Essential Oils in Postharvest Treatment against Microbial Spoilage of the Rosaceae Family Fruits
Essential Oils in Postharvest Treatment against Microbial Spoilage of the Rosaceae Family Fruits
Magri, Anna;Curci, Martina;Battaglia, Valerio;Fiorentino, Antonio;Petriccione, Milena
2023-04-03 00:00:00
Review Essential Oils in Postharvest Treatment against Microbial Spoilage of the Rosaceae Family Fruits 1 , 2 , 2 3 1 2 Anna Magri * , Martina Curci , Valerio Battaglia , Antonio Fiorentino and Milena Petriccione Department of Environmental, Biological and Pharmaceutical Sciences and Technologies (DiSTABiF), University of Campania Luigi Vanvitelli, Via Vivaldi 43, 81100 Caserta, Italy; antonio.fiorentino@unicampania.it Council for Agricultural Research and Economics (CREA), Research Center for Olive, Fruits, and Citrus Crops, Via Torrino 3, 81100 Caserta, Italy; martina.curci@crea.gov.it (M.C.); milena.petriccione@crea.gov.it (M.P.) Council for Agricultural Research and Economics (CREA), Research Centre for Cereal and Industrial Crops, Via Torrino 3, 81100 Caserta, Italy; valerio.battaglia@crea.gov.it * Correspondence: anna.magri@unicampania.it Abstract: Fruits undergo numerous chemical, physical, and microbiological changes during storage that shorten their postharvest life, reducing shelf-life and boosting food loss. Food quality and safety are seriously threatened by postharvest infections, one of the factors behind postharvest deterioration and mycotoxin contamination in fruits. The control of postharvest deterioration is a big concern because there are few management methods available. Several attempts have been undertaken to prevent the microbial degradation of fresh food at the postharvest stage without using synthetic fungicides, which are dangerous for the environment and people’s health. A good substitute for synthetic fungicides among them is the use of natural plant compounds, such as essential oils included or not included in the edible coatings. This review’s aim was to collect information from the scientific literature on the biological activity of essential oil, with or without edible coatings, against pathogens that cause the postharvest spoilage of many fruit belonging to Rosaceae family in order to develop appropriate substitute tactics for synthetic fungicides in the treatment of postharvest fruit Citation: Magri, A.; Curci, M.; diseases. Advances and obstacles surrounding emerging methods that may be useful for enhancing Battaglia, V.; Fiorentino, A.; the effectiveness and dependability of essential oils were evaluated. Petriccione, M. Essential Oils in Postharvest Treatment against Keywords: postharvest fungi; phytochemicals; botanicals; plants extract; postharvest disease stone Microbial Spoilage of the Rosaceae and pome fruits Family Fruits. AppliedChem 2023, 3, 196–216. https://doi.org/10.3390/ appliedchem3020013 Academic Editors: Antony C. 1. Introduction Calokerinos, Nikos Lydakis-Simantiris Fruits are abundant in vitamins, minerals, fiber, and other nutrients that the hu- and Spyros Grigorakis man body needs, and their consumption is growing worldwide. However, postharvest Received: 9 March 2023 pathogens appear to be the primary cause of the fruit’s reduced shelf-life and loss, also Revised: 24 March 2023 leading to financial losses [1]. Most postharvest pathogens are unable to penetrate directly Accepted: 28 March 2023 through the fruit cuticle, requiring a wound for their penetration. In general, these wounds Published: 3 April 2023 are inflicted during harvest, transport, packing, and storage. Several postharvest pathogens have been recorded in pome and stone fruit of the Rosaceae family, including rot caused by Rhizopus stolonifer [2–5], brown rot caused by Monilinia fructicola, M. fructigena, and M. laxa [2,5–10], gray mold caused by Botrytis cinerea [2,3,9–11], anthracnose caused by Copyright: © 2023 by the authors. Colletotrichum gloeosporioides and C. acutatum [2–4,12,13], blue mold caused by Penicillium Licensee MDPI, Basel, Switzerland. expansum [2,11,14,15], and bull’s eye rot caused by Neofabraea vagabunda [14], which are the This article is an open access article main aggressive diseases that cause postharvest rot. In addition, other pathogens have distributed under the terms and been described, such as Diaporthe spp, Neofusicoccum yunnanense, and Diplodia spp. in conditions of the Creative Commons apple [16], Penicillium spp., Talaromyces minioluteus, and T. rugulosus in pear [15], Mucor Attribution (CC BY) license (https:// piriformis, Rhizoctonia solani, Phytophthora spp., and Fusarium oxysporum in strawberry [3,12], creativecommons.org/licenses/by/ Gibberella avenacea and Alternaria alternata in apple and peach [12], Pestalotiopsis clavispora in 4.0/). AppliedChem 2023, 3, 196–216. https://doi.org/10.3390/appliedchem3020013 https://www.mdpi.com/journal/appliedchem AppliedChem 2023, 3 197 loquat [17], and Aspergillus niger, Cladosporium herbarum, Stemphylium spp., Geotrichum spp., Sclerotinia spp., M. piriformis, Coryneum carpophilum, and Fusarium spp. in stone fruits [18]. Synthetic antimicrobial substances are usually used to eradicate or slow the growth of pathogens that cause postharvest food deterioration. Since they present several toxicologi- cal challenges and are not safe for human consumption, the use of chemical antimicrobial agents as food preservatives has worried consumers [19]. For this reason, during the past two decades, the scientific community aimed to implement knowledge of natural antimicrobial agents such as essential oils (EOs), studying their distinctive physicochemical characteristics and wide range of biological activities [20]. As a potential alternative for industrial chemicals, the U.S. Food and Drug Administration (USFDA) classifies them as Generally Recognized as Safe (GRAS) [21]. EOs can be extracted from different plant organs, including fruits, bark, seeds, pulp, peel, roots, and, in some cases, entire plants [22]. Currently, EOs are used in the food industry as flavoring agents, but their potential as a natural food-grade preservative has not been properly investigated. Several studies have been carried out in in vitro settings. The concentration of EOs and associated con- stituents, which are necessary to prevent microbial development, is often greater in food than in culture media. Often, the quantities that are beneficial against the disease may nevertheless occasionally be phytotoxic and lead to the additional chemical degradation of treated food. New strategies, such as inclusion in nanoparticles or coatings, to reduce the impact of essential oils on the fruit should be investigated. Due to their inherent antimi- crobial characteristics, EOs could offer a useful tool for food preservation [23]. The EOs are made up of several different classes of chemicals, containing aromatic hydrocarbons (phenylpropanoids), terpenes (monoterpenes and sesquiterpenes), esters, lactones, alcohols, aldehydes, and ketones [24]. Due to the intricacy of their chemical composition, it is difficult to identify the most effective molecule responsible for the antimicrobial action of EOs; this propriety is probably correlated to the synergistic effect of all of the EO chemicals, and not just one molecule [25]. Many EOs were discovered to be helpful against gray mold and brown rot on nectarines, peaches, plums, pears, and strawberries [9,26–28]. Several studies reported the effectiveness of EOs against blue mold (P. expansum) [29,30], gray mold [31–34], and bitter rot [35], as well as Alternaria spp. and Fusarium spp. rots [36]. In this paper, the use of EOs, incorporated or not incorporated in edible coating, to prevent the microbial deterioration of fruits belonging to Rosaceae family during postharvest was reviewed. The results presented in this review could help the formulation of new and sustainable solutions for fruit storage based on EOs. 2. Postharvest Pathogens The main postharvest pathogens responsible for serious damage to stone and pome fruit, against which the antimicrobial activities of EOs have been assayed in vitro and in vivo, are shown in Table 1. Table 1. The main postharvest pathogens and their abbreviation and types of fruits and diseases [2–18]. Pathogens Abbreviations Fruit Disease Alternaria alternata (Fr.) Keissl. (Basionym: Torula alternata Fr.) AA Stone Brown spot Aspergillus carbonarius (Bainier) Thom (Basionym: Sterigmatocystis carbonaria Bainier) AC Pome Bunch rot Aspergillus flavus Link AF Stone Green mold Aspergillus niger Tiegh. AN Stone Black mold Botrytis cinerea Pers. BC Pome–Stone Gray mold Colletotrichum acutatum J.H. Simmonds CA Pome–Stone Antrachnose Colletotrichum gloeosporioides (Penz.) Penz. & Sacc. (Basionym: Vermicularia gloeosporioides Penz.) CG Pome–Stone Antrachnose Colletotrichum nymphaeae (Pass.) Aa (Basionym: Ascochyta nymphaeae Pass.) CN Stone Antrachnose Monilinia fructicola (G. Winter) Honey (Basionym: Ciboria fructicola G. Winter) MFC Stone Brown rot Monilinia fructigena (Pers.) Honey (Basionym: Torula fructigena Pers.) MFG Pome–Stone Brown rot Monilinia laxa (Aderh. & Ruhland) Honey (Basionym: Sclerotinia laxa Aderh. & Ruhland) ML Pome–Stone Brown rot Mucor piriformis A. Fisch. MP Stone Brown