Cycloheximide Efficacy Varies Temporally

in Improving Postharvest Performance of

Cool Wet Stored Dianthus chinensis L. Cut
Sprays
Riyaz Ahmad Dar, Inayatullah Tahir &
Syed Sabhi Ahmad

Proceedings of the National

Academy of Sciences, India Section B:
Biological Sciences
ISSN 0369-8211
Proc. Natl. Acad. Sci., India, Sect. B Biol.
Sci.
DOI 10.1007/s40011-015-0584-z

1 23

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Author's personal copy

Proc. Natl. Acad. Sci., India, Sect. B Biol. Sci.
DOI 10.1007/s40011-015-0584-z

RESEARCH ARTICLE

Cycloheximide Efficacy Varies Temporally in Improving

Postharvest Performance of Cool Wet Stored Dianthus chinensis
L. Cut Sprays
Riyaz Ahmad Dar1 Inayatullah Tahir1

Syed Sabhi Ahmad1

Received: 8 February 2015 / Revised: 15 April 2015 / Accepted: 9 June 2015

 The National Academy of Sciences, India 2015

Abstract An experiment was designed to determine the

synergistic effect of cold temperature and cycloheximide
(CHI) on the postharvest performance of cut sprays of
Dianthus chinensis. Sprays were harvested at 800 h with
the oldest bud at paint brush stage, cut to a length of 27 cm
and wet stored at 5 C for 72 h before or after 1 h 0.1 mM
CHI pulse. Another set of sprays was pulsed with 0.1 mM
CHI for 1 h during storage (after 35 h). A set of
unpulsed sprays stored at 5 C represented control. The
treatment effects were evaluated by placing the sprays in
holding solution comprising of distilled water and sucrose
(SUC) (0.05 M) at room temperature (21 2 C) in the
laboratory. The sprays wet stored at 5 C and pulse treated
with 0.1 mM CHI after 72 h wet storage showed an
increment of vase life by 3 days; besides sustained rate of
blooming, higher fresh and dry mass, floral diameter and
sugar fractions, but maintained lower soluble protein content in comparison to the control. Vase life, fresh mass, dry
mass and floral diameter of sprays wet stored at 5 C and
1 h pulse treated with 0.1 mM CHI before and during
storage was considerably lower than the sprays 1 h pulse
treated with 0.1 mM CHI after storage, but significantly
higher than that of the control. The study revealed that 1 h
pulse treatment with 0.1 mM CHI after 72 h wet storage at
5 C, followed by transfer to SUC can be used as an
effective postharvest storage protocol to bring about the
transportation of this beautiful cut flower within 72 h.

& Inayatullah Tahir

inayatullahtahirku@gmail.com;
inayatullahtahir@gmail.com
1

Plant Physiology and Biochemistry Research Laboratory,

Department of Botany, University of Kashmir,
Srinagar 190006, India

Keywords Dianthus chinensis
Cycloheximide

Postharvest storage
Proteins

Introduction
Caryophyllaceae, a dicotyledonous family belonging to
order Caryophyllales comprises wide range of important
ornamentals. Among these, Dianthus chinensis is grown
throughout the world as an ornamental plant and has a vast
potential in global cut flower industry. The vase life of
various species of Dianthus varies greatly not only among
each other but also with the growing season and it has been
reported that the same Dianthus species grown in two
different seasons showed a significant difference in the
vase life varying from 8 to 10 days in FebMar to 45 days
in AprilMay [1]. Dianthus chinensis blooms from May to
August in Kashmir valley and possesses flowers of various
hues (from pink to white) borne on medium sized sprays
(3040 cm). Each individual spray bears about 45 flowers. The average field life of an individual flower is about
4 days. However the sprays held in distilled water (DW)
last for 7 days after harvesting with the oldest bud at paint
brush stage.
Flower senescence in plants is chiefly regulated by an
interaction between various growth regulators like ethylene, gibberellins, cytokinins, abscisic acid and auxins.
Ethylene is the main candidate for senescence in ethylene
sensitive flowers while as in ethylene insensitive flowers it
has little or no role to play [25]. The genus Dianthus falls
under the ethylene sensitive group as the senescence is
chiefly governed by ethylene [6, 7]. Factors like pollination, drought and temperature also influence senescence by
altering the balance of hormones produced by the flowers
[8]. Temperature affects rate of respiration, response to

