The addition of glucose in holding solution enhances vase life and inflorescence quality of cut hydrangea flower over the application of sucrose or mannitol

Abstract

This study aimed to investigate the effect of different sugar types on the vase life of cut hydrangea flowers. There were 19 treatments based on concentrations and sucrose, glucose, and mannitol combinations. The results showed that the vase solution with 5% glucose provided the most extended vase life, which was 12.4 d, while the control solution with distilled water presented 8.86 d. The results were correlated with the total solution uptake, the number of days for reaching maximum inflorescene diameter, maximum sepal hardness score, chlorophyll content, and sepal electrolyte leakage. It could be implied that glucose alone extends hydrangea vase life by inactivating the ethylene signaling pathway. Based on the sepal size and colour, 3% glucose treatment which generated the second-highest vase life, could be the appropriate concentration for improving flower quality and longevity. This study provides essential information to understand hydrangea flower senescence and develop better vase solutions for cut hydrangea flowers.

Full text

Journal of Applied Horticulture (www.horticultureresearch.net)
Journal of Applied Horticulture, 25(1): 25-31, 2023
DOI: https://doi.org/10.37855/jah.2023.v25i01.04
The addition of glucose in holding solution enhances vase life
and inorescence quality of cut hydrangea ower over the
application of sucrose or mannitol
Piyatida Amnuaykan
Faculty of Agricultural Production, Maejo University, Chiang Mai, Thailand. E-mail: piyatida.chou@gmail.com
Abstract
This study aimed to investigate the eect of dierent sugar types on the vase life of cut hydrangea owers. There were 19 treatments
based on concentrations and combinations of sucrose, glucose, and mannitol. The results showed that the vase solution with 5%
glucose provided the most extended vase life, which was 12.4 d, while the control solution with distilled water recorded 8.86 d. The
results were correlated with the total solution uptake, the number of days for reaching maximum inorescent diameter, maximum sepal
hardness score, chlorophyll content, and sepal electrolyte leakage. It could be implied that glucose alone extends hydrangea vase life
by inactivating the ethylene signalling pathway. Based on the sepal size and colour, 3% glucose treatment, which generated the second-
highest vase life, could be the appropriate concentration for improving ower quality and longevity. This study provides the essential
information that will lead to understanding hydrangea ower senescence and developing better vase solutions for cut hydrangea owers.
Key words: Hydrangea macrophylla, glucose, sucrose, mannitol, vase life
Introduction
Hydrangea macrophylla is an important cut ower worldwide,
and increasing its vase life for the market requirement is
challenging (Olsen et al., 2015). There were several studies to
enhance cut hydrangea vase life. It has been found that the holding
solution with sucrose, 8-hydroxyquinoline (8-HQ), and citric acid
can prolong its vase life for ve days (Pei et al., 2013). Later,
surfactant and biocide in holding solution were introduced but did
not prolong the vase life of antique hydrangea (Aros et al., 2016).
In recent years, the eective application of 8-hydroxyquinoline
sulfate (8-HQS) in a vase solution has been observed (Kazaz et
al., 2019), followed by the nding of its suitable concentration
used (Kazaz et al., 2020). The current studies have been mainly
focusing on extending hydrangea vase life by reducing vascular
occlusion by utilizing the combination of sucrose and thymol
(Kazaz et al., 2020; Kiliç et al., 2020) and the application of
chrysal professional Ⅲ (CPIII) together with sucrose (Yang et
al., 2021). According to these studies, there is no variation of
sugar types used in the vase solution until the investigation of
1% glucose together with 8-HQS that can eectively prolong cut
hydrangea vase life (Suntipabvivattana et al., 2020). Interestingly,
sugar type might be the critical factor involving hydrangea vase
life extension.
Sugar supply is necessary for extended life of ower (Chuang and
Chang, 2013). There have been applications of sucrose to increase
the longevity of many owers such as Dendrobium (Chandran
et al., 2006), Paeonia lactiora (Xue et al., 2018), Gerbera sp.
