Preservative Solutions Containing Boric Acid Delay Senescence of Carnation Flowers

Postharvest Biology and Technology 23 (2001) 133–142 www.elsevier.com/locate/postharvbio

Preservative solutions containing boric acid delay senescenceof carnation flowers

Marıa Serrano a,*, Asuncion Amoros a, Marıa Teresa Pretel a,Marıa Concepcion Martınez-Madrid a, Felix Romojaro b

a Escuela Politecnica Superior, Uni�ersidad Miguel Hernandez, Carretera Beniel-Orihuela Km 3.2, 03312 Orihuela (Alicante), Spainb Centro de Edafologıa y Biologıa 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 100mM boric acid or continuous treatment with 1 mM boric acid resulted in strong inhibition of the climacteric ethyleneproduction. Both the pulse and continuous treatments significantly increased flower longevity. Free and conjugated1-aminocyclopropane-1-carboxylic acid (ACC) and ACC oxidase activity increased in carnation petals duringsenescence, although significantly less in boric acid-treated carnations than in control flowers. The levels of putrescineincreased as senescence progressed in both control and boric acid-treated carnations and an increase in spermidinelevels was higher in treated carnations. Abscisic acid levels in petals also increased during senescence, but much lessin boric acid-treated carnations. It is concluded that boric acid prevents the early rise in ethylene production andconsiderably improves carnation vase life. © 2001 Elsevier Science B.V. All rights reserved.

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

1. Introduction

Senescence of carnation flowers is associatedwith a climacteric-like increase in ethylene pro-duction. Preclimacteric flowers produce a lowconstant rate of ethylene. During the climacteric,there is a co-ordinated increase in the activities of1-aminocyclopropane-1-carboxylic acid synthase

(ACC synthase) and ACC oxidase (Peiser, 1986;Serrano et al., 1991), which convert S-adenosyl-methionine (SAM) to ACC and ACC to ethylene,respectively. SAM is also a precursor for thesynthesis 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 ofethylene action, has become an essential tool forthe delay of senescence of climacteric flowers andhas been applied in the cut flower industry (Reidand Wu, 1992). However, STS is a potential envi-

* Corresponding author. Tel.: 34-96-6749616; fax: 34-96-6749619.

E-mail address: [email protected] (M. Serrano).

0925-5214/01/$ - see front matter © 2001 Elsevier Science B.V. All rights reserved.

PII: S 0925 -5214 (01 )00108 -9

M. Serrano et al. / Posthar�est Biology and Technology 23 (2001) 133–142134

ronmental hazard and many countries currentlyprohibit its use. At present, there are very fewalternatives to STS. Some synthetic cyclopropeneshave been shown to bind to the ethylene receptorand so prevent the physiological action of ethylenefor extended periods. They have, for example, beenshown useful in prolonging petunia (Serek et al.,1995) and carnation (Sisler et al., 1996; Sisler andSerek, 1997) longevity. Ethanol has also beenshown to be effective in extending the vase life ofcarnations by inhibiting ethylene production(Podd and Van Staden, 1999).

Aminotriazole (AT) is another compound thatinhibits 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 putativecarcinogen (Sine et al., 1991) and therefore it isdifficult to use as a cut flower preservative com-mercially. Aminooxoacetic acid (AOA), a knowninhibitor of ACC synthase activity (Van Altvorstand Bovy, 1995) is currently being used in Hollandas a pulse treatment in the cut flower industry, inparticular for carnations. However, studies withAOA indicate certain toxicological risks (Wolter-ing et al., 1987) and it is rather expensive. Boricacid, already included in a mixture of chemicalsused to improve flower vase life (Odom, 1954),may be a good competitor as far as price isconcerned.

