Journal of the Science of Food and Agriculture
J Sci Food Agric 81:1244±1249 (online: 2001)
DOI: 10.1002/jsfa.933
Methyl jasmonate reduces chilling injury
symptoms and enhances colour development
of ‘Kent’ mangoes
GA González-Aguilar,1,2 JG Buta1 and CY Wang1*
1
Horticultural Crops Quality Laboratory, Agricultural Research Service, US Department of Agriculture, 10300 Baltimore Avenue, Beltsville,
MD 20705-2350, USA
2
Centro de Investigación en Alimentación y Desarrollo, AC, Dirección de Tecnologı́a de Alimentos de Origen Vegetal, Apartado Postal
1735, Hermosillo, Sonora 83000, Mexico
Abstract: Treatment of mango (Mangifera indica cv `Kent') fruits with methyl jasmonate (MJ) vapour
for 20 h at 20 °C reduced chilling injury (CI) symptoms and enhanced skin colour development. MJ at
10 4 M was the most effective concentration for reducing CI and decay in fruits stored at 5 °C followed by
7 days at 20 °C (shelf-life period). The use of 10 5 M MJ enhanced yellow and red colour development of
mangoes stored at 20 °C. These fruits possessed higher L*, a* and b* values than untreated fruits and
those treated with 10 4 M MJ. Ripening processes were inhibited by cold storage (5 °C) in control fruits.
After cold storage and shelf-life period, fruits treated with 10 5 M MJ ripened normally and contained
the highest total soluble solids (TSS). These fruits also maintained higher sugar and organic acid levels
than fruits subjected to other treatments. We concluded that MJ treatment could be used to reduce
decay and CI symptoms and also to improve colour development of mango fruits without adversely
affecting quality.
# 2001 Society of Chemical Industry
Keywords: methyl jasmonate; Mangifera indica; sugars; organic acids; chilling injury; decay
INTRODUCTION
Mango fruits are susceptible to a number of physiological disorders during cold storage. Like many other
tropical and subtropical fruits, mangoes develop
chilling injury (CI) when held below the critical
chilling temperature.1,2 The most common visual
symptoms of CI in mango fruits are dark, scald-like
discolourations and pitting or sunken lesions in the
peel.1 In addition to the surface symptoms, the chilled
tissues are weakened and, as a result, susceptible to
decay and pathogen invasion. Therefore postharvest
life of the fruits is limited. Various methods such as
heat treatment, storage in modi®ed and controlled
atmospheres, intermittent warming, and application of
plant growth regulators have been developed to
alleviate CI symptoms of mangoes.3±5
It has been reported that treatments with methyl
jasmonate (MJ) effectively reduce CI in grapefruit6 as
well as in other chilling-susceptible commodities.6,7
Recently, it was reported that MJ can also be applied,
as a postharvest treatment, to effectively suppress the
grey mould rot caused by Botrytis cinerea in strawberries8 and cut roses9 and to reduce decay caused by the
green mould Penicillium digitatum in grapefruit.10 Also,
MJ extends the shelf-life and reduces microbial
contamination of fresh-cut celery and peppers.11
Recently, we observed that vapour treatment with
10 4 M MJ reduces CI during storage at 7 °C without
altering the ripening process of `Tommy Atkins'
mangoes.12 However, the biochemical and colour
changes induced by MJ during cold storage and
shelf-life period were not studied. Also, bene®cial
effects of MJ on other mango varieties and other
tropical fruits have not been studied. A major problem
associated with CI occurrence is the lack of colour
development after transfer of the fruits to higher
temperatures. Skin colour is a major determinant of
consumer appeal in most mango cultivars. Fruits with
no green colour and free of defects, such as CI
symptoms, often command higher prices. It has been
observed that postharvest illumination and treatment
with ethylene can increase both the yellow and red
colours of mango skin.13 It has been found that
application of MJ vapour improved skin colour by
promoting b-carotene synthesis and chlorophyll degradation in Golden Delicious apples.14
Since there was a lack of information on the
in¯uence of MJ on mango fruit quality during low-
* Correspondence to: CY Wang, Horticultural Crops Quality Laboratory, Agricultural Research Service, US Department of Agriculture, 10300
Baltimore Avenue, Beltsville, MD 20705-2350, USA
E-mail: [email protected]
Contract/grant sponsor: Consejo Nacional de Ciencia y Technologı́a (CONACYT), Mexico
(Received 3 January 2001; revised version received 23 April 2001; accepted 22 June 2001)
# 2001 Society of Chemical Industry. J Sci Food Agric 0022±5142/2001/$30.00
1244
Effect of methyl jasmonate on chilling injury and colour of mangoes
temperature storage and its effect on skin colour, the
present study was initiated to determine the ef®cacy of
MJ vapour application in (1) reducing deterioration
and CI symptoms, (2) improving the skin colour of
mango fruits and (3) the biochemical changes associated with quality loss.
