{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,30]],"date-time":"2026-09-30T06:03:59Z","timestamp":1790748239937,"version":"4.1.0"},"reference-count":104,"publisher":"MDPI AG","issue":"10","license":[{"start":{"date-parts":[[2020,5,17]],"date-time":"2020-05-17T00:00:00Z","timestamp":1589673600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Molecules"],"abstract":"<jats:p>Cucumber mosaic cucumovirus (CMV) is a deadly plant virus that results in crop-yield losses with serious economic consequences. In recent years, environmentally friendly components have been developed to manage crop diseases as alternatives to chemical pesticides, including the use of natural compounds such as glycine betaine (GB) and chitosan (CHT), either alone or in combination. In the present study, the leaves of the cucumber plants were foliar-sprayed with GB and CHT\u2014either alone or in combination\u2014to evaluate their ability to induce resistance against CMV. The results showed a significant reduction in disease severity and CMV accumulation in plants treated with GB and CHT, either alone or in combination, compared to untreated plants (challenge control). In every treatment, growth indices, leaf chlorophylls content, phytohormones (i.e., indole acetic acid, gibberellic acid, salicylic acid and jasmonic acid), endogenous osmoprotectants (i.e., proline, soluble sugars and glycine betaine), non-enzymatic antioxidants (i.e., ascorbic acid, glutathione and phenols) and enzymatic antioxidants (i.e., superoxide dismutase, peroxidase, polyphenol oxidase, catalase, lipoxygenase, ascorbate peroxidase, glutathione reductase, chitinase and \u03b2-1,3 glucanase) of virus-infected plants were significantly increased. On the other hand, malondialdehyde and abscisic acid contents have been significantly reduced. Based on a gene expression study, all treated plants exhibited increased expression levels of some regulatory defense genes such as PR1 and PAL1. In conclusion, the combination of GB and CHT is the most effective treatment in alleviated virus infection. To our knowledge, this is the first report to demonstrate the induction of systemic resistance against CMV by using GB.<\/jats:p>","DOI":"10.3390\/molecules25102341","type":"journal-article","created":{"date-parts":[[2020,5,20]],"date-time":"2020-05-20T10:37:38Z","timestamp":1589971058000},"page":"2341","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":109,"title":["Improving Regulation of Enzymatic and Non-Enzymatic Antioxidants and Stress-Related Gene Stimulation in Cucumber mosaic cucumovirus-Infected Cucumber Plants Treated with Glycine Betaine, Chitosan and Combination"],"prefix":"10.3390","volume":"25","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-2685-5851","authenticated-orcid":false,"given":"Ahmed R.","family":"Sofy","sequence":"first","affiliation":[{"name":"Botany and Microbiology Department, Faculty of Science, Al-Azhar University, Nasr City, Cairo 11884, Egypt"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rehab A.","family":"Dawoud","sequence":"additional","affiliation":[{"name":"Department of Virus Research, Plant Pathology Research Institute, ARC, Giza 12619, Egypt"},{"name":"Department of Biology, Faculty of Science, Jazan University, Box 114, Jazan 45142, Saudi Arabia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1727-430X","authenticated-orcid":false,"given":"Mahmoud R.","family":"Sofy","sequence":"additional","affiliation":[{"name":"Botany and Microbiology Department, Faculty of Science, Al-Azhar University, Nasr City, Cairo 11884, Egypt"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Heba I.","family":"Mohamed","sequence":"additional","affiliation":[{"name":"Department of Biological and Geological Sciences, Faculty of Education, Cairo 11566, Egypt"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ahmed A.","family":"Hmed","sequence":"additional","affiliation":[{"name":"Botany and Microbiology Department, Faculty of Science, Al-Azhar University, Nasr City, Cairo 11884, Egypt"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Noha K.","family":"El-Dougdoug","sequence":"additional","affiliation":[{"name":"Botany and Microbiology Department, Faculty of Science, Benha University, Benha 13518, Egypt"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2020,5,17]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"59","DOI":"10.1046\/j.1364-3703.2001.00058.x","article-title":"Cucumber mosaic virus, a model for RNA virus evolution","volume":"2","author":"Roossinck","year":"2001","journal-title":"Mol. Plant. Pathol."