The objective of our research was to investigatethe role of a novel preservative solution containingboric acid in retarding the senescence of carnationflowers. This solution was applied in pulse andcontinuous treatments. The levels of ACC andsome polyamines were also determined in order toestablish whether the inhibition of ethylene biosyn-thesis by boric acid would increase polyaminecontent, which would indicate that both pathways,ethylene and polyamine biosynthesis, compete fortheir common precursor. In addition, we analyzedthe levels of abscisic acid (ABA), a hormonerelated to maturation, especially in non-climactericfruit (Serrano et al., 1995; Martınez-Madrid et al.,1996; Kondo and Inoue, 1997), in order to deter-mine the role of ABA in controlling carnationsenescence when ethylene biosynthesis is inhibited.

2. Materials and methods

2.1. Plant material

Carnations (Dianthus caryophyllus L. cv.Master) were obtained from a local greenhousein Puerto Lumbreras (Murcia, Spain) and wereharvested at the commercial opening stage(the petals forming an angle of 120° with the baseof the calyx). In the laboratory, the flowerswere trimmed to a 15-cm stem length, randomizedand 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 carnationflowers for 24 h in the preservative solution (PS)containing 2% sucrose as an energy source,Roquat BL-80 (benzalconyl chloride) as amicrobicide and 25, 50, 75 or 100 mM of boricacid, in 25 mM citrate buffer pH 4.5. Afterpulsing, carnation flowers were transferred to dis-tilled water. Flowers pulsed with the PS withoutboric acid served as controls. In the continuoustreatment, carnations were maintained in the samePS containing 1 mM boric acid until the end ofsenescence and flowers maintained in PS withoutboric acid served as controls. The volumes of PSor distilled water were maintained at constantlevels by replenishing the test tubes daily.Each treatment consisted of ten flowers andeach experiment was repeated twice. Since theygave similar results, the results of only one exper-iment are presented. The environmental con-ditions maintained throughout the experimentwere temperature 20–22°C, relative humidity(RH) 75–80% and a 12 h photoperiod usingwhite fluorescent light (74.5 �mol s−1 m−2). Thecontinuous PS plus 1 mM boric acid treatment wasrepeated with 40 flowers using another 40flowers as control. Every 2 days, three car-nations were taken from each treatment (the PSand the PS plus boric acid). Carnation petals wereremoved from the first, second and third whorls ofthe flower and strictly randomized into fourgroups of 15, nine, six and six petals, in whichACC content, ACC oxidase activity andpolyamine and ABA levels, respectively, were ana-lyzed.

M. Serrano et al. / Posthar�est Biology and Technology 23 (2001) 133–142 135

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 siliconseptum for 1 h. After this time, a 1 ml sample ofthe 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 stainlesssteel column with 3.17 mm inner diameter,filled with 80/100 activated alumina. Resultswere expressed as nanoliters of ethylene producedper gram of fresh weight per hour (nl g−1 h−1)and are the mean�S.E. of ten flowers. Another 1ml gas sample of the same jar was used todetermine CO2 concentration in a Shimadzu 14-Bgas chromatograph (Kyoto, Japan). Respirationrate was expressed as milligram of CO2 re-leased by a kilogram of fresh weight per hour (mgkg−1 h−1). Results are the mean�S.E. of tenflowers.

2.3. ACC extraction and assay

Total (free and conjugated ACC) was extractedas previously described (Serrano et al., 1991).Fifteen petals were macerated in a mortar with apestle in 10 ml of 0.2 M trichloroacetic acid. Themacerate was centrifuged at 7000×g for 10 minand the supernatant was used to determine its freeACC content by chemical conversion of ACC toethylene which was then quantified as describedabove. Conjugated ACC was hydrolyzed to freeACC with 2 N HCl. In both cases, measurementswere made in triplicate. A relative calibrationprocedure was used to determine the amount ofACC in samples using a standard curve of ACCfrom Sigma (Poole, Dorset, UK). Results wereexpressed as nmol per gram fresh weight (nmolg−1 f.w.) and are the mean�S.E. of triplicatemeasurements on each of three carnations.