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ethylene, loss of moisture and apparent damages in various

flowers such as Narcissus, Iris and Consolida [912]. The
upsurge in ethylene production has been found to regulate
the expression of genes for protease, thereby leading to
overall protein degradation in senescing floral parts [9, 13].
In addition, the expression of proteins leading to death of
floral tissues has been reported during senescence in various flowers. Proteolytic cleavage seems to play an important role, because the expression of the protease genes
manifests the initial senescence related changes in the
genes [14]. Flower senescence in various plants such as Iris
and Consolida have been shown to be delayed by various
protein synthesis inhibitors, suggesting that senescence in
these flowers is initiated by proteins synthesized de novo
[11, 15]. It is in this perspective that a novel strategy
involving the combined effect of protein synthesis inhibitor, cycloheximide (CHI) along with cool wet storage
was employed which may result in maintenance of flower
quality for adjustability in the floriculture market and significantly curtail the postharvest loss. Moreover, the study
was aimed to simulate a long term transportation protocol
using CHI along with cool wet storage in the cut sprays of
D. chinensis.

Material and Methods

Uniform and healthy cut sprays of D. chinensis grown in
Kashmir University Botanic Garden (KUBG) were utilized
for the study. The sprays were harvested at 800 h, when the
oldest bud was at paint brush stage (optimal harvest
maturity stage) in the latter half of May. The cut sprays
were brought to laboratory with the cut ends immersed in
DW, defoliated, cut to a uniform length of 27 cm and
divided into four sets. One set of sprays was pulsed with
0.1 mM CHI for 1 h before processing for cool wet storage
(5 C) for 72 h. Another set of cut sprays was pulsed with
0.1 mM CHI for 1 h after 72 h cool wet storage. The third
set of cut sprays was 1 h pulsed with 0.1 mM CHI during
(after 35 h) 72 h cool wet storage. The fourth set of
sprays kept unpulsed and wet stored at 5 C for 72 h designated the control (Fig. 1). For 72 h cool wet storage, the
cut sprays were held in 500 ml of DW and kept in incubator at 5 C with relative humidity of 60 10 %. The
samples were taken out of the incubators after 72 h and
held at room temperature after transferring them to 100 ml
Erlenmeyer flasks containing 75 ml of either DW or
0.05 M sucrose (SUC). Each test solution was represented
by 5 replicates, with each flask holding two cut sprays. All
the samples were maintained under cool white florescent
lamp with a mix of diffused light (10 W/m-2) for 12 h a
day and a relative humidity of 60 10 %. The day of
transfer of cut sprays to holding solutions was designated

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Fig. 1 Sprays of Dianthus chinensis held in distilled water before

(a) and after (b) 72 h wet storage at 5 C. Control: samples kept at
5 C for 72 h without 0.01 mM cycloheximide (CHI) pulse treatment.
BWS: Samples 1 h pulse treated with 0.01 mM CHI before 72 h wet
storage at 5 C. DWS: Samples 1 h pulse treated with 0.01 mM CHI
during 72 h wet storage at 5 C. AWS: Samples 1 h pulse treated with
0.01 mM CHI after 72 h wet storage at 5 C. (BWS: before wet
storage, DWS: during wet storage, AWS: after wet storage)

as day zero (D0). The average vase life of the cut sprays
was counted from day 1 and was assessed to be terminated
when the final flower lost its ornamental value. Fresh and
dry mass, floral diameter, number of blooms, volume of
holding solution absorbed per spray, soluble proteins, aamino acids, phenols and sugar fractions were estimated on
day 4 and 12 of transfer to holding solutions. Estimation of
dry mass was done by drying the material in an oven for
48 h at 70 C.
Protein extraction was performed on 1 g petal tissue
drawn separately from ten different flowers. The tissue was
homogenized in 5 ml of 5 % sodium sulphite (w/v) adding
100 mg of polyvinylpyrrolidone and centrifuged. Proteins
were precipitated from a suitable volume of cleared
supernatant with equal volume of 20 % trichloroacetic
acid, centrifuged at 20009g for 15 min and the pellet redissolved in 0.1 N NaOH. Proteins were estimated by the
method of Lowry et al. [16] from suitable aliquot using
BSA as the standard. Reducing sugars were estimated by
Nelsons method [17] using glucose as standard. Total
sugars were estimated as reducing sugars by conversion of
non-reducing sugars to reducing sugars by using invertase.
Non-reducing sugars were computed as the difference
between total and reducing sugars.
For the estimation of specific protease activity, 1 g of
prechilled petal tissue was homogenized in 15 ml cold
0.1 M phosphate buffer (pH 6.5) in a pre-cooled glass
pestle and mortar. Protease activity was determined by the
modification of the method as described by Tayyab and
Qamar [18]. One ml of reaction mixture (0.1 % BSA dissolved in 0.1 M phosphate buffer, pH 6.5) was mixed with

