REVIEW OF LITERATURE

Flower is an actively metabolizing system and carries out all its metabolic
activities at the expense of stored food in the form of carbohydrates, proteins
and fats (Nowak and Rudnicki 1990, Singh et al 2001, Bhattacharjee and De
2003). Besides, high level of turgidity and sensitivity towards ethylene
contribute to potential vase life of the flowers (Bhattacharjee 1999, Singh et al
2002, Singh and Srivastava 2006). Cut flowers detiorate very quickly and
hence, to maintain freshness of flowers, they have to be handled with utmost
care. While petal senescence is clearly a degenerative process, the upregulation
of new genes and synthesis of new proteins appear to be necessary for this
process (Suttle and Kende 1980, Woodson et al 1989, Singh and Singh 2002).
Senescence is considered to be internally programmed, because it is specific
and orderly in terms of when, where and how it occurs (Nooden and Leopold
1978).

FLOWER SENESCENCE

Senescence of petals is marked by changes in membrane properties

especially, membrane phospholipids and hence, membrane permeability,
eventually leading to loss of cellular compartmentation and mixing of
metabolites thereby, causing death of cells (Beutelmann and Kende 1977,
Suttle and Kende 1980, Halevy and Mayak 1981, Borochov et al 1982,
Singh and Srivastava 2006). Involvement of free radicals during petal
senescence has also attracted considerable attention (Dhindsa et al 1981,
Leshem et al 1986, Singh and Jegadheesan 2003, Kumar et al 2007).
Kumar et al (2008) reported that in rose flowers, activities of catalase,
peroxidase and ascorbate peroxidase increased till petals completely
unfolded and declined thereafter concomitant with petal senescence.
Unlike fruits and vegetables, the flower is an intricate system

which is constituted of various parts viz. sepals, petals, stamen and

carpel, each of which coordinates the process of its senescence. The most
important barrier in the marketing and commercialization of many cut

flowers is their short vase life and their inability to withstand stresses

during storage or transit (Halevy and Mayak 1981, Nowak and Rudnicki

1990).

A degradation of macromolecules during petal senescence in the

carnation flowers was observed ( Bovy et al 1995 ). Starch the dominant

carbohyderate in alstromeria petals and glucose, the dominant

carbohyderate in tulip teplas during senescence ( collier 1997 ). Hussain

et al ( 2001 ) observed changes in soluble proteins and carbohuyderates in

cut gladiolus under different physiological stages ( 0, 3 and 6 days ). The

amount of soluble carbohyderate present in single floret of cut gladiolus

increased with floret opening and gradually decreased with senescence.

Soluble proteins were found to be higher in bud stage then at fully spend

stage. Pulsing with 20 % sucrose , significantly increased the amount of

soluble carbohyderates and soluble proteins. Spikes stored for 3 days

showed higher content of carbohyderate and protein than those stored for

6 days.

Petal senescene was also found to be associated with decrease in fluidity

of membrane of all flowers in roses (Borochov et al 1983) and other

flowers (Borochov and Woodson 1989).Quantitative estimations of

microviscisity indicated an appropriate two fold increase with age in petal

membrane of rose (Borochov et al 1976),carnation (Thompson et al

1982),and petunia (Borochov and Woodson 1989).

 Flower provides an excellent system for the study of

senescence. Different flower parts senesce at different rates. In the

commercial use of cut flowers, it is usually the life span of the petals, an

ornamental part of the flower which determines its effective life.

Therefore, the study of petals senescence should provide insight into the

methods to improve the post harvest longevity of cut flowers and insight

into the mechanisms involved in the control of plant senescence.

Many factors such as deficiency of carbon source, water stress

conditions caused by bacterial contamination of vase water or air

embolism and ethylene sensitivity contribute towards senescence of

foliage and bud on the cut stems. Flower bud acts as a strong sink thereby,

drawing sugars, nutrients and water from the leaves or the stem (Halevy

and Mayak 1981, Nowak and Rudnicki 1990).

Senescence of petals is accompanied by decline in the levels of

protein and nucleic acids but the tissue also retains the capacity for
continued synthesis of proteins and RNA until very late in senescence
( Matile and Winkenbach, 1971; Woodson and Handa, 1987; Woodson,
1991 ). Woodson (1987 ) and Woodson and Lawton ( 1988 ) have reported
that ethylene induced senescence was accompanied by changes in mRNA
and protein profiles indicating the involvement of ethylene-induced gene
expression both at the transcription and translational level.

The senescence of flower petals is associated with a series of

physiological and biochemical changes. These include an increase in
hydrolytic enzymes, degradation of macromolecules, increased respiratory
activity and loss of cellular compartmentalization.

Variation in vase life, weight loss and flower diameter among different
 cultivars studied may be due to difference in the senescing behavior by
producing higher amount of ACC,ethylene forming enzymes and ethylene
( Accati and Jona 1989;Wu et al 1991 ).

