Vol. 65 No. 2 (2026)
Articles

Identification of antifungal essential oils for control of brown spot pathogens of ‘Rocha’ pear

Inês MENDONÇA
INIAV, IP – National Institute for Agrarian and Veterinary Research, Polo de Vairão, Rua dos Lagidos, Lugar da Madalena, 4485-655, Vairão, Vila do Conde, Portugal
Beatriz FERNANDES
INIAV, IP – National Institute for Agrarian and Veterinary Research, Avenida da República, Quinta do Marquês, 2780-157, Oeiras, Portugal
Carmo SERRANO
INIAV, IP – National Institute for Agrarian and Veterinary Research, Avenida da República, Quinta do Marquês, 2780-157, Oeiras, Portugal
Ana C. MARQUES
CERENA, DEQ, Instituto Superior Técnico, University of Lisbon, Avenida Rovisco Pais, No. 1, 1049-001, Lisbon, Portugal
Ricardo OLIVEIRA
INIAV, IP – National Institute for Agrarian and Veterinary Research, Polo de Vairão, Rua dos Lagidos, Lugar da Madalena, 4485-655, Vairão, Vila do Conde, Portugal
Carina ALMEIDA
INIAV, IP – National Institute for Agrarian and Veterinary Research, Polo de Vairão, Rua dos Lagidos, Lugar da Madalena, 4485-655, Vairão, Vila do Conde, Portugal
Miguel LEÃO DE SOUSA
INIAV, IP – National Institute for Agrarian and Veterinary Research, Polo de Alcobaça, Estrada de Leiria, 2460-059, Alcobaça, Portugal
Armando VENÂNCIO
CEB – Centre of Biological Engineering, University of Minho, Campus de Gualtar, 4710-057, Braga, Portugal
Sónia SILVA
CEB – Centre of Biological Engineering, University of Minho, Campus de Gualtar, 4710-057, Braga, Portugal

Published 2026-09-10

Keywords

  • Alternaria arborescens,
  • Stemphylium vesicarium,
  • fungicide sensitivity,
  • in vitro screening

How to Cite

[1]
MENDONÇA I., “Identification of antifungal essential oils for control of brown spot pathogens of ‘Rocha’ pear”, Phytopathol. Mediterr., vol. 65, no. 2, pp. 267–286, Sep. 2026.

Abstract

Brown spot of pear (BSP) is a major concern in ‘Rocha’ pear production. Essential oils were identified with in vitro antifungal properties against fungi isolated from BSP, and encapsulated formulations were developed via spray-drying. Four fungal isolates from symptomatic leaf and fruit tissues of ‘Rocha’ pears were identified as Stemphylium vesicarium (isolates MUM 26.01 and MUM 26.02) and Alternaria arborescens (isolates MUM 26.03 and MUM 26.04) through multilocus sequencing. Sensitivity of these isolates was assessed to five fungicides commonly used in Portuguese orchards. Tebuconazole and cyprodinil were the most effective across isolates, while trifloxystrobin gave limited inhibition, indicating low sensitivity of these isolates. To assess sustainable alternatives, 18 essential oils (EOs) were screened, showing that lemongrass and lemon thyme EOs were the most promising candidates, completely inhibiting mycelium growth and spore germination of the fungi at low concentrations. Encapsulation of these two EOs on HI-CAP® 100, using spray-drying, preserved their antifungal activity. Chemical characterization showed that lemongrass and lemon thyme EOs consisted primarily of oxygenated monoterpenes, which were probably responsible for the high antifungal activity observed. These results highlight the potential of lemongrass and lemon thyme EOs as promising candidates for in vivo validation as sustainable alternatives to synthetic fungicides for control of S. vesicarium and A. arborescens, and management of BSP in ‘Rocha’ pear production.

Downloads

Download data is not yet available.

