological (2) the distinct
treatment implications
technologies; (3) of studies on physical,
dose‒response chemical,condition-de-
(i.e., treatment and biological
pendent efficacies) of technologies to estimate the optimal operational methods;efficacies)
treatment technologies; (3) dose-response (i.e., treatment condition-dependent and (4)
ofadvantages
the technologies to disadvantages
and estimate the optimal operational
of the combined methods;compared
treatment and (4) the to advantages
singular treat- and
disadvantages
ments. of the combined treatment compared to singular treatments.

Figure
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using the data from FAOSTAT [20]).

2. Quality Factors of the Pre- and Post-Harvest Treatment of Peach Fruits

2. Quality Factors of the Pre- and Post-Harvest Treatment of Peach Fruits
The characteristics of fruit include sensory properties (e.g., texture, taste, aroma, and
The characteristics of fruit include sensory properties (e.g., texture, taste, aroma, and
appearance), chemical constituents, functional values, safety factors (e.g., the concentra-
appearance), chemical constituents, functional values, safety factors (e.g., the concentration
of toxigenic substances, and population level of contaminated microorganisms), nutri-
tional value, mechanical properties, and defects caused by the growth or metabolism of
pathogens [21]. Major quality factors of peaches investigated from previous research re-
garding pre- and post-harvest treatments can be categorized as the stability of fruit quality,
microbial deterioration, and antioxidant capacity. As shown in Table 1, the quality factors
can be estimated from the quantitatively measured values by the indicators related to taste,
Horticulturae 2023, 9, 315 3 of 41

color, and nutrients. Examples of the measurable parameters are as follows: firmness,
weight loss, volume, total soluble solid (TSS), titratable acidity (TA), ethylene production,
vitamin C content, activation of enzymes related to antioxidant ability, antioxidant content,
decay incidence, and antimicrobial function. Taste, one of the sensory characteristics, is
mainly divided into a sweet and sour flavor that can be indicated by the values of TSS and
TA, respectively.

Table 1. Quality factors as the indicator of the effects of pre- and/or post-harvest treatments for peach
fruits.

Category Examples of the Factors

Weight, volume, length, width or diameter, total soluble solid (TSS), soluble
solid content (SSC), titratable acidity (TA), ethylene emission (production),
Stability of fruit quality
malondialdehyde (MDA) content, firmness, color, vitamin C content, pectin
content
Microbial deterioration and damage Infected wounds, decay, disease incidence, lesion diameter
Antioxidants content (phenolics, flavonoids), activity of enzymes (catalase
(CAT), superoxide dismutase (SOD), ascorbate peroxidase (APX), peroxidase
Antioxidant capacity (POD), pectin methyl esterase (PME), phenylalanine ammonia-lyase (PAL),
enzymes of ascorbate-glutathione (AsA-GSH) cycle, polyphenol oxidase
(PPO), lipoxygenase (LOX))

The stability of fruit quality allows for the maintenance of the product value of peaches
from cultivation to harvest followed by storage, and various factors correlated with the
changes in physiological characteristics of products have been identified. Quantitative
values of morphological characteristics (e.g., volume, length, and width or diameter) can
belong to quality factors. One of the major goals of the pre-harvest treatment of fruits is
the modulation of metabolisms related to fruit growth and development [22,23]. Whereas,
preventing a decrease in weight during fruit storage under refrigerated temperatures due
to the loss of water and the response to cold stress is an additional intended function of
both pre- and post-harvest treatments [24]. Since an increase in sugar content is closely
related to the ripening of peach fruits, TSS can also be an indicator of the product quality
of fruit [25]. The measurement of soluble solid content (SSC) is an index of the flavor of
peaches to determine the appropriate time for harvest and storage from the perspective of
fruit maturation [26]. TA, determined by the titration of internal acid, is a measurement of
total acid, which can be used to analyze the growth level and taste of fruits [27]. The ratio
of SSC to TA (SSC/TA) is negatively related to the maturity of the fruit and is also used as
the representative ripening index [28]. Peach produces ethylene inducing fruit ripening,
and the level of ethylene emission or production can be a parameter of maturity [29].
Malondialdehyde (MDA) produced by reactive oxygen species (ROS) is known as an
indicator of fruit damage [30]. The color of peach skin and flesh is generally considered a
factor in the stability and value of commodities, but is also related to antioxidant capacity
because of phytochemicals [31].
The incidence of fungal decay is an important index for shelf life and commercial
value because damage to peach fruits is critical from the perspectives of economic loss
for producers and retailers. The major fungi reported as the cause of deterioration from
peach fruits are brown rot fungi [32], Rhizopus rot fungi [33], blue mold [34], and gray
mold [35]. Brown rot is commonly caused by fungal species including Monilinia fructicola,
Monilinia laxa, and Monilinia fructigena [36]. Rhizopus rot, blue mold disease, and gray
mold disease are generally caused by the fungal species of Rhizopus stolonifer, Penicillium
expansum, and Botrytis cinerea, respectively [37]. The level of fungal infection has been
generally measured by the visual observation of indicators for the severity of fruit diseases
(e.g., infected wounds, decay, disease incidence, and lesion diameter).
Antioxidant capacity is the ability to remove ROS, which is a cause of undesirable
quality changes in fruits due to weakened stress tolerance and can be evaluated by factors
including the contents of phenolics and/or flavonoid compounds [38]. The activity of
major enzymes involved in the production or removal of ROS in peach fruits can also
Horticulturae 2023, 9, 315 4 of 41

be reported as indicators of the antioxidant capacity as follows: catalase (CAT) [39], su-
peroxide dismutase (SOD) [40], ascorbate peroxidase (APX) [41], peroxidase (POD) [42],
pectin methyl esterase (PME) [43], phenylalanine ammonia-lyase (PAL) [44], enzymes that
comprise the ascorbate-glutathione (AsA-GSH) cycle [45], polyphenol oxidase (PPO) [46],
and lipoxygenase (LOX) [47].

3. Strategic Approach to the Application of Pre-Harvest Treatment Technologies for

Peach Fruits
Pre-harvest treatments for general fruits have been used as measures for enhancing
the quality and safety of fruits with the treatment of chemical agents (e.g., spraying and
spreading, fertilizing) or physical treatments (e.g., bagging fruits, setting a canopy on the
fruit tree, and irrigating the field of the orchard) before harvest [16]. Most relevant research
using pre-harvest peach fruits reported the application of chemical substances to improve
the product quality at harvest and to prevent the deterioration of harvested fruits during
storage [48–57]. This indicates that the effects expected from the post-harvest treatment can
also be achieved by pre-harvest technologies. Thus, as shown in Table 2, this review mainly
analyzed the previous research regarding the evaluation of the efficacies of pre-harvest treat-
ment technologies with the perspectives of not only harvesting but also storage time [58].
Key chemical agents that can be utilized as pre-harvest treatments for peaches by spreading
and/or spraying are the following: calcium salts [48–52], acids [53–55], sodium nitroprus-
side (SNP) [56], and putrescine (PUT) [57]. The use of fertilizer has also been regarded as
an effective method to control the quality of peach fruit during cultivation [59,60].

3.1. Spraying and Spreading Chemical Agents to Fruits

3.1.1. Calcium Salts
Spraying salts is a popular pre-treatment for fruits, and salt composed of calcium
ions is a representative agent used to treat peaches before harvesting [48,50,52]. Calcium
treatment functions in various ways, such as delaying the ageing of fruits by increasing
calcium concentrations in the epidermis or slowing hydrolysis by strengthening cell walls
to prevent external intrusion [61,62]. Although calcium treatment cannot be an alternative
to disinfectants, it increases the resistance of fruit against fungal infection by strengthening
cell walls. The composition of calcium salts (e.g., calcium chloride and calcium nitrate) and
working concentrations have been reported as important factors that determine effective-
ness in the pre-harvest process [48]. The results vary depending on the concentration of
the substance, and higher concentrations are generally more effective: in particular, values
representing the morphological characteristics (e.g., weight and size) of peaches increase as
the content of calcium gradually increases [49,50]. To improve the pre-harvest treatment
effects using calcium salts, a combined treatment strategy has generally been adopted to
achieve additional or synergistic effects rather than the use of a single compound. Improve-
ment of the stability of fruit quality during the post-harvest storage of peach fruits treated
with the agent complex organized with calcium as a basis material (i.e., combining calcium
salts with other substances to make the pre-harvest treatment agent: potassium silicate
(K2 SiO3 ) + Ca2+ EDTA, chitosan + calcium chloride (CaCl2 ), magnesium and titanium +
calcium, and potassium sulfate (K2 SO4 ) + CaCl2 ) has been reported. Agents that can
improve the stress-modulating capacity of peach fruit can also be combined with calcium
salts. Aziz et al. [49] showed the effects of Ca2+ EDTA with K2 SiO3 on the growth and
storability of fruit quality from Ca2+ EDTA with K2 SiO3 , which support the enzymatic ac-
tivity encouraging the antioxidant defense mechanism. El-Badawy [50] combined calcium
salts with chitosan as the coating material available for the stimulation of plant immune
systems supporting fruit growth and protection against microbial attack, and the highest
treatment concentration showed the greatest effect. Supplementation of calcium with other
activating nutrients (e.g., magnesium and titanium) related to the growth of fruits helps
maintain the quality of peaches during storage [51]. Although the combination of various
pre-treatment agents is generally expected to show better effects than a singular agent,
Horticulturae 2023, 9, 315 5 of 41

agents can have antagonistic effects (i.e., inhibition of the efficacies from agents when used
in combination). El-Dengawy et al. [52] reported that the decrease in TSS and anthocyanin
with CaCl2 treatment can be complemented by the addition of K2 SO4 ; however, this study
also showed that combined treatment (CaCl2 + K2 SO4 ) resulted in lower firmness than
the individual treatment of CaCl2 . Therefore, the validation of the presence of undesirable
interactions of agents in the pre-treatment complex should be regarded as a prerequisite for
the application of combination technology to design appropriate operational conditions.

3.1.2. Acids
Salicylic acid is an organic acid with antimicrobial effects and is regarded as one of
the key phenolics naturally present in plants to regulate the growth and stress responses
of fruits [63,64]. Erogul and Özsoydan [53] compared the effects of 1 mM and 2 mM
salicylic acid solutions spread on peaches at the 23rd and 15th days before harvest on the
quality changes during cold storage (2 ◦ C, 8 days), followed by exposure to a simulated
environment for shelf life (20 ◦ C, 2 days): the results showed the smallest quality changes
(e.g., weight loss, reduction in flesh firmness, and reduction in acidity) with the higher
concentration (2 mM) [53].
Oxalic acid, universally present in plants, acts as a natural antioxidant, inducing
an increase in membrane integrity and a delay in fruit ripening by controlling enzymes
associated with quality changes of peaches [65,66]. Razavi and Hajilou [55] showed dose-
dependent enhancement of the antioxidant capacity of the fruit surface treated with oxalic
acid (up to 5 mM treatment concentration) 15 days before harvest to elicit an increase in the
antioxidant content (phenolics and flavonoids) and the activities of antioxidant enzymes as
the key mechanism for the extension of shelf life. Although pre-harvest oxalic acid spraying
of peaches with higher concentrations can be expected to improve efficacy, it is necessary to
consider the problematic health risk factors in the case of overtreatment due to the potential
toxicity of oxalic acid [67].
Gibberellic acid is a diterpenoid carboxylic acid that can function as a plant growth
hormone to extend the period of fruit availability by controlling the ability of the fruit to
respond to abiotic stress factors and to support plant development during ripening [68].
Pegoraro et al. [54] emphasized the beneficial effects of spraying gibberellic acid on peach
fruits at the beginning of pit hardening as a pre-harvest treatment in regard to woolliness
(i.e., the prevention of woolliness during long-term post-harvest cold storage) and quality
factors (i.e., higher values of fruit size and mass); although a delay in ripening did not
occur. Gene expression (mRNA abundance) analysis performed to understand the mecha-
nism of the effects of spraying gibberellic acid also showed the metabolism of beneficial
functionalities of cell wall structure, intracellular transport, heat shock proteins associated
with the homeostasis of the metabolism of peach ripening, and ethylene biosynthesis [54].