EFFECT OF STAGES OF HARVEST

The optimum stage of harvest vary with species and depends upon
the post harvest ability of the buds to open. The flowers of gladiolus
can be harvested at colour stage when the buds are yet to open. The
buds of gladiolus continue to expand in water and exibit acceptable vase
life. Flowers of gladiolus should be harvested at three stages - S 1 - when

1-2 buds are fully open, S 2 - when 5-6 florets show colour, S 3 – when
basal floret is half open.
The vase life of gladiolus (Gladiolus sp.) spikes treated with ethanol
(0.1, 1.0 or 10.0%), sodium chloride (0.005, 0.01 or 0.1%), sucrose (2, 4 or
8%), boric acid (0.001, 0.01 or 0.1%), citric acid (0.03, 0.02 or 0.01%), or
aluminium sulfate (0.02, 0.05 or 0.1%) was studied . The highest number of
flowers opening simultaneously (2.0) was recorded for citric acid at all
concentrations. Sucrose (4 and 8%) gave the highest number of flowers that
remain open at a time (6.00). The longest useful life (14 days) was obtained
with 4% sucrose and 0.1% sodium chloride. The longest rachis (65.16 cm) and
the highest number of flowers that opened perfectly (14.66%) were recorded for
4% sucrose. Sucrose at all concentrations was most effective in preventing the
occurrence of partially opened flowers..

Waithaka et al ( 2001 ) observed that opening of gladiolus florets was

accompanied by a substantial increase in fresh and dry weight and
carbohyderate content of the perianths. The principal soluble carbohyderate
found was fructose, with substantially lower concenterations of glucose and
sucrose. Stored carbohyderate, starch , disappeared during floret opening and
its contribution to total carbohyderate content of the open flower was minor.
During development of inflorescence, it appeared that there was a transfer of
carbohyderate from senescing lower floret to those developing acropetally.

Otsubo and lwaya ( 2000 ) observed severe wilting in florets of cut gladiolus 4 days
after flower opening. Treatment with 0.1 M trehalose prolonged vase life by 2 days
and it was seen that upper florets also opened properly in trehalose treated spikes.
Trehalose treated spikes maintained water content more effectively and parenchyma
adjacent to vascular bundles in the petals of trehalose treated spikes maintained
inability for 4 days thus suggesting that trehalose preserves cell viability in gladiolus
spikes, therby enhancing water uptake into petal tissues.

Barman and Rajni made an attempt to standardize the stage of harvest of spike and the spike
length to achieve maximum vase life of gladiolus (cv. Eighth Wonder).Field grown spikes of
Eighth Wonder of 60 cm length at various stages (half developed bud, fully swollen bud not
showing colour, first bud showing colour, first floret open and two florets open) were cut. Spikes
were also cut at 60, 70, 80, 90 and 100 cm when the lowest floret was half open. Spikes
harvested at various stages showed statistically significant effect in terms of water uptake, fresh
weight of spikes, floret size, days taken to first floret withering, percentage of open floret and
vase life. Water uptake increased as the stage of harvest advanced. Total water uptake was
highest in spikes harvested at the two florets open stage. Fresh weight of spike recorded higher
values as the maturity advanced. Spike weight was highest on the sixth day at the two florets
open stage. Spikes harvested at the first floret open stage showed the largest floret diameter.
Days to first floret withering were lowest in spikes harvested at the half swollen bud stage and
highest in spikes harvested at the fully swollen bud stage. The percentage of open floret and vase
life were highest in spikes harvested at the first floret open stage, followed by those harvested at
two florets open stage. Spike length had positive effect on total water uptake, fresh weight of
spike on the third day, floret size and percentage of open florets. Water uptake was higher on the
third day than the sixth and ninth days. Total water uptake was highest when spike length was
100 cm. Spike weight was highest on the sixth day when spike length cut was 100 cm. Spike
length of 100 cm also recorded the highest floret diameter, days to first floret withering and vase
life. The percentage of open floret was highest when spike length cut was 90 cm, followed by 70
cm..

STORAGE OF FLOWERS

The viability of the cut flower trade depends upon the regular flow of
flowers in the market. Storage of flowers is an important post harvest operation
for regulating supply in the market. Storage is required to hold the flowers
during periods of overproduction and low demand. A successful cut flower
species must be able to withstand post harvest storage (Singh et al 2001).
Different flowers and varieties are reported to differ in their vase life and also
with regard to their response towards refrigerated storage due to genetic,
physiological or anatomical characteristics (Nowak and Rudnicki 1990, Singh
et al 2001, Redman et al 2002, Bala et al 2008). Flowers are stored under
refrigerated conditions because low temperatures reduce utilization of
substrates during respiration, lower water loss, production of ethylene and also
check excessive and undesired opening of the buds (Nowak and Rudnicki 1990,
Singh et al 2001). Besides, low temperatures significantly reduce microbial
growth rate (van Doorn and De Witte 1991). Different storage techniques such
as Normal Refrigerated Storage, Controlled Atmosphere (CA) storage and Low
pressure Storage (LPS) can be employed for storage of flowers (Goszczynska
and Rudnicki 1988).
Normal refrigerated storage is the most commonly employed method for
storage of flowers. Flowers in normal refrigerated storage can be stored by
placing the stems in water or chemical solutions (wet storage) or by wrapping
them in water-retentive plastic sleeves (dry storage) which tend to create
suitable gaseous environment for storage (Nowak and Rudnicki 1990, Grover
2001). An appropriate storage temperature and harvest at an appropriate
maturity influences storage period and longevity of cut flowers (Cevallos and
Reid 2001, Zencirkiran and Menguc 2003). A temperature range of 0 O to 1 O C
has been suggested for flowers, however, flowers originating from tropical and
sub-tropical regions require higher temperatures for storage (Nowak and
Rudnicki 1990). Gladiolus can be best stored at 2-4 O C (Kofranek and Halevy
1976). Cevallos and Reid (2001) reported that there was no significant
difference between the vase life of flowers of Iris cv. Telstar, stored under dry
as well as wet condition at temperatures ranging from 0 to 10 O C. Optimum
stage of flower development for harvesting cut flowers, however, depends upon
the species, cultivar, season, and consumer preference (Nowak and Rudnicki
1990, Singh et al 2001, Redman et al 2002, Zencirkiran and Menguc 2003).
Zencirkiran and Menguc (2003) also reported that cut flowers of Alstroemeria
harvested at different harvest maturities responded differently to storage
duration. The stems harvested with fully open flower were found to exhibit
short vase life. The flowers showed decrease in the contents of water soluble
solids in petals and the decrease was prominent in fully open flowers.
Zencirkiran (2002) also reported decrease in respiration rate of both dry and
wet-stored flowers of Freesia refracta, the decline being more rapid in wet
stored flowers. The flowers were successfully stored for 14 and 21 days in wet
and dry storage, respectively. The advantage of using bud cut carnations in
prolonging the period of storage and the improvement of flower quality and
longevity has also been reported (Goszczynsa and Rudnicki 1982). The storage
supremacy of bud cut over full grown carnations were attributed to lower
respiration, lower sensitivity to ethylene, lower susceptibility of petals to
fungal diseases and reduced petal surface area which reduces water loss
(Goszczynsa and Rudnicki 1988).