2.4. ACC oxidase acti�ity

ACC oxidase activity was measured in triplicatein each carnation flower. Three carnation petals

were cut into small pieces and enclosed in vialswith 25 mM Tris–Hepes buffer, pH 7.5, contain-ing 1 mM ACC. After 2 h at 30°C and continuousshaking, a 1-ml gas sample of the vial atmospherewas withdrawn and monitored for its ethylenecontent. ACC oxidase activity was expressed asnanolitres of ethylene released per gram freshweight 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 previouslyreported (Serrano et al., 1991). Extracts forpolyamine analysis were prepared by homogeniz-ing six carnation petals in 10 ml of 5% HClO4

using a mortar and pestle. The homogenate wasthen centrifuged for 30 min at 20 000×g and thesupernatant was used to quantify the polyaminecontent in duplicate. A total of 2 ml of thesupernatant were mixed with 2 ml of 4 N NaOHand 20 �l of benzoyl chloride in a glass tube.After vortexing for 15 s, the mixture was incu-bated for 20 min at room temperature. SaturatedNaCl (4 ml) and 4 ml of cold diethyl etherwere then added. The tube content was vortexedfor 15 s and incubated for 30 min at −18°C.Finally, 2 ml of the ether phase (containingbenzoyl-polyamines) were evaporated under nitro-gen and redissolved in 1 ml of methanol (HPLCgrade). Benzoyl-polyamines were analyzed byHPLC using a Hewlett–Packard system, series1100 (Waldbrom, Germany). The elutionsystem consisted of methanol:water (64:36, v/v assolvent), run isocratically with a flow rate of 0.8ml min−1. The benzoyl-polyamines wereeluted through a reverse-phase column(LiChroCart 250-4, 5 �m, Merck, Darmstadt,Germany) and detected by absorbance at 254 nm.A relative calibration procedure was used to de-termine the amounts of polyamines in samplesusing standard curves of putrescine, spermidineand spermine from Sigma and adding hexa-nediamine as the internal standard. Results wereexpressed as nmol per gram fresh weight (nmolg−1 f.w.) and are the mean�S.E. of two mea-surements made independently on three carna-tions.


2.6. Abscisic acid determination

ABA was extracted from six petal samples with5 ml of a solution of 80% acetone containing 100mg l−1 of butylated hydroxytoluene and 0.5 g l−1

of citric acid. The extracts were diluted with a 50mM Tris buffer, pH 7.8, containing 1 mM MgCl2and 150 mM NaCl and then quantified by anenzyme-linked immunosorbent assay (ELISA), us-ing an IgG monoclonal antibody (Idetek Inc., SanBruno, CA), as previously reported (Martınez-Madrid et al., 1996). ABA content was estimatedfrom the standard curve prepared for each partic-ular plate using a spectrophotometer StarFax2100 (Awareness Technology Inc., Palm City,FL). The absorbance was fixed at 405 nm. Foreach extract, four dilutions were made and at leastthree fell on the standard curve. All determina-tions were carried out in dim light. The ABAlevels were consistent with the dilution made andno interference from impurities was detected whenABA standards were added to diluted extracts.Results are expressed as pmol per gram freshweight (pmol g−1 f.w.) and are the mean�S.E.of the extractions made independently from eachof three flowers and each extract was measured inquadruplicate.

2.7. Statistics

Experimental data are the mean�S.E. A vari-ance analysis using Student’s t-test was performedto determine if treatments showed significant dif-ferences (P�0.05).

