Postharvest conservation of lisianthus inflorescences with bioregulators

ABSTRACT

Cut flowers are known for their beauty and variety of colors and shapes. However, they quickly lose their commercial value after harvest due to the intensity of physiological processes that result in senescence. This work aimed to evaluate the effects of bioregulators on the postharvest longevity of lisianthus flowers (Eustoma grandiflorum cv. Flare Deep Rose) by applying a pulsing solution for 24 hours. The treatments consisted of 70 µM of 6-benzylaminopurine (BAP); 5 µM of gibberellic acid (GA3); 10 µM of abscisic acid (ABA); and deionized water as control. Turgidity, floral development, total and reducing sugar contents, respiration rate, colorimetry, anthocyanin contents, phenolic compound contents, and phenylalanine ammonia-lyase (PAL) enzyme activity were evaluated. BAP resulted in higher PAL enzyme activity and greater accumulation of anthocyanins when compared to the other treatments. The treatment GA3 resulted in the highest increase in respiration rate during storage, causing a larger number of inflorescences with wilting and senescence symptoms, reducing postharvest quality. The treatment ABA resulted in greater turgidity and floral opening, delayed senescence, and maintained respiration rate due to the greater total sugar contents on the fourth and 12th days. The application of ABA contributes to the maintenance of inflorescence quality for lisianthus at postharvest, but it reduces anthocyanin contents, providing petals with lighter colors.

Key words
Eustoma grandiflorum ; cut flower; abscisic acid; gibberellic acid; 6-benzylaminopurine

Introduction

Lisianthus (Eustoma grandiflorum) is an important cut flower widely used in floral arrangements due to the beauty of its inflorescences and variety of colors (^{Chuang and Chang 2013}). The inflorescences are large with long stems, which provides easy handling. However, flowers are highly perishability products due to the ephemeral nature of their tissues; physiological processes occur intensively and, thus, they lose their commercial value in a short period after harvest (^{Nowak and Rudnicki 1990}).

The stem cutting triggers senescence processes that are irreversible in flowers accelerates water loss and loss of color and brightness of petals, which are the main causes of product depreciation (^{Dias 2016}, ^{Ma et al. 2018}). Color loss is detrimental to the decorative quality of floral stems, resulting in reduced commercial value, and it is related to reductions in accumulation of pigments during senescence, such as anthocyanins and carotenoids (^{Mattiuz et al. 2010}, ^{Park et al. 2021}).

Senescence is a process highly controlled by bioregulators, as well as the other stages of plant development; thus, these compounds can be effective when used as preservative solutions at post-harvest of flowers (^{Favero et al. 2020}). The bioregulator 6-benzylaminopurine (BAP) can induce resistance to biotic and abiotic stress because it can reduce free radical activity in the plant (^{Kamran et al. 2021}). Thus, good results have been obtained in pulsing treatments with BAP delaying senescence in chrysanthemum, and lisianthus (^{Kaur and Singh 2015}, ^{Musembi et al. 2013}).

Gibberellic acid (GA3) has shown good results for postharvest maintenance of several inflorescences, such as anthurium and lisianthus. It can maintain the stability of cell membranes and, thus, reduce senescence of cut flowers (^{Emongor 2004}, ^{Musembi et al. 2013}, ^{Simões et al. 2018}). Postharvest application of abscisic acid (ABA) can induce tolerance to water stress in petunia and pelargonium, and reduce leaf chlorosis in lily (Lilium oriental cv. Sorbonne), which depreciates the stems (^{Runkle et al. 2007}, ^{Geng et al. 2015}).

Carbohydrate balance is an important factor in flower postharvest; its effects on quality maintenance and inhibition of senescence symptoms are attributed to the importance of carbohydrates as substrate for respiration, structural function, and maintenance of osmotic pressure (^{Halevy and Mayak 1979}, ^{Gupta and Dubey 2018}). Reductions in carbohydrate levels are connected to the consumption of plant energy reserves; the application of postharvest treatments can alter these levels and assist in maintaining the quality of inflorescences after cutting (^{Cavasini et al. 2018}).

