Postharvest Biology and Technology 23 (2001) 133– 142 www.elsevier.

com/locate/postharvbio

Preservative solutions containing boric acid delay senescence

of carnation flowers
Marı́a Serrano a,*, Asunción Amorós a, Marı́a Teresa Pretel a,
Marı́a Concepción Martı́nez-Madrid a, Félix Romojaro b
a
Escuela Politécnica Superior, Uni6ersidad Miguel Hernández, Carretera Beniel-Orihuela Km 3.2, 03312 Orihuela (Alicante), Spain
b
Centro de Edafologı́a y Biologı́a Aplicada del Segura (CSIC), Campus de Espinardo, 30100 Murcia, Spain

Received 21 November 2000; accepted 27 February 2001

Abstract

We investigated the effect of a preservative solution containing boric acid on the senescence of cut carnation flowers
(Dianthus caryophyllus L. cv. Master). A 24-h pulse treatment with the preservative solution containing 50, 75 or 100
mM boric acid or continuous treatment with 1 mM boric acid resulted in strong inhibition of the climacteric ethylene
production. Both the pulse and continuous treatments significantly increased flower longevity. Free and conjugated
1-aminocyclopropane-1-carboxylic acid (ACC) and ACC oxidase activity increased in carnation petals during
senescence, although significantly less in boric acid-treated carnations than in control flowers. The levels of putrescine
increased as senescence progressed in both control and boric acid-treated carnations and an increase in spermidine
levels was higher in treated carnations. Abscisic acid levels in petals also increased during senescence, but much less
in boric acid-treated carnations. It is concluded that boric acid prevents the early rise in ethylene production and
considerably improves carnation vase life. © 2001 Elsevier Science B.V. All rights reserved.

Keywords: Abscisic acid; Boric acid; Dianthus caryophyllus; Ethylene; Polyamines; Senescence

1. Introduction (ACC synthase) and ACC oxidase (Peiser, 1986;

Senescence of carnation flowers is associated
with a climacteric-like increase in ethylene pro-
duction. Preclimacteric flowers produce a low
constant rate of ethylene. During the climacteric,
there is a co-ordinated increase in the activities of
1-aminocyclopropane-1-carboxylic acid synthase
* Corresponding author. Tel.: 34-96-6749616; fax: 34-96-
6749619.
E-mail address: m.serrano@umh.es (M. Serrano).
Serrano et al., 1991), which convert S-adenosyl-
methionine (SAM) to ACC and ACC to ethylene,
respectively. SAM is also a precursor for the
synthesis of the polyamines spermidine and sper-
mine, which are related to young or actively grow-
ing tissues (Tiburcio et al., 1997).
Silver thiosulfate (STS), a known inhibitor of
ethylene action, has become an essential tool for
the delay of senescence of climacteric flowers and
has been applied in the cut flower industry (Reid
and Wu, 1992). However, STS is a potential envi-