3. Results and discussion

3.1. Ethylene production and respiration rate

Ethylene production was very low in controlflowers, during the first 7 days at 20°C (Fig. 1A).Autocatalytic ethylene production started on day8, reaching a maximum of 15.6�1.5 nl g−1 h−1

on day 12. Maximum ethylene production coin-cided with the appearance of visible senescencesymptoms, such as petal in-rolling and withering.Pulse treatment of cut carnation flowers with the

PS containing 25 mM boric acid significantlydiminished ethylene production, which reached amaximum of 6.8�1.0 nl g−1 h−1, 4 days laterthan in controls. Pulse treatment with the samesolution containing 50, 75 and 100 mM of boricacid resulted in almost complete inhibition of theclimacteric ethylene production (Fig. 1A). Therespiration rate also increased during carnationsenescence, showing a peak of respiration thatcoincided with the peak of ethylene production(Fig. 1B). The maximum respiration rate of con-trol carnations was significantly higher (P�0.05)than in pulse-treated carnations (Fig. 1B).

Fig. 1. Ethylene production (A) and respiration rate (B) duringsenescence of Master carnation flowers at 20°C after pulsetreatment with PS containing 25, 50, 75 and 100 mM boricacid. Carnation pulsed with PS without boric acid served ascontrol. Each value is the mean�S.E. of ten flowers.


Fig. 2. Ethylene production (A) and respiration rate (B) duringsenescence at 20°C of continuously-treated carnations with PS(control) and PS containing 1 mM boric acid. Data are themean�S.E. of ten flowers.

potent inhibitor of ethylene action (Cook andVan Staden, 1987; Altman and Solomos, 1995)and suggest that the respiratory climacteric maybe regulated by even very low ethylene levels,being suppressed only when the ethylene produc-tion is completely inhibited.

3.2. Flower longe�ity

Flower longevity, defined as the time that car-nation flowers maintain their decorative proper-ties (that is, the time until visible symptoms ofsenescence appear), was significantly increased byboric acid treatment. In pulse-treated flowers withthe PS containing boric acid, longevity increasedwith the strength of the boric acid concentration(Table 1). Visible senescence was also delayed inthe continuous treatment with PS plus 1 mMboric acid (Table 1). In both pulse and continuoustreatments, carnations used as controls weretreated with the PS without boric acid to ensurethat differences between treated and control car-nations were exclusively due to boric acid. Never-theless, in a previous experiment we found thatwith carnations kept in distilled water, longevitywas slightly shorter than in those treated with thePS without boric acid, while ethylene productionwas similar (data not shown).

Continuous and pulse treatments with PS plusboric acid were also effective in inhibiting ethyl-ene production and in prolonging flower longevityin other carnation cultivars, such as Dover,Eroico, Omagio and Oriana (unpublished data).However, no additional increase in longevity wasfound in carnations continuously treated with thePS containing 10 mM boric acid, in which anundesirable leaf chlorosis was observed, probablydue to boric toxicity (unpublished data).

Until recently, it was assumed that ethyleneproduction and petal in-rolling in carnation wereregulated in concert, since these phenomena coin-cided. However, we found that petal in-rollingwas suppressed in carnations flowers pulsed withthe PS containing 75 and 100 mM boric acid, aswell as in those kept continuously in PS plus 1mM boric acid, while it was observed in thosepulsed with a lower concentration (50 mM) ofboric acid, which produced a similar low level of

Continuous treatment with the PS containing 1mM boric acid also drastically inhibited the ethyl-ene production of carnation flowers (Fig. 2A).The peak of respiration rate was significantly(P�0.05) lower in the treated flowers and oc-curred later (Fig. 2B).

Thus, a peak of respiration was observed in allcarnation flowers treated with boric acid (Fig. 1Band Fig. 2B). However, boric acid treatments(except in the pulse with PS containing 25 mMboric acid) virtually prevented the climactericpeak of ethylene production (Fig. 1A and Fig.2A). These results are in agreement with thosefound in carnation flowers treated with STS, a


ethylene. These findings are consistent with theproposal that petal in-rolling and withering areethylene independent. They agree with the resultsof 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 cysteineproteinase (related to withering of petals) wereregulated differently in carnation petals.

