Postharvest Biology and Technology 173 (2021) 111429

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Postharvest Biology and Technology

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

The influence of pre- and postharvest 1-MCP application and oxygen

regimes on textural properties, cell wall metabolism, and physiological
disorders of late-harvest ‘Bartlett’ pears
Meng Li a, 1, Huanhuan Zhi a, 1, Yu Dong b, *
a
College of Food and Bioengineering, Zhengzhou University of Light Industry, Zhengzhou, Henan, 450002, China
b
Department of Horticulture, Oregon State University, Mid-Columbia Agricultural Research and Extension Center, 3005 Experiment Station Drive, Hood River, OR,
97031, USA

A R T I C L E I N F O A B S T R A C T

Keywords: Late-harvest (LH) pears are prone to postharvest disorders and eating-quality deterioration in storage and
Pyrus communis retailing. The purpose of this work was to evaluate the effects of pre- and post-harvest 1-methylcyclopropene (1-
Late harvest MCP, Harvista (H) and SmartFresh (SF)) on LH ‘Bartlett’ pears under the various O2 regimes. Spraying 320 μL L− 1
1-Methylcyclopropene
H delayed fruit maturation and suppressed ethylene production rate (EPR) when pears were harvested at 70.51 N
O2 regimes
Melting texture
(LH, whereas commercial harvest (CH) at 75.04 N). However, the H-treated LH fruit had 100 % decay after
Physiological disorders 5 months of regular-air (RA) storage. The 0.15 μL L− 1 SF and SF + 160 μL L− 1 H extended melting texture life of
LH fruit to 5 months with high levels of water-soluble polyuronides (WSP) and CDTA-soluble polyuronides (CSP)
and activities of pectin methylesterase (PME), pectate lyase (PL), and α-arabinofuranosidase (α-ARF). Raising H
application concentration from 160 to 320 μL L− 1 in H + SF treatments resulted in blockage of ripening capacity.
Decreasing O2 concentration from 2 to 1 % did not impact LH pears’ ripening, but effectively curtailed the
development of melting texture in H-treated LH fruit by suppressing EPR, degradation of pectin polyuronides,
and activities of PL and β-galactosidase (β-GAL). Furthermore, applying SF in H-treated LH pears stored in 1 or 2
% O2 resulted in the loss of ripening capacity. Results indicated that 160 μL L− 1 H + SF and H at 160− 320 μL L− 1
extended the melting period and controlled physiological disorders in LH ‘Bartlett’ pears for 5 and 7 months of
storage in RA and 2 % O2, respectively.

1. Introduction
‘Bartlett’ pear is the second largest European pear cultivar (Pyrus
communis L.) grown in Pacific Northwest (PNW) region of the United
States. The recommended harvest maturity for ‘Bartlett’ pear, as indi­
cated by flesh firmness (FF), is FF between 83.6 to 75.6 N (Wang et al.,
2016). However, labor shortages and warmer growing seasons, espe­
cially in high elevation orchards, result in a large amount of pears being
harvested at lower FF values. Additionally, ‘Bartlett’ pears have a short
storage life and a rapid ripening process due in part to high ethylene
generated (Wang et al., 2016). Therefore, late harvest (LH) pears are
more susceptible to loss due to reduced FF and yellowing, are susceptible
to postharvest disorders (i.e. decay, senescent scald, and senescent core
breakdown), and rapid softening in storage and on the retail shelf
(Villalobos-Acuña et al., 2011; Wang and Sugar, 2015). This inspired us
to develop strategies to extend storage life of LH ‘Bartlett’ pears.
Retention of desirable eating quality in pears delivered to market could
give higher returns to growers.
Ethylene continues to be a major factor influencing softening be­
haviors of European pears (Villalobos-Acuña and Mitcham, 2008).
Well-ripened ‘Bartlett’ pears develop a buttery (melting) mouth feel
with characteristics of sweetness, juiciness, and full flavor (Raffo et al.,
2010). However, the accumulated ethylene contributes to increase
ethylene-related responses including ripening, senescence, and physio­
logical disorders (Villalobos-Acuña and Mitcham, 2008). 1-Methylcyclo­
propene (1-MCP) was used to bind ethylene receptors and counteracted

* Corresponding author at: Department of Horticulture, Oregon State University, Mid-Columbia Agricultural Research and Extension Center, Hood River, OR,
97031, USA.
E-mail address: dongyu@oregonstate.edu (Y. Dong).
1
These authors contributed equally to this work.