MATERIALS AND METHODS
Materials and experiments
Mangoes (Mangifera indica cv `Kent') were obtained in
April and May 1999 from a wholesale market near
Jessup, MD, USA. Fruits used for this experiment
initially had a ®rmness of 120±125 N and a soluble
solids content of 8.2 °Brix. Fruits were sorted, cleaned
and randomised. Fruits of uniform size, shape and
maturity and free from defects were used. Mangoes
were then washed with a 200 ppm solution prepared
from 5% sodium hypochlorite (Chlorox), air-dried
and randomly divided into three lots of 135 fruits each.
Lot 1 was used as control; lots 2 and 3 were treated
with MJ vapour.
Fruits from lots 2 and 3 were placed in 120 l
containers together with 10 5 or 10 4 M MJ applied on
®lter paper, then incubated for 20 h at 20 °C. Afterwards, jars were opened and ventilated for 6 h, then the
fruits were stored at 5, 10 or 20 °C and 85±88, 78±80
or 70±75% relative humidity. Fruits from lot 1 were
handled in the same way but without MJ treatment
(control). The MJ concentrations which achieved
higher reductions in physiological disorders were
selected based on preliminary studies carried out in
the laboratory. In addition, we observed that concentrations of MJ higher than 10 4 M signi®cantly
increased changes in colour and abnormal ripening.
After 14 days at 5 or 10 °C, fruits were transferred for
an additional 7 days at 20 °C (shelf-life period). Fifteen
fruits from each treatment and storage temperature
were sampled after 7 and 14 days at 5, 10 or 20 °C and
shelf-life period. Fruits were evaluated for changes in
weight loss, ¯esh ®rmness, colour (L*, a*, b* and
°Hue), pH, total soluble solids (TSS) and titratable
acidity (TA). Decay, ripening, overall quality and CI
injury symptoms were evaluated after 14 days at 5 or
10 °C and shelf-life period. Four samples (2 g) of ¯esh
tissue were randomly taken from each treatment
initially, after 14 days at 20 °C and after 14 days at
5 °C ‡ 7 days at 20 °C for analysis of organic acids and
sugars.
Quality attributes
Mangoes were weighed before and after the storage
period to calculate the percentage of fresh weight loss.
Flesh ®rmness was determined at the same four
positions on each fruit (skin removed) using a ®rmness
tester (Pressure Tester Model EPT-1R, Lake City
Technical Products, Inc, Kelowa, BC, Canada) with
an 8 mm plunger. Skin colour was assessed with a
tristimulus colour difference meter (Minolta CR 300)
and expressed as L*, a* and b* values. These values
were used to calculate °Hue values. Negative a* values
indicate green and higher positive a* values red colour.
Higher positive b* values indicate a more yellow skin
colour. Six locations around the circumference of each
fruit were evaluated. Extent of decay was assessed
based on the area of decay. Degree of ripeness was
based on red and yellow colour development. Overall
quality was rated on a scale of 1±5, where 1 = very
Figure 1. Percentage of weight loss of ‘Kent’ mango fruits treated with methyl jasmonate (MJ) vapour for 20 h at 20 °C and then stored for 14 days at 5, 10 and
20°C ‡ shelf-life period. Vertical bars indicate standard error of the mean (n = 15).