(Muraleedharan et al., 2019), and Rosa sp. (Kshirsagar et al.,
2021). Additionally, sucrose and glucose can improve the vase
life extension activity. The study in cut Lilium suggested that
sucrose and glucose levels are upregulated during ower opening
(Arrom and Munné-Bosch, 2012). In cut Dendrobium, sucrose
and glucose suppress the abscission of the ower which results
in vase life extension (Pattaravayo et al., 2013). Moreover, the
investigation into Antirrhinum majus has suggested that sucrose,
glucose, and preservatives can signicantly increase vase life
(Ichimura et al., 2022). Glucose alone can enhance water solution
uptake to retain water content in petals (Hirose et al., 2019). The
sugar can improve the vase life of Chamelaucium sp. (Dung et
al., 2017), Rosa sp. (Sudaria et al., 2017), Protea sp. (Vardien et
al., 2017), Dahlia variabilis (Azuma et al., 2018), Hypericum sp.
(Oguta, 2019), and Dianthus caryphyllus (Kaviani and Sharafshah
Rostami, 2021). However, it is not only metabolizable but also
non-metabolizable sugar that is relevant in vase life extension.
Mannitol has been recognized as an important non-metabolizable
sugar that increases the vase life of some cut owers such as
Delphinium (Norikoshi et al., 2015), and Antirrhinum (Ichimura
et al., 2016).
Previous studies about the eect of dierent sugar types on the
vase life of several other cut owers have proposed that sugar
type can also be essential to improving cut hydrangea vase life.
Likewise, many types of sugar content are discovered in the
hydrangea ower (Kao, 1963). This investigation aims to oer
a novel and potentially more ecient and cost-eective vase
solution for enhancing the longevity of cut hydrangea owers
by understanding the impact of various sugar types, including
sucrose, glucose, mannitol, and their combinations at dierent
concentrations, on the vase life of cut hydrangea flowers.
Additionally, how do these sugar solutions induce physiological
changes that contribute to extending the hydrangea vase life.
Materials and methods
Plant materials: H. macrophylla ‘031’ owers were harvested
in the morning from a commercial grower (Khun Pae Royal
Project Foundation, Chiangmai, Thailand) at the commercial
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ISSN: 0972-1045
Journal of Applied Horticulture, 25(1): , 2023

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stage of ower development (approximately 50% of the fully
open decorative orets). The quality, including inorescence
size (approximately 15 cm in diameter) and stem diameter
(approximately 1.2 cm), were selected consistently. Flowers were
placed in tap water and transported to the postharvest laboratory
within half an hour.
Treatments with dierent sugar types: The stems were re-cut
underwater to 30 cm long to prevent an air embolism. Only two
upper leaves were retained on each stem. After that, the owers
were individually placed in a 1000-mL glass bottle containing
700 mL of different freshly prepared sugar solutions. There
were three types of sugars used such as glucose (KemAusTM,
CAS:50-99-7), sucrose (KemAusTM, CAS: 57-50-1), and
mannitol (HiMedia, CAS: 69-65-8), and there were 19 treatments,
including control based on dierent sugar combinations (Table 1).
The seven replicates per treatment were placed randomly under
25 ±1 °C with a 12-h photoperiod.
Table 1. Treatments with dierent sugar types and combinations used
in the experiment.
