Plant Science 158 (2000) 87 – 96

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Delayed ripening of banana fruit by salicylic acid

Manoj K. Srivastava, Upendra N. Dwivedi*
Department of Biochemistry, Lucknow Uni6ersity, Lucknow 226 007, India

Received 30 July 1999; received in revised form 26 May 2000; accepted 2 June 2000

Abstract

Salicylic acid treatment has been found to delay the ripening of banana fruits (Musa acuminata). Fruit softening, pulp:peel
ratio, reducing sugar content, invertase and respiration rate have been found to decrease in salicylic acid treated fruits as
compared with control ones. The activities of major cell wall degrading enzymes, viz. cellulase, polygalacturonase and xylanase
were found to be decreased in presence of salicylic acid. The major enzymatic antioxidants namely, catalase and peroxidase, were
also found to be decreased in presence of salicylic acid during banana fruit ripening. © 2000 Published by Elsevier Science Ireland
Ltd. All rights reserved.

Keywords: Ripening; Salicylic acid; Banana (Musa acuminata); Catalase

1. Introduction derivative acetyl salicylic acid (ASA) have been

Salicylic acid, a ubiquitous plant phenolic, has
been reported to regulate a number of processes in
plants [1]. One of its earliest known in vivo func-
tions in plants is heat production during flower
induction in some angiosperm species [2]. Later it
was shown to regulate expression of pathogenesis
related protein genes, which suggested its role as a
signal molecule in providing resistance against
pathogen attack. Salicylic acid mediated hypersen-
sitive and systemic acquired resistance against
pathogen attack are proposed to be mediated
through the inhibition of catalase, which subse-
quently raises intracellular H2O2 concentration.
This increased intracellular H2O2 concentration
has been proposed to act as a second messenger in
activation and expression of defense related genes
[1]. Recently, it has been proposed to be a new
kind of plant hormone [3]. Thus, salicylic acid has
been shown to interfere with the biosynthesis and/
or action of ethylene [1], abscisic acid [4] and
cytokinins [5] in plants. Salicylic acid and its
* Corresponding author. Tel.: +91-522-389290.
E-mail address: upendradwivedi@hotmail.com (U.N. Dwivedi).
reported to inhibit ethylene production in pear
[6,7], carrot cell suspension cultures [8], in apple
and pear discs and mung bean hypocotyls [9]
suggesting the role of salicylic acid as an antago-
nist to ethylene action. However, salicylic acid and
ethylene have been reported to have some similar
functions also. For example, both can induce cer-
tain pathogenesis-related proteins and the alterna-
tive pathway of respiration in many plant tissues
[10]. Moreover, an increase in endogenous ethyl-
ene biosynthesis at low concentrations of salicylic
acid has been reported in suspension cultures of
carrot [11]. However, at higher salicylic acid con-
centration (\10 − 4 M) an inhibition of ethylene
production was observed in this study.
The plant hormone ethylene has been reported
to affect a diverse array of plant growth and
developmental processes including germination,
senescence and abscission of both flowers and
leaves, fruit ripening as well as the response to a
wide variety of stresses such as pathogen attack
and drought [12]. The induction of ethylene
biosynthesis by a wide variety of stimuli, including
wounding, pathogen attack, various stresses, me-
chanical stimulus and by hormones such as auxins,

