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Review

The Potential Postharvest Treatments to Delay Flower Senescence and Improve Botrytis Resistance in Cut Peony Flowers

by
Toan Nguyen
1,2 and
Suong Ha
1,2,*
1
Institute of Research and Development, Duy Tan University, Da Nang 550000, Vietnam
2
School of Engineering & Technology, Duy Tan University, Da Nang 550000, Vietnam
*
Author to whom correspondence should be addressed.
Horticulturae 2024, 10(12), 1352; https://doi.org/10.3390/horticulturae10121352
Submission received: 25 October 2024 / Revised: 3 December 2024 / Accepted: 11 December 2024 / Published: 16 December 2024
(This article belongs to the Special Issue Pre/Post-harvest Treatments to Improve Quality and Longevity of Cut Flowers)
Versions Notes

Abstract

:
The herbaceous peony (Paeonia lactiflora Pall.) flower is one of the most important ornamental plants in current international flower markets and is widely used for festive occasions because of its bright colors, large flowers, pleasant scent, and plump flower shape. However, the cut peony flowers have a relatively short postharvest life in vases compared to other flower species. The short vase life and susceptibility to gray mold disease caused by Botrytis cinerea significantly influence the commercial value of cut peonies. Here, we reviewed the main factors for postharvest quality reduction in cut peony flowers and the various postharvest treatments aiming for vase life extension and enhancing gray mold disease resistance in cut peonies. Chemical treatments, including ethylene inhibitors, antimicrobial agents, and other preservatives, have been assessed for their effectiveness in improving vase life. Moreover, physical treatments, such as temperature management, controlled atmosphere storage, and dry storage methods, were also discussed for their effectiveness in delaying flower senescence and reducing gray mold disease infection. The review also highlighted the importance of cultivar-specific responses to gray mold disease, ethylene, and water stress, which is important for the development of new effective and specific postharvest practices to improve the vase life of cut peonies.

1. Introduction

Herbaceous peony (Paeonia lactiflora Pall.) flowers hold an important place in the ornamental plant industry because of their elegant forms and unique colors [1,2]. More than 3000 cultivars and 30 species of peonies are found and developed nowadays, and more than 20 countries are actively involved in the production and trade of cut peony flowers [1,3,4]. Peony flowers are popular in European countries and North America. The Netherlands has a major share of cut peony production and wholesale distribution in the European flower market. North American countries (the United States and Canada) also exhibit substantial demand and are one of the important cut peony flower markets. In addition, Asia is emerging as a growing flower market due to increasing demand in Japan and China [2,5,6]. The long-term breeding strategies and the garden variety of peony selections have resulted in various flower colors and types [2,5]. Among peony flower types, the double type is the most popular as cut flowers because of their longer vase life compared to other types (single, anemone, bomb, and semi-double) [2,7].
Flower shape and color, uprightness, and fragrance are the main selection criteria of customers for cut peony flowers [2]. However, short postharvest life and seasonal production of cut peony flowers are the main factors limiting customer satisfaction and availability [8,9]. The longevity of cut peonies is influenced by various factors such as genetic factors, preharvest conditions, harvested stage, and postharvest handling [2,5,10]. The main senescence symptoms in cut peony flowers are botrytis blight, flaccid bud and/or flaccid flower, stem bending (bent of flower neck), petal abscission, petal wilting, leaf desiccation, and yellow and/or brown leaves [11]. Cut peony flowers rely on stored carbohydrate content accumulated from mother plants reserves for flower freshness and opening [12]. Moreover, postharvest water relations, hormone balance, lignin content, and oxidative stresses influence cut peony flowers’ quality and vase life [13,14,15]. Water balance determined by water uptake, transpiration, and evaporation is an important factor for the vase life of cut flowers [16]. The negative water balance (more water loss than water uptake) due to unsuitable environmental conditions or blockage caused by microbial growth in the vase solution causes early wilting of flowers and leaves, improper opening, bent neck (bending of pedicel), and ultimate death of cut flowers due to lack of water [16]. Low lignin content in flower stems also contributes to the incidence rate of the bent neck in cut flowers, reducing the quality and ornamental value of cut flowers [14,15]. Stem straightness is considered one of the most important criteria for the postharvest quality of cut peonies [11]. The flower stems of many peony cultivars readily break or bend and cannot support the fresh weight of cut flowers due to low stem strength [14,15]. Previous studies showed that lignin content provides mechanical support to cut peony flower stems, and lignin content is correlated well with the stem strength and incidence rate of the bending stem [13,14]. Exposure to ethylene decreased the vase life and ornamental values of the ethylene-sensitive peony flowers [16,17]. Tree peony flowers exposed to ethylene (10 µL L−1 for 6 h) exhibited premature wilting of petals and petal discoloration [17,18]. In flowers, ethylene-induced senescence can lead to a reduction in longevity. In peony flowers, for instance, ethylene-sensitive cultivars showed increased susceptibility to gray mold disease. The application of ethylene inhibitors can prevent ethylene damage and prolong the longevity of flowers. Ethylene inhibitors can block ethylene production and receptors, preventing the downstream signaling of ethylene and delaying senescence and disease progression [16,17,18].
In addition to the above factors, peony flowers are susceptible to a wide range of fungal diseases, including powdery mildew, phytophthora, leaf spots, root rots, and botrytis (gray mold or botrytis light) [2,19]. Among these fungal pathogens, peony botrytis is considered the most economically significant postharvest disease of peony flowers. Several species of Botrytis can infect peony flowers, including Botrytis cinerea (B. cinerea), B. pseudocinerea, B. euroameriana, and B. paeoniae [19]. Generally, B. cinerea and B. pseudocinerea are responsible for gray mold and late blight, while B. paeoniae are associated with early blight and are known as specific to peony flowers [19]. Botrytis can develop and infect all plant organs, such as flowers, young shoots, leaves, roots, and stems. Among them, flowers are the most susceptible organ to botrytis infection and can be infected from the earliest stages of flower bud development [19,20]. Botrytis infection in buds causes them to fail to open or partially open. Individual peony petals may also become infected with botrytis, initiating as small brown lesions before developing and spreading to the entire petal, eventually brown, dry symptom appearance [18,20]. Severe gray mold infection can develop rapidly down to the flower receptacle and pedicel, and after time, the whole flower becomes macerated and falls away [20]. If the disease is not controlled effectively in the fields, the peony crop losses from botrytis infection are approximately 20% [20]. Postharvest losses of cut peonies are also a serious problem where disease symptoms develop on flower buds or blooms during transport and storage, in retail flower shops, or in customers’ homes. Wholesale and retail market rejection due to botrytis infection is one of the main commercial issues for peony flower producers [11]. Therefore, to maintain and promote the development of the cut peony flower industry, mechanisms/postharvest treatments need to be improved and developed that are capable of extending vase life while maintaining high cut flower quality during the vase period. To gain this, the mechanisms of the flower senescence and the factors that affect the postharvest quality and vase life of cut peony flowers should be deeply understood, and based on this knowledge, the high-efficiency postharvest treatments will be developed for each cut peony cultivar group.
This review aims to provide an overview of the postharvest treatments to improve postharvest quality, vase life, and botrytis resistance in cut peony flowers. We also discussed the advantages and disadvantages of the postharvest treatment method applied to cut peony flowers. A better understanding of these postharvest treatments will be useful to improve higher postharvest quality and longer vase life of cut peonies and speed up the development of new eco-friendly and suitable preservative solutions for peonies. Additionally, we propose an effective method for reducing botrytis infection and increasing the longevity of cut peonies based on the classification of cut peonies susceptible to ethylene, water stress, and botrytis.

