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Talent Training in Zhejiang University","award":["31800386"],"award-info":[{"award-number":["31800386"]}]},{"name":"Collaborative Innovation Center for Modern Crop Production","award":["2022C02034"],"award-info":[{"award-number":["2022C02034"]}]},{"name":"Collaborative Innovation Center for Modern Crop Production","award":["LY21C130006"],"award-info":[{"award-number":["LY21C130006"]}]},{"name":"Collaborative Innovation Center for Modern Crop Production","award":["32072127"],"award-info":[{"award-number":["32072127"]}]},{"name":"Collaborative Innovation Center for Modern Crop Production","award":["31800386"],"award-info":[{"award-number":["31800386"]}]}],"content-domain":{"domain":["www.mdpi.com"],"crossmark-restriction":true},"short-container-title":["Antioxidants"],"abstract":"<jats:p>Chromium (Cr) is an important environmental constraint effecting crop productivity. Spermine (SPM) is a polyamine compound regulating plant responses to abiotic stresses. However, SPM-mediated tolerance mechanisms against Cr stress are less commonly explored in plants. Thus, current research was conducted to explore the protective mechanisms of SPM (0.01 mM) against Cr (100 \u00b5M) toxicity in two rice cultivars, CY927 (sensitive) and YLY689 (tolerant) at the seedling stage. Our results revealed that, alone, Cr exposure significantly reduced seed germination, biomass and photosynthetic related parameters, caused nutrient and hormonal imbalance, desynchronized antioxidant enzymes, and triggered oxidative damage by over-accretion of reactive oxygen species (ROS), malondialdehyde (MDA) and electrolyte leakage in both rice varieties, with greater impairments in CY927 than YLY689. However, seed priming with SPM notably improved or reversed the above-mentioned parameters, especially in YLY689. Besides, SPM stimulated the stress-responsive genes of endogenous phytohormones, especially salicylic acid (SA), as confirmed by the pronounced transcript levels of SA-related genes (OsPR1, OsPR2 and OsNPR1). Our findings specified that SPM enhanced rice tolerance against Cr toxicity via decreasing accumulation of Cr and markers of oxidative damage (H2O2, O2\u2022\u2212 and MDA), improving antioxidant defense enzymes, photosynthetic apparatus, nutrients and phytohormone balance.<\/jats:p>","DOI":"10.3390\/antiox11091704","type":"journal-article","created":{"date-parts":[[2022,8,30]],"date-time":"2022-08-30T21:25:18Z","timestamp":1661894718000},"page":"1704","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":35,"title":["Seed Priming with Spermine Mitigates Chromium Stress in Rice by Modifying the Ion Homeostasis, Cellular Ultrastructure and Phytohormones Balance"],"prefix":"10.3390","volume":"11","author":[{"given":"Farwa","family":"Basit","sequence":"first","affiliation":[{"name":"Institute of Crop Science, Zhejiang Key Laboratory of Crop Germplasm, Zhejiang University, Hangzhou 310058, China"},{"name":"Hainan Research Institute, Zhejiang University, Sanya 572025, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Javaid","family":"Bhat","sequence":"additional","affiliation":[{"name":"International Genome Center, Jiangsu University, Zhenjiang 212013, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zaid","family":"Ulhassan","sequence":"additional","affiliation":[{"name":"Institute of Crop Science, Zhejiang Key Laboratory of Crop Germplasm, Zhejiang University, Hangzhou 310058, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Muhammad","family":"Noman","sequence":"additional","affiliation":[{"name":"Institute of Crop Science, Zhejiang Key Laboratory of Crop Germplasm, Zhejiang University, Hangzhou 310058, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Biying","family":"Zhao","sequence":"additional","affiliation":[{"name":"International Genome Center, Jiangsu University, Zhenjiang 212013, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Weijun","family":"Zhou","sequence":"additional","affiliation":[{"name":"Hainan Research Institute, Zhejiang University, Sanya 572025, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3145-2849","authenticated-orcid":false,"given":"Prashant","family":"Kaushik","sequence":"additional","affiliation":[{"name":"Independent Researcher, 46022 Valencia, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ajaz","family":"Ahmad","sequence":"additional","affiliation":[{"name":"Department