Prolonged duration of storage is reported to decrease vase life of flowers

(Xu et al 1987, Joyce et al 2000, Waithaka et al 2001, Redman et al 2002). This
could be due to high levels of respiration during storage leading to depletion of
carbohydrates and other storage materials in plant tissues, increase in
weight/water loss of the stems during storage or decrease in water absorption
following storage (Nowak and Rudnicki 1990, Singh et al 2007). Bhattacharjee
(1999) suggested that senescence processes continue during cold storage of
flowers resulting in shorter vase life. Due to higher water uptake and slow bud
development, a 20% increase in fresh weight was observed during wet storage
of carnation flowers (Goszczynska et al 1982). The premature senescence of cut
rose flowers after 17 days of storage was associated with an advanced and
stimulated rise in ethylene production (Faragher and Mayak 1984). Carnation
flowers also exhibited very high respiration rates during wet storage
(Hardenburg et al 1969). It has also been reported that the upper florets of
gladiolus lose substantial ability to open after storage (Singh et al 2002, 2003).

Pulsing or conditioning of flowers before storage has been used to extend

the storage period and improve the quality and longevity of the stored flowers
(Goszczynska and Rudnicki, 1988). Sucrose has been reported to be an
important ingredient of pulsing solution. Treatment with sucrose is reported to
help in the continuation of normal metabolic activity after storage, and
retardation of processes associated with senescence (Bhattacharjee and De
2003, Singh and Srivastava 2006). Sucrose is also reported to delay
degradation of proteins, ribonucleic acids, maintain membrane integrity and
mitochondrial structure and function (Halevy and Mayak, 1979). Song et al
(1992) observed that the vase life and flower quality of cut gladiolus spikes in
cultivars Hunting Song and Spic and Span was extended by the application of
sucrose (5 per cent) + 8- HQC, 200 ppm after 4 mM silver thiosulphate pulsing
for 30 minutes. Pal et al (2003) observed that the combination of sucrose (4 per
cent) + 8-HQC, 200 ppm effectively extended vase life of cut spike of gladiolus
cv. Pink Friendship. Pruthi et al (2001) reported that the treatment with sucrose
(5 and 8 per cent), calcium chloride (4 mM) and cobalt sulphate (2 mM) also
increased opening of gladiolus florets. Singh and Singh (2009) reported that
gladiolus cultivars differ in their vase life. Polymeric film sleeves exerted
considerable influence on post storage life of gladiolus spikes (Grover et al
2008). Naidu and Reid (1989) found that substantial demand for carbohydrates
in the developing inflorescence of tuberose caused bud abortion and failure of
florets to open. A sugar containing vase preservative and/or pre-treatment with
sugar improved display life of the stems. Rudnicki et al (1989) observed large
differences between cultivars and stages of bud development in keeping quality
of flowers after storage. Carnation buds of cultivars ‘Red Ivette’ and ‘Sancho’
stored dry at tight bud stage exhibited opening after storage and maintained
good keeping quality after storage for three and one month, respectively. The
best flower quality was achieved when preservative solution (200 mg/l HQC +
50 g/l sucrose) was used as continuous treatment following storage. Mor (1989)
reported that vase life of rose cv. Gabriella stored dry at 1 O C for 3 weeks was
shortened as compared to that of fresh flowers. Sucrose and silver thiosulphate
(STS) had beneficial effect on vase life of stored rose flowers. Grover (2001)
reported that modified atmosphere storage of flowers depends upon the gaseous
permeability properties of the polymeric film packages in which the flowers
have been previously dry-stored. The films which tend to maintain elevated
levels of CO 2 and low levels of O 2 inside the packages are considered suitable
for modified atmosphere storage of flowers. Pulsing of gladiolus spikes with
sucrose (20 per cent) + aluminium sulphate (400 mg/l) + GA 3 (100 and 200
mg/l) prior to wet storage significantly improved post storage opening of florets
and vase life (Namita et al 2006). The treatment of spikes before modified
atmosphere storage also showed similar effects (Grover et al 2008). Studies
also showed that during storage, gladiolus florets maintained relatively high
levels of total soluble and reducing sugars. Therefore, loss of vase life in
gladiolus following refrigerated storage did not appear to be associated with
depletion of sugars in florets (Grover et al 2005). Hence, the role of
endogenous levels of carbohydrates with respect to storage of spikes of
gladiolus needs further investigations.

