Application of Chitosan in Plant Defense Responses to Biotic and Abiotic Stresses
Keywords:Abiotic stress, Biotic stress, Chitosan, Plant response
Chitosan, a copolymer of N-acetyl-D-glucosamine and D-glucosamine, which possesses properties that make it useful in various fields, is produced by the deacetylation of chitin derivatives. It is used in agriculture as a biostimulant for plant growth and protection, it also induces several responsive genes, proteins, and secondary metabolites in plants. Chitosan elicits a signal transduction pathway and transduces secondary molecules such as hydrogen peroxide and nitric oxide. Under biotic stress, chitosan can stimulate phytoalexins, pathogenesis-related proteins, and proteinase inhibitors. Pretreatment of chitosan before exposure to abiotic stresses (drought, salt, and heat) induces plant growth, production of antioxidant enzymes, secondary metabolites, and abscisic acid (ABA). It also causes changes in physiology, biochemistry, and molecular biology of the plant cells. However, plant responses depend on different chitosan-based structures, concentrations, species, and developmental stages. This review collects updated information on chitosan applications, particularly in plant defense responses to biotic and abiotic stress conditions.
 K. Ohta, S. Morishita, K. Suda, N. Kobayashi, and T. Hosoki, “Effect of chitosan soil mixture treatment in the seedling stage and fIowering of several ornamental plants,” Journal of the Japanese Society for Horticultural Science, vol. 73, pp. 66–68, 2004.
 P. Pornpienpakdee, R. Singhasurasak, P. Chaiyasap, R. Pichyangkura, R. Bunjongrat, S. Chadchawan, and P. Limpanavech, “Improving the micropropagation efficiency of hybrid Dendrobium orchids with chitosan,” Scientia Horticulturae, vol. 124, pp. 490– 499, 2010.
 D. Elieh-Ali-Komi and M. R. Hamblin, “Chitin and chitosan: Production and application of versatile biomedical nanomaterials,” International Journal of Advanced Research, vol. 4, no. 3, pp. 411–427, 2016.
 P. K. Dutta, J. Dutta, and V. S. Tripathi, “Chitin and chitosan: Chemistry, properties and applications,” Journal of Scientific and Industrial Research, vol. 63, pp. 20–31, 2003.
 F. Shahidi, J. K. V. Arachchi, and Y. J, Jeon, “Food applications of chitin and chitosans,” Trends in Food Science and Technology, vol. 10, no. 2, pp. 37–51, 1999.
 S. Islam, M. A. R. Bhuiyan, and M. N. Islam, “Chitin and chitosan: Structure, properties and applications in biomedical engineering,” Journal of polymers and the environment, vol. 25, pp. 854–866, 2017.
 I. Aranaz, M. Mengíbar, R. Harris, I. Panos, B. Miralles, N. Aeosta, G. Galed, and A. Heras, “Functional characterization of chitin and chitosan,” Current Chemical Biology, vol. 3, pp 203–230, 2009.
 M. Malerba and R.Cerana, “Reactive oxygen and nitrogen species in defense/stress responses activated by chitosan in sycamore cultured cells,” International journal of molecular sciences, vol. 16, pp. 3019–3034, 2015.
 A. Hidangmayum, P. Dwivedi, D. Katiyar, and A. Hemantaranjan, “Application of chitosan on plant responses with special reference to abiotic stress,” Physiology and Molecular Biology of Plants, vol. 25, pp. 313–326, 2019.
 H. P. Chen and L. L. Xu, “Isolation and characterization of a novel chitosan-binding protein from non-heading Chinese cabbage leaves,” Journal of Integrative Plant Biology, vol. 47, pp. 452–456, 2005.
 B. E. Amborabe´, J. Bonmort, P. Fleurat-Lessard, and G. Roblin, “Early events induced by chitosan on plant cells,” Journal of Experimental Botany, vol. 59, pp. 2317–2324, 2008.