Considering this information and the need for using technologies that reduce the impact of senescence symptoms, which depreciate inflorescences and, consequently, generate large postharvest losses and economic losses to growers, the use of preservative solutions is an effective alternative for maintaining the quality and reducing the effects of senescence (^{Nowak and Rudnicki 1990}, ^{Gupta and Dubey 2018}).

The application of bioregulators favors anthocyanin biosynthesis and, as a result, preserves the color of petals for a longer storage period after harvest. Thus, this work aimed to evaluate changes, caused by bioregulators (BAP, GA3, and ABA), in petals of open flowers of lisianthus (Eustoma grandiflorum cv. Flare Deep Rose).

METHODS

Stems of lisianthus (Eustoma grandiflorum cv. Flare Deep Rose) were harvested in an area of the Monalisa Flores company, in Paranapanema, SP, Brazil (23º23’19”S, 48º43’22”W, and 610 m of altitude). The established harvest point was two or three open flowers per inflorescence; the flowers were harvested, packed, and transported, under refrigeration, to the Postharvest Physiology and Biochemistry Laboratory of the Universidade de São Paulo (Piracicaba, SP, Brazil). In the laboratory, the stems were standardized at 50 cm in length, and leaves were removed from the first 15 cm of the lower part of the stem. Then, they were randomized for the application of treatments.

The bioregulators BAP (CAS 1214-39-7), GA₃ (CAS 77-06-5), and ABA (CAS 21293-29-8) were obtained from the Sigma-Aldrich^® company.

The treatments consisted of bioregulators pulsing solutions for 24 hours at the rates 0 μM (deionized water; control), 70 μM BAP, 5 μM GA3, and 10 μM ABA, which were determined through preliminary tests. After applying the treatments, the stems were kept in acrylic pots containing 500 mL of a solution of deionized water and sodium dichloroisocyanurate at the concentration of 0.25%. The pots were stored under air temperature of 20°C and relative air humidity of 80 ± 2%.

A completely randomized experimental design was used, in a factorial arrangement, with four bioregulators pulsing solutions and four days of evaluation. Four replications of three stems were used, and evaluations were carried out every four days (zero, four, eight, and 12 days). Petal samples were collected, frozen in liquid nitrogen, freeze-dried, ground, and stored at -80°C for further biochemical analysis. Freeze-drying of the samples was carried out using a device operating at temperature of -55 ± 5°C and pressure of 1×10^-2 to 3×10^-2 mbar (0.001 to 0.003 kPa) (Liotop L108, São Paulo, Brazil), and then milled in an analytical mill (A11 Basic; IKA® Werke GmbH & Co. KG, Staufen, Germany).

According to visual analysis, buds that were at suitable developmental stage for full opening and flowers that were open at the time of harvest were quantified. Grades were assigned as follows:

• Bud;

• Early anthesis;

• Full opening;

• Senescent flowers.

Using the same structures, the following grades were assigned to determine the turgidity:

• Wilted;

• Slightly wilted;

• Turgid.

Total and reducing sugar contents–25 mg of petals, 25 mg of leaves, and 250 mg of stem–were used for sugar extraction. Each sample was immersed in 5 mL of 80% ethanol, stirred, and placed in a water bath at 60°C for 30 minutes. The samples were then centrifuged at 6,440 × g (Jouan BR4i, Paris, France) for 5 minutes; the supernatant was collected, transferred to 15-mL tubes, and then 5-mL ethanol was added, and the procedure was repeated. Then, 80% ethanol was added to complete the volume to a final extract of 10 mL.

Total sugars were quantified according to the phenol-sulfuric methodology of ^{Dubois et al. (1956)}, and reducing sugars were quantified according to the Somogyi-Nelson methodology (^{Nelson 1944}). Readings were carried out in a spectrophotometer (Biochrom Libra S22 UV-Vis, Cambrige, United Kingdom) at 490 nm for total sugars and 540 nm for reducing sugars. The results were expressed as mg glucose per g of dry matter.