https://doi.org/10.1016/j.postharvbio.2020.111429
Received 26 May 2020; Received in revised form 12 November 2020; Accepted 29 November 2020
Available online 2 December 2020
0925-5214/© 2020 Elsevier B.V. All rights reserved.
M. Li et al.
ethylene biosynthesis; as a result, the occurrence of postharvest disor­
ders and senescence processes were delayed (Watkins, 2006). Investi­
gation into the commercial use of postharvest 1-MCP (SmartFresh™, SF)
applications on ‘Bartlett’ pears prior to storage/shipping has been con­
ducted (Ekman et al., 2004; Trinchero et al., 2004; Acuna et al., 2011;
Wang et al., 2016). SF prevents softening and fruit fail to develop a
desirable texture after being subjected to warmer temperature (Acuna
et al., 2011; Wang and Sugar, 2015; Zhi et al., 2019). Harvista™ (H,
1-MCP) is designed as a pre-harvest spray to respond to harvest labor
shortages and prevent fruit drop (Defilippi et al., 2010; DeEll and
Ehsani-Moghaddam, 2010; Villalobos-Acuña et al., 2010; Sakaldas et al.,
2016; Escribano et al., 2017). While H did not interfere with softening,
its efficacy in reducing disorders began to weaken after excessive storage
(Li et al., 2020). Recent research indicated a tendency to use a
pre-harvest H spray combined with a postharvest SF treatment to
manipulate ripening process in apple and peach (Varanasi et al., 2013;
Yoo et al., 2019). However, the response of pears to 1-MCP was affected
by application concentration, timing, harvest maturity, storage tem­
perature, storage condition, and cultivars (Ekman et al., 2004; Trinchero
et al., 2004; Acuna et al., 2011; DeEll and Ehsani-Moghaddam, 2011;
Villalobos-Acuña et al., 2011; Wang and Sugar, 2015; Escribano et al.,
2017; Li et al., 2020). Therefore, a successful commercial strategy for
retaining softening behavior of PNW pears produced in the PNW region
must optimize pre- and post-harvest 1-MCP applications.
Controlled atmosphere (CA) has been recommended to extend pears’
storage life. The commercial standard CA regime for ‘Bartlett’ pears in
Oregon’s Hood River Valley is 2–2.5 % O2 with 0.8–1 % CO2 at − 1.1 ◦ C
(Hansen and Mellenthin, 1979). Under these conditions, the lifespan of
dessert-quality ‘Bartlett’ can be extended for up to 6 months (Bai et al.,
2006). Reducing O2 concentration had greater effects on extending
storage life and retaining the capacity for ripening. But, extremely low
O2 levels (i.e. ≤ 0.5 %) caused severe disorders (i.e. pithy core brown,
black speck) with loss of ripening capacity and off flavors (Chen, 2004;
Li et al., 2019). Additionally, various O2 regimes could be imposed to
regulate the pears’ softening behaviors by influencing the depolymer­
ization and solubilization of pectic polyuronides and activities of cell
wall-modifying enzymes (Chen and Borgic, 1985; Li et al., 2019). Little
information is available to assess the changes in textural properties and
cell wall metabolism of LH ‘Bartlett’ pears in response to pre- and
post-harvest 1-MCP treatment under the various O2 regimes.
In the PNW region, 0.3 μL L− 1 SF effectively inhibited ethylene-
induced ripening of ‘Bartlett’ pears when fruit were harvested at 83.6-
75.6 N (Bai et al., 2006; Wang and Sugar, 2015; Zhi et al., 2019).
Reducing the dose of SF resulted a part of ethylene receptors without
being occupied and permitted initiation of the ripening processes (Bai
et al., 2009; Wang and Sugar, 2015). Therefore, the first objective of this
study was to evaluate the effects of SF at 0.15 μL L− 1 combined with H at
160 and 320 μL L− 1 on textural attributes, pectic polyuronides, cell
wall-modifying enzymes, ethylene production, and physiological disor­
ders of LH ‘Bartlett’ pears under regular-air (RA) condition. The second
objective was to investigate the effect of 1 and 2 % O2 on LH pears
treated with H alone or in combination with SF.
2. Materials and methods
2.1. Fruit and experimental designs
‘Bartlett’ trees were selected in a commercial orchard in Parkdale,
OR, USA (45.30 ◦ N, 121.34 ◦ W, elevation 576 m). Trees were grafted on
OHxF97 rootstock. Orchard spacing was 4.5 m × 4.5 m. All trees
received standard fertilizers, herbicides, and pesticides.
Experiment 1: Harvista™ (1-MCP; AgroFresh, Spring House, PA,
USA) at concentrations of 160 and 320 (manufacture recommended
concentration) μL L− 1 plus 0.05 % (v/v) Xiameter OFX-0309 silicone
surfactant (Norac Concepts Inc., Guelph, ON, Canada) was applied 10
d before the projected commercial harvest (CH) date. A turbomist