rot Penicillium expansum Link PE Pome–Stone Blue mold Penicillium notatum Westling (current name: Penicillium chrysogenum Thom) PN Stone Green mold Rhizopus microsporus Tiegh. RM Stone Rotten spots Rhizopus stolonifer (Ehrenb.) Vuill. (Basionym: Mucor stolonifer Ehrenb.) RS Pome–Stone Rotten spots Trichothecium roseum (Pers.) Link (Basionym: Trichoderma roseum Pers.) TR Pome Core rot AppliedChem 2023, 3 198 3. EOs’ Chemical Composition Botanical active ingredients for plant-based goods are obtained differently from syn- thetic chemical active ingredients. Whereas botanical active ingredients are obtained via the processing of biologically derived material, synthetic chemicals are created through chemical processes [37]. In botanicals, there is a subcategory that includes EOs, defined according to the U.S. Environmental Protection Agency (EPA): “Essential oils are mixtures of natural substances that come from various parts of plants, such as flowers, fruits, and wood. They are responsible for the distinctive odor or flavor of the plant they come from”. Plant-derived EOs are thought to be non-phytotoxic substances and may be useful as phytochemicals to protect crops. They include intricate combinations of active secondary metabolites that have antibacterial, allelopathic, antioxidant, and bioregulatory activities [38]. The number of molecules, the stereochemical characteristics, and the extraction method affect the chemical composition of EOs. The quality, quantity, and content of the extraction products may vary according to the environment, soil composition, plant organs, age, and vegetative cycle stage (chemotypes) [39]. Due to their verified safety, widespread public acceptability, and potential for several purposes, EOs from various plants are garnering interest. Over 300 distinct chemicals constitute EOs, although most of them are volatile sub- stances with molecular weights below 1000 Da (often 300 Da). In contrast to other com- pounds, which are present in trace levels, a select few chemicals are essentially present as large ones at roughly 20–70% (Table 2) [40]. A large class of chemicals in EOs that are known to have antimicrobial action are terpenes and terpenoids. Terpenes represent a wide group of hydrocarbons, characterized by different chemical structures, constructed from isoprene units C . They are classified into numerous categories based on the number of isoprenic units, including emiterpenes (C ), monoterpenes (C ), sesquiterpenes (C ), diterpenes (C ), sesterterpenes (C ), triterpenes 10 15 20 25 (C ), and tetraterpenes (C ). 30 40 Terpenoids are also found in EOs. They are defined as terpene molecules with a methyl group that has been transposed or eliminated. The terpenoids can be oxidized and found as esters, aldehydes, ketones, alcohols, ethers, and epoxides. Approximately 90% of EOs include monoterpenes, which make up most of their constituent parts. These often have a nice odor, and they are volatile. The monoterpenes are significantly involved in the synthesis of a wide range of structures. Carvacrol, geraniol, eugenol, thymol, linalyl acetate, linalool, citronellal, citronellol, and terpineol are examples of terpenes and terpenoids (Figure 1). Thymol and carvacrol are the two main components of thyme and oregano EOs. Bouchra et al. [41] demonstrated that the essential oil of Origanum compactum Benth., which contains carvacrol (58.1%), p-cymene (11.4%), and thymol (9.0%) as major components, was more toxic against BC. Additionally, according to Pérez-Alfonso et al. [42], thymol and carvacrol demonstrated substantial antifungal efficacy against postharvest pathogens such as Penicillium digitatum, P. italicum, Fusarium spp., and Aspergillus spp. Citronellol, octanal, citral, decanal, nonanal, b-pinene, linalool, and g-terpinene, present at a high quantity in citrus EOs, showed antifungal properties against P. italicum, whereas octanal, decanal, nonanal, limonene, citral, a-terpineol, and linalool showed antifungal properties against P. digitatum [43]. According to Cai et al. [44], menthol caused serious membrane lipid peroxidation and the generation and accumulation of reactive oxygen species in Geotrichum candidum var. citri-aurantii, which led to the breakdown of the integrity of the plasma membrane of the pathogen. AppliedChem 2023, 3, FOR PEER REVIEW 4 found as esters, aldehydes, ketones, alcohols, ethers, and epoxides. Approximately 90% of EOs include monoterpenes, which make up most of their constituent parts. These often have a nice odor, and they are volatile. The monoterpenes are significantly involved in the synthesis of a wide range of structures. Carvacrol, geraniol, eugenol, thymol, linalyl ace- tate, linalool, citronellal, citronellol, and terpineol are examples of terpenes and terpenoids (Figure 1). Thymol and carvacrol are the two main components of thyme and oregano EOs. Bouchra et al. [41] demonstrated that the essential oil of Origanum compactum Benth., which contains carvacrol (58.1%), p-cymene (11.4%), and thymol (9.0%) as major compo- nents, was more toxic against BC. Additionally, according to Pérez-Alfonso et al. [42], thy- mol and carvacrol demonstrated substantial antifungal efficacy against postharvest path- ogens such as Penicillium digitatum, P. italicum, Fusarium spp., and Aspergillus spp. Cit- ronellol, octanal, citral, decanal, nonanal, β-pinene, linalool, and γ-terpinene, present at a high quantity in citrus EOs, showed antifungal properties against P. italicum, whereas octanal, decanal, nonanal, limonene, citral, α-terpineol, and linalool showed antifungal properties against P. digitatum [43]. According to Cai et al. [44], menthol caused serious membrane lipid peroxidation and the generation and accumulation of reactive oxygen species in Geotrichum candidum var. citri-aurantii, which led to the breakdown of the integ- rity of the plasma membrane of the pathogen. AppliedChem 2023, 3 199 Figure 1. The chemical structures of significant and major terpenoids and monoterpenes contained in EOs: (a) carvacrol, (b) ( )-limonene, (c) thymol, (d) geraniol, (e) ( )-linalool, (f) (+)-citronellol, Figure 1. The chemical structures of significant and major terpenoids and monoterpenes contained (g) ( )-a-terpineol, (h) b-caryophyllene, (i) p-cymene, (l) ( )-menthol, (m) (+)-sabinene, and (n) 1,8- in EOs: (a) carvacrol, (b) (−)-limonene, (c) thymol, (d) geraniol, (e) (−)-linalool, (f) (+)-citronellol, (g) cineole. (−)-a-terpineol, (h) β-caryophyllene, (i) p-cymene, (l) (−)-menthol, (m) (+)-sabinene, and (n) 1,8-cine- The second category of compounds mainly present in EOs was phenylpropanoids, ole. which consist of aromatic substances such as benzyl, phenylethyl, and phenylpropyl. They are biosynthesized by the shikimate pathway. Typically, they have a benzene ring (C H ) 6 6 The second category of compounds mainly present in EOs was phenylpropanoids, bonded to a three-carbon chain. Important phenylpropanoids and derivatives are anethole, estragole, cinnamaldehyde, eugenol, safrole, apiol, vanillin, and benzaldehyde (Figure 2). which consist of aromatic substances such as benzyl, phenylethyl, and phenylpropyl. Kalleli et al. [45] demonstrated that fennel essential oil (composed of 78% trans-anethole) They are biosynthesized by the shikimate pathway. Typically, they have a benzene ring reduced the severity of the infection caused by Fusarium oxysporum f. sp. lycopersici by (C6H6) bonded to a three-carbon chain. Important phenylpropanoids and derivatives are 98% in treated plants with the oil, eight weeks after inoculation. The cinnamaldehyde anethole, estragole, cinn andam estragole aldehhad yde, inhibitory eugenol activity , safro against le, api Pseudomonas ol, vanillisyringae n, and pv ben . actinidiae zaldehyde at a con- centration of 1.250 and 2.500 ppm [46]. The Ocimum sanctum L. essential oil, which has (Figure 2). Kalleli et al. [45] demonstrated that fennel essential oil (composed of 78% trans- a wide range of fungitoxic properties and was also found to be effective in lowering the anethole) reduced the severity of the infection caused by Fusarium oxysporum f. sp. lyco- incidence of AF in raw materials (roots) of Rauvolfia serpentina (L.) Benth. Ex Kurz during persici by 98% in treated plants with the oil, eight weeks after inoculation. The cinnamal- storage, was found to be primarily composed of eugenol, followed by b-caryophyllene and germacrene D [47]. The growth of CA, CG, and Botryodiplodia theobromae, three important dehyde and estragole had inhibitory activity against Pseudomonas syringae pv. actinidiae at postharvest pathogens of fruits, was reduced in vitro by the EOs of Piper auritum Kunth a concentration of 1.250 and 2.500 ppm [46]. The Ocimum sanctum L. essential oil, which and P. holtonii C.DC., which were primarily composed of safrole and apiol [48]. Vanillin has a wide range of fungitoxic properties and was also found to be effective in lowering treatment in vivo and in vitro increased the BC and AA membrane permeability, inhibiting the pathogenicity-related enzyme activities and reducing the infection ability of fungi [49]. the incidence of AF in raw materials (roots) of Rauvolfia serpentina (L.) Benth. Ex Kurz dur- ing storage, was