Respiration rate in inflorescences was monitored by gas chromatography. The CO₂ evolution was determined by placing three stems in acrylic vases containing 500 mL of water and maintaining them in an airtight container for 1 hour. Gas samples of 0.5 mL were collected from each container and injected into a gas chromatograph device (Thermofinigan, model Trace GC Ultra) equipped with a flame ionization detector (FID) and a stainless-steel column of 1/8” and 4-m long, prepared with Porapak N 50/80. The temperatures of the column, injector, and detector were 110, 140, and 200°C, respectively; hydrogen was used as carrier gas at a flow rate of 25 mL min^-1. Respiration rate was expressed as mL of CO₂ per kg of fresh matter per hour (mL CO₂.kg^-1.h^-1).

About colorimetry, readings were carried out using a Minolta CR-400 colorimeter in six flowers per replication. Five readings were carried out on the inner side of each flower. The results were calculated based on the parameters L, a*, and b* and expressed as luminosity (light-to-dark range) and chromaticity (color intensity). Chromaticity was calculated according to the Eq. 1 (^{Gonnet 1998}):

$$\mathrm{C}={({\mathrm{a}}^{\ast 2}+{\mathrm{b}}^{\ast 2})}^{0.5}$$ (1)

Regarding total anthocyanin contents, the method described by ^{Lee and Francis (1972)}, with modifications, was used. Anthocyanin was extracted from a dry sample of 100 mg using 1% methanol-acidified HCl, and the extract was kept at 4°C for 12 hours. Absorbance readings were carried out at 510 nm (Biochrom Libra S22 UV-Vis, Cambrige, United Kingdom). The results were expressed in mg per g of dry matter.

Phenylalanine ammonia-lyase (PAL) enzyme activity (EC 4.3.1.5) was evaluated using the methodology proposed by ^{Peixoto et al. (1999)}, with modifications. PAL was extracted from a dry sample of 100 mg and homogenized in 10 mL of cooled sodium borate buffer (0.1 mol.^L-1, pH 8.8) containing polyvinylpyrrolidone 10 (50 g.L^-1) and β-mercaptoethanol (0.0002 mol.L^-1). The solution was centrifuged at 6,440 × g for 15 min at 4°C (Jouan BR4i, Paris, France), and the supernatant was collected, resulting in an enzyme extract. The whole process was conducted at 17°C. The reaction was carried out at room temperature (25 ± 1°C); 1 mL of the enzyme extract was incubated at 37°C for 1 hour in 1 mL sodium borate buffer (0.2 mol.L^-1, pH 8.8) and 1 mL L-phenylalanine (0.1 mol.L^-1). The reactions were stopped by the addition of 0.1 mL of hydrochloric acid (6 mol.L^-1), and the amount of t-cinnamic acid was estimated in triplicate by measuring the absorbance at 290 nm. The data were expressed as µkatals per g of dry matter.

Total phenolic compounds were quantified using the Folin-Ciocalteu reagent, according to ^{Randhir et al. (2002)}, with adaptations: 10 mL of 80% methanol was added to 0.1 g of freeze-dried lyophilized sample; the samples were stored at room temperature, protected from light, for 2 hours. The extracts were then separated, and the supernatant was collected. The reaction was carried out by adding 0.2 mL of the sample extract, 1.5 mL of distilled water, 0.1 mL of Follin-Ciocauteu reagent, and 0.2 mL of 20% calcium carbonate. It was incubated for 2 hours in the dark. Spectrophotometric readings were then carried out at 765 nm, and the results were expressed as mg of gallic acid equivalents (standard) per g of petal dry matter (EAG g.kg.^L-1).

Longevity was determined when the treatments presented 50% of wilted inflorescences and/or senescence symptoms such as wilting, peduncle curvature, abscission, and petal darkening. Statistical analyses were performed using the program R 3.2.5. The results were subjected to analysis of variance and expressed as mean ± standard error. The normality and homogeneity of the data were verified by the Shapiro-Wilk and Bartlett’s tests, respectively. Statistical differences between means were calculated by the Tukey’s test (p = 0.05).