0925-5214/01/$ - see front matter © 2001 Elsevier Science B.V. All rights reserved.
PII: S 0 9 2 5 - 5 2 1 4 ( 0 1 ) 0 0 1 0 8 - 9
134 M. Serrano et al. / Posthar6est Biology and Technology 23 (2001) 133–142
ronmental hazard and many countries currently
prohibit its use. At present, there are very few
alternatives to STS. Some synthetic cyclopropenes
have been shown to bind to the ethylene receptor
and so prevent the physiological action of ethylene
for extended periods. They have, for example, been
shown useful in prolonging petunia (Serek et al.,
1995) and carnation (Sisler et al., 1996; Sisler and
Serek, 1997) longevity. Ethanol has also been
shown to be effective in extending the vase life of
carnations by inhibiting ethylene production
(Podd and Van Staden, 1999).
Aminotriazole (AT) is another compound that
inhibits the climacteric peak of ethylene produc-
tion and prolongs the vase life of carnation flowers
(Altman and Solomos, 1994; Serrano et al., 1999).
However, AT has been classified as a putative
carcinogen (Sine et al., 1991) and therefore it is
difficult to use as a cut flower preservative com-
mercially. Aminooxoacetic acid (AOA), a known
inhibitor of ACC synthase activity (Van Altvorst
and Bovy, 1995) is currently being used in Holland
as a pulse treatment in the cut flower industry, in
particular for carnations. However, studies with
AOA indicate certain toxicological risks (Wolter-
ing et al., 1987) and it is rather expensive. Boric
acid, already included in a mixture of chemicals
used to improve flower vase life (Odom, 1954),
may be a good competitor as far as price is
concerned.
The objective of our research was to investigate
the role of a novel preservative solution containing
boric acid in retarding the senescence of carnation
flowers. This solution was applied in pulse and
continuous treatments. The levels of ACC and
some polyamines were also determined in order to
establish whether the inhibition of ethylene biosyn-
thesis by boric acid would increase polyamine
content, which would indicate that both pathways,
ethylene and polyamine biosynthesis, compete for
their common precursor. In addition, we analyzed
the levels of abscisic acid (ABA), a hormone
related to maturation, especially in non-climacteric
fruit (Serrano et al., 1995; Martı́nez-Madrid et al.,
1996; Kondo and Inoue, 1997), in order to deter-
mine the role of ABA in controlling carnation
senescence when ethylene biosynthesis is inhibited.
2. Materials and methods
2.1. Plant material
Carnations (Dianthus caryophyllus L. cv.
Master) were obtained from a local greenhouse
in Puerto Lumbreras (Murcia, Spain) and were
harvested at the commercial opening stage
(the petals forming an angle of 120° with the base
of the calyx). In the laboratory, the flowers
were trimmed to a 15-cm stem length, randomized
and placed individually in test tubes con-
taining the various treatments. This was consid-
ered to be day 0 of the experiment. Pulse treat-
ments were performed, holding the carnation
flowers for 24 h in the preservative solution (PS)
containing 2% sucrose as an energy source,
Roquat BL-80 (benzalconyl chloride) as a
microbicide and 25, 50, 75 or 100 mM of boric
acid, in 25 mM citrate buffer pH 4.5. After
pulsing, carnation flowers were transferred to dis-
tilled water. Flowers pulsed with the PS without
boric acid served as controls. In the continuous
treatment, carnations were maintained in the same
PS containing 1 mM boric acid until the end of
senescence and flowers maintained in PS without
boric acid served as controls. The volumes of PS
or distilled water were maintained at constant
levels by replenishing the test tubes daily.
Each treatment consisted of ten flowers and
each experiment was repeated twice. Since they
gave similar results, the results of only one exper-
iment are presented. The environmental con-
ditions maintained throughout the experiment
were temperature 20–22°C, relative humidity
(RH) 75 –80% and a 12 h photoperiod using
white fluorescent light (74.5 mmol s − 1 m − 2). The
continuous PS plus 1 mM boric acid treatment was
repeated with 40 flowers using another 40
flowers as control. Every 2 days, three car-
nations were taken from each treatment (the PS
and the PS plus boric acid). Carnation petals were
removed from the first, second and third whorls of
the flower and strictly randomized into four
groups of 15, nine, six and six petals, in which
ACC content, ACC oxidase activity and
polyamine and ABA levels, respectively, were ana-
lyzed.
 M. Serrano et al. / Posthar6est Biology and Technology 23 (2001) 133–142
2.2. Ethylene production and respiration rate
To measure ethylene production and respira-
tion rate, cut carnations were individually en-
closed in 500 ml glass jars fitted with a silicon
septum for 1 h. After this time, a 1 ml sample of
the jar atmosphere was withdrawn and the ethyl-
ene concentration determined using a Hewlett–
Packard model 5890 gas chromatograph
(Wilmington, DE) equipped with a flame ioniza-
tion detector (FID) and a 3 m length stainless
steel column with 3.17 mm inner diameter,
filled with 80/100 activated alumina. Results
were expressed as nanoliters of ethylene produced
per gram of fresh weight per hour (nl g − 1 h − 1)
and are the mean9S.E. of ten flowers. Another 1
ml gas sample of the same jar was used to
determine CO2 concentration in a Shimadzu 14-B
gas chromatograph (Kyoto, Japan). Respiration
rate was expressed as milligram of CO2 re-
leased by a kilogram of fresh weight per hour (mg
kg − 1 h − 1). Results are the mean9 S.E. of ten
flowers.
2.3. ACC extraction and assay
Total (free and conjugated ACC) was extracted
as previously described (Serrano et al., 1991).
Fifteen petals were macerated in a mortar with a
pestle in 10 ml of 0.2 M trichloroacetic acid. The
macerate was centrifuged at 7000×g for 10 min
and the supernatant was used to determine its free
ACC content by chemical conversion of ACC to
ethylene which was then quantified as described
above. Conjugated ACC was hydrolyzed to free
ACC with 2 N HCl. In both cases, measurements
were made in triplicate. A relative calibration
procedure was used to determine the amount of
ACC in samples using a standard curve of ACC
from Sigma (Poole, Dorset, UK). Results were
expressed as nmol per gram fresh weight (nmol
g − 1 f.w.) and are the mean9 S.E. of triplicate
measurements on each of three carnations.
2.4. ACC oxidase acti6ity
ACC oxidase activity was measured in triplicate
in each carnation flower. Three carnation petals
135
were cut into small pieces and enclosed in vials
with 25 mM Tris– Hepes buffer, pH 7.5, contain-
ing 1 mM ACC. After 2 h at 30°C and continuous
shaking, a 1-ml gas sample of the vial atmosphere
was withdrawn and monitored for its ethylene
content. ACC oxidase activity was expressed as
nanolitres of ethylene released per gram fresh
weight per hour (nl g − 1 h − 1).
2.5. Polyamine extraction and quantification
Polyamines were extracted with HClO4 and an-
alyzed by the benzoylation method, as previously
reported (Serrano et al., 1991). Extracts for
polyamine analysis were prepared by homogeniz-
ing six carnation petals in 10 ml of 5% HClO4
using a mortar and pestle. The homogenate was
then centrifuged for 30 min at 20 000× g and the
supernatant was used to quantify the polyamine
content in duplicate. A total of 2 ml of the
supernatant were mixed with 2 ml of 4 N NaOH
and 20 ml of benzoyl chloride in a glass tube.
After vortexing for 15 s, the mixture was incu-
bated for 20 min at room temperature. Saturated
NaCl (4 ml) and 4 ml of cold diethyl ether
were then added. The tube content was vortexed
for 15 s and incubated for 30 min at − 18°C.
Finally, 2 ml of the ether phase (containing
benzoyl-polyamines) were evaporated under nitro-
gen and redissolved in 1 ml of methanol (HPLC
grade). Benzoyl-polyamines were analyzed by
HPLC using a Hewlett–Packard system, series
1100 (Waldbrom, Germany). The elution
system consisted of methanol:water (64:36, v/v as
solvent), run isocratically with a flow rate of 0.8
ml min − 1. The benzoyl-polyamines were
eluted through a reverse-phase column
(LiChroCart 250-4, 5 mm, Merck, Darmstadt,
Germany) and detected by absorbance at 254 nm.
A relative calibration procedure was used to de-
termine the amounts of polyamines in samples
using standard curves of putrescine, spermidine
and spermine from Sigma and adding hexa-
nediamine as the internal standard. Results were
expressed as nmol per gram fresh weight (nmol
g − 1 f.w.) and are the mean9S.E. of two mea-
surements made independently on three carna-
tions.
136 M. Serrano et al. / Posthar6est Biology and Technology 23 (2001) 133–142