Treatments Combinations
Glu 1 1% Glucose
Glu 3 3% Glucose
Glu 5 5% Glucose
Suc 1 1% Sucrose
Suc 3 3% Sucrose
Suc 5 5% Sucrose
Man 1 1% mannitol
Man 3 3% mannitol
Man 5 5% mannitol
Glu + Suc 1 0.5% Glucose + 0.5% Sucrose
Glu + Suc 3 1.5% Glucose + 1.5% Sucrose
Glu + Suc 5 2.5% Glucose + 2.5% Sucrose
Glu + Man 1 0.5% Glucose + 0.5% Mannitol
Glu + Man 3 1.5% Glucose + 1.5% Mannitol
Glu + Man 5 2.5% Glucose + 2.5% Mannitol
Suc + Man 1 0.5% Sucrose + 0.5% Mannitol
Suc + Man 3 1.5% Sucrose + 1.5% Mannitol
Suc + Man 5 2.5% Sucrose + 2.5% Mannitol
Control Distilled water
Determination of vase life, relative fresh weight, daily solution
uptake, and total solution uptake: The vase life of each
ower was investigated by observing the number of days from
the rst day placed in sugar treatment solutions to the day the
ower exhibited sepal browning and wilting. To obtain relative
fresh weight (RFW), each glass bottle with and without ower
stem was weighed at a three-day interval. At the same time, the
measurements were carried out daily during vase life for daily
solution uptake (DSU) and total solution uptake (TSU). The
formula for RFW (%) and DSU (g stem-1 day-1) were (Wt/W0) ×
100 and St-1-St, respectively. The used formulas consisted of Wt,
the ower weight (g) at day t; W0, the ower weight (g) at day
0; St, sugar solution weight (g) at day t; and St-1, sugar solution
weight (g) on the previous day. TSU (g stem-1) and the sum of the
entire DSU of each treatment were estimated (Kazaz et al., 2020).
Determination of maximum flower diameter, days until
maximum ower diameter, and maximum sepal hardness:
Each ower diameter (DIA) was measured using a 600 mm
vernier calliper (RS Components Co., Ltd., Thailand) at three-
day interval during vase life. Then, the maximum diameter of
the replicates was used to calculate the average number of the
maximum diameter (DIAmax) for each treatment. In addition, the
number of days each ower used to reach its DIAmax (DtDIAmax)
was also recorded. For maximum sepal hardness (SHmax), the
observation was completed at three-day intervals during vase
life. A ve-score rating criterion was created to evaluate the sepal
hardness (SH) of hydrangea inorescence from the softest (score
1) to the hardest (score 5).
Determination of total leaf chlorophyll content: The leaf
chlorophyll content of hydrangea inorescence was determined
using SPAD-502 Plus (Konica Minolta Optics, Japan). Three
measurements were taken per leaf to obtain an average SPAD
value of the leaf for each ower.
Determination of sepal electrolyte leakage: Ten sepal disks (0.5
cm in diameter) from each ower were cleaned in distilled water
three times. Then, the disks were placed in a test tube containing
10 mL of double-distilled water. After that, the tube was incubated
in a 40 °C water bath WNE 45 (Memmert, Germany) for 30 min.
The electrical conductivity (EC1) of the solution in the tube was
measured. After obtaining EC1, the tube was autoclaved at 121 °C
for 15 min, followed by incubation under 25 °C for 24 h. Then,
the EC2 was measured. The sepal electrolyte leakage (%) was
calculated using the formula: (EC1/EC2) × 100.
Statistical analysis: This experiment was carried out in a
completely randomized design (CRD) with 19 treatments,
including control and seven replicates per treatment. Statistical
signicance between mean values was evaluated using one-way
ANOVA and the Tukey test (Assaad et al., 2014).