 E. K. Petutschnig, A. M. E. Jones, L. Serazetdinova, U. Lipka, and V. Lipka, “The lysin motif receptorlike kinase (LysM-RLK) CERK1 is a major chitin-binding protein in Arabidopsis thaliana and subject to chitin-induced phosphorylation,” Journal of Biological Chemistry, vol. 285, pp. 28902–28911, 2010.
 G. Povero, E. Loreti, C. Pucciariello, A. Santaniello, D. D. Tommaso, G. D. Tommaso, D. Kapetis, F. Zolezzi, A. Piaggesi, and P. Perata, “Transcript profiling of chitosan-treated Arabidopsis seedings,” Journal of Plant Research, vol. 124, pp. 619–629, 2011.
 R. Pichyangkura and S. Chadchawan, “Biostimulant activity of chitosan in horticulture,” Scientia Horticulturae, vol. 196, pp. 49–65, 2015.
 W. Pongprayoon, S. Roytrakul, R. Pichayangkura, and S. Chadchawan, “The role of hydrogen peroxide in chitosan-induced resistance to osmotic stress in rice (Oryza sativa L.),” Plant Growth Regulation, vol. 70, pp. 159–173, 2013.
 H. Kohle, W. Jeblick, F. Poten, W. Blaschek, and H. Kauss, “Chitosanelicited callose synthesis in soybean cells as a Ca2+-dependent process,” Plant Physiology, vol. 77, pp. 544–551, 1985.
 F. Faoro, D. Maffi, D. Cantu, and M. Iriti, “Chemical-induced resistance against powdery mildewin barley: The effects of chitosan and benzothiadiazole,” BioControl, vol. 53, pp. 387– 401, 2008.
 A. Zuppini, B. Baldan, R. Millioni, F. Favaron, L. Navazio, and P. Mariani, “Chitosan induces Ca2+ mediated programmed cell death in soybean cells,” New Phytologist, vol. 161, pp. 557–568, 2003.
 L. A. Hadwiger, “Multiple effects of chitosan on plant systems: Solid science or hype,” Plant Science, vol. 208, pp. 42–49, 2013.
 S. H. Doares, T. Syrovets, E. W. Wieler, and A. Ryan, “Oligogalacturonides and chitosan activate plant defensive gene through the octadecanoid pathway,” Proceedings of the National Academy of Sciences of the United States of America, vol. 92, pp. 4095–4098, 1995.
 M. Iriti, V. Picchi, M. Rossoni, S. Gomarasca, N. Ludwig, M. Gargano, and F. Faoro, “Chitosan antitranspirant activity is due to abscisic aciddependent stomatal closure,” Environmental and Experimental Botany, vol. 66, pp. 493–500, 2009.
 H. Yin, S. Li, X. Zhao, Y. Du, and X. Ma, “cDNA microarray analysis of gene expression in Brassica napus treated with oligochitosan elicitor,” Plant Physiology and Biochemistry, vol. 44, pp. 910–916, 2006.
 R. Rakwal, S. Tamogami, G. K. Agrawal, and H. Iwahashi, “Octadecanoid signaling component ‘‘burst’’ in rice (Oryza sativa L.) seedling leaves upon wounding by cut and treatment with fungal elicitor chitosan,” Biochemical and Biophysical Research Communications, vol. 295, no. 5, pp. 1041–1045, 2002.
 M. Iriti and F. Faoro, “Abscisic acid mediates the chitosan-induced resistance in plant against viral disease,” Plant Physiology and Biochemistry, vol. 46, pp. 1106–1111, 2008.
 A. Koornneef and C. M. Pieterse, “Cross talk in defense signaling,” Plant Physiology, vol. 146, no. 3, pp. 839–844, 2008.
 Y. Heng, Z. Xiaoming, and D. Yuguang, “Oligochitosan: A plant diseases vaccine— A review,” Carbohydrate Polymers, vol. 82, pp. 1–8, 2010.