2. Chemical Treatments Improve Longevity and Quality of Peony Flowers

2.1. Application of Ethylene Antagonists Affected Flower Quality

The perception of ethylene by ethylene receptors is a requirement to initiate and maintain ethylene responses during senescence. The perception of ethylene by the receptors inactivates Constitutive Triple Response1, a Raf-like MAPK kinase, thereby relieving the suppression of the downstream signaling elements. In the nucleus, Ethylene Insensitive 2 (EIN2) activates the transcription factor EIN3 or EIN3-like 1 to initiate transcriptional responses to ethylene. The progression of senescence induced by ethylene can be delayed by inhibiting the binding and synthesis of ethylene in flowers and fruits [21,22,23,24,25]. Ethylene inhibitors such as aminoethoxyvinylglycine (AVG), silver thiosulfate (STS), and 1-methylcyclopropene (1-MCP) have been applied in the cut flower industry as postharvest treatments for reducing ethylene damage and improving the vase life of various cut flowers [21,22,23,24,25]. Previous studies have shown that ethylene sensitivity differed among peony cultivars [17,18]. Ethylene exposure decreased the vase life and promoted flower opening and senescence in cut peonies ‘Luo Yang Hong’ but did not influence the postharvest quality of the ‘Xue Ying Tao Hua’ flowers [17,18]. The response of cut peony flowers to ethylene inhibitors treatment was also different among cultivars. Hoffman et al. [26] showed that AVG, STS, and 1-MCP applications negatively influenced the postharvest quality and vase life of two cut peony flowers, ‘Karl Rosenfield’ and ‘Sarah Bernhardt’. AVG treatment and storage duration failed to affect the petal openness level, fresh weight, chroma, and petal color hue angle of cut peony ‘Sarah Bernhardt’ flowers. In addition, AVG treatment increased the incidence rate of gray mold disease and reduced the vase life of cut peony flowers during storage time [26]. STS as a pre-storage treatment was ineffective in increasing postharvest life and total quality of cut flowers compared to the control group [26]. Whereas the effectiveness of 1-MCP on cut peony flowers depends on the cultivars, the working concentrations of this ethylene-binding inhibitor, and the treatment duration. Treatment with 1-MCP (0.6 µL L−1 for 2 h) shortened the vase life of cut ‘Karl Rosenfield’ and ‘Sarah Bernhardt’ peony flowers [26]. Other previous studies have shown that treatment with 1-MCP (1 µL L−1 for 12, 24, 36, 48, 60, and 72 h) retarded the flower opening process, prevented petal abscission, and resulted in a longer vase life and better postharvest quality of the ethylene-sensitive (‘Luo Yang Hong’) peony cultivar [17,18] (Table 1). Moreover, 1-MCP effectively prevented ethylene-induced senescence in cut peony flowers by inhibiting ethylene production in tree peony flowers and suppressing the activity of two main ethylene biosynthesis enzymes (ACS and ACO) and ACC content in petals [17,18]. However, 1-MCP treatment did not influence the postharvest performance, vase life, ethylene production, and the activities of PlACS and PlACO enzymes in the ethylene-insensitive tree peony ‘Xue Ying Tao Hua’ cultivar [17,18]. These results indicate that the responses of the cut peony flowers to ethylene may depend on the cultivars. Ethylene inhibitors application in ethylene-insensitive peony flowers may cause decreased ornamental values and postharvest quality of cut flowers. Thus, the application of ethylene inhibitors in cut peony flowers should be clarified based on the ethylene sensitivity of the peony cultivars.