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Postharvest Biology and Technology 173 (2021) 111429
sprayer (Slimline Manufacturing Ltd., Penticton, BC, Canada) was used
to achieve uniform, complete coverage of the entire canopy. Fruit were
harvested on the commercial harvest date (CH) and FF of control fruit
was 75.04 ± 0.39 N. Pears were then harvested after 10 d (late harvest,
LH) and FF of control fruit was 70.51 ± 1.26 N. Fruit were selected for
their freedom from any visible signs of damage or fungal infection, then
packed in wooden boxes (~80 fruit per box) with standard polyethylene
(PE) bags. In all there were 12 treatments (control CH, SF CH, 160 μL
L− 1 Harvista CH (160 H CH), 160 μL L− 1 Harvista + SF CH (160 H + SF
CH), 320 μL L− 1 Harvista H1 (320 H CH), 320 μL L− 1 Harvista + SF H1
(320 H + SF CH), control LH, SF LH, 160 μL L− 1 Harvista H2 (160 H LH),
160 μL L− 1 Harvista + SF LH (160 H + SF LH), 320 μL L− 1 Harvista LH
(320 H LH), 320 μL L− 1 Harvista + SF LH (320 H + SF LH)) with 3
replications per treatment and one box as a replicate. The 0.15 μL L− 1 SF
was generated using SmartFresh™ tablet (AgroFresh, Spring House, PA)
according to the application procedures recommended by the manu­
facturer. All pears were stored in RA condition at − 1.1 ◦ C and 95 %
relative humidity (RH). After 3 and 5 months of storage, one box from
each treatment per replicate was transferred to 20 ◦ C and 90 % RH for a
5-d ripening period. The fruit firmness, ethylene production rate, and
decay were evaluated on day 1; ripening capacity, sensory quality,
pectic polyuronides, cell wall-modifying enzymes, and senescence dis­
orders were measured on day 5. Flesh tissue was quick-frozen in liquid
nitrogen (N2), then stored at − 80 ◦ C.
Experiment 2: The 160 and 320 μL L− 1 Harvista™ plus 0.05 % (v/v)
Xiameter OFX-0309 silicone surfactant was applied 10 d before the
harvest date. After selecting and packing, fruit were divided into 8
treatments (2 % O2 control, 2 % O2 160 μL L− 1 Harvista (2 % O2 160 H),
2 % O2 320 μL L− 1 Harvista (2 % O2 320 H), 2 % O2 160 μL L− 1 Har­
vista + SF (2 % O2 160 H + SF), 1 % O2 control, 1 % O2 160 μL L− 1
Harvista (1 % O2 160 H), 1 % O2 320 μL L− 1 Harvista (1 % O2 320 H), 1
% O2 160 μL L− 1 Harvista + SF (1 % O2 160 H + SF)) with 3 replications
per treatment and one box as a replicate. SmartFresh (0.15 μL L− 1) was
applied as described as above. Two boxes of each treatment were loaded
into each of 24 gas-tight cabinets at − 1.1 ◦ C, then sealed and flushed
with N2 generated from a membrane gas generator (CPA-5, Permea, St.
Louis, MO, USA). Each box of fruit per cabinet represented one replicate
of the treatment. CO2 concentrations were maintained at < 0.5 % by
adding hydrated lime (0.5 kg per cabinet). Concentrations of O2 and CO2
were monitored every 2-d using an O2/CO2 analyzer (Storex, Grave­
ndeel, Netherlands). The 2 and 1 % O2 were established within 6 d. After
5 and 7 months of CA storage, one box from each cabinet were trans­
ferred to 20 ◦ C for a 5-d ripening period. CA conditions were re-
established within 3 h by flushing cabinets with N2. Physiological and
biochemical evaluations were performed as described in Experiment 1.
2.2. Evaluations of flesh firmness (FF), ripening capacity, and sensory
quality
FF of 10 fruit from each box was determined at a penetration speed of
10 mm s− 1 using a texture analyzer (GS-14, Güss Manufacturing Ltd.,
Strand, South Africa) with an 8-mm probe. The measurement was per­
formed at two points on opposite sides of the equator of each fruit after
2-mm peel had been removed. Maximum force was recorded. Data were
averaged per 10 fruit and expressed in newtons (N). Ripening capacity
was determined as competency of fruit softening when fruit FF < 24 N;
ten fruit from each box were ripened for 5 d at 20 ◦ C. Sensory quality
(melting/mealy texture score) was evaluated on a 5-point hedonic scale
(Dong et al., 2018). Fruit with highly, moderately, or slightly melting
texture were rated as 5, 4, or 3, respectively, and those rated as
moderately or very firm (i.e., underripe pears) or moderately or very
mealy flesh texture (i.e., overripe pears) were rated as 2, or 1, respec­
tively. Ten taste-testers were oriented to scale anchor points and defi­
nitions before the first evaluation session. An average score of 3 or
higher was defined as commercially acceptable. Each panelist tasted one
small fruit slice from each treatment of 5 fruit.
M. Li et al. Postharvest Biology and Technology 173 (2021) 111429