found to be primarily composed of eugenol, followed by β-caryophyllene AppliedChem 2023, 3 200 Table 2. Plant material for EOs extraction and main components [50]. Scientific Name Common Name Abbreviation Organs Principal Compounds Acorus calamus L. Calamus oil CaEO Flowers and leaves a-asarone, (E)-methylisoeugenol, methyleugenol and b-asarone Allyl methyl trisulfide, diallyl disulfide, diallyl trisulfide, and allyl Allium sativum L. Garlic oil GEO Bulb methyl disulfide Aloysia citriodora Kunth. Lemon Verbena oil LVEO Leaves Geranial, neral, spathulenol, and limonene Alpinia officinarum Hance Galangal oil GalEO Rhizomes 1,8-cineole, a-fenchyl acetate, b-myrcene, and b-ocimene Artemisia argyi H.Lév. & Vaniot Artemisia oil ArEO Leaves Eucalyptol, b-pinene, camphor, and artemisia ketone Cinnamomum zeylanicum L. Cinnamon oil CEO Barks Cinnamaldehyde, benzaldehyde, and trans-cinnamyl acetate Citrus aurantiifolia L. Lime oil LiEO Fruits Limonene, a-terpineol, terpinen-4-ol, and 1,8-cineole Citrus limon L. Lemon oil LeEO Fruits Limonene, citrale, citronellal, geranyl acetate Germacrene D, terpinolene, (Z)-b-ocimene, and Cortex dictamni Turcz. Cortex oil CoEO Leaves b-caryophyllene (7.74%) Cymbopogon citratus L. Lemongrass oil LgEO Leaves Citral, isoneral, isogeranial, and geraniol Cymbopogon winterianus Jowitt Cymbopogon martini (Roxb.) Wats. Palmarosa oil PalEO Flowers and leaves Geraniol, geranylacetate, farnesol, and nerolidol Eucalyptus globulus Labill. Eucalyptus camaldulensis Dehnh. Eucalyptus oil EuEO Leaves Eucalyptol, p-cymene, a-pinene, and b-limonene Eucalyptus staigeriana F.Muell. ex Bailey Foeniculum vulgare Mill. Fennel oil FEO Seeds and fruits Trans-anethol, fenchone, estragole, and limonene Laurus nobilis L. Laurel oil LaEO Leaves 1,8-cineole, sabinene, -pinene, and linalool Lavandula officinalis L. Lavander oil LEO Flowers and leaves Linalyl acetate, a-terpineol, and borneol Pepper-rosmarin Lippia sidoides Cham. PREO Leaves 1,8-cineole, sabinene, a-terpineol, and -pinene oil Malaleuca ericifolia Sm. Rosalina oil RosEO Leaves 1,8-cineole, -pinene,
-terpinene, and terpinen-4-ol Mentha piperita L. Peppermint oil PEO Leaves Menthol, menthone, menthofuran, and cis-carane Mucuna pruriens (E)-2-hexenal, linalool, 1-hexanol, and Mucuna pruriens (L.) DC. MPEO Leaves oil trans-dehydroxylinalool oxide Ocimum Ocimum americanum L. OAEO Leaves 1,8-cineole, (E)-
-bisabolene, -bisabolene, and eugenol americanum Origanum majorana L. Majorana oil MEO Flowers and leaves 4-terpineol, cis-thujan-4-ol, -terpinene, and -terpinene Origanum vulgare L. Oregano oil OEO Flowers and leaves Carvacrol, p-cymene, and linalool Pelargonium graveolens Geranium oil GaEO Flowers and leaves Citronellol, geraniol, citronellyl formate, and linalool Pimenta pseudocaryophyllus Pimenta oil PPEO Leaves Methyl eugenol, neral, geranial, and (E)–methyl isoeugenol (Gomes) L.R. Landrum Applied Rosmarinus Chem 202 officinalis 3, 3, FOR L. PEER REVIEW Rosemary oil REO Flowers and leaves p-cymene, camphor, limonene, and myrcene 5 Salvia officinalis L. Sage oil SaEO Flowers and leaves a-thujone, b-thujone, camphor, and camphene Satureja hortensis L. Summer savory oil SSEO Flowers and leaves Carvacrol, o-cymene, linalool, and caryophyllene oxide Sesamum indicum L. Sesamum oil SeEO Leaves and seeds Linoleic acid, oleic acid, and palmitic and stearic acid Syringa vulgaris L. Lilac oil LLEO Flowers and leaves Ocimene, benzyl methyl ether, 1,4-dimethoxybenzene, and indole Syzygium aromaticum ((L.) Merr. & and germacrene D [47]. The growth of CA, CG, and Botryodiplodia theobromae, three im- L.M.Perry, 1939) Clove oil ClEO Leaves and seeds Eugenol, -caryophyllene, -humulene, and eugenyl acetate Eugenia caryophyllata portant postharvest pathogens of fruits, was reduced in vitro by the EOs of Piper auritum Thymus vulgaris L. Thyme oil TEO Flowers and leaves Thymol, cis-3-hexenyl acetate, p-cymene, and carvacrol Kunth and P. holtonii C.DC., which were primarily composed of safrole and apiol [48]. Verbena officinalis L. Vervain oil VEO Flowers and leaves Limonene, 1,8-cineole, pathuleno1, and caryophyllene oxide Zanthoxylum Vanillin treatment in vivo and in vitro increased the BC and AA membrane permeability, Zanthoxylum bungeanum Maxim ZBEO Flowers and leaves Terpineol-4-ol, (-)- -pinene,
-terpinene, and terpinyl acetate bungeanum oil inhibiting the pathogenicity-related enzyme activities and reducing the infection ability Zataria multiflora Boiss Zataria oil ZMEO Flowers and leaves Thymol, carvacrol, p-cymene, and g-terpinene Mentha spicata L. Mint oil MiEO Flowers and leaves Linalyl acetate, linalool, carvone, and limonene of fungi [49]. Litsea cubeba Pers. Litsea oil LitEO Flowers and leaves Geranial, neral, limonene, and -thujene Figure 2. The chemical structures of significant and major phenylpropanoids and derivatives con- Figure 2. The chemical structures of significant and major phenylpropanoids and derivatives con- tained in EOs: (a) anethole, (b) estragole, (c) cinnamaldehyde, (d) eugenol, (e) safrole, (f) apiol, tained in EOs: (a) anethole, (b) estragole, (c) cinnamaldehyde, (d) eugenol, (e) safrole, (f) apiol, (g) (g) vanillin, and (h) benzaldehyde. vanillin, and (h) benzaldehyde. Table 2. Plant material for EOs extraction and main components [50]. Scientific Name Common Name Abbreviation Organs Principal Compounds Flowers and α-asarone, (E)-methylisoeugenol, Acorus calamus L. Calamus oil CaEO leaves methyleugenol and β-asarone Allyl methyl trisulfide, diallyl disul- Allium sativum L. Garlic oil GEO Bulb fide, diallyl trisulfide, and allyl methyl disulfide Lemon Verbena Geranial, neral, spathulenol, and limo- Aloysia citriodora Kunth. LVEO Leaves oil nene 1,8-cineole, α-fenchyl acetate, β-myr- Alpinia officinarum Hance Galangal oil GalEO Rhizomes cene, and β-ocimene Artemisia argyi H.Lév. & Eucalyptol, β-pinene, camphor, and ar- Artemisia oil ArEO Leaves Vaniot temisia ketone Cinnamomum zeylanicum Cinnamaldehyde, benzaldehyde, and Cinnamon oil CEO Barks L. trans-cinnamyl acetate Limonene, α-terpineol, terpinen-4-ol, Citrus aurantiifolia L. Lime oil LiEO Fruits and 1,8-cineole Limonene, citrale, citronellal, geranyl Citrus limon L. Lemon oil LeEO Fruits acetate Germacrene D, terpinolene, (Z)-β-oci- Cortex dictamni Turcz. Cortex oil CoEO Leaves mene, and β-caryophyllene (7.74%) Cymbopogon citratus L. Citral, isoneral, isogeranial, and gera- Cymbopogon winterianus Lemongrass oil LgEO Leaves niol Jowitt AppliedChem 2023, 3 201 4. Essential Oils’ Effects on Human Health and the Environment In addition to increasing costs, handling risks, worries about plant protection product (PPP) residues in food, and threats to human health and the environment, the extensive use of PPPs has serious negatives [51]. Finding better substitutes for PPPs made of synthetic chemicals has become more popular because of growing public awareness of these con- cerns. In line with the current trend of investigating new options for extending storage life, techniques that reduce horticultural product degradation while minimizing negative effects on human health or the environment must be prioritized [52]. Natural plant additives with pesticidal and antimicrobial action are one such alternative; they tend to be less poi- sonous to mammals, have fewer negative impacts on the environment, and are well-liked by the general population. EOs and other natural plant products are regarded as more biodegradable than synthetic substances. There are significant worries about the potential negative impacts of EOs on human health and the environment, which are still under inves- tigation. EOs are extracted using a variety of contemporary and conventional procedures, including steam distillation, hydrodistillation, ultrasonic, ohmic, and microwave-aided hydrodistillation, and supercritical fluid extraction [53,54]. Conventional techniques have a few drawbacks, including a lengthy extraction process, low efficiency, solvent toxicity, and a detrimental effect on the environment. Modern techniques are now preferred over traditional approaches or extraction methods because they are quicker, more effective, use less energy, and produce more EOs with a low negative effect on the environment [55]. To date, there are more studies describing the beneficial effects of EO consumption on human health than there are studies attesting to their toxicity. EOs are typically known to be non-toxic for humans and other mammals, as they are quickly absorbed and processed in the liver and eliminated by the kidneys [56]. Ezeorba et al. [57] reported that garlic EOs possessed interesting medical properties, such as antimicrobial, cardio-protective, anti-cancer, anti-Alzheimer, anti-diabetic, and immunomodulatory activities. According to studies by Loizzo et al. [58] and Koul et al. [59], limonene, which is the compound that most characterizes lemon and citron EOs, had lymphotonic, powerful diuretic, healing, scalping tonic, antifungal, anti-moth decongestant, antiseptic respiratory, purgative, relaxing, and comforting actions. Menthol and menthone, mainly found in mint essential oil, has shown tonic, stimulant, decongestant, anesthetic and analgesic, cooling, and anti-inflammatory properties [60]. Cinnamon essential oil, particularly cinnamaldehyde, was described as a potent antibacterial, antifungal, antiviral, anticoagulant, and uterine tonic [61,62]. 