Changes of antioxidant enzymes in ‘Thomson-Navel’ orange during induction of resistance to green mold (Penicillium digitatum (Pers.) Sacc.) as provoked by jasmonic acid, epibrassinolide, chitosan and cinnamon essential oil


  • Behrooz ALIJOO Department of Horticultural science, Science and Research Branch, Islamic Azad University, Tehran, Iran.
  • Vahid ABDOSSI Department of Horticultural science, Science and Research Branch, Islamic Azad Universiy, Tehran, Iran.
  • Vahid ZARRINNIA Department of plant protection, Science and Research Branch, Islamic Azad University, Tehran, Iran.
  • Sepideh KALATEH JARI Department of Horticultural science, Science and Research Branch, Islamic Azad Universiy, Tehran, Iran.
  • Mohamad CHAMANI Department of animal science, Science and Research Branch, Islamic Azad Universiy, Tehran, Iran.



peroxidase, catalase, superoxide dismutase, ascorbate peroxidase, citrus green mold, inoculation, lesion diameter


Pathogenic agents are one of the causes of post-harvest citrus fruit loss. Therefore, the aim of this study was to evaluate the effect of post-harvest treatments with jasmonic acid (ja), epibrassinolide (epiBL), chitosan (chi) and cinnamon essential oil (cin) on induction of resistance to the citrus green mold (Penicillium digitatum (Pers.) Sacc.) and reduction of fungal growth by improving the activity of some antioxidant enzymes in ‘Thomson-Navel’orange. For this purpose, a factorial experiment was conducted in a completely randomized design. Treatments included positive and negative control, jasmonic acid (5, 10, 20 and 40 μl l-1), epibrassinolide (1, 4, 7 and 10 μmol l-1), chitosan (2.5, 5, 7.5 and 10 g l-1) and cinnamon essential oil (250, 500, 750 and 1000 ppm). Characteristics such as lesion diameter and activities of antioxidant enzymes including SOD, APX, CAT and POD were evaluated for a period of 96 hours with 24 hour intervals. The results indicated that all treatments significantly decreased the lesion diameter of fruits. Consequently, chitosan treatments (7.5 and 10 g l-1) and cinnamon essential oil (750 and 1000 ppm) inhibited the spread of fungal infection better than other treatments, and therefore reduced the growth of green mold. Also, different concentrations of (ja) and (chi) increased the activity of SOD and APX enzymes, while different concentrations of (epiBR) and (cin) stimulated the activity of POD and CAT enzymes. Finally, the present study proposes using natural products as an appropriate alternative to fungicides in order to reduce the citrus green mold rot


Abdolahi, A., Hassani, A., Ghosta, Y., Javadi, T.and Meshkatalsadat, M. (2010). Essential oils as control agents of postaharvest Alternaria and Penicillium rots on tomato fruits, Journal of Food Safety, 30, 341-352.

Aebi, H. E. (1984). Catalase in vitro. Methods in Enzymology, 105, 121-126.

Asghari, M., Hasanlooe, A.R. (2015). Interaction effects of salicylic acid and methyl jasmonate on total antioxidant content, catalase and peroxidase enzymes activity in 'Sabrosa' strawberry fruit during storage. ScientiaHorticulturae, 197, 490-495.

Asghari, M., Hasanlooe, A.R. (2016). Methyl jasmonate effectively enhanced some defense enzymes activity and Total Antioxidant content in harvested "Sabrosa" strawberry fruit. Food science & nutrition, 4(3), 377-383.

Badawy, M.E.I., Rabea, E.I. (2009). Potential of the biopolymer chitosan with different molecular weights to control postharvest gray mold of tomato fruit. Postharvest Biology and Technology, 51(1), 110-117.

Bautista-Banos, S., Hernandez-Lauzardo, A.N., Velazquez-del Valle, M. G., Hernandez-Lopez, M., AitBarka, E., Bosquez-Molina, E. and Wilson, C. L. (2006). Chitosan as Potential Natural Compounds to Control Pre and Postharvest Diseases of Horticultural Commodities. Crop Protection, 25: 108-118.

Bendahou, M., Muselli, A., Grignon-Dubois, M., Benyoucef, M., Desjobert, J.M., Bernardini, A.F. (2008). Antimicrobial activity and chemical composition of Origanum glandulosum Desf. essential oil and extract obtained by microwave extraction: Comparison with hydro distillation. Food Chemistry, 106(1), 132-139.

Champa, W.A.H., Gill1, MIS., Mahajan, BVC., Aror, NK., BediS. (2015). Brassinosteroids improve quality of table grapes (Vitis vinifera 'Flame Seedless'. Tropical Agricultrucal Research, 26(2):368 - 379.

Chien, P.J., Sheu, F., Lin, H.R. (2007). Coating citrus ('Murcott Tangor') fruit with low molecular weight chitosan increases postharvest quality and shelf life. Food chemistry, 100(3), 1160-1164.