The pulse treatments of the spikes with silver thiosu;fate improved floret
opening but not the life of individual florets ( Farhoomand 1978, Mor et al
1981 ). Beura and singh ( 2002 ) studied the post harvest life of gladiolus .
spikes dipped for 30 minutes in 4 mM silver thiosulfate or 1000 ppm silver
nitrate followed by 20 hour in 10 or 20 percenr sucrose. Silver nitrate + 20
percent sucrose gave the highest percentage of flowers ( 95.55 % ) and also
increased floret life and vase life.greatest solution uptake was observed in 1000
ppm silver nitrate 20 % sucrose.

VASE LIFE

Variation in vase life among different cultivars attributed to the differences

in physiological, anatomical, biochemical and genetic make up.

Mayak et al ( 1974 ) observed decline in fresh weight in the last phase of

vase life which occurred earlier in short lived cultivars than in long lived ones.

A flower begins to age as soon as the stem is cut from the plant.
Meanwhile, it may travel far and wide and change hands numerous times
between grower, wholesaler and rewholesaler until it finally reaches the retail
store where we purchase it.
Vase life of cut spikes of gladiolus is not more than 6 days. Preservative
solutions combination with sucrose, sucrose + 8-HQS and sucrose + GA3
increased the floret opening and vase life by increasing the water uptake,
reducing the transpirational loss of water, maintaining better water balance and
lowered ratios of water loss to water uptake thereby maintaining higher fresh
weight of the gladiolus (Gladiolus hybridus) cv. Nova Lux spikes. These
preservatives showed beneficial effects in terms of better water relations,
increased floret opening and vase life. These preservative solutions exerted a
dual effect in delaying senescence of gladiolus by increasing water uptake and
reducing water loss, thereby improving water balance and preventing water
stress in the cut flower, leading to increased fresh weight and vase life. Sucrose
 + 8-HQS solution was found to be best among the different preservative
solutions which extended vase life up to 11 days at room temperature (Rekha,-
M-K; Shankaraiah,-V; Reddy,-K-C; Srihari,-D; Sarma,-P-S, 2001 ).

Cut spikes of Gladiolus cultivars were transferred to a vase solution

containing STS [silver thiosulfate] (1.0 mM), sucrose (4%), sulfosalicylic acid
(100 ppm) with or without sucrose, or STS + sucrose. The effects of
sulfosalicylic acid on membrane stability and vase life of flowers were studied.
All the vase solutions significantly enhanced the quality of the cut flowers.
Sulfosalicylic acid phosphorus + sucrose significantly improved membrane
stability in cut flowers, and prolonged the vase life of the cut flowers from 4.80
to 12.00 days (Narendra-Kumar; Singh,-R-K, 2006 ).

According to Sward et al ( 1986 ) the vase life, general appearance, fresh

mass, volume of medium uptake of gladiolus inflorescences improved with
sucrose treatment : 30 gm sucrose/ dm 3 concenteration was the most effective
treatment. Sucrose uptake from the vase solution replenished intracellular
respirable carbohyderates allowing a sustained high respiration rate and
prolonged vase life.

The treatment with sucrose improved the quality of cut gladiolus. The quality of
flowers stored at 0 degree C with sucrose treatment was as high as that of fresh
flowers ( jiang et al 1989 ) suggesting that a decrease in sugar content of the spikes is
the factor leading to poor ornamental quality of cut flowers and that an assessment of
sugar content could provide an indication of keeping quality. Murali and reddy
( 1993 ) studied the influence of sucrose and metal salts on the post harvest life of
gladiolus Cv. Friendship. The number of fully opened florets was increased and
percent unopenend florets was decreased. A vase life of 12 days was recorded with
sucrose and cobalt compared with 7 days for control.

The vase life of carnation flowers has been generally determined by observing
senescence profiles, i.e., in-rollingof petal margin and wilting of whole petals as well
as ethylene production. In the present study, we tried to determine the vase life of
spray type carnation flowers by observing the number of open flowers, i.e., the
percentage of open flowers to the total number of initialflower buds. The vase life
determined by this method was similar to that determined by the former method, and
was useful to
evaluate the action of preservatives on carnation flowers.

After storage at different temperatures for a simulated transportation period, the vase lives at 20
degrees C (68 degrees F) of carnations (Dianthus caryophyllus cv. Imperial White), daffodils
(Narcissus pseudonarcissus cv. King Alfred), iris (Iris hollandica cv. Telstar), killian daisies
(Chrysanthemum maximum [Leucanthemum maximum]), paperwhite narcissus (Narcissus
tazetta cv. Paperwhite), roses (Rosa hybrida cv. Ambiance), and tulips (Tulipa gesneriana)
decreased with increasing storage temperature. There were no significant differences between the
vase life of flowers stored dry and flowers stored in water when storage temperatures were from
0 to 10 degrees C (32 to 50 degrees F). The vase life after wet storage at temperatures of 12.5
degrees C (54.5 degrees F) and greater was significantly higher than vase life after dry storage at
those temperatures for all the flowers studied. Iris and carnation flowers survived storage at 15
and 20 degrees C (59 and 68 degrees F) only when stored in water.. (Cevallos,-J-C; Reid,-M-
S,2001).

Murali ( 1990 ) studied the mode of action of metal salts and sucrose extending the

vase life of cut gladioli. The cut gladioli flower spikes were placed in solutions

containing 4 percent sucrose and silver thiosulfate , Co 2 + , Ni 2 + ,Zn 2 + ,Ca 2 + or Al 3 + or

sucrose alone. All ions except Zn maintained petal membrane integerity. Co 2 + ,

STS and Ni 2 + treatments markedly reduced ethylene evolution. Among all the

treatments , Co 2 + + sucrose were most effective in prolonging vase life,

restricting lipid peroxidation and reducing ethylene evolution. Cut flowers

supplied with water progressively loose most of their proteins ( paulin 1985 ).