3.1.3. Other Chemical Agents for Pre-Harvest Treatment

Sodium nitroprusside (SNP), as a pre- or post-harvest treatment agent for fruits, has
been used as a donor of nitric oxide (NO), which contributes to the enhancement of disease
resistance and the suppression of ethylene production for fruit quality control [69]. The
modulation of fruit metabolic pathways related to antioxidant and antifungal metabolites
by SNP can also result in the maintenance of fruit quality through the suppression of ROS
formation and the prevention of pathogenic fungal decay, respectively [70,71]. The research
from Saba and Moradi [56] showed the efficacy of pre-harvest spraying of SNP on peach
fruits (25 and 50 µmol/L, 14 days before harvest) in regards to chilling tolerance, which
results in an improvement of the stability of fruit quality during cold storage, with the
suppression of ethylene emission and the activation of antioxidative enzymes being the key
mechanisms. However, excessive treatment of SNP can exhibit toxic effects on peach fruits
(e.g., increase in weight loss, decrease in firmness, accumulation of reactive nitrogen oxide
species, decrease in antioxidant activity, and acceleration of peroxidation), highlighting the
importance of investigating adequate treatment concentrations [56].
Horticulturae 2023, 9, 315 6 of 41

Putrescine (PUT) is a polyamine involved in the physiological activities of fruits (e.g.,

growth, softening, senescence, maturation, and regulation of the stress response) [72]. The
pre-storage exogenous application of PUT is expected to improve the shelf life and the
stability of fruit quality during cold storage by increasing endogenous polyamine levels
and inducing the acclimation of fruits [73]. Abbasi et al. [57] revealed the substantial
alleviation of chilling injury in peach fruit with desirable effects on quality factors (e.g.,
weight, firmness, ethylene production, and fruit skin color) during low-temperature storage
with the exogenous spray of PUT (1–3 mM) before harvest, but the greatest effect was
obtained from the 2 mM PUT treatment; thus, finding the optimal treatment concentration
should be regarded as a prerequisite for actual application to fields [57].

3.2. The Use of Fertilizers

Irrational fertilization can cause problems including soil acidification, salinization,
compaction, and a lack of fruit moisture. To prevent indiscriminate abuse of fertilizers,
the determination of a desirable formulation of ingredients and fertilization period is es-
sential [59,60]. Xiao et al. [59] showed the impact of formulated fertilization (containing
urea, organic fertilizer, potassium sulfate, mono ammonium phosphate, and superphos-
phate) applied to a peach orchard on the physical and chemical characteristics of soil as
the increase in the organic matter with the decrease of nitrogen content, highlighting the
potential of fertilizers to significantly improve fruit quality. Liang et al. [60] practically
demonstrated that the treatment of sprouting fertilizer (i.e., 10–15 days before flowering)
and pre-harvest fertilizer (i.e., 15 days before harvest) could increase the quality factors
(e.g., weight, firmness, and TSS) of peach fruits.
Horticulturae 2023, 9, 315 7 of 41

Table 2. Pre-harvest treatments on peach fruit.

Method Cultivar Method and Treatment Conditions Results and Implications Reference

- Calcium concentration of control

* Dry weight: 59 mg/100 g
Foliar spray of salts six times with 2-week - Treatment conditions
- Increase of calcium concentrations in epidermis
intervals before harvest followed by cold Scarlett O’Hara * Calcium chloride, 4.7–17.9 g/tree - Decrease of brown rot infections [48]
storage after harvest * Calcium nitrate, 9.6 g/tree
* Calcium glycine chelate, 10.4 g/tree
- Storage: 2–4 ◦ C, 3 weeks

- Improvement in the stability of fruit quality

* Higher physical characteristics (weight, volume,
- Control: tap water length, diameter) and lower weight loss
- Treatment conditions * Higher SSC 2 and TA 3
Foliar spray of salts three times with 15-day * Decrease of vitamin C contents
intervals before harvest followed by cold Dessert Red * Combined treatment of Potassium silicate 0.1–0.3% + [49]
* Increase of anthocyanin content
storage after harvest Ca2+ EDTA 0.1–0.3%
- Combined treatment with the highest concentrations of each
- Storage: 0 ± 1 ◦ C, 90–95% RH 1 , 28 days agent showed the highest effects.
- Beneficial effects of pre-harvest treatment can be observed at
both the harvesting date and during storage

- Single treatment
- Improvement in the stability of fruit quality
* Chitosan 0.5–1.0%
* Calcium chloride 2–4% - Lower weight loss, TSS 4
Foliar spray of salts three times with 2-week
intervals before harvest followed by cold Florida Prince - Combined treatment * Higher firmness and TA 3 [50]
storage after harvest - Combined treatment with the highest concentrations of each
* Chitosan 0.5–1.0% + calcium chloride 2–4%
agent showed the highest effects
- Storage: 0 ± 2 ◦ C, 90–95% RH 1 , 35 days

- Composition of spray solution

- Effects on fruits at harvest time
* Calcium 0.1 mM (4 mg/L) + Magnesium 0.103 mM
* Quality improvement: higher firmness and weight
Foliar spray of calcium supplemented with (2.5 mg/L) + Titanium 0.042 mM (2 mg/L)
other activating nutrients 10 days after - Effects on fruits during cold storage
Sevilla 2 - Treatment condition [51]
anthesis followed by cold storage after * Improvement in the stability of fruit quality: lower
harvest * Three times, 5 L per tree
weight loss, increase of ripening index (TSS 4 /TA 3
- Storage: 2 ± 0.5 ◦ C, 90% RH 1 , 28 days ratio) and ethylene production, higher firmness
- Post-storage: 20 ◦ C, 5 days
Horticulturae 2023, 9, 315 8 of 41

Table 2. Cont.

Method Cultivar Method and Treatment Conditions Results and Implications Reference

- Improvement in the stability of fruit quality

- Control: tap water
- Single treatment * Lower weight loss, TSS 4 /acid value
* Potassium sulphate (K2 SO4 ) 1.5% * Higher firmness, lower decay incidence
* Calcium chloride (CaCl2 ) 2% * Higher pectin and vitamin C content
Foliar spray of salts 6 weeks after blooming - Improvement of nutritional quality
Medium Sultani - Combined treatment [52]
followed by cold storage after harvest
* Potassium sulphate (K2 SO4 ) 1.5% + Calcium chloride * Higher anthocyanin
(CaCl2 ) 2% - Combined treatment showed diversity in the increase or the
- Storage: 5 ± 1 ◦ C, 80–85% RH 1 , 24 days decrease of the effects for the quality factors compared with
the effects by singular treatments

- Improvement in the stability of fruit quality

* Increase of weight, width, and length of fruit
- Improvement of fruit taste factors
- Treatment conditions
* Increase of TA 3 (whereas no changes in TSS 4 )
Spreading of the salicylic acid solution 23 and * Salicylic acid 1–2 mM * Higher firmness
15 days before harvest followed by cold Cresthaven [53]
storage - Storage: 2 ◦ C, 85–90% RH 1 , 8 days - Improvement of antioxidant activity
- Post-storage: 20 ◦ C, 2 days
* Increase of phenol content
- Effects on fruits during cold storage
* Decrease in color changes of fruit flesh

- Improvement of antioxidant activity

- Treatment conditions * Increase of phenol and flavonoids contents
* Oxalic acid 1–5 mmol/L * Activation of catalase (CAT), peroxidase (POD), and
Spray of oxalic acid on fruit surface 15 days
Anjiry maleki superoxide dismutase (SOD) [55]
before harvest followed by cold storage - Storage: 1 ◦ C, 90% RH 1 , 28 days
- Improvement of the stability of the quality
- Post-storage: 20 ◦ C, 24 h
* Higher firmness

- No effect to delay the fruit ripening

* Ripening without differences in color, firmness of
flesh, TSS 4 , or ethylene production
- Treatment conditions - Improvement in the stability of fruit quality
* Gibberellic acid solution 400 L/ha (gibberellic acid 50 * Increase of fruit size and mass from the treatment at
Spray of gibberellic acid before harvest (at the mg/L + surfactant Silwet® 0.05% (v/v; pH 4.5) at the the beginning of pit hardening stage (whereas,
beginning or the end of pit hardening) Chiripa beginning or the end of the pit hardening stage treatment at the end of pit hardening stage did not [54]
followed by cold storage show differences)
- Storage: 1 ± 1 ◦ C, 92 ± 5% RH 1 , 30 days
- Decrease of the woolliness
- Post-storage: 23 ± 3 ◦ C, 75 ± 5% RH 1 , 2 days
* The prevention of woolliness from the treatment at
the beginning of pit hardening stage (whereas,
treatment at the end of pit hardening stage did not
show differences)
Horticulturae 2023, 9, 315 9 of 41

Table 2. Cont.

Method Cultivar Method and Treatment Conditions Results and Implications Reference

- Improvement in the stability of fruit quality

* Lower weight loss
* Decrease of ethylene production
* Increase of firmness
- Treatment conditions - Alleviation of CI 5
Spray of sodium nitroprusside (SNP) 14 days * SNP 25–100 µmol/L * Decrease of CI 5 index
GH Hill [56]
before harvest followed by cold storage - Storage: 4 ◦ C, 1
80–90% RH , 28 days - All beneficial effects were not observed from samples treated
- Post-storage: 20 ◦ C, 1–4 days with an excess level of SNP (100 µmol/L)
* Increase of weight loss, increase of ethylene
production, decrease of firmness, increase of CI5
index, accumulation of ROS 6 , decrease of the
antioxidant activity, acceleration of peroxidation

- Improvement in the stability of fruit quality

* Lower weight loss
* Increase of firmness
* Decrease of ethylene production
- Treatment conditions * Lower SSC 2 /TA 3 ratio
Spray of putrescine (PUT) three times before * PUT 1–3 mM - Maintenance of fruit skin color
Flordaking [57]
harvest followed by cold storage - Alleviation of CI 5
- Storage: 1 ± 1 ◦ C, 90 ± 2% RH 1 , 6 weeks
* Decrease of CI 5 index
- Treatment condition for the highest effects was not the
maximum concentration analyzed in this study (i.e., 2 mM
PUT showed a higher effect than 3 mM PUT)

- Soil condition
* Annual average precipitation: 895.6 mm
* Annual average pressure: 956.4 hpa
* Annual average RH 1 : 81%
* Average annual sunshine hours: 1032.9 h - Improvement in the stability of fruit quality
* Annual average wind speed: 1 m/s
Application of fertilizer 10–15 days before * Higher weight
Beijing 2 - Fertilizer [60]
flowering and 15 days before harvest * Higher firmness and TSS 4
* Urea (N content 46%), organic fertilizer (main
ingredient is dried chicken manure), potassium
sulfate (K2 O content 50%), monoammonium
phosphate (P2 O5 content 46%, N content 12%),
superphosphate (P2 O5 content 12%)

1 RH: relative humidity; 2 SSC: soluble solid content; 3 TA: total (titratable) acidity; 4 TSS: total soluble solid; 5 CI: chilling injury; 6 ROS: reactive oxygen species.
Horticulturae 2023, 9, 315 10 of 41

4. Strategic Approach of the Application of Post-Harvest Treatment Technologies for

Peach Fruits
4.1. Physical Treatments
Since consumers’ concerns about the hazard of residue when using chemical compounds
have increased, physical treatment technology emerges as an alternative to protect the quality
and safety of fruits [74]. Major examples of physical treatment technologies are as follows:
temperature control [75–82], modified atmosphere [83–86], and irradiation [87–93] (Table 3).