Results
Vase life, total solution uptake, maximum ower diameter,
days until reaching maximum ower diameter, and maximum
sepal hardness of hydrangea cut ower: Vase life, TSU, DIAmax,
DtDIAmax, and SHmax of hydrangea diered depending on the vase
solution’s sugar combination (Table 2). Glu 5 exhibited the most
extended vase life, approximately 12.4 days and 3.54 days longer
than the control. However, colour change in the inorescence of
this treatment was clearly evident. The colour was lighter than
all treatments in this study (data not shown). Glu 3 provided the
second most extended vase life. It lasted approximately ten days,
which was 1.14 days longer than the control. However, it was
not signicantly dierent with the Glu+Suc 3, Glu+Suc 5, and
Glu+Man 1 treated samples, which were also more prolonged
than the control. At the same time, Man 5 provided the shortest
vase life, which was approximately 2.29 days. Noticeably, the
accumulation of white powder was observed on the sepals of all
owers treated with mannitol. Glu 5, with the highest vase life,
also obtained the highest DtDIAmax and SHmax. However, the
highest DIAmax was not found in such treatment. DIAmax of Glu
5 was shorter than Glu 1. While Glu 5 generated longer vase life
than Glu 3, the other factors, such as DIAmax, DtDIAmax, TSU,
and SHmax values, were not signicantly dierent. Although the
vase life was not signicantly dierent among Glu 3, Glu+Suc
3, Glu+Suc 5, and Glu+Man 1 treated samples, the DIAmax,
DtDIAmax, TSU, and SHmax values were dierent. Among these
four treatments, Glu 3 presented the longest DIAmax and the
highest SHmax. Also, Glu 3 and Glu+Suc 3 provided the highest
DtDIAmax and TSU. Comparing all treatments together, the high
TSU was found in the Glu 1, Glu 3, Glu 5, Glu+Suc 1, Glu+Suc
26 Glucose is the most eective sugar for extending hydrangea vase life

Journal of Applied Horticulture (www.horticultureresearch.net)
3, and control. In contrast, the lowest one was detected in Man 5.
Interestingly, the treatments with sucrose, the most popular sugar
used for hydrangea vase life extension, did not show outstanding
results in prolonging the vase life. The results indicate that glucose
inclusion without mannitol could prolong the vase life of cut
hydrangea owers, which can be linked to TSU, DtDIAmax, and
SHmax.
Eect of sugar types on daily solution uptake of hydrangea cut
ower: From the results above, the treatments with a combination
of sugars did not strongly support the vase life extension, and
one sugar-type treatment provided better results. Therefore,
only Glu 1 (provided the longest DIAmax and the highest TSU),
Glu 3 (provided the longest DIAmax), Glu 5 (provided the most
extended vase life, the highest TSU, and the highest SHmax),
Man 5 (provided the shortest vase life, the shortest DIAmax, the
lowest TSU, and the lowest SHmax), and control were selected
to present in the graphs for the clearer vision (Fig. 1). Since
the vase life of Man 5 was the lowest, DSU could be recorded
only on the second day. DSU of all treatments apart from Man
5 decreased dramatically after the second day of the experiment
and slightly uctuated to the end of their vase life. On the second
day, DSU in Glu 1 seemed the highest. However, there was no
signicant dierence between Glu 1, Glu 3, Glu 5, and the control.
A signicant dierence was only observed between Glu 1 and
Man 5. On the third to fth days, all treatments had no statistical
dierence. Glu 1, 3, and 5 exhibited small peaks of DSU around
the sixth day, while there was no peak from the control. The
dierence was observed, and the control provided the lowest
DSU compared to the others. After the sixth day, DSU from Glu
1, 3, and 5 were reduced, which made the dierence disappear.
Although the patterns of DSU were dierent, the highest TSU was
found commonly in Glu 1, 3, 5, and control. The peaks and the
higher rates of DSU in Glu 1, 3, and 5 exhibited the improvement
of hydrangea solution uptake by glucose.
Eect of sugar types on relative fresh weight of hydrangea
cut ower: There was no signicant dierence in RFW in all
treatments on the rst day. From the second day, there were
two dierent types of RFW patterns for all treatments (Fig. 2).
The rst pattern was found in Glu 1, Glu 3, Glu 5, and control.