References

  1. Akbari Dana M., Hashemi S.J., Daei Ghazvini R., Khodavesi S., Modiri M., … Rezaie S., 2019. Effect of benomyl and diazinon on acquired azole resistance in Aspergillus flavus and expression of mdr1 and cyp51c genes. Current Medical Mycology 5: 27–32. https://doi.org/10.18502/cmm.5.2.1158.
  2. Akhtari A., Davari M., Habibi-Yangjeh A., Ebadollahi A., Feizpour S., 2022. Antifungal Activities of Pure and ZnO-Encapsulated Essential Oil of Zataria multiflora on Alternaria solani as the Pathogenic Agent of Tomato Early Blight Disease. Frontiers in Plant Science 13: 932475. https://doi.org/10.3389/fpls.2022.932475.
  3. Alberoni G., Cavallini D., Collina M., Brunelli A., 2010. Characterisation of the first Stemphylium vesicarium isolates resistant to strobilurins in Italian pear orchards. European Journal of Plant Pathology 126: 453–457. https://doi.org/10.1007/s10658-009-9559-3.
  4. Altay Ö., Köprüalan Ö., İlter I., Koç M., Ertekin F.K., Jafari S.M., 2024. Spray drying encapsulation of essential oils; process efficiency, formulation strategies, and applications. Critical Reviews in Food Science and Nutrition 64: 1139–1157. https://doi.org/10.1080/10408398.2022.2113364.
  5. Andrade-Ochoa S., Sánchez-Aldana D., Rodríguez-Valdez L.M., Nevárez-Moorillón G.V., 2023. In vitro and Quantitative and Structure Activity Relationship (QSAR) evaluation of the antifungal activity of terpenoid constituents of essential oils against Alternaria alternata and Fusarium oxysporum. Biomédica 43: 156–169. https://doi.org/10.7705/biomedica.6883.
  6. Antonioli G., Fontanella G., Echeverrigaray S., Longaray Delamare A.P., Fernandes Pauletti G., Barcellos T., 2020. Poly(lactic acid) nanocapsules containing lemongrass essential oil for postharvest decay control: In vitro and in vivo evaluation against phytopathogenic fungi. Food Chemistry 326: 126997. https://doi.org/10.1016/j.foodchem.2020.126997.
  7. Asgarian Z.S., Palou L., Souza R.F.L. de, Quintanilla P.G., Taberner V., … Pérez-Gago M.B., 2023. Hydroxypropyl Methylcellulose and Gum Arabic Composite Edible Coatings Amended with Geraniol to Control Postharvest Brown Rot and Maintain Quality of Cold-Stored Plums. Foods 12: 2978. https://doi.org/10.3390/foods12152978.
  8. Bajac J., Nikolovski B., Petrović L., Nemeš T., Kostić M., … Mitrović I., 2025. Antimicrobial and insecticidal activity of spray dried juniper berry (Juniperus communis L.) essential oil microcapsules prepared by using gum arabic and maltodextrin. International Journal of Biological Macromolecules 306: 141128. https://doi.org/10.1016/j.ijbiomac.2025.141128.
  9. Bakkali F., Averbeck S., Averbeck D., Idaomar M., 2008. Biological effects of essential oils – A review. Food and Chemical Toxicology 46: 446–475. https://doi.org/10.1016/j.fct.2007.09.106.
  10. Baranauskiene R., Venskutonis P.R., 2009. Comparison of wall materials for the microencapsulation of thyme essential oil. In: XVIIth International Conference on Bioencapsulation, Groningen, Netherlands, 1–4.
  11. Beg M.A., Aktaruzzaman Md., Lewis K.J., Oliver J.E., 2025. Fungicide resistance profiles of Alternaria spp. associated with fruit rot of blueberry in Georgia, USA. Frontiers in Plant Science 16: 1524586. https://doi.org/10.3389/fpls.2025.1524586.
  12. Borkow G., Gabbay J., 2005. Copper as a Biocidal Tool. Current Medicinal Chemistry 12: 2163–2175. https://doi.org/10.2174/0929867054637617.
  13. Burt S., 2004. Essential oils: their antibacterial properties and potential applications in foods ‒ a review. International Journal of Food Microbiology 94: 223–253. https://doi.org/10.1016/j.ijfoodmicro.2004.03.022.