4.1.1. Temperature Control

Post-harvest treatment based on the exposure of peach fruits to the temperature change
can be categorized as refrigeration (i.e., cooling and/or storage at low temperature) or
heating (i.e., thermal processing at high temperature).
Cooling and/or cold storage is one of the simplest methods for extending the shelf life
of fruits with minimal quality change through inhibitory effects on not only the enzymatic
activities related to the maturation but also microbial infectious metabolism [94–96]. Since
fruit rot and microbial growth are mainly caused by the degree and mode of temperature
fluctuation, previous research focused on the discovery of determinant factors (e.g., the
condition of raw materials, storage temperature and time) for maintaining fruit quality and
novel operational methods of temperature control (e.g., hydrocooling). Ceccarelli et al. [75]
explored the influence of fruit maturity categorized as IAD (index of maturity defined as a
measure of flesh and skin chlorophyll content) and the period of refrigeration on the quality
factor and aromatic characteristics of peaches; although chilling injury occurred when fruits
were stored for over 4 weeks, the relationship between harvest maturity stages (immature,
mature, and ripe) and storage time could be estimated to determine the time required for
desirable quality changes during storage (e.g., aroma development, and ripening). In the
case of the operational method of cooling, the impact of the decrease in respiration rate and
the related decrease in carotenoid content by hydrocooling (i.e., dipping of fruits in H2 O at
low temperature; 1 ◦ C for 1 h) on storability was evaluated through research by Caprioli
et al. [76], and the results of their comparative analysis confirmed the remarkable efficacy of
hydrocooling with various types of gas treatment (1-methylcyclopropene (1-MCP), carbon
dioxide, and nitrogen).
To apply thermal post-harvest technologies to peach fruit, hot air and hot water treat-
ments have been adopted as applicable methods operated by air circulation and dipping in
solution, respectively. Both hot air and hot water treatments can contribute to not only the
improvement of fruit quality (e.g., firmness, ripening, and decay) but also the prevention
of chilling injury during cold storage by inhibiting the loss of membrane integrity and the
accumulation of ROS [97–99]. However, when tested on the same peach fruit samples, the
comparison of hot air and hot water as post-harvest treatment technologies demonstrated
that hot water had a higher efficacy than hot air. Huan et al. [77] showed the differences
between the heat transfer methods (hot water and hot air) in regard to the efficacy against
chilling injury (higher efficiency on the heat transmission of hot water treatment, which
is useful for the inhibition of internal browning compared to hot air treatment) and an-
tioxidant activity (enhancement of AsA-GSH metabolisms by hot water treatment, but
not by hot air treatment). The effects of hot water treatment were also assessed at room
temperature to understand the response of peach fruit during ripening after treatment.
Zhang et al. [78] conducted a proteomic analysis of peach fruit treated with hot water (48 ◦ C,
10 min) followed by storage at 25 ◦ C and showed the distinct heat-shock protein expression
linked to the resistance to stress responses or self-defense capability and the activation
of multiple antioxidant metabolic pathways (e.g., AsA-GSH). The intervention effect of
hot water treatment as an effective decontamination technology on peach fruit artificially
inoculated with fungi (e.g., Monilinia sp.) was also assessed at room temperature [79] and
low temperature [80]. Liu et al. [79] revealed the mode of action for hot water treatment
against the post-harvest decay caused by brown rot fungi (M. fructicola) during the exposure
to room temperature by both direct antifungal effects (the dysfunction of mitochondria and
Horticulturae 2023, 9, 315 11 of 41

inhibition of spore germination of M. fructicola) and the host defense mechanism (induction
of chitinase, β-1,3-glucanase, and phenylalanine ammonia lyase enzymes). M. laxa, capable
of germinating at low temperatures, was also adopted as a target fungus inactivated by
hot water treatment (48 ◦ C, 12 min) of peach fruit by Jemric et al. [80], and a decrease in
microbial deterioration could also be achieved during cold storage (0 ◦ C, 90% RH, 20 days).

4.1.2. Modified Atmosphere

The methods for the modification of atmosphere during the storage of peaches to
extend their shelf life and to improve the fruit quality can be represented as controlled
air [82] and hypobaric treatment controlling the atmospheric pressure [84–86].
Supplying controlled air can maintain the intended composition of gas filled in the
area for the storage of fruits. Liu et al. [83] showed the effects of sealing peach fruits in a
50 L container with a strictly controlled atmosphere (5% O2 , 10% CO2 , and 85% N2 ) during
cold storage to alleviate chilling injury and the accumulation of aroma volatiles preferred
by consumers.
The hypobaric treatment allows the generation of a low oxygen environment by decreasing
atmospheric pressure. Wu et al. [84] suggested the beneficial effect of hypobaric treatment as
the regulation of defense-related enzymes governing antioxidant activities (e.g., stimulation of
SOD and PPO, and suppression of CAT). Whereas, the major problems in applying modified
atmospheric control technology (e.g., high operating costs, the difficulty of maintaining a stable
hypobaric environment, and the loss of quality of firmness) are expected to be solved by using
short-term hypobaric treatment [85,86]. Zhang et al. [85] placed peaches in the hypobaric tank
(0.45 atm for 4 h at 20 ◦ C) before cold storage and enhanced chilling tolerance of fruits by the
activation of antioxidant enzymes along with the suppression of membrane oxidation to prevent
the accumulation of MDA was shown. Zhan, et al. [86] also emphasized the protective effect
of short-term hypobaric treatment against chilling injury during subsequent cold storage and
revealed the major working mechanism as the regulation of fruit metabolisms linked to the
membrane fatty acids (e.g., activation of fatty acid synthetase (the synthesis of fatty acid) and
fatty acid desaturase (the desaturation of fatty acid), but suppression of LOX (the degradation
of unsaturated fatty acid)).

4.1.3. Irradiation
Light Irradiation
Light-based antimicrobial photoinactivation has been applied in the food industry
for the maintenance of quality, especially for fresh-cut fruits and vegetables that cannot
be processed with heat [100]. The irradiation of fruits with light can also affect various
physiological metabolic pathways related to growth, development, ripening, softening,
stress-response, and disease tolerance [101]. Both visible light and ultraviolet light (UV) are
expected to achieve decontamination and to modulate metabolism, but UV treatment has
been preferred due to the probability of inducing ripening (e.g., synthesis of ethylene) and
the lower decontamination effects of visible light [102]. The post-harvest effects of light
irradiation are known to be wavelength-dependent within the area of UV (categorized
as UV-A (315–400 nm), UV-B (280–315 nm), and UV-C (100–280 nm)) [103–105]. Previous
research regarding the application of UV irradiation to harvested peaches has been reported
as a singular treatment of UV-B [88] or UV-C [87], and the combined treatment of UV-B
with UV-C [89].
Santin et al. [88] analyzed changes in plant secondary metabolism through non-
targeted fruit metabolomics after a low level of UV-B irradiation (2.3134 W/m2 , 60 min,
24 ◦ C), and showed noteworthy metabolomic changes in most phenolics according to the
storage time after treatment (e.g., the decrease and accumulation of phenolics after 24 h
and 36 h storage, respectively), emphasizing the importance of time after UV-B irradiation
as the key determining factor of the efficacy [88]. The transcriptome-based investigation of
the effects of UV-C (4 kJ/m2 , 30 min, 10 ◦ C) irradiation on the control of fruit metabolism
associated with softening and senescence (cell wall, antioxidant, secondary metabolism,
Horticulturae 2023, 9, 315 12 of 41

lipid, and energy) during storage was also conducted by Kan et al. [87] and revealed
the upregulation of genes linked to defense systems with the activation of antioxidation
enzymes and the downregulation of genes inducing undesirable quality deterioration (e.g.,
ethylene biosynthesis, oxidative stress, lipid peroxidation, and cell wall decomposition).
In the case of combined treatment (UV-B + UV-C), Abdipour et al. [89] demonstrated a
relatively lower effect of UV-B than UV-C by the direct comparison of the effects of UV
irradiation with different wavelengths on the storability of the same peach samples, but
the combined treatment can be considered a strategy to complement the limited effects of
UV-B and to increase the overall effects, improving fruit quality parameters (TSS, firmness,
total phenolic compounds, TA, vitamin C content, and acidity).

Gamma Irradiation
Gamma ray that has greater energy than X-ray or UV photons has been generally used
to destroy covalent bonds in the DNA of microbial contaminants by penetrating target
foods [106]. Whereas, Khan et al. [91] observed that irradiating peaches with gamma rays
(2.5 and 5 kGy) in an ambient temperature did not induce a significant difference in the qual-
ity of fruits. However, excessive exposure of peaches to gamma ray causes fruit softening
(>1 kGy). Melo et al. [92] also demonstrated that irradiation-induced immediate softening
is associated with cell wall modifications, pectin hydrolysis, and pectin methylesterase
activity by evaluating the influence of gamma irradiation (ca. 1 kGy) to peaches on ripening
parameters (e.g., changes in color, weight, and contents of antioxidants).

Microwave Irradiation
Microwaves, which transform electromagnetic field energy into thermal energy, have
frequencies and wavelengths in the range of 0.3–300 GHz and 0.001–1 m, respectively [107].
Wang et al. [93] observed that the quality change of peach fruits exposed to a microwave
and the longest irradiation time exhibited the most effective results (e.g., decrease of inter-
nal browning, inhibition of the phenolic accumulation, and improvement of membrane
stability) among treatment conditions (45.5 W for 3, 5, and 7 min). This study also sug-
gested that a protective mechanism to internal browning induced by chilling injuries was
considered as a non-thermal effect because 45.5 W microwave irradiation could not increase
the core temperature of peach fruit [93].
Horticulturae 2023, 9, 315 13 of 41

Table 3. Post-harvest treatments of physical technologies on peach fruit.

Method Cultivar Treatment Conditions Results and Implications Reference

- Maturity class of fruit samples

- Fruit ripening and quality change were affected by the harvest
* Pre-climacteric (CI; immature): IAD 1 1.6–1.3 maturity stage according to the period of cold storage
* Onset of climacteric (CM; mature): IAD 1 1.2–0.8 * Different pattern of the changes in firmness, SSC 3 ,
Cold storage August Flame [75]
* Climacteric (CR; ripen): IAD 1 0.7–0.0 TA 4 , ethylene emission, and aroma development
- Storage: 0 ◦ C, 95% RH 2 , 4 weeks among samples (CI, CM, CR)
- Post-storage: 18 ◦ C, 6 days

- Treatment conditions - Improvement in the stability of fruit quality

* Dipping in 1 ◦C water, 1 h * Higher firmness and carotenoid content

Hydrocooling followed by cold storage Spring Belle * Lower level of respiration rate and ethylene [76]
- Storage: 0 ◦ C, 7 days production
- Post-storage: 20 ◦ C, 50–60% RH 2 , 1 day - No effect on SSC 3 and skin color

- Hot air treatment - Higher efficacy from hot water than hot air treatment
* Air-circulation (38 ◦ C, 3 h) * Improvement in the stability of fruit quality: Higher
Hot air or hot water treatment followed by - Hot water treatment firmness and membrane integrity, lower respiration
Xiahui 5 rate and ethylene production, lower internal [77]
cold storage
* Immersion (48 ◦ C, 10 min) and air-drying (1 h) browning index
- Storage: 4 ± 0.5 ◦ C, 85–90% RH 2 , 35 days * Improvement of antioxidant capacity

- Improvement of antioxidant capacity

- Treatment conditions * Lower ROS 5 content
Hot water treatment followed by room * Immersion (48 ◦ C, 10 min) and air-drying (1 h) - Improvement of self-capability of the defense
Huiyulu [78]
temperature storage
- Storage: 25 ± 1 ◦ C, 80–90% RH 2 , 5 days * Expression of proteins related to the stress-response
and antioxidant metabolism

- Decrease of microbial deterioration and damage

- Treatment conditions * Lower disease incidence and lesion diameter
* Inoculation before the treatment: Monilinia fructicola - Induction of the expression of defense-related genes
Hot water treatment followed by room (10 µL, ca. 4 log spores/mL) * Chitinase, β-1,3-glucanase, phenylalanine ammonia
June Prince * Immersion (40 ◦ C, 5–10 min) and air-drying (25 ◦ C, 10 [79]
temperature storage lyase
min)
- No adverse effects on product quality
- Storage: 25 ◦ C, 3 days
* Firmness, SSC 3 , and TA 4
Horticulturae 2023, 9, 315 14 of 41

Table 3. Cont.