of Clinical Pharmacy, College of Pharmacy, King Saud University, Riyadh 11451, Saudi Arabia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2734-4180","authenticated-orcid":false,"given":"Parvaiz","family":"Ahmad","sequence":"additional","affiliation":[{"name":"Department of Botany, GDC, Pulwama 192301, Jammu and Kashmir 192301, India"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yajing","family":"Guan","sequence":"additional","affiliation":[{"name":"Institute of Crop Science, Zhejiang Key Laboratory of Crop Germplasm, Zhejiang University, Hangzhou 310058, China"},{"name":"Hainan Research Institute, Zhejiang University, Sanya 572025, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2022,8,30]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","unstructured":"Basit, F., Chen, M., Ahmed, T., Shahid, M., Noman, M., Liu, J., An, J., Hashem, A., Fahad Al-Arjani, A.-B., and Alqarawi, A.A. (2021). Seed priming with brassinosteroids alleviates chromium stress in rice cultivars via improving ROS metabolism and antioxidant defense response at biochemical and molecular levels. Antioxidants, 10.","DOI":"10.3390\/antiox10071089"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"8","DOI":"10.1016\/j.wace.2014.06.003","article-title":"Droughts in Asian least developed countries: Vulnerability and sustainability","volume":"7","author":"Miyan","year":"2015","journal-title":"Weather Clim. Extrem."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"190","DOI":"10.1007\/s11270-017-3356-y","article-title":"Chromium in agricultural soils and crops: A review","volume":"228","author":"Ertani","year":"2017","journal-title":"Water Air Soil Pollut."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"829","DOI":"10.1016\/j.jhazmat.2008.07.034","article-title":"Toxicity assessment of soil amended with tannery sludge, trivalent chromium and hexavalent chromium, using wheat, oat and sorghum plants","volume":"163","year":"2009","journal-title":"J. Hazard. Mater."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"8814","DOI":"10.1007\/s11356-017-8603-7","article-title":"Promotive role of 5-aminolevulinic acid on chromium-induced morphological, photosynthetic, and oxidative changes in cauliflower (Brassica oleracea botrytis L.)","volume":"24","author":"Ahmad","year":"2017","journal-title":"Environ. Sci. Pollut. Res."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"44768","DOI":"10.1007\/s11356-021-15087-8","article-title":"Brassinosteroids as a multidimensional regulator of plant physiological and molecular responses under various environmental stresses","volume":"28","author":"Basit","year":"2021","journal-title":"Environ. Sci. Pollut. Res."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"142","DOI":"10.1016\/j.plaphy.2019.10.035","article-title":"Selenium mitigates the chromium toxicity in Brassicca napus L. by ameliorating nutrients uptake, amino acids metabolism and antioxidant defense system","volume":"145","author":"Ulhassan","year":"2019","journal-title":"Plant Physiol. Biochem."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"128547","DOI":"10.1016\/j.chemosphere.2020.128547","article-title":"Hexavalent chromium induced heart dysfunction via Sesn2-mediated impairment of mitochondrial function and energy supply","volume":"264","author":"Yang","year":"2021","journal-title":"Chemosphere"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"103789","DOI":"10.1016\/j.envexpbot.2019.06.002","article-title":"Regulation of chromium toxicity tolerance in tomato and brinjal by calcium and sulfur through nitric oxide: Involvement of enzymes of sulfur assimilation and the ascorbate-glutathione cycle","volume":"166","author":"Singh","year":"2019","journal-title":"Environ. Exp. Bot."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"180","DOI":"10.1016\/j.envexpbot.2018.11.009","article-title":"Selenium modulates dynamics of antioxidative defence expression, photosynthetic attributes and secondary metabolites to mitigate chromium toxicity in Brassica juncea L. plants","volume":"161","author":"Handa","year":"2019","journal-title":"Environ. Exp. Bot."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"1001","DOI":"10.1016\/j.plaphy.2021.07.013","article-title":"Salicylic acid underpins silicon in ameliorating chromium toxicity in rice by modulating antioxidant defense, ion homeostasis and cellular ultrastructure","volume":"166","author":"Yang","year":"2021","journal-title":"Plant Physiol. Biochem."