},{"key":"ref_2","doi-asserted-by":"crossref","unstructured":"Agrios, G.N. (2005). Plant disease caused by viruses. Plant Pathology, Elsevier-Academic Press.","DOI":"10.1016\/B978-0-08-047378-9.50020-8"},{"key":"ref_3","first-page":"714","article-title":"Attenuation of viral symptoms and yield reduction responses to CMV infection in tomato crops subjected to plant growth promoting rhizobacteria","volume":"25","author":"Montasser","year":"2011","journal-title":"Fed. Am. Soc. Exp. Biol. J."},{"key":"ref_4","doi-asserted-by":"crossref","unstructured":"Sofy, A.R., Sofy, M.R., Hmed, A.A., and El-Dougdoug, N.K. (2019). Potential effect of plant growth-promoting rhizobacteria (PGPR) on enhancing protection against viral diseases. Field Crops: Sustainable Management by PGPR, Springer International Publishing.","DOI":"10.1007\/978-3-030-30926-8_15"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"480","DOI":"10.1016\/j.plaphy.2007.03.002","article-title":"Physiological and metabolic changes of Cucurbita pepo leaves in response to Zucchini yellow mosaic virus (ZYMV) infection and salicylic acid treatments","volume":"45","author":"Radwan","year":"2007","journal-title":"Plant Physiol. Biochem."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"891","DOI":"10.1007\/s11738-010-0477-y","article-title":"Modifications of antioxidant activity and protein composition of bean leaf due to Bean yellow mosaic virus infection and salicylic acid treatments","volume":"32","author":"Radwan","year":"2010","journal-title":"Acta Physiol. Plant."},{"key":"ref_7","first-page":"2019","article-title":"Metabolic changes of cucumber plants due to two CMV Egyptian isolates","volume":"26","author":"Farahat","year":"2018","journal-title":"Arab Univ. J. Agric. Sci."},{"key":"ref_8","first-page":"61","article-title":"Physiological and biochemical responses in Cucurbita pepo leaves associated with some elicitors-induced systemic resistance against Zucchhini yellow mosaic virus","volume":"4","author":"Sofy","year":"2014","journal-title":"Int. J. Mod. Bot."},{"key":"ref_9","first-page":"1010","article-title":"Isozyme markers for response of wild potato species to potato spindle tuber viroid Egyptian isolate","volume":"27","author":"Sofy","year":"2013","journal-title":"World Appl. Sci. J."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"297","DOI":"10.1046\/j.1439-0434.2000.00503.x","article-title":"Role of reactive oxygen species and antioxidant enzymes in systemic Virus infections of plants","volume":"148","year":"2000","journal-title":"J. Phytopathol."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"16203","DOI":"10.1038\/nplants.2016.203","article-title":"ROS accumulation and antiviral defence control by microRNA528 in rice","volume":"3","author":"Wu","year":"2017","journal-title":"Nat. Plants"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"581","DOI":"10.1055\/s-2005-873009","article-title":"Significance of flavonoids in plant resistance and enhancement of their biosynthesis","volume":"7","author":"Treutter","year":"2005","journal-title":"Plant Biol."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1007\/s10343-017-0387-7","article-title":"Association of increase in some biochemical components with flax resistance to powdery mildew","volume":"69","author":"Aly","year":"2017","journal-title":"Gesunde Pflanzen"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"353","DOI":"10.3329\/bjb.v43i3.21613","article-title":"Use of phenols, peroxidase, and polyphenoloxidase of seed to quantify resistance of cotton genotypes to fusarium wilt disease","volume":"43","author":"Asran","year":"2014","journal-title":"Bangladesh J. Bot."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"149","DOI":"10.5423\/PPJ.2012.28.2.149","article-title":"Examination of correlations between several biochemical components and powdery mildew resistance of flax cultivars","volume":"28","author":"Aly","year":"2012","journal-title":"Plant Pathol. J."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"386","DOI":"10.1590\/S1982-56762012000600002","article-title":"Association of oxidative stress components with resistance to flax powdery mildew","volume":"37","author":"Mohamed","year":"2012","journal-title":"Trop. Plant Pathol."