RFW suddenly increased before it dropped later, and the RFW
of all glucose treatments were considerably higher than that of
the control. For the peaks of RFW on day two, Glu 3 and Glu 5
provided the highest value, followed by Glu 1 and the control,
respectively, while there was no signicant dierence between
Glu 1 and the control. After the second day, all RFW were
dropped. The control provided the lowest, and Glu 3 exhibited the
highest RFW until the sixth day, when the signicant dierence
among all treatments disappeared. Although the trends of Glu 1,
3, and 5 were similar, Glu 3 showed the best RFW. The second
pattern was observed in Man 5. This pattern showed a sharp
reduction in RFW until the end of their vase life. Interestingly,
Table 2 The effects of different sugar types and their combinations on vase life and other physiological characteristics in cut hydrangea flower
TreatmentVase life (d) DIAmax (cm) DtDIAmax (d) TSU (g) SHmax (score)
Glu 1 7.43 ± 0.297cd 23.5 ± 0.408a3 ± 0.309ab 277 ± 8.62a4.00 ± 0.309ac
Glu 3 10.00 ± 0.436b22.3 ± 0.596ab 3.29 ± 0.286ab 255 ± 15.1a4.43 ± 0.297ab
Glu 5 12.4 ± 0.369a21.6 ± 0.484b4.14 ± 0.8a272 ± 12.7a4.86 ± 0.143a
Suc 1 4.71 ± 0.286ef 19.7 ± 0.214cd 2.29 ± 0.184b 108 ± 5.42df 3.00 ± 0.218ce
Suc 3 4.00 ± 0.309fh 18.3 ± 0.214de 2.43 ± 0.202b90.9 ± 2.89df 2.71 ± 0.184def
Suc 5 3.57 ± 0.202fh 18.1 ± 0.297e2.29 ± 0.184b129 ± 9.36d2.29 ± 0.184eg
Man 1 6.43 ± 0.429de 19.4 ± 0.385ce 2.14 ± 0.143b170 ± 2.57c2.14 ± 0.261eg
Man 3 4.86 ± 0.261ef 19.8 ± 0.286cd 2.14 ± 0.143b85.6 ± 3.27ef 1.57 ± 0.202fg
Man 5 2.29 ± 0.184h15.9 ± 0.0922f2.00 ± 0b41.7 ± 3.89g1.29 ± 0.184g
Glu+Suc 1 9.00 ± 0.535bc 22.1 ± 0.254ab 3.29 ± 0.522ab 275 ± 12.4a3.29 ± 0.184bce
Glu+Suc 3 9.71 ± 0.522b19.9 ± 0.18c3.14 ± 0.829ab 263 ± 10.2a3.86 ± 0.261acd
Glu+Suc 5 9.29 ± 0.36b19.9 ± 0.18c2.29 ± 0.184b193 ± 5.3bc 4.14 ± 0.261ac
Glu+Man 1 9.86 ± 0.261b19.4 ± 0.335ce 2.14 ± 0.143b210 ± 5.2b3.14 ± 0.34ce
Glu+Man 3 4.71 ± 0.286ef 17.9 ± 0.17e2.14 ± 0.143b116 ± 6.68de 3.14 ± 0.261ce
Glu+Man 5 4.29 ± 0.184fg 21.6 ± 0.18b2.14 ± 0.143b73.9 ± 2.45fg 1.57 ± 0.202fg
Suc+Man 1 4.14 ± 0.34fg 19.9 ± 0.261c2.14 ± 0.143b103 ± 4.19df 1.43 ± 0.202g
Suc+Man 3 4.57 ± 0.297fg 16.4 ± 0.143f2.00 ± 0b89.1 ± 1.87ef 1.71 ± 0.184fg
Suc+Man 5 2.86 ± 0.34gh 18 ± 0.218e2.00 ± 0b76.1 ± 5.37fg 1.14 ± 0.143g
Control 8.86 ± 0.459bc 18.5 ± 0.189ce 2.00 ± 0b253 ± 6.62a3.71 ± 0.184acd
Values are means ± SEM, n = 7 per treatment group. As analysed by one-way ANOVA and the TUKEY test, means in a column without a common
superscript letter differ (P<0.05). Abbreviations: DIAmax, maximum flower diameter; DtDIAmax, days until reaching maximum flower diameter; TSU,
total solution uptake; and SHmax, maximum sepal hardness
Fig. 1. Effects of different sugar treatments on daily solution uptake of
a cut hydrangea flower. Daily solution uptake was expressed as mean ±
SE (n = 7 flowering stems from each experiment).