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"329","DOI":"10.1016\/j.chemosphere.2019.03.028","article-title":"Dual behavior of selenium: Insights into physio-biochemical, anatomical and molecular analyses of four Brassica napus cultivars","volume":"225","author":"Ulhassan","year":"2019","journal-title":"Chemosphere"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"111758","DOI":"10.1016\/j.ecoenv.2020.111758","article-title":"Jasmonic acid-mediated enhanced regulation of oxidative, glyoxalase defense system and reduced chromium uptake contributes to alleviation of chromium (VI) toxicity in choysum (Brassica parachinensis L.)","volume":"208","author":"Kamran","year":"2021","journal-title":"Ecotoxicol. Environ. Saf."},{"key":"ref_14","doi-asserted-by":"crossref","unstructured":"Takahashi, T. (2022, July 08). Plant Polyamines. Available online: https:\/\/www.mdpi.com\/2223-7747\/9\/4\/511.","DOI":"10.3390\/plants9040511"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"35","DOI":"10.1016\/j.plaphy.2018.05.017","article-title":"Exogenously applied spermidine alleviates photosynthetic inhibition under drought stress in maize (Zea mays L.) seedlings associated with changes in endogenous polyamines and phytohormones","volume":"129","author":"Li","year":"2018","journal-title":"Plant Physiol. Biochem."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"1815","DOI":"10.1007\/s12298-020-00869-0","article-title":"The protective effects of polyamines on salinity stress tolerance in foxtail millet (Setaria italica L.), an important C4 model crop","volume":"26","author":"Rathinapriya","year":"2020","journal-title":"Physiol. Mol. Biol. Plants"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"1807722","DOI":"10.1080\/15592324.2020.1807722","article-title":"Polyamine biosynthetic pathways and their relation with the cold tolerance of maize (Zea mays L.) seedlings","volume":"15","author":"Gao","year":"2020","journal-title":"Plant Signal. Behav."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"7467","DOI":"10.1038\/s41598-020-64468-5","article-title":"The alleviating effect of exogenous polyamines on heat stress susceptibility of different heat resistant wheat (Triticum aestivum L.) varieties","volume":"10","author":"Jing","year":"2020","journal-title":"Sci. Rep."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"125308","DOI":"10.1016\/j.chemosphere.2019.125308","article-title":"2-Hydroxymelatonin mitigates cadmium stress in cucumis sativus seedlings: Modulation of antioxidant enzymes and polyamines","volume":"243","author":"Shah","year":"2020","journal-title":"Chemosphere"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"21206","DOI":"10.1007\/s11356-016-7295-8","article-title":"Physiological and biochemical mechanisms of spermine-induced cadmium stress tolerance in mung bean (Vigna radiata L.) seedlings","volume":"23","author":"Nahar","year":"2016","journal-title":"Environ. Sci. Pollut. Res."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"22338","DOI":"10.1007\/s11356-019-05555-7","article-title":"Polyamines modulate growth, antioxidant activity, and genomic DNA in heavy metal\u2013stressed wheat plant","volume":"26","author":"Taie","year":"2019","journal-title":"Environ. Sci. Pollut. Res."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"285","DOI":"10.1016\/j.plaphy.2020.12.022","article-title":"Combined ability of salicylic acid and spermidine to mitigate the individual and interactive effects of drought and chromium stress in maize (Zea mays L.)","volume":"159","author":"Naz","year":"2021","journal-title":"Plant Physiol. Biochem."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"185","DOI":"10.1007\/s10534-006-9026-y","article-title":"Polyamines and heavy metal stress: The antioxidant behavior of spermine in cadmium-and copper-treated wheat leaves","volume":"20","author":"Groppa","year":"2007","journal-title":"Biometals"},{"key":"ref_24","doi-asserted-by":"crossref","unstructured":"Paul, S., Banerjee, A., and Roychoudhury, A. (2018). Role of polyamines in mediating antioxidant defense and epigenetic regulation in plants exposed to heavy metal toxicity. Plants under Metal and Metalloid Stress, Springer.","DOI":"10.1007\/978-981-13-2242-6_8"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"272","DOI":"10.1016\/j.jhazmat.2017.07.016","article-title":"Polyamines may influence phytochelatin synthesis during Cd stress in rice","volume":"340","author":"Szalai","year":"2017","journal-title":"J. Hazard. Mater."