2.6. Abscisic acid determination PS containing 25 mM boric acid significantly

diminished ethylene production, which reached a
ABA was extracted from six petal samples with maximum of 6.89 1.0 nl g − 1 h − 1, 4 days later
5 ml of a solution of 80% acetone containing 100 than in controls. Pulse treatment with the same
mg l − 1 of butylated hydroxytoluene and 0.5 g l − 1 solution containing 50, 75 and 100 mM of boric
of citric acid. The extracts were diluted with a 50 acid resulted in almost complete inhibition of the
mM Tris buffer, pH 7.8, containing 1 mM MgCl2 climacteric ethylene production (Fig. 1A). The
and 150 mM NaCl and then quantified by an respiration rate also increased during carnation
enzyme-linked immunosorbent assay (ELISA), us- senescence, showing a peak of respiration that
ing an IgG monoclonal antibody (Idetek Inc., San coincided with the peak of ethylene production
Bruno, CA), as previously reported (Martı́nez- (Fig. 1B). The maximum respiration rate of con-
Madrid et al., 1996). ABA content was estimated
trol carnations was significantly higher (PB 0.05)
from the standard curve prepared for each partic-
than in pulse-treated carnations (Fig. 1B).
ular plate using a spectrophotometer StarFax
2100 (Awareness Technology Inc., Palm City,
FL). The absorbance was fixed at 405 nm. For
each extract, four dilutions were made and at least
three fell on the standard curve. All determina-
tions were carried out in dim light. The ABA
levels were consistent with the dilution made and
no interference from impurities was detected when
ABA standards were added to diluted extracts.
Results are expressed as pmol per gram fresh
weight (pmol g − 1 f.w.) and are the mean9S.E.
of the extractions made independently from each
of three flowers and each extract was measured in
quadruplicate.