  14. Camiletti B.X., Lichtemberg P.S.F., Paredes J.A., Carraro T.A., Velascos J., Michailides T.J., 2022. Characterization, pathogenicity, and fungicide sensitivity of Alternaria isolates associated with preharvest fruit drop in California citrus. Fungal Biology 126: 277–289. https://doi.org/10.1016/j.funbio.2022.02.003.
  15. Castresana J., 2000. Selection of Conserved Blocks from Multiple Alignments for Their Use in Phylogenetic Analysis. Molecular Biology and Evolution 17: 540–552. https://doi.org/10.1093/oxfordjournals.molbev.a026334.
  16. Cavina F., Baiocco S., Tomba L., Ravaglia F., Moretti C., … Bugiani R., 2024. A New Source of Inoculum for Stemphylium vesicarium: Consequences for the Management of Brown Spot of Pear. Agronomy 14: 2522. https://doi.org/10.3390/agronomy14112522.
  17. Chen Y., Zeng Y., Li Y., Ye Z., Li L., Zhou Z., 2025. Antifungal effects of citral against Alternaria alternata in postharvest pear fruit and its potential mechanism. Postharvest Biology and Technology 223: 113424. https://doi.org/10.1016/j.postharvbio.2025.113424.
  18. Collina M., Alberoni G., Brunelli A., 2007. First occurrence of strobilurin-resistant isolates of Stemphylium vesicarium in an Italian pear orchard. Communications in Agricultural and Applied Biological Sciences 72: 735–738.
  19. Contreras Martínez O.I., Angulo Ortíz A., Santafé Patiño G., 2022. Mechanism of Antifungal Action of Monoterpene Isoespintanol against Clinical Isolates of Candida tropicalis. Molecules 27: 5808. https://doi.org/10.3390/molecules27185808.
  20. Cordero R.J.B., Casadevall A., 2017. Functions of fungal melanin beyond virulence. Fungal Biology Reviews 31: 99–112. https://doi.org/10.1016/j.fbr.2016.12.003.
  21. Corkley I., Fraaije B., Hawkins N., 2022. Fungicide resistance management: Maximizing the effective life of plant protection products. Plant Pathology 71: 150–169. https://doi.org/10.1111/ppa.13467.
  22. de Sousa D.P., Damasceno R.O.S., Amorati R., Elshabrawy H.A., de Castro R.D., … Lima T.C., 2023. Essential Oils: Chemistry and Pharmacological Activities. Biomolecules 13: 1144. https://doi.org/10.3390/biom13071144.
  23. Dereeper A., Guignon V., Blanc G., Audic S., Buffet S., … Gascuel O., 2008. Phylogeny.fr: robust phylogenetic analysis for the non-specialist. Nucleic Acids Research 36: W465–W469. https://doi.org/10.1093/nar/gkn180.
  24. DGAV, 2024. Caderno Técnico: A Estenfiliose da Pereira - Estratégia de Gestão de Resistência a Fungicidas, Direção Geral da Alimentação e Veterinária, Lisboa, Portugal. Available at: https:/www.dgav.pt/. Accessed May 5, 2026.
  25. Donne I., Higgins D.S., Brisco-McCann E., Hausbeck M.K., 2020. Limiting Fungal Foliar Diseases on Carrots for Organic and Conventional Markets. Plant Health Progress 21: 217–223. https://doi.org/10.1094/PHP-12-19-0096-RS.
  26. Dorjee L., Gogoi R., Kamil D., Kumar R., Mondal T.K., … Gurung B., 2023. Essential oil-grafted copper nanoparticles as a potential next-generation fungicide for holistic disease management in maize. Frontiers in Microbiology 14: 1204512. https://doi.org/10.3389/fmicb.2023.1204512.
  27. European Commission, 2011. Commission Implementing Regulation (EU) No. 540/2011 of 25 May 2011 implementing Regulation (EC) No 1107/2009 of the European Parliament and of the Council as regards the list of approved active substances. Official Journal of the European Union L153: 1‒186. Available at: http://data.europa.eu/eli/reg_impl/2011/540/oj. Accessed May 26, 2026.
  28. European Commission, 2012. Commission Implementing Regulation (EU) No 872/2012 of 1 October 2012 adopting the list of flavouring substances provided for by Regulation (EC) No 2232/96 of the European Parliament and of the Council, introducing it in Annex I to Regulation (EC) No 1334/2008 of the European Parliament and of the Council and repealing Commission Regulation (EC) No 1565/2000 and Commission Decision 1999/217/EC. Official Journal of the European Union L267: 1‒161. Available at: http://data.europa.eu/eli/reg_impl/2012/872/oj. Accessed May 26, 2026.