RESULTS AND DISCUSSION

The initial floral development was advanced by the application of GA₃ and ABA bioregulators. Despite presenting the highest percentage of senescent flowers on the fourth day of storage (10%), the treatment ABA resulted in less visual quality loss after 12 days, maintaining 63% of the flowers open. Contrastingly, the application of GA₃ and BAP maintained the visual quality only during the first eight days of storage, and BAP did not differ from the control. GA₃ resulted in accelerated anthesis at four days and in a higher percentage of senescent flowers after 12 days of storage when compared to the control and the other bioregulators, resulting in discarding of stems on the 12th day (Figs. 1a, 1b, and 2).

Figure 1
(a) Development of inflorescences of lisianthus (Eustoma grandiflorum cv. Flare Deep Rose); (b) percentage of lisianthus inflorescences with wilting symptoms. Inflorescences treated with bioregulators and stored for 12 days at 20°C and 80% relative air humidity.

The maintenance of turgidity using ABA may be connected to its ability to regulate the water balance by controlling stomatal opening, which reduces transpiration and, consequently, the intense water loss that occurs after harvest (^{Halevy et al. 1974}, ^{Shimizu-Yumoto et al. 2010}).

Figure 2
Monitoring of opening of lisianthus flowers (Eustoma grandiflorum cv. Flare Deep Rose) treated with bioregulators and stored for 12 days at 20°C and 80% relative air humidity.

In addition to present a larger number of senescent flowers, the treatment ABA resulted in a greater percentage of flowers with pronounced wilting symptoms, more than the control on the fourth day of storage. However, this trend did not continue; at the end of 12 days of storage, ABA resulted in the lowest wilting percentages. The application of GA3 markedly accelerated the loss of visual quality of flowers in the last four days of storage; in this period, there was a 45% increase in wilted flowers when compared to day 8, which is equivalent to that one found for the control. BAP presented no efficacy in controlling the flower wilting, showing no differences from the control (Figs. 1a and 1b).

Sugar contents are involved with flower turgescence. Reducing osmotic potential through carbohydrate accumulation can increase the influx of water into flowers, maintaining the pressure potential, which is connected to cell expansion and floral opening. Moreover, flowers are the main carbohydrate drain during the inflorescence development (^{Waithaka et al. 2001}). The inflorescences treated with ABA (310.2 mg.g^-1) and GA3 (322.2 mg.g-1) showed the highest contents of total sugars at four days of storage. The treatment ABA showed higher total sugar contents (305.9 mg.g^-1) than the other treatments at 12 days of storage. The treatment BAP did not show significant difference from the control (Fig. 3).

Unlike the control, which showed a constant reduction in total sugar contents, the application of ABA and GA₃ resulted in the highest contents in petals throughout the storage period. However, the visual quality result was different between these two treatments. The stems with GA₃ did not maintain the commercial quality by the 12th day of storage. In contrast to other bioregulators, the application of BAP resulted in reductions in total sugars over the first four days of storage and did not differ from the control on any day of analysis (Fig. 3a).

The metabolism of sugars was affected by the application of bioregulators, with decreases in reducing sugar contents in all treatments during storage. However, the treatment ABA had the highest contents in the first eight days of evaluation. GA₃ showed the highest decrease in reducing sugar contents on the fourth day, but, on the eighth day of evaluation, it was the only treatment that showed increases in these contents throughout the storage period. BAP had the same result as the control, with less effect on reducing sugars when compared to the other bioregulators (Fig. 3b).

ABA is a bioregulator connected to the plant physiological response to stress, conferring adaptation to the environment. It induces the formation of reactive oxygen species that act as abiotic stress signaling molecules, which is important for the adjustment of metabolism, causing stomatal closure. Reactive oxygen species cause cell damages. However, these damages are reduced when cells have high energy reserves (^{Mittler and Blumwald 2015}, ^{Choudhury et al. 2017}).