2.2. Antimicrobial Agents in Vase Solutions

In addition to ethylene damage, the proliferation of bacteria in the vase solution and the stem ends of cut flowers is the principal reason for the decreased vase life of cut flowers [16,27]. The growth of bacteria blocks the cut flower stems and disturbs the transportation and absorption of water and nutrient factors to flower heads [27]. In addition to blockage, the bacterial growth in the vase solution produces enzymes and toxic compounds, and these metabolic products can accelerate flower senescence in cut flowers [27]. Many antimicrobial agents, such as ClO2, rapamycin, nano silver, and herbaceous peony polyphenols, have been applied to suppress microbial proliferation in the vase solution to improve vase life and postharvest quality of cut peony flowers.

2.2.1. ClO2 and Rapamycin Treatments Improve Quality of Peony Flowers

Adding ClO2 with various concentrations (25, 50, and 100 mg L−1) in a basic vase solution improved the postharvest quality of cut ‘Luoyanghong’ peony flowers [28]. Among treated concentrations, the dose of 50 mg L−1 was the most effective in inhibiting bacterial growth in the vase solution, improving water balance, and increasing the vase life of ‘Luoyanghong’ peony flowers by 2.12 d compared to the control (Table 1) [28]. Additionally, ClO2 reduced the respiration rate and ethylene production, inhibited the increase in product malonaldehyde content and membrane lipid peroxidation, and increased the antioxidase activities and the soluble protein content in tree peony flowers [28], contributing to improving the total quality of flowers [28]. Pretreatment with rapamycin (0.01 µM L−1 for 2 h) also delayed the senescence and extended the postharvest life of ‘Luoyanghong’ tree peony flowers by 0.8 d compared to non-treated flowers (Table 1) [29]. However, long-term use of these chemicals harms human health and the environment.

2.2.2. Nanotechnology Application in Peony Flowers

In recent years, nanotechnology has been widely used in the floral industry to improve the postharvest quality of various cut flowers due to its antimicrobial and ethylene inhibitor effects [30,31,32,33,34]. In peony flowers, nanotechnology can potentially increase the vase life of cut flowers. Treatment with 10 mg L−1 of the green synthesis of Ag-NPs (Ag-NPs) using the leaf isolation of E. ulmoides inhibited bacterial growth in the xylem vessels of tree peony ‘Luoyanghong’ during the vase period, resulting in a higher water absorption rate, better postharvest quality, and a longer vase life (Table 1) [35]. Applying 20 µg L−1 nanosilver (NS) to cut peony ‘Bartzella’ flowers increased water absorption and extended vase life compared to untreated flowers (Table 1) [36]. The previous study also showed that pretreatment with NS significantly decreased oxidative stresses and the susceptibility of the cut stem end of peony flowers to microbial infection [8]. Additionally, NS induced the mRNA levels of three aquaporin genes in peony petals, including the NOD26-like intrinsic protein gene (PlNIP) and plasma intrinsic protein genes (PlPIP1;2 and PlPIP2;1), which influence the maintenance of the positive water balance of cut peony flowers [8]. Consequently, the floral freshness and flower diameter were retained, and the vase life of cut peonies was prolonged by 4 d (Table 1).
The combination of NS with other preservative solutions or storage methods also extended the postharvest life of cut peony flowers. Pretreatment with NS by spraying and combination with cold storage or a combination of both cold storage and low oxygen prolonged the vase life of cut ‘Qihua Lushuang’ peony flowers up to 5.6 and 5.8 d, compared to control groups (Table 1) [37]. Combined treatment of NS (1 mg L−1) and sucrose (20 g L−1) (NS+S) was effective in extending the vase life of cut peony flowers in four cultivars by 15% (‘Hania’), 17% (‘Duchese de Nemorous’), 22% (‘Ursynów’), and 35% (‘Wiesbaden’) compared to control and other preservative solutions (Table 1) [38]. NS+S treatment also reduced tylose formation and bacterial blockages in the cut flower stems and increased the accumulation of the carbohydrate content in peony petals [38]. However, in other peony cultivars, NS+S did not positively influence the postharvest longevity and quality of cut flowers during vase periods or storage duration [39]. This indicates that NS postharvest treatment should be individually tailored to each cultivar.

