Journal of Experimental Botany

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Journal of Experimental Botany, Vol. 51, No. 353, pp. 2067-2073, December 2000
© 2000 Oxford University Press

Original Papers

Role of the gynoecium in natural senescence of carnation (Dianthus caryophyllus L.) flowers

Kenichi Shibuya, Toshihito Yoshioka, Teruyoshi Hashiba and Shigeru Satoh¹

Laboratory of Bioadaptation, Graduate School of Agricultural Science, Tohoku University, Tsutsumidori-amamiyamachi 1-1, Aoba-ku, Sendai 981-8555, Japan

Received 21 March 2000; Accepted 25 July 2000

	Abstract

Top Abstract Introduction Materials and methods Results Discussion References

 
Although the role of the gynoecium in natural senescence of the carnation flower has long been suggested, it has remained a matter of dispute because petal senescence in the cut carnation flower was not delayed by the removal of gynoecium. In this study, the gynoecium was snapped off by hand, in contrast to previous investigations where removal was achieved by forceps or scissors. The removal of the gynoecium by hand prevented the onset of ethylene production and prolonged the vase life of the flower, demonstrating a decisive role of the gynoecium in controlling natural senescence of the carnation flower. Abscisic acid (ABA) and indole-3-acetic acid (IAA), which induced ethylene production and accelerated petal senescence in carnation flowers, did not stimulate ethylene production in the flowers with gynoecia removed (–Gyn flowers). Application of 1-aminocyclopropane-1-carboxylate (ACC), the ethylene precursor, induced substantial ethylene production and petal wilting in the flowers with gynoecia left intact, but was less effective at stimulating ethylene production in the –Gyn flowers and negligible petal in-rolling was observed. Exogenous ethylene induced autocatalytic production of the gas and petal wilting in the –Gyn flowers. These results indicated that ethylene generated in the gynoecium triggers the onset of ethylene production in the petals of carnation during natural senescence.

Key words: Carnation, Dianthus caryophyllus, ethylene production, gynoecium, senescence.

	Introduction

Top Abstract Introduction Materials and methods Results Discussion References

 
Ethylene is the primary plant hormone involved in the senescence of cut carnation flowers (Abeles et al., 1992; Borochov and Woodson, 1989; Reid and Wu, 1992). A large amount of ethylene is synthesized several days after full opening of the flower during natural senescence (Manning, 1985; Peiser, 1986; Woodson et al., 1992), or several hours after compatible pollination (Nichols, 1977; Nichols et al., 1983; Larsen et al., 1995) or treatment with exogenous ethylene (Borochov and Woodson, 1989; Wang and Woodson, 1989; ten Have and Woltering, 1997). The increased ethylene production accelerates in-rolling of petals resulting in wilting of the flower. Ethylene is synthesized through the following pathway: L-methionine S-adenosyl-L-methionine ACC ethylene. ACC synthase and ACC oxidase catalyse the last two reactions (Yang and Hoffman, 1984; Kende, 1993).

Although the petals make a substantial contribution to the ethylene that is produced during natural or pollination-induced senescence in carnation flowers, the gynoecium produces a significant amount of ethylene before the onset of production of the gas in the petals (Nichols, 1977; Woodson et al., 1992; Jones and Woodson, 1997; ten Have and Woltering, 1997). Thus, it has been hypothesized that the gynoecium plays an important role in controlling ethylene production in the flower during natural senescence and senescence induced by pollination. In the pollination-induced senescence of the carnation flower, a pollination signal is produced by the style, most probably on the stigma surface, and this signal is translocated through the ovary to the petals, where it induces autocatalytic ethylene production resulting in wilting of the petals (Jones and Woodson, 1997, 1999a). Concomitant with the transport of the signal, sequential expression of the genes for ACC synthase and ACC oxidase occurs in the respective floral organs (Jones and Woodson, 1997, 1999b). It was recently shown that the primary signal is ethylene, although ACC may partly act as a secondary stimulus (Jones and Woodson, 1999a).