 F. Chen, Q. Li, and Z. He, “Proteomic analysis of rice plasma membrane-associated proteins in response to chitooligosaccharide elicitors,” Journal of Integrative Plant Biology, vol. 49, pp. 863–870, 2007.
 M. Ferri, A. Tassoni, M. Franceschetti, L. Righetti, M. J. Naldrett, and N. Bagni “Chitosan treatment induces changes of protein expression profile and stilbene distribution in Vitis vinifera cell suspensions,” Proteomics, vol 9, no. 3, pp. 610–624, 2009.
 N. Chamnanmanoontham, W. Pongprayoon, R. Pichayangkura, S. Roytrakul, and S. Chadchawan, “Chitosan enhances rice seedling growth via gene expression network between nucleus and chloroplast,” Plant Growth Regulation, vol. 75, pp. 101–114, 2015.
 Z. Li, Y. Zhang, X. Zhang, E. Merewitz, Y. Peng, X. Ma, and Y. Yan, “Metabolic pathways regulated by chitosan contributing to drought resistance in white clover,” Journal of Proteome Research, vol. 16, no. 8, pp. 3039–3052, 2017.
 L. A. Hadwiger and J. M. Beckman, “Chitosan as a component of pea-Fusarium solani interactions,” Plant Physiology, vol. 66, pp. 205–211, 1980.
 U. Conrath, A. Domard, and H. Kauss, “Chitosanelicited synthesis of callose and coumarin derivatives in parsley cell suspension cultures,” Plant Cell Reports, vol. 8, no. 8, pp. 152–155, 1989.
 H. Kauss, W. Jeblik, and A. Domard, “The degree of polymerization and N-acetylation of chitosan determine its ability to elicit callose formation in suspension cells and protoplasts of Catharantus roseus,” Planta, vol. 178, pp. 385–392, 1989.
 W. Wang, S. Li, X. Zhao, B. Lin, and Y. Du, “Determination of six secondary metabolites including chlorogenic acid in tobacco using high performance liquid chromatography with coulometric array detection,” Chinese Journal of Chromatography, vol. 25, no. 6, pp. 848–852, 2007.
 H. Köhle, W. Jeblick, F. Poten, W. Blaschek, and H. Kauss, “Chitosan-elicited callose synthesis in soybean cells as a Ca-dependent process,” Plant Physiology, vol. 77, no. 3, pp. 544–551, 1985.
 P. Vander, K. M. Varum, A. Domard, N. E. E. Gueddari, and B. M. Moerschbacher, “Comparison of the ability of partially N-acetylated chitosans and chitooligosaccharides to elicit resistance reactions in wheat leaves,” Plant Physiology, vol. 118, no. 4, pp. 1353–1359, 1998.
 M. Ghasemnezhad, M. A. Shiri, and M. Sanavi, “Effect of chitosan coatings on some quality indices of apricot (Prunus armeniaca L.) during cold storage,” Caspian Journal Environmental Sciences, vol 8, pp. 25–33, 2010.
 G. Kerch, M. Sabovics, Z. Kruma, S. Kampuse, and E. Straumite, “Effect of chitosan and chitooligosaccharide on vitamin C and polyphenols contents in cherries and strawberries during refrigerated storage,” European Food Research and Technology, vol. 233, pp. 351–358, 2011.
 A. Ali, N. Zahid, S. Manickam, Y. Siddiqui, P. G. Alderson, and M. Maqbool, “Effectiveness of submicron chitosan dispersions in controlling anthracnose and maintaining quality of dragon fruit,” Postharvest Biology and Technology, vol. 86, pp. 147–153, 2013.
 C. R. Allan and L. A. Hadwiger, “The fungicidal effect of chitosan on fungi of varying cell wall composition,” Experimental Mycology, vol. 3, pp. 285–287, 1979.