  29. Fernandes B., Mendonça I., Silva S., Sousa S., Matos C., … Serrano C., 2026. Encapsulation and Release Profiling of Antifungal Essential Oils via Spray Drying Analyzed by HS-SPME-GC-MS/MS. Food Chemistry.
  30. Fincheira P., Jofré I., Espinoza J., Levío-Raimán M., Tortella G., … Rubilar O., 2023. The efficient activity of plant essential oils for inhibiting Botrytis cinerea and Penicillium expansum: Mechanistic insights into antifungal activity. Microbiological Research 277: 127486. https://doi.org/10.1016/j.micres.2023.127486.
  31. Fogarty R. V., Tobin J.M., 1996. Fungal melanins and their interactions with metals. Enzyme and Microbial Technology 19: 311–317. https://doi.org/10.1016/0141-0229(96)00002-6.
  32. Fontaine K., Fourrier-Jeandel C., Armitage A.D., Boutigny A.-L., Crépet M., … Aguayo J., 2021. Identification and pathogenicity of Alternaria species associated with leaf blotch disease and premature defoliation in French apple orchards. PeerJ 9: e12496. https://doi.org/10.7717/peerj.12496.
  33. FRAC, 2025. FRAC Code List 2025: Fungal control agents sorted by cross-resistance pattern and mode of action (including coding for FRAC Groups on product labels). Fungicide Resistance Action Committee. Available at: https://www.frac.info/. Accessed January 21, 2026.
  34. Gangavarapu Y., Palwai S., 2022. Antifungal Activity of Lemongrass Oil Against Pathogenic Fungi. International Journal of High School Research 4: 34–38. https://doi.org/10.36838/v4i1.7.
  35. Garibaldi A., Bertetti D., Franco Ortega S., Martini P., Gullino M.L., 2016. Powdery Mildew Caused by Golovinomyces biocellatus on Lemon Thyme ( Thymus × citriodorus ) ‘Aureus’ in Italy. Plant Disease 100: 2168. https://doi.org/10.1094/PDIS-04-16-0451-PDN.
  36. Głos H., Bryk H., Michalecka M., Puławska J., 2023. First report of Stemphylium vesicarium, a new pathogen causing brown spot of pear in Poland. Journal of Plant Pathology 105: 1161–1162. https://doi.org/10.1007/s42161-023-01365-6.
  37. Gossen B.D., Tayviah C.S., McDonald M.R., 2021. The Role of Ascospores and Conidia, in Relation to Weather Variables, in the Epidemiology of Stemphylium Leaf Blight of Onion. Plant Disease 105: 1912–1918. https://doi.org/10.1094/PDIS-06-20-1283-RE.
  38. Griffith G.W., Easton G.L., Detheridge A., Roderick K., Edwards A., … Perkins W.T., 2007. Copper deficiency in potato dextrose agar causes reduced pigmentation in cultures of various fungi. FEMS Microbiology Letters 276: 165–171. https://doi.org/10.1111/j.1574-6968.2007.00923.x.
  39. Habib W., Masiello M., El Ghorayeb R., Gerges E., Susca A., … Moretti A., 2021. Mycotoxin Profile and Phylogeny of Pathogenic Alternaria Species Isolated from Symptomatic Tomato Plants in Lebanon. Toxins 13: 513. https://doi.org/10.3390/toxins13080513.
  40. He M.-H., Wang Y.-P., Wu E.-J., Shen L.-L., Yang L.-N., … Zhan J., 2019. Constraining Evolution of Alternaria alternata Resistance to a Demethylation Inhibitor (DMI) Fungicide Difenoconazole. Frontiers in Microbiology 10: 1609. https://doi.org/10.3389/fmicb.2019.01609.
  41. Hyldgaard M., Mygind T., Meyer R.L., 2012. Essential Oils in Food Preservation: Mode of Action, Synergies, and Interactions with Food Matrix Components. Frontiers in Microbiology 3: 12. https://doi.org/10.3389/fmicb.2012.00012.
  42. Kaur G., Ganjewala D., Bist V., Verma P.C., 2019. Antifungal and larvicidal activities of two acyclic monoterpenes; citral and geraniol against phytopathogenic fungi and insects. Archives of Phytopathology and Plant Protection 52: 458–469. https://doi.org/10.1080/03235408.2019.1651579.