Reid and lukazewski ( 1989 ) observed that the flowers of day lilies treated with

cycloheximidine , an inhibitor of protein synthesis, lasted as long as 6 days in

the vase as compared with one .gladiolus florets fed with cycloheximidine

showed double vase life ( jones et al 1994 ) .

Yamane and ogata ( 1995 ) observed that by placing gladiolus spikes in

a vase containing cycloheximide, there was a decrease in perianth length ,

fresh weight, dry weight, total soluble sugar and protein contents. Florets

on treated spikes remain folded whereas those on control spikes unfolded.

CHI did not affect perianth acid invertase activity but it inhibited other
protein synthesis and lowered perianth TSS content.

Various concentrations of polyamines like spermine, spermidine and

putrescine were applied alone (50, 100 and 150 ppm) or in combination

with sucrose 2% in vase solution to study their effect on the vase life and

other associated parameters in gladiolus (cultivars Dhanwantari and Snow

Princess) . All the three polyamines at 100 ppm concentration in

combination with sucrose were effective in extending vase life of

gladiolus. Of these three, 100 ppm spermine along with sucrose in vase

solution was also able to keep fresh weight of gladiolus spikes higher for

longer duration in comparison to other treatments due to higher uptake of

holding solution. Aesthetic look of cut spikes in terms of floret diameter,

turgidity and freshness was also better in this treatment( Singh,-V-P;

Kiran,-D; Arora,-A, 2005 ).

Baweja studied the effect of sulfur deficiency on the vase life of

Gladiolus cut flowers, an experiment was conducted in two sets with sand

pot culture (i) complete nutrient supplied with sulfur, (ii) nutrient solution

supplied without sulfur. Spikes of cut flowers were treated with: water,

6% sucrose solution, 650 ppm Al2SO4 solution. The longest vase life was

recorded in the 650 ppm Al2SO4 solution (15.0 days), followed by the 6%

sucrose solution (12.5 days). The water recorded the shortest vase life of

cut Gladiolus flowers (10.5 days). However, in sulfur-deficient plants, the

vase life was 10.0, 11.5 and 13.0 days in water, 6% sucrose solution and

650 ppm Al2SO4 solution, respectively.

Cytokinins are involved in slowing down many of the process that

contribute to plant senescence ( Nooden and Leopold 1978, Kaminek et al

1992, mok and Mok 1994 ). Cytokinins enhanced water uptake in roses

( mayek and Halevy 1974 ). Pretreatment of detached carnation petals for

24 hrs with 0.1mM of cytokinins benzyladenine ( BA ), kinetin and zeatin

blocked the conversion of externally supplied ACC ( 1-

aminocyclopropane-1-carboxylic acid ) to ethylene and delayed petal

senescence by 8 days ( Halevy et al 1983 ). Treatment with BA improved

the longevity of opened carnation flowers and quality.

Cut flowers of gladiolus ( gladiolus grandiflorus ) treated with certain

beverages such as 7-UP and anti-ethylene such as silver nitrate, silver

thiosu;fate ( STS ), benzylaminopurine ( benzyladenine ) ( BAP ) in

different concenterations had long vase life and best visual quality were

also achieved ( premawardena et al 2000 ).

6- BA treatment promoted the flowering and increased the flower fresh

weight of china rose , the water absorption capacity of cut flowers was

higher than that of control

( Chen 1996 ).

In general, the endogenous levels of auxin decrease before or during

senescence, although in some cases, they do not change ( Nooden and

Leopold 1978 ). The auxin level decreased with age in two poinsettia
cultivars , but the decline was faster in the short lived one ( Gilbert and

Sink 1971 ). Auxin induced ethylene production has been reported in a

wide variety of experimental systems, however, the relationship between

the two on the senescence process is unclear ( abeles et al 1992 ).

A postharvest storage study with different treatments was carried out to

investigate a procedure in prolonging the vase life of the cut flower

gladiolus. Various chemical and nonchemical treatments which are locally

available were used. Compared with the control (distilled water),

beverages such as 7-UP in different concentrations, specific anti-ethylene

compounds, silver nitrate and silver thiosulfate (STS), benzylaminopurine

[benzyladenine] (BAP) in different concentrations, hot water treatment to

the base of the flower stalk as a sterilizing method, maintaining the pH at

3.5 using HCl, ethylene oxidation with KMnO4 and ethylene absorption

with activated charcoal were tested. The longest vase life and the best

visual quality were achieved by gladiolus flowers treated with 25% 7-UP.

Next best visual quality was achieved by silver nitrate, 2.25% 7-UP, 4 mg

KMnO4/litre and STS in the descending order. Flowers treated with 2 mg

KMnO4/litre and the control (distilled water) had the lowest vase life. The

visual quality was lowest in flowers treated with 2 mg KMnO4/litre. When

the costs of compounds used were considered, the study indicates that the

treatment 7-UP and KMnO4 were found to be the cheapest, while silver

nitrate and silver thiosulfate are comparatively expensive

 (Premawardena,-P-S; Peiris,-B-C-N; Peiris,-S-E, 2000 ).