0168-9452/00/$ - see front matter © 2000 Published by Elsevier Science Ireland Ltd. All rights reserved.
PII: S 0 1 6 8 - 9 4 5 2 ( 0 0 ) 0 0 3 0 4 - 6
88 M.K. Sri6asta6a, U.N. Dwi6edi / Plant Science 158 (2000) 87–96
cytokinins and ethylene itself during certain devel-
opmental processes is well documented [12–14].
Among all these effects of ethylene, its role in fruit
ripening has been studied in great detail including
isolation and characterization of gene encoding a
key enzyme of ethylene biosynthesis [15] and de-
velopment of transgenic tomato having delayed
ripening [16]. Treatment of green banana fruits
with ethylene during the preclimacteric phase, has
been shown to accelerate their ripening. All the
climacteric fruits are characterized by a transient
increase in both ethylene synthesis and respiration
at an early stage of ripening. Recently, Vendrell et
al. [17] have attributed the ethylene as a trigger for
ripening in banana.
As salicylic acid has been shown to affect the
biosynthesis and action of ethylene, a well known
fruit ripening hormone, in the present paper, we
have investigated the effects of salicylic acid on
banana fruit ripening by measuring respiration
rate, pulp and peel contents and their ratio, reduc-
ing and non-reducing sugars and their ratio and
invertase, antioxidant enzymes such as catalase
and peroxidase, cell wall degrading enzymes such
as cellulase, polygalacturonase and xylanase.
2. Materials and methods
2.1. Fruit tissue
Banana fingers (Musa acuminata cv Hari chhal)
were purchased from local market.
2.2. Treatment of fruits
Banana fingers were randomized and divided
into groups, containing ten fruits in each, accord-
ing to the following treatments:
Group 1. Water
Group 2. Teepol (0.2%) [Control]
Group 3. Teepol (0.2%)+500 mM salicylic acid
Group 4. Teepol (0.2%)+1000 mM salicylic
acid
Banana fingers were dipped in 2 l of distilled
water (group 1) or teepol (group 2) or teepol
containing salicylic acid (groups 3 and 4) under
laboratory conditions for 6 h with occasional
shaking. After draining the water/solution fingers
were washed, air dried and kept in plastic contain-
ers at room temperature and covered with poly-
thene sheets containing small holes.
2.3. Determination of pulp/peel ratio
Two banana fruits from each treatment were
peeled off and the pulp and peel portions of each
finger were weighed separately on a laboratory
balance. The ratio of pulp to peel of each finger
was calculated and mean value was recorded.
2.4. Determination of respiration rate
Respiration was measured as CO2 production.
A single banana fruit was taken in a chamber and
air was passed through the chamber. The effluent
air was connected to an ADC 225 MK3 Infrared
Gas Analyzer (IRGA) and respiratory rate was
measured. The IRGA was earlier calibrated with
standard CO2. The results were expressed as ml
h − 1 kg − 1 fresh weight.
2.5. Determination of soluble sugars
Banana fruit pulp (1 g) was homogenized in 5.0
ml 90% ethanol by Ultra Turrax T25 for 2 min
and then refluxed for 30 min. The sample was
centrifuged at 10 000×g for 30 min. The residue
was again subjected to ethanol extraction. The
extracts were combined and alcohol was removed
by evaporation. An aliquot was taken and total
sugars were measured by phenol–sulfuric acid
method [18]. Reducing sugars were measured as
described [19]. Non reducing sugar content was
determined by the difference between total sugar
and corresponding reducing sugar value. Glucose
was used as standard for sugar estimation.
2.6. Enzyme extraction and assay
A 10% homogenate was prepared by homoge-
nizing 2 g of pulp in 15 ml sodium phosphate
buffer (20 mM, pH 7.0) containing cysteine–HCl
(20 mM), EDTA (20 mM) and Triton X-100
(0.05%) by Ultra Turrax T25 for 2 min at high
speed. The homogenate was passed through two
layers of muslin cloth to remove cell debris and
the volume was made to 20 ml with the homoge-
nization medium. The homogenate was cen-
trifuged at 15 000×g for 30 min at 4°C in a
Sorvall RC 5C refrigerated centrifuge. The clear
supernatant was used for enzyme assay.
