Stewart Postharvest Review

Stewart Postharvest Review An international journal for reviews in postharvest biology and technology

Preharvest application of synthetic fungicides and alternative treatments to control postharvest decay of fruit Erica Feliziani and Gianfranco Romanazzi* Department of Agricultural, Food and Environmental Sciences, Marche Polytechnic University, Ancona, Italy

Abstract Purpose of the review: This article provides a state-of-the-art review of fungicides that are commonly used before harvest in conventional agriculture to prolong the storage of fresh commodities, and of the alternatives to fungicides recently made available for plant protection. Findings: Considering the high percentage of postharvest loss of fruit due to pathogen spoilage and the frequent development of pathogen isolates that are resistant to one or more active ingredients, alternatives to synthetic fungicides are needed. This review compares the current practices in conventional agriculture that are used to control postharvest rot of fruit with the alternatives to synthetic fungicides that are now available. The review summarizes the different fungicides and the corresponding alternatives, such as natural compounds, decontaminating agents that are ‘generally recognized as safe’, and biological control agents that have been applied in smallscale and large-scale tests. For some cultivated crops, including strawberries, table grapes, and stone and pome fruits, we include the time and method of application of preharvest treatments that can be applied to preserve fruit quality during storage. Limitations/implications: Even considering the research efforts in the search for alternatives to fungicides, at present there are few natural compounds that are as effective as fungicides. However, according to integrated pest management, to overcome the drawbacks that can arise with the use of a single strategy, an integration of methods might provide additive or synergistic effects for disease control. Directions for future research: Further insight at the molecular level into the interactions between host plants and chemicals applied will help us to better understand the changes that occur in host plants following treatments, or the effects of the treatments on the pathogens. This new knowledge will optimize the treatment application to provide the greatest effects with the minimum number/ time/ concentration of treatments applied.

Abbreviations: Biological Control Agent BCA Generally Recognized as Safe GRAS *Correspondence to: Gianfranco Romanazzi, Department of Agricultural, Food and Environmental Sciences, Marche Polytechnic University Via Brecce Bianche, 60131 Ancona, Italy. Tel: +39-0712204336; Fax: +39-0712204856; Email: [email protected] Stewart Postharvest Review 2013, 3:4 Published online October 2013 doi: 10.2212/spr.2013.3.4

© 2013 Stewart Postharvest Solutions (UK) Ltd. Online ISSN:1945-9656 www.stewartpostharvest.com

Introduction Postharvest rot is the main problem in the storage of fruit and vegetables. In developing countries, 50% or more of the fresh produce can be lost because of fungal spoilage. For this reason, field application of synthetic fungicides is common practice in conventional agriculture. For some crops, such as kiwi, apple, pear and citrus fruit, treatments with fungicides are also carried out after harvest. Even if the use of fungicides is widely accepted, their continuous application can also have several negative side effects. The appearance of pathogen strains that are resistant to synthetic fungicides has been reported for many fungi [1]. For example, several strains of Botrytis cinerea, which is considered the most important postharvest pathogen [2**], are resistant to almost all of the commonly used fungicides [3]. For many crops, current legislation does not allow the use of fungi-