Method Cultivar Treatment Conditions Results and Implications Reference

- Treatment conditions - Decrease of microbial deterioration and damage

* Inoculation before the treatment: Monilinia laxa (2 µL, * Lower decay index
Hot water treatment followed by cold storage Roig 5 log conidia/mL) [80]
- Improvement in the stability of fruit quality
* Immersion (48 ◦ C, 12 min)
4 3 4
* Lower TA and SSC /TA ratio
- Storage: 0 ◦ C, 90% RH 2 , 20 days

- Improvement in the stability of fruit quality

- Treatment conditions
* Inhibition of ethylene emission and internal
Controlled air treatment during cold storage Hujingmilu * 5% O2 and 10% CO2 , and 85% N2 browning [83]
* Higher firmness
- Storage: 0 ◦ C, 90% RH 2 , 28 days
- Higher accumulation of aroma volatiles

- Improvement in the stability of fruit quality

* Higher firmness
- Treatment conditions * Lower malondialdehyde (MDA) content
Hypobaric treatment during cold storage Xiahui-8 * 55 ± 5 kPa in a hypobaric chamber * Delay of respiration peak [84]
- Storage: 4 ± 0.5 ◦ C, 16 days * Activation of the antioxidant enzymes (superoxide
dismutase (SOD), polyphenol oxidase)
* Suppression of catalase (CAT) activity

- Improvement in the stability of fruit quality

- Treatment conditions * Lower respiration rate
Short-term hypobaric treatment followed by * 0.45 atm (standard atmospheric pressure) for 4 h at 20 * Higher TSS (total soluble sugar) concentration
Yingshuanghong ◦ C in a hypobaric tank * Delay of climacteric peak [85]
cold storage
- Storage: 5 ± 1 ◦ C, 2
85 ± 5% RH , 24 days * Activation of the antioxidant enzymes (SOD, CAT,
peroxidase (POD), ascorbate peroxidase)

- Improvement in the stability of fruit quality

* Lower MDA and electrolyte leakage
* Higher membrane fluidity and fatty acid
- Treatment conditions unsaturation
Short-term hypobaric treatment followed by * 45.6 kPa at 20 °C for 2.5 h in a hypobaric chamber - Regulation of the enzymes involved in membrane fatty acid
Feicheng [86]
cold storage metabolisms
- Storage: 0 ± 0.5 ◦ C, 90 ± 5% RH 2 , 35 days
* Suppression of lipoxygenase activity
* Activation of fatty acid synthetase and fatty acid
desaturase
Horticulturae 2023, 9, 315 15 of 41

Table 3. Cont.

Method Cultivar Treatment Conditions Results and Implications Reference

- Improvement in the stability of fruit quality

* Higher firmness
* Lower ethylene production
- Treatment conditions - Transcriptomic analysis
UV-C irradiation followed by cold storage Xiahui 5 * UV-C irradiation: 4 kJ/m2 , 30 min, 10 ◦ C * Upregulation of genes related to pectin content, [87]
◦ C, 2
secondary metabolism, lipid or energy metabolism,
- Storage: 10 85% RH , 9 days and stress resistance
* Upregulation of genes related to antioxidant enzymes
(SOD, POD)
* Upregulation of genes related to signal transduction

- Improvement in the stability of fruit quality

- Treatment conditions * Higher structural lipid
UV-B irradiation followed by room * UV-B irradiation: 2.3134 W/m2 , 60 min, 24 ◦ C - Improvement of antioxidant capacity
Fairtime [88]
temperature storage
- Storage: 24 ◦ C, 85% RH 2 , 24–36 h * Higher phenolic contents (dihydroflavonols,
anthocyanins, and flavones), alkaloid (pteridine)

- Improvement in the stability of fruit quality

* Lower weight loss
- Singular treatment * Higher firmness, vitamin C content, and TA 4
* UV-B irradiation: 0.72 kJ/m2 , 20 min, 8 ◦ C * Lower TSS 6
* UV-C irradiation: 0.72 kJ/m2 , 20 min, 8 ◦ C - Decrease of microbial deterioration and damage
UV-C and UV-Birradiation followed by cold
- - Combined treatment [89]
storage * Lower decay rate
* 2
UV-B + UV-C irradiation: 1.44 kJ/m , 20 min, 8 ◦C - Improvement of antioxidant capacity
- Storage: 4 ◦ C, 80–85% RH 2 , 25 days * Higher total phenolics content
- Synergistic effects by the combined treatments of UV-B with
UV-C

- Treatment conditions
Gamma irradiation followed by room * Co-60 gamma ray: 71.4 krad/h - No significant difference between control and irradiated
- * Irradiation: 2.5, 5 kGy peach samples [91]
temperature storage
- Storage: 25 ± 2 ◦ C, 14 days
Horticulturae 2023, 9, 315 16 of 41

Table 3. Cont.

Method Cultivar Treatment Conditions Results and Implications Reference

- Treatment conditions
* Prior to irradiation: dipping in 100 mg/L active
- Demonstration of the mechanism for irradiation-induced
chlorine solution at 3 ◦ C for 10 min
Gamma irradiation followed by cold storage Mid pride immediate loss of firmness as pectin hydrolysis, cell wall [92]
* Irradiation: average absorbed dose as 1030 Gy
modifications, and pectin methylesterase activity
- Storage: 1 ◦ C,7 days
- Post-storage: 23 ± 2 ◦ C, 6 days

- Treatment conditions - Improvement in the stability of fruit quality

Microwave irradiation followed by cold * Power: 45.5 W, 25 ◦ C, 3–7 min * Decrease of internal browning
* Inhibition of phenolic accumulation [93]
storage - Storage: 5 ◦ C, 30 days
- Post-storage: 25 ◦ C, 1 day * Increase of membrane stability

1 IAD : index of absorbance difference; 2 RH: relative humidity; 3 SSC: soluble solid content; 4 TA: total (titratable) acidity; 5 ROS: reactive oxygen species; 6 TSS: total soluble solid.
Horticulturae 2023, 9, 315 17 of 41

4.2. Chemical Treatments

Using chemical agents that can enhance the quality and safety of fruits by controlling
fruit maturation (e.g., naturally occurring compounds extracted from growing plants; pu-
trescine, spermidine) and decontamination (e.g., bactericidal and/or fungicidal substances)
has been regarded as an efficient post-harvest treatment method. Treatment strategies can
be categorized as liquid (solution) and gas (vaporized materials) phases of the chemical
agents according to the application methods, as shown in Table 4.

4.2.1. Spraying or Dipping Treatment Methods Using Solutions of Chemical Agents

Treatment solutions containing active chemical agents can be mainly applied to fruits
by spraying or dipping. Dipping treatment enables the even exposure of fruits to chemical
agents, and thus is likely to be preferable as the post-harvest treatment to fruits rather than
spraying treatment.

Spraying Treatment
Citric acid spraying can enhance the stability of fruit quality factors (firmness, TA, and
TSS) during room temperature storage, and the considerable reduction of decay incidence
is likely due to the potential antifungal capability of citric acid [108].
Glucose oxidase (GOx) is a natural anti-browning and antimicrobial agent that can
be used as an alternative to synthetic chemicals [109]. Batool et al. [110] immobilized
GOx by using zinc oxide nanoparticles (ZnONPs) to improve not only the stability but
also the activity of the enzyme, and GOx/ZnONP bioconjugation spray resulted in the
maintenance of the physiological appearance of peach fruits and a decrease in undesirable
quality changes of peach fruits (e.g., firmness, and TSS). These effects occurred through the
expected mechanisms as follows: (1) antioxidant (i.e., scavenging oxygen) and antimicrobial
effects; and (2) formation of H2 O2 layer to slow fruit metabolism associated with ripening
and to protect the fruit from fungal contamination.
Spraying essential oils (EOs) on the surface of fruits has been adopted as the rep-
resentative decontamination treatment against pathogenic and/or spoilage bacteria and
fungi [111–113]. In the case of peaches, research by Elshafie et al. [114] on antifungal effects
of the major constituents in Greek oregano (Origanum vulgare L. ssp. hirtum) EO revealed
that thymol and carvacrol showed strong efficacy against fruit pathogenic fungi (Monilinia
spp.; M. laxa, M. fructigena, and M. fructicola).

Dipping Treatment
EO is one of the most popular antifungal agents and has been generally applied
directly to food products by spraying or dipping methods. Dipping peaches in EOs showed
antifungal effects against fruit pathogens equivalent to those of commercial fungicide
products, highlighting the value of EOs as natural antifungals that are feasible alternatives
to synthetic fungicides; however, the species-dependent efficacy from each source of EOs
indicates the importance of identifying the spectrum of EOs to be used [115]. However,
the peculiar fragrance of EOs and relatively high cost compared with synthetic chemical
antifungals used for fruits have been regarded as the major limitations [116,117]. Thus, the
combined treatment of EO with other antifungal agents is expected to improve the overall
effects on the product quality as a countermeasure for those limitations. Rahimi et al. [118]
reported that dipping treatment with a solution of EO and chitosan considerably prevented
fungal decay with desirable effects on sensory characteristics as well.
Glycine betaine (GB) can act as an osmotic adjustment substance to enhance the tolerance
of fruits against cold stress factors by preventing membrane damage [119]. Shan et al. [120]
evaluated the effects of exogenous GB treatment from the perspective of reducing the chilling
injury of cold-stored peaches; their findings suggested that the key mechanism is an increase in
the contents of endogenous substances involved in protective responses to cold stresses (GB,
g-aminobutyric acid (GABA), and proline) through the induction of relevant enzymes (betaine
Horticulturae 2023, 9, 315 18 of 41

aldehyde hydrogenase (BADH), glutamate decarboxylase (GAD), D 1 -pyrroline-5-carboxylate