},{"key":"ref_17","doi-asserted-by":"crossref","unstructured":"Boobis, A.R., Ossendorp, B.C., Banasiak, U., Hamey, P.Y., Sebestyen, I., and Moretto, A. (2008). Cumulative risk assessment of pesticide residues in food. Toxicol. Lett., 137\u2013150.","DOI":"10.1016\/j.toxlet.2008.06.004"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"117","DOI":"10.1016\/j.pestbp.2017.04.001","article-title":"Changes in antioxidants potential, secondary metabolites and plant hormones induced by different fungicides treatment in cotton plants","volume":"142","author":"Mohamed","year":"2017","journal-title":"Pest Biochem Physiol."},{"key":"ref_19","first-page":"8585","article-title":"The role of systemic and non-systemic fungicides on the physiological and biochemical parameters in Gossypium hirsutum plant, implications for defense responses","volume":"27","author":"Mohamed","year":"2018","journal-title":"Fresenius Environ. Bull."},{"key":"ref_20","doi-asserted-by":"crossref","unstructured":"Aly, A.A., Mohamed, H.I., Mansour, M.T.M., and Omar, M.R. (2013). Suppression of powdery mildew on flax by foliar application of essential oils. J. Phytopathol., 376\u2013381.","DOI":"10.1111\/jph.12080"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"1","DOI":"10.2478\/v10102-009-0001-7","article-title":"Impact of pesticides use in agriculture: Their benefits and hazards","volume":"2","author":"Aktar","year":"2009","journal-title":"Interdiscip. Toxicol."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"197","DOI":"10.1016\/S0308-8146(03)00063-3","article-title":"Glycine betaine and glycine betaine analogues in common foods","volume":"83","author":"Slow","year":"2003","journal-title":"Food Chem."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"206","DOI":"10.1016\/j.envexpbot.2005.12.006","article-title":"Roles of glycine betaine and proline in improving plant abiotic stress resistance","volume":"59","author":"Ashraf","year":"2007","journal-title":"Environ. Exp. Bot."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"891","DOI":"10.1007\/s00253-007-1222-3","article-title":"Co-ordinate expression of glycine betaine synthesis genes linked by the FMDV 2A region in a single open reading frame in Pichia pastoris","volume":"77","author":"Wang","year":"2007","journal-title":"Appl. Microbiol. Biotechnol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"45","DOI":"10.1016\/j.ijfoodmicro.2012.04.010","article-title":"Pretreatment of the yeast antagonist, Candida oleophila, with glycine betaine increases oxidative stress tolerance in the microenvironment of apple wounds","volume":"157","author":"Sui","year":"2012","journal-title":"Int. J. Food Microbiol."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"55","DOI":"10.1016\/j.biocontrol.2017.02.011","article-title":"Mechanisms of glycine betaine enhancing oxidative stress tolerance and biocontrol efficacy of Pichia caribbica against blue mold on apples","volume":"108","author":"Zhang","year":"2017","journal-title":"Biol. Control."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"706","DOI":"10.1023\/A:1013144000718","article-title":"Influence of the chitosan oligomer on the phage particles and reproduction of phage 1-97A in the culture of Bacillus thuringiensis","volume":"70","author":"Kochkina","year":"2001","journal-title":"Microbiology"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1023\/A:1013206517442","article-title":"The antiviral activity of chitosan (review)","volume":"38","author":"Chirkov","year":"2002","journal-title":"Appl. Biochem. Microbiol."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1016\/0885-5765(89)90012-X","article-title":"Chitin oligosaccharides elicit lignification in wounded wheat leaves","volume":"34","author":"Barber","year":"1989","journal-title":"Physiol. Mol. Plant Pathol."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"307","DOI":"10.1046\/j.1365-313X.1993.04020307.x","article-title":"Specific perception of subnanomolar concentrations of chitin fragments by tomato cells: Induction of extracellular alkalinization, changes in protein phosphorylation, and establishment of a refractory state","volume":"4","author":"Felix","year":"1993","journal-title":"Plant J."