5. Use of Essential Oils to Control Postharvest Microbial Spoilage The antimicrobial properties of EOs have been described in several studies. EOs have been shown to have a wide range of antifungal properties and have the advantage of being environmentally friendly [59,63,64]. EOs are natural compounds that show antifungal activity, and they can be used instead of synthetic fungicides. Several studies demonstrated that EO activity appears to be stronger in vitro than in vivo, which is necessary to confirm the in vitro results with in vivo tests. There are several modes of action of EOs that make them natural fungicides, e.g., they interact with ergosterol, which is essential for maintaining cell integrity as well as fungal function, viability, and growth; they can pass through the lipid bilayer of the cell membrane, increasing permeability and causing cell death or the inhibition of fungal germination and sporulation; and phenolic compounds interact with porins, causing ion leakage and cell breakdown [65–67]. Due to their low permeability, uncontrolled volatility, and low solubility, the commer- cialization of EOs is restricted. To avoid these limitations, there are different strategies, such as developing proper formulation by the encapsulation of EOs in nanoemulsion or developing edible coatings [68,69]. EO concentrations require careful optimization de- pending on fruit crops. Although, in in vitro tests, the efficacy is directly correlated with the EO concentration, when applied to fruit, EO vapors could have phytotoxic effects, which reduce the treatment efficacy [9]. Many factors, such as the chemical composition, AppliedChem 2023, 3 202 environmental factors, targeted pathogens, botanical sources, and type of food to be treated can affect the antimicrobial action of EOs [70,71]. Below are studies on the effectiveness of EOs in vitro and in vivo on postharvest pathogens responsible for fruit decay and food wastage, divided by pome (apple and pear) and stone fruits (sweet cherry, apricot, peach, strawberry, and plum). 5.1. Pome Fruits Fresh fruits are a favorable habitat for pathogenic and spoilage fungi and bacteria due to their concentration of nutrients, such as sugars, and the high moisture [72,73]. All bioassays cited in this paragraph are reported in Table 3. Table 3. Overview of antimicrobial activity of essential oils, with or without edible coating, in pome fruits. Fruit Pathogens EO Coating EOs Application Method Reference Aspergillus carbonarius Cinnamon Na-alginate coating Inclusion in coating [74] Oregano, savory, and thyme Dropped [27] Lime, lemon, and lemongrass Thermofogging treatment [31] Thyme and savory Dropped [26] Thyme Starch–gellan coating Dropped [75] Fennel, thyme, lavender, neem, Dropped [76] pennyroyal, salvia and asafetida cinnamon, pimento, and laurel Spray [33] Macro-sized bacterial Botrytis cinerea Cinnamomum zeylanicum, Zataria cellulose coating [77] multiflora, and Satureja khuzestanica emulsion Pelargonium graveolens Volatile activity test [78] Films were prepared Fennel oil Vapor [79] with SSOS, chitosan Thyme and savory Fumigation [34] Colletotrichum acutatum and Poly(lactic acid) Lemongrass Inclusion in nanocapsules [80] C. gloeosporioides nanocapsules Apples Eucalyptus radiata ssp. Mentha pulegium, Rosmarinus officinalis, Origanum Monilinia laxa and M. compactum, Lavandula angustifolia, Wound application [70] fructigena Syzygium aromaticum, Thymus vulgaris, Citrus aurantium, and Citrus sinensis Macro-sized bacterial Cinnamomum zeylanicum, Zataria cellulose coating [77] multiflora, and Satureja khuzestanica emulsion Pelargonium graveolens Volatile activity test [78] Penicillium expansum Films were prepared Fennel Vapor [79] with SSOS, chitosan Oregano, savory, and thyme Dropped [27] Chitosan and sodium Cinnamon Coating [81] alginate Exogenous EO decanal Dropped [82] Rhizopus stolonifer Pelargonium graveolens Volatile activity test [78] Films were prepared Trichothecium roseum Fennel Vapor [79] with SSOS, chitosan Aspergillus carbonarius Cinnamon Na-alginate coating Inclusion in coating [74] Botrytis cinerea Cinnamon, rosemary, and marjoram EOs Dropped [83] Pears Cinnamon, rosemary, and marjoram EOs Dropped [83] Penicillium expansum Eucalyptus and rosemary Vapor [28] Exogenous EO decanal Dropped [82] The major fungal pathogens that have been associated with losses in the quality of pectin-rich fruits, such as pear and apple fruits, include PE, the causal agent of blue mold, and BC, the causing factor of gray mold [84–87]. The antifungal effects, both in vitro and in vivo, of the rosemary, eucalyptus, thyme, cinnamon, and marjoram EOs and their combination against BC and PE, as well as their ef- fects on quality attributes, were highlighted, and their possible use as natural fungicides in the fresh fruit industry for postharvest disease control was encouraged [28,83]. Specifically, 1 1 the in vitro treatment with 300 L L eucalyptus EO (EuEO,) 300 L L rosemary EO (REO), and their mixture revealed a decrease in spores germination and fungal colonies of PE. Their application in vapor form on fresh pears in vivo led to an increase in the AppliedChem 2023, 3 203 respiration rate and a reduction in the growth of PE lesions [28]. Nikkhah [83] investi- gated the antifungal activity of thyme, cinnamon, rosemary, and marjoram EOs and the synergistic activity of their combinations. In vitro, thyme EO (TEO) and cinnamon EO (CEO) demonstrated the highest antifungal activity and the lowest minimum inhibitory concentration (MIC) against PE and BC. A remarkable synergistic interaction was observed in the thyme/cinnamon dual combinations against both mold species. Fractional inhibitor concentration analysis (FICi) and in vivo treatments indicated that the triple combination of cinnamon/majoran/thyme produced the most synergistic antifungal effect. In accordance with the previously mentioned study, in vitro experiments demonstrated that the CEO significantly inhibits PE. In vivo, the multi-layer coating, formed by chitosan (1% w/v), sodium alginate (1% w/v), and CEO (4% v/v), showed a more significant and lasting inhibition of PE in two varieties of apples, ‘Guoguang’ and ‘Huangyuanshuai’, compared to a single-layer coating with chitosan and CEO or sodium alginate and CEO [81]. PE produces patulin, a mycotoxin hazardous to consumer health. Zhou et al. [82] demonstrated that exogenous EO decanal in in vitro tests (0.12 g L ) reduced the PE growth and, in in vivo tests (0.24 g L ), controlled the effect against blue mold rot on apples and pears; moreover, EO decanal inhibited patulin production. Apple is one of the most important fruits in international trade and can suffer severe postharvest losses during transport and storage. BC, which causes gray mold, is one of the main pathogens causing postharvest losses [88]. The use of TEO at a 1% v/v concentration showed high efficiency in reducing the damage and incidence of BC symptoms and could be used as an effective postharvest treatment in stored apples. Furthermore, treatment with TEO increases the expression of the specific pathogenesis-related gene PR-8, suggesting an induction of resistance against BC through the priming of defense responses in apple fruit [26]. In vitro, the application of Zataria EO (ZMEO, 300–1200 L L ) and CEO (300–1200 L L ) presented inhibitory effects on BC and PE. This result was confirmed by in vivo tests, where the lesion diameter in inoculated apples was reduced by the ZMEO 1 1 (600–1200 L L ) and CEO (600–1200 L L ) [77]. Sadat Razavi et al. [77] demonstrated that BCNCs/GelA-CEO (75–2400 L L ) emulsions decreased BC and PE lesions on AppliedChem 2023, 3, FOR PEER REVIEW 11 inoculated apples, with a better performance of non-encapsulated EO. The strongest effect was shown at the highest concentration of CEO, with which lesions on the fruit skin were barely detectable (Figure 3). Figure 3. Botrytis cinerea and Penicillium expansum lesions on inoculated apples treated with Figure 3. Botrytis cinerea and Penicillium expansum lesions on inoculated apples treated with cin- cinnamon essential oil. Reproduced with permission from Sadat Razavi, M.; Golmohammadi, A.; namon essential oil. Reproduced with permission from Sadat Razavi, M.; Golmohammadi, A.; Nem- Nematollahzadeh, A.; Ghanbari, A.; Davari, M.; Carullo, D.; Farris, S. Foods 2022. atollahzadeh, A.; Ghanbari, A.; Davari, M.; Carullo, D.; Farris, S. Foods 2022. The incorporation of fennel EO (FEO) in a starch sodium octenyl succinate/chitosan film (mass ratio SSOS/glycerol/FEO/chitosan was 1:0:2:1:0.5) for apple fruit showed good The incorporation of fennel EO (FEO) in a starch sodium octenyl succinate/chitosan antifungal activity, in vivo and in vitro, against BC, TR, and PE. Moreover, the coating film (m reduced ass ratio the r S espiration SOS/glyrate cerol/FEO/ch and weight loss itosa wher n w eas as the 1:firmness, 0:2:1:0.5) peel for a lightness, pple solub fruitle showed good solids, and titratable acid content were maintained [79]. antifungal activity, in vivo and in vitro, against BC, TR, and