Temperature also plays an important role in extending vase life. Gladiolus

prefers mild climate and sunny situation for their proper growth and flowering.
Temperatureis considered as a major factor influencing the number of days taken for
flowering. Optimum growth ofgladioli occurs at temperatures between 10 and 25OC
where the night temperature is not above 16OC.However, it can tolerate temperature
upto 40OC only if the relative humidity is high and soil moisturelevels are optimum.
Light levels affect initiation of flower. The period of flower initiation
commenceswhen the third leaf becomes visible and ends when the sixth and seventh
leaf appears. Quality offlower spikes and yield is better in a long day condition than
short days. Minimum illumination of 8hours/day is essential for most of the gladiolus
varieties

Cut flowers containing relatively high amount of carbohyderates, especially

mobile sugars, have longer vase life.

Humidity plays an important role as humid air may provide conditions for
bacterial and fungal diseases especially gray mould ( Botrytis cinerae ) which
can cause losses during storage and transport ( Nowak and Rudnicki 1990 ).
Therefore during growth period, moderate atmospheric humidity ( 50-60% )
beneficial for optimum vase life.

Dissease and pests of flowers adversely affect the keeping quality. Protecting
plants in the nursery against diseases and pest is necessary for producing cut
flowers of good quality and long vase life ( Nowak and Rudnicki 1990 ). To
produce flowers of acceptable quality it is necessary to maintain an optimal
fertilization program until harvest. Excessive nitrogen decreases the vase life
and increases susceptibility to infection. High salinity and high chloride levels
in growing media also tend to reduce vase life ( Nowak and Rudnicki 1990 ).

EFFECT OF FLORAL PRESERVATIVES

Any chemical formulation, which is used for extending vase life of flowers is
known as floral preservatives. Floral preservatives help to improve flower
opening, flower size, shape and colour.

Floral preservatives have two basic components, sugar and biocide. Sugar
provides food source and energy. Besides they maintain water balance of cut
stems ( Halevy and Mayak 1981 ). Sugar provides an additional food to cut
flowers. Generally sugar is the most commonly and frequently used sugar source
in floral preservatives. Biocides are the compounds, which control the growth of
microorganism in the vase water as well as on the stems.biocides check the
growth of bacteria, fungi and other microorganisms. They are helpful in
preventing blockage of stem tissues and maintaining water supply to the flowers
and leaves. Besides floral preservatives may alos have other ingredients such as
inorganic salts, growth regulators and antiethylene compounds ( halevy and
mayak 1981 ). Many commercial formulations are also available in the form of
tablets and powder form in advanced countries like U.K, U.S.A, holland for
extending vase life ( Nowak and rudnicki 1990; Singh et al 2001 ).

Kofranek and havely ( 1976 ) reported that sucrose improves the water uptake
by stem through decreasing water potential of tissue and limiting the
transpiration through the closure of stoamta.

The improvement in vase life in gladiolus by GA 3 and NAA might be due to its
prevention in increasing endogenous level of ABA thereby delaying senescence (
Borochov et al 1976 ).

Ktaller and Steponkus ( 1976 ) reported that sugar in preservative solutions

not only provide substrate for respiration but are also important in extending the
longevity of flowers by maintaining mitochondrial structure and functions. Chin
and Sacalis ( 1977 ) reportyed that addition of carbohyderates to holding
solutions resulted in exrension of vase life of flowers.

From the studies on preservative solutions in combination with sucrose,

sucrose + 8-HQS and sucrose + GA3 increased the floret opening and vase life
by increasing the water uptake, reducing the transpirational loss of water,
maintaining better water balance and lowered ratios of water loss to water
uptake thereby maintaining higher fresh weight of the gladiolus (Gladiolus
hybridus) cv. Nova Lux spikes. These preservatives showed beneficial effects in
terms of better water relations, increased floret opening and vase life. These
preservative solutions exerted a dual effect in delaying senescence of gladiolus
by increasing water uptake and reducing water loss, thereby improving water
balance and preventing water stress in the cut flower, leading to increased fresh
weight and vase life. Sucrose + 8-HQS solution was found to be best among the
different preservative solutions which extended vase life up to 11 days at room
temperature..( Rekha,-M-K; Shankaraiah,-V; Reddy,-K-C; Srihari,-D; Sarma,-P-
S, 2001 ).

Anjupal and Santosh kumar found that the vase life of cut flowers of gladiolus
cv. Pink Friendship was highest (11.06 days) in the solution of sucrose (4%) and
8-HQC (200 ppm). However, combinations such as 2% sucrose + 8-HQC(200
ppm), 4% sucrose + Al2(SO4)3 (300 ppm) and 4% sucrose + CoSO4(400 ppm)
were also better in increasing vase life and quality of flower over control
significantly. Whereas longevity of each floret was highest (4.57 days) in 4%
sucrose + Al2(SO4)3 (300 ppm). An experiment was conducted to determine the
suitable packaging material and preservative solutions at different storage
conditions of cut gladiolus spikes. The preservatives used for treatment were 3%
sucrose, 200 ppm 8-HQS, and 100 ppm gibberellic acid and the storage
conditions were room temperature (RT), cool chamber (CC), and cold storage
(CS). The packaging materials were corrugated fibreboard box and
polypropylene. The results show that an increase in vase life was significantly
more with spikes held in PM+sucrose+8-HQS for 22 days at CS. Tabulated data
on the effect of storage conditions on different physical parameters in gladiolus
flower spikes are presented..

An experiment was conducted to determine the suitable packaging material and

preservative solutions at different storage conditions of cut gladiolus spikes.
The preservatives used for treatment were 3% sucrose, 200 ppm 8-HQS, and 100
ppm gibberellic acid and the storage conditions were room temperature (RT),
cool chamber (CC), and cold storage (CS). The packaging materials were
corrugated fibreboard box and polypropylene. The results show that an increase
in vase life was significantly more with spikes held in PM+sucrose+8-HQS for
22 days at CS ( Rekha,-M-K; Shankaraiah,-V, 2000 ).