The total and reducing sugars in stems treated with ABA in the first days of storage (Fig. 3) may have reduced damages caused by senescence. Soluble sugars can have antioxidant action, protecting the cell against oxidative stress (^{Keunen et al. 2013}, ^{Peshev et al. 2013}). In addition, ABA can increase translocation of endogenous carbohydrates to flowers and buds, making the osmotic potential more negative and intensifying water uptake, which aids in maintaining turgidity and opening of buds (^{Emongor 2004}, ^{Shimizu-Yumoto et al. 2010}).

Figure 3
(a) Total sugars and (b) reducing sugars in petals of lisianthus flowers (Eustoma grandiflorum cv. Flare Deep Rose) treated with bioregulators and stored for 12 days at 20°C and 80% relative air humidity.

On the fourth day of storage, the treatment ABA showed an advance in the opening of the floral buds, with an increase in total sugar contents (Fig. 3a). Floral opening is a result of the development and expansion of petal cells, which are processes regulated by the presence of soluble carbohydrates in the tissues (^{Pun and Ichimura 2003}, ^{In et al. 2006}).

The increase in sugars caused by application of ABA in cut flowers may reflect the ability of the bioregulator to interfere with the metabolism of these compounds present in petals and leaves (^{Geng et al. 2015}). Studies have reported the ability of bioregulators to affect the activity of the enzyme invertase, which is responsible for the hydrolysis of sucrose into hexoses (^{Trouverie et al. 2004}, ^{Pan et al. 2006}). Sucrose is translocated from leaves through the phloem to other tissues, in which it is hydrolyzed by the invertase, and ABA can increase the activity of this enzyme (^{Tauzin and Giardina 2014}).

The application of GA3 also affected sugar contents in lisianthus petals. There were an increase in total soluble sugar contents and a decrease in reducing sugars on the fourth day of storage (Fig. 3). These results may be connected to the increase in respiration of plants in this treatment throughout the 12 days of storage (Fig. 3). Thus, unlike ABA, GA3 probably did not accumulate enough sugars to supply the respiration rate, consuming sugar reserves of petals and inducing senescence.

Accelerated metabolism may be involved in the anticipation of floral bud opening in the treatment GA₃, as the transport of soluble sugars from leaves to petals was probably promoted by exogenous GA₃, inducing carbohydrate loading in the flowers (^{Aloni et al. 1986}, ^{Murcia et al. 2018}). Gibberellins can increase hydrolysis of starch, fructans, and sucrose into glucose and fructose, reducing the osmotic potential of tissues, facilitating the influx of water to upper parts of the stem (^{Emongor 2004}). Thus, floral opening occurred as a response to the maintenance of turgidity caused by GA₃ until the eighth day of storage (Figs. 1b, 2, and 4).

Figure 4
Respiration rate of inflorescences of lisianthus (Eustoma grandiflorum cv. Flare Deep Rose) treated with bioregulators and stored for 12 days at 20°C and 80% relative air humidity*.

The effect of sugars on the senescence of different plant organs has not yet been fully elucidated, but it is known that carbohydrates can delay senescence when they are present at high levels in inflorescences, and that low levels of these sugars in petals and leaves can induce senescence (^{Wojciechowska et al. 2018}).

The application of bioregulators showed different effects on the color chromaticity and luminosity of lisianthus petals (Figs. 5a and 5b). The treatment BAP showed a significant difference in anthocyanin contents only on the 12th day of storage (Fig. 5c).

The BAP treatment showed higher PAL activity, although the activity of this enzyme decreased over time in all treatments (Fig. 5d). Exogenous BAP can induce the expression of genes linked to the anthocyanin biosynthesis pathway, causing accumulation of this pigment (^{Das et al. 2012}). However, there was no significant difference in luminosity and chromaticity between BAP and the control (Figs. 5a and 5b).