PE. Moreover, the coating The treatment of the apple cultivar ‘Opal’ with 1% TEO and summer savory EO reduced the respiration rate and weight loss whereas the firmness, peel lightness, soluble (SSEO) by fumigation shows significant gray mold control after 60 days of storage at 1 C, solids, and titratable acid content were maintained [79]. The treatment of the apple cultivar ‘Opal’ with 1% TEO and summer savory EO (SSEO) by fumigation shows significant gray mold control after 60 days of storage at 1 °C, confirming the results obtained in vitro, where the application of thyme and savory EOs significantly reduced the growth of BC [34]. An application that allows for a uniform distribution of EOs and enables a small quantity to be used is the thermofogging treatment. Applying lime EO (LiEO), lemon EO (LeEO), and lemongrass EO (LgEO) on the ‘Golden Delicious’, ‘Pink Lady’, and ‘Granny Smith’ cultivars at a concentration of 0.016–1% results in a significant inhibition of the mycelial growth and spore germination of BC, resulting in a reduction in the decay of the apples [31]. In contrast, Sapper [75] evaluated that adding a starch–gellan (80:20 w/w) coating formulation with the incorporation of TEO (0.25 and 0.5 per g of polymer) directly or encapsulated in lecithin promotes the respiration rates and quotient in apples, but no antifungal action against BC was observed in vivo, despite in vitro tests and other studies proving the opposite. Geranium essential oil exhibits significant antifungal activity against RS, PE, and BC in vitro using poisoned food [78]. Despite in vitro tests demonstrating the effectiveness of CEO, pimenta EO (PPEO), and laurel EO (LaEO) against BC, these had no significant ef- fects in in vivo tests carried out on ‘Connel Red’ apples during storage, except for cinna- mon extract even at high concentrations [33]. Several studies have shown that the higher the concentration of the essential oil, the greater the inhibition of the mycelium growth of pathogenic fungi, but a concentration of 10% was found to be phytotoxic to fruit [27,80]. Furthermore, the efficiency of EOs in in- hibiting BC and PE depends not only on the concentration and type of oil but also on the cultivar and storage time [27]. The treatment of ‘Golden Delicious’, ‘Granny Smith’, ‘Red Chief’, and ‘Royal Gal’ apple cultivars with the emulsion at 1% of oregano EO (OEO), SSEO, and TEO dropped into the artificial wound inoculated with BC and PE showed a decrease in the lesion compared to the control [27]. Other pathogenic fungi causing postharvest disease in pome fruit are ML and MFG, causal agents of brown rot. These fungi cause blossom blight, twig cankers, fruit rot, and fruit mummification in the orchard [7]. EuEO, mint EO (MiEO), REO, OEO, lavander EO (LEO), clove EO (ClEO), TEO, Zan- thoxylum bungeanum EO (ZBEO), and OEO were tested, in vitro and in vivo, against ML and MFG. In vitro tests showed significant antifungal activity of EOs that reduced myce- −1 lial growth and spore germination. In in vivo tests, a concentration of 100 µ L mL of ClEO, TEO, ZBEO, and OEO reduced the brown rot incidence, spore germination, and lesion AppliedChem 2023, 3 204 confirming the results obtained in vitro, where the application of thyme and savory EOs significantly reduced the growth of BC [34]. An application that allows for a uniform distribution of EOs and enables a small quantity to be used is the thermofogging treatment. Applying lime EO (LiEO), lemon EO (LeEO), and lemongrass EO (LgEO) on the ‘Golden Delicious’, ‘Pink Lady’, and ‘Granny Smith’ cultivars at a concentration of 0.016–1% results in a significant inhibition of the mycelial growth and spore germination of BC, resulting in a reduction in the decay of the apples [31]. In contrast, Sapper [75] evaluated that adding a starch–gellan (80:20 w/w) coating formulation with the incorporation of TEO (0.25 and 0.5 per g of polymer) directly or encapsulated in lecithin promotes the respiration rates and quotient in apples, but no antifungal action against BC was observed in vivo, despite in vitro tests and other studies proving the opposite. Geranium essential oil exhibits significant antifungal activity against RS, PE, and BC in vitro using poisoned food [78]. Despite in vitro tests demonstrating the effectiveness of CEO, pimenta EO (PPEO), and laurel EO (LaEO) against BC, these had no significant effects in in vivo tests carried out on ‘Connel Red’ apples during storage, except for cinnamon extract even at high concentrations [33]. Several studies have shown that the higher the concentration of the essential oil, the greater the inhibition of the mycelium growth of pathogenic fungi, but a concentration of 10% was found to be phytotoxic to fruit [27,80]. Furthermore, the efficiency of EOs in inhibiting BC and PE depends not only on the concentration and type of oil but also on the cultivar and storage time [27]. The treatment of ‘Golden Delicious’, ‘Granny Smith’, ‘Red Chief’, and ‘Royal Gal’ apple cultivars with the emulsion at 1% of oregano EO (OEO), SSEO, and TEO dropped into the artificial wound inoculated with BC and PE showed a decrease in the lesion compared to the control [27]. Other pathogenic fungi causing postharvest disease in pome fruit are ML and MFG, causal agents of brown rot. These fungi cause blossom blight, twig cankers, fruit rot, and fruit mummification in the orchard [7]. EuEO, mint EO (MiEO), REO, OEO, lavander EO (LEO), clove EO (ClEO), TEO, Zanthoxylum bungeanum EO (ZBEO), and OEO were tested, in vitro and in vivo, against ML and MFG. In vitro tests showed significant antifungal activity of EOs that reduced mycelial growth and spore germination. In in vivo tests, a concentration of 100 L mL of ClEO, TEO, ZBEO, and OEO reduced the brown rot incidence, spore germination, and lesion diameter. Fruit quality parameters such as weight loss, total soluble solids, titratable acidity, firmness, and the maturity index were preserved during storage at 4 C for 20 days [69]. LgEO has shown antifungal activity against the two fungi responsible for apple bitter rot CA and GC. In vitro tests demonstrated that the use of poly(lactic acid) nanocap- sules containing LgEO at a dosage of 0.1% (v/v) resulted in fungicidal activity against both phytopathogenic fungi [80]. In vivo tests confirmed that apples treated with encap- sulated essential oil show significantly lower bitter rot lesions than those treated with non-encapsulated EO. This is because the active compounds are released more slowly if the EO is contained in nanocapsules and there is a reduction in the necrotic effects by essential oil terpenoids [80]. Adding Na-alginate coating supplemented with 0.3% and 0.9% v/v CEO to pear and apple slices or wounded skin limits the growth of the fungus AC and the production of ochratoxin A. This could be a strategy for limiting spoilage losses and preserving consumer health [74]. Additional studies showed the antimicrobial effect of EOs on total molds, yeast, and mesophilic microorganisms of pome fruit, the main results of which are reported in Table 4. Further analysis may be necessary to evaluate the efficacy against the postharvest pathogens mentioned in this review. AppliedChem 2023, 3 205 Table 4. Antimicrobial effect of EOs on microorganism responsible for fruit spoilage in pome fruits. Fruit Pathogen Coating EOs Results Reference Coatings were effective in inhibiting Alginate 1% and 2% or Citral 0.15, 0.3% Eugenol 0.1, 0.2% yeasts and molds and reducing ~2 logs Pectin 1% and 2% Total molds, yeast and [89] Ascorbic acid 1% Eugenol 0.1% + Citral CFU/g mesophilic bacteria during 8 days mesophilic Calcium chloride 1% 0.15% of storage compared to uncoated fruit. microorganisms Only EC with cinnamon 0.3% was capable Cassava starch 2% Cinnamon 0.1, 0.3% of inhibiting pathogens in vitro. EOs at [90] Glycerol 0.5% Fennel 0.1, 0.3% 0.3% provided better color in 4 days Tapioca starch 0.3% EO treatments reduced 4 log cycles of Apples + dHG1 0.3% mesophilic and psychrophilic bacteria and Glycerol 1.7% Cinnamon 0.2% [91] 3 log cycles reduction in yeasts and molds Ascorbic acid 1%/ after 12 days of storage. Calcium chloride 1% Alginate 1.29% Lower concentrations of EOs were more Total aerobic bacteria, Glycerol 1.5% effective in reducing total aerobic bacteria, Thyme 0.05%, 0.35% coliforms, yeasts and [92] Calcium chloride 2% coliforms, yeasts, and molds, increasing and 0.65% molds Ascorbic acid 1% shelf life in 8 days. Citric acid 1% Pectin 2%, Whey Sweet orange 0.1% and EOs at 0.15% slightly reduced (>1 log Total mesophilic protein 1%, Calcium [93] 0.15% cycle) total mesophilic counts. chloride 2% Treated pears displayed lower counts of Pears Total mould and yeasts - Seaweed 0.50% (w/v) [94] mold and yeasts. 