An experiment was conducted to compare the natural (90 or 100% coconut

water) and chemical floral preservatives (KMS [potassium metabisulfite] (100,
200 or 400 ppm), Al2(SO4)3 (100, 200 or 500 ppm), ascorbic acid (1 or 2%)) in
cut Gladiolus spikes to increase vase life. Cut spikes treated with tap or distilled
water served as the controls. Among the natural floral preservatives, 90%
coconut water as holding solution recorded the maximum solution uptake (41.87
ml) over the tap water-treated control (18.77 ml). Among the chemical
preservatives, 100 ppm Al2(SO4)3 absorbed the maximum solution (61.83 ml).
Among all preservative types, the maximum uptake of holding solution was
recorded with 100 ppm Al2(SO4)3, followed by 200 ppm Al2(SO4)3. Overall,
Al2(SO4)3 recorded the longest vase life, followed by ascorbic acid (2%) and
coconut water (100%) compared to the control treatments. Although 100 ppm
Al2(SO4)3 was the best in increasing the vase life among the holding solutions,
its cost was 21.26% higher than that of 100% coconut water. Ascorbic acid (2%)
recorded the highest percent increase in the cost, 49.51% higher compared to
90% coconut water (Nair,-S-A; Shiva,-K-N; Attri,-B-L; Sharma,-T-V-R-S,
2002 ).

WATER RELATIONS OF CUT FLOWERS

Water relations are the main enviormental determinants of flower petal

senescence rate in cut flowers ( Halevy and Mayak 1981, Singh et al 2001,
2002). The termination of vase life of many cut flowers is characterized by
wilting of petals even though the stems are placed in water. After cut stems are
placed in water, they exhibit marked changes in fresh weight which intially
increases due to expansion of buds and subsequently decreases due to
senescence of bud or foliage ( rogers 1973, Halevy and Mayak 1981, Singh et al
2001 ). In roses , loss of petal turgidity and fresh weight was caused by
decreased rate of water uptake ( Durkin and Kuc 1966, Burdett 1970 ). The
senescence of carnation flower was also accomplished by the loss of petal turgor
( Nichols 1966, Trippi and Paulin 1984, Whitehead et al 1984 ).

Exposure of cut flowers to drought, even for short periods, leads to an earlier
appearence of senescence symptoms ( borochov et al 1982, mayak et al 1985,
Paulin et al 1985 ). Water stress also caused carnation flowers to exhibit an
earlier loss of membrane integerity, decline in membrane phospholipid content
( Mayak et al 1985, paulin et al 1985 ) and decrease of fluidity ( coker et al
1985, mayak et al 1985 ).

Reduction in water uptake coupled with continous transpiration leads to water

deficit and reduced turgidity of cut flowers ( Nowak and Rudnicki 1990 ). The
decreased water uptake by the stem has been reported to be mainly due to
plugging of xylem vessels caused by bacteria, fungi and other microorganisms
which grow in vase water or on the dipped portion of the stem( Singh et al
2000,2001,2003 ). Microbial growth paralled the increase in stem resistance to
water flow and decrease in vase life ( Zagory and Reid 1989, Louband and van
Doorn 2004 ). Sterile water and germicides controlled microbial growth and
partially decreased the resistance to water flow ( Larsen and Cromarty 1967,
Marousky 1969, Burdett 1970, van Meeteren 1978a,b ).

Cut roses held in sterile water had decreased rates of water uptake ( Marousky
1969 ). It was suggested to be due to vascular blockage which wasthe result of
oxidative process induced from harvesting injury ( Halevy and Mayak 1981,
Nowak and Rudnicki 1990 ). Physiological stem plugging occuring due to a
wound or injury in cut stem alsocauses reduced water transport ( Bhattacharjee
1994 ).

Water uptake by cut chrysenthmum flowers was found to vary with the season
and the lignification of stem ( Marousky 1973 ). Stems harvested close to the
soil level were more lignified and absorbed less water than those cut higher on
the plant ( Singh et al 2002 ). Disruptions of water columns in the stem vessels
by air embolism has been considered to be one of the major factors causing
water deficit in chrysenthmum ( Singh and Moore 1992a ). The continous water
loss primarily through leaves during storage or transit is also reported to create
empty coloumnin the xylem thereby causing air to enter the stems ( van
Meeteren 1989 ).the air bubbles block the xylem vessels and make them non
functional ( van Meeteren 1989, Singh and Moore 1992a ). Thye physiologial
blockage of xylem vessels also leads to decrease in water uptake by the stem
( Singh and Moore 1994 ).

Recutting of lower 2-3 cm portion of stem exposes fresh tissue and is found
beneficial to improve vase life and water absorption ( nowak and Rudnicki 1990,
Singh et al 2001 ). Besides, water quality also plays an important role in vase
life. Water which contains impurities with high mineral contents and high pH
were found to reduce vase life of flowers. Total soluble solutes more than 200
ppm in water adversely affected vase life of chrysenthmum stems ( Nowak and
Rudnicki 1990 ).