2.3. Isolation and determination of pectin polyuronides the mixture. One unit of β-GAL or α-ARF activity was defined as 1 mg of
p-nitrophenol released per min at 400 nm on a fresh weight basis.
Flesh samples of 10 fruit from each box were homogenized in 30 mL
of 80 % ethanol using a homogenizer, then boiled for 30 min, as describe
2.5. Measurement of ethylene production rate (EPR)
by Murayama et al. (2002). After centrifugation at 8,000g for 15 min at
20 ◦ C, the residue was treated with 30 mL of ethanol and again centri­
Five fruit from each box were sealed in a 3.8-L jar at 20 ◦ C for 1 h.
fuged. Next, the residue was suspended in acetone, and centrifuged; this
Each jar represented one replicate of the treatment. One mL of gas
process was repeated. An alcohol insoluble residue (AIR) fraction was
sample was withdrawn and injected into a gas chromatograph (GC-8A,
obtained by air drying the residue at 20 ◦ C for 48 h, then weighted. AIR
Shimadzu, Tokyo, Japan) equipped with a flame ionization detector and
were fractionated with distilled water and continuously shaken for 1 h,
a Porapack Q column (80/100 mesh, 3-mm diameter, 2-m long). The
then centrifuged. This procedure was repeated and supernatants were
injection and detector port temperatures were 90 ◦ C and 140 ◦ C,
collected as water-soluble polyuronides (WSP). The residue was then
respectively. The carrier gas was nitrogen at a flow rate of 0.8 mL s− 1.
treated with 50 mM sodium acetate (pH 6.5) containing 50 mM
EPR was expressed as ng kg-1 s− 1 on a fresh weight basis.
cyclohexanetrans-1,2-diaminetetra (CDTA) and shaken for 1 h. This
procedure was repeated, and the resultant supernatants were collected
as CDTA-soluble polyuronides (CSP). Finally, the residue was suspended 2.6. Evaluations of decay and senescence disorders
in 50 mM sodium carbonate containing 10 mM sodium borohydride,
then shaken for 1 h and centrifuged. The two supernatants were After 1 d of storage at 20 ◦ C, decay was evaluated in thirty fruit from
collected as sodium carbonate-soluble polyuronides (SSP). each box and expressed as percentage of fruit with any pathological
Uronic acids were determined as described by Blumenkrantz and lesion in the skin tissue. After 5 d of storage at 20 ◦ C, above fruit were
Asboe-Hansen (1973). Each extracted polyuronides solution (200 μL) evaluated for senescent scald, then cut longitudinally and transversely to
was added to H2SO4 with 75 mM sodium borate. After boiling, 0.15 % assess senescent core breakdown. The incidence of senescence disorders
(w/v) m-phenylphenol was added to the mixture and loaded into a was expressed as percentage of fruit with either senescent scald or se­
96-well plate. Absorbance was measured at 550 nm using a plate reader nescent core breakdown or both regardless of severity (Zhi et al., 2019).
(ELx800, Bio-Tek Instruments Co., Winooski, VT, USA). A standard
calibration curve was constructed using galacturonic acid and the data 2.7. Statistical analysis
were expressed as mg kg− 1 on a AIR basis.
Experiments were performed using a completely randomized design.
2.4. Assessing of cell wall-modifying enzymes The data were subjected to analysis of variance (ANOVA) using IBM
SPSS Statistics (IBM Co., Armonk, NY, USA). When appropriate, means
Flesh samples of 10 fruit from each box were ground into powder were separated by Fisher’s protected least significant difference (LSD)
with liquid N2, then stored at − 80 ◦ C. test at p < 0.05. The data of sensory quality was analyzed using Kruskal-