5.2. Stone Fruits All bioassays cited in this paragraph are reported in Table 5. Table 5. Overview of antimicrobial activity of EOs, with or without edible coating, in stone fruits. Fruit Pathogens EOs Coating EOs Application Method Reference Monilinia fructicola Apricots Thymus vulgaris Eugenia caryophyllata - Spray [10] Botrytis cinerea Syringa EO - Inclusion in microcapsules [95] Alternaria alternata Alpinia officinarum, Cortex dictamni, and - Inclusion in microemulsion [96] Artemisia argyi Alpinia officinarum, Cortex dictamni, and Aspergillus flavus - Inclusion in microemulsion [96] Artemisia argyi Lemongrass and Thymus vulgaris - Wound application [97] Syringa vulgaris - Inclusion in microcapsules [95] Botrytis cinerea Arabic gum 10% and Coating Mint and thyme [98] Tween 20 Alpinia officinarum, Cortex dictamni, - Inclusion in microemulsion [96] and Artemisia argyi Rosewood EO - Vapor-phase and fumigation [99] Thymus vulgaris and verbena - Spray [100] Monilinia laxa - Vapor Thymus vulgaris and cinnamon [101] Thymus vulgaris and verbena - Spray [100] Thymus vulgaris and savory - Biofumigation [9] Chitosan 1.8%-Arabic Thymus vulgaris Coating and fumigation [102] gum 0.6% (w/v) M. fructicola Peaches/nectarines Chitosan 1.8%-Arabic Coating Thymus vulgaris [103] gum 0.6% (w/v) Tea tree EO - Vapor [104] Aloysia citriodora, Cymbopogon winterianus, Lippia alba, and Ocimum [105] americanum EOs Alpinia officinarum, Cortex dictamni, - Inclusion in microemulsion [96] and Artemisia argyi Thymus vulgaris and verbena - Spray [100] Mucor piriformis Rosewood EO - Vapor-phase and fumigation [99] Lemongrass and thyme EOs - Wound application [97] Arabic gum 10% and Penicillium expansum Mint and thyme EOs Coating [98] Tween 20 Alpinia officinarum, Cortex dictamni, Inclusion in microemulsion [96] and Artemisia argyi Rosewood EO - Vapor-phase and fumigation [99] Lemongrass and thyme EOs - Wound Application [97] Arabic gum 10% and Rhizopus stolonifer Mint and thyme EOs Coating [98] Tween 20 Alpinia officinarum, Cortex dictamni, - Inclusion in microemulsion [96] and Artemisia argyi Rosewood EO - Vapor-phase and fumigation [99] Aspergillus flavus Aspergillus niger - Fumigation [106] Plums Allium sativum Penicillium notatum Rhizopus microsporus AppliedChem 2023, 3 206 Table 5. Cont. Fruit Pathogens EOs Coating EOs Application Method Reference Aspergillus flavus Polyvinyl alcohol-b- Incorporation in nanofibers Botrytis cinerea Zanthoxylum bungeanum Cyclodextrin [107] Sweet cherries Penicillium (CGMCC films nanofiber film 3.13650) Collelotrichum Clove - Filter paper discs [108] gloeosporioides Polyvinyl alcohol-b- Aspergillus flavus Zanthoxylum bungeanum Cyclodextrin [107] nanofibers-film Salvia officinalis, [109] Zataria multiflora, and Cinnamomum Chitosan Dipping [110] zeylanicum Cinnamon Emulsion and nanoemulsion [111] Eucalyptus staigeriana and Eucalyptus Botrytis cinerea Carboxymethylcellulose Dipping [112] urograndis Basil Aloe vera gel coating Dipping [113] Aloysia citriodora, Cymbopogon winterianus, Lippia alba, and Ocimum [105] americanum Cymbopogon citratus, Thymus vulgraris, Vapor [114] and Origanum heracleoticum Eucalyptus camaldulensis, Mentha Vapor [115] piperita, Moringa oleifera FO CT thymol lemongrass, litsea, lavender, peppermint, mint, [116] petitgrain, sage, and thyme Polyvinyl alcohol-b- Zanthoxylum bungeanum Cyclodextrin [107] nanofibers-film Colletotrichum Allium sativum and Rosmarinus Vapor [117] Strawberries nymphaeae officinalis Thyme, cinnamon bark, and clove bud Fumigation [3] C. acutatum Aloysia citriodora, Cymbopogon winterianus, Lippia alba, and Ocimum [105] americanum Eucalyptus staigeriana, Lippia sidoides, Carboxymethylcellulose Dipping [118] and Pimenta pseudocaryophyllus Aloysia citriodora, Cymbopogon M. fructicola winterianus, Lippia alba, and Ocimum [105] americanum Polyvinyl alcohol-b- Penicillium (CGMCC Incorporation in nanofibers Zanthoxylum bungeanum Cyclodextrin [107] 3.13650) films nanofiber film Eucalyptus staigeriana, Lippia sidoides, Dipping Carboxymethylcellulose [119] and Pimenta pseudocaryophyllus Rhizopus stolonifer Cinnamon Emulsion and nanoemulsion [111] Eucalyptus staigeriana and Eucalyptus Carboxymethylcellulose Dipping [112] urograndis Mentha spicata, Mentha piperita, Thymus vulgaris CT carvacrol, and Fumigation [120] Thymus vulgaris CT thymol Monilinia spp. are responsible for brown rots in stone fruits, which are postharvest diseases that cause significant fruit losses [121]. TEO and vervain EO (VEO) showed fungi- cidal activity against MFC, ML, and MFG in the bioassay on peaches, where a concentration of 1000 ppm and 500 ppm for VEO and TEO, respectively, reduced the brown rot lesion on tested peach fruits [100]. Thyme essential oil is also effective against other postharvest pathogens. In fact, Alamri et al. demonstrated the efficacy of a new coating material to protect peaches against postharvest fungal infections, which contains EOs of mint and thyme (1 mL L ), gum arabic (10% w/v), and Tween 20. The preparations succeeded in slowing down disease development in vivo and reducing fungal growth in vitro at different temperatures. The fruits were kept in cold storage for up to one month, with very low levels of disease incidence and severity due to the application of the developed materials [98]. Naturally contaminated nectarines and peaches were exposed to TEO and SSEO by biofumigation in order to control postharvest diseases and maintain fruit quality. These EO vapors showed high antifungal activity against MFC, reducing fruit rot incidence during shelf-life. In contrast, BC was less sensitive to the treatments with EOs and in vitro tests confirmed the in vivo results. Moreover, EO vapors reduced weight loss and maintained the carotenoid and ascorbic acid content [9]. The effectiveness of the spike lavender, lavender absolute, dill weed, rosewood, FEO, and benchmark TEO against BC, MP, PE, and RS was examined in vitro using both the solid-phase diffusion method and the vapor-phase diffusion method, and in vivo on peach fruits. The results of in vitro and in vivo tests AppliedChem 2023, 3 207 showed that rosewood EO, used in the vapor-phase, prohibited the hypha growth and spore germination of BC, MP, PE, and RS at a concentration of 300 L L , keeping the quality of the peach fruit intact [99]. TEO and LgEO inhibited the growth of BC, PE, and RS mycelia by 25 and 50%, respectively, in vitro. Furthermore, the efficacy of a delivery system helps to reduce an unpleasant odor and taste resulting from the use of EOs. The EOs combined with the fungi antagonist Bacillus amyloliquefaciens in peach fruits were evaluated. The current investigation showed the potential of adopting B. amyloliquefaciens in a biodegradable modified atmosphere (MAP) pad delivery system in conjunction with LgEO to improve the postharvest life of peach fruits and reduce the growth of BC, PE, and RS [97]. Tea tree EO has a strong antifungal activity against MFC in vitro and in inoculated peaches. Tea tree EO changes the mycelial morphology, membrane permeability, and levels of intracellular ROS [103]. O. americanum and C. winterianus acted as antifungals, drastically reducing the growth of MFC and BC on peach fruits [105]. The use of coating or polysaccharide-based film avoids the problem of the uncontrolled volatilization of EOs and inhibits the growth of bacteria and fungi, allowing for a longer preservation of postharvest fruits [122–125]. Lian et al. [102] tested different chitosan-based films, also adding other polysaccharides, in order to improve the film matrix composition and the antifungal proprieties; specifically, they used films composed of: chitosan 1.8% (w/v), chitosan 1.2% and xanthan gum 0.6% (w/v), chitosan 1.8% and pullulan 0.6% (w/v), chitosan 1.8% and gum tragacanth 0.6% (w/v), and chitosan 1.8% and Arabic gum 0.6 (w/v). The use of thyme essential oil chitosan–Arabic gum composite film on peach fruit by coating or fumigation was the most effective in improving the hydrogen bonding interaction among the film matrix and stabilized the antifungal activity of TEO against MFC. Similarly, chitosan–Arabic gum composite films (1.8–0.6% w/v) with the incorporation of thyme EO reduced the lesion diameter of brown mold caused by MFC on the nectarine [103]. Treating ‘Spring Princes’ and ‘Sonnet’ peaches with TEO vapor leads to an increase in the total phenolic content and activities of defense-related enzymes and a reduction in brown rot lesion disease [101]. Among 26 EOs, only M. spicata, M. piperita, TEO CT carvacrol, and TEO CT thymol exhibited a strong inhibition on RS growth in the screening in vitro. Regarding the in vitro results, only peppermint EO (PEO) showed a consistent and substantial effect in limiting lesion growth on peach fruit inoculated with RS, whereas all four essential oil treatments reduced disease development [120]. The microcapsules of lilac EO (LLEO) showed strong antifungal activity against AA and BC, and the peaches’ quality attributes were maintained [95]. EOs extracted from galangal (GaEO), cortex EO (CoEO), and artemisia EO (ArEO) showed antifungal activity against pathogens causing postharvest diseases in peaches, such as PE, BC, AF, AA, RS, and MFC. A compound preparation based on the optimized concentration of three EOs was made into a microemulsion and was demonstrated to have greater antifungal activity [96]. The use of TEO and ClEO on apricots had a significant effect on the fruit quality and reduced the incidence and disease severity of MFC and BC, causal agents of gray and brown mold, respectively [10]. A key factor in fruit storage deterioration is pathogen attack. One of the most sig- nificant postharvest