},{"key":"ref_26","first-page":"41","article-title":"Identification of chilling-tolerance in maize inbred lines at germination and seedling growth stages","volume":"32","author":"Zheng","year":"2006","journal-title":"J. Zhejiang Univ. (Agric. Life Sci.)"},{"key":"ref_27","first-page":"591","article-title":"Determinations of total carotenoids and chlorophylls a and b of leaf extracts in different solvents","volume":"11","author":"Lichtenthaler","year":"1983","journal-title":"Analysis"},{"key":"ref_28","unstructured":"Antonio Pasqualini, A., Tadeu Paulino, V., Maria Duarte, K., Hansen Rosolen, F., and Menegante Neri, A. (2013). Chlorophyll evaluation methods in Brachiaria brizantha brs Piat\u00e3 added with poultry litter compost. B. Ind\u00fastr. Anim., 70."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"41","DOI":"10.1023\/A:1006004921265","article-title":"Uniconazole-induced alleviation of freezing injury in relation to changes in hormonal balance, enzyme activities and lipid peroxidation in winter rape","volume":"26","author":"Zhou","year":"1998","journal-title":"Plant Growth Regul."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"4151","DOI":"10.1128\/AEM.70.7.4151-4157.2004","article-title":"Effect of substratum surface chemistry and surface energy on attachment of marine bacteria and algal spores","volume":"70","author":"Ista","year":"2004","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"248","DOI":"10.1016\/0003-2697(76)90527-3","article-title":"A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of protein-dye binding","volume":"72","author":"Bradford","year":"1976","journal-title":"Anal. Biochem."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"167","DOI":"10.1038\/168167a0","article-title":"A colorimetric method for the determination of sugars","volume":"168","author":"Dubois","year":"1951","journal-title":"Nature"},{"key":"ref_33","doi-asserted-by":"crossref","unstructured":"Kim, Y.-H., Khan, A.L., Kim, D.-H., Lee, S.-Y., Kim, K.-M., Waqas, M., Jung, H.-Y., Shin, J.-H., Kim, J.-G., and Lee, I.-J. (2014). Silicon mitigates heavy metal stress by regulating P-type heavy metal ATPases, Oryza sativa low silicon genes, and endogenous phytohormones. BMC Plant Biol., 14.","DOI":"10.1186\/1471-2229-14-13"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"21","DOI":"10.1023\/A:1006299414481","article-title":"GC-MS-SIM analysis of abscisic acid and indole-3-acetic acid in shoot bark of apple rootstocks","volume":"28","author":"Kamboj","year":"1999","journal-title":"Plant Growth Regul."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"430","DOI":"10.1007\/s004250050210","article-title":"Herbivory and caterpillar regurgitants amplify the wound-induced increases in jasmonic acid but not nicotine in Nicotiana sylvestris","volume":"203","author":"McCloud","year":"1997","journal-title":"Planta"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"372","DOI":"10.1007\/BF00201815","article-title":"Ultraviolet light and ozone stimulate accumulation of salicylic acid, pathogenesis-related proteins and virus resistance in tobacco","volume":"193","author":"Yalpani","year":"1994","journal-title":"Planta"},{"key":"ref_37","unstructured":"Fang, Y., Umasankar, Y., and Ramasamy, R.P. (June, January May). Plant volatile sensor: Enzymatic transducer for selective and sensitive determination of methyl salicyalte. Proceedings of the ECS Meeting Abstracts, Vancouver, BC, Canada."},{"key":"ref_38","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_39","doi-asserted-by":"crossref","first-page":"185","DOI":"10.1016\/j.jplph.2012.09.017","article-title":"Accumulation of peroxidase-related reactive oxygen species in trichoblasts correlates with root hair initiation in barley","volume":"170","author":"Kwasniewski","year":"2013","journal-title":"J. Plant Physiol."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"1265","DOI":"10.1093\/pcp\/pce162","article-title":"Effect of abscisic acid on active oxygen species, antioxidative defence system and oxidative damage in leaves of maize seedlings","volume":"42","author":"Jiang","year":"2001","journal-title":"Plant Cell Physiol."