Feliziani and Romanazzi / Stewart Postharvest Review 2013, 3:4

cides after fruit harvest, and in organic agriculture, the use of any synthetic active ingredient is not allowed. In addition, over the last few years, there has been increasing numbers of requests from consumers of fresh fruit and vegetables that these are free from fungicide residues. This review reports on the most common practices applied in conventional agriculture, and the alternatives to synthetic fungicides that have been introduced more recently or that are been studied to control postharvest rot of fruit and to preserve fruit quality during storage through the application of preharvest treatments. Synthetic fungicides In conventional agriculture, several fungicides that are active against postharvest pathogens are commercially available. These can be classified into categories according to their biochemical modes of action: fungicides affect fungal respiration, microtubule function, osmoregulation, methionine biosynthesis, or sterol biosynthesis [4]. However, as pathogen strain resistant to fungicides is now very common, this emphasizes the need for accurate adoption of the Fungicide Resistance Action Committee (FRAC) guidelines that suggest a limited number of applications per year. To prevent fungicide resistance, it is necessary to combine or alternate compounds with different modes of action, especially between pre- and postharvest treatments. Fungicides should also be used as preventive rather than curative treatments, when the levels of contamination or infection are low. Therefore, according to this approach, it is fundamental to continuously monitor the field to control the eventual development of pathogenresistant strains. Some of the fungicides that have been most used recently belong to the anilinopyrimidines, which include cyprodinil, mepanipyrim, and pyrimethanil, where the mode of action consists of inhibition of methionine biosynthesis and secretion of hydrolytic enzymes. Boscalid is a contact fungicide that belongs to the pyridine-carboxamides, which prevents energy production, as it is a succinate dehydrogenase inhibitor. Boscalid blocks the synthesis of essential cell components, and hence it disrupts fungal growth through deleterious effects on spore germination, germ-tube elongation, mycelial growth, and sporulation. Similarly, the mode of action of the active principle of fluopyram, which is a pyridinyl-ethylbenzamide fungicide, mainly consists of inhibition of succinate dehydrogenase, and thus fungal respiratory chain complex II. Pyraclostrobin acts through inhibition of mitochondrial respiration by binding at the ubiquinol oxidation center (the Qo site) of cytochrome bc1 in complex III. Fenhexamid is a hydroxyanilide that inhibits the C3-keto-reductase step in ergosterol biosynthesis, which prevents fungal germ-tube elongation. As fenhexamid, fenpyrazamine, belonging to the same class, inhibits the sterol biosynthesis, preventing fungal germ tube and mycelium elongation. Beside fenhexamid and fenpyrazamine, the sterol biosynthesis inhibitors are a wide class of fungicides that include imidazoles, triazoles (eg,

teboconazole, fenbuconazole), and morpholin, some of which are used to control decay causing agents. Iprodione belongs to the class of dicarboximides and it prevents the germination of spores and the growth of mycelia through inhibition of DNA and RNA synthesis and cell division in fungi. Other fungicides used to control fungi able to cause postharvest losses are the phenylpyrroles fludioxonil and fenpiclonil, whose mode of action is interference with the transport processes of sugars and aminoacids in the plasma membrane of fungi. Thiabendazole belongs to the class of benzimidazoles, and it is a systemic fungicide that interferes with the mitotic process [1]. Alternatives to synthetic fungicides Efforts of researchers have led to the development of novel control tools that can be used as alternatives to synthetic fungicides. This kind of preharvest application can be grouped into three categories: (i) natural compounds; (ii) decontaminating agents that are ‘generally recognized as safe’ (GRAS); and (iii) biological control agents (BCAs). All of these tools can be used alone or as combinations, to benefit from additive or synergistic effects [5, 6**]. Natural and GRAS compounds Natural and GRAS compounds are substances that are known not to be harmful to the environment and to human health, and these are used for their antimicrobial properties and their induction of plant defenses. Among the natural compounds, plant extracts and essential oils have been reported to control postharvest diseases, both in vitro and in vivo, and to prolong the overall quality and storage life of fresh commodities [7** –10]. Inorganic salts have been shown to be active antimicrobial agents against a range of phytopathogenic fungi, and among these agents, the bicarbonates have been proposed as a safe and effective alternative means of controlling postharvest rot of fruit and vegetables. In addition, as well as these salts being nontoxic and having minor environmental impact at their effective concentrations, they are inexpensive [11]. Several sanitizers that are classified as GRAS have been applied to extend postharvest storage of various produce, including acetic acid, electrolyzed oxidizing water, and ethanol [6**]. However, decontaminating agents are more commonly used in postharvest applications rather than in field applications. Resistance inducers are plant or pathogen constituents, or their analogs, that act as plant elicitors, as they can activate the plant defense mechanisms, and thus simulate the presence of pathogens. Among the resistance inducers, the natural biopolymer chitosan and the synthetic elicitor benzothiadiazole have been reported to activate systemic acquired resistance in horticultural produce [12**–14]. Biological control agents BCAs are mainly bacteria and yeasts that are ‘antagonists’ to the pathogens that cause postharvest fruit spoilage. These can act through different mechanisms: competition for nutrients and space; antibiosis; parasitism; induction of resistance in the host tissue; and production of volatile metabolites [15**, 2