Author's personal copy

Cycloheximide Efficacy Varies Temporally in Improving Postharvest Performance of Cool Wet

1 ml of enzyme extract. The activity of enzyme has been

expressed as microgram tyrosine equivalents liberated per
milligram protein per minute (lg tyr. mg protein-1 min-1).
a-amino acids were determined by Rosens method [19]
using glycine as standard. Total phenols were estimated by
the method of Swain and Hillis [20] with gallic acid as the
standard.
Completely randomized experimental design was followed during the experiment. The experiment was repeated
thrice for reproducibility of the results obtained. The data
has been statistically analyzed and LSD computed at P0.05
using MINITAB (v 15. 1.2-EQUINOX_Softddl.net)
software.

Results and Discussion

Fig. 2 Sprays of Dianthus chinensis held in distilled water (DW) or
0.1 M sucrose (SUC) after 72 h wet storage at 5 C at day 2 (a), day 8
(b) and day 20 (c) of transfer. From left to right are arranged flasks
containing sprays (previously 72 h wet stored at 5 C) held in either
distilled water (DW) or sucrose (SUC). The digits 1 to 8 represents
the nature of the treatment as (1) 5 C ? DW; (2) 5 C ? SUC; (3)
CHI 0.01 mM 1 h pulse BWS ? DW; (4) CHI 0.01 mM 1 h pulse
BWS ? SUC; (5) CHI 0.01 mM 1 h pulse DWS ? DW; (6) CHI
0.01 mM 1 h pulse DWS ? SUC; (7) CHI 0.01 mM 1 h pulse
AWS ? DW and (8) CHI 0.01 mM 1 h pulse AWS ? SUC. (DW:
distilled water, SUC: sucrose, CHI: cycloheximide, BWS: before wet
storage, DWS: during wet storage, AWS: after wet storage)

25

20

Vase life (days)

The greenish buds of D. chinensis open into a wide range

of colored flowers (from pink to white). The flower lasts for
about 4 days after it opens fully on mature spray in DW in
the laboratory and field. Senescence starts with loss of petal
turgidity and is followed by petal inrolling and wilting
which is the characteristic feature of the members of family
Caryophyllaceae [3, 21]. The average life of cut sprays
harvested with their oldest bud at paint brush stage is about
7 and 10 days in DW and SUC respectively. The synergistic effect of cold storage along with the CHI treatment
showed an increase in vase life of the cut sprays of D.
chinensis (Figs. 2, 3). Cool temperature has been found to
minimize microbial growth, curtail the metabolic processes
and increase the vase life in various other flowers such as
Iris, Consolida, Hemerocallis, Ranunculus and Narcissus
thereby delaying senescence [2225]. CHI, that inhibits
protein synthesis at translational level, has been reported to
exhibit an important role in delaying senescence of various
flowers like Consolida, Iris and Hemerocallis [9, 12, 26].
The vase life of the cut sprays pulsed with 0.1 mM CHI
either before, during or after 72 h cool wet storage showed
increment in the vase life as compared to the control
(without CHI pulse treatment) where it was 14 and 16 days
in DW and SUC. Maximum vase life of about 21 days was
recorded in the sprays pulse treated with CHI after storage
and transferred to SUC as compared to 18 days in the
corresponding sprays transferred to DW followed by the
cut sprays treated with CHI during storage which showed
the vase life of 16 and 17 days in DW and SUC respectively (Figs. 2, 3). The enhanced vase life in CHI post
storage treated cut sprays could be attributed to the fact that
in the sprays treated with CHI before or during storage, the
uptake of CHI may be retarded because of storage at low
temperature (5 C). At low temperature (5 C) the water
uptake is decreased and as such CHI is not transported to
flowers effectively. While as, in case of the sprays treated