  43. Koka J.A., Bhat M.Y., Wani A.H., 2021. In vitro efficacy of fungicides on mycelial growth and spore germination of Alternaria alternata and Mucor plumbeus. Journal of Drug Delivery and Therapeutics 11: 17–22. https://doi.org/10.22270/jddt.v11i3.4692.
  44. Kumar S., Stecher G., Suleski M., Sanderford M., Sharma S., Tamura K., 2024. MEGA12: Molecular Evolutionary Genetic Analysis Version 12 for Adaptive and Green Computing. Molecular Biology and Evolution (F.U. Battistuzzi, ed.) 41(12):1‒9. https://doi.org/10.1093/molbev/msae263.
  45. La Torre A., Iovino V., Caradonia F., 2018. Copper in plant protection: Current situation and prospects. Phytopathologia Mediterranea, Firenze University Press 57: 201–236. https://doi.org/10.14601/Phytopathol_Mediterr-23407.
  46. Lamichhane J.R., Osdaghi E., Behlau F., Köhl J., Jones J.B., Aubertot J.-N., 2018. Thirteen decades of antimicrobial copper compounds applied in agriculture. A review. Agronomy for Sustainable Development 38: 28. https://doi.org/10.1007/s13593-018-0503-9.
  47. Llorente C., Bárcena A., Vera Bahima J., Saparrat M.C.N., Arambarri A.M., … Balatti P.A., 2012a. Cladosporium cladosporioides LPSC 1088 Produces the 1,8-Dihydroxynaphthalene-Melanin-Like Compound and Carries a Putative pks Gene. Mycopathologia 174: 397–408. https://doi.org/10.1007/s11046-012-9558-3.
  48. Llorente I., Montesinos E., 2006. Brown Spot of Pear: An Emerging Disease of Economic Importance in Europe. Plant Disease 90: 1368–1375. https://doi.org/10.1094/PD-90-1368.
  49. Llorente I., Moragrega C., Ruz L., Montesinos E., 2012b. An update on control of brown spot of pear. Trees 26: 239–245. https://doi.org/10.1007/s00468-011-0607-1.
  50. Llorente I., Vilardell A., Vilardell P., Pattori E., Bugiani R., … Montesinos E., 2010. Control of brown spot of pear by reducing the overwintering inoculum through sanitation. European Journal of Plant Pathology 128: 127–141. https://doi.org/10.1007/s10658-010-9637-6.
  51. Loebler M., Sánchez C., Muchagato Maurício E., Diogo E., Santos M., … Duarte M.P., 2020. Potential Application of Propolis Extracts to Control the Growth of Stemphylium vesicarium in “Rocha” Pear. Applied Sciences 10: 1990. https://doi.org/10.3390/app10061990.
  52. Lomba-Viana X., Raymundo A., Prista C., Alegria M.J., Sousa I., 2022. Clean Label “Rocha” Pear (Pyrus communis L.) Snack Containing Juice By-Products and Euglena gracilis Microalgae. Frontiers in Nutrition 9: 825999. https://doi.org/10.3389/fnut.2022.825999.
  53. Luo D., Ye S., Qu G., Ba L., 2024. Inhibitory effect and action mechanism of citral against black rot in pitaya fruit. Physiological and Molecular Plant Pathology 131: 102275. https://doi.org/10.1016/j.pmpp.2024.102275.
  54. Luz J.P., Amaro C., Isabel R., Jambrek A., Barrios D., 2018. Efficacy of fungicides in vitro for Stemphylium vesicarium of pear. Revista de Ciências Agrárias, Especial 41: 97–101. https://doi.org/10.19084/RCA.17072.
  55. McGrath M.T., 2009. Fungicides and other Chemical Approaches for use in Plant Disease Control. In: Encyclopedia of Microbiology (M. Schaechter, ed.), 3rd ed. Elsevier/Academic Press, Amsterdam, The Netherlands, 412–421. https://doi.org/10.1016/B978-012373944-5.00357-6.
  56. Minozzo M., de Souza M.A., Bernardi J.L., Puton B.M.S., Valduga E., … Cansian R.L., 2023. Antifungal activity and aroma persistence of free and encapsulated Cinnamomum cassia essential oil in maize. International Journal of Food Microbiology 394: 110178. https://doi.org/10.1016/j.ijfoodmicro.2023.110178.
  57. Nazzaro F., Fratianni F., Coppola R., Feo V. De, 2017. Essential Oils and Antifungal Activity. Pharmaceuticals 10: 86. https://doi.org/10.3390/ph10040086.