pathogens in terms of economic impact is gray mold fungi, which is brought on by BC [125]. Chitosan (0.15%) and ZMEO and CEO (0.15%) exhibited antimicrobial power against BC. Furthermore, the synergy between chitosan and essential oil (1:1 ratio) also showed antimicrobial power in vitro against BC. Similarly, strawberries treated with chitosan, ZMEO, CEO, and their combinations showed a significant reduction in the severity of BC compared to control fruits at the end of storage (Figure 4) [110]. Next, the antimicrobial power against BC was tested using an Aloe vera coating embedded with two different concentrations (500 and 1000 L L ) of basil essential oil. After 12 days of cold storage, strawberries treated with A. vera and the higher concentration of basil essential oil displayed AppliedChem 2023, 3, FOR PEER REVIEW 17 EOs extracted from galangal (GaEO), cortex EO (CoEO), and artemisia EO (ArEO) showed antifungal activity against pathogens causing postharvest diseases in peaches, such as PE, BC, AF, AA, RS, and MFC. A compound preparation based on the optimized concentration of three EOs was made into a microemulsion and was demonstrated to have greater antifungal activity [96]. The use of TEO and ClEO on apricots had a significant effect on the fruit quality and reduced the incidence and disease severity of MFC and BC, causal agents of gray and brown mold, respectively [10]. A key factor in fruit storage deterioration is pathogen attack. One of the most signif- icant postharvest pathogens in terms of economic impact is gray mold fungi, which is brought on by BC [125]. Chitosan (0.15%) and ZMEO and CEO (0.15%) exhibited antimicrobial power against BC. Furthermore, the synergy between chitosan and essential oil (1:1 ratio) also showed antimicrobial power in vitro against BC. Similarly, strawberries treated with chitosan, ZMEO, CEO, and their combinations showed a significant reduction in the severity of BC compared to control fruits at the end of storage (Figure 4) [110]. Next, the antimicrobial power against BC was tested using an Aloe vera coating embedded with two different con- AppliedChem 2023, 3 −1 208 centrations (500 and 1000 μL L ) of basil essential oil. After 12 days of cold storage, straw- berries treated with A. vera and the higher concentration of basil essential oil displayed the suppression of BC mycelium growth in comparison to the control, with no change in the suppression of BC mycelium growth in comparison to the control, with no change in the physico-chemical and sensory properties of the fruit [113]. the physico-chemical and sensory properties of the fruit [113]. Figure 4. Strawberries fruit treated with chitosan, Zataria multiflora, and Cinnamomum zeylanicum EOs. Figure 4. Strawberries fruit treated with chitosan, Zataria multiflora, and Cinnamomum zeylanicum Reproduced with permission from Mohammadi, A.; Hashemi, M.; Hosseini, S.M. Innovative Food EOs. Reproduced with permission from Mohammadi, A.; Hashemi, M.; Hosseini, S.M. Innovative Science and Emerging Technologies 2015. Food Science and Emerging Technologies 2015. Lemon verbena oil (LVEO), LgEO, Pepper–rosemarin oil (PREO), and Ocimum ameri- canum EO (OAEO) have shown antifungal activity against Colletotrichum sp., BC, and MFC. Lemon verbena oil (LVEO), LgEO, Pepper–rosemarin oil (PREO), and Ocimum amer- In vitro tests demonstrated that the use of these EOs with a concentration of 1.2 mL L icanum EO (OAEO) have shown antifungal activity against Colletotrichum sp., BC, and completely constrained CG mycelial growth. LVEO completely stopped BC at a dosage 1 1 1 of 0.8 mL L , LgEO and PREO at 1 mL L , and OAEO at 1.4 mL L . LVEO and LgEO MFC. In vitro tests demonstrated that the use of these EOs with a concentration of 1.2 mL 1 1 prevented the development of MFC with a dose of 0.4 mL L , PREO at 1.0 mL L , and −1 L completely constrained CG mycelial growth. LVEO completely stopped BC at a dosage OAEO at 0.8 mL L . The four EOs significantly reduced the growth of Colletotrichum sp. −1 −1 −1 of 0.8 mL L , LgEO and PREO at 1 mL L , and OAEO at 1.4 mL L . LVEO and LgEO mycelium on strawberries [105]. −1 −1 PEO and EuEO used in combination (1:1 v/v) and the peppermint essential oil com- prevented the development of MFC with a dose of 0.4 mL L , PREO at 1.0 mL L , and pletely suppressed the in vitro growth of BC mycelium, whereas the EuEO and the Moringa −1 OAEO at 0.8 mL L . The four EOs significantly reduced the growth of Colletotrichum sp. FO coating were less effective. The outcomes of the in vitro tests were confirmed in vivo, mycelium on strawberries [105]. with moderate fungus growth when EuEO and Moringa fatty oil were used as treatments, and BC was completely inhibited in strawberry samples treated with a combination of PEO PEO and EuEO used in combination (1:1 v/v) and the peppermint essential oil com- and EuEO, or without PEO [115]. pletely suppressed the in vitro growth of BC mycelium, whereas the EuEO and the Furthermore, sage oil (SaEO) at 1000 L L was able to completely inhibit the growth Moringa FO coating were less effective. The outcomes of the in vitro tests were confirmed of BC mycelium in vitro. With bioassay studies, it was confirmed that the concentration of 60 L L of SEO was a valid tool for controlling fungal infection, with a low effect on the quality of strawberry fruits [108]. At a concentration of 150 L L , TEO, Rosalina EO (RosEO), GaEO, PEO, palmarosa EO (PalEO), LgEO, and OEO completely inhibited BC’s mycelial growth, whereas, at a concentration of 100 L L , only LgEO, TEO, and OEO showed complete mycelial inhibition. The in vitro findings reveal that TEO and OEO were particularly effective at preventing postharvest gray mold on strawberry fruit. Exposure to OEO also preserved the fruit quality by preventing the natural deterioration in strawberry [114]. The CEO emulsion and nanoemulsion both exhibited a 100% deterrent 1 1 percentage in vitro against BC at 500 L L and against RS at 1000 L L . The fruit AppliedChem 2023, 3 209 infection rate was lowest (5.43%) in the strawberry fruit treated with CEO nano-emulsion at a concentration of 0.2%; moreover, it had a significant impact on preventing fruit degra- dation [111]. The in vitro growth of BC was fully inhibited by LgEO, litsea EO (LitEO), LEO, PEO, MiEO, LeEO, SaEO, and TEO at a concentration of 625 L L . Strawberry fruits that had been purposefully inoculated with BC, TEO, PEO, LgEO, and LitEO were examined in vivo. At concentrations of 250 or 500 L L , all four EOs exhibited high inhibition; however, at a dose of 125 L L , only PEO was able to prevent the growth of BC lesions. However, these EOs changed the sensory quality of strawberries. Nevertheless, LgEO and LitEO EOs seem to be acceptable for consumers when applied at a 125 L L concentration [116]. The name of the disease produced by RS, soft rot, already describes the distinctive symptom: a soft, watery rot with juice leakage that, in conditions of high moisture, is covered by mycelium and contains pathogen structures [126]. PEO displayed an antifungal activity on strawberries artificially inoculated with RS at 86, 84, and 59% after 24, 48, and 72 h of inoculation, respectively. Furthermore, MiEO, TEO CT carvacrol, and TEO CT thymol also exhibited a fair amount of antifungal power, but much less than PEO [120]. The essential oil of EuEO showed great antifungal capacity in vitro against BC and RS at concentrations of 2000 and 1500 L L , respectively. In the in vivo strawberry test, EuEO essential oil, when applied curatively in association with carboxymethylcellulose, inhibited the proliferation of RS mycelium, whereas it had no antifungal effect against BC [112]. On the contrary, Oliveira et al. (2019) [118,119] tested the effectiveness of EuEO, PREO, and PPEO in vitro and in vivo against RS and CA on strawberry fruits. The PREO presented the highest antifungal activity in vitro (62.5 < MIC 125). In vivo, the EO incorporated in the CMC coating showed efficiency in disease severity reduction in strawberry fruits. CN is the causative agent of strawberry black spots. Hosseini and coworkers [116] studied the effects of six different concentrations of garlic oil (GEO) (20, 155, 290, 430, 560, 1 1 and 700 mL L ) and REO (100, 420, 740, 1060, 1380, and 1700 mL L ) in vitro and in vivo on strawberry fruit. They argued that GEO and REO significantly inhibited the mycelial growth and conidial germination in vitro of CN in contact and vapor assays compared with the untreated control. However, the EC50 assay indicated that GEO was more effective than REO against the pathogen [117]. The results of the in vitro test were confirmed in vivo and in greenhouse conditions; in fact, these EOs significantly reduced the anthracnose disease incidence and severity compared with untreated controls [117]. Collelotrichum gloeosporioids, but, in general, all fungi of this family, is a significant and dangerous saprophytic pathogen of sweet cherries that frequently results in postharvest losses [127]. In vitro tests with TEO, CEO, and ClEO in the contact phase demonstrated the highest inhibitory effect on CA mycelial growth at concentrations of 467 and 667 L L , respectively. TEO volatiles totally controlled the