Postharvest Biology and Technology 114 (2016) 6975

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Postharvest Biology and Technology

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

1-Methylcyclopropene (1-MCP) inhibits ethylene production of durian

fruit which is correlated with a decrease in ACC oxidase activity in the
peel
Siriporn Amornputtia , Saichol Ketsaa,b,* , Wouter G. van Doornc,1
a
Department of Horticulture, Faculty of Agriculture, Kasetsart University, Bangkok 10900, Thailand
b
Academy of Science, The Royal Society of Thailand, Dusit, Bangkok 10300, Thailand
c
Mann Laboratory, Department of Plant Sciences, University of California, Davis 95616, CA, USA

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

Article history: Fruit of cv. Monthong durian (Durio zibethinus Murr.) were treated for 12 h with 0 (control) and 500 nL L 1
Received 18 August 2015 1-methylcyclopropene (1-MCP) at 25  C and were then stored at a temperature of 15 C. The fruit were
Received in revised form 29 November 2015 transferred from a temperature of 15  C to a temperature of 25  C within a 3-day interval. 1-MCP
Accepted 30 November 2015
treatment did not affect the relatively low rate of ethylene production in fruit stored at a temperature of
Available online 5 January 2016
15  C. In contrast, 1-MCP delayed the dramatic increase in ethylene production in fruit that were
transferred from a temperature of 15  C to a temperature of 25  C. It is well known that the peel (husk)
Keywords:
produces much more ethylene than the pulp, hence, the effects of 1-MCP on the peel will likely
Ethylene production
Durian
overshadow any effect on the pulp. In the peel, 1-MCP had no effect on 1-aminocyclopropene-1-
Pulp carboxylic acid (ACC) synthase activity, but decreased ACO activity. ACC levels in the peel were not
Peel affected by the treatment at 25  C. In the pulp, by contrast, ACO activity was not affected, but ACS activity
ACC was not inhibited at 25  C and ACC levels were found to be lower, after the 1-MCP treatment. It was
ACS concluded that 1-MCP seems to inhibit ethylene production of durian mainly by preventing an increase in
ACO ACC oxidase activity in the peel.
Ripening temperature 2015 Elsevier B.V. All rights reserved.

1. Introduction delayed for a day, while ethylene production was considerably

Durian (Durio zibethinus Murr.) is a climacteric fruit with a short
shelf life (Tongdee et al., 1990; Booncherm and Siripanich, 1991).
The storage temperature cannot be lower than 15  C because lower
temperatures induce chilling injury, whereby, the peel (husk) turns
dark brown, the pulp looses its aroma, and the fruit softening is
delayed (Booncherm and Siripanich, 1991; Ketsa and Paull, 2008).
The peel of durian fruit shows a much higher rate of ethylene
production than the pulp. Ethylene from the peel is required for
normal softening, as the pulp of durian fruit without the peel stays
hard and does not develop its aroma (Booncherm and Siripanich,
1991; Chayprasat, 1993). Ketsa and Pangkool (1995) also showed
that durian ripened above 30  C, climacteric respiration peak was
Abbreviations: ACC, 1-aminocyclopropane-1-carboxylic acid; ACS,
synthase; ACO, ACC oxidase; SAM, S-adenosyl methionine.
* Corresponding author. Fax: +66 2 579 1951x112.
E-mail addresses: agrsck@ku.ac.th, agrsck26@yahoo.com (S. Ketsa).
1
Deceased.
inhibited. Ethylene production of durian appeared to be more
sensitive to high temperature than respiration.
Ethylene derived from S-adenosyl methionine (SAM) is con-
verted to 1-aminocyclopropene-1-carboxylic acid (ACC), the direct
precursor of ethylene. The last two enzymes in the ethylene
synthesis pathway are ACC synthase and ACC oxidase. Each of
these can limit the rate of ethylene production (Yang, 1985). 1-
Methylcyclopropene (1-MCP) is an effective inhibitor of ethylene
action (Blankenship and Dole, 2003). We previously reported that
1-MCP extended the shelf life of durian fruit, if held at room
temperature in the tropics (about 25  C). We also reported that the
storage life of fruit held at a temperature of 15  C is increased from
18 to 30 days, after 1-MCP treatment, that is, treatment with
500 nL L 1 1-MCP for 12 h (at 25  C) prior to storage at 15  C,
increased the storage life of durian cv. Monthong fruit (Amornputti
et al., 2014). Fruit kept at a temperature of 15  C did not show
ACC adequate eating quality until at least, day 18 of storage. Fruit were,
therefore, transferred to 25  C, within a 3 day storage interval to
hasten ripening, in order to obtain an optimum combination of
storage and shelf life duration. To induce ripening even further,