2.3. Other Chemicals Used as Holding Solutions

In addition to ethylene inhibitors and antimicrobial agents, other preservative solutions such as Chrysal, Floralife, 8-hydroxyquinoline citrate (8-HQC), carbon source fresh-keeping solution (glucose, sucrose, and trehalose), silicon, and melatonin were also applied to improve the ornamental performance and vase life of cut peony flowers.

2.3.1. Chrysal and Floralife Treatments

Chrysal and Floralife, two popular commercial preservative solutions, are widely used in the cut flower industry. The Chrysal sachet was applied for 14 cut peony cultivars but extended the postharvest longevity of only two cultivars by approximately 2 d [38]. Whereas the treatment with Floralife 300 was effective only in one cut peony cultivar among 14 tested cultivars [38]. However, an initial hydration pulse by Floralife after harvest and before low-temperature storage and transport positively affects the floral freshness and vase life of cut peony flowers. Cut peony flowers treated with a combination of Foralife® Express 200 (FloraLife®, Walterboro, SC, USA) (10 mL L−1) and Foralife® Bulb 100 (FloraLife®, Walterboro, SC, USA) (2 mL L−1) exhibited a longer vase life by 1.3 d in ‘Jules Elie’ and 2.8 d in ‘Festiva Max’ compared to water only (Table 1) [40]. Treatments with 0.5% and 1% of Floralife as holding solutions after dry storage also effectively prolonged the vase life of cut peony flowers by 1.2–1.7 d compared to non-treated flowers by improving the water balance of cut flowers [41].

2.3.2. Effectiveness of 8-HQC in Vase Life Extension in Peony Flowers

Recently, 8-HQC has been used as a preservative solution for cut peonies during the vase period, storage, or shipping [38,39]. In vase solution, 8-HQC is often combined with sucrose to enhance the postharvest quality of cut flowers. However, 8-HQC showed less effectiveness in the improvement of vase life and quality of cut peonies compared to other preservative solutions [38,39]. The previous observation indicated that the vase solution composed of 8-HQC and sucrose extended the longevity of only one peony cultivar (by 32%, >2 d) among 14 tested cultivars relative to control flowers [38]. In cut peony ‘Sarah Bernhardt’ flowers, 8-HQC plus 2% of sucrose in the holding solution increased the vase life of cut flowers by 0.6 d in non-stored conditions and 0.7 d in stored conditions (stored for 12 weeks) [39]. 8-HQC treatment under cold storage and low oxygen conditions prolonged the vase life of cut ‘Qihua Lushuang’ peony flowers by 5.7 d compared to control [37].

2.3.3. Carbohydrate Sources Significantly Prolonged Vase Life of Cut Peony Flowers

The single treatments of glucose, sucrose, and trehalose were also applied to supply energy for flower organ respiration and help to maintain the osmotic balance in cut flowers to improve flower opening and floral freshness [37]. Glucose, sucrose, and trehalose treatments promoted the flowering of cut flowers and effectively improved the ornamental value of peony ‘Hong Feng’ flowers. Glucose, sucrose, and trehalose applications significantly prolonged the vase life of cut peonies by 1.4–2.3 d (Table 1) and the flowering period by approximately 4 d compared to control flowers. These treatments also increased the flower diameter and relative fresh weight of cut peonies during the vase period [37].

2.3.4. Silicon Treatment Improved Lignin Content and Delayed Flower Senescence

Silicon is a nontoxic and useful element that participates in various plant activities. It has been shown that silicon helps plants respond to biotic and abiotic stresses and increases the rigidity of the plant cell walls [42,43,44,45]. Silicon can also stimulate the defense system in plants against the infection of Fusarium wilt and P. ultimum, which cause plant tissue susceptibility to wilt symptoms and damping off [45]. Previous studies also showed that silicon function in plant cell walls was similar to lignin, which plays an important role in the ability of the cell wall to resist external force [46]. Pre-harvest application of silicon (500 µg mL−1) significantly enhanced the accumulation of lignin content in peony flower stems by up-regulating the expression levels of the key genes related to lignin biosynthesis pathways (PlPAL, Pl4CL, and PlCCoAOMT) [46]. Higher lignin content in flower stems prevents the breaking or bending of peony inflorescence stems, contributing to maintaining good ornamental value of cut flowers [46]. Postharvest treatment with silicon (75 mg L−1) improved the total quality of cut peonies through the flower stem mechanical strength and antioxidant enzyme activities [47,48]. Silicon treatment effectively maintained the flower’s fresh weight, delayed flower senescence, and prolonged the vase life by 2 d in ‘Taebaek’ and 4 d in ‘Euiseong’ peony flowers [47] (Table 1).