synthetase (P5CS), and ornithine d-aminotransferase (OAT)).
Exogenous melatonin treatment extends the shelf life of peach fruits through the
activation of antioxidant enzymes capable of enzymatic ROS control during both room
temperature [121] and cold storage [122]. Gao et al. [121] reported that dipping peaches in
a melatonin solution can maintain fruit quality (e.g., firmness and decay incidence) and
decrease weight loss, with the key mechanism being the activation of antioxidant enzymes
(SOD, POD, CAT, and APX). Research by Cao et al. [122] also showed an increase in the
activity of antioxidant enzymes (POD, SOD, and CAT) in peaches during cold storage,
and suggested that antioxidant systems were activated by the upregulated transcription
of genes involved in the production of not only those enzymes but also reductants (AsA
and GSH), serving as the mode of action of the increased tolerance of peaches against cold
stress.
Putrescine, a poly-amine substance widely used as an antiaging compound for fruit
skin, can contribute to the control of post-harvest loss of the quality and nutrition of
peaches by preventing chilling injuries and the breakdown of biochemical compounds (e.g.,
phenolic compounds, vitamin C, and organic acids), respectively [123].
Endogenous GABA is involved in the defense system against cold stress and the
effects of exogenous GABA treatment as post-harvest interventions of chilling injuries
were reported as the result of the accumulation of endogenous GABA with proline linked
to stress adaptation to the cold environment. Shang et al. [124] reported those effects on
peaches treated with 10 min of dipping in GABA solution; although, the effects were not
concentration-dependent (i.e., 5 mM was the most effective treatment concentration rather
than the highest treatment concentration in this study (10 mM)).
Salicylic acid is a phenolic compound that plays a role in the regulation of the ripening
and growth of fruits. The effects of salicylic acid dipping treatment as the post-harvest
intervention against the quality change of peach fruits during cold storage have also been
consistently reported, with in-depth examinations indicating that the major mechanisms
of these effects are both the activation of vital antioxidant enzymes (SOD, POD, and CAT)
and the inhibition of the browning enzyme (PPO) [125,126].
CaCl2 has been reported as one of the most common post-harvest treatment agents that
can stabilize cellular membranes and delay senescence by inhibiting enzymes responsible
for the deterioration of products [127]. The dipping of peaches in 6% CaCl2 solution for
10 min showed a delay in spoilage and various undesirable quality changes (decrease in
firmness, acidity, and reduction of sugar content) accompanied by minimized PME activity
to ensure the long-term storage (3 weeks under ambient temperature and 3 days under
cold temperature as storage and post-storage conditions, respectively) of fruits in an edible
state, which was also validated by the palatability test [128].
The direct validation of antifungal effects has been conducted by using peaches
artificially inoculated with pathogenic or spoilage fungi and a case study of the applicability
test for peach fruits has been reported with the following agents: yeast saccharides (YS) and
benzo-thiadiazole-7-carbothioic acid S-methyl ester (BTH) [129,130]. YS from the cell wall
can induce the defense responses of products due to their antifungal activities (activation of
chitinase and β-1,3-glucanase, which can degrade chitin and β-1,3-glucan in the cell wall of
fungi, respectively) with enhanced phenolic synthesis (higher PAL and POD activities) and
these activities were linked to the role of YS as a trigger for increasing endogenous nitric
oxide (NO) levels of the product [129]. However, according to research by Yu et al. [129],
optimizing the treatment conditions of dipping products in YS based on endogenous
NO levels is necessary because those effects are not treatment concentration-dependent:
treatment with 0.5 mg/L YS was more effective (higher NO levels and lower decay) than
treatment with 0.1 or 1.0 mg/L YS. For BTH treatment, the expected mechanisms of
antifungal effects against P. expansum in the product are the production of ROS followed by
the strengthening of systemic acquired resistance through the activation of host defense
enzymes (e.g., PAL, PPO, and POD) [130].
Horticulturae 2023, 9, 315 19 of 41

Fruit surface can be coated by dipping in the solution of nanoparticles, which are
nontoxic and available for the targeted localization. Calcium nanoparticles combined
with ascorbic acid (9 mM/L) suppressed the incidence of chilling injury during the cold
storage of peach fruits with a stable preservation of skin color and moisture [131]. Gad and
Ibrahim [132] suggested nano-chitosan as a coating agent, which allowed the maintenance
of fruit quality (e.g., lower weight loss, a decrease of decay incidence, and higher firmness)
of peaches, and showed better effects obtained from a specific treatment condition (400
ppm) than the maximum concentration tested in this study (800 ppm) to highlight the
importance on the exploration of the optimal condition. Since chitosan nanoparticles are
effective and eco-friendly, the enhancement of marketability and storability of peach fruits
is expected [132].
Films and coatings applied to fruits and vegetables with edible agents (e.g., gum)
by dipping can protect foods from environmental stress factors (e.g., moisture migration,
microbial contamination, light exposure, and oxidation), which can result in product quality
changes [133]. Peach-gum coating (i.e., dipping in 1–10% gum solution) was suggested
as a novel strategy for the prevention of ageing, which can result in the softening of the
products, and the mechanism of this effect was revealed by transcriptomic analysis to be
the downregulation of genes related to the deterioration of product quality (e.g., ethylene
synthesis and cell wall degradation) [134].

Sequential Dipping and Spraying Treatments

The combined treatment of dipping followed by the spraying of antifungal agents
can be adopted as the post-harvest sanitation strategy with durable fruit decontamination
ability. The effects of the dipping treatment of near-neutral (pH = 6.3–6.5) electrolyzed
oxidizing water (NEO water), which inactivates brown rot fungi (M. fructicola) to mitigate
the potential infection on the surface of peaches (i.e., reducing the incidence and severity of
brown rot), was improved by the combined treatment of daily spraying of NEO water after
NEO dip [32]. The use of electrolyzed water as the post-harvest antifungal treatment has
been regarded as practical and has commercial traits due to its economic feasibility (e.g.,
low cost of raw materials and the maintenance of electrolyzed water generators) [135].

4.2.2. Gas Treatment of Vaporized Chemical Agents

Volatile organic compounds (VOCs) produced by microorganisms have been used
for post-harvest pathogen control and Zhou et al. [136] showed that the fumigation of
benzothiazole as a VOC (from Bacillus subtilis), which is known as an antifungal agent
against M. fructicola, can be used not only for the decontamination of pathogenic fungi but
also for the activation of antioxidant enzymes.
Fumigation of NO can prevent the quality change of peaches during storage through
the reduction of ethylene production and/or the improvement of antioxidant capacity sup-
ported by higher activities of antioxidant enzymes (e.g., CAT and SOD) [137–139]. NO can
also reduce the activity of LOX and 1-aminocyclopropane-1-carboxylic acid (ACC) oxidase
through nitrosation to inhibit membrane lipid peroxidation and the ethylene biosynthesis
pathway, respectively [138]. Since controversial results regarding the effects of NO fumiga-
tion on product quality factors (e.g., color, TA, or TSS) were also observed, optimization of
treatment conditions based on various quality factors is also required [137,138]. In the case
of changes in the lipid composition associated with ion leakage and membrane integrity,
the most desirable effects were achieved by treatment with an intermediate concentration
(10 µL/L), whereas opposite effects according to the excessive exposure of peaches to
fumigated NO were observed at the highest concentration (15 µL/L), highlighting the
importance of determining the optimum conditions [139]. Mechanisms for the beneficial ef-
fects of NO treatment during ripening were also revealed by proteomic analysis as follows:
(1) induction of antioxidant enzymes (SOD, enzymes of the AsA-GSH cycle); (2) decrease in
ethylene production by the production of complex 1-aminocyclopropane-1-carboxylic acid
oxidase (ACO–NO–ACC) supported by the upregulation of S-adenosylmethionine syn-
Horticulturae 2023, 9, 315 20 of 41

thetase (SAMS) to promote the generation of the precursor of ACC (S-adenosylmethionine

(SAM)); (3) the reduction of ATP supply generated by substrate oxidation by the application
of alternative pathways for energy production (TCA cycle and glycolysis) through the regu-
lation of cytochrome c oxidase synthesis with electron transport and oxygen consumption;
(4) suppression of the degradation of the cell wall by upregulation of proteins associated
with both the loss of Ca2+ ions and the structural components of the cell wall; and (5) the
induction of defense capacity by upregulating the heat-shock protein 70 (HSP70) [140].
Peach fruits can be exposed to volatile EOs in a gas state as the post-harvest treatment
for the control of fungi responsible for the deterioration of products (e.g., B. cinerea and
Alternaria alternata) [141,142]. However, the requirement of high EO concentrations to
achieve the suppression of brown rot and Rhizopus rot limits their application in the food
industry from the perspective of marketability due to relatively higher cost of EOs as natural
agents compared that of conventional gaseous fumigants as synthetic chemical agents;
furthermore, EOs have potential phototoxicity [141]. Thus, the encapsulation of EOs and
the combined treatment with another decontamination technology can be adopted as the
countermeasure for those limitations of EO fumigation treatment [142]. Cyclodextrin-based
(CD-based) microencapsulation could protect EOs from environmental factors that decrease
their stability (e.g., temperature, light, and oxygen) and the simultaneous treatment with
1-MCP could contribute to the maintenance of fruit quality (e.g., firmness, acidity, and
decay incidence) through its inhibitory effects against ethylene releases.
1-MCP is an ethylene-antagonizing compound controlling endogenous and exogenous
ethylene to prevent the senescence of fruits, and the ephemeral microcirculation of gaseous
post-harvest agents during the storage of peach fruits showed effective preservative effica-
cies. Du et al. [143] sequentially treated peach fruits fumigated with 1-MCP with ozone
during cold storage (8 ppm O3 at 0 ◦ C for 45 days) and reported the effects on storability
(reduction of the decay rate, reduction of MDA content, the maintenance of fruit quality
(firmness, SSC, TA, and color)).
Horticulturae 2023, 9, 315 21 of 41

Table 4. Post-harvest treatments of chemical technologies on peach fruits.

Method Cultivar Method and Treatment Conditions Results and Implications Reference

- Improvement in the stability of fruit quality

- Treatment conditions
Spraying citric acid followed by room * Even spraying with 1.5 L (10 g/L) citric acid * Lower TSS 2 and decay incidence
Hujingmilu [108]
temperature storage * Increase of TA 3
- Storage: 20 ± 0.5 ◦ C, 85 ± 1% RH 1 , 14 days * Higher firmness

- Treatment conditions - Improvement in the stability of fruit quality

Spraying glucose oxidase immobilized on * Catalytic activity of GOx/ZnONPs: 23.3 ± 2.08 * Lower weight loss
ZnO nanoparticles (GOx/ZnONPs) followed - U/mL * Higher firmness, lower TSS 2 [110]
by room temperature storage
- Storage: 25 ± 1 ◦ C, 15 days - Higher DPPH free radical scavenging activity

- Treatment conditions - Antifungal effects

Spraying the essential oil (EO) compounds
emulsion followed by room temperature Springcrest * Carvacrol and thymol emulsion: 150, 500 ppm * Effective against Monilinia laxa, Monilinia fructigena, [114]
storage - Storage: 16–24 ◦ C (room temperature), moist chamber, 5 days and Monilinia fructicola

- Treatment conditions
* Dipping time: 10 min
* Essential oil: 0.15, 0.20% extracts from Eucalyptus
- Antifungal effects
globulus, Cinnamomum camphora, and Cymbopogum
Dipping in EO solution followed by cold or citratus * Effectiveness equivalent to fungicide (orthocide)
Chimarrita [115]
room temperature storage * Fungicide: Orthocide 2.4 g/L
C. camphora and C. citratus against Colletotrichum gloeosporioides
- Storage : C. citratus against M. fructicola
* Cold condition: 20 days
* Room temperature: 10 days

- Treatment conditions - Improvement in the stability of fruit quality

* Dipping time: 15 min * Lower weight loss
* Single treatment * Higher firmness
: 0.5% chitosan * Lower TSS 2 and decay incidence
Dipping in EO and/or chitosan followed by : 200 mg/L thymol
Zaferani - Improvement of nutritional quality [118]
cold storage
* Combined treatment * Higher anthocyanin and carotenoid
: 0.5% Chitosan + 200 mg/L thymol - Sensory properties
◦ C, 1
* Preferable sensory characteristics
- Storage: 6 80% RH , 30 days
Horticulturae 2023, 9, 315 22 of 41

Table 4. Cont.