http://dx.doi.org/10.1016/j.postharvbio.2015.11.020
0925-5214/ 2015 Elsevier B.V. All rights reserved.
70 S. Amornputti et al. / Postharvest Biology and Technology 114 (2016) 6975
ethephon at 480 mL L 1 was applied with brush at the cut surface
of the cut stalk. Adequate storage life was dened as storage
leading to a time of ripeness (adequate eating quality), determined
at 25  C, within 3 days or more. The storage life of control fruit at
15  C was 18 days, whilst after 1-MCP treatment, the storage life
was 30 days. Additionally, treatment with 1-MCP extended shelf
life. The time interval to ripeness (adequate eating quality) at 25  C
in controls was 5 days. After 500 nL L 1 1-MCP treatment for 12 h at
25  C, the shelf life at 25  C was 11 days (Amornputti et al., 2014).
The objective of the present work was to determine if 1-MCP
inhibits ethylene production through ACC synthase or ACC oxidase.
We tested the hypothesis that the activity of one of these two
enzymes (or both enzymes) is reduced by 1-MCP. We also used
500 nL L 1 1-MCP prior to storage at 15  C, and the fruit was again
transferred from 15  C to 25  C within a 3-day interval, but an
ethephon treatment of the cut fruit stalk was not included, after
storage at 15  C.
2. Materials and methods
2.1. Plant material
Durian (Durio zibethinus Murr.) fruit cv. Monthong were
harvested 123  2 days after full bloom, from a commercial
orchard in Chanthaburi province of Eastern Thailand. The maturity
of the durian fruits was determined by combined techniques: day
count, character of fruit spines, tapping the fruit, colour and shape
Fig. 1. Ethylene production of durian cv. Monthong treated at 25  C with 0 (
) or of ACC g 1.
500 nL L 1 1-MCP (*) for 12 h. Fruit was stored at 15  C for 18 days (A). Other fruit
was taken out of 15  C storage at 3-day intervals and was transferred to 25  C (B) 1-
MCP treatment started on day 0. Data are means  SD of 10 fruit each (whole fruit).
Statistical differences between treatments (P < 0.05) for each day are indicated by a
different letter.
of the fruit (Yaacob and Subhadrabandhu, 1995). Fruit were
selected based on uniformity in size about 2.5 kg each. They were
dipped, in the packing house, in imazalil at 0.5 mL L 1 for 20 s to
control fruit rot, caused by Phytophthora palmivora. They were then
transported in a temperature-controlled truck (25  C) to the
laboratory, where they arrived within a day of harvest. Transpor-
tation took about 6 h.
2.2. Treatment with 1-MCP; storage at 15  C and transfer to 25  C
Fruit were placed in a 71 L sealed chamber and subjected to
1-MCP at 500 nL L 1, for 12 h at 25  C, as described by Palapol et al.
(2015). 1-MCP was generated by adding water to the 1-MCP
powder (EthylBloc1, Floralife Inc., Walterboro, SC, USA), placed in a
vial. Introducing water into the vial generated 1-MCP gas, resulting
in a nal concentration of 500 nL L 1 of 1-MCP. Fans were used to
maintain air circulation. The control batch of fruit was placed in an
identical plastic chamber without 1-MCP treatment.
After the 1-MCP treatment, the fruit (four per carton box) were
stored at 15  C and 8590% RH. For every 3 days of storage at 15  C,
18 fruit for each treatment (18 fruit for control and 18 fruit for
15  C), each half was used immediately to analyse ACC, ACS and
ACO. The second half of 18 fruit were left at 25  C for 3 days and
ACC, ACS and ACO were analysed), were transferred to 25  C and
8090% RH and was left at that temperature.
2.2. Ethylene production
Ethylene production was measured using the method of Palapol
et al. (2015). At intervals, fruit were taken from each treatment and
assessed for ethylene production. Five fruit of each treatment were
placed individually into 18.5 L jars (that were closed airtight) for
30 min, A 5 mL gas sample was taken from the head space and
injected into a gas chromatograph, equipped with a Shimadzu GC-
14 ame ionization detector (Shimadzu, Tokyo, Japan).
2.3. ACC content; activities of ACS and ACO
The peel and pulp were separated and cut into 0.5  0.5  0.5
cm3 segments within a 3-day interval period. The material was
immediately frozen in liquid nitrogen and stored at a temperature
of 70  C until the time of use. The ACC content, ACS and ACO
activities were determined in frozen samples.
The ACC content and ACS activity were measured using the
method of Hoffman and Yang (1982), though with slight
modication. For ACC assessments, samples of 4 g pulp or peel
were homogenized with a 10 mL of 9% trichloroacetic acid (TCA),
and kept at a temperature of 4  C for 24 h, and then, the
homogenate was centrifuged at 18,190  g for 25 min at 4  C. The
supernatant solution was adjusted to pH 78 range with 1 N NaOH
and the supernatant volume was recorded. The sample was placed
in capped 10 mL vials, which contained, apart from the sample
solution (500 mL), 100 mL of 10 mM HgCl2 and 300 mL of distilled
water respectively. The internal standard solution contained
500 mL of ACC solution, 100 mL, 10 mM of HgCl2, 50 mL of the
0.04 mM ACC standard and 250 mL of distilled water. One hundred
microliters of a cold mixture of 5.25% NaOCl and saturated NaOH
(2:1 v/v) was injected into the vial through the serum cap. The
mixture was vortexed and incubated on ice. After incubation, a
1 mL gas sample was taken, using a syringe. The gas was injected
into a gas chromatograph and the results were expressed as nmol
For ACS activity, frozen samples (containing 4 g of pulp or peel)
were homogenized with 0.2 g of polyethyleneglycol (PEG) and
10 mL extraction buffer (100 mM phosphate, 10% glycerol, 5 mM
DTT and 5 mM pyridoxal-5-phosphate) with an addition of 10 mL
 S. Amornputti et al. / Postharvest Biology and Technology 114 (2016) 6975 71