2.7. Statistics

Experimental data are the mean9S.E. A vari-

ance analysis using Student’s t-test was performed
to determine if treatments showed significant dif-
ferences (PB 0.05).

3. Results and discussion

3.1. Ethylene production and respiration rate

Ethylene production was very low in control

flowers, during the first 7 days at 20°C (Fig. 1A).
Autocatalytic ethylene production started on day
8, reaching a maximum of 15.691.5 nl g − 1 h − 1
Fig. 1. Ethylene production (A) and respiration rate (B) during
on day 12. Maximum ethylene production coin-
senescence of Master carnation flowers at 20°C after pulse
cided with the appearance of visible senescence treatment with PS containing 25, 50, 75 and 100 mM boric
symptoms, such as petal in-rolling and withering. acid. Carnation pulsed with PS without boric acid served as
Pulse treatment of cut carnation flowers with the control. Each value is the mean 9S.E. of ten flowers.
 M. Serrano et al. / Posthar6est Biology and Technology 23 (2001) 133–142
Fig. 2. Ethylene production (A) and respiration rate (B) during
senescence at 20°C of continuously-treated carnations with PS
(control) and PS containing 1 mM boric acid. Data are the
mean9 S.E. of ten flowers.
Continuous treatment with the PS containing 1
mM boric acid also drastically inhibited the ethyl-
ene production of carnation flowers (Fig. 2A).
The peak of respiration rate was significantly
(PB 0.05) lower in the treated flowers and oc-
curred later (Fig. 2B).
Thus, a peak of respiration was observed in all
carnation flowers treated with boric acid (Fig. 1B
and Fig. 2B). However, boric acid treatments
(except in the pulse with PS containing 25 mM
boric acid) virtually prevented the climacteric
peak of ethylene production (Fig. 1A and Fig.
2A). These results are in agreement with those
found in carnation flowers treated with STS, a
137
potent inhibitor of ethylene action (Cook and
Van Staden, 1987; Altman and Solomos, 1995)
and suggest that the respiratory climacteric may
be regulated by even very low ethylene levels,
being suppressed only when the ethylene produc-
tion is completely inhibited.
3.2. Flower longe6ity
Flower longevity, defined as the time that car-
nation flowers maintain their decorative proper-
ties (that is, the time until visible symptoms of
senescence appear), was significantly increased by
boric acid treatment. In pulse-treated flowers with
the PS containing boric acid, longevity increased
with the strength of the boric acid concentration
(Table 1). Visible senescence was also delayed in
the continuous treatment with PS plus 1 mM
boric acid (Table 1). In both pulse and continuous
treatments, carnations used as controls were
treated with the PS without boric acid to ensure
that differences between treated and control car-
nations were exclusively due to boric acid. Never-
theless, in a previous experiment we found that
with carnations kept in distilled water, longevity
was slightly shorter than in those treated with the
PS without boric acid, while ethylene production
was similar (data not shown).
Continuous and pulse treatments with PS plus
boric acid were also effective in inhibiting ethyl-
ene production and in prolonging flower longevity
in other carnation cultivars, such as Dover,
Eroico, Omagio and Oriana (unpublished data).
However, no additional increase in longevity was
found in carnations continuously treated with the
PS containing 10 mM boric acid, in which an
undesirable leaf chlorosis was observed, probably
due to boric toxicity (unpublished data).
Until recently, it was assumed that ethylene
production and petal in-rolling in carnation were
regulated in concert, since these phenomena coin-
cided. However, we found that petal in-rolling
was suppressed in carnations flowers pulsed with
the PS containing 75 and 100 mM boric acid, as
well as in those kept continuously in PS plus 1
mM boric acid, while it was observed in those
pulsed with a lower concentration (50 mM) of
boric acid, which produced a similar low level of
138 M. Serrano et al. / Posthar6est Biology and Technology 23 (2001) 133–142