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"111","DOI":"10.1111\/j.1399-3054.1997.tb03460.x","article-title":"N-acetylchitooligosaccharides elicit expression of a single (13)-beta-glucanase gene in suspension-cultured cells from barley (Hordeum vulgare)","volume":"100","author":"Kaku","year":"1997","journal-title":"Physiol. Plant."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"405","DOI":"10.1271\/bbb.57.405","article-title":"Induction of phytoalexin formation in suspension-cultured rice cells by N-Acetyl-chitooligosaccharides","volume":"57","author":"Yamada","year":"1993","journal-title":"Biosci. Biotechnol. Biochem."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"138","DOI":"10.1007\/BF01276805","article-title":"EPR evidence for generation of hydroxyl radical triggered by N-acetylchitooligosaccharide elicitor and a protein phosphatase inhibitor in suspension-cultured rice cells","volume":"188","author":"Kuchitsu","year":"1995","journal-title":"Protoplasma"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"387","DOI":"10.1104\/pp.110.2.387","article-title":"Involvement of jasmonic acid in elicitor-induced phytoalexin production in suspension-cultured rice cells","volume":"110","author":"Nojiri","year":"1996","journal-title":"Plant Physiol."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"240","DOI":"10.3923\/ijv.2012.240.257","article-title":"A new Egyptian satellite strain of Cucumber mosaic cucumovirus","volume":"8","author":"Megahed","year":"2012","journal-title":"Int. J. Virol."},{"key":"ref_36","first-page":"178","article-title":"Molecular identification of a Cucumber mosaic virus subgroup I Egyptian isolate from geranium based on bioinformatics analysis of CP gene sequence Egypt","volume":"8","author":"Sofy","year":"2011","journal-title":"Egypt. J. Virol."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"91","DOI":"10.1016\/j.aoas.2012.08.001","article-title":"Induced systemic resistance against Cucumber mosaic cucumovirus and promotion of cucumber growth by some plant growth-promoting rhizobacteria","volume":"57","author":"Oraby","year":"2012","journal-title":"Ann. Agric. Sci."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"70","DOI":"10.3923\/ijpp.2014.70.83","article-title":"Molecular identification and analysis of coat protein gene of Cucucmber mosaic cucumovirus sugar beet Egyptian isolate","volume":"5","author":"Megahed","year":"2014","journal-title":"Int. J. Plant Pathol."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"326","DOI":"10.21608\/nrmj.2019.30612","article-title":"Detection of Cucumber mosaic cucumovirus in infected cowpea plants (Vigna unguiculata L.) from northern Egypt","volume":"3","author":"Younes","year":"2019","journal-title":"Nov. Res. Microbiol. J."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"473","DOI":"10.1007\/s11103-008-9435-0","article-title":"Role of plant hormones in plant defence responses","volume":"69","author":"Bari","year":"2009","journal-title":"Plant Mol. Biol."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"6889","DOI":"10.3390\/ijms14046889","article-title":"Auxin and cytokinin metabolism and root morphological modifications in Arabidopsis thaliana seedlings infected with Cucumber mosaic virus (CMV) or exposed to cadmium","volume":"14","author":"Vitti","year":"2013","journal-title":"Int. J. Mol. Sci."},{"key":"ref_42","unstructured":"Lavanya, N.S., and Amruthesh, N.K. (2017). Glycine betaine mediated disease resistance against Sclerospora graminicola in Pearl Millet. J. Appl. Biol. Biotechnol., 5."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"1001","DOI":"10.2307\/3870211","article-title":"Decreased susceptibility to viral disease of [beta]-1,3-glucanase-deficient plants generated by antisense transformation","volume":"8","author":"Beffa","year":"1996","journal-title":"Plant Cell"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"1520","DOI":"10.3389\/fpls.2016.01520","article-title":"Trichoderma harzianum T-22 induces systemic resistance in tomato infected by Cucumber Mosaic Virus","volume":"7","author":"Vitti","year":"2016","journal-title":"Front. Plant Sci."