There was less synthesis of anthocyanins in the treatment ABA (Fig. 5c), which affected color parameters, causing an increase in luminosity and reduction in chromaticity (Figs. 5a and 5b). The higher the luminosity, the lighter the analyzed surface. Chromaticity is a parameter related to color purity. Thus, the higher the chromaticity, the more intense the color of petals, which can be connected to anthocyanin contents present in the lisianthus petals (^{Uddin et al. 2001}).

In this context, the reduction in anthocyanin contents resulted in a lighter hue in petals in inflorescences treated with ABA. This result was probably due to the deviation of the biosynthesis route, with formation of other types of flavonoids instead of anthocyanins. Flavonoids are among the main non-enzymatic antioxidants produced during oxidative stress (^{Baskar et al. 2018}), whereas ABA induces formation of reactive species for signaling and plant adaptation to this stress (^{Choudhury et al. 2017}).

Figure 5
(a) Chromaticity; (b) luminosity; (c) anthocyanin contents; and (d) phenylalanine ammonia-lyase activity in petals of lisianthus (Eustoma grandiflorum cv. Flare Deep Rose) treated with bioregulators and stored for 12 days at 20°C and 80% relative air humidity*.

These results explain the reduction in anthocyanin accumulation and the greater formation of total phenolics in the first days of storage in the treatment ABA (Fig. 6), with no effect on PAL enzyme activity (Fig. 5d). In addition, plants exposed to oxidative stress increase endogenous ABA and may form proanthocyanidins at the expense of anthocyanins, resulting in flowers with lighter petals. Proanthocyanidins are involved in induction of tolerance and expression of stress response genes (^{Luo et al. 2016}, ^{Li et al. 2019}).

The postharvest application of GA₃ in lisianthus increased anthocyanin contents until the fourth day and probably accelerated the development of inflorescences, leading to senescence (Figs. 1 and 5c). The same result was found for chrysanthemums cultivars (Flippo, Recital, and Bronze Repim); the application of increasing rates of GA₃ in a preservative solution accelerated the senescence of leaves and flowers, reducing the vase life of inflorescences (^{Brackmann et al. 2005}).

Although PAL showed the lowest activity in the treatment GA₃, there was an increase in anthocyanin contents in the first days of storage (Fig. 5c and 5d). However, total phenolic compounds did not change (Fig. 6), indicating that the treatment induced formation of anthocyanins to the detriment of other phenolics in the first days of storage. Gibberellins act indirectly in the expression of genes linked to the anthocyanin biosynthesis pathway; however, the expression of these genes begins to decrease after anthesis (^{Weiss et al. 1995}).

Figure 6
Total phenolic contents of lisianthus (Eustoma grandiflorum cv. Flare Deep Rose) treated with bioregulators and stored for 12 days at 20°C and 80% relative air humidity*.

CONCLUSION

The application of ABA during postharvest of lisianthus contributes to maintenance of inflorescence quality. However, it reduces anthocyanin contents, resulting in petals with lighter colors. The application of GA₃ accelerates the senescence, reducing postharvest quality of inflorescences.

ACKNOWLEDGMENTS

We acknowledge Cooperflora cooperative members for providing the plant material for this work.

• How to cite: Guzzo, E. C., Vendramim, J. D., Corrêa, O. M. B. and Lourenção, A. L. (2023). Calaboni, C., Kluge, R. A., Preczenhak, A. P. and Mattiuz, F. M. (2023). Postharvest conservation of lisianthus inflorescences with bioregulators. Bragantia, 82, e20220207. https://doi. org/10.1590/1678-4499.20220207

• FUNDING

  Fundação de Amparo à Pesquisa do Estado de São Paulo
  [https://doi.org/10.13039/501100001807]
  Grant no. 2019/01156-0
  Coordenação de Aperfeiçoamento de Pessoal de Nível Superior
  [https://doi.org/10.13039/501100002322]
  Finance Code 001.

DATA AVAILABILITY STATEMENT

All dataset were generated and analyzed in the current study.

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Edited by

Section Editor: Juliana Sanches

Publication Dates

• Publication in this collection
  22 May 2023
• Date of issue
  2023

History

• Received
  17 Oct 2022
• Accepted
  09 Mar 2023