During natural senescence, ACC content and ethylene production increase in the gynoecium earlier than in the petals (Nichols, 1977; Hsieh and Sacalis, 1986; Veen and Kwakkenbos, 1982/1983, 1984). Recently, ten Have and Woltering have shown that the expression of ACC synthase and ACC oxidase genes and ethylene production first started in the ovary followed by the style and petals in the senescing carnation flower (ten Have and Woltering, 1997). They also showed that removal of the petals before the ethylene production in the petal had become autocatalytic suppressed ethylene production in the detached petals and greatly prolonged the life of the petals. These findings strongly support a role for the gynoecium in controlling the natural senescence of the carnation flower. However, it has been reported that the removal of the gynoecium did not delay the petal senescence of the cut carnation flower held in water (Mor et al., 1980; Sacalis and Lee, 1987; Woodson and Brandt, 1991), and isolated petals senesced at the same time as those in the intact flower (Mor et al., 1980). These results seem to be in conflict with the proposed role of the gynoecium in controlling the senescence of the petals. Thus, the role of the gynoecium in the natural senescence of carnation, whether ethylene production in petals occurs independently of or is mediated by the gynoecium, remains to be revealed. In the present study, the excision method was improved and it was shown that the removal of the gynoecium markedly prolonged the vase life of the carnation flower. In addition, using the flower with gynoecium removed, it was demonstrated that the gynoecium is involved in the production of ethylene after application of ABA, IAA or ACC to the flower.

	Materials and methods

Top Abstract Introduction Materials and methods Results Discussion References

 
Plant materials
Flowers of carnation (Dianthus caryophyllus L. cv. Reiko) were harvested in the morning at the usual commercial stage of flowering at a nursery of a local grower. They were transported to the laboratory within the day of harvest. Stems were trimmed to 30 cm, placed with their cut end in distilled water and held for 1–2 d in a room set at 16 h light (40 µmol m^-2 s^-1), 8 h darkness at 23 °C. The flowers were used when their outermost petals were at right angles to the stem of the flower.

Treatment of flowers
Flowers at their full opening stage (day 0) were trimmed to 5 cm in stem length and placed with their stem end in 20 ml distilled water in 50 ml glass vials (one flower per vial). Usually ten flowers were used per treatment. To remove the gynoecium, the calyx of each flower was slit to its base and the petals carefully opened to expose the gynoecium. Then the gynoecium was snapped off at the junction of ovary and receptacle using fingers. Control flowers were treated in a similar manner, except that the gynoecium was left intact. The petals were then placed carefully in their original position, and the calyx was closed and held in place with a narrow strip of cellophane tape. After the treatment, the flowers were left under the conditions described above. Flowers were examined for the development of petal in-rolling symptoms and ethylene production in flowers was determined each day.

After removal of the gynoecium, the carnation flower was treated with ABA, IAA or ACC. The stem end was placed for 24 h from day 0 to day 1 in 20 ml of 0.1 mM ABA, 0.1 mM IAA or 0.1 mM ACC solution in 50 ml glass vials (one flower per vial). Then, the flowers were transferred to distilled water and left under the same conditions as above. The flowers were left for the observation of petal senescence, and ethylene production was measured at intervals.

For the treatment with ethylene, the flowers with gynoecium retained and removed were incubated in a 53.2 l glass chamber with ethylene at 10 µl l^-1 for 24 h at 23 °C under continuous white fluorescent light. The flowers were held in air for 1 h to allow the accumulated ethylene to diffuse from the tissues prior to the measurement of ethylene production.

Assay for ethylene production
Ethylene production by the whole flower was measured by enclosing the flowers in 350 ml glass containers (one flower per container) for 1 h at 23 °C (Kosugi et al., 1997), and that by the petals of ethylene-treated flowers was determined by enclosing the detached petals in 50 ml glass vials for 30 min. A 1 ml gas sample was taken by hypodermic syringe from the inside of the container through a rubber septum of a sampling port on the lid of the container, and analysed for ethylene with a gas chromatograph (Model 263-30, Hitachi, Japan) equipped with an alumina column and a flame ionization detector.