  58. Nene Y.L., Thapliyal P.N., 1979. Fungicides in Plant Disease Control. Oxford & IBH Publishing Co., New Delhi, India.
  59. Oli N., Singh U.K., Jha S.K., 2019. Antifungal Activity of Plant’s Essential Oils against Post Harvest Fungal Disease of Apple Fruit. Forestry: Journal of Institute of Forestry, Nepal 16: 86–100. https://doi.org/10.3126/forestry.v16i0.28361.
  60. Olita T., Stankovic M., Sung B., Jones M., Gibberd M., 2024. Growers’ perceptions and attitudes towards fungicide resistance extension services. Scientific Reports 14: 6821. https://doi.org/10.1038/s41598-024-57530-z.
  61. Oliveira A.S., Rolo J., Gaspar C., Cavaleiro C., Salgueiro L., … Palmeira-de-Oliveira A., 2022. Chemical characterization and bioactive potential of Thymus × citriodorus (Pers.) Schreb. preparations for anti-acne applications: Antimicrobial, anti-biofilm, anti-inflammatory and safety profiles. Journal of Ethnopharmacology 287: 114935. https://doi.org/10.1016/j.jep.2021.114935.
  62. Parikh L., Agindotan B.O., Burrows M.E., 2021. Antifungal Activity of Plant-Derived Essential Oils on Pathogens of Pulse Crops. Plant Disease 105: 1692–1701. https://doi.org/10.1094/PDIS-06-20-1401-RE.
  63. Paušič A., Roškarič M., Lešnik M., 2023. Preharvest Treatments with Low-Risk Plant Protection Products Can Help Apple Growers Fulfill the Demands of Supermarket Chains Regarding Pesticide Residues and Marketing Apples under 0-Residue Brands. Agronomy 13: 1151. https://doi.org/10.3390/agronomy13041151.
  64. Perina F.J., Amaral D.C., Fernandes R.S., Labory C.R., Teixeira G.A., Alves E., 2015. Thymus vulgaris essential oil and thymol against Alternaria alternata (Fr.) Keissler: effects on growth, viability, early infection and cellular mode of action. Pest Management Science 71: 1371–1378. https://doi.org/10.1002/ps.3933.
  65. Puig M., Ruz L., Montesinos E., Moragrega C., Llorente I., 2015. Combined morphological and molecular approach for identification of Stemphylium vesicarium inoculum in pear orchards. Fungal Biology 119: 136–144. https://doi.org/10.1016/j.funbio.2014.11.006.
  66. Raveau R., Fontaine J., Lounès-Hadj Sahraoui A., 2020. Essential Oils as Potential Alternative Biocontrol Products against Plant Pathogens and Weeds: A Review. Foods 9: 365. https://doi.org/10.3390/foods9030365.
  67. Sattary M., Amini J., Hallaj R., 2020. Antifungal activity of the lemongrass and clove oil encapsulated in mesoporous silica nanoparticles against wheat’s take-all disease. Pesticide Biochemistry and Physiology 170: 104696. https://doi.org/10.1016/j.pestbp.2020.104696.
  68. Shukla R., Singh P., Prakash B., Dubey N.K., 2012. Antifungal, aflatoxin inhibition and antioxidant activity of Callistemon lanceolatus (Sm.) Sweet essential oil and its major component 1,8-cineole against fungal isolates from chickpea seeds. Food Control 25: 27–33. https://doi.org/10.1016/j.foodcont.2011.10.010.
  69. Singh P., Bugiani R., Cavanni P., Nakajima H., Kodama M., … Kohmoto K., 1999. Purification and Biological Characterization of Host-Specific SV-Toxins from Stemphylium vesicarium Causing Brown Spot of European Pear. Phytopathology 89: 947–953. https://doi.org/10.1094/PHYTO.1999.89.10.947.
  70. Soares C., Morales H., Faria J., Figueiredo A.C., Pedro L.G., Venâncio A., 2016. Inhibitory effect of essential oils on growth and on aflatoxins production by Aspergillus parasiticus. World Mycotoxin Journal 9: 525–534. https://doi.org/10.3920/WMJ2015.1987.
  71. Sobreiro J., Batalha Neto C.S., 2024. Insights into brown spot disease: causal agents and host interactions in agroecosystems. In: Estudos em Ciências Agrárias e Ambientais I (E. Spers, ed), Editora Artemis, Curitiba, PR, Brazil, 88–100. https://doi.org/10.37572/EdArt_30072419210.