 M. Malerba and R. Cerana, “Reactive oxygen and nitrogen species indefense/stress responses activated by chitosan in sycamore cultured cells,” International Journal of Molecular Sciences, vol. 16, pp. 3019–3034, 2015.
 D. Katiyar, A. Hemantaranjan, B. Singh, and N. A. Bhanu, “A future perspective in crop protection: Chitosan and its oligosaccharides,” Advances in Plants and Agriculture Research, vol. 1, no. 1, pp. 23–30, 2014.
 M. V. B. Reddy, P. Angers, F. Castaigne, and J. Arul, “Chitosan effects on blackmold rot and pathogenic factors produced by Alternaria alternata in postharvest tomatoes,” Journal of the American Society for Horticultural Science, vol. 125, pp. 742–747, 2000.
 P. Trotel-Aziz, Mn. Couderchet, G. Vernet, and A. Aziz, “Chitosan stimulates defense reactions in grapevine leaves and inhibits development of Botrytis cinereal,” European Journal of Plant Pathology, vol. 114, pp. 405–413, 2006.
 G. Lizama-Uc, I. A. Estrada-Mota, M. G. Caamal-Chan, R. Souza-Perera, C. Oropeza- Salín, I. Islas-Flores, and J. J. Zúniga-Aguilar, “Chitosan activates a MAP-kinase pathway and modifies abundance of defense-related transcripts in calli of Cocos nucifera L.,” Physiological and Molecular Plant Pathology, vol. 70, pp. 130–141, 2007.
 Z. Ma, L. Yang, H. Yan, J. F. Kennedy, and X. Meng, “Chitosan and oligochitosan enhance the resistance of peach fruit to brown rot,” Carbohydrate Polymers, vol. 94, pp. 272–277, 2013.
 A. Ali, N. Zahid, S. Manickam, Y. Siddiqui, P.G. Alderson, and M. Maqbool, “Induction of lignin and pathogenesis related proteins in dragon fruit plants in response to submicron chitosan dispersions,” Crop Protection, vol. 63, pp. 83–88, 2014.
 W. Lin, X. Hu, W. Zhang, W. J. Rogers, and W. Cai, “Hydrogen peroxide mediates defence responses induced by chitosans of different molecular weights in rice,” Journal of Plant Physiolology, vol. 162, pp. 937–944, 2005.
 M. Walker-Simmons, L. Hadwiger, and C. A. Ryan, “Chitosans and pectic polysaccharides both induce the accumulation of the antifungal phytoalexin pisatin in pea pods and antinutrient proteinase inhibitors in tomato leaves,” Biochemical and Biophysical Research Communications, vol. 110, pp. 194–199, 1983.
 M. Walker-Simmons and C. A. Ryan, “Proteinase inhibitor synthesis in tomato leaves,” Plant Physiology, vol. 76, pp. 787–790, 1984.
 F. Yang, J. Hu, J. Li, X. Wu, and Y. Qian, “Chitosan enhances leaf membrane stability and antioxidant enzyme activities in apple seedlings under drought stress,” Plant Growth Regulation, vol. 58, pp. 131– 136, 2009.
 Y. J. Guan, J. Hu, X. Wang, and C. Shao, “Seed priming with chitosan improves maize germination and seedling growth in relation to physiological changes under low temperature stress,” Journal of Zhejiang University. Science. B, vol. 10, no. 6, pp. 427–433, 2009.
 K. Górnik, M. Grzesik, and B. Romanowska-Duda, “The effect of chitosan on rooting of grapevine cuttings and on subsequent plant growth under drought and temperature stress,” Journal of Fruit and Ornamental Plant Research, vol. 16, pp. 333– 343, 2008.
 S. Boonlertnirun, E. Sarobol, S. Meechoui, and I. Sooksathan, “Drought recovery and grain yield potential of rice after chitosan application,” Kasetsart Journal, vol. 41, pp. 1–6, 2007.
 M. Bittelli, M. Flury, G. S. Campbell, and E. J. Nichols, “Reduction of transpiration through foliar application of chitosan,” Agricultural and Forest Meteorology, vol. 107, pp. 167–175, 2001.