RNA gel blot analysis
Total RNA was isolated by the SDS-phenol method (Palmiter, 1974) from petals of the flowers with gynoecium retained and removed. Probes for mRNAs of DC-ACS1 (formerly CARACC3; Park et al., 1992; Jones and Woodson, 1999b) and DC-ACO1 (pSR120; Wang and Woodson, 1991; Jones and Woodson, 1999b) were constructed by PCR amplification with appropriate primers. The plasmids pBluescript-ACS1(OR) and pBluescript-ACO1(OR) (Kosugi et al., 2000) were used as templates. The probe for DC-ACS1 was 560 bp in size and contained the sequence ranging from 1 bp to 560 bp of the coding region of DC-ACS1 cDNA (Park et al., 1992). The probe for DC-ACO1 was 560 bp in size and contained the sequence ranging from 261 bp to 820 bp of the coding region of DC-ACO1 cDNA (Wang and Woodson, 1991). The cDNAs were labelled with ³²P-dCTP by random priming using Multiprime DNA labelling systems (Amersham Pharmacia Biotech) according to the manufacturer's instructions.

Thirty µg of total RNA was denatured at 65 °C for 15 min in 10 mM MOPS, pH 7.0, containing 2.2 M formaldehyde, and 50% (v/v) formamide. The denatured RNA was separated on a 1.0% agarose gel containing 2.2 M formaldehyde and transferred to nylon membranes (Hybond N⁺, Amersham Pharmacia Biotech). Hybridization was performed by the method of Church and Gilbert (Church and Gilbert, 1984). Blots were prehybridized at 68 °C for 1 h in 0.5 M NaHPO₄, pH 7.2 (1 M NaHPO₄, pH 7.2 stock is composed of 134 g of Na₂HPO₄.7H₂O and 4 ml of 85% H₃PO₄ l^-1), 7% (w/v) SDS, 1 mM EDTA, followed by hybridization at 68 °C for 18 h in the solution of the same composition but containing 5x10⁵ cpm ml^-1 of ³²P-labelled probes. Membranes were washed twice in 2xSSPE (1xSSPE is 0.15 M NaCl, 1.25 mM EDTA, 8.65 mM NaH₂PO₄, pH 7.4) and 0.1% SDS for 10 min at room temperature, followed by two washes for 30 min each at 68 °C. Hybridization signals were detected using an imaging plate and an image analyser (FLA 2000, Fuji Photo Film, Japan). Blots were used for multiple hybridization after stripping in boiling 0.1% SDS.

	Results

Top Abstract Introduction Materials and methods Results Discussion References

 
Effects of removal of gynoecium on the vase life of carnation
First, the method of excision in relation to the vase life of treated carnation flowers was compared. Some of the flowers with the gynoecium removed at the junction of ovary and receptacle using forceps or surgical scissors had a vase life longer than the control flowers, but others had a shorter vase life. The vase life of the flowers with part of the ovary tissue remaining on the receptacle was similar to that of the control flowers. When the gynoecia were snapped off with fingers to remove all the ovary tissue, all –Gyn flowers showed a longer vase life than the controls (Fig. 1). Figure 2 shows the profile of flowers 8 d after the procedure. Petals of the –Gyn flowers remained fully turgid up to day 16, whereas those of the control plants completely wilted with in-rolling of the petals beginning at day 6. After day 16 the –Gyn flowers showed desiccation and browning at the rim of petals. Corresponding to the in-rolling and wilting of the petals, the fresh weight of the control flowers started to decrease after day 6, and reached 59% of the initial weight at day 8. The fresh weight of –Gyn flowers decreased only slightly (<14% at day 8).

View larger version (105K): [in this window] [in a new window]   Fig. 1. Carnation flower before (Control) and after (–Gyn) removal of the gynoecium.

 

View larger version (129K): [in this window] [in a new window]   Fig. 2. Profile of flowers 8 d after the removal of gynoecia. Control flowers had gynoecia left intact.

 

Ethylene production in –Gyn flowers
Figure 3 shows the changes in ethylene production between the control and –Gyn flowers for 10 d after the gynoecium excision procedure. Ethylene was not produced in –Gyn flowers throughout the experimental period, whereas it was produced in the control flower from day 5 reaching a maximum at day 8, and declining rapidly thereafter.