Fig. 2. ACC content in the pulp of durian cv. Monthong treated at 25  C with 0 ( ) or 500 nL L 1 1-MCP (*) for 12 h and stored at 15  C (A) and then transferred to 25  C at
3-day interval (B). 1-MCP treatment started on day 0. Data are means  SD of three replications (912 fruit). Statistical differences between treatments (P < 0.05) for each day
are indicated by a different letter.

cooled to 70  C. The mixture was vortexed. The homogenate was
centrifuged at 18,190  g for 20 min at 4  C and the pellet was lled
with 5 mL extraction buffer and incubated for 20 min at 4  C. The
homogenate was centrifuged at 26,640  g for 20 min at 4  C. The
solution was placed in a dialysis tube, immersed in a dialysis buffer
(10 mM phosphate, 10% glycerol, 5 mM DTT and 5 mM pyridoxal-5-
phosphate) and incubated for 2 h at 4  C. The supernatant volume
was recorded. ACC synthase activity was determined in a reaction
mixture containing 400 mL enzyme solution, 600 mM EPPS (50 mL)
(pH 8.5), and distilled water (90 mL). After incubation for 3 h at
30  C, 0.5 mM SAM (60 mL) was added to the solution. The samples
were held in capped 12 mL vials containing 100 mL sample, 10 mM
HgCl2 (100 mL) and distilled water (200 mL). The internal standard
sample contained 10 mM HgCl2 (100 mL), 0.04 mM ACC (50 mL) and
distilled water (150 mL) respectively. The ACS synthase activity was
determined in a reaction mixture containing 400 mL of the sample,
50 mL assay buffer, 60 mL 0.2 mM SAM and 90 mL distilled water.
After incubation for 30 min at 30  C, the samples were kept in
capped 12 mL vials and 100 mL of 10 mM HgCl2 was added to stop
the reaction. The internal standard containing 150 mL distilled and
50 mL 0.04 mM ACC and closed tubes with a serum cap made on ice,
afterwards, 100 mL of a cold mixture of 5.25% NaOCl was added and
72 S. Amornputti et al. / Postharvest Biology and Technology 114 (2016) 6975
Fig. 3. ACC content in the peel of durian cv. Monthong treated at 25  C with 0 (
) or
500 nL L 1 1-MCP (*) for 12 h and stored at 15  C (A) and then transferred to 25  C at
3-day interval (B). 1-MCP treatment started on day 0. Data are means  SD of three
replications (912 fruit). Statistical differences between treatments (P < 0.05) for
each day are indicated by a different letter.
saturated NaOH (2:1 v/v) was injected into the vial through the
serum cap. The mixture was vortexed and incubated on ice for
5 min. A 1 mL gas sample was taken, using a syringe and the gas
was injected into a gas chromatograph. A conversion factor was
determined by comparison with ethylene production from an
internal ACC standard. The results were expressed as mg 1 protein
of ACC produced. The protein level was determined using the
Bradford method, with serum albumin as a standard.
ACO activity was measured using the method of Hoffman
and Yang (1982), though, with slight modication. Samples
were homogenized in a mortar with 1 mL of extraction buffer
(2 mL g 1 for pulp and 5 mL g 1 for peel) containing 0.1 M Tris-
(hydroxymethyl)-aminomethane, 5 mM DTT, 30 mM L(+)-ascor-
bic acid, sodium salt and 30% glycerol; pH 7.2 with HCl. The
homogenate was centrifuged for 40 min at 18,190  g and the
supernatant was used for the ACO assay. The ACO activity was
assayed by a gas chromatograph, equipped with a Shimadzu GC-
14 ame ionization detector (Shimadzu, Tokyo, Japan) for
determination of the ethylene produced for 30 min after
incubation at a temperature of 30  C in capped 12 mL vials,