Method Cultivar Method and Treatment Conditions Results and Implications Reference

- Alleviation of CI 4
* Lower CI 4 index
* Higher level of endogenous GB, γ-aminobutyric acid
(GABA), and proline contents
* Activation of enzymes related to the response to cold
- Treatment conditions stress: betaine aldehyde hydrogenase (BADH),
Dipping in glycine betaine (GB) solution * Dipping time: 10 min glutamate decarboxylase (GAD),
YuhuRa No.2 * 10 mM exogenous GB D1-pyrroline-5-carboxylate Synthetase (P5CS), and [120]
followed by cold storage
ornithine d-aminotransferase (OAT)
- Storage: 0 ◦ C, 5 weeks
- Improvement in the stability of fruit quality
* Lower firmness, higher extractable juice
- Higher energy status
* Higher energy charge, ATP, and ADP contents

- Improvement in the stability of fruit quality

* Lower weight loss
- Treatment conditions
* Higher firmness, lower decay incidence
Dipping in melatonin solution followed by Shahong, * Dipping time: 10 min * Lower malondialdehyde (MDA), decreased activity
* 0.1 mM/L melatonin of lipoxygenase (LOX) [121]
room temperature storage Qinmi
* Activation of the antioxidant enzymes: superoxide
- Storage: 25–28 ◦ C, 60–70% RH 1 , 7 days
dismutase (SOD), peroxidase (POD), catalase (CAT),
ascorbate peroxidase (APX)

- Alleviation of CI 4

- Treatment conditions * Lower CI 4 index

* Improvement in the stability of fruit quality
Dipping in melatonin followed by cold * Dipping time: 120 min * Lower MDA content
Hujing * 100 µM melatonin * Activation of antioxidant enzymes (CAT, SOD, APX) [122]
storage
- Storage: 4 ◦ C, 80% RH 1 , 28 days * Upregulation of the genes associated with the
modulation of reductants (ascorbate acid (AsA),
glutathione (GSH)), which can directly detoxify ROS 5

- Improvement in the stability of fruit quality

- Treatment conditions
* Lower weight loss
Dipping in putrescine (PUT) solution * Dipping time: 2 min * Lower decay incidence
Monley * 0.4, 0.8, 1.2, 1.6 mM PUT (1 L) * Higher firmness, higher fruit density [123]
followed by room temperature storage
* Prevention of the degradation of phenolic
- Storage: 0 ± 0.5 ◦ C, 90 ± 5% RH 1 , 40 days
compounds, vitamin C, and organic acid
Horticulturae 2023, 9, 315 23 of 41

Table 4. Cont.

Method Cultivar Method and Treatment Conditions Results and Implications Reference

- Alleviation of CI 4

- Treatment conditions * Lower CI 4 index

* Dipping time: 10 min - Increase in the activities of enzymes related to the

Dipping in GABA solution followed by cold * 1, 5, 10 mM GABA accumulation of endogenous GABA and proline: glutamate
Baifeng decarboxylase (GAD), ∆1-pyrroline-5-carboxylate synthetase [124]
storage
- Storage: 1 ◦ C, 85–95% RH 1 , 35 days (P5CS), and ornithine δ-aminotransferase (OAT)
- Post-storage: 20 ◦ C, 3 days - Overall effects were not concentration-dependent: 5 mM
GABA showed the highest effects (compared with 1 mM and
10 mM GABA)

- Improvement in the stability of fruit quality

* Lower weight loss
- Treatment conditions
* Lower pH and TSS 2
Dipping in salicylic acid solution followed by * Dipping: 0.5, 1, 1.5 mM, 10 min
Anjiry maleky * Higher firmness and TA [126]
cold storage
- Storage: 1 ◦ C, 80–90% RH 1 , 28 days - Improvement of nutritional quality
* Higher contents of total phenol and flavonoid

- Improvement in the stability of fruit quality

* Lower weight loss
- Treatment conditions * Higher firmness, lower pH, lower decay incidence
Dipping in salicylic acid solution followed by - Dipping: 0.5, 1, 1.5, 2.0 mM, 5 min * Activation of antioxidant enzymes (CAT, SOD, POD)
Flordaking [125]
cold storage - Storage: 0 ◦ C, 90% RH 1 , 5 weeks - Decrease in the activity of fruit browning enzyme: polyphenol
oxidase (PPO)
- Higher radical scavenging activity

- Treatment conditions - Improvement in the stability of fruit quality

Dipping in CaCl2 solution followed by - Dipping: 4, 6% CaCl2 , 10 min * Lower spoilage incidence, higher firmness, higher
ambient temperature storage after cold Earli Grande acidity, higher reducing sugar, minimum pectin [128]
* Storage: 0–2 ◦ C, 85–90% RH 1 , 21 days
storage methyl esterase (PME) activity
- Post-storage: 28–30 ◦ C, 1
65–70% RH , 72 h * Higher palatability rating
Horticulturae 2023, 9, 315 24 of 41

Table 4. Cont.

Method Cultivar Method and Treatment Conditions Results and Implications Reference

- Improvement in the stability of fruit quality

* Lower decay incidence and lesion diameter
- Treatment conditions
◦ C, - Induction of defense responses of products against fungal
* Dipping: 0.1, 0.5, 1.0 mg/L (in 10 L), 20 5 min
infection based on the increase of endogenous nitric oxide
* Drying: 3 h
Dipping in yeast saccharide solution followed (NO) levels
Baifeng - Inoculation [129]
by room temperature storage * Activation of the antifungal enzymes (chitinase,
* Penicillium expansum β-1,3-glucanase)
* Enhancement of phenolic synthesis by higher
- Storage: 20 ◦ C, 95% RH 1 , 6 days
activities of phenylalanine ammonia-lyase (PAL) and
POD

- Improvement in the stability of fruit quality

- Treatment conditions
* Lower weight loss
* Dipping: 200 mg/L BTH solution, 5 min * Lower disease incidence rate
Dipping in benzo-thiadiazole-7-carbothioic
acid S-methyl ester (BTH) solution followed Jiubao - Inoculation - Induction of defense responses of product against fungal [130]
by room temperature storage * P. expansum (after 60 h of dipping treatment) infection
* Enhancement of the activities of PAL, PPO, and POD
- Storage: 22 ◦ C, 85–95% RH 1 , 16 days
* Increase of total phenolic compounds and H2 O2

- Improvement in the stability of fruit quality

- Treatment conditions
* Soaking: 9 mM calcium nanoparticles with ascorbic * Lower CI 4 index, SSC 6 /TA 3 ratio, ethylene
Soaking in calcium nanoparticles with production, and respiration rate
Florida Prince acid, 15 min, 4 ◦ C [131]
ascorbic acid followed by cold storage * Higher firmness
- Storage: 4 ± 1 ◦ C, 1
95 ± 1% RH , 30 days * Lower MDA content

- Treatment conditions - Improvement in the stability of fruit quality

Dipping in nano-chitosan followed by cold * Dipping: 200, 400, 800 ppm nano-chitosan * Lower weight loss
Florida Prince * Higher firmness [132]
storage - Storage: 0 ± 1 ◦ C, 90–95% RH 1 , 3 weeks
* Lower decay incidence
- Post-storage: 20 ± 2 ◦ C, 70–75% RH 1 , 3 days

- Improvement in the stability of fruit quality

* Lower weight loss
* Lower ethylene production
- Treatment conditions
* Higher firmness
Dipping in gum followed by refrigerated * Dipping: 1, 5, 10% peach gum solution, 10 min
Jinxiu * Reduction of sorbitol breakdown [134]
storage
- Storage: 8 ◦ C, 25 days - Transcriptomic analysis
* Downregulation of genes governing ethylene
synthesis, softening, ageing, and stress responses
Horticulturae 2023, 9, 315 25 of 41

Table 4. Cont.

Method Cultivar Method and Treatment Conditions Results and Implications Reference

- Inoculation
* M. fructicola
- Treatment conditions
- Antifungal effects
Dipping and daily spraying of electrolyzed * Near-neutral electrolyzed oxidizing water (pH
oxidizing water followed by room - 6.5–6.7; oxidation/reduction potential 800–900 mV; * Effective against M. fructigena [32]
temperature storage 250 ppm total residual chlorine species) : Lower incidence of infection and diseases severity
* Dipping: 10 min
* Spraying: 6 mL, daily
- Storage: 25 ◦ C, 90% RH 1 , 20 days

- Improvement in the stability of fruit quality

- Treatment conditions * Decrease of enzyme (cellulase, pectinase) activities
Fumigation of volatile organic compounds which destroy the plant tissues
* Addition of 200 µL of CF-3 24h FB on filter paper
from Bacillus subtilis followed by room Zhaohui * Activation of antioxidant enzymes (POD, PPO, CAT, [136]
sheet
temperature storage SOD) to decrease MDA content and cell membrane
- Storage: 25 ◦ C, 4 days permeability by the protection from product damage
by oxidants

- Improvement in the stability of fruit quality

* Higher firmness, lower ion leakage, lower rot index
- Treatment conditions
Fumigation of NO gas followed by cold - Changes of lipid composition
Feicheng * 5, 10, 15 (µL/L), 3 h [139]
storage * Decrease of unsaturated C-18 fatty acid (18:3)
- Storage: 5 ◦ C, 40 days
associated with ion leakage and the integrity of the
membrane

- Treatment conditions - Improvement in the stability of fruit quality

* 5, 10, 15 (µL/L), 3 h * Higher firmness, lower TSS 2
Fumigation of NO gas followed by cold or - Storage (Cold): 5 ◦ C, 40 days
Feicheng - Lower ethylene production [138]
room temperature storage - Storage (Room temperature) - Lower activity of LOX and 1-aminocyclopropane-1-carboxylic
: 25 ◦ C, 7 days acid (ACC) oxidase

- Improvement in the stability of fruit quality

- Treatment conditions * Lower respiratory rate and ethylene production
Fumigation of NO gas followed by room * 5 µL/L, 7 min * Higher activities of the antioxidant enzymes (CAT,
Rojo Rito [137]
temperature storage SOD)
- Storage: 20 ◦ C, 14 days
- No effects to TSS 2 , TA 3 , and color
Horticulturae 2023, 9, 315 26 of 41

Table 4. Cont.

Method Cultivar Method and Treatment Conditions Results and Implications Reference

- Treatment conditions
Fumigation of NO gas followed by room * 10 µL/L, 3 h - Mode of action for the effects of NO gas fumigation was
Xiahui no.5 revealed by the proteomic analysis [140]
temperature storage
- Storage: 20–25 ◦ C, 1
80–90% RH , 5 days

- Treatment conditions - Suppression of brown rot and Rhizopus rot diseases

Fumigation of EO followed by room * Peppermint and sweet basil EOs, 1–4 mL/box - Phytotoxicity of fruits with the treatment of a high
Early grand [141]
temperature storage concentration of EO (4 mL/box)
- Storage: 27 ◦ C, 22 days

- Singular and combined treatment

* Syringa EO microencapsulation (SEOM): attachment - Improvement in the stability of fruit quality: Delaying fruit
Exposure to volatile EO (microencapsulated) of 1.5 g powder sealed in non-woven bag (5 cm × 5 ripening, decrease of ethylene production, higher firmness
and/or 1-methylcyclopropene (1-MCP) Yanhong cm) with holes to the headspace inside fruit packages - Synergistic effects by the combined treatments of SEOM with [142]
followed by cold storage * *1-MCP: 1 µL/L, 20 h 1-MCP
- Storage: 1 ± 0.5 ◦ C, 90% RH 1 , 35 days

- Singular treatment
* 1-MCP: Fumigating, 5 µL/L, 24 h - Improvement in the stability of fruit quality
* Storage: 0 ◦ C, 45 days * Decrease of MDA content and ethylene emission
Fumigation of 1-MCP followed by cold - Combined treatment * Higher firmness, SSC 6 , and TA 3
storage with the flow microcirculation of Jinqiuhong * 1-MCP: Fumigating, 5 µL/L, 24 h * Color retention [143]
ozone (O3 ) * Storage with O3 flow microcirculation - Decrease of microbial deterioration and damage
: O3 0.08 ppm, 0 ◦ C, 45 days * Reduction of decay rate, increase of PPO resistance