J Sci Food Agric 81:1244±1249 (online: 2001)
1245
GA GonzaÂlez-Aguilar, JG Buta, CY Wang
Figure 2. Firmness of ‘Kent’ mango fruits treated with methyl jasmonate (MJ) vapour for 20h at 20°C and then stored for 14 days at 5, 10 and 20 °C ‡ shelf-life
period. Vertical bars indicate standard error of the mean (n = 15).
poor, 2 = poor, 3 = fair, 4 = good and 5 = excellent. The
CI score was based on the percentage of total surface
area affected by sheet pitting, where 0 = no injury,
1 = slight, 2 = moderate and 3 = severe, according to
GonzaÂlez-Aguilar et al. 15 TSS was determined using a
temperature-adjusted refractometer, TA (as % citric
acid) using 0.1 M NaOH, and pH using a pH meter,
according to AOAC16 procedures. Organic acids and
sugars were determined according to Wang.17
Analysis of variance (ANOVA) and Tukey's multiple-range test for comparison of means and least
signi®cant differences (p < 0.05) were performed on
the data using the SAS 6.03 system.18
MJ treatment had little effect on the ®rmness of
mangoes during storage at 5 or 20 °C (Fig 2). After the
shelf-life period, softening occurred at higher storage
temperatures and was not affected by MJ treatment.
However, 10 5 M MJ treatment reduced the loss of
®rmness of fruits stored for 14 days at 10 °C ‡ shelf-life
period. MJ treatment slightly decreased the loss of
®rmness of strawberries20 but enhanced softening of
apples.21
Fig 3 shows the effect of MJ treatment on mango
quality after storage for 14 days at 5 °C ‡ 7 days at
20 °C. In general, decay of mangoes was signi®cantly
reduced by MJ treatment, and ef®cacy was increased
RESULTS AND DISCUSSION
While weight loss was generally greater at higher
storage temperatures (Fig 1), MJ-treated fruits showed
less weight loss than control fruits when stored for 14
days at 10 °C ‡ 7 days at 20 °C (shelf-life). The reduction in weight loss observed in fruits treated with
10 5 M MJ and stored at 20 °C could be associated with
stomatal closure induced by MJ, which had reduced
transpiration of fruits as observed in other tissues.19
Weight loss was also signi®cantly reduced by MJ
treatment in radishes, and this reduction was associated with MJ-induced inhibition of sprouting and
decreased transpiration.17 However, 10 4 M MJ enhanced ripening and decay in mangoes at 20 °C,
resulting in increased transpiration and moisture loss
and the appearance of shrivelling symptoms. Similar
concentrations did not signi®cantly affect water loss
and ®rmness of `Keitt' mangoes during storage at
7 °C.12
1246
Figure 3. Decay, ripeness and overall quality of ‘Kent’ mango fruits treated
with methyl jasmonate (MJ) vapour for 20 h at 20 °C and then stored for 14
days at 5 °C ‡ shelf-life period. Vertical bars indicate standard error of the
mean (n = 15).
J Sci Food Agric 81:1244±1249 (online: 2001)
Effect of methyl jasmonate on chilling injury and colour of mangoes
with the lower concentration. MJ treatment at 10 5 M
considerably improved ripening and overall quality of
mangoes after storage at 5 °C ‡ shelf-life period. A
delay in colour development was observed in untreated fruits (controls), possibly as a consequence of
the chilling temperature (5 °C) used. It has been
observed that abnormal or failure in ripening can
occur in mango fruits after exposure to chilling
temperatures.3 Better quality was usually observed in
MJ-treated mangoes. Fruits treated with 10 5 M MJ
developed more intense red and yellow colours and
retained overall quality compared with controls.
MJ treatment protected against CI during both 5 or
10 °C storage and subsequent shelf-life period (Fig 4).
Chilling injury, manifested as surface pitting, was
evident on control fruits kept in cold storage and
progressed rapidly during the shelf-life period. The
conditions of time and temperature that cause CI
symptoms in various mango cultivars are highly
variable, but CI symptoms were greater on fruits
stored at 5 °C than on those stored at 10 °C. It has been
reported that temperatures below 13 °C for a period of
10 days produced CI symptoms in fully ripened `Kent'
mangoes.22 It is well known that low-temperature
stresses render fruits more susceptible to attack by
pathogens.23 In the present study we observed that
chilled control fruits were more susceptible to decay
and pathogen invasion, whereas MJ treatment signi®cantly increased tolerance to CI and fungal development (Figs 3 and 4). MJ at 10 4 M reduced CI
symptoms of mango fruits more than 10 5 M MJ.