Polygalacturonase (PG) was measured as described by Gross (1982). Wallis one-way ANOVA and Fisher’s LSD test was used for pairwise
Powder samples were homogenized in 0.3 M NaCl and centrifuged at 10, comparison.
000g for 15 min at 4 ◦ C. The supernatant (50 μL) was added to a solution
of 0.1 % (w/v) polygalacturonic acid and 50 mM acetate buffer (pH 4.5), 3. Results
then incubated at 37 ◦ C for 2 h. After adding 0.1 M borate buffer (pH
9.0) and 1 % (w/v) 2-cyanoacetamide, the mixture was boiled for 3.1. Effect of H on fruit maturity and EPR of ‘Bartlett’ pears
10 min. One unit of PG activity was defined as 1 mg of galacturonic acid
released per min at 276 nm absorbance using a UV/visible spectro­ In Exp 1, the control LH ‘Bartlett’ pears had lower FF and higher EPR
photometer (Ultrospec 3100 Pro, Biochrom Ltd, Cambridge, UK). Data compared to the control CH fruit (Table 1). Pre-harvest H sprays resulted
were expressed on a fresh weight basis. in the retention of FF and reduction of EPR in CH and LH fruit. The effect
Pectin methylesterase (PME) was measured as described by Basak of H treatments on FF and EPR in Exp. 2 was similar to the effect
and Ramaswamy (1996). Powder samples were homogenized in 0.3 M observed in Exp. 1. Increasing H application concentration from 160 to
NaCl and centrifuged. The supernatant was collected and adjusted to pH 320 μL L− 1 had great effects on delaying fruit maturity and reducing EPR
8.1 with 0.05 M NaOH, then 4 mL of the supernatant was added to in LH fruit.
36 mL of 0.5 % (w/v) pectin adjusted to pH 8.1 and incubated at 37 ◦ C
for 1 h. After the mixture was titrated to pH 8.1 using a titration system
Table 1
(DL15, Mettler-Toledo Inc., Columbus, OH, USA), the consumption Effect of Harvista (H) sprays at 160 and 320 μL L− 1 on flesh firmness (FF) and
volume of NaOH solution was recorded. One unit of PME activity was ethylene production rate (EPR) of ‘Bartlett’ pears in Exp. 1 and 2. Measurements
defined as 1 mM of ester hydrolyzed released per min on a fresh weight were made on day of harvest.
basis. 1 − 1
Treatments FF (N) EPR (ng kg− s )
Pectate lyase (PL) was measured as described by Chourasia et al.
(2006). Powder samples homogenized in 50 mM Tris-HCl buffer (pH Exp. 1
Control CH 75.04 ± 0.39 b 0d
8.5) containing 0.6 mM CaCl2, 5 mM EDTA and 0.05 % (v/v) Triton 160 H CH 75.81 ± 0.19 b 0d
X-100 and centrifuged. The supernatant (0.2 mL) was added to 50 mM 320 H CH 78.93 ± 0.71 a 0d
Tris-HCl (pH 8.5) containing 0.6 mM CaCl2 and 0.24 % (w/v) poly­ Control LH 70.51 ± 1.26 d 0.13 ± 0.02 a
galacturonic acid, then incubated at 37 ◦ C for 30 min and boiled for 160 H LH 70.54 ± 0.76 d 0.04 ± 0.03 b
320 H LH 72.72 ± 0.71 c 0.02 ± 0.01 c
10 min. One unit of PL activity was defined as 1 mM of 4, 5-unsaturated
Exp. 2
product at 232 nm on a fresh weight basis. Control 72.13 ± 0.54 b 0.05 ± 0.02 a
β-galactosidase (β-GAL) and α-arabinofuranosidase (α-ARF) were 160 H 72.50 ± 0.85 b 0.03 ± 0.01 b
measured as described by Brummell et al. (2004) and Sozzi et al. (2002). 320 H 74.74 ± 0.76 a 0.01 ± 0.01 c
Powder samples were homogenized in 0.3 M NaCl, then centrifuged. Values are presented as the means ± standard deviation (SD). Different letters
The supernatant (0.4 mL) was added to 0.04 % (w/v) p-nitro­ indicate significant differences among treatments in Exp.1 or 2 according to
phenyl-β-D-galactoside or p-nitrophenyl-α-D-arabinofuranoside. After Fisher’s protected LSD test at p < 0.05. CH, commercial harvest date; LH, 10
incubation at 37 ◦ C for 30 min, 0.2 M sodium carbonate was added to d after the commercial harvest date.