 M.K. Sri6asta6a, U.N. Dwi6edi / Plant Science 158 (2000) 87–96
2.7. Assay of polygalacturonase (EC 3.2.1.15)
Polygalacturonase activity was assayed by mea-
suring the formation of reducing groups using the
methods of Nelson [20] and Somogyi [21]. The
reaction mixture contained 0.2 ml sodium acetate
buffer (200 mM, pH 4.5), 0.1 ml NaCl (200 mM),
0.3 ml polygalacturonic acid (PGA) (1%, pH 4.5),
and an appropriate amount of enzyme in a total
volume of 1.0 ml. The reaction was initiated by the
addition of substrate. The reaction mixture was
incubated at 37°C for 1 h. The reaction was
terminated by heating the reaction mixture in a
boiling waterbath. In the control tubes, the sub-
strate was added after the heat treatment. The
formation of reducing group was calculated using
D-galacturonic acid as a standard. One unit of
polygalacturonase activity is defined as the
amount of enzyme producing 1 mmol of reducing
groups per min at 37°C.
2.8. Assay of cellulase (EC 3.2.1.4)
Cellulase activity was measured as described by
Ahmed and Labavitch [22] with slight modifica-
tions. The reaction mixture consisted of sodium
acetate buffer (100 mM, pH 5.0), carboxy methyl
cellulose (1.5% w/v) and enzyme in a final volume
of 1.0 ml. The reaction mixture was incubated at
37°C for 16 h. After 16 h, the substrate was added
to the control tubes and colour was developed
according to Miller [19] using dinitrosalicylic acid
(DNS). The tubes were boiled on a waterbath for
10 min and the colour was read at 540 nm using
Spectronic 20D spectrophotometer. Amount of re-
ducing sugar released was calculated from a cali-
bration curve drawn using glucose as standard.
One unit of cellulase activity was defined as the
amount of enzyme liberating 1 mmol of reducing
sugar per min at 37°C.
2.9. Assay of xylanase (EC 3.2.1.8)
Xylanase activity was assayed as described by
Singh and Singh [23] with slight modifications.
The assay mixture consisted of sodium acetate
buffer (100 mM, pH 5.0), xylan (0.1%) and en-
zyme preparation in a total volume of 1.0 ml. The
mixture was incubated for 1 h at 37°C. The re-
leased reducing sugars were measured using DNS
as for cellulase (described above). One unit of
89
xylanase activity was defined as 1 mmol of reduc-
ing sugar released per min at 37°C.
2.10. Assay of in6ertase (EC 3.2.1.26)
Invertase activity was measured as described by
Moriguchi et al. [24] with slight modifications. The
assay mixture contained acetate buffer (100 mM,
pH 4.5), sucrose (100 mM) and enzyme prepara-
tion in a total volume of 1.0 ml. The reaction
mixture was incubated for 1 h at 37°C. The sub-
strate was added to control tubes after the incuba-
tion and colour was developed using DNS (as
described for cellulase). Amount of reducing sugar
released was calculated from the calibration graph.
One unit of invertase activity was defined as
micromoles of reducing sugars equivalent released
per min at 37°C.
2.11. Assay of peroxidase (EC 1.11.1.7)
The peroxidase activity was assayed as de-
scribed by Pütter [25] with slight modifications.
The assay mixture consisted of sodium phosphate
buffer (50 mM, pH 7.0), H2O2 (4mM), guaiacol
(3.33 mM) and 0.1 ml enzyme in a final volume of
3.0 ml. The increase in absorbance at 470 nm was
measured using a Spectronic-2000 spectrophoto-
meter. One unit of enzyme activity was defined as
the amount of enzyme catalyzing the production
of 1 mmol tetraguaiacol per min at 30°C.
2.12. Assay of catalase (EC 1.11.1.6)
The catalase activity was assayed as described
by Aebi [26] with some modifications. The assay
medium consisted of sodium phosphate buffer (50
mM, pH 7.0), (20 mM) and 0.1 ml enzyme in a
final volume of 3.0 ml. The disappearance of H2O2
was measured as decrease in absorbance at 240 nm
using a Spectronic-2000 spectrophotometer. One
unit of enzyme activity was defined as the amount
of the enzyme catalyzing the decomposition of 1
mmol H2O2 per min at 30°C.
3. Results
3.1. Effect of salicylic acid on the rate of
respiration during banana fruit ripening
The effects of salicylic acid on the rate of respi-
90 M.K. Sri6asta6a, U.N. Dwi6edi / Plant Science 158 (2000) 87–96