2.3.5. Melatonin Treatment Improved Flower Stem Strength and Retarded Flower Senescence

Melatonin is a multifaceted plant hormone with a structure similar to auxin (IAA) [49]. Melatonin functions are related to various plant physiological processes, including germination, growth and development regulation, photosynthesis enhancement, abiotic stress resistance, flowering, plant organ senescence delay, and postharvest physiology improvement [50,51,52,53]. Recently, melatonin has been widely used in the postharvest of horticultural products due to its safety and stable nature [54,55,56]. Postharvest application of melatonin positively affected vase life and ornamental value in many cut flower species [57,58,59]. In peony flowers, the application of exogenous melatonin effectively enhanced flower stem strength by elevating the ratio of lignin content and the S/G lignin composition by influencing the mRNA levels of genes involved in lignin biosynthesis (PlPAL, PlCCR, PlCAD, PlCOMT, and PlPOD) [60]. Postharvest treatment with melatonin (50 µM L−1) delayed petal senescence and extended the vase life of cut peony flowers by 1.6 d (‘Qi Hua Lu Shang’) and 1.2 d (‘Da Fu Gui’), respectively, compared to non-treated flowers [61] (Table 1). Melatonin treatment also increased flower opening and improved the water balance of cut peony flowers during vase life [61]. The malondialdehyde content in peony petals was reduced by melatonin treatment [61]. Additionally, the activities of antioxidant enzymes were enhanced in the melatonin-treated peony petals during vase periods [61]. The combination of melatonin (0.5 mM) and sucrose (0.5%) in the vase solution also prolonged the vase life of cut peony flowers by 2 d compared to control flowers after 4 weeks of cold storage [62]. Combined treatment of melatonin, sucrose, NaCl, acid citric, and BA also extended the vase life of the cut double peony flower to 7.5–8.9 d by enhancing water absorption and antioxidant capacity in petals and delaying the senescence process and damage to cell membranes [62]. These results show that melatonin application improves flower stem strength, delays petal senescence, and preserves the postharvest quality of cut peony flowers, offering valuable insight into the cut flower industry.

2.3.6. Polyphenols Extracted from Herbaceous Peony Improved the Postharvest Quality of Peonies

Polyphenols, a natural plant extract, have strong activities, including antibacterial, antioxidant, and anti-inflammatory properties [63,64,65,66,67]. Recently, polyphenols extracted from plants have been used in the cut flower industry to improve the postharvest quality of cut flowers [68]. The extracted polyphenols from the petals of herbaceous peonies were used as the vase preservative solution for cut peony ‘Hongyan Zhenghui’ flowers [69]. Polyphenols significantly increased the flower diameter and prolonged the vase life of cut peonies by 2 d compared with the control flowers [69]. Moreover, the polyphenol treatment enhanced the activities of the antioxidant enzymes CAT, POD, and SOD and the content of soluble protein in the petals of cut peony flowers [69]. The growth of Aspergillus spp. at the stem ends of cut peonies was inhibited by the polyphenol treatment [69]. Additionally, the polyphenol treatment upregulated the expression of the aquaporin genes (PlPIP1;2, PlPIP2;1, and PlPIP2;2) in peony petals during vase life [69]. Therefore, the polyphenol treatment effectively maintained positive water balance, initial fresh weight, and flower diameter of cut flowers and prolonged the postharvest life of peonies.
Table 1. Improved vase life of cut flowers upon chemical application in different peony cultivars.
Table 1. Improved vase life of cut flowers upon chemical application in different peony cultivars.
Chemicals
(Single or Combined Treatment)		ConcentrationsCultivarsExtended Vase Life (Days)References
1-MCP1 µL L−1‘Luo Yang Hong’0.4[17,18]
ClO250 mg L−1‘Luoyanghong’2.12[28]
Rapamycin0.01 µM L−1‘Luoyanghong’0.8[29]
Ag-NPs (green synthesis)10 mg L−1‘Luoyanghong’1[35]
Nanosilver (NS)20 µg L−1‘Bartzella’1.9[36]
Nanosilver (NS)-‘Hongyan Zhenghui’4.0[8]
Nanosilver (Cold storage + Low O2)2.3 mg L−1‘Qihua Lushuang’5.8[37]
Nanosilver + Sucrose10 mg L−1‘Hania’1.5
‘Duchese de Nemorous’1.5[38]
‘Ursynów’1.8
‘Wiesbaden’2.4
Chrysal Clear Sachet5 g L−1‘Hania’1.3
‘Königin Wilhelmina’1.9[38]
‘Ursynów’2.3
Floralife 30010 mL L−1‘Hania’1.5[38]
Floralife0.5%‘Mons Jules Elie’1.2[41]
‘John C. Lee’1.4
1.0%‘Mons Jules Elie’1.7[41]
‘John C. Lee’1.6
Foralife® Express 200+ Foralife® Bulb 10010 mL L−1 + 2 mL L−1‘Jules Elie’1.3
‘Festiva Max’2.8[40]
8-HQC + Sucrose200 mg L−1 + 20 g L−1‘Graziella’2[38]
‘Wiesbaden’2.2
8-HQC + Sucrose (non-stored condition)200 mg L−1 + 20 g L−1‘Sarah Bernhardt’0.6[39]
8-HQC + Sucrose (stored condition)200 mg L−1 + 20 g L−1‘Sarah Bernhardt’0.7[39]
8-HQC (Cold storage + Low O2)0.05 mg mL−1‘Qihua Lushuang’5.7[37]
Glucose20 g L−1‘Hong Feng’1.5[37]
Sucrose20 g L−1‘Hong Feng’1.7[37]
Trehalose20 g L−1‘Hong Feng’2.2
Silicon (Na2SiO3)75 mg L−1‘Taebaek’2[47]
‘Euiseong’4
Melatonin50 µM‘Da Fu Gui’1.2[61]
‘Qi Hua Lu Shang’1.6
75 µM‘Da Fu Gui’0.8[61]
25 µM‘Qi Hua Lu Shang’1[61]
Melatonin + Sucose0.5 mM + 0.5%‘Sarah’2[62]
Polyphenols8%‘Hongyan Zhenghui’2[69]
“-”, not given.