containing 400 mL of reaction buffer (pH 7.2; 0.1 M Tris-
(hydroxymethyl)-aminomethane, 30 mM Na-ascorbate, 30 mM
NaHCO3, 100 mM FeSO4 and 30% glycerol) and 10 mM ACC
(100 mL). The reaction was initiated by the addition of 500 mL of
the supernatant. In controls, the reactions used no enzymes or
boiled ones. In neither case was an ethylene production
detected. The protein concentration was determined using the
Bradford method, with serum albumin as a standard.
2.4. Statistical analysis
Ethylene was determined using 10 replications (single fruit),
while ACC, ACS and ACO were determined using three biological
replications per treatment, each containing 34 fruit. Data (1-MCP
effect) were analyzed by ANOVA (analysis of variance) and means
were compared using least signicance difference (LSD) at
P  0.05 and t test.
3. Results
3.1. Ethylene production
The ethylene production of fruit stored at 15  C with 1-MCP
treatment increased gradually from day 0 to day 18 (Fig. 1A), while
that of fruit with 1-MCP treatment slightly increased particularly
from day 0 to day 3 and day 12 to day 18. Ethylene production of
fruit without and with 1-MCP treatment after transfer to a
temperature of 25  C, rapidly increased to a maximum peak on day
12 and day 15, respectively and thereafter, it declined sharply. The
maximum peak of ethylene in the 1-MCP-treated fruit was
signicantly higher and later than that of the control fruit (Fig. 1B).
3.2. ACC content
For fruit stored at 15  C, 1-MCP delayed the peak production of
ACC in the pulp at 15  C (Fig. 2A) and almost suppressed it at 25  C
(Fig. 2B). 1-MCP signicantly reduced ACC content in the peel on
day 3, 9, and 12 at 15  C (Fig. 3A) but did not reduce ACC content at
25  C.
3.3. ACS activity
In pulp at 15  C, 1-MCP delayed the onset of ACS activity peak
and signicantly reduced the maximum peak compared with the
untreated fruit (Fig. 4A). 1-MCP completely suppressed the peaks
of ACS activity at 25  C Fig. 4B). In the peel at 15  C, non-signicant
differences in ACS activity were found (Fig. 5A), while ACS activity
peaked after 12 days at 25  C irrespective of the treatments
(Fig. 5B).
3.4. ACO activity
In pulp, 1-MCP did not delay the onset of ACO activity peak and
signicantly reduced the maximum peak compared with the
untreated fruit at both 15  C (Fig. 6A) and 25  C (Fig. 6B). In peel, 1-
MCP reduced signicantly the ACO activity peak on day 9 at 15  C
(Fig. 7A), while 1-MCP reduced signicantly the ACO activity peak
on day 9 and 12 at 25  C (Fig. 7B).
4. Discussion
Little effect of the 1-MCP treatment was found during fruit
storage at 15  C. This might be explained by the relatively low
ethylene production at 15  C. As soon as the fruit was transferred to
25  C, the ethylene production showed a large increase. This
indicates that ethylene production was inhibited at 15  C.
According to the law of Arrhenius, the Q10 of many reactions is
23. The production of ethylene by durian fruit, when compared at
15  C and 25  C, showed a much higher difference. The inhibition,
therefore, is larger than is expected as shown from the effect of
temperature on enzyme activity. This inhibition might be due to
slight chilling injury, already at 15  C. Durian is a tropical fruit and
considered to be sensitive to chilling injury even though stored at
an optimum temperature of 15  C longer than 2 weeks (Siripha-
nich, 1993). Similarly other tropical fruits such as custard apple
 S. Amornputti et al. / Postharvest Biology and Technology 114 (2016) 6975 73