3.2. Effect of salicylic acid on softening of banana

fruits during its ripening

Softening of fruits is one of the most common

physical parameter to assess the progress of ripen-
ing. Therefore, the effect of salicylic acid on soft-
ening of banana fruits, as a physical parameter to
assess the extent of ripening, has been investigated.
It was observed that salicylic acid treatment inhib-
ited the process of banana fruit softening during
ripening. Another physical parameter, yellowing
of peel was also found to be less in salicylic acid
treated than those of control fruits. Thus, the
control fruits were soft enough at day 8 of treat-
ment while the salicylic acid treated fruits were
still hard.

Fig. 1. Effect of salicylic acid on rate of respiration during
ripening of banana fruit in the absence ( — ) and presence
of 500 (
—
) and 1000 (—) mM salicylic acid. Each
value represents a mean of three independent replicates.
ration during banana fruit ripening have been
investigated. Results are shown in Fig. 1. In
control fruits (in the absence of salicylic acid) the
rate of respiration was found to increase with
ripening exhibiting a peak on day 4 (climacteric
peak). Rate of respiration has been reported to
increase with ripening of all climacteric fruits.
Salicylic acid treatment resulted in a decrease in
the rate of respiration as well as a delay in
appearance of climacteric peak, in a concentration
dependent manner, as compared to control. There
was a gradual increase in respiration rate of
salicylic acid treated fruits up to 6th day of
treatment suggesting a delay in ripening
(respiratory burst).
3.3. Effect of salicylic acid on pulp and peel
contents of banana fruit during its ripening
Effect of salicylic acid on pulp and peel contents
of banana fruits have been investigated during its
ripening. Pulp content increased throughout the
experimental period both in control and salicylic
acid treated banana fruits. However, salicylic acid
treatment resulted in decreased pulp content of
fruits in a concentration dependent manner. On
the other hand, peel content decreased throughout
the ripening both in the absence as well as pres-
ence of salicylic acid. However, the decrease in
peel content was less in salicylic acid treated fruits
as compared with that of control. Because the
pulp to peel ratio of fruits is taken as an important
parameter for fruit ripening, we have presented the
data in the form of their ratio. (Table 1). It is
noteworthy that pulp to peel ratio increased with

Table 1
Effect of salicylic acid on ratio of pulp to peel of banana fruits during its ripeninga

Concentration of salicylic acid (mM) Pulp to peel ratio

Days

0 2nd 4th 5th 6th

0 (control) 1.4090.22 1.549 0.13 1.709 0.12 1.90 9 0.17 2.0090.18

500 1.4090.22 1.5090.13 1.58 9 0.18 1.60 9 0.20 1.6490.14
1000 1.4090.22 1.469 0.08 1.52 90.07 1.56 9 0.07 1.6390.15