- Post-storage: 20 ± 0.1 ◦ C, 1
95% RH , 10 days

1 RH: relative humidity; 2 TSS: total soluble solid; 3 TA: total (titratable) acidity; 4 CI: chilling injury; 5 ROS: reactive oxygen species; 6 SSC: soluble solid content.
Horticulturae 2023, 9, 315 27 of 41

4.3. Biological Treatments

Fungal diseases in peach fruits are usually caused by latent infection via wounds dur-
ing handling in fields, processing, and storage [144,145]. Antagonists are microorganisms
that control pathogens by colonization on fruit surfaces or flesh exposed by the wound (i.e.,
competitive exclusion) and resource competition for nutrients [146]. Moreover, previous
research regarding the inoculation of antagonists on peaches also showed an increase in
the activities of antioxidant enzymes (e.g., APX, CAT, PAL, POD, and SOD) as an indirect
effect of post-harvest treatments [147,148]. Since the antagonists used on fruits have been
validated as safe for consumption, there is no concern about the residue, which is not the
case for the chemical treatment technologies using toxic fungicides [149]. Major examples
of antagonists reported as applicable for peach fruits are as follows: Pichia caribbica [147], B.
subtilis [148], Cryptococcus laurentii [150], and Aureobasidium pullulans [151]. The function of
antagonism can be diverse according to the determinant factors including region, cultivar,
and environment (Table 5).
Rapid growth on peaches can be a key characteristic of antagonists from the per-
spective of colonization competition. Xu et al. [147] revealed the competitive growth of
a fast-growing antagonist (P. caribbica) against a fungal pathogen (R. stolonifera), which
can also grow rapidly in the flesh of peaches by penetration of the wound, and showed
the effects of biocontrol capability by decreasing the level of fungal decay (lower disease
incidence and lesion diameter) during room temperature storage.
The identification of post-harvest pathogens isolated from stone fruits and the sub-
sequent characterization of their ability to cause decay in fresh fruit can indicate the
microbial strain that should be targeted for control [152,153]. Zhang et al. [148] isolated
peach-decaying fungal strains (Alternaria tenuis and B. cinerea, which showed 100% and
92.33% disease incidence for fresh peach fruits, respectively) and revealed competitive
inhibition as the antagonistic mechanism of B. subtilis by co-culture with those pathogens
on peach wounds under room temperature storage. This study also showed that ca. 7 log
CFU/mL antagonist suspension achieved the highest inhibitory effects against pathogenic
fungal growth compared to other concentrations (from 5 to 9 log CFU/mL), suggesting the
importance of the specific concentration optimized for each type of antagonist.
Since the changes in physicochemical and/or organoleptic characteristics (e.g., odor,
flavor, and color) from the results of the use of synthetic fungicides on fruits have been
regarded as one of the major obstacles for the commercialization of chemical post-harvest
treatment technologies, antagonists can be an alternative that does not result in quality
changes after treatment [146,154]. Zhang et al. [150] showed the broad spectrum of treat-
ment concentration-dependent inhibitory effects of C. laurentii (6–9 log CFU/mL) against
fungal decay, such as gray mold (B. cinerea), blue mold (P. expansum), and Rhizopus rot (R.
stolonifer) during room temperature storage (at 25 ◦ C up to 5 days) without changes in other
undesirable quality parameters.
Although most co-culture growth experiments of antagonist pathogens have been
conducted at room temperature within a few days due to the limited growth range of
the antagonist and short shelf life of peach fruits [147,148,150], antagonists active at low
temperatures are also needed for the preservation of fruits during long-term storage
under refrigeration conditions. A. pullulans is a well-known cold-tolerant antagonist that
can be used to control fungi decaying the fruit during cold storage [155–158]. Zhang
et al. [151] demonstrated the antifungal effects of A. pullulans PL 5 treatment of peach fruit
in regards to the reduction of M. laxa incidence and the diameter of decay and showed
that the mechanisms were both direct (competition during the co-culture of antagonist
pathogens) and indirect (activation of enzymes governing the host defense (chitinase and
β-1,3-glucanase)).
Horticulturae 2023, 9, 315 28 of 41

Table 5. Post-harvest treatments of biological technologies on peach fruits.

Method Cultivar Treatment Conditions Results and Implications Reference

- Treatment conditions - Improvement of antioxidant activity

* Antagonist: Pichia carbic (spot-inoculation, 30 µL, 6–9 * Activation of peroxidase (POD), catalase (CAT), and
Inoculation of antagonist on peach followed log cells/mL) phenylalanine ammonia-lyase (PAL)
Dajiubao * Target fungi: Rhizopus stolonifer (spot-inoculation, 30 [147]
by room temperature storage - Decrease of microbial deterioration and damage
µL, ca. 4 log spore/mL)
* Decrease of disease incidence and lesion diameter
- Storage: 20 ◦ C, 95% RH 1 , 4 days

- Treatment conditions
- Improvement of antioxidant activity
* Antagonist: Bacillus subtilis JK-14 (spot-inoculation,
30 µL, 5–9 log CFU/mL) * Activation of superoxide dismutase (SOD), POD,
Inoculation of antagonist on peach followed * Target fungi: Alternaria tenuis (spot-inoculation, 15 µL CAT, and ascorbate peroxidase (APX)
Baifeng [148]
by room temperature storage each, 6 log CFU/mL, Botrytis cinerea - Decrease of microbial deterioration and damage
(spot-inoculation, 15 µL each, log CFU/mL) * Decrease of disease incidence and lesion diameter
- Storage: 20 ◦ C, 85% RH 1 , 5 days

- Treatment conditions
* Antagonist: Cryptococcus laurentii (spot-inoculation,
30 µL, 6–9 log CFU/mL) - Decrease of microbial deterioration and damage
* Target fungi: B. cinerea (spot-inoculation, 15 µL, 5 log * Decrease of disease incidence
Inoculation of antagonist on peach followed spore/mL), Penicillium expansum (spot-inoculation, 15
Baihua - No effects on other quality parameters [150]
by room temperature storage µL, ca. 4 log spore/mL), R. stolonifer
(spot-inoculation, 15 µL, ca. 4 log spore/mL) * Firmness, ascorbate content, TA 2
- Storage: 25 ◦ C for 4 days (B. cinerea, P. expansum), 25 ◦ C for 5
days (R. stolonifer)

- Treatment conditions - Decrease of microbial deterioration and damage

* Antagonist: Aureobasidium pullulans PL5 (dipping * Decrease of disease incidence, lesion diameter
Inoculation of antagonist on peach followed inoculation, 8 log cell/mL, 1 min)
Redhaven - Improvement of pathogenesis-related proteins linked to the [151]
by cold storage * Target fungi: Monilinia laxa (spray inoculation, 1 mL,
host defense
4 log conidia/mL)
* Activation of chitinase and β-1,3-glucanase
- Storage: 1 ◦ C, 95% RH 1 , 28 days

1 RH: relative humidity; 2 TA: total (titratable) acidity.

Horticulturae 2023, 9, 315 29 of 41

4.4. Combined Treatments

Many previous studies conducted to enhance the quality of peach fruits combined
multiple post-harvest technologies categorized as physical (e.g., heating and irradiation),
chemical (e.g., dipping in treatment solution and spraying or fumigating solution), and
biological (e.g., co-culture of antagonistic organisms) treatments. The major aim of the
combination of technologies is to achieve a greater effect than the sum of individual
technologies (i.e., synergistic effect) and/or to complement the limitation of each technology
with perspectives on the intervention mechanism of quality changes in peaches. The
findings of studies on the development of the combination method of technologies and the
establishment of optimal treatment conditions highlight novel effects that were unexpected
based on the results of the application of individual technologies for peach fruit post-
harvest treatments. Since the combination of technologies results in antagonistic effects
(i.e., a lower effect than the sum of individual technologies or the inhibition of the intended
effects by the interaction among the combined technologies), the exploration of adequate
treatment concentrations is also needed to avoid inefficiency. Table 6 summarizes the
findings of the previous research focused on combined treatment methods applied for the
post-harvest quality and safety control of peach fruits.

4.4.1. Combination of Physical and Chemical Treatment

Since there are concerns about the toxicity and environmental pollution derived from
the excessive use of chemical post-harvest agents, the combination of physical treatment
technologies has been attempted to reduce the treatment concentration of agents [159,160].
In the case of peach fruits, salicylic acid treatment combined with ultrasonication or ther-
mal pre-treatment are representative examples [161,162]. Ultrasound treatment for fresh
fruits and vegetables has been used to clean surfaces and to inactivate microbial contam-
inants with the disruption of cells; however, direct antimicrobial efficacy is likely to be
insufficient to achieve a reduction in fungal decay, and thus the combination with other
decontamination technology is generally needed [163]. Yang et al. [161] combined salicylic
acid (dipping in solution (0.05 mM)) and ultrasonication at 20 ◦ C for 10 min followed
by room temperature storage (20 ◦ C, 6 days) of peach fruits infected with blue mold (P.
expansum); all treatments (both singular and combined) did not affect the quality factors
(weight loss, firmness, TSS, and vitamin C content), but the beneficial effects from the
singular salicylic acid treatment (activation of enzymes governing antioxidant and host
defense mechanisms and decrease of the fungal decay) were synergistically improved by
the combined treatment with ultrasonication; although, individual ultrasound treatment
did not influence those effects. The sequential approach of thermal pre-treatment before
the application of chemical agents has also been adopted as a combined treatment strategy
for post-harvest preservative processing [164]. According to the research by Cao et al. [162],
exposure to heat (hot air at 38 ◦ C, 12 h) prior to salicylic acid treatment (dipping in salicylic
acid 1 mM at 20 ◦ C for 5 min) is expected to induce the expression of heat-shock proteins
related to fruit tolerance, and thus showed desirable effects on internal browning as an
indicator of cold stress response capability, antioxidant activities, and polyamine levels
during long-term cold storage (0 ◦ C, 35 days).
Radical irradiation has been widely used for the extension of the shelf life of vegetables
and fruits due to its cold nature and strong decontamination effects [165]. Edible coating
with polysaccharides can preserve the quality parameters of fruits by inhibiting fungal
growth on the coated surface [166–168]. As reported by Hussain et al. [169], gamma
irradiation (1.2 kGy) of peach fruits coated with carboxymethyl cellulose (1%) can improve
the effects on the inhibition of fungal infection, the prevention of quality changes (firmness,
TSS, and TA), and the delay of ripening or senescence during cold storage.

4.4.2. Combination of Physical and Biological Treatment

Although it is difficult to predict the effect of commonly used fungicide products
through only physical or biological treatment, the support of antifungal biocontrol agents
Horticulturae 2023, 9, 315 30 of 41

in combination with physical treatments (e.g., heat treatment) has achieved desirable
pathogen control efficacies for fruits [170,171]. Biocontrol agents are generally ineffective
against the micro-organisms infecting fruits prior to the application of those agents, and
thus the complementation of the antifungal efficacy by support from physical treatment
is needed. Microwave treatment enables the rapid heating of food products to efficiently
inactivate the microbial cause of decay of fruits and vegetables [172,173]. Zhang et al. [170]
suggested the biocontrol strategy of the inoculation of an antagonist (C. laurentii) into
infected peach fruit (R. stolonifera) after microwave heating (2450 MHz, 2 min) for fungal
inactivation, and the persistent protective effects of the antagonist were also validated by
the decrease in infected wounds on fruits without quality changes (firmness, TA, and TSS).
Zhang et al. [171] also showed that the exposure of peach fruit to heated air (37 ◦ C, 48 h)
before the application of the antagonist (C. laurentii) also ensured its active competitive
effects against fungal contaminants causing the decay of peaches (decrease in the ratio of
infected wounds by 22.5% and 5% for the infectious disease caused by P. expansum and R.
stolonifer, respectively) without remarkable differences in physicochemical characteristics
of fruits (TSS, TA, and vitamin C content).