Fig. 4. ACS activity in the pulp of durian cv. Monthong treated with 0 ( ) or 500 nL L 1 1-MCP (*) for 12 h and stored at 15  C (A) then after transfer to 25  C at 3-day interval
(B). 1-MCP treatment started on day 0. Data are means  SD of three replications (912 fruit). Statistical differences between treatments (P < 0.05) for each day are indicated
by a different letter.

(Brown and Scott, 1985), pineapple (Paull and Rohrbach, 1985), and
rambutan (Ketsa and Klaewkasetkorn, 1995) have been reported to
be very sensitive to chilling injury even at 15  C.
It was previously reported that ethylene production from
durian peel is up to a factor 100 higher than that from the pulp. This
means that the ethylene production in the whole fruit is mainly
determined by the production in the peel (Booncherm and
Siripanich, 1991; Chayprasat, 1993). In our present experiments,
ACO activity in peel was about 5 fold higher than that in pulp, while
their ACC content and ACS activity were comparable. ACO involves
in the last step of ethylene biosynthesis catalyzing the conversion
of ACC to ethylene (Yang, 1985). This may explain why peel
produced ethylene much higher than pulp in durian.
Many fruits show an autocatalytic increase in ethylene
production, which involves the perception of an initial increase
in ethylene production (and/or increased sensitivity to existing
ethylene levels), which leads to a further increase in ethylene
production. Increased expression of genes encoding ACS and/or
ACO, and increased activity of these two enzymes is responsible, at
least in part, for the increase in ethylene production rates (Watkins,
2006). The autocatalytic rise in the ethylene production rate is
inhibited by 1-MCP, which blocks the ethylene receptor.
Although in the present tests, there was no noticeable effect of
1-MCP on fruit ethylene production at 15  C, but it affected peel
ACO activity involved in the last step of ethylene synthesis. 1-MCP
did not affect ACS activity but affected ACC content in peel. ACS
74 S. Amornputti et al. / Postharvest Biology and Technology 114 (2016) 6975
) or
Fig. 5. ACS activity in the peel of durian cv. Monthong treated at 25  C with 0 (
) or (*) 1500 nL L 1 MCP for 12 h and then stored at 15  C (A) and then after transfer
500 nL L 1 1-MCP (*) for 12 h and stored at 15  C (A) and then transferred to 25  C at
3-day interval (B). 1-MCP treatment started on day 0. Data are means  SD of three
replications (912 fruit). Statistical differences between treatments P < 0.05) for
each day are indicated by a different letter.
catalyses the conversion of SAM to ACC, and ACC is the direct
precursor of ethylene (Yang, 1985). During storage at 15  C,
treatment with 1-MCP resulted in lower pulp ACS activity and
the peel ACC concentrations and peel ACO activity, while did not
affect pulp ACO activity. One possible explanation for the inhibition
of peel ACO in the absence of a noticeable decrease in ethylene
production of durian fruit at 15  C, suggesting that 1-MCP efcacy
is less at lower temperatures (Acuna et al., 2011).
For fruit transferred from 15  C to 25  C, the ethylene production
drastically increased, by a factor of about 10 at its maximum. This
increase in production was delayed, not attenuated, by 1-MCP.
Lower ethylene production was observed after treatment with 1-
MCP on day 6-day12. The data suggested that the concentration of
applied 1-MCP (500 nL L 1), effectively inhibited the mechanism of
autocatalytic ethylene production for about 3 days (from day 3 to
day 6), after which the same kinetics were found as observed in the
controls. In other experiments, we found a 12 h treatment of durian