 S. Farouk and A. R. Amany, “Improving growth and yield of cowpea by foliar application of chitosan under water stress,” Egyptian Journal of Biology, vol. 14, no. 1, pp. 14–16, 2012.
 Z. E. Bistgani, S. A. Siadat, A. Bakhshandeh, A. G. Pirbalouti, and M. Hashemi, “Interactive effects of drought stress and chitosan application on physiological characteristics and essential oil yield of Thymus daenensis Celak,” The Crop Journal, vol. 5, no. 5, pp. 407–415, 2017.
 A. G. Pirbalouti, F. Malekpoor, A. Salimi, and A. Golparvar, “Exogenous application of chitosan on biochemical and physiological characteristics, phenolic content and antioxidant activity of two species of basil (Ocimum ciliatum and Ocimum basilicum) under reduced irrigation,” Scientia Horticulturae, vol. 217, pp. 114–122, 2017.
 N. Jabeen and R. Ahmad, “The activity of antioxidant enzymes in response to salt stress in safflower (Carthamus tinctorius L.) and sunflower (Helianthus annuus L.) seedlings raised from seed treated with chitosan,” Journal of The Science of Food and Agriculture, vol. 93, no. 7, pp. 1699–1705, 2013.
 B. Mahdavi and A. Rahimi, “Seed priming with chitosan improves the germination and growth performance of ajowan (Carum copticum) under salt stress,” Eurasian Journal of Biosciences, vol. 7, pp. 69–76, 2013.
 B. Mahdavi, “Seed germination and growth responses of Isabgol (Plantago ovata Forsk) to chitosan and salinity,” International Journal of Agriculture and Crop Sciences, vol. 5, pp. 1084– 1088, 2013.
 G. Martı´nez, G. Reyes, R. Falco´n, and V. Nu´n˜ez, “Effect of seed treatment with chitosan on the growth of rice (Oryza sativa L.) seedlings cv. INCA LP-5 in saline medium,” Cultivos Tropicales, vol. 36, no. 1, pp. 143–150, 2015.
 S. R. Ray, M. J. H. Bhuiyan, M. A. Hossain, A. K. Hasan, and S. Sharmin, “Chitosan ameliorates growth and biochemical attributes in mungbean varieties under saline condition,” Research in Agriculture Livestock and Fisheries, vol. 3, no. 1, pp. 45–51, 2016.
 A. R. Al-Tawaha, M. A. Turk, A. R. M. Al- Tawaha, M. H. Alu’datt, M. Wedyan, E. Al-D. M. Al-Ramamneh, and A. T. Hoang, “Using chitosan to improve growth of maize cultivars under salinity conditions,” Bulgarian Journal of Agricultural Science, vol. 24, no. 3, pp. 437–442, 2018.
 L. Ma, Y. Li, C. Yu, Y. Wang, X. Li, N. Li, and N. Bu, “Alleviation of exogenous oligochitosan on wheat seedlings growth under salt stress,” Protoplasma, vol. 249, no. 2, pp. 393–399, 2012.
 B. D. McKersie and Y. Lesheim, Stress and Stress Coping in Cultivated Plants. Berlin, Germany: Springer, 2013.
 Y. Ishibashi, H. Yamagguchi, T. Yuasa, M. Iwaya-Inoue, S. Arima, and S. Zheng, “Hydrogen peroxidase spraying alleviates drought stress in soybean plants,” Journal of Plant Physiology, vol. 168, pp. 1562–1567, 2011.
 Y. S. Choi, Y. M. Kim, O. J. Hwang, Y. J. Han, S. Y. Kim, and J. I. Kim, “Overexpression of Arabidopsis ABF3 gene confers enhanced tolerance to drought and heat stress in creeping bentgrass,” Plant Biotechnology Reports, vol. 7, pp. 165–173, 2013.