15

10

0
Control

CHI
BWS

CHI
DWS

CHI Control CHI

AWS
BWS

DW

CHI
DWS

SUC

CHI
AWS

LSD
P 0.05

Treatments

Fig. 3 Effect of 1 h pulse treatment with 0.01 mM cycloheximide

before, during and after 72 h wet storage (BWS, DWS, AWS) at 5 C
and subsequent transfer to distilled water (DW) and sucrose (SUC) on
vase life in cut sprays of Dianthus chinensis. (DW: distilled water,
SUC: sucrose, CHI: cycloheximide, BWS: before wet storage, DWS:
during wet storage, AWS: after wet storage)

with CHI after storage, the sprays are kept at room temperature and as such pulse of CHI solution gets transported
up to xylem which becomes quickly available in eliciting

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the desired effect efficiently and thereby enhancing vase

life. However, the earlier studies on Consolida and Iris
reported that treatment of CHI before storage enhanced
vase life effectively than the post storage CHI treatment [9,
12].
The enhancement of vase life by CHI has been suggested to be because of the prevention in upregulation of so
called death proteins [15, 27]. Treatment of flowers with
various inhibitors of protein synthesis has been found to
increase the time to apparent symptoms of senescence
suggesting that active protein synthesis is needed for cell
death execution [28, 29].
Samples from the cut sprays treated with CHI after 72 h
wet storage at 5 C maintained a higher floral diameter and
fresh or dry mass (Table 1). These parameters showed a
significant decrease with the progression in time from D4
to D12. Low temperature along with CHI has been found to
improve the fresh and dry mass in the flowers of rose and
Consolida [9, 30]. Increased fresh and dry mass and floral
diameter manifested by low temperature has been found to
be due to decreased respiratory loss in some flowers like
Hemerocallis and rose [31, 32]. Since uptake of CHI was
more in the cut sprays treated after storage, as such maximum fresh and dry mass was recorded in those cut sprays.
The increase in fresh mass, dry mass and floral diameter

was augmented when SUC was present as vase solution

(Table 1). This effect of SUC could be because SUC
mainly acts as a metabolite; besides as a signaling molecule. Once the sprays are detached from the parent plant,
they are to be supplied with the carbohydrate source for
continuation of the cellular metabolism [15]. SUC alone or
in combination with various biocides has been shown to
enhance the postharvest performance of various cut flowers
like Amaryllis, Consolida, Iris, Delphinium and Nerine [11,
3337].
The cut sprays treated with CHI after 72 h wet storage
showed improved rate of blooming (92 %) as compared to
control where it was only 52 %, besides showing least
absorption of the vase solution as against the control and
other treatments (Table 1). The increased rate of blooming
and reduced absorption of vase solution by CHI post
treated sprays could be because of the property of CHI to
minimize the respiration and water loss through the plant
parts [3, 36]. Decreased water uptake from the vase solution can be partly attributed to the already saturated state of
sprays with water.
Lower soluble protein content was observed in all the
samples treated with CHI before, during or after 72 h wet
storage. The content of soluble protein registered a
decrease with the progression in time from D4 to D12,

Table 1 Effect of pretreatment with cycloheximide (0.01 mM CHI,

1 h pulse) before, during and after 72 h wet storage at 5 C (BWS,
DWS and AWS) and subsequent transfer to (DW) and (SUC) on fresh

and dry mass, floral diameter, number of blooms and volume of

holding solution absorbed per spray at day 4 and 12 (D4 and D12) of
transfer in samples from cut sprays of Dianthus chinensis

Days after transfer (D) Set A (DW)

Set B (SUC)

Control CHI
(-CHI) (BWS)

CHI
(DWS)

CHI
(AWS)

Control
(-CHI)

CHI
(BWS)

CHI
(DWS)

CHI
(AWS)

LSD
P B 0.05

Fresh mass flower-1(g)

D4

0.228

0.228

0.235

0.298

0.232

0.226

0.237

0.301

0.05

D12

0.212

0.215

0.229

0.29

0.228

0.224

0.234

0.301

0.054

Dry mass flower-1 (g)

D4

0.042

0.045

0.056

0.061

0.045

0.04

0.043

0.063

0.02

D12

0.032

0.034

0.039

0.053

0.041

0.038

0.038

0.059

0.01

Floral diameter (cm)