},{"key":"ref_45","first-page":"107","article-title":"Salix alba extract induces systemic resistance in Cucumis sativus infected by Cucumber mosaic virus","volume":"16","author":"Sofy","year":"2018","journal-title":"Nature and Science"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"286","DOI":"10.1034\/j.1399-3054.2000.110220.x","article-title":"Inhibition of photosynthesis by viral infection: Effect on PSII structure and function","volume":"110","author":"Rahoutei","year":"2000","journal-title":"Physiol. Plant."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"203","DOI":"10.1111\/j.1399-3054.1997.tb04776.x","article-title":"Concepts of plant biotic stress. Some insights into the stress physiology of virus-infected plants, from the perspective of photosynthesis","volume":"100","author":"Balachandran","year":"1997","journal-title":"Physiol. Plant."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"4522","DOI":"10.3390\/ijms12074522","article-title":"Antiviral activity and mechanism of action of novel thiourea containing chiral phosphonate on Tob","volume":"12","author":"Fan","year":"2011","journal-title":"Mosaic Virus. Int. J. Mol. Sci."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"9","DOI":"10.1016\/j.plaphy.2014.12.009","article-title":"Identification and expression of a stearoyl-ACP desaturase gene responsible for oleic acid accumulation in Xanthoceras sorbifolia seeds","volume":"87","author":"Zhao","year":"2015","journal-title":"Plant Physiol. Biochem."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"373","DOI":"10.3389\/fpls.2015.00373","article-title":"Anatomy of a nonhost disease resistance response of pea to Fusarium solani: PR gene elicitation via DNase, chitosan and chromatin alterations","volume":"6","author":"Hadwiger","year":"2015","journal-title":"Front. Plant Sci."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"225","DOI":"10.1007\/s12298-015-0292-4","article-title":"Influence of exogenous application of glycinebetaine on antioxidative system and growth of salt-stressed soybean seedlings (Glycine max L.)","volume":"21","author":"Malekzadeh","year":"2015","journal-title":"Physiol. Mol. Biol. Plants"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"837","DOI":"10.1111\/j.1365-3040.2009.01962.x","article-title":"Reassessing the role of phospholipase D in the Arabidopsis wounding response","volume":"32","author":"Bargmann","year":"2009","journal-title":"Plant Cell Env."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"379","DOI":"10.1094\/MPMI-09-14-0259-R","article-title":"Salicylic acid inhibits the replication of Tomato bushy stunt virus by directly targeting a host component in the replication complex","volume":"28","author":"Tian","year":"2015","journal-title":"Mol. Plant Microbe Interact."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.plantsci.2015.01.011","article-title":"Is modulating virus virulence by induced systemic resistance realistic?","volume":"234","author":"Faoro","year":"2015","journal-title":"Plant Sci."},{"key":"ref_55","doi-asserted-by":"crossref","unstructured":"Jin, L., Qin, Q., Wang, Y., Pu, Y., Liu, L., Wen, X., Ji, S., Wu, J., Wei, C., and Ding, B. (2016). Rice dwarf virus P2 protein hijacks auxin signaling by directly targeting the rice OsIAA10 protein, enhancing viral infection and disease development. PLoS Pathog., 12.","DOI":"10.1371\/journal.ppat.1005847"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"1012","DOI":"10.1104\/pp.113.220541","article-title":"Strigolactones stimulate internode elongation independently of gibberellins","volume":"163","author":"Ligerot","year":"2013","journal-title":"Plant Physiol."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"1999","DOI":"10.1093\/pcp\/pct137","article-title":"Jasmonic acid negatively regulates resistance to Tobacco mosaic virus in tobacco","volume":"54","author":"Oka","year":"2013","journal-title":"Plant Cell Physiol."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"355","DOI":"10.1146\/annurev.arplant.48.1.355","article-title":"Biosynthesis and action of jasmonates in plants","volume":"48","author":"Creelman","year":"1997","journal-title":"Annu. Rev. Plant Physiol. Plant Mol. Biol."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"177","DOI":"10.1094\/MPMI-08-13-0216-R","article-title":"The abscisic acid pathway has multifaceted effects on the accumulation of Bamboo Mosaic Virus","volume":"27","author":"Alazem","year":"2014","journal-title":"Mol. Plant Microbe Interact."