2.4. Chemical Treatments to Reduce the Susceptibility of Peony Flowers to Botrytis

The application of the chemical to decrease gray mold disease infection in peony flowers is shown in Table 2. Peony flowers are often harvested at the bud stage and can be infected by botrytis during cold storage, and the disease symptoms can be more severe with long-term cold storage [20]. Peony flowers can be stored successfully for a long period (several weeks) if the gray mold disease is controlled effectively [20]. Fungicides are often applied to reduce botrytis infection in cut peonies during cold storage [20]. Many fungicides have been used to treat peony botrytis based on the conditions and cultivar susceptibility [20]. The previous study showed that treatment with paradichlorobenzene effectively suppressed the disease infection in cut peony ‘Felix Crousse’ flowers [70]. The treatment with 30% pyrimethanil for 8h under normal and controlled atmosphere storage strongly decreased the gray mold disease incidence rate on leaves and petals of cut peony flowers after 10 weeks of dry storage [71]. However, the downside of fungicide treatment was the stimulation of the senescence symptoms, such as browning of the leaves during storage, and the effectiveness of the fungicide was decreased under long-term cold storage. Moreover, the use of fungicides causes fungicide resistance in botrytis since botrytis is susceptible to developing fungicide-resistant strains [20].
Pre-storage treatment with STS (0.463 mM for 2 h at room temperature) significantly reduced the disease infection rate and symptom severity in all plant parts in cut peony ‘Shawnee Chief’ flowers after 6 and 8 weeks of storage [72]. However, the disease symptoms and incidence rate had increased after 10 weeks of storage [72]. In addition, the use of STS causes environmental pollution and is harmful to humans.
Other alternative treatments using elicitors of defense response, such as methyl jasmonate (MeJA), were applied to enhance the botrytis resistance in cut peonies [72]. The pre-storage application of 0.2 mL vapor MeJA (working concentration is 130 mM m−3) to fresh-cut peonies reduced incidence and symptom severity in cut flowers [72]. However, MeJA treatment did not affect the botrytis infection rate and flower opening of cut peony ‘Shawnee Chief’ flowers after 8 or 10 weeks of long-term cold storage [72]. Therefore, an effective and stable method for enhancing the botrytis resistance in cut peony flowers is necessary.
Table 2. Chemical treatments reduced Botrytis susceptibility in peony flowers.
Table 2. Chemical treatments reduced Botrytis susceptibility in peony flowers.
Chemical TreatmentConcentrationsCultivarsResponse to Botrytis References
1-MCP1 µL L−1‘Luo Yang Hong’Reducing Botrytis susceptibility[17,18]
Silver nanoparticles
(Ag-NPs)		10 mg L−1‘Luoyanghong’Inhibiting Botrytis infection[35]
Nanosilver (NS)20 µg L−1‘Bartzella’Reducing Botrytis infection[36]
Nanosilver
(Green Synthesize)		20 µg L−1‘Hongyan Zhenghui’Prevents Botrytis infection by suppressing ethylene response and microbial proliferation[8]
Chlorine Dioxide
(ClO2)		50 mg L−1‘Luoyanghong’Reducing Botrytis infection and microbial growth[28]
8-HQC200 mg L−1‘Graziella’Reducing Botrytis susceptibility through antimicrobial action[38]
Fluazinam25 mg L−1‘Sarah Bernhardt’Fungicide activity reduced Botrytis infection[20]
Prochloraz50 mg L−1‘Hania’Fungicide activity inhibited Botrytis development[20]
Azoxystrobin10 mg L−1‘Wiesbaden’Fungal inhibitor inhibiting Botrytis cinerea growth[20]
Iprodione10 mg L−1‘Ursynów’Fungicide with proven effects against Botrytis infection[20]
Pyrimethanil30%‘Sarah Bernhardt’
‘The Fawn’		Decreasing the gray mold disease incidence rate [71]
STS0.5 mM‘Hania’Preventing ethylene-mediated senescence and reducing Botrytis infection[72]
MeJA100 µM‘Wiesbaden’Inducing resistance through the jasmonic acid pathway, reducing Botrytis infection[72]