16]. These microorganisms have been selected not to be toxic for the plant and not to produce secondary metabolites that are harmful for human health. Several products against some of the main postharvest pathogens are based on microorganisms and are registered and commercially available. Some examples of antagonistic microorganisms that are currently applied preharvest or postharvest are: Pseudomonas syringae (BioSave, JET Harvest Solutions, Longwood, FL, US), Cryptococcus albidus (YieldPlus, Lallemand, Montreal, Canada), Bacillus subtilis (Serenade, Bayer, Leverkusen, Germany), Candida sake (Candifruit, IRTA, Lleida, Spain), Pantoea agglomerans (Pantovital, Domca, Granada, Spain), Aurobasidium pullulans (Boni Protect, Bio-Ferm, Tulln, Austria; Botector, Manica, Rovereto, Italy), Candida oleophila (Nexy, BioNext, Paris, France), Bacillus amyloliquefaciens (AmyloX, Biogard CBC, Grassobbio, Italy), and Metschnikowia fructicola (Shemer, Bayer, Leverkusen, Germany) [17–21]. It is essential that a formulated product based on antagonistic microorganisms retains the properties of the initial laboratory -grown cultures, despite the mass production of large quantities. These properties include species purity, genetic stability, cell viability, attributes as colonizers on fruit surfaces, as well as other aspects of their mechanisms of action [19, 20, 22]. To be successful for preharvest applications, putative BCAs must tolerate low nutrient availability, UV radiation, high temperatures and dry conditions [15**, 23]. A promising approach for improving the effectiveness of antagonistic microorganisms in decay control is the combination of BCAs with various other substances, such as salts, and decontaminating agents with other microorganisms, or in some cases with fungicides [6**, 23–26]. These combinations both reduce the fluctuation and increase the level of decay control, as they are based on the use of different mechanisms of action. In addition, the integration of fungicides with BCAs offers the opportunity to reduce the amounts of fungicides in preharvest application, thus lowering the risk of development of pathogen resistance and of residues remaining on marketed products [23]. Preharvest treatments to prevent postharvest rot To increase the effectiveness of preharvest treatments with either fungicides or alternatives, it is necessary to know the right time and method of application. Moreover, it is important to analyze the interactions of a specific crop with a specific pathogen, by studying the ways through which a fungus can infect its host, the time of the greatest risk of contamination/ infection, and the environmental factors that can affect the onset and development of disease. Compared to postharvest treatments, applications before the harvest can be suitable for fruit, such as table grapes and strawberries, which have a bloom on the surface and/or can suffer postharvest wetting or handling. Moreover, preharvest treatment might have a preventive effect other than a curative one, as the development of postharvest decay often arises from an inoculum that survives and accumulates on the fruit in the field, or

during postharvest storage and shelf life. On some commodities, eg, table grapes, strawberry, early season preharvest treatments with fungicides or alternatives can reduce the fungal inoculum, then allow a better control of postharvest decay [27]. The fungicides recently available on markets have often a preharvest interval that is very short, sometimes of 1–3 days on some crops. In this way they could be applied very close to the harvest, and maintain their efficacy even during fruit storage. Some of the alternatives to synthetic fungicides have the advantage that they do not have preharvest intervals, so they can be used right before the harvest. The sections below report on the control of postharvest decay and preservation of fruit quality through preharvest applications for some important crops, including strawberries, table grapes, and stone and pome fruit. Strawberries In the control of strawberry postharvest gray mold caused by B. cinerea, it is necessary to carry out preharvest treatments to prevent infection of the pathogen, which occurs mainly on the flowers in the field. B. cinerea remains latent on infected stamens underneath the sepals until fruit ripening, and then close to or after harvest it turns from a saprophyte into a parasite [28]. Fungicides are applied around flowering, and are repeated up to harvest, depending on weather conditions and the preharvest interval of the formulations used. Because of the frequent applications of fungicides, the development of resistant strains is quite common in strawberry fields [29]. In organic agriculture, some trials have reported that strawberries treated with chitosan at full bloom, or at the greenfruit or whitening-fruit stages have shown decreased gray mold and Rhizopus rot infections, and that this decay control was equal to or better than that achieved by standard fungicides [30–32]. Recently, some experimental trials were carried out at the postharvest stage to select natural compounds other than chitosan. These have included benzothiadiazole, oligosaccharides, soybean lecithin, calcium and organic acids, and Abies sibirica and Urtica dioica extracts, for their use as alternatives to synthetic fungicides for the control of postharvest rot of strawberry, and these might be used in future preharvest trials [13]. Table grapes In table grapes grown according to the standards of conventional agriculture, preharvest treatments are usually carried out four times during the season: at berry set, before bunch closure, at veraison, and three weeks before harvest. Preharvest fungicide regimes can significantly reduce subsequent postharvest decay, as the residual fungicide content deposited on berries has an influence on their postharvest rot [14]. Usually, table grapes are stored in the presence of sulfur dioxide pads, which allow the storage to be prolonged. However, for postharvest use on grapes, in the USA, sulfur dioxide is clas3