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"1760","DOI":"10.3389\/fpls.2017.01760","article-title":"Antiviral roles of abscisic acid in plants","volume":"8","author":"Alazem","year":"2017","journal-title":"Front. Plant Sci."},{"key":"ref_61","first-page":"138","article-title":"Callose homeostasis at plasmodesmata: Molecular regulators and developmental relevance","volume":"5","author":"Geelen","year":"2014","journal-title":"Front. Plant Sci."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"1187","DOI":"10.1111\/nph.12436","article-title":"A novel role of PR2 in abscisic acid (ABA) mediated, pathogen-induced callose deposition in Arab","volume":"200","author":"Oide","year":"2013","journal-title":"Thaliana. New Phytol."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"10328","DOI":"10.1073\/pnas.95.17.10328","article-title":"Defense gene induction in tobacco by nitric oxide, cyclic GMP, and cyclic ADP-ribose","volume":"95","author":"Durner","year":"1998","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"493","DOI":"10.4161\/psb.4.6.8392","article-title":"Biosynthesis of salicylic acid in plants","volume":"4","author":"Chen","year":"2009","journal-title":"Plant Signal. Behav."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"643","DOI":"10.1111\/j.1365-313X.1992.tb00133.x","article-title":"Salicylic acid and plant disease resistance","volume":"2","author":"Malamy","year":"1992","journal-title":"Plant J."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"1004","DOI":"10.1126\/science.250.4983.1004","article-title":"Increase in salicylic acid at the onset of systemic acquired resistance in cucumber","volume":"250","author":"Metraux","year":"1990","journal-title":"Science"},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"5487","DOI":"10.1093\/jxb\/ers212","article-title":"Analysis of the antioxidant response of Nicotiana benthamiana to infection with two strains of Pepper Mild Mottle Virus","volume":"63","author":"Hakmaoui","year":"2012","journal-title":"J. Exp. Bot."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"155","DOI":"10.1080\/10715769900301451","article-title":"Elevation of glutathione level and activation of glutathione-related enzymes affect virus infection in tobacco","volume":"31","author":"Gullner","year":"2016","journal-title":"Free Radic. Res."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"1291","DOI":"10.1104\/pp.105.067686","article-title":"Ascorbic acid deficiency activates cell death and disease resistance responses in Arab","volume":"139","author":"Pavet","year":"2005","journal-title":"Plant Physiol."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"212","DOI":"10.3923\/ppj.2015.212.222","article-title":"Biochemical Characterization of compatible plant virus interaction: A case study with Bunchy top virus-banana host-pathosystem","volume":"14","author":"Anuradha","year":"2015","journal-title":"Plant Pathol. J."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"73","DOI":"10.1016\/0048-4059(82)90025-X","article-title":"Association of enhanced peroxidase activity with induced systemic resistance of cucumber to Colletotrichum Lagenarium","volume":"20","author":"Hammerschmidt","year":"1982","journal-title":"Physiol. Plant Pathol."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"73","DOI":"10.1034\/j.1399-3054.2002.1140111.x","article-title":"Involvement of putative chemical wound signals in the induction of phenolic metabolism in wounded lettuce","volume":"114","author":"Saltveit","year":"2002","journal-title":"Physiol. Plant."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"110","DOI":"10.1016\/j.postharvbio.2008.09.002","article-title":"Defense responses of harvested tomato fruit to burdock fructooligosaccharide, a novel potential elicitor","volume":"52","author":"Wang","year":"2009","journal-title":"Postharvest Biol. Technol."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"1439","DOI":"10.1007\/BF00016484","article-title":"The salicylic acid signal in plants","volume":"26","author":"Klessig","year":"1994","journal-title":"Plant Mol. Biol."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"909","DOI":"10.1016\/j.plaphy.2010.08.016","article-title":"Reactive oxygen species and antioxidant machinery in abiotic stress tolerance in crop plants","volume":"48","author":"Gill","year":"2010","journal-title":"Plant Physiol. Biochem."