a
Each value represents a mean 9S.D. of three independent replicates.
 M.K. Sri6asta6a, U.N. Dwi6edi / Plant Science 158 (2000) 87–96
Fig. 2. Effect of salicylic acid on reducing (A), nonreducing
(B) sugars and invertase (C) activity during ripening of ba-
nana fruit in the absence ( — ) and presence of 500
(
—
) and 1000 (—) mM salicylic acid. Each value
represents a mean of three independent replicates.
the progress of ripening both in the absence and
presence of salicylic acid. However, salicylic acid
treatment decreased the pulp to peel ratio in a
concentration dependent manner. Our data of
pulp to peel ratio suggested a delay in ripening of
banana fruit in the presence of salicylic acid.
3.4. Effect of salicylic acid on soluble sugar
content of banana fruit during ripening
Effect of salicylic acid on reducing and nonre-
ducing sugar contents of banana fruit has been
studied during ripening. The results are shown in
Fig. 2. Reducing sugar content increased both in
control and salicylic acid treated fruits throughout
the ripening (Fig. 2A). However, salicylic acid
91
treatment resulted in, a concentration dependent
manner, decreased levels of reducing sugar than
those in the absence of salicylic acid (control)
throughout the ripening period of banana fruits.
On the other hand, the measurement of nonreduc-
ing sugar content of banana fruit revealed an
opposite trend to that of reducing sugar, during
ripening (Fig. 2B). Thus, on day 4 of ripening,
salicylic acid treatment resulted in a decrease of
total reducing sugar by 25 and 42% at 500 and
1000 mM salicylic acid concentration, respectively,
while it resulted in an increase of nonreducing
sugar by 9 and 20% at 500 and 1000 mM salicylic
acid, respectively.
3.5. Effect of salicylic acid on in6ertase during
banana fruit ripening
Effect of salicylic acid on invertase activity has
been investigated during ripening of banana fruit.
The results are shown in Fig. 2C. Invertase activ-
ity increased throughout the ripening both in con-
trol and in salicylic acid treated fruits. However,
salicylic acid treatment resulted in decreased levels
of invertase than those of controls, in a concentra-
tion dependent manner, during banana ripening.
Thus, on day 4, salicylic acid treatment resulted in
a decrease of invertase activity by 51 and 71% of
the control value at 500 and 1000 mM salicylic acid
concentrations, respectively, during the banana
fruit ripening.
3.6. Effect of salicylic acid on cell wall degrading
enzymes during banana fruit ripening
Effect of salicylic acid on the developmental
profiles of major cell wall degrading enzymes
namely, cellulase, polygalacturonase (PG) and xy-
lanase in the absence and presence of salicylic acid
have been investigated during ripening of banana
fruits. Results are shown in Fig. 3. It is notewor-
thy that the activity of all the three major cell wall
degrading enzymes increased gradually both in the
absence and presence of salicylic acid during the
ripening of banana fruits. However, the salicylic
acid treatment resulted in decreased levels of all
these cell wall degrading enzymes in a concentra-
tion dependent manner, during the ripening of
banana. Thus, on day 6, the activities of cellulase,
polygalacturonase and xylanase decreased by 55
and 82, 33 and 45, 35 and 50% of their respective
92 M.K. Sri6asta6a, U.N. Dwi6edi / Plant Science 158 (2000) 87–96
controls at 500 and 1000 mM salicylic acid, respec-
tively during banana fruit ripening. Thus, out of
the three cell wall degrading enzymes studied,
cellulase seems to be most sensitive to salicylic acid
inhibition. Furthermore, the two cell wall degrad-
ing enzymes namely, cellulase and polygalactur-
onase seems to play a major role in banana fruit
ripening as their activities increased several folds
during ripening in comparison to that of xylanase.
Similarly, salicylic acid inhibited cellulase and
polygalacturonase activity more strongly than that
of xylanase.
Fig. 3. Effect of salicylic acid on cellulase (A), polygalactur-
onase (B) and xylanase (C) activity during ripening of banana
fruit in the absence ( — ) and presence of 500 (
—
) by an increase in the reducing sugar content and
and 1000 (—) mM salicylic acid. Each value represents a
mean of three independent replicates.
3.7. Effect of salicylic acid on catalase and
peroxidase during banana fruit ripening
As catalase and peroxidase have also been re-
ported to be affected during fruit ripening, the
effect of salicylic acid on these two enzymes has
been investigated during banana fruit ripening.
Results are shown in Table 2. Salicylic acid treat-
ment resulted in decreased levels of catalase and
peroxidase, than their respective controls, in a
concentration dependent manner, during the
ripening of banana fruits. Thus, the effect of sali-
cylic acid on these two enzymes, during banana
fruit ripening, revealed a similar response as ob-
served for that of the pea during seedling growth
[27].
4. Discussion
The data presented in this paper suggest that
salicylic acid delays banana fruit ripening. Thus,
the rapid rise in respiration during ripening of
banana fruits, an observation typical of climacteric
fruits, was found to be delayed by salicylic acid in
a concentration dependent manner.
The ripening of banana fruit was accompanied
by increase in pulp to peel ratio. Similar observa-
tion has been reported by others [28,29]. Rise in
pulp to peel ratio during fruit ripening was sug-
gested to be due to change in sugar concentration
in the two tissues. A rapid increase in sugar con-
tents in the pulp than those in the peel leads to a
change in osmotic pressure, as a result of which
water is withdrawn from the peel and hence pulp
to peel ratio increases accordingly. Salicylic acid
treatment reduced this increase in pulp to peel
ratio, in a concentration dependent manner, lead-
ing to a delay in banana fruit ripening. Delay in
ripening of banana is reported by ABA [30] and
GA3 [31]. GA3 is also reported to delay ripening in
tomato pericarp disc [31] and mango [32]. Post
harvest dips of mature unripe Alphonso (variety
of mango) fruits in solution of GA3 (10 − 6 M),
IAA (10 − 6 M) and kinetin (10 − 5 M) are reported
to delay ripening [33].
The data on soluble sugar content and invertase
suggested that banana ripening was accompanied
invertase activity concomitant with decrease in
nonreducing sugar content. Salicylic acid treat-
Table 2
Effect of salicylic acid on catalase and peroxidase activities of banana fruits during ripeninga

Concentration of Catalase activity (U/ml) Peroxidase activity (U/ml)

salicylic acid (mM)

Number of days Number of days

2nd 4th 6th 8th 0 2nd 4th 6th 8th

0 (control) 0.091 90.015 0.04790.011 0.0279 0.006 0.1299 0.006 0.0279 0.007 0.0099 0.004 0.012 9 0.005 0.014 9 0.003 0.036 9 0.004
500 0.091 90.015 0.03790.007 0.01590.006 0.0759 0.012 0.0279 0.007 0.004 90.002 0.006 9 0.001 0.011 9 0.003 0.016 9 0.006
1000 0.09190.015 0.02990.011 0.0129 0.004 0.0679 0.010 0.02790.007 0.0029 0.001 0.003 9 0.001 0.008 9 0.003 0.012 9 0.004