4.4.3. Combination of Chemical and Biological Treatment

Since biological control using antagonists is generally not as effective as the direct
application of disinfectants from the perspective of immediate killing effects, antagonists
for peaches have been combined with organic or inorganic additives (e.g., antifungal effects
and/or plant growth regulators) to complement the limited effects of antagonists [37,174].
Zhang et al. [174] used antagonist suspensions (Rhodotorula glutinis; 8 log cells/mL)
amended with salicylic acid (100 µg/mL) for 30 s of dipping treatment before artificial
inoculation with gray mold (Botrytis cinerea) on peaches, and showed that limited effects of
individual treatment with salicylic acid or an antagonist (R. glutinis) in reducing the lesion
diameter could be significantly improved by the combined treatment without impairing
the quality of wounded fruits (TSS, TA, ascorbic acid content, and firmness) [174].
The antagonist Bacillus spp. can protect the fruit surface by producing extracellular
polysaccharides (e.g., glycocalyx) in biofilm as a physical barrier against the colonization
of fungal pathogens, and this biofilm-forming ability can be supported by Eos. Arrebola
et al. [37] designed combined treatment technologies between post-harvest spraying of an
antagonist on peach fruits followed by packaging with the delivery system of EOs (e.g.,
thymol and lemongrass oil) and showed a broad spectrum of the disease control, including
gray mold (B. cinerea), blue mold (P. expansum), and Rhizopus rot (R. stolonifer). This
combined treatment technology can also reduce the burden of the application concentration
of EO, which is known to have a potential phytotoxic effect causing the browning of fruits
and an unpleasant odor [37,113].
Horticulturae 2023, 9, 315 31 of 41

Table 6. Post-harvest treatments of combined technologies (physical, chemical, and biological technologies) on peach fruits.

Method Cultivar Treatment Conditions Results and Implications Reference

- Inoculation - Improvement of antioxidant activity

* Penicillium expansum (spot-inoculation, 15 µL, 5 log * Activation of phenylalanine ammonia-lyase (PAL),
spores/mL) polyphenol oxidase (PPO), and peroxidase (POD)
- Singular treatment - Activation of the production of pathogenesis-related proteins
Dipping in salicylic acid solution with * Dipping: salicylic acid 0.05 mM, 20 ◦ C, 10 min linked to the host defense
ultrasound treatment followed by room Baifeng * Ultrasonication (40 kHz, 350 W): 20 ◦ C, 10 min * Activation of chitinase and β-1,3-glucanase [161]
temperature storage
- Combined treatment - Decrease of microbial deterioration and damage
* Dipping in salicylic acid (0.05 mM) + ultrasonication * Decrease of lesion diameter and disease incidence
(40 kHz, 350 W) at 20 ◦ C for 10 min - Synergistic effects by the combined treatments of salicylic acid
- Storage: 20 ◦ C, 95% RH 1 , 6 days with ultrasonication

- Improvement in the stability of fruit quality

* Decrease of internal browning
- Singular treatment - Improvement of antioxidant activity
* HA: 38 ◦ C, 12 h * Activation of superoxide dismutase (SOD), catalase
* Dipping: salicylic acid 1 mM, 20 ◦ C, 5 min (CAT), ascorbate peroxidase (APX), and glutathione
Dipping in salicylic acid solution after hot air reductase (GR)
Baifeng - Combined treatment [162]
treatment followed by the cold storage * Decreased activity of lipoxygenase (LOX)
* Dipping in salicylic acid 1 mM at 20 ◦ C for 5 min after
- Increase of polyamine levels
hot air (38 ◦ C, 12 h) treatment
* Higher levels of putrescine, spermidine, and
- Storage: 0 ◦ C, 90–95% RH 1 , 35 days
spermine
- Synergistic effects by the combined treatments of salicylic acid
with hot air

- Singular treatment
* Carboxymethyl cellulose coating: 0.5–1.0% - Improvement in the stability of fruit quality
* Irradiation: γ-ray, 1.2 kGy * Lower weight loss
Gamma irradiation treatment after - Combined treatment * Decrease of ascorbic acid, TSS 2 , and total sugar
carboxymethyl cellulose coating followed by - - Irradiation (γ-ray, 1.2 kGy) after the carboxymethyl cellulose * Delay of ripening, senescence, and respiration rate [169]
cold or room temperature storage coating (0.5–1.0%) - Improvement of antioxidant activity
* Storage (cold): 3 ± 1 ◦ C, RH 1 80%, 35 days * Increase of total phenol compounds
- Storage (room temperature): 25 ± 2 ◦ C, RH 1 70%, 15 days
Horticulturae 2023, 9, 315 32 of 41

Table 6. Cont.

Method Cultivar Treatment Conditions Results and Implications Reference

- Treatment conditions for the experiments of the application of

antagonist after the microwave treatment of peach inoculated - Decrease of microbial deterioration and damage
with pathogen * Decrease of infected wounds
* Target fungi: Rhizopus stolonifer (spot-inoculation, 20 - Antifungal effects
Inoculation of antagonist after microwave µL, 5 log spore/mL)
Baihua * Effective against R. stolonifer [170]
treatment * Microwave treatment: 2450 MHz, 2 min
* Antagonist: Cryptococcus laurentii (spot-inoculation, - No effects to other quality parameters
30 µL, 8 log CFU/mL) * Firmness, TSS 2 , TA 3 , vitamin C content
* Storage: 25 ◦ C, 4 days

- Treatment conditions for the experiments of the application of

antagonist after the hot air treatment of peach inoculated with - Improvement in the stability of fruit quality
pathogen
* Higher firmness
* Target fungi: R. stolonifer (spot-inoculation, 20 µL, 5
- Decrease of microbial deterioration and damage
Inoculation of antagonist after hot air log spore/mL), P. expansum (spot-inoculation, 20 µL,
Baihua ca. 4 log spore/mL) * Decrease of infected wounds [171]
treatment
* HA: 37 ◦ C, 48 h - No effects to other quality parameters
* Antagonist: C. laurentii (spot-inoculation, 30 µL, 8 log
CFU/mL) * Firmness, TSS 2 , TA 3 , vitamin C content
- Storage: 25 ◦ C, 7 days

- Treatment conditions for the experiments of the application of

antagonist and salicylic acid treatment before inoculation of
pathogen - Decrease of microbial deterioration and damage
* Antagonist: Rhodotorula glutinis (spot-inoculation, 30 * Decrease of decay incidence, lesion diameter
Treatment of salicylic acid with antagonist Jiubao µL, 8 log CFU/mL) - No effects to other quality parameters [174]
* SA: 100 µg mL (spot-inoculation, 30 µL)
* Target fungi: B. cinerea (spot-inoculation, 30 µL, 8 log * TSS 2 , TA 3 , ascorbic acid, firmness
CFU/mL)
- Storage: 20 ◦ C, 95% RH 1 , 5 days
Horticulturae 2023, 9, 315 33 of 41

Table 6. Cont.

Method Cultivar Treatment Conditions Results and Implications Reference

- Singular treatment
* Antagonist: Bacillus amyloliquefaciens PPCB004 (spray
inoculation, 10–15 min, 8 log CFU/mL)
* EO: lemongrass or thyme (75 µL impregnated for
each package bag)
- Combined treatment

Treatment of essential oils (EOs) with * Packaging of peach fruits sprayed with antagonist (B. - Decrease of microbial deterioration and damage
antagonist followed by room temperature and Transvaal amyloliquefaciens PPCB004; spray inoculation, 10–15 [37]
min, 8 log CFU/ mL) in bag impregnated with EO * Decrease of the incidence and severity of the diseases
cold storage
(lemongrass or thyme; 75 µL)
- Inoculation of target pathogen after the treatment: B. cinerea
(spraying for 10–15 min, 6 log spore/mL), P. expansum
(spraying for 10–15 min, 6 log spore/mL), R. stolonifer
(spraying for 10–15 min, 6 log spore/mL)
- Storage (room temperature): 25 ◦ C, 5 days
- Storage (cold): 4 ◦ C, 90% RH 1 , 14 days

1 RH: relative humidity; 2 TSS: total soluble solid; 3 TA: total (titratable) acidity.
Horticulturae 2023, 9, 315 34 of 41

5. Conclusions
This review provides comprehensive information based on the findings from studies
regarding pre- and post-harvest treatment strategies optimized for peach fruits to extend
the durable intake. Since peaches are vulnerable to environmental stresses under room
temperature, most relevant studies aim to ensure fruit quality during long-term cold
storage. Recent research has mainly focused on the development of new technologies
and the design of novel combined treatment, whereas the in-depth study of pre- and post-
harvest processes previously reported as applicable for stone fruits to optimize operational
conditions for peaches should also be consistently conducted due to the diversity in the
efficacies of treatment methods according to various determinant factors (e.g., a cultivar of
fruits, processing environments, storage temperature, and time). Major implications from
the analysis of the literature can be summarized as follows: (1) the discovery of side-effects
from the overuse of treatment agents (chemical and biological technologies) or severe
treatment conditions (physical technology) highlights the importance of the determination
of the adequate criteria for the limitation of operational conditions; (2) since the result of the
combined treatment is generally unexpectable (e.g., synergistic, additive, and antagonistic
effects), the establishment of strategies which can harmonize both the efficacy and efficiency
should be followed; and (3) pre-harvest treatment technologies generally aim to achieve
sustentative effects allowing the improvement in the stability of fruit quality during the
long-term cold storage, and thus the combined (sequential) treatment with subsequent post-
harvest treatment is expected to enhance overall efficacies. This focused review suggests
practical information for the design of advanced pre- and post-harvest treatments for
peach fruits based on insights into advantages and disadvantages of currently reported
technologies. As a future perspective on the research area in peaches, the quality control
system based on the technologies in the Fourth Industrial Revolution era is expected to
be integrated into pre- and post-harvest treatment strategies for peach fruit by sensing
the fruit quality, strict pre-harvest quality control in smart farms, and web cloud-based
precise quality management during the storage and/or distribution. The sensor-based
analysis of the changes in the fruit quality factor can be a promising countermeasure
for undesirable antagonistic effects derived from the combined treatment of pre- and
post-harvest technologies described in this study.

Author Contributions: Contributions for each author are as follows: conceptualization, H.W.K.
and T.J.C.; investigation, J.S.S. (Jin Song Shin), H.S.P., K.W.L. and H.Y.H.; writing—original draft
preparation, J.S.S. (Jin Song Shin), H.S.P. and J.S.S. (Ji Seop Song); writing—review and editing, H.W.K.
and T.J.C.; supervision, T.J.C.; funding acquisition, T.J.C. All authors have read and agreed to the
published version of the manuscript.
Funding: This research was supported by the Regulatory Science Research Grant of the Ministry of
Food and Drug Safety (MFDS, Republic of Korea). This research was also funded by BK21 FOUR
(Fostering Outstanding Universities for Research) of the Ministry of Education (MOE, Republic of
Korea) and National Research Foundation of Korea (NRF, Republic of Korea).
Data Availability Statement: No new data were created or analyzed in this study. Data sharing is
not applicable to this article.
Acknowledgments: The authors also thank the Department of Food Regulatory Science and the
Department of Food and Biotechnology of Korea University for access to equipment and facilities.
Conflicts of Interest: The authors declare no conflict of interest. The funders had no role in the design
of the study; in the collection, analyses, or interpretation of data; in the writing of the manuscript; or
in the decision to publish the results.

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