D4

2.685

3.83

4.1

4.2

3.86

4.06

4.16

4.5

0.41

D12

2.693

3.87

4.11

3.231

3.86

4.07

4.16

4.52

0.17

4.2 (42) 4.7 (42.4)

4.62 (43)

5.42 (49.4)

5.14 (42.5)

4.62 (42.2)

4.62 (48.92)

5.92 (57.92) 0.72

4.7 (52) 5 (52.5)

5.2 (54.5) 6.5 (91.3)

6.01 (63)

4.09 (55.2)

5.33 (64.2)

7.04 (92)

0.28

Blooms/spray
D4
D12

Volume of holding solution absorbed/spray (ml)

D4

0.66

6.78

6.98

5.45

0.33

5.54

5.67

4.93

0.51

D12

11

14.54

12.89

9.66

10.83

12.98

11.49

11.04

0.42

Each value is the mean of 10 independent replicates

The values in the parenthesis represent percent blooms
DW: distilled water, SUC: sucrose, CHI: cycloheximide, -CHI: without CHI, BWS: before wet storage, DWS: during wet storage, AWS: after
wet storage

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Cycloheximide Efficacy Varies Temporally in Improving Postharvest Performance of Cool Wet
Table 2 Effect of pretreatment with cycloheximide (0.01 mM CHI,
1 h pulse) before, during and after 72 h wet storage at 5 C (BWS,
DWS and AWS) and subsequent transfer to (DW) and (SUC) on

soluble protein content, a-amino acids, total phenols, reducing, nonreducing and total sugars at day 4 and 12 (D4 and D12) of transfer in
samples from cut sprays of Dianthus chinensis

Days after transfer Set A (DW)

Control
(-CHI)

Set B (SUC)
CHI
(BWS)

CHI
(DWS)

CHI
(AWS)

Control
(-CHI)

CHI
(BWS)

CHI
(DWS)

CHI
(AWS)

LSD
P B 0.05

Soluble proteins (mg g-1 FM)

D4

2.08 (0.56)

1.86 (0.53)

1.36 (0.32)

1.04 (0.31)

2.12 (0.49)

1.2 (0.27)

0.78 (0.18)

1.02 (0.30) 0.041

D12

1.96 (0.41)

1.82 (0.39)

1.29 (0.29)

0.98 (0.28)

2.21 (0.50)

1.27 (0.28)

0.98 (0.23)

1.24 (0.37) 0.019

Specific protease activity (lg tyr mg protein-1 min-1)

D4

0.45

0.44

0.42

0.40

0.43

0.43

0.39

0.38

0.07

D12

0.57

0.55

0.54

0.55

0.54

0.52

0.51

0.49

0.04

a-amino acids (mg g-1 FM)

D4

2.1

3.451

4.107

5.751

5.1

5.54

5.422

6.572

0.1

D12

2.18

3.54

4.97

6.32

6.43

6.64

6.03

7.36

0.25

1.44
1.975

1.22
2.03

1.388
1.98

1.6
2.13

1.611
2.1

1.611
2.43

1.444
2.38

0.1
0.11

Total phenols (mg g-1 FM)

D4
D12

1.60
2.09

Reducing sugars (mg g-1 FM)

D4

11.25

20.53

22.32

31.4

30.23

27.68

34.02

40.53

0.32

D12

9.87

18.09

15.04

27.54

28.09

24.43

29.76

37.89

0.42

Non reducing sugars (mg g-1 FM)

D4

6.26

7.15

13.39

15.63

6.49

12.17

13.48

17.65

0.54

D12

5.39

6.76

16.03

26.5

3.34

26.77

11.91

13.51

0.32

Total sugars (mg g-1 FM)

D4

17.71

27.68

35.71

47.5

36.72

39.85

47.5

57.65

0.41

D12

14.65

24.85

31.07

42.13

31.43

35.98

41.67

51.4

0.65

Each value is the mean of 10 independent replicates

The figures in the parenthesis represent proteins on mg g-1 dry mass basis (mg g-1 DM)
The expiations are the same that are under the Table 1