View larger version (16K): [in this window] [in a new window]   Fig. 3. Ethylene production in the control and –Gyn flowers for 10 d after the removal of gynoecia. Ethylene released from flowers was measured. (•) Control flowers; () –Gyn flowers. Each point is the mean±SE obtained with 10 flowers.

 
In the petals of control flowers, mRNAs for DC-ACS1 encoding ACC synthase and for DC-ACO1 encoding ACC oxidase were detected 5 d after treatment corresponding to the time when ethylene production increased (Fig. 4). Neither the mRNA for DC-ACS1 nor the mRNA for DC-ACO1 was detected in the petals of –Gyn flowers during incubation.

View larger version (55K): [in this window] [in a new window]   Fig. 4. RNA gel blot analysis of the transcripts for ACC synthase and ACC oxidase. Total RNAs were prepared from the petals of the flowers with gynoecia retained (Control) and removed (–Gyn). Petals from three flowers were pooled for RNA extraction.

 

Ethylene production in –Gyn flowers treated with ABA, IAA, ACC or ethylene
ABA, IAA, ACC and ethylene stimulate ethylene production and wilting of petals in cut carnation flowers (Hall and Forsyth, 1967; Mayak and Dilley, 1976a, b; Nowak and Veen, 1982; Onoue et al., 2000; Ronen and Mayak, 1981; Sacalis and Nichols, 1980; Sacalis, 1989). The possible involvement of the gynoecium in the action of these chemicals using –Gyn flowers was investigated. ABA, IAA or ACC was applied from the cut end of stem of the flower by immersing the stem in a 0.1 mM solution of the chemical for 24 h (from day 0 to day 1). Administration of the chemicals to the flowers of the control and –Gyn flowers was secured by measuring the volume of test solutions (and water), taken up by the flowers during 24 h.

ABA did not induce ethylene production in the –Gyn flower during the incubation period of 8 d (Fig. 5), but enhanced the onset of ethylene production by 2 d in the control flowers (with the gynoecium retained). IAA showed a similar effect (Fig. 6). ACC caused some ethylene production and a slight wilting of petals in the –Gyn flower (Fig. 7), and caused a rapid induction of ethylene production and in-rolling of petals in the control flowers.

View larger version (18K): [in this window] [in a new window]   Fig. 5. Effect of removal of the gynoecium on the ethylene production in carnation flowers treated with ABA. Control and –Gyn flowers were treated with 0.1 mM ABA for 24 h (from day 0 to day 1). Control flowers were treated with water. Ethylene released from the flowers was measured every day until day 8. Each point is the mean±SE obtained with ten flowers. (•) Control flowers; () control flowers+ABA; () –Gyn flowers+ABA.

 

View larger version (18K): [in this window] [in a new window]   Fig. 6. Effect of removal of the gynoecium on the ethylene production in carnation flowers treated with IAA. Procedures were similar to those described in the legend for Fig. 5 except that 0.1 mM IAA was applied instead of 0.1 mM ABA. (•) Control flowers; () control flowers+IAA; () –Gyn flowers+IAA.

 

View larger version (18K): [in this window] [in a new window]   Fig. 7. Effect of removal of the gynoecium on the ethylene production in carnation flowers treated with ACC. Procedures were similar to those described in the legend for Fig. 5 except that 0.1 mM ACC was applied instead of 0.1 mM ABA. (•) Control flowers; () control flowers+ACC; () –Gyn flowers+ACC.

 
Finally, the control and –Gyn flowers were treated with 10 µl l^-1 ethylene for 24 h. Then ethylene production in detached petals was measured (Fig. 8). Exogenous ethylene induced ethylene production and wilting of petals in both –Gyn and control flowers.

View larger version (31K): [in this window] [in a new window]   Fig. 8. Effect of removal of the gynoecium on the ethylene production in petals of carnation flowers treated with ethylene. Control or –Gyn flowers were treated with 10 µl l-1 ethylene for 24 h (from day 0 to day 1). After flowers were held in air for 1 h, ethylene production in the detached petals was measured. Each point is the mean±SE obtained with flowers, each with five petals. ND, not detectable.