Similar concentrations were found to reduce CI
symptoms of `Hass' avocado, grapefruit, peppers,
zucchini squash and `Tommy Atkins' mangoes.6,7,12
The mechanism by which MJ induced tolerance to
CI and resistance against different pathogens is
unclear. It has been suggested that jasmonates play
an important role as signal molecules in mobilisation
of plant defence responses to pathogen attack. Meir et
al 6 suggested that MJ might mediate the natural
response of fruits to chilling stress. Droby et al 10
concluded that the mode of action of MJ in reducing
green mould decay in citrus fruits could be associated
with the induction of natural resistance against
pathogens. Wang and Buta7 suggested that MJ delays
the onset of chilling symptoms by mechanisms that
involve an increase in abscisic acid and polyamine
levels. MJ activates genes encoding antifungal proteins24,25 and other genes involved in phytoalexin
biosynthesis.26 Further studies are necessary, however, to elucidate the mechanisms by which MJ
reduces CI and pathogen invasion in mango fruits.
No signi®cant differences in pH and % TA were
found among the treatments evaluated (data not
shown). Treatment with 10 5 M MJ signi®cantly increased L*, a* and b* values during storage at 20 °C
(Table 1). This treatment improved red and yellow
colour development of mangoes compared with
controls. No signi®cant differences were observed in
°Hue values among treatments evaluated at 20 °C.
Only higher b* values were observed on fruits treated
with 10 5 M MJ after 14 days at 5 °C ‡ shelf-life period.
Fruits treated with 10 5 M MJ contained lower TSS
than controls. It appears that this treatment enhanced
skin colour changes without affecting the rate of
ripening and maintained the quality of mangoes. Cold
storage ‡ shelf-life period suppressed colour (a* and
b*) development of mango fruits compared with those
stored at 20 °C. No signi®cant differences were found
in skin colour (L*, a* and °Hue) among the treatments. However, the highest b* values were observed
on fruits treated with 10 5 M MJ after 14 days at
5 °C ‡ shelf-life period.
Control fruits had the lowest TSS after cold storage
and shelf-life period, indicating that fruit ripening was
partially retarded or inhibited (Table 1). Lederman et
al 27 found that `Keitt' mangoes with severe chilling did
not reach normal TSS and citric acid levels compared
with non-chilled fruits stored at higher temperatures.
Table 1. Changes in colour (L*, a*, b* and °Hue) and total soluble solids
(TSS) of mango fruits treated with methyl jasmonate (MJ) vapour for 20 h
prior to storage at 5 and 20°Ca
Colour value
Treatment
Figure 4. Chilling injury score of ‘Kent’ mango fruits treated with methyl
jasmonate (MJ) vapour for 20 h at 20 °C and then stored for 14 days at 5 and
10°C ‡ shelf-life period. Vertical bars indicate standard error of the mean
(n = 15).
J Sci Food Agric 81:1244±1249 (online: 2001)
L*
a*
b*
°Hue
TSS
14 days at 20 °C
Control
53.9b
59.6a
MJ 10 5 M
55.8ab
MJ 10 4 M
16.9b
19.1a
16.1b
43.1b
47.3a
44.8ab
68.6a
71.2a
66.9a
14.8a
13.8b
15.5a
14 days at 5 °C ‡ 7 days
Control
53.7a
MJ 10 5 M
56.9a
54.1a
MJ 10 4 M
at 20 °C
6.2a
8.1a
7.0a
39.2b
43.2a
40.1b
78.3a
82.1a
80.2a
11.8b
14.4a
13.1b
a
Means with the same letter within columns at the same storage period are
not signi®cantly different according to Tukey's test (p < 0.05).