  72. Soltanzadeh M., Peighambardoust S.H., Ghanbarzadeh B., Mohammadi M., Lorenzo J.M., 2021. Chitosan nanoparticles encapsulating lemongrass (Cymbopogon commutatus) essential oil: Physicochemical, structural, antimicrobial and in-vitro release properties. International Journal of Biological Macromolecules 192: 1084–1097. https://doi.org/10.1016/j.ijbiomac.2021.10.070.
  73. Soylu E.M., Kurt Ş., Soylu S., 2010. In vitro and in vivo antifungal activities of the essential oils of various plants against tomato grey mould disease agent Botrytis cinerea. International Journal of Food Microbiology 143: 183–189. https://doi.org/10.1016/j.ijfoodmicro.2010.08.015.
  74. Szczygieł T., Koziróg A., Otlewska A., 2024. Synthetic and Natural Antifungal Substances in Cereal Grain Protection: A Review of Bright and Dark Sides. Molecules 29: 3780. https://doi.org/10.3390/molecules29163780.
  75. Tang X., Shao Y.-L., Tang Y.-J., Zhou W.-W., 2018. Antifungal Activity of Essential Oil Compounds (Geraniol and Citral) and Inhibitory Mechanisms on Grain Pathogens (Aspergillus flavus and Aspergillus ochraceus). Molecules 23: 2108. https://doi.org/10.3390/molecules23092108.
  76. Tao N., OuYang Q., Jia L., 2014. Citral inhibits mycelial growth of Penicillium italicum by a membrane damage mechanism. Food Control 41: 116–121. https://doi.org/10.1016/j.foodcont.2014.01.010.
  77. Temperini C.V., Tudela M.A.A., Gimenez G.N., Di Masi S.N., Pardo A.G., Pose G.N., 2022. Brown spot of pear, an emerging disease in Argentina: identification and pathogenicity characterization of Argentinean Stemphylium vesicarium isolates. European Journal of Plant Pathology 163: 529–544. https://doi.org/10.1007/s10658-022-02493-y.
  78. Toledo E., Félix C., Vicente T.F.L., Augusto A., Félix R., … Lemos M.F.L., 2023. Seaweed Extracts to Control Postharvest Phytopathogenic Fungi in Rocha Pear. Journal of Fungi 9: 269. https://doi.org/10.3390/jof9020269.
  79. Trombetta D., Castelli F., Sarpietro M.G., Venuti V., Cristani M., … Bisignano G., 2005. Mechanisms of Antibacterial Action of Three Monoterpenes. Antimicrobial Agents and Chemotherapy 49: 2474–2478. https://doi.org/10.1128/AAC.49.6.2474-2478.2005.
  80. Tudela M.A.A., Gimenez G.N., Di Masi S.N., Pose G.N., Basanta M., 2023a. Susceptibility of South American pear cultivars to brown spot of pear caused by Stemphylium vesicarium. Journal of Plant Diseases and Protection 130: 1347–1356. https://doi.org/10.1007/s41348-023-00783-z.
  81. Tudela M.A.A., Lutz M.C., Giménez G.N., Del Brío D., Di Masi S.N., … Molina J.P.E., 2023b. Efficacy of fungicides against brown spot of pear in Argentina. Crop Protection 174: 106425. https://doi.org/10.1016/j.cropro.2023.106425.
  82. Wang H., Yang Z., Ying G., Yang M., Nian Y., … Kong W., 2018. Antifungal evaluation of plant essential oils and their major components against toxigenic fungi. Industrial Crops and Products 120: 180–186. https://doi.org/10.1016/j.indcrop.2018.04.053.
  83. Xylia P., Chrysargyris A., Ahmed Z.F.R., Tzortzakis N., 2021. Application of Rosemary and Eucalyptus Essential Oils and Their Main Component on the Preservation of Apple and Pear Fruits. Horticulturae 7: 479. https://doi.org/10.3390/horticulturae7110479.
  84. Yang J.H., Brannen P.M., Schnabel G., 2015. Resistance in Alternaria alternata to SDHI Fungicides Causes Rare Disease Outbreak in Peach Orchards. Plant Disease 99: 65–70. https://doi.org/10.1094/PDIS-04-14-0387-RE.
  85. Yin Y., Miao J., Shao W., Liu X., Zhao Y., Ma Z., 2023. Fungicide Resistance: Progress in Understanding Mechanism, Monitoring, and Management. Phytopathology 113: 707–718. https://doi.org/10.1094/PHYTO-10-22-0370-KD.