Coll, Y., Coll, F., Amorós, A., Pujol, M. (2015). Brassinosteroids roles and applications: an up-date.Biologia, 70(6):726-732 doi: 10.1515/biolog-2015-0085.

Ding, C.K., Wang, C.Y., Gross, K.C., Kenneth, C.S., David, L. (2002). Jasmonate and salicylate induce the expression of pathogenesis-related-protein genes and increase resistance to chilling injury in tomato fruit. Planta 214, 895-901.

Droby, S., Wisniewski, M., Macarisin, D. and Wilson, C. (2009). Twenty years of postharvest biocontrol research: is it time for a new paradigm. Postharvest Biology and Technology, 52,137e145.

El guilli, M., Hamza, A., Clement, C., Ibriz, M., Aitbarak, E. (2016). Effectiveness of postharvest treatment with chitosan to control citrus green mold. Agriculture, 6(2), 12.

FAO/WHO (2019).

Ge, Y.H., Li, C.Y., Tang, R.X., Sun, R.H., Li, J.R. (2015). Effects of postharvest brassinolide dipping on quality parameters and antioxidant activity in peach fruit. In III International Symposium on Postharvest Pathology, Using Science to Increase Food Availability, 1144.377-384.

Giannopolitis, S. (1977). Superoxide dismutase. I. Occurrence in higher plants. Plant Physiology, 59, 309-314.

González-Aguilar, G., Buta, J., Wang, C. (2003). Methyl jasmonate and modified atmosphere packaging (MAP) reduce decay and maintain postharvest quality of papaya 'Sunrise'. Postharvest Biology and Technology, 28, 361-370.

Guo, J., Fang, W., Lu, H., Zhu, R., Lu, L., Zheng, X., Yu, T. (2014). Inhibition of green mold disease in mandarins by preventive applications of methyl jasmonate and antagonistic yeast Cryptococcus laurentii. Postharvest biology and technology, 88, 72-78.

Han, J. H., Tao, W.Y., Hao, H.K., Zhang, B.L., Jiang, W.B., Niu, T.G., et al. (2006). Physiology and quality responses of fresh-cut broccoli florets pretreated with ethanol vapor. Journal of Food Science, 71, 385-389.

Jayaraj, J., Rahman, M., Wan, A., Punja, Z.K. (2009). Enhanced resistance to foliar fungal pathogens in carrot by application of elicitors. Annals of Applied Biology, 155, 71-80.

Jiménez-Reyes, M.F., Carrasco, H., Olea, A., Silva-Moreno, E. (2018). Natural compounds: A sustainable alternative for controlling phytopathogens. PeerJ Reprints, 6, e26664v1.

Jongsri, P., Wangsomboondee, T., Rojsitthisak, P., Seraypheap, K. (2016). Effect of molecular weights of chitosan coating on postharvest quality and physicochemical characteristics of mango fruit. LWT-Food Science and Technology, 73, 28-36.

Ladanyia, M., Ladaniya, M. (2010). Citrus Fruit: Biology, Technology and Evaluation. Academic Press: San Diego, CA, USA.

Liu, Q., Xi, Z., Gao, J., Meng, Y., Lin, S., Zhang, Z. (2016). Effects of exogenous 24 epibrassinolide to control grey mould and maintain postharvest quality of table grapes. International Journal of Food Science and Technology, 51(5), 1236-1243.

Luan, L. Y, Zhang. Z. W, Xi, Z. M, Huo, S. S, Ma, L. N. (2013). Brassinosteroids regulate anthocyanin biosynthesis in the ripening of grape berries. South African Journal of Enology and Viticulture, 34(2):196-203.

Mac-Adam, J.W., Nelson, C.J., Sharp, R.E. (1992). Peroxidase activity in the leaf elongation zone of tall fescue I. Spatial distribution of ionically bound peroxidase activity in genotypes differing in length of the elongation zone. Plant Physiology, 99(3), 872-878.

N. G. Tzortzakis, C. M. Economakis. (2007). Innovative Food Science and Emerging Technologies, 8, 253.

Peng, J., Zheng, Y., Tang, S., Rui, H., Wang, C. (2009). Enhancing disease resistance in peach fruit with methyl jasmonate. Journal of the Science of Food and Agriculture, 89(5), 802-808.

Pretorius, J.C., Zietsman, P.C., Eksteen, D. (2002). Fungitoxic properties of selected South African plant species against plant pathogens of economic importance in agriculture. Annals of Applied Biology, 141(2), 117-124.

Qin, G.Z., Tian, S.P., Xu, Y., Wan, Y.K. (2003). Enhancement of biocontrol efficacy of antagonistic yeasts by salicylic acid in sweet cherry fruit. Physiological and Molecular Plant Pathology, 62(3), 147-154.