sified as a pesticide. Furthermore, at high rates, sulfur dioxide fumigation can cause grape injury, as seen by bleaching of the berry skin. In organic agriculture, field applications with alternatives to synthetic fungicides have been studied in terms of natural antimicrobials, decontaminating agents, and antagonistic microorganisms. Calcium chloride and potassium or sodium salts applied in the field are among the compounds known to reduce table grape postharvest rot [14, 33]. Chitosan applied alone in the field [14, 34] or in combination with the microorganism Cryptococcus laurentii [35] can significantly reduce natural decay of table grapes stored at 0°C. In other trials, over 6 weeks of cold storage, the losses due to gray mold rots were reduced by 50% in ethanol-treated and calcium-chloride-treated bunches, compared to untreated controls. The treatments did not induce significant changes in the fruit quality or damage to the vines and clusters [36]. Stone fruit For stone fruit, field treatments carried out to control postharvest rot that is mainly caused by Monilinia spp. consist of one to three treatments with synthetic fungicides: the first around flowering, for varieties that are very susceptible and in areas prone to infection, and the second and/or third treatment before harvest, also considering climate conditions, the intervals between treatments, and the preharvest interval of the formulation use. Field trials have been carried out to test the effectiveness of compounds that are alternatives to synthetic fungicides. For example, with sweet cherries treated 7 days before the harvest with 0.1%, 0.5% and 1% chitosan there was decreased gray mold and brown rot after 2 weeks of storage at 0°C followed by 7 days of shelf life, as compared to the untreated control. At the highest chitosan concentration, the decay reduction was no different with respect to that seen after application of tebuconazole [37]. Similar results were obtained when 1% chitosan was applied 3 days before harvest, as this reduced sweet cherry postharvest disease to a level comparable to that obtained with the synthetic fungicide fenhexamid [8]. Although less effective, fir and nettle extracts reduced the postharvest rots of sweet cherry. Pome fruit For pome fruit, treatments with the systemic fungicide thiabendazole are the main means of defense against the pathogen Phlyctema vagabunda, which causes lenticel rot. These treatments are carried out one week before harvest, when this systemic fungicide can control the pathogen that is quiescent in the lenticels. However, thiabendazole is no longer effective in reducing green mold, which is another important pathogen of pome fruit, because of the development of pathogen resistance. Instead, two or three weeks before the harvest, contact fungicides that are active against green mold can be used to prevent the penetration of the pathogen P. vagabunda into the lenticels. Alternatively, selected biocontrol yeasts are very interesting candidates for use in integrated control strategies that are aimed at reducing the use of fungicides [25]. Among antimicrobial microorganisms that are effective for pome fruit, some are commercially available or have been

studied for many years, such as: C. oleophila, B. amyloliquefaciens, P. syringae, C. laurentii, and Rhodotorula glutinis [18, 38].

Conclusions Depending on the characteristics of the commodity and on the specific situation, one strategy of controlling postharvest decay of fruit can be more appropriate than another. To overcome the drawbacks that can arise with the use of a unique strategy, integration of the methods used might provide additive or synergistic effects for disease control [39]. The ideal formulation, either as a fungicide or a natural compound, that can be applied at the preharvest stage to control postharvest rot of fruit and vegetables should meet the following criteria [6]: • Consistent effectiveness against the pathogen(s); • No injury or phytotoxic effects on the fruit; • Low toxicity to human health and the environment; • Residues that persist after harvest to protect the fruit during storage; • Compatible with standard practices, and affordable and easy to implement. In future, new insights, even at the molecular level, into the modes of action of the chemicals applied, either as fungicides, natural antimicrobials or resistance inducers, will be helpful to better understand the changes that occur in the host plants following treatments, or the effects of the treatments on the pathogen and on the interaction of the host-plant chemicals. In addition, the recent availability of the sequence of the entire genome of some of the postharvest pathogens [40] could open a new scenario in our understanding of the mechanisms of infection of fungi. This new knowledge could optimize the application of chemical treatments to get the highest effects with the minimum number/ time/ concentration of the applications.

Acknowledgment The financial support was carried out within the projects “Pre and postharvest treatments to control storage decay of sweet cherries”, granted by Marche Polytechnic University and “EUBerry Project” (EU FP7 KBBE 2010-4, Grant Agreement No. 265942).

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