3. Physical Treatments Delayed Flower Senescence and Reduced Botrytis Susceptibility in Peony Flowers

Physical methods include management of the environmental conditions (temperature and humidity) during storage, modified atmosphere packing, and alternative transport methods (dry transport), which have been applied to delay the flower senescence process and inhibit the gray mold disease development in cut peony flowers (Table 3).

3.1. Temperature Management During Storage

Cold (low-temperature) storage technology is important during the postharvest handling and transport of perishable cut flowers to maintain good quality and minimize biotic and abiotic stress that can decrease ornamental values and the vase life of cut flowers [73,74]. Holding cut flowers (except for chilling-damaged flower species) at low temperatures (2–4 °C) reduces transpiration, respiration, and carbohydrate loss and limits ethylene production, thus maintaining the vase life of cut flowers [41,75,76,77,78]. The temperature between 0 and 2 °C has been determined as the industry standard temperature for storing cut peony flowers (>25 cultivars) with a storage time of 1 to 3 months [41,75,79,80,81]. The postharvest life of cut flowers stored for a month near 0 °C was decreased by 1–2 d compared to non-cold storage flowers [82]. An additional one or two days is decreased as storage periods increase to two or three months of cold storage along with an increase in the incidence rate of gray mold disease infection in cut peonies [82]. Sub-zero temperatures have been applied to edible crops for improved storage duration, and these methods helped maintain better postharvest attributes of the fruits [83,84,85,86]. Cut peony flowers showed multiple characteristics of tolerating near-freezing temperatures [87]. In peonies, the tight bud stage tolerated a lower temperature than the full development of flowers. Exposure to −3.8 and −4.0 °C did not freeze the sepal tissues of tree peony flowers ‘Luoyanhong’ and ‘Fengdanbai’ [87,88,89,90,91]. However, long-term storage at sub-zero temperatures caused the hydrolysis of starch to soluble solids content, thus decreasing the tolerance of freezing and sub-zero temperatures in peony buds [92,93,94]. To prevent this, pre-treatment of the cut peony stems with sucrose or commercial hydrator pulses before sub-zero temperature storage successfully extended the cut peonies’ storage duration. However, this innovative method reduced the quality of cut peony flowers, such as failing to open and flowers deformed [95]. Whereas dry (where cut flower stems were not held in water or solution) and cold storage (0–4 °C) preserved the bud stage and vase life of cut peonies and is the standard method of holding cut peony stems in both short- and long-term cold storage [81]. The improvement of the vase life of cut peonies by this cold storage method may be due to faster carbohydrate consumption during short storage and carbohydrate hydrolysis at the initial days of the vase life and the higher levels of fructose and glucose content in petals at the end of the long storage [81].

3.2. Controlled Atmosphere Storage

Controlled atmosphere (CA) storage of cut flowers is based on three main factors, temperature, O2, and CO2, which significantly influence the vase life and ornamental values of cut flowers [96,97,98,99]. CA storage prolongs the postharvest life of the cut flowers by inhibiting the synthesis and action of ethylene, reducing respiration rate, and preventing gray mold disease and bacterial growth [99]. This method has been applied in the floricultural industry and extends the storage life of various cut flower species [96,97,98,99]. The atmosphere was modified using a commercial pallet box, which resulted in average concentrations of CO2 and O2 of approximately 3% applied to two peony cultivars, ‘Sarah Bernhardt’ and ‘The Fawn’ [71]. After 10 weeks of storage, B. cinerea infection rate on the cut peony flowers was significantly lower under a modified atmosphere compared to a normal atmosphere [71]. The vase life of stored cut flowers under a modified atmosphere was shorter than that of non-stored cut flowers but still commercially acceptable [71]. The maximum quantum efficiency of the PSII value was significantly lower for cut flowers stored under modified atmosphere conditions but did not affect the vase life of cut flowers [71].
Table 3. Physical treatments reducing Botrytis susceptibility in peony flowers.
Table 3. Physical treatments reducing Botrytis susceptibility in peony flowers.
Physical TreatmentsConditionsCultivarsBotrytis ResistanceReferences
Cold Storage4 °C for 7 days‘Sarah Bernhardt’Moderate[39]
Low temperature
pretreatment		1 °C for 12 h before vase‘Hongyan Zhenghui’Moderate[38]
Modified Atmosphere PackagingLow O2 (2–3%), high CO2‘Qihua Lushuang’High[37]
Modified AtmosphereAverage concentrations of CO2 and O2 of approximately 3%‘Sarah Bernhardt’
‘The Fawn’		Moderate[71]