fruit with 1000 nL L 1 at 25  C, after which the fruit was kept at
25  C, similarly, it delayed the increase in ethylene production,
while it did not attenuate the production. The delay lasted 56 days
(from day 0 to day 56), after this treatment (Palapol et al., 2015).
In explaining the inhibition of ethylene production of the whole
fruit by 1-MCP, after its transfer from 15  C to 25  C, day 612 of the
data should be analyzed. The ACC concentrations in the peel were
not affected by the treatment, while in the pulp the ACC levels were
observed to be lower on day 6 and 12, after the treatment. In both
Fig. 6. ACO activity in the pulp of durian cv. Monthong treated at 25  C with 0 (
stored at 25  C at 3-day interval (B). 1-MCP treatment started on day 0. Data are
means  SD of three replications (912 fruit). Statistical differences between
treatments (P < 0.05) for each day are indicated by a different letter.
the peel and pulp, the ACS activities seemed unaffected by the
transference, if judged by t test on each sampling day and the
overall LSD values. Transfer from 15  C to 25  C had no signicant
effects on ACS in both pulp and peel. It means that the transfer does
not inuence the effect of 1-MCP acting at both temperatures. The
reason for this lack of response after transference is not known.
One possibility is that low temperatures may affect a variety of
factors inuencing the interaction of 1-MCP with the receptors.
Enzymatic 1-MCP metabolism has been shown to be important in
plant tissues and would be slowed at lower temperatures (Huber
et al., 2010). Lower temperatures might decrease 1-MCP afnity or
interaction with the ethylene receptors as suggested previously
(Mir et al., 2001; Jiang et al., 2004).
The ACO activity in the pulp was not affected by the transfer to
warmer temperature. In contrast, in the peel, the ACO activities had
decreased on day 6 and 9. This seems to be the main explanation
for the low ethylene production of the fruit. In fruit other than
durian, 1-MCP also inhibits the activities of ACS and/or ACO. In
peach fruit, for example, 1-MCP also inhibits ethylene production
through a reduced ACO activity, while ACS activity was not affected
(Mathooko et al., 2001). In whole banana fruit, Pathak et al. (2003)
reported the inhibition of the activities of both ACS and ACO after
1-MCP treatment. When studying only the banana peel, it was
found that 1-MCP inhibited ethylene mainly through the inhibition
of ACO activity (Ketsa et al., 2013). This same difference between
peel and pulp is reported here for durian.
It can be concluded that a 12-h treatment of durian with
500 nL L 1 1-MCP inhibited ethylene production after transfer from
 S. Amornputti et al. / Postharvest Biology and Technology 114 (2016) 6975
Fig. 7. ACO activity in the peel of durian cv. Monthong treated at 25  C with 0 (o) or
500 nL L 1 (*) 1-MCP for 12 h and stored at 15  C and then transferred to 25  C at 3-
day interval (B). 1-MCP treatment started on day 0. Data are means  SD of three
replications (912 fruit). Statistical differences between treatments (P < 0.05) for
each day are indicated by a different letter.
a temperature of 1525  C, due to a reduction of ACO activity in the
peel. For fruit that remained at 15  C throughout the experiment,
any inhibition of ethylene production due to the 1-MCP treatment
was not detectable, but fruit peel exhibited a lower ACO activity.
Acknowledgements
The research was nancially supported by the Thailand
Research Fund (TRF-DPG528002) and the Kasetsart University
Research and Development Institute (KURDI-ORP-1.52).
75
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