when the cut sprays were transferred to DW but an increase

in the soluble protein content was registered with the
progression in time from D4 to D12 when sprays were held
in SUC (Table 2). In some flowers, specific proteins
referred to so called death proteins have been observed
to be upregulated within the flowers to initiate senescence
[15, 27]. The increase in proteins in this experiment with
senescence could be due to the manifestation of the death
proteins which regulate the process of flower senescence,
but requires a thorough investigation. Specific protease
activity has been shown to increase as the time progressed
from day 4 to day 12 in the control and various CHI treated
(before, during and after 72 h wet storage) sprays irrespective of the vase solutions used. The sprays pulse
treated with CHI after 72 h wet storage showed lowest
specific protease activity than the control and other treatments (Table 2). Decreased specific protease activity in
these sprays can be attributed to their delayed flower
senescence and longer vase life. In various flowers such as
Dianthus, Iris, Hemerocallis, Petunia and Consolida

protease activity has been shown to increase towards

senescence [9, 14, 3840]. Application of proteolysis
inhibitors like phenyl methane sulfonyl fluoride (PMSF)
has been found to decrease the activity of specific proteases
and thereby increase the flower longevity of various
flowers like Sandersonia and Iris [38, 41]. The treatment of
CHI also resulted in an increase in the total a-amino acid
pool. Maximum a-amino acid content was recorded in the
samples from cut sprays treated with CHI after 72 h wet
storage (Table 2). During flower senescence of Dianthus,
ovary becomes the substantial sink and as such the amino
acid pool present in the senescing petal tissues is transported back to ovary [15]. During the present study maximum a-amino acids were reported in the petal tissue of
sprays treated with CHI after storage. This could be
attributed to the fact that in fresh flowers amino acids are
retained within the petals for biochemical processes.
However, in order to understand the complex mechanism
of senescence detailed biochemical investigation is
required to analyze individual amino acids at various stages

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of senescence. The total phenolic content recorded on D4

and D12 was almost comparable in all the treatments along
with the control (Table 2). Phenolic content has generally
been shown to increase in various flowers such as Hemerocallis and Dianthus with senescence [25, 36].
The present investigation revealed that higher concentration of various sugar fractions was maintained in the cut
sprays treated with CHI as compared to the control. Maximum sugar fractions were recorded in the samples treated
with CHI after 72 h wet storage and transferred to 0.05 M
SUC. Various sugar fractions exhibited a continuous
decrease as the time progressed from D4 to D12. The cut
sprays transferred to SUC showed higher content of sugar
fraction than those transferred to DW irrespective of various CHI treatments (Table 2). The beneficial effects of
sugars on flower senescence have been attributed to the
decline in ethylene biosynthesis or sensitivity to ethylene
[9, 34, 4244]. Various sugars have been reported to delay
the biosynthesis of mRNA that code for the senescence
associated genes [45]. The research progress made in
recent years has demonstrated the extensive and complex
molecular crosstalk between the sugar and hormone signaling pathways which have opened new vistas to study the
complex role of sugars (exogenous and endogenous pool)
at biochemical and molecular level [46].

Conclusion
The results suggest that the cut sprays of D. chinensis
picked at the appropriate stage (paint brush stage) can
prove to be a good model for market supply for export of
this cut flower. The sprays may be pulse treated with
0.1 mM CHI for 1 h after cool wet storage for 72 h at 5 C
and then transferred to vase solution without interfering
with their postharvest life. As such, it can be used as an
effective protocol for storage treatment of this beautiful cut
flower. Moreover, senescence in D. chinensis has been
shown to be governed by a turnover and interplay between
various biomolecules like proteins, carbohydrates, phenols
and amino acids. In addition, specific protease activity was
found to slow down under cool wet storage at near freezing
temperature.
Future Perspectives
Studying the role of CHI in flower senescence at molecular
level can open new vistas in this field. Maintaining lower
content of proteins by the application of CHI by monitoring the expression of so called death proteins can
open a new window for unraveling this fascinating
mechanism of flower senescence. Besides such biotechnological approaches can help in future to target death

123

proteins and lead to various cues about the complex

regulation of flower death. The specific death proteins
can be suppressed to delay senescence and increase the
vase life of this prized ornamental flower by improved
biotechnological methods.
Acknowledgments Prof. A. Q. John is acknowledged for the correct identification of the plant material used in present experimental
work. The authors acknowledge Prof. S. Farooq for his influence
through the opportunities provided by him and insights conveyed.
Syed Sabhi Ahmad thanks University Grants Commission (UGC),
Govt. of India for providing JRF under (UGC-BSR) JRF scheme.

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27:11001102
3. van Doorn WG (2004) Is petal senescence due to sugar starvation? Plant Physiol 134:3542
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