1247
GA GonzaÂlez-Aguilar, JG Buta, CY Wang
Low temperatures inhibit mango ripening and enhance senescence.28 However, the ripening of MJtreated fruits stored at 5 °C and then transferred for 7
days at 20 °C was not altered, as indicated by TSS
values. These results agree with those reported
previously for `Tommy Atkins' mangoes.12 It has been
observed that during ripening of mangoes a loss of
lutein occurred concomitant with an increase in
carotenoid content responsible for yellow and red
colour development.29 Olias et al 21 reported that MJ
promotes b-carotene synthesis and chlorophyll degradation in apple peel. The colour change of ripe fruits
from green to yellow may be partly explained by the
unmasking of existing carotenoids as chlorophylls are
degraded. A stimulatory effect of anthocyanin accumulation by MJ treatment was also observed in peach
shoots.30 MJ promoted apple ripening as indicated by
increased ethylene synthesis, acceleration of yellowing
and softening.31 It appears that the response of mango
fruits to MJ depends upon the maturity stage, cultivar
and subsequent storage temperature.
Glucose, fructose and sucrose are major sugars in
mangoes, with sucrose being the predominant one
(Table 2). Treatment with 10 4 M MJ increased the
glucose and sucrose levels of ¯esh tissue of mangoes
stored at 20 °C. Sugar levels of controls and MJtreated fruits are in agreement with TSS contents
found in these fruits (Tables 1 and 2). After cold
storage and shelf-life period, we observed that 10 5 M
MJ treatment signi®cantly increased the sugar and
organic acid contents of mangoes (Tables 2 and 3).
Fruits stored at 20 °C also contained the highest levels
of organic acids (Table 3). Citric acid is the
predominant organic acid in mangoes. The higher
level of organic acids, especially citric acid, in fruits
treated with 10 5 M MJ was correlated with the
maintenance of fruit quality (Fig 3). These fruits also
showed fewer symptoms of CI and deterioration than
control fruits and those treated with 10 4 M MJ. In
mangoes, higher levels of carbohydrates and organic
acids as well as development of yellow and red colours
Table 2. Effect of methyl jasmonate (MJ) vapour treatment
for 20h at 20°C on sugar content of mango fruits after storage
at 5 and 20°Ca
Sugar content (mg g
Treatment
1
FW)
Fructose Glucose Sucrose
14 days at 20 °C
Control
22.3a
15.8b
MJ 10 5 M
25.4a
MJ 10 4 M
68.2b
69.6b
91.7a
91b
85b
118a
14 days at 5 °C ‡ 7 days at 20 °C
Control
24.3b
0.49b
73.4b
MJ 10 5 M
29.4a
0.89a
97.3a
MJ 10 4 M
21.6b
0.68b
92.3a
98c
127a
114b
a
0.49b
0.34b
0.94a
Total
Means with the same letter within columns at the same
storage period are not signi®cantly different according to
Tukey's test (p < 0.05).
1248
Table 3. Effect of methyl jasmonate (MJ) vapour treatment for 20 h at 20°C
on organic acid content of mango fruits after storage at 5 and 20 °Ca
Organic acid content (mg g
Treatment
Malic
1
FW)
Shikimic
Citric
Quinic
Total
14 days at 20 °C
Control
0.033b
0.065a
MJ 10 5 M
0.041b
MJ 10 4 M
0.20b
0.27a
0.19b
0.65b
1.46a
1.23a
0.23a
0.15b
0.22a
1.12c
1.94a
1.68b
14 days at 5 °C ‡ 7 days
Control
0.025b
MJ 10 5 M
0.063a
MJ 10 4 M
0.068a
at 20 °C
0.12b
0.38a
0.33a
0.32b
0.97a
0.40b
0.24a
0.25a
0.19b
0.70c
1.67a
0.99b
a
Means with the same letter within columns at the same storage period are
not signi®cantly different according to Tukey's test (p < 0.05).
are necessary to give the desired ¯avour characteristics
for consumer appeal and acceptance.32
CONCLUSIONS
We found that MJ when applied exogenously to
mango fruits can induce resistance to CI and fungal
development. Treatment with 10 5 M MJ could be
used to improve colour development and maintain
postharvest quality of fruits stored at 20 or 5 °C ‡ shelflife period. The higher concentration of MJ (10 4 M)
could be used to reduce CI symptoms of `Kent'
mangoes.
ACKNOWLEDGEMENTS
The authors wish to thank to David Spaulding and
Hilarine Repace for their technical assistance. Dr
GonzaÂlez-Aguilar received a sabbatical fellowship
from Consejo Nacional de Ciencia y TecnologõÂa
(CONACYT), Mexico.
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