3.3. Dry Storage Influenced Postharvest Quality of Peony Flowers

Wet storage (WS) and dry storage (DS) are two common methods used in the cut flower industry during cold storage and are associated with different advantages [100]. The disadvantages of WS are that it requires more space in the cold stores, shorter storage duration, and the development of fungal diseases and bacterial proliferation in water and flower stems [100]. DS has various advantages over WS, but this method limits cut flowers sensitive to water stress. DS reduces the fungal disease infection and enhances water uptake, thus improving the postharvest performance of cut flowers [100]. The gray mold disease index on peony petals of ‘Sarah Bernhardt’ and ‘The Fawn’ was significantly reduced after 10 weeks of DS under a normal and controlled atmosphere [71]. In peony ‘Yang Fei Chu Yu’ flowers, DS in short-term cold storage effectively improved the vase quality of the cut flowers by enhancing the water uptake rate through aquaporin regulation [101]. DS also increased the cut peonies’ fresh weight and flower diameter during the vase period. DS treatment increased the water uptake rate of cut flowers, thus inducing the expression levels of aquaporin genes (PlPIP1;3, PlTIP2;1, and PlNIP1;2-like) in peony petals [101]. The upregulated expression of aquaporin genes by DS influenced the cell expansion in peony petals, resulting in improved flower opening and maintaining flower freshness during the vase period [101]. In addition, the DS peony flower showed faster starch consumption and sucrose hydrolysis during the initial days of short storage and the vase period, resulting in higher levels of glucose and sucrose at the last days of the storage, contributing to the higher postharvest quality of cut flowers [81].

4. Conclusions and Future Direction

Herein, we have summarized and discussed how the potential postharvest treatments work to improve vase life and botrytis resistance in cut peony flowers. Enhancing ethylene and botrytis resistance, lignin content in flower stems, and water uptake through postharvest treatments is necessary to extend vase life and improve the postharvest performance of cut peony flowers. Moreover, a deep study of flower senescence characteristics in peonies needs to be conducted to clarify the peony flowers based on their susceptibility to botrytis, ethylene, and water stress (including due to bacterial growth and low lignin content in flower stems). This classification in cut peony flowers greatly aids in developing effective postharvest treatments for each specific cultivar group. Careful consideration of the sensitivity of cut peony flowers to ethylene, whether ethylene-sensitive or not, is crucial to avoid using ethylene inhibitors on ethylene-insensitive cultivars, thereby reducing undesirable ornamental value losses and economic costs. Since NS has the potential to inhibit bacterial growth in vase solutions, preventing B. cinerea infection, and suppressing the ethylene response in ethylene-sensitive flowers, we believe that NS (especially green-synthesized NS) can provide a commercially viable and environmentally friendly method applied to all botrytis-susceptible, ethylene-sensitive, and water stress-sensitive cut peony flowers. Further research will be conducted for each peony cultivar group to optimize the postharvest treatments and improve the quality of cut peony flowers.

Author Contributions

T.N.; writing—original draft. S.H.; writing—original draft, writing—review and editing. All authors have read and agreed to the published version of the manuscript.

Funding

This research received no external funding.

Data Availability Statement

The original contributions presented in the study are included in the article, further inquiries can be directed to the corresponding author.

Conflicts of Interest

The authors declare no conflicts of interest.

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MDPI and ACS Style

Nguyen, T.; Ha, S. The Potential Postharvest Treatments to Delay Flower Senescence and Improve Botrytis Resistance in Cut Peony Flowers. Horticulturae 2024, 10, 1352. https://doi.org/10.3390/horticulturae10121352

AMA Style

Nguyen T, Ha S. The Potential Postharvest Treatments to Delay Flower Senescence and Improve Botrytis Resistance in Cut Peony Flowers. Horticulturae. 2024; 10(12):1352. https://doi.org/10.3390/horticulturae10121352

Chicago/Turabian Style

Nguyen, Toan, and Suong Ha. 2024. "The Potential Postharvest Treatments to Delay Flower Senescence and Improve Botrytis Resistance in Cut Peony Flowers" Horticulturae 10, no. 12: 1352. https://doi.org/10.3390/horticulturae10121352

APA Style

Nguyen, T., & Ha, S. (2024). The Potential Postharvest Treatments to Delay Flower Senescence and Improve Botrytis Resistance in Cut Peony Flowers. Horticulturae, 10(12), 1352. https://doi.org/10.3390/horticulturae10121352

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