Glucose is the most eective sugar for extending hydrangea vase life 27

Journal of Applied Horticulture (www.horticultureresearch.net)
the treatment with the highest vase life provided the rst pattern
of RFW, and the treatment with the lowest vase life showed the
second. The results suggested that glucose aected cut hydrangea
fresh weight alteration, linked to its vase life extension.
Total leaf chlorophyll content: The leaf chlorophyll content
is represented in the SPAD value. The SPAD values were
not significantly different among all treatments on the first
day. After that, the value reduction occurred in all treatments
presented in the graph, and Glu 5 provided the highest SPAD
value throughout the vase life (Fig. 3). There were two graph
patterns. The rst pattern contained a gradual drop line (Glu 1,
Glu 3, Glu 5, and control), whereas the second pattern provided
a sudden fall (Man 5). For the rst pattern on the second day, the
SPAD values of control were not signicantly dierent to Glu 3
and 5. However after that, the value was gradually reduced and
showed a statistical dierence compared to Glu 5. Interestingly,
the total leaf chlorophyll content in every treatment with the rst
pattern apart from Glu 1 was higher than that of the control. This
pattern also included the treatment with the most extended vase
life. Hence, it can be implied that glucose might be necessary in
delaying hydrangea leaf senescence by maintaining hydrangea
leaf chlorophyll content.
Sepal electrolyte leakage: Overall, sepal electrolyte leakages
(SEL) were slightly increased throughout the experiment (Fig. 4).
On the rst day, the SEL of Man 5 was the highest, followed by
control. The values of all glucose treatments were not signicantly
dierent and were the lowest until day four. After that, there was
a uctuation in SEL value comparisons. The values of all glucose
treatments were signicantly dierent on the sixth and tenth days,
not day eight. However, all glucose treatments exhibited lower
SEL than the control throughout the vase life. The lowest SEL
was found in Glu 5, with the most extended vase life, whilst the
highest SEL was detected in Man 5, which had the shortest vase
life. Therefore, SEL, aected by sugar types in vase solution, was
one of the factors inuencing cut hydrangea vase life.
Discussion
Sugar, including monosaccharides and disaccharides, involves
stress recovery and can protect the plant from drought and
senescence (Martínez-Noël and Tognetti, 2018). Therefore, it has
been used as an ingredient in vase solutions to prolong the vast
life of several cut owers, including hydrangea. Although many
studies have evaluated the appropriate vase solution for hydrangea
ower, there are not many variations of sugar type used. In
this study, three types of sugar and their combinations were
used: metabolizable monosaccharide (glucose), metabolizable
Fig. 4. Effects of different sugar treatments on sepal electrolyte leakages
of cut hydrangea flower. Sepal electrolyte leakages were expressed as
mean ± SE (n = 7 flowering stems from each experiment).
Fig. 2. Effects of different sugar treatments on relative fresh weight of
cut hydrangea flower. Relative fresh weight was expressed as mean ±
SE (n = 7 flowering stems from each experiment).
Fig. 3. Effects of different sugar treatments on leaf chlorophylls content
(SPAD) of cut hydrangea flower. Leaf chlorophylls content was expressed
as mean ± SE (n = 7 flowering stems from each experiment).
28 Glucose is the most eective sugar for extending hydrangea vase life

Journal of Applied Horticulture (www.horticultureresearch.net)
disaccharide (sucrose), and non-metabolizable sugar alcohol
(mannitol). The positive changes in vase life and other
physiological characteristics occurred when the appropriate
amounts of glucose were added to vase solutions. Glucose alone
was more eective in extending the vase life of cut hydrangea
ower than sucrose, mannitol, and the combinations of glucose
with sucrose and mannitol. Besides extending vase life, glucose
could also improve sepal strength and prolong the days to full
opening ower—the results related to the total amount of vase
solution uptake.