PHYSIOLOGICAL AND BIOCHEMICAL ASPECTS OF PETAL SENESCENCE

The senescence of flower petals is associated with a series of physiological

and biochemical changes. Endogenous carbohyderate status of cut flowers is an
important factor which regulates their vase life ( Coorts 1973, Rao et al 1986,
Bhattacharjee 1994 ). Nowak ( 1979 ) correlated flower senescence of gerbera
jamesonii inflorescence with decrease of carbohyderate levels in the flower and
flower stalk. Ferreira and Swardt ( 1980 ) reported decrease in starch
concenteration from 77.19 to 31.45 mg g - 1 dry weight during the first stage of
senescence in the petals of cv. Sonia roses kept in deionized water. Petal
senescence generally accompanied loss of dry matter , apparently due to
hydrolysis of macromolecules like starch, protein and nucleic acids and the
redistribution of carbon and nitrogen compounds to other parts of the flower
( Borochov and Woodson 1989, bovy et al 1995 ). Gulzar et al ( 2005 ) also
reported decline in total soluble and reducing sugars i9n petals of petunia
hybrida during senescence. Decline in total soluble sugar content was also
reported with floret senescence in gladiolus (kaur et al 2006). Mayak and Halevy
(1980) reported that the hydrolysis of cell components viz. Macromolecular
components such as starch and cell wall polysaccharides was a major metabolic
event occurring in the senescing petals. Mayak and tirosh

(1988) reported that the starch content in the senescing flowers of carnation cv.
Improved white sim, decline gradually during senescence. Starch, the dominant
carbohydrate in alstromeria petals and glucose the dominant carbohyderate in
tulip tepals also showed decrease during senescence (Collier 1997). Hussain et
al (2001) observed changes in soluble proteins and carbohyderates in cut
gladiolus cvs. Rose Moureuto and Melody under different physiological stages .
the amount of soluble carbohyderates present in single floret of cut gladiolus
increased with floret opening and gradually decreased with senescence. Soluble
proteins were found to be higher in bud stage than in fully expanded stage.
Senescent tissues of carnation petals were also found to contain as much as 1 to
2 % reducing sugars ( nicholos 1973 ).

The applied sugar extends longevity by maintaining motochondrial

structure and functions ( Halevy and Mayak 1981, Kaur et al 2006 ). Halevy and
Mayak ( 1974 ) have reported taht sucrose decrease the water potential of petals
and enhanced their ability to absorb water. Cut flowers with high sugar contents
usually lasted longer ( Halfacre and Barden 1979 ). Rao et al ( 1980 ) presumed
that the hydrolysis of starch to osmotically active reducing sugars caused water
intake and corolla tissue expansion in gladiolus.

The process of flower senescence also involves an increase in the efficiency of

hydrolyzing enzymes such as RNAase, DNAase, protease , hydrolases of cell
wall polysaccharides as well as increase in ethylene production and respiration
rate ( Halevy and Mayak 1981, Borochov and Woodson 1989, Singh 1994, 2000,
Singh et al 2001, Singh and Singh 2002b ).

In a number of commercially omportant flowers especially, belonging to

families compostae, Iridaceae and Liliaceae, ethylene is not a primary
coordinating agent in inducing senescence ( Woltering and van Doorn 1988,
Lukazewski and Reid 1989, Singh and Moore 1994, Sultan and Farooq 2000).
Molecular basis of so called non-climateric ethylene insensitive flowers is not
fully understood. Efforts were hence, made to understand the process of flower
senescence in such flowers. It was reported that cycloheximide, an inhibitor of
protein synthesis inhibited meembrane detioration in day lily and narcissus
flowers (Bieleski and Reid 1991, Sultan and Farooq 2000). Lay-Yee et al (1992)
demonsterated that wilting of day lily flowers was also associated with rapid
lossof proteins.

Cycloheximide increased flower vase life and prevented disappearance of

proteins, suggesting that senescence required synthesis of new proteins.
Woodson and Handa (1987) have also suggested that onset of senescence in
hibiscus petals was associated with the increased protein synthesis as evidenced
by the increased incorporation of radioactive amino acids into proteins. The
senescence of cut gladiolus ,iris and narcissus flowers was also significantly
inhibted base cycloheximide,a plant protein inhibitor ( Jones et al 1994, Sultan
and Farooq 1977,2000, Singh and Singh 2002b). Carnation petal senescence was
also found to be associated with high level of mRNA until the final stage of
senescence ( Woodson 1987 ).

Flower petals , like other terminally differentiated sink tissues, often contain
active invertase ( Winkenbach and Matile 1970,Hawker et al 1976, Woodson and
Wang 1987). An increase in ratio of sucrose to reducing sugars occured
concominant with petal senescence in carnation (Nichols 1973,1976). This is
indicative of a loss of invertase activity with petal senescence. Thsi loss in
invertase activity has been linked to the de novo synthesis of a prteinaceous
invertase inhibitor (Winkenbach and Matile 1970, Halaba and Rudnicki 1983,
1986). A marked increase in RNAase and peroxidase activity occured during the
corolla senescence of Mirabilis jalapa (Yin and Shao 1994). Bovy et al (1995)
observed an increase in the hydrolytic enzymes during the petal senescence in
carnation flowers.
Stephenson and Rubinstein (1998) recorde a marked increase in the enzyme
activitiescapable of digesting native daylily (Hamerocallis) proteins and
suggested that the programmed cell death in daylily petals might involve
increased activity of atleast three classes of proteinases.

Involovement of free radicals during petal senescence has also attracted

considerable attention (Dhindsa et al 1981, Leshem et al 1986, Singh and
Jegadheesan 2003,Kumar et al 2007). Kumar et al (2008) reported that in rose
flowers, activities of catalase, proxidase and ascorbate peroxidase increased till
petals completely unfolded and declined thereafter concominant with petal
senescence.