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"589","DOI":"10.1111\/mpp.12034","article-title":"Pepino mosaic virus triple gene block protein 1 (TGBp1) interacts with and increases tomato catalase 1 activity to enhance virus accumulation","volume":"14","author":"Mathioudakis","year":"2013","journal-title":"Mol. Plant Pathol."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"1673","DOI":"10.1104\/pp.108.4.1673","article-title":"Hydrogen peroxide stimulates salicylic acid biosynthesis in tobacco","volume":"108","author":"Leon","year":"1995","journal-title":"Plant Physiol."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"381","DOI":"10.1111\/j.1365-313X.2004.02142.x","article-title":"Plant growth-promoting rhizobacteria systemically protect Arabidopsis thaliana against Cucumber mosaic virus by a salicylic acid and NPR1-independent and jasmonic acid-dependent signaling pathway","volume":"39","author":"Ryu","year":"2004","journal-title":"Plant J."},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"543","DOI":"10.1111\/j.1744-7348.1996.tb05775.x","article-title":"Resistance of tomato infected with cucumber mosaic virus satellite RNA to potato spindle tuber viroid","volume":"129","author":"Yang","year":"1996","journal-title":"Ann. Appl. Biol."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"127","DOI":"10.1023\/A:1009923702103","article-title":"Induction of resistance in tomato against Cucumber mosaic cucumovirus by plant growth-promoting rhizobacteria","volume":"45","author":"Zehnder","year":"2000","journal-title":"BioControl"},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"134","DOI":"10.1016\/j.jbiotec.2019.10.003","article-title":"A new strategy to control Cucumber mosaic virus using fabricated NiO-nanostructures","volume":"306","author":"Derbalah","year":"2019","journal-title":"J. Biotechnol."},{"key":"ref_82","doi-asserted-by":"crossref","unstructured":"Wang, X., Tang, C., Zhang, G., Li, Y., Wang, C., Liu, B., Qu, Z., Zhao, J., Han, Q., and Huang, L. (2009). cDNA-AFLP analysis reveals differential gene expression in compatible interaction of wheat challenged with Puccinia striiformis f. sp. tritici. Bmc Genom., 10.","DOI":"10.1186\/1471-2164-10-289"},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"142","DOI":"10.1111\/1574-6968.12461","article-title":"Analysis of the defence-related mechanism in cucumber seedlings in relation to root colonization by nonpathogenic Fusarium oxysporum CS-20","volume":"355","author":"Pu","year":"2014","journal-title":"Fems Microbiol. Lett."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"402","DOI":"10.1006\/meth.2001.1262","article-title":"Analysis of relative gene expression data using real-time quantitative PCR and the 2\u2212\u0394\u0394CT method","volume":"25","author":"Livak","year":"2001","journal-title":"Methods"},{"key":"ref_85","unstructured":"Vernon, L., and Seely, G. (1966). The Chlorophylls, Academic Press."},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"350","DOI":"10.1016\/0076-6879(87)48036-1","article-title":"Chlorophylls and carotenoids: Pigments of photosynthetic biomembranes","volume":"Volume 148","author":"Lichtenthaler","year":"1987","journal-title":"Methods in Enzymology"},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"189","DOI":"10.1016\/0003-9861(68)90654-1","article-title":"Photoperoxidation in isolated chloroplasts: I. Kinetics and stoichiometry of fatty acid peroxidation","volume":"125","author":"Heath","year":"1968","journal-title":"Arch. Biochem. Biophys."},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"753","DOI":"10.1016\/0031-9422(73)80672-7","article-title":"A rapid, sensitive and accurate determination of indolyl-3-acetic acid","volume":"12","author":"Knegt","year":"1973","journal-title":"Phytochemistry"},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"73","DOI":"10.1111\/j.1399-3054.1989.tb05980.x","article-title":"The effect of sucrose on the levels of abscisic acid, indoleacetic acid and zeatin\/zeatin riboside in wheat ears growing in liquid culture","volume":"77","author":"Lee","year":"1989","journal-title":"Physiol. Plant."