3
M. Li et al.
3.2. Textural quality
In Exp. 1, the control CH fruit had lower FF compared to the treated
fruit during the 5 months of RA storage (Fig. 1A). After a 5-d ripening at
20 ◦ C, the control CH fruit developed a melting texture at month 3, but
pears failed to ripen (FF on day 5 > 24 N) and a mealy texture was
observed at month 5 (Fig. 1C and E). H and SF, applied separately or in
combination, significantly suppressed the loss in FF of CH fruit during
storage, and the 320 H + SF treatment showed the greatest effect.
Regardless of H application concentration, the H-treated CH fruit soft­
ened < 24 N with a melting texture over the entire storage period. In
contrast, the H + SF resulted in the losing of ripening capacity. For LH
fruit, the control and 160 and 320 H-treated fruit developed a melting
texture after 3 months of storage. No result of FF, sensory scores of
4
Postharvest Biology and Technology 173 (2021) 111429
textural quality, pectic polyuronides, cell wall-modifying enzymes, or
senescence disorders was shown in this study due to 100 % decay
observed in above treatments. However, the SF and H + SF treatments
prolonged the lifespan of LH pears to 5 months. Notably, the develop­
ment of a melting pattern of softening was observed in SF- and 160
H + SF-treated LH fruit, but not in 320 H + SF-treated LH fruit.
Because H and 160 H + SF treatments did not affect LH pears’
ripening, the effects of 1 and 2 % O2 was addressed in Exp. 2. Decreasing
the concentration of O2 from 2 to 1 % maintained higher FF after 7
months of storage (Fig. 1B). The control fruit stored in either O2 con­
centration could soften < 24 N and develop a melting texture after 5 and
7 months of storage plus 5 d at 20 ◦ C (Fig. 1D and F). The 2 and 1 % O2 H
treatments maintained higher FF compared to the control, but no dif­
ference was observed between the two application concentrations.
Fig. 1. Effects of Harvista (H) sprays at 160 and
320 μL L− 1 and 0.15 μL L− 1 SmartFresh (SF),
applied separately or in combination, on flesh
firmness (FF) on day 1 (A and B) and 5 (C and
D) at 20 ◦ C and sensory scores of textural
quality (E and F) of ‘Bartlett’ pears after 3 and
5 months of regular-air (RA) storage and 5 and
7 months of storage in 1 and 2 ± 0.1 % O2 at
− 1.1 ◦ C. Values are presented as the
means ± SD. Different letters indicate signifi­
cant differences among treatments according to
Fisher’s protected LSD test at p < 0.05. CH,
commercial harvest date; LH, 10 d after the
commercial harvest date; 160 H, 160 μL L− 1
Harvista; 160 H + SF, 160 μL L− 1 Harvis­
ta + SF; 320 H, 320 μL L− 1 Harvista; 320
H + SF, 320 μL L− 1 Harvista + SF.
M. Li et al.
Softening of 1 % O2 H-treated fruit was impaired; no fruit attained the
ripening capacity after 7 months of storage. Irrespective of O2 concen­
trations, the 160 H + SF-treated treatments resulted in the firmest fruit
throughout the entire storage period.
3.3. Pectic polyuronides
In Exp. 1, the control and H-treated CH and LH fruit produced high
levels of WSP and CSP with no difference among treatments after 3
months of RA storage plus 5 d at 20 ◦ C (Fig. 2A and C). The levels of WSP
and CSP in the control and H-treated CH fruit decreased at month 5, but
H delayed the losses in WSP and CSP. The SF and H + SF treatments
significantly suppressed the accumulations of WSP and CSP in CH and
LH fruit after 3 months of storage. However, increases of WSP and CSP
5
Postharvest Biology and Technology 173 (2021) 111429
were observed in SF- and H + SF-treated LH treatments after 5 months
of storage. Relatively lower SSP was observed after 5 months, while no
treatment affected SSP (Fig. 2E).
In Exp. 2, the 2 % O2 control and H-treated fruit had higher WSP and
CSP than that in 1 % O2 after 5 months of storage (Fig. 2B and D). After 7
months of storage, decreased WSP and CSP were observed in the 1 and 2
% O2 control fruit and 2 % O2 H treatments. Slightly elevated WSP and
CSP were observed in 1 % O2 H treatments. The 1 and 2 % O2 160 H + SF
treatments completely inhibited the accumulations of WSP and CSP. No
treatment affected SSP (Fig. 2F).
3.4. Cell wall-modifying enzymes
In Exp. 1, compared to CH fruit, LH fruit displayed greater PG
Fig. 2. Effects of H sprays at 160 and 320 μL
L− 1 and SF, applied separately or in combina­
tion, on water-soluble polyuronides (WSP, A
and B), CDTA-soluble polyuronides (CSP, C and
D), and sodium carbonate-soluble polyuronides
(SSP, E and F) of ‘Bartlett’ pears on day 5 at
20 ◦ C after 3 and 5 months of RA storage and 5
and 7 months of CA storage in 2 and 1 ± 0.1 %
O2 at − 1.1 ◦ C. Values are presented as the
means ± SD. Different letters indicate signifi­
cant differences among treatments according to
Fisher’s protected LSD test at p < 0.05.
M. Li et al.
activity. No significant difference was observed among treatments after
3 or 5 months of RA storage (Fig. 3A). Compared to the control CH fruit,
the control LH fruit had higher PME, PL, β-GAL, and α-ARF activity after
3 months of storage (Fig. 3C, E, G, and I); prolonging storage period to
5 months resulted in losses in PME, PL, and β-GAL activity of control CH
fruit and an increase in α-ARF activity. Regardless of H application
concentration, H had no effect on PME, β-GAL, and α-ARF activity in CH
or LH fruit compared to the control fruit. H slightly increased PL activity
in CH fruit, but not in LH fruit. SF and H + SF significantly reduced PME,
PL, β-GAL, and α-ARF activity in CH and LH fruit after 3 months of
storage; elevated PL, β-GAL, and α-ARF activity were observed after
5 months.
In Exp. 2, PG activity was unaffected by O2 concentration, H, or SF
during 7 months of storage (Fig. 3B). Additionally, O2 concentration and
H did not affect PME and α-ARF activity. However, 2 and 1 % O2 160
H + SF treatments significantly reduced PME and α-ARF activity
(Fig. 3D and J). The 1 % O2 H treatments had lower PL and β-GAL ac­
tivity than the 2 % O2 H treatments (Fig. 3F and H). No difference was
observed between the two H application concentrations under 2 or 1 %
O2. However, 2 or 1 % O2 160 H + SF treatment further reduced PL and
β-GAL activity.
3.5. EPR
In Exp. 1, EPR in control CH fruit decreased from 233.70 to
162.50 ng kg− 1 s− 1 after 5 months of RA storage (Fig. 4A). H reduced
EPR in CH and LH fruit with no difference between two application
concentrations. SF significantly inhibited EPR in CH and LH fruit treated
with or without H at month 3. Extending the storage period to 5 months