 

	Discussion

Top Abstract Introduction Materials and methods Results Discussion References

 
When the gynoecium was removed from the carnation flower at the junction of ovary and receptacle using forceps or surgical scissors, a reproducible extension of vase life was not obtained. However, after the gynoecia were snapped off with fingers to remove all traces of ovary tissue, all –Gyn flowers showed a vase life longer than the control flowers. The –Gyn flowers had a vase life about twice that of the control flowers, before undergoing desiccation and turning brown at the margins of their petals. This phenomenon is typical for the ethylene-independent senescence in carnation flowers. Actually, no ethylene production was detected in –Gyn flowers throughout the period of incubation. The lack of ethylene production in the flowers was accompanied by the absence of transcripts for ACC synthase and ACC oxidase, indicating no induction of transcription of the genes in the petals of –Gyn flowers. These findings are in agreement with the results of ten Have and Woltering, who showed that when petals were detached from carnation flowers before the autocatalytic ethylene production had started, they exhibited a suppressed ethylene production and a prolonged petal life (ten Have and Woltering, 1997).

Interestingly, the onset of ethylene production was not suppressed by incomplete removal of gynoecium in the carnation flower, in which a part of the ovary tissue remained on the receptacle. It is probable that the remaining tissue still functions as an ovary, having its influence on timing of the initiation of senescence. In a previous study, gynoecia were excised from carnation flowers using forceps (Sacalis and Lee, 1987). The procedure might have caused incomplete removal of ovary tissue which could obscure the effect of gynoecium excision on the extension of vase life. The present results have demonstrated a decisive role of the gynoecium in natural senescence of the carnation flower.

Exogenous ABA or IAA accelerated the senescence of the cut carnation flower with the intact gynoecium through the stimulation of ethylene biosynthesis (Figs 5, 6). In the –Gyn flowers, however, ABA or IAA treatment induced neither ethylene production nor wilting of the petals, indicating that these phenomena depend on the action of ABA or IAA on the gynoecium. The effect of IAA treatment presented here was similar to that reported previously (Sacalis, 1989). ACC application induced ethylene production in the –Gyn flowers, but the magnitude of ethylene production was far lower than that in the control flowers. This indicates that ethylene production induced by applied ACC was also mediated by the gynoecium. Exogenous ethylene induced ethylene production and wilting in the petals of –Gyn flowers, indicating the direct action of ethylene on the petals of carnation flowers. Thus, it is considered that the exogenous ACC induced ethylene production in the petals of the control flowers after its conversion to ethylene in the gynoecium.

In conclusion, the present study showed that only exogenous ethylene, not ACC, ABA or IAA, induced the ethylene production in –Gyn flowers. These observations indicate that the gynoecium plays a decisive role in the induction of ethylene production in petals of carnation, which undergoes natural senescence or senescence caused by application with ABA, IAA or ACC, as previously shown in the pollination-induced senescence (Jones and Woodson, 1999a) and in the natural senescence (ten Have and Woltering, 1997) of the flower. Ethylene has been revealed as the inter-organ signal in natural and in pollination-induced senescing flowers of carnation (ten Have and Woltering, 1997; Jones and Woodson, 1999a). The present results confirmed its role as the inter-organ signal in carnation flower senescence.

It has been shown previously that ABA content increases transiently in the gynoecium and petals before the onset of ethylene production during natural senescence of the carnation flower and a chemical that prevents the increase in ABA content suppresses ethylene production resulting in the delay of wilting of the flower (Onoue et al., 2000). Therefore, combining the present and previous findings, the following model is proposed for the regulation of ethylene production in carnation flowers during natural senescence. First, some endogenous factors, like ABA, IAA or other unidentified factor(s), initiate ethylene production in the gynoecium. Then the ethylene produced acts as a diffusible signal and is perceived by the petals. As a result of ethylene perception, the genes for ACC synthase and ACC oxidase are expressed in the petals, resulting in autocatalytic ethylene burst and rapid wilting of the petals.

	Notes

 
¹ To whom correspondence should be addressed. Fax: +81 22 717 8834. E-mail: ssatoh{at}bios.tohoku.ac.jp

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