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"205","DOI":"10.1007\/BF00018060","article-title":"Rapid determination of free proline for water-stress studies","volume":"39","author":"Bates","year":"1973","journal-title":"Plant Soil"},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"55","DOI":"10.1111\/j.1399-3054.1992.tb08764.x","article-title":"Water stress induced changes in concentrations of proline and total soluble sugars in nodulated alfalfa (Medicago sativd) plants","volume":"84","author":"Irigoyen","year":"1992","journal-title":"Physiol. Plant."},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"303","DOI":"10.1007\/BF02374789","article-title":"Rapid assay for determination of water soluble quaternary ammonium compounds","volume":"70","author":"Grieve","year":"1983","journal-title":"Plant Soil"},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"166","DOI":"10.1111\/j.1399-3054.1983.tb04162.x","article-title":"Implications of water stress-induced changes in the levels of endogenous ascorbic acid and hydrogen peroxide in Vigna seedlings","volume":"58","author":"Mukherjee","year":"1983","journal-title":"Physiol. Plant."},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"207","DOI":"10.1016\/0003-2697(80)90139-6","article-title":"Determination of glutathione and glutathione disulfide using glutathione reductase and 2-vinylpyridine","volume":"106","author":"Griffith","year":"1980","journal-title":"Anal. Biochem."},{"key":"ref_95","unstructured":"Galicia, L.N., Rosales, E., and A Palacios Rojas, N. (2009). Laboratory Protocols: Maize Nutrition Quality and Plant Tissue Analysis Laboratory, CIMMYT."},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"469","DOI":"10.1111\/j.1432-1033.1974.tb03714.x","article-title":"Involvement of the superoxide anion radical in the autoxidation of pyrogallol and a convenient assay for superoxide dismutase","volume":"47","author":"Marklund","year":"1974","journal-title":"Eur. J. Biochem."},{"key":"ref_97","unstructured":"Bergmeyer, H.U., and Gawehn, K. (1974). Methods of Enzymatic Analysis, Academic Press."},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"1225","DOI":"10.1104\/pp.94.3.1225","article-title":"Characteristics of a membrane-associated lipoxygenase in tomato fruit","volume":"94","author":"Todd","year":"1990","journal-title":"Plant Physiol."},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"357","DOI":"10.1007\/s001280000136","article-title":"Effects of rare earth metal ions and their EDTA complexes on antioxidant enzymes of fish liver","volume":"65","author":"Chen","year":"2000","journal-title":"Bull. Environ. Contam."},{"key":"ref_100","first-page":"867","article-title":"Hydrogen peroxide is scavenged by ascorbate-specific peroxidase in Spinach chloroplasts","volume":"22","author":"Nakano","year":"1981","journal-title":"Plant Cell Physiol."},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"2401","DOI":"10.1093\/jxb\/erf090","article-title":"Water stress-induced abscisic acid accumulation triggers the increased generation of reactive oxygen species and up-regulates the activities of antioxidant enzymes in maize leaves","volume":"53","author":"Jiang","year":"2002","journal-title":"J. Exp. Bot."},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"689","DOI":"10.1139\/m69-122","article-title":"The chitinase of Serratia marcescens","volume":"15","author":"Monreal","year":"1969","journal-title":"Can. J. Microbiol."},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"395","DOI":"10.1104\/pp.45.4.395","article-title":"Temporal and hormonal control of \u03b2-1,3-glucanase in Phaseolus vulgaris L","volume":"45","author":"Abeles","year":"1970","journal-title":"Plant Physiol."},{"key":"ref_104","unstructured":"Gomez, K.A., and Gomez, A.A. (1984). Statistical Procedures for Agricultural Research, John Wiley & Sons Inc."}],"container-title":["Molecules"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1420-3049\/25\/10\/2341\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T09:29:40Z","timestamp":1760174980000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1420-3049\/25\/10\/2341"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2020,5,17]]},"references-count":104,"journal-issue":{"issue":"10","published-online":{"date-parts":[[2020,5]]}},"alternative-id":["molecules25102341"],"URL":"https:\/\/doi.org\/10.3390\/molecules25102341","relation":{},"ISSN":["1420-3049"],"issn-type":[{"value":"1420-3049","type":"electronic"}],"subject":[],"published":{"date-parts":[[2020,5,17]]}}}