resulted in an increase of EPR in LH fruit. For example, the SF- and 160
H + SF-treated LH fruit increased in EPR levels to 56.33 and 50.30 ng
kg− 1 s− 1, respectively, at month 5. However, increasing the H applica­
tion concentration from 160 to 320 μL L− 1, combined with SF, reduced
the level of EPR.
In Exp. 2, the 1 % O2 control fruit had lower EPR than 2 % O2 control
fruit (Fig. 4B). Regardless of H application concentration, H significantly
reduced EPR. The 1 % O2 resulted in a further reduction in EPR of 160
and 320 H-treated fruit. EPR of 1 % O2 160 H + SF treated fruit was
inhibited over the entire storage period.
3.6. Decay and senescence disorders
In Exp. 1, the control CH and LH fruit showed 4 and 13 % of fruit with
decay after 3 months of RA storage; this increased to 39 and 100 % of
fruit with decay after 5 months, respectively (Fig. 5A). Regardless of H
application concentration, no decay was observed in H or H + SF CH
treatments at month 3, while elevated rates of decay were observed in H
CH treatments at month 5. For LH fruit, the 160 and 320 H treatment
had 2 and 2 % of fruit with decay, respectively, after 3 months, then
increased to 100 % at month 5. No decay was observed in SF and H + SF
treatments after 3 months of storage, although increased rates of decay
were observed in these treatments after 5 months of storage. The effect
of H and SF on senescence disorders were similar to those on decay in CH
fruit (Fig. 5C). For LH fruit, the control and H-treated fruit developed
senescence disorders at month 3; we showed no results at month 5 due to
the 100 % decay. The SF and H + SF LH treatments did not display
senescence disorders after 3 months of storage, while senescence dis­
orders increased at month 5; the 320 H + SF LH treatment showed a
significantly lower rate of senescence disorders.
In Exp. 2, the 2 % O2 control fruit first expressed decay after
5 months of storage (Fig. 5B). After 7 months of storage, the 2 and 1 %
O2 control fruit developed 6 and 1 % of fruit with decay, respectively. No
decay was observed in 2 and 1 % O2 H and 160 H + SF treatments during
the entire 7-month storage period. No senescence disorders were
observed in any treatment after 5 months of storage plus 5 d at 20 ◦ C
(Fig. 5D). At month 7, the 2 % O2 control and 160 and 320 H-treated
6
Postharvest Biology and Technology 173 (2021) 111429
fruit displayed 23, 19, and 19 % of fruit with senescence disorders,
respectively. Regardless of treatment, fruit stored in 1 % O2 were free of
senescence disorders.
4. Discussion
4.1. Impacts of pre- and post-harvest 1-MCP on LH ‘Bartlett’ pears under
RA storage
LH pears are more susceptible to quality deterioration and post­
harvest disorders during storage (Boonyakiat et al., 1987), in accordance
with our results showing that the control LH ‘Bartlett’ pears developed
100 % decay after 5 months of RA storage. Additionally, a rapid loss in
the capacity of EPR after excess storage contributed to the impaired fruit
softening pattern (Chen and Borgic, 1985; Dong et al., 2018). This ex­
plains why in this study EPR and FF on day 5 and sensory scores in
control LH fruit were lower than the control CH fruit after 3 months.
After 5 months of storage, the decline in EPR paralleled development of
mealy texture in control CH fruit. To extend the life of storage and
melting texture in LH pears, ethylene synthesis should be managed
before or after storage. In this study, H significantly reduced EPR in CH
and LH fruit after 3 months of RA storage. While decreased EPR did not
affect the development of melting texture during ripening, that might be
because H did not bind entirely with ethylene receptors when it was
applied in an open environment, as a result, the residual receptors
induced the softening process (Li et al., 2020). However, H failed to
control decay in LH fruit after 5 months, indicating that pre-harvest
1-MCP sprays were inadequate to prolong the storage and melting life
of LH pears under RA condition. Postharvest application of SF in LH fruit
with or without H treatment undoubtedly extended the storage life to
5 months. Notably, SF- and 160 H + SF-treated LH fruit developed a
melting texture. Wang and Sugar (2015) found that 0.3 μL L− 1
SF-treated pears harvested at 75.6 N took 10–14 d to ripen < 24 N.
Certainly, reducing the SF dose recovered fruit ripening, but physio­
logical disorders could not be controlled (Bai et al., 2006); similar results
were observed in SF-treated LH fruit. Raising H concentration from 160
to 320 μL L− 1 did not suppress EPR or reduce disorders, while the 320
H + SF treatment resulted in firmer CH and LH fruit. Taken together,
results indicated that the 160 H + SF treatment was the optimized
application to retain ripening capacity and reduce the incidences of
disorders in LH ‘Bartlett’ pears during RA storage.
Softening of pears is associated with the solubilization and depoly­
merization of cell wall polyuronides and activity of cell wall-modifying
enzymes during storage and ripening (Brummell and Harpster, 2001).
Previous studies indicated that increases in WSP, CSP, and activities of
cell wall-modifying enzymes were strongly associated with the devel­
opment of melting texture in European pears (Chen and Borgic, 1985;
Murayama et al., 2002, 2006; Dong et al., 2018; Li et al., 2019). Post­
harvest application of 1-MCP slowed WSP accumulation and PME ac­
tivity, as a result, 1-MCP-treated fruit did not achieve a melting texture
(Dong et al., 2018). Similar to this study, SF and H + SF treatments
significantly suppressed the accumulation of WSP and CSP in CH fruit. In
contrast, SF- and 160 H + SF-treated LH fruit developed melting texture
with high levels of WSP and CSP and activities of PME, PL, and α-ARF
after 5 months, suggesting that the delay in harvest or the length of the
storage period impaired the efficacy of SF. In this study, H did not hinder
the degradation of pectin polyuronides chains or modification of cell
wall-modifying enzymes in CH or LH fruit, regardless of application
concentration. However, 320 H + SF-treated LH fruit remained firmer
with lower WSP and CSP over the 5-month storage period, indicating
that increasing H concentration in H + SF application would reduce the
solubilization of pectin polyuronides and the collapse of cell adhesive­
ness associated with softening.
M. Li et al. Postharvest Biology and Technology 173 (2021) 111429