a
Each value represents a mean 9S.D. of three independent replicates.
M.K. Sri6asta6a, U.N. Dwi6edi / Plant Science 158 (2000) 87–96
93
94 M.K. Sri6asta6a, U.N. Dwi6edi / Plant Science 158 (2000) 87–96
ment resulted in decreased levels of invertase and
reducing sugar contents while an opposite effect
on nonreducing sugar content, suggesting that
salicylic acid delays banana fruit ripening. This
accumulation of reducing sugars may be due to
increased breakdown of starch during ripening as
reported by Beaudry et al. [34]. Increased starch
phosphorylase activity has been reported during
development and ripening of banana fruits
[35,36]. Recently, an increase in sucrose phos-
phate synthase and decrease in sucrose synthase
activity have been reported during ripening of
banana fruits [37]. Increase in invertase activity
has been shown during papaya fruit ripening [38].
The data on cell wall degrading enzymes sug-
gested that banana ripening was accompanied by
an increase in all the three cell wall degrading
enzymes namely, cellulase, polygalacturonase and
xylanase. Levels of these cell wall degrading en-
zymes were found to be decreased, in a concen-
tration dependent manner, suggesting that
salicylic acid delays ripening of banana. Poly-
galacturonase is reported to be primarily respon-
sible for ripening associated pectin degradation
and fruit softening [39]. Level of polygalactur-
onase activity has been positively correlated with
fruit ripening and softening in banana [29]. Poly-
galacturonase activity [40], mRNA [41] and gene
transcription [42,43] are reported to increase at
the onset of ripening of tomato fruits. Further-
more, tomato fruit ripening mutants, with de-
layed ripening exhibited decreased levels of
polygalacturonase activity [44], mRNA [45] and
gene transcription [43], while the activities of sev-
eral other cell wall enzymes appeared to be af-
fected to a lesser extent. Also, physical and
chemical treatments which suppress ripening, in-
hibit polygalacturonase gene expression [46–48].
Carboxymethyl cellulase (Cm –cellulase) activity
increases during ripening of tomato [49,50],
strawberry, pear, peach [51] and avocado [52].
Modifications of hemicellulose structure of cell
wall, during fruit ripening have been reported in
tomato, pepper, strawberry and melons. The sizes
of hemicellulose polymers are found to decrease
during ripening of these fruits [39,53,54]. Ripen-
ing associated changes in hemicellulose structure
are proposed to be responsible for the textural
change in fruit.
Data on catalase and peroxidase revealed that
ripening of banana fruit was accompanied by an
increase in the activities of catalase and perox-
idase while levels of these enzymes were found to
be decreased in presence of salicylic acid, in a
concentration dependent manner. Ripening of pa-
paya fruit has been reported to be accompanied
by an increase in the catalase and peroxidase
activity [38]. Salicylic acid has been demonstrated
to inhibit catalase and peroxidase in pea [26].
The inhibition of catalase by salicylic acid is sug-
gested to be one of modes of action of salicylic
acid in providing defense against pathogen attack
[27,55].
Thus, the data presented here unequivocally
suggest that salicylic acid delays banana fruit
ripening. Based on these observations salicylic
acid may be considered as antagonist of ethylene,
a major fruit ripening hormone. Salicylic acid is
reported to inhibit biosynthesis of ethylene [7,8].
Recently, salicylic acid has also been shown to
inhibit ACC oxidase activity [56]. Even in the
studies of Nissen [11] in which salicylic acid (at
low concentration) has been shown to stimulate
ethylene biosynthesis, an inhibition of ethylene
production occurs at higher concentrations of
salicylic acid (\10 − 4 M). Therefore, in the
present study since we have taken a much higher
concentration of salicylic acid than those of Nis-
sen [11] (\10 − 4 M), only the inhibition of ethyl-
ene biosynthesis is expected to occur in our case.
Ethylene synthesis is normally limited by the sup-
ply of the immediate precursor amino cyclo-
propane-1-carboxylic acid (ACC). There is a
sharp rise in ACC synthase activity and the con-
tent of ACC during ripening [57]. Thus, present
studies on the effect of salicylic acid in delaying
the ripening of banana fruits may be thought of
due to inhibition of ethylene biosynthesis and/or
action. However, this needs further investiga-
tions.
Acknowledgements
The financial assistance (in the form of SRF to
M.K. Srivastava) of the Council of Scientific and
Industrial Research (CSIR), New Delhi, is grate-
fully acknowledged. The grants from the Depart-
ment of Biotechnology, New Delhi; University
Grants Commission, New Delhi under the Spe-
cial Assistance Programme and COSIST Pro-
gramme to the Department of Biochemistry is
also acknowledged.
 M.K. Sri6asta6a, U.N. Dwi6edi / Plant Science 158 (2000) 87–96
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