ethylene. These findings are consistent with the
proposal that petal in-rolling and withering are
ethylene independent. They agree with the results
of Satoh et al. (2000) and Onoue et al. (2000),
who reported that the expression of ACC syn-
thase and ACC oxidase genes and that of cysteine
proteinase (related to withering of petals) were
regulated differently in carnation petals.
The continuous treatment with the PS contain-
ing 1 mM boric acid was selected to study the
effect of boric acid on ethylene, polyamines and
ABA biosynthesis.
3.3. ACC le6els and ACC oxidase acti6ity
Free and total ACC levels were very low in
recently harvested flowers and rose slowly in con-
trol flowers during the first 8 days of senescence at
20°C (Fig. 3A). After day 8, free and total ACC
levels rose sharply (Fig. 3A). In carnations main-
tained in the PS with 1 mM boric acid, free and
total ACC levels in the petals were significantly
lower than in control flowers (Fig. 3A). Although
ACC synthase was not directly measured, results
show that boric acid treatment probably inhibited
this enzyme by \50%, but did not inhibit ACC
conjugation, since only a small fraction of the
ACC synthesized remained in a free state (Fig.
3A).
ACC oxidase activity in control carnations was
very low during preclimacteric stages but in-
creased along with ethylene production, peaking
on day 12 (Fig. 3B). However, in carnation flow-
ers kept continuously in PS with 1 mM boric acid,
ACC oxidase remained very low, only a small
increase being observed on day 16 (Fig. 3B).
These results show that the lack of ethylene pro-
duction in boric acid-treated carnations could be
due to a combination of two causes: a low
availability of free ACC, probably resulting from
low ACC synthase activity, and the failure to
convert ACC into ethylene because of decreased
ACC oxidase activity.
In untreated carnations, the ability of petal
tissue to convert exogenous ACC into ethylene in
the presence of boric acid was assayed at two
developmental stages. Ethylene production in-
creased with increasing ACC concentrations in
both preclimacteric and climacteric carnations
(Table 2). The higher ethylene production rate in
climacteric petals shows that ACC oxidase activ-
ity increased during senescence (as can also be
inferred from Fig. 3B). However, no significant
differences in ethylene production were found
when 10 and 20 mM boric acid were added to the
incubation medium. This finding shows that boric
acid does not directly inhibit ACC oxidase activ-
ity, as do cobalt ions and a-aminoisobutyric acid
(Serrano et al., 1990). Thus, the low ACC oxidase
activity and ACC levels found in boric acid-
treated carnation petals during senescence may be
due to the effect of boric acid on the synthesis of
both ACC synthase and ACC oxidase, which
normally occurs during carnation senescence

Table 1
Longevity (days) of Master carnation flowers after pulse treatment with a pretreatment solution containing various concentrations
of boric acid or continuously kept in the solution containing 1 mM boric acida

Pulse treatments (boric acid (mM) concentration in the PS)

0 25 50 75 100

Longevity 12.2 90.4a 16.79 1.0b 20.8 9 0.7c 21.7 9 0.7c 22.29 0.5c
Continuous treatment (boric acid (mM) concentration in the PS)