Ranieri, A., Castagna, J., Pacini, B., Baldan, A., Mensuali. Sodi, G.F. (2003). Soldatini Early production and scavenging of hydrogen peroxide in the apoplast of sunflowers plants exposed to ozone. Journal of Experimental Botany, 54, 2529-2540.

Reddy, M.V.B., Angers, P., Gosselin, A., Arul, J. (1997). Characterization and use of essential oil from Thymus vulgaris against Botrytis cinerea and Rhizopus stolonifer in strawberry fruits. Phytochemistry, 47, 1515-1520.

Romanazzi, G.; Murolo, S.; Feliziani, E. (2013). Effects of an innovative strategy to contain grapevine Bois noir: Field treatment with resistance inducers. Phytopathology, 103, 785-791.

Shan, B., Cai, Y.Z., Brooks, J.D., Corke, H. (2007). The in vitro antibacterial activity of dietary spice and medicinal herb extracts. International Journal of food microbiology, 117(1), 112-119.

Sharma, R.R., Singh, D., Singh, R. (2009). Biological control of postharvest diseases of fruits and vegetables by microbial antagonists: A review. Biological control, 50, 205-221.

T, M., Ibrahim, S., Cliver, D. (2010).Antimicrobial herb and spice compounds in food. Food Control, 21(9), 1199-1218.

Walters, D., Wash, D., Newton, A., Lyon, G. (2005). Induced resistance for plant disease control: Maximizing the efficacy of resistance elicitors. Phytopathology, 95, 1368-1373.

Wang Q, Lai TF, Qin GZ, Tian SP. (2009). Response of jujube fruits to exogenous oxalic acid treatment based on proteomic analysis. Plant Cell Physiol, 50:230-242.

Wang, C.Y., Fung, R., Ding, C. (2003). Reducing chilling injury and enhancing transcript levels of heat Shock proteins, pr-proteins and alternative oxidase by methyl jasmonate and methyl salcylate in tomatoes and peppers. Meeting Abstract, 38, 860.

Wang, K.T., Jin, P., Han, L., Shang, H.T., Tang, S.S., Rui, H.J., et al. (2014). Methyl jasmonate induces resistance against Penicillium citrinum in Chinese bayberry by priming of defense responses. Postharvest Biology and Technology, 98, 90-97.

Wasternack, C., Hause, B. (2013). Jasmonates-Biosynthesis and Role in Stress Responses and Developmental Processes. Annals of Botany, 111, 1021-1058.

Win, N.K.K., Jitareerat, P., Kanlayanarat, S., Sangchote, S. (2007). Effects of cinnamon extract, chitosan coating, hot water treatment and their combinations on crown rot disease and quality of banana fruit. Postharvest biology and technology, 45(3), 333-340.

Wu, Y., Duan, X., Jing, G., Ouyang, Q., Tao, N. (2017). Cinnamaldehyde inhibits the mycelial growth of Geotrichum citri-aurantii and induces defense responses against sour rot in citrus fruit. Postharvest Biology and Technology, 129, 23-28.

Xu, W.T., Peng, X.L., Luo, Y.B., Wang, J., Guo, X., Huang, K.L. (2009). Physiological and biochemical responses of grape fruit seed extract dip on 'Redglobe' grape. LWT-Food Science and Technology, 42, 471-476.

Yao, H.J., Tian, S.P. (2005). Effects of a biocontrol agent and methyl jasmonate on postharvest diseases of peach fruit and the possible mechanisms involved. Journal of Applied Microbiology, 98(4), 941-950.

Zeng, K., Deng, Y., Ming, J., Deng, L. (2010). Induction of disease resistance and ROS metabolism in navel oranges by chitosan. Scientia Horticulturae, 126, 223-228.

Zhang, H., Li, R. and Liu, W. (2011). Effects of Chitin and Its Derivative Chitosan on Postharvest Decay of Fruits: A Review. International Journal of Molecular Sciences, 12(2), 917-934.

Zhu, F., Yun, Z., Ma, Q. (2015). Effects of exogenous 24-epibrassinolide treatment on postharvest quality and resistance of Satsuma mandarin (Citrus unshiu). Postharvest Biology and Technology, 100, 8-15.

Zhu, Z., Zhang. Z.Q., Qin, G.Z., Tian, S.P. (2010). Effects of brassinosteroids on postharvest disease and senescence of jujube fruit in storage. Postharvest Biology and Technology, 56, 50-55.



13. 12. 2019



Agronomy section

How to Cite

ALIJOO, B., ABDOSSI, V., ZARRINNIA, V., KALATEH JARI, S., & CHAMANI, M. (2019). Changes of antioxidant enzymes in ‘Thomson-Navel’ orange during induction of resistance to green mold (Penicillium digitatum (Pers.) Sacc.) as provoked by jasmonic acid, epibrassinolide, chitosan and cinnamon essential oil. Acta Agriculturae Slovenica, 114(2), 239–250.

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