Although sucrose can delay senescence in many owers (Arrom
and Munné-Bosch, 2012), its ability on vase life extension
can still be found to be inferior to glucose in several owers,
including Antirrhinum (Ichimura and Hisamatsu, 2006) and
Paeonia suruticosa (Wang et al., 2014). It can be implied that
dierent types of sugar work dierently depending on ower
species. These positive eects of glucose on hydrangea vase life
extension and ower quality will likely involve sugar degradation.
Glucose is a monosaccharide derived from sucrose cleavage
(Stein and Granot, 2019). Hence, its molecule is smaller, easier
to metabolize, and quicker to act than sucrose. Besides that,
glucose incorporates antioxidant activity in plants (Gangola and
Ramadoss, 2018; Li et al., 2020; Osakabe et al., 2014). Since the
shortening of ower vase life is due to the overproduction of ROS,
which leads to oxidative stress (Jędrzejuk et al., 2018; Zhang and
Becker, 2015), and antioxidant activity can protect the plants
from ROS damage followed by increasing postharvest longevity
(Mohammadi et al., 2020; Pourzarnegar et al., 2020; Safari
Motlagh et al., 2021; ul Haq et al., 2021), glucose; undoubtedly,
provides good quality of hydrangea ower during the vase life.
Although a high amount of glucose (5%) suggested the
longest vase life, this treatment did not exhibit the maximum
inorescence diameter. Interestingly, the biggest, inorescence
was found in treatment with a slightly smaller amount of glucose
(1%), with shorter vase life. In fact, glucose aects ower opening
by controlling petal cell enlargement (Arrom and Munné-Bosch,
2012). As reported in the present study, 1% glucose might be the
optimum concentration for the ower opening of hydrangea. The
vase life of owers with 1% glucose that provided the bigger size
of inorescence was reduced. This is likely due to the higher-rated
sepal transpiration occurring when more orets are opened. A
similar result was reported that the decorative orets removal
led to the vase life extension of cut hydrangea ower (Kitamura
and Ueno, 2015). However, the sepal transpiration rate might not
have a major eect on vase life, considering that the vase life in
1% glucose treatment was not the shortest one. Types of sugar
and its combinations should take into account.
Mannitol acts as an osmoprotectant (Sickler et al., 2007),
antioxidant (Parvaiz and Satyawati, 2008), and antibiotic (da
Silva Brandão et al., 2019) that inuences ower longevity.
However, the application of mannitol in hydrangea signicantly
accelerated ower senescence and reduced vase life. The visible
mannitol white powder was also accumulated on the sepal surface.
These ndings might be caused by the toxicity of mannitol (da
Silva Brandão et al., 2019), and hydrangea is more susceptible
to that. Moreover, the major carbohydrate content in petals can
be considered. Mannitol extends Antirrhinum vase life because
it is the major soluble carbohydrate the petal (Ichimura and
Hisamatsu, 2006). In contrast, glucose is reported as the most
abundant carbohydrate found in hydrangea sepal. The ndings
were related to the positive changes in hydrangea vase life due to
the application of glucose. They emphasized the dierent eects
of sugar types on the dierent types of owers.
According to the results, glucose provided a different daily
solution uptake pattern than other sugar types. Without mannitol,
glucose could eectively increase the ower’s daily and total
solution uptake. Moreover, adding glucose together with other
sugars, even with the ineective mannitol, could increase ower
daily solution uptake. The higher solution uptake could be linked
to the reduction of xylem occlusion.
Glucose positively affected the relative fresh weight, while
mannitol posed a negative one. Therefore, glucose decelerated
the reduction of relative fresh weight. The outcomes correlate
with vase life, undoubtedly.
The leaf’s lower chlorophyll content is one of the processes
involved with plant senescence (López-Fernández et al., 2015).