Fig. 3. Effects of H sprays at 160 and 320 μL L− 1 and SF, applied separately or
in combination, on polygalacturonase (PG, A and B), pectin methylesterase
(PME, C and D), pectate lyase (PL, E and F), β-galactosidase (β-GAL, G and H),
and α-arabinofuranosidase (α-ARF, I and J) of ‘Bartlett’ pears on day 5 at 20 ◦ C
after 3 and 5 months of RA storage and 5 and 7 months of storage in 2 and
1 ± 0.1 % O2 at − 1.1 ◦ C. Values are presented as the means ± SD. Different
letters indicate significant differences among treatments according to Fisher’s
protected LSD test at p < 0.05.

7
M. Li et al. Postharvest Biology and Technology 173 (2021) 111429

Fig. 4. Effects of H sprays at 160 and 320 μL L− 1 and SF, applied separately or in combination, on ethylene production rate (EPR, A and B) of ‘Bartlett’ pears on day 1
at 20 ◦ C after 3 and 5 months of RA storage and 5 and 7 months of storage in 2 and 1 ± 0.1 % O2 at − 1.1 ◦ C. Values are presented as the means ± SD. Different letters
indicate significant differences among treatments according to Fisher’s protected LSD test at p < 0.05.

Fig. 5. Effects of H sprays at 160 and 320 μL L− 1 and SF, applied separately or in combination, on decay (A and B) and senescence disorders (C and D) on day 5 at
20 ◦ C of ‘Bartlett’ pears after 3 and 5 months of RA storage and 5 and 7 months of storage in 2 and 1 ± 0.1 % O2 at − 1.1 ◦ C. Values are presented as the means ± SD.
Different letters indicate significant differences among treatments according to Fisher’s protected LSD test at p < 0.05.

4.2. Impact of O2 regimes and 1-MCP on LH ‘Bartlett’ pears combined with 160 μL L− 1 H might produce the minimum ethylene
ripening requirement to recover fruit softening after long-term CA
1
According to the results from Exp. 1, we hypothesized 0.15 μL L− SF storage. Unfortunately, no combination-treated fruit ripened after 7

8
M. Li et al.
months of storage in either O2 condition. One possible explanation for
the disparity in pear softening was attributed to low O2 concentrations;
as it was previously observed that reducing O2 concentration to 0.5 %
showed a greater effect on removing internal ethylene production and,
as a result fruit lost ripening capacity (Li et al., 2019). Similar results
were observed in 1 % O2 H-treated pears; regardless of application
concentration, H-treated pears attained ripening capacity and developed
a melting texture after 5 and 7 months of storage in 2 % O2. Conversely,
1 % O2 control fruit extended melting texture life and reduced post­
harvest disorders for 7 months. In practice, significant amounts of pears
were harvested later than the control fruit. Therefore, the 1 % O2 and 2
% O2 H treatment would appear effective strategies to develop ripening
capacity and to control disorders in LH ‘Bartlett’ pears during CA
storage.
In this study, decreasing O2 concentration did not affect the levels of
WSP and CSP or activity of cell wall-modifying enzymes in LH pears.
This could be because ‘Bartlett’ pears are inherently less sensitive to O2
regime (Zlatić et al., 2016). On the other hand, the relatively short
storage life and susceptibility to fruit senescence of ‘Bartlett’ pears are
associated with high ethylene production during storage
(Villalobos-Acuña et al., 2011; Wang and Sugar, 2015). Although
decreasing O2 concentration resulted in lower internal ethylene pro­
duction in fruit, the level of ethylene receptors continually increased,
and permitted recovery of the softening pattern during shelf life (Tatsuki
et al., 2007). In this study, H significantly decreased the levels of WSP
and CSP and activity of PL and β-GAL. Further, 1 % O2 H-treated fruit
showed significantly lower solubilization of WSP and CSP and activity of
PL and β-GAL, suggesting that ethylene receptors generated at harvest
contributed to cell wall loosening, disaggregation and activation of
pectin-degrading enzymes, such as PL and β-GAL, after removal from
CA. However, lowering O2 concentration or SF treatment would increase
the risk of losing ripening capacity in H-treated pears by reducing EPR,
delaying pectin chain breakdown, and suppressing cell wall-modifying
enzymes. Whether reducing application concentration of SF enabled
recovery of LH pears softening capability requires further exploration
and clarification.
5. Conclusion
In this study, we investigated the changes in melting texture devel­
opment, pectin polyuronides, cell wall-modifying enzymes, ethylene
production, and physiological disorders of LH ‘Bartlett’ pears as influ­
enced by pre- and post-harvest 1-MCP application during ripening after
storage in RA or at 1 and 2 % O2 conditions. Overall, the 160 H + SF
treatment might be the optimal strategy to extend storage and melting
texture life of LH ‘Bartlett’ pears to 5 months at RA condition by accu­
mulating WSP and CSP, maintaining high activities of PME, PL, and
α-ARF, and reducing physiological disorders. However, raising H
application concentration to 320 μL L− 1 in the combination treatment
resulted in blockage of ripening, reduction of solubilization of pectin
polyuronides and activity of cell wall-modifying enzymes. Reducing O2
concentration to 1 % or H treatment plus 2 % O2 storage condition
provided great benefits in maintaining ripening capacity with melting
texture for up to 7 months, retaining high levels of WSP and CSP and
activities of PL and β-GAL, and reducing the rates of decay and senes­
cence disorders. Our findings have led us to conclude that the maximum
melting texture life of LH ‘Bartlett’ pears can be extended by pre- and
post-harvest 1-MCP and O2 regime management during labor shortages
at harvest and warmer growing seasons.
CRediT authorship contribution statement
Meng Li: Methodology, Software, Formal analysis, Writing - original
draft, Visualization. Huanhuan Zhi: Methodology, Software, Formal
analysis, Writing - original draft, Investigation, Resources, Visualization.
Yu Dong: Conceptualization, Validation, Investigation, Resources,
9
Postharvest Biology and Technology 173 (2021) 111429
Writing - review & editing, Supervision, Project administration.
Declaration of Competing Interest
The authors report no declarations of interest.
Acknowledgements
We are grateful to the Columbia Gorge Fruit Growers Association for
financial support and Duckwall Fruit for providing the fruit. The authors
are grateful for the expert assistance provided by Dr. Paul Chen and
technical assistance of Caixia Li. Also, the authors pay tribute to Dr. Yan
Wang, a professor in Oregon State University who died in 2017. This
work would have been impossible without him.
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