disease, whereas CEO volatiles decreased the pathogen penetration and development in strawberry fruit under disease-favorable conditions [3]. Wang and coworkers tested the efficacy of nine EOs (clove, thyme, palmarosa, lemon- grass, lemon, cinnamon, and laurel leaf) in vitro in vapor and contact phases (100 and 1000 L L , respectively). Of these, CEO and ClEO showed the greatest inhibition of growth against CG. The in vivo findings demonstrated that clove oil was the most effective inhibitor against CG in artificially infected sweet cherries. Its fungitoxic activities may be attributed to the destruction of the cell wall and cell membrane, as well as the leakage of intracellular components [108]. Against AF, BC, and PE, Zhang and coworkers [107] tested a new polyvinyl alcohol/b- cyclodextrin (PVA/b-CD, 6:1) nanofibers with ZBEO (10%) incorporated. Both in vitro and in vivo tests on strawberries and sweet cherries showed a significant antifungal activity of film/ZBEO, which increased the shelf-life of fruits, inhibiting the fungi growth without changing their sensory characteristics (Figure 5) [107]. AppliedChem 2023, 3, FOR PEER REVIEW 19 volatiles decreased the pathogen penetration and development in strawberry fruit under disease-favorable conditions [3]. Wang and coworkers tested the efficacy of nine EOs (clove, thyme, palmarosa, lemongrass, lemon, cinnamon, and laurel leaf) in vitro in vapor and contact phases (100 −1 and 1000 µ L L , respectively). Of these, CEO and ClEO showed the greatest inhibition of growth against CG. The in vivo findings demonstrated that clove oil was the most effective inhibitor against CG in artificially infected sweet cherries. Its fungitoxic activities may be attributed to the destruction of the cell wall and cell membrane, as well as the leakage of intracellular components [108]. Against AF, BC, and PE, Zhang and coworkers [107] tested a new polyvinyl alcohol/β- cyclodextrin (PVA/β-CD, 6:1) nanofibers with ZBEO (10%) incorporated. Both in vitro and in vivo tests on strawberries and sweet cherries showed a significant antifungal activity of film/ZBEO, which increased the shelf-life of fruits, inhibiting the fungi growth without AppliedChem 2023, 3 210 changing their sensory characteristics (Figure 5) [107]. Figure 5. Sweet cherries treated with Zanthoxylum bungeanum essential oil incorporated or added to Figure 5. Sweet cherries treated with Zanthoxylum bungeanum essential oil incorporated or added to polyvinyl alcohol/b-cyclodextrin film. The red circles indicated the presence of fungi. Reproduced polyvinyl alcohol/β-cyclodextrin film. The red circles indicated the presence of fungi. Reproduced with permission from Zhang, H.; Zhang, C.; Wang, X.; Huang, Y.; Xiao, M.; Hu, Y.; Zhang, J. LWT 2022. with permission from Zhang, H.; Zhang, C.; Wang, X.; Huang, Y.; Xiao, M.; Hu, Y.; Zhang, J. LWT The EOs from Acorus calamus, Allium sativum, Mucuna pruriens, and Sesamum indicum were particularly successful in in vitro tests in preventing the development of several postharvest fungi. In particular, PN was the target of GEO’s strongest antifungal action among the EOs tested. GEO also considerably slowed the development of other fungal The EOs from Acorus calamus, Allium sativum, Mucuna pruriens, and Sesamum indicum species that were applied in the test, including AN, AF, and RM. In vivo, the GEO also were particularly successful in in vitro tests in preventing the development of several significantly diminished the growth of AF and AN on plum fruit, reducing the deterioration postharvest fungi. In particular, PN was the target of GEO’s strongest antifungal action of these fruits [106]. among the Dif E fer Os enttes papers ted. rGEO eported also antimi con crobial siderab effects ly of slow EOsed onttotal he deve yeast,lo molds, pment and of other fungal mesophilic bacteria of stone fruit. The antimicrobial activity reported in this review could species that were applied in the test, including AN, AF, and RM. In vivo, the GEO also be analyzed further. The main results of these studies are shown in Table 6. significantly diminished the growth of AF and AN on plum fruit, reducing the deteriora- Table 6. Antimicrobial effect of EOs on microorganisms responsible for fruit spoilage in stone fruits. tion of these fruits [106]. Different papers reported antimicrobial effects of EOs on total yeast, molds, and Fruit Pathogen Coating EOs Results Reference Basil seed gum 5% When increasing OEO concentration, there is mesophilic bacteria of stone fruit. The antimicrobial activity reported in this review could Mesophilic aerobic Glycerol 30% w/w an incremental reduction to 31.81% and 25.65% Apricots bacteria and for mold Oregano [128] Tween-20 15% of the total plate counts and yeast and mold be analyzed further. The main results of these studies are shown in Table 6. and yeast counts Oregano EO (0–6% v/v) counts, respectively Chitosan 1:3.2 (w/v) Tween 80 0.5 wt% Eryngium campestre EO Sweet Mesophilic aerobic Reduction in total bacteria population of Table 6. Antimicrobial effect of EOs on microorganisms responsible for fruit spoilage in stone pentasodium (800, 1600, 2400, and [129] cherries bacteria around 4 log CFU mL tripolyphosphate 3200 mg L ) fruits. 0.3% (w/v) Total molds, yeast, and Aliginate 2% Eugenol (0.1–0.2%) Decrease in aerobic mesophilic microrganisms, mesophilic [130] Fruit Pathogen Coating EOs Results Reference Pectin 2% Citral (0.15–0.3%) yeasts, and molds development microorganisms Aloe vera gel (20–40%) Lower total aerobic mesophilic bacteria and Basil seed gum 5% When increasing OEO Lemongrass 1% [131] Tween 80 (0–01% w/v) yeast and mold counts Mesophilic aerobic Geraniol 1.2–2.4 L mL Psychrophilic bacteria, yeast, mold, and Strawberries Gellan gum (0.5% w/v) Glycerol 30% w/w concentration, there is an Pomegranate extract mesophilic bacteria counts were lower and [132] Glycerol (0.56% w/v) bacteria and for 1 Total molds, yeast, and 360–720 g mL more constant than the control Apricots Tween-20 15% Oregano incremental reduction to [128] mesophilic Red thyme, peppermint Chitosan and Tween 80 Preservative action on the stored strawberries [133] mold and y microorganisms east and limonene Oregano Gelatin 1:25 EO (w/v) (0–6% The incorporation 31.81% ofand mint EO 25.65% into of the counts Sorbitol w/w Peppermint 0.5–1% gelatin-based edible coating inhibited total [134] Tween 80 aerobic mesophilic flora, molds, and yeasts. v/v) total plate counts and yeast 6. Conclusions and Perspectives EOs could contribute to the goal of sustainable production because, to date, the acquisition of tolerance by the studied pathogens in treated fresh products has not been AppliedChem 2023, 3 211 demonstrated. They are considered safe for both the environment and human health, so interest in their use as antimicrobial agents for fresh produce has increased in recent years. Several research studies have been conducted to demonstrate the benefits of EOs in containing many fruit postharvest diseases. In general, EOs extracted from different plant organs have been found to be non-harmful to fresh food, human health, and the environment if the correct doses and timing are observed. Several research studies have been conducted under in vitro conditions. Research on the use of EOs on fresh fruit has shown that the EOs concentration and their compounds, required to inhibit microbial growth, are generally higher in food than in culture media. Unfortunately, in some cases, the concentrations that are effective against the disease can be phytotoxic and cause the further chemical deterioration of treated food. Botanicals can be applied in the form of dipping, but several studies report their use as vapors due to their advantage in preventing product contamination. Many of these EOs have already been registered as flavoring agents, so they can easily be registered for postharvest use. Despite all of the obvious advantages, much work remains to be carried out on the detailed examination of the biological activity and dispersion of EOs and their compounds in fruit tissues in order to develop a formulation that maintains antimicrobial activity without causing undesirable effects on the foods. A viable alternative to increasing the half-life of the EOs and reducing the contact between the essential oil and the food, so as to reduce the possibility of undesirable effects on the fruit, is the inclusion of the EOs in nanoparticles, coatings, or films. The inclusion of EOs in film, coating, and/or nanoparticles represents a valid strategy for increasing the water solubility, stability, and bioavailability and decreasing the volatility of EOs. To date, there are few scientific contributions reporting the active inclusion of EOs as a treatment against postharvest pathogens in fruits of the Rosaceae family. Author Contributions: Conceptualization, A.M.; investigation, A.M. and M.C.; resources, A.M. and M.C.; data curation, A.M.; writing—original draft preparation, A.M. and M.C.; writing—review and editing, A.M., V.B., A.F. and M.P.; visualization, A.F. and M.P.; supervision, A.F. and M.P. All authors have read and agreed to the published version of the manuscript. 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