The continuous treatment with the PS contain-ing 1 mM boric acid was selected to study theeffect of boric acid on ethylene, polyamines andABA biosynthesis.

3.3. ACC le�els and ACC oxidase acti�ity

Free and total ACC levels were very low inrecently harvested flowers and rose slowly in con-trol flowers during the first 8 days of senescence at20°C (Fig. 3A). After day 8, free and total ACClevels rose sharply (Fig. 3A). In carnations main-tained in the PS with 1 mM boric acid, free andtotal ACC levels in the petals were significantlylower than in control flowers (Fig. 3A). AlthoughACC synthase was not directly measured, resultsshow that boric acid treatment probably inhibitedthis enzyme by �50%, but did not inhibit ACCconjugation, since only a small fraction of theACC synthesized remained in a free state (Fig.3A).

ACC oxidase activity in control carnations wasvery 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 smallincrease being observed on day 16 (Fig. 3B).These results show that the lack of ethylene pro-duction in boric acid-treated carnations could bedue to a combination of two causes: a lowavailability of free ACC, probably resulting fromlow ACC synthase activity, and the failure toconvert ACC into ethylene because of decreasedACC oxidase activity.

In untreated carnations, the ability of petaltissue to convert exogenous ACC into ethylene inthe presence of boric acid was assayed at twodevelopmental stages. Ethylene production in-creased with increasing ACC concentrations inboth preclimacteric and climacteric carnations(Table 2). The higher ethylene production rate inclimacteric petals shows that ACC oxidase activ-ity increased during senescence (as can also beinferred from Fig. 3B). However, no significantdifferences in ethylene production were foundwhen 10 and 20 mM boric acid were added to theincubation medium. This finding shows that boricacid does not directly inhibit ACC oxidase activ-ity, as do cobalt ions and �-aminoisobutyric acid(Serrano et al., 1990). Thus, the low ACC oxidaseactivity and ACC levels found in boric acid-treated carnation petals during senescence may bedue to the effect of boric acid on the synthesis ofboth ACC synthase and ACC oxidase, whichnormally occurs during carnation senescence

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

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

0 25 1007550

Longevity 22.2�0.5c21.7�0.7c20.8�0.7c16.7�1.0b12.2�0.4a

Continuous treatment (boric acid (mM) concentration in the PS)

0 1

Longevity 11.8�0.6a 18.59�0.7b

a Each experiment consisted of ten flowers. Means within each treatment group followed by different letters are significantlydifferent (P�0.05).


Fig. 3. Free and total ACC content (A) and ACC oxidaseactivity (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�S.E. of mea-surements made in triplicate in each of three flowers.

the first 8 days of senescence and from day 8 theyincreased sharply (Fig. 4A). In boric acid-treatedcarnations putrescine also increased during senes-cence, although slowly, reaching a final valuewhich was not significantly different from thelevels in controls (Fig. 4A).

Spermidine content was significantly higherthan the putrescine content (Fig. 4A,B), and de-creased until day 10 in both control and treatedcarnation petals. Then, spermidine content in-creased again until the end of senescence. A simi-lar pattern of spermidine behaviour has beenreported in other carnation cultivars (Roberts etal., 1984; Serrano et al., 1991). In carnationstreated with the PS plus 1 mM boric acid, sper-midine levels increased more than in controls(Fig. 4B).

Thus, taking into account the antisenescentproperties of the polyamines spermidine and sper-mine (Tiburcio et al., 1997), the high spermidinelevels could be responsible for the greaterlongevity of the boric acid-treated carnations.However, the rise in spermidine seems to cometoo late to account for the delay in the ethylenerise. The sharp increase in spermidine levels in thepetals of boric acid-treated carnation (in which

Table 2Ethylene production (nl g−1 h−1) of carnations petals at twostages of development after incubation for 2 h with variousACC concentrations plus 10 or 20 mM boric acida