Interestingly, this factor is also an eective indicator of ower
senescence (Liao et al., 2012; Lim et al., 2007). SPAD value
is used in this study to indicate the quality of hydrangea leaf
chlorophyll content. As expected, SPAD values of all treatments
decreased from the beginning to the end of their vase life due to
the ageing process. The reduction patterns of SPAD value were
slower by glucose and quicker by mannitol.
Moreover, treatment with the highest glucose concentration
containing the longest vase life exhibited the highest leaf
chlorophyll concentration at the end. It can be suggested that
leaf chlorophyll content influences the vase life quality of
hydrangea. It is well-documented that ethylene plays a role in
plant chlorophyll breakdown (Jibran et al., 2013). Also, many
investigations conrm that glucose activates the degradation of
the ethylene signalling regulator and plays a crucial role in the
senescence of ethylene-sensitive owers (Kakhki et al., 2009).
Because hydrangea is an ethylene-sensitive ower (Lauridsen
et al., 2015), glucose is possibly involved with ethylene activity
in hydrangea.
Degradation of the petal membrane, which leads to electrolyte
leakage, is one of the common metabolic changes during ower
senescence (Arora, 2008; Shahri and Tahir, 2011). Accordingly,
glucose reduced sepal electrolyte leakage in hydrangea. The
lower electrolyte leakage correlated with sepal hardness and
vase life extension. This study accentuates the ability of glucose
to prevent plant cells from protein degradation, phospholipids
breakdown and the competence of ethylene. Moreover, there are
clear associations between electrolyte leakage and ethylene. In
ethylene-sensitive owers, ethylene activates enzymes involving
protease and nuclease activity during senescence (Hoeberichts
et al., 2007) through ETHYLENE-INSENSITIVE3 (EIN3)
(Yanagisawa et al., 2003) and ETHYLENE-INSENSITIVE3
‐like 1 (EIL1) (Kakhki et al., 2009), the transcriptional regulator
in the ethylene signalling pathway. Also, an increase in ethylene
increases ROS within the cells (Gan, 2008), and ROS contributes
to the loss of membrane stability (Rogers, 2012). This data
interestingly links to the ability of glucose to act as an antioxidant
in plants mentioned earlier.
Glucose is the most eective sugar for extending hydrangea vase life 29

Journal of Applied Horticulture (www.horticultureresearch.net)
This study provides a new finding essential for improving
hydrangea vase life. The type of sugar is an important factor in
prolonging hydrangea cut ower life. Although sucrose is widely
used in hydrangea vase solutions, this study observed that glucose
was more eective in prolonging ower vase life than sucrose.
Moreover, potent antioxidants like mannitol are unsuitable for
making hydrangea vase life solutions. Glucose acts rapidly
as an osmoprotectant and increases the solution uptake of the
ower. Since hydrangea is a climacteric ower, it is evident that
glucose might generate sepal strength and vase life extension via
interrupting ethylene signalling. As a result, ethylene’s ability to
activate chlorophyll degradation, electrolyte leakage, and ROS
accumulation is disturbed. However, the investigation of ethylene
and antioxidant activity in hydrangea owers treated with glucose
is required for further research to emphasize the relationship
between glucose and ethylene in hydrangea.
Another point to consider is the ornamental value of owers
treated with glucose solution. The appropriate concentration
should be analyzed carefully. Although the solution with 5%
glucose delivered the most extended vase life, the sepal colour
of owers in this treatment was changed to be lighter compared
with others, and inorescence size was the smallest among three
concentration of glucose treatments. Therefore, the solution with
3% glucose could be applied as an eective holding solution if
the sepal colour or inorescence size is concerned.
Acknowledgments
The author would like to thank the Royal Project Foundation
Thailand for the funding.
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Received: November, 2022; Revised: December, 2022; Accepted: January, 2023
Glucose is the most eective sugar for extending hydrangea vase life 31