0 1

Longevity 11.8 90.6a 18.59 90.7b

a
Each experiment consisted of ten flowers. Means within each treatment group followed by different letters are significantly
different (PB0.05).
 M. Serrano et al. / Posthar6est Biology and Technology 23 (2001) 133–142
Fig. 3. Free and total ACC content (A) and ACC oxidase
activity (B) evolution in petals from continuously-treated car-
nations with PS (control) and PS containing 1 mM boric acid,
during senescence at 20°C. Data are the mean 9 S.E. of mea-
surements made in triplicate in each of three flowers.
(Jones and Woodson 1999; Satoh et al., 2000).
Another possibility is that the low ACC oxidase
activity could be due to substrate limitation (Fig.
3). Since ACC synthase and ACC oxidase activity
are in an autocatalytic loop, the present data do
not suggest which part of the chain is broken.
3.4. Polyamine le6els
The most abundant polyamines in the carna-
tion petals of the cultivar Master were putrescine
and spermidine. Spermine levels were very low,
just at or below the detection limit. In control
flowers, putrescine levels increased slowly during
139
the first 8 days of senescence and from day 8 they
increased sharply (Fig. 4A). In boric acid-treated
carnations putrescine also increased during senes-
cence, although slowly, reaching a final value
which was not significantly different from the
levels in controls (Fig. 4A).
Spermidine content was significantly higher
than the putrescine content (Fig. 4A,B), and de-
creased until day 10 in both control and treated
carnation petals. Then, spermidine content in-
creased again until the end of senescence. A simi-
lar pattern of spermidine behaviour has been
reported in other carnation cultivars (Roberts et
al., 1984; Serrano et al., 1991). In carnations
treated with the PS plus 1 mM boric acid, sper-
midine levels increased more than in controls
(Fig. 4B).
Thus, taking into account the antisenescent
properties of the polyamines spermidine and sper-
mine (Tiburcio et al., 1997), the high spermidine
levels could be responsible for the greater
longevity of the boric acid-treated carnations.
However, the rise in spermidine seems to come
too late to account for the delay in the ethylene
rise. The sharp increase in spermidine levels in the
petals of boric acid-treated carnation (in which
Table 2
Ethylene production (nl g−1 h−1) of carnations petals at two
stages of development after incubation for 2 h with various
ACC concentrations plus 10 or 20 mM boric acida
ACC Boric acid concentration (mM)
concentration
(mM) 0 10 20
Preclimacteric stage
0.1 22.3 9 3.7a 19.1 93.1a 25.4 9 3.2a
0.5 157.4 911.6b 132.6 912.9b 149.6 914.8b
1 258.1 912.9c 282.7 925.0c 294.0 921.5c
Climacteric stage
0.1 229.9 919.2a 226.5 926.8a 204.0 919.0a
0.5 267.2 98.7b 285.4 922.2b 269.1 921.0b
1 367.8 924.3 386.5 925.7c
c
361.1 917.0c
a
Data are the mean 9S.E. of two replicates of three petal
samples per treatment. Means within each row and column
(for each development stage) followed by different letters are
significantly different (PB0.05).
140 M. Serrano et al. / Posthar6est Biology and Technology 23 (2001) 133–142

previously (Eze et al., 1986; Hanley and Bram-

lage, 1989). Application of exogenous ABA to
carnation flowers (through the cut stem end)
caused a rapid increase in the ABA content in
flower tissues and promoted ethylene production
in the flowers (Onoue et al., 2000). These findings
are in agreement with the proposal that ABA may
play a role in the induction of ethylene production
during natural senescence in carnation flowers.
In boric-acid treated carnations (Fig. 5), as well
as in those treated with AT (Serrano et al., 1999),
the ABA content increased to half of the level
reached in control carnations, whereas the ethyl-
ene production of treated carnations was very
low, which might indicate a separation of the
relationship between these two hormones. In ad-
dition, the ABA levels reached in boric acid-
treated carnations were lower than that in control
flowers, which may point to an inhibition of ABA
biosynthesis by boric acid either directly or by
some intermediate effect.

4. Conclusions

We conclude that a pulse or continuous treat-

ment with a preservative solution containing boric
acid was effective in prolonging carnation flower
Fig. 4. Putrescine (A) and spermidine (B) levels in petals of
continuously-treated carnations with PS (control) and PS con-
taining 1 mM boric acid, during senescence at 20°C. Data are
the mean 9S.E. of quantification made in duplicate in each of
three flowers.

ethylene production was inhibited) may also indi-

cate competition between the ethylene and
polyamine pathways for their common precursor.

3.5. ABA le6els

The ABA content in freshly harvested carna-

tions was low and levels accumulated from day 6
onwards (Fig. 5). In carnations continuously
treated with the PS containing 1 mM boric acid,
ABA started to increase from day 12 (Fig. 5) and
Fig. 5. Evolution of petal ABA content during senescence at
reached values significantly lower than those in 20°C of continuously-treated carnations with PS (control) and
control flowers. ABA thus accumulated before the PS containing 1 mM boric acid. Data are the mean 9S.E. of
onset of ethylene production, as has been reported measurements made in quadruplicate in each of three flowers.
 M. Serrano et al. / Posthar6est Biology and Technology 23 (2001) 133–142
longevity and could be useful in the cut flower
industry. Boric acid treatments apparently inhib-
ited ethylene production in carnation flowers by
lowering ACC synthase activity and consequently
the availability of ACC and probably also by
inhibiting the synthesis of ACC oxidase. In addi-
tion, spermidine levels were higher in the boric
acid treated carnations, whereas the increase of
ABA levels was lower. Thus, increased spermidine
levels and lower ABA levels may also be responsi-
ble for the inhibition of ethylene production of
boric acid treated carnations.
Acknowledgements
This study was funded by the Comisión Inter-
ministerial de Ciencia y Tecnologı́a (CICYT),
project PETRI-95-0271-OP.
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