ACC Boric acid concentration (mM)concentration

10 20(mM) 0

Preclimacteric stage19.1�3.1a22.3�3.7a 25.4�3.2a0.1

157.4�11.6b0.5 132.6�12.9b 149.6�14.8b

282.7�25.0c258.1�12.9c 294.0�21.5c1

Climacteric stage0.1 226.5�26.8a 204.0�19.0a229.9�19.2a

267.2�8.7b0.5 285.4�22.2b 269.1�21.0b

1 367.8�24.3c 386.5�25.7c 361.1�17.0c

a Data are the mean�S.E. of two replicates of three petalsamples per treatment. Means within each row and column(for each development stage) followed by different letters aresignificantly different (P�0.05).

(Jones and Woodson 1999; Satoh et al., 2000).Another possibility is that the low ACC oxidaseactivity could be due to substrate limitation (Fig.3). Since ACC synthase and ACC oxidase activityare in an autocatalytic loop, the present data donot suggest which part of the chain is broken.

3.4. Polyamine le�els

The most abundant polyamines in the carna-tion petals of the cultivar Master were putrescineand spermidine. Spermine levels were very low,just at or below the detection limit. In controlflowers, putrescine levels increased slowly during


Fig. 4. Putrescine (A) and spermidine (B) levels in petals ofcontinuously-treated carnations with PS (control) and PS con-taining 1 mM boric acid, during senescence at 20°C. Data arethe mean�S.E. of quantification made in duplicate in each ofthree flowers.

previously (Eze et al., 1986; Hanley and Bram-lage, 1989). Application of exogenous ABA tocarnation flowers (through the cut stem end)caused a rapid increase in the ABA content inflower tissues and promoted ethylene productionin the flowers (Onoue et al., 2000). These findingsare in agreement with the proposal that ABA mayplay a role in the induction of ethylene productionduring natural senescence in carnation flowers.

In boric-acid treated carnations (Fig. 5), as wellas in those treated with AT (Serrano et al., 1999),the ABA content increased to half of the levelreached in control carnations, whereas the ethyl-ene production of treated carnations was verylow, which might indicate a separation of therelationship between these two hormones. In ad-dition, the ABA levels reached in boric acid-treated carnations were lower than that in controlflowers, which may point to an inhibition of ABAbiosynthesis by boric acid either directly or bysome intermediate effect.

4. Conclusions

We conclude that a pulse or continuous treat-ment with a preservative solution containing boricacid was effective in prolonging carnation flower

Fig. 5. Evolution of petal ABA content during senescence at20°C of continuously-treated carnations with PS (control) andPS containing 1 mM boric acid. Data are the mean�S.E. ofmeasurements made in quadruplicate in each of three flowers.

ethylene production was inhibited) may also indi-cate competition between the ethylene andpolyamine pathways for their common precursor.

3.5. ABA le�els

The ABA content in freshly harvested carna-tions was low and levels accumulated from day 6onwards (Fig. 5). In carnations continuouslytreated with the PS containing 1 mM boric acid,ABA started to increase from day 12 (Fig. 5) andreached values significantly lower than those incontrol flowers. ABA thus accumulated before theonset of ethylene production, as has been reported


longevity and could be useful in the cut flowerindustry. Boric acid treatments apparently inhib-ited ethylene production in carnation flowers bylowering ACC synthase activity and consequentlythe availability of ACC and probably also byinhibiting the synthesis of ACC oxidase. In addi-tion, spermidine levels were higher in the boricacid treated carnations, whereas the increase ofABA levels was lower. Thus, increased spermidinelevels and lower ABA levels may also be responsi-ble for the inhibition of ethylene production ofboric acid treated carnations.

Acknowledgements

This study was funded by the Comision Inter-ministerial de Ciencia y Tecnologıa (CICYT),project PETRI-95-0271-OP.

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Preservative Solutions Containing Boric Acid Delay Senescence of Carnation Flowers

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