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"309","DOI":"10.1104\/pp.59.2.309","article-title":"Superoxide dismutases: I. Occurrence in higher plants","volume":"59","author":"Giannopolitis","year":"1977","journal-title":"Plant Physiol."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"121","DOI":"10.1016\/S0076-6879(84)05016-3","article-title":"Catalase in vitro","volume":"105","author":"Aebi","year":"1984","journal-title":"Methods Enzymol."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"764","DOI":"10.1016\/S0076-6879(55)02300-8","article-title":"Assay of catalases and peroxidase","volume":"2","author":"Change","year":"1955","journal-title":"Methods Enzym."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"867","DOI":"10.1093\/oxfordjournals.pcp.a076232","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_45","doi-asserted-by":"crossref","first-page":"3129","DOI":"10.4236\/ajps.2014.521329","article-title":"Rapid and reliable method of high-quality RNA extraction from diverse plants","volume":"5","author":"Sah","year":"2014","journal-title":"Am. J. Plant Sci."},{"key":"ref_46","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_47","doi-asserted-by":"crossref","first-page":"58","DOI":"10.1016\/j.envexpbot.2017.02.007","article-title":"Seed polyamines metabolism induced by seed priming with spermidine and 5-aminolevulinic acid for chilling tolerance improvement in rice (Oryza sativa L.) seedlings","volume":"137","author":"Sheteiwy","year":"2017","journal-title":"Environ. Exp. Bot."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"1413","DOI":"10.1007\/s00344-018-9831-x","article-title":"Role of zinc\u2013lysine on growth and chromium uptake in rice plants under Cr stress","volume":"37","author":"Hussain","year":"2018","journal-title":"J. Plant Growth Regul."},{"key":"ref_49","unstructured":"Bewley, J.D., and Black, M. (2012). Physiology and Biochemistry of Seeds in Relation to Germination: Volume 2: Viability, Dormancy, and Environmental Control, Springer Science & Business Media."},{"key":"ref_50","first-page":"702","article-title":"Enhancement effect of biofertilizers on germination percentage and plant height in maize grown under chromium toxic soil","volume":"9","author":"Devi","year":"2020","journal-title":"J. Pharmacogn. Phytochem."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"116","DOI":"10.3389\/fpls.2016.00116","article-title":"Physiological and biochemical mechanisms of seed priming-induced chilling tolerance in rice cultivars","volume":"7","author":"Hussain","year":"2016","journal-title":"Front. Plant Sci."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"159","DOI":"10.1016\/j.colsurfb.2007.11.016","article-title":"Chromium stress induced alterations in biochemical and enzyme metabolism in aquatic and terrestrial plants","volume":"63","author":"Ganesh","year":"2008","journal-title":"Colloids Surf. B BioInterfaces"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"932","DOI":"10.1016\/j.bbapap.2016.02.020","article-title":"Toxicity of heavy metals and metal-containing nanoparticles on plants","volume":"1864","author":"Mustafa","year":"2016","journal-title":"Biochim. Biophys. Acta (BBA)-Proteins Proteom."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"112401","DOI":"10.1016\/j.ecoenv.2021.112401","article-title":"Zinc oxide nanoparticles alleviates the adverse effects of cadmium stress on Oryza sativa via modulation of the photosynthesis and antioxidant defense system","volume":"220","author":"Faizan","year":"2021","journal-title":"Ecotoxicol. Environ. Saf."},{"key":"ref_55","doi-asserted-by":"crossref","unstructured":"Mostofa, M.G., Rahman, M., Ansary, M., Uddin, M., Fujita, M., and Tran, L.-S.P. (2019). Interactive effects of salicylic acid and nitric oxide in enhancing rice tolerance to cadmium stress. Int. J. Mol. Sci., 20.","DOI":"10.3390\/ijms20225798"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"11","DOI":"10.1007\/s00425-019-03304-1","article-title":"Chromium effects on photosynthetic electron transport in pea (Pisum sativum L.)","volume":"251","author":"Todorenko","year":"2020","journal-title":"Planta"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"972","DOI":"10.3389\/fpls.2019.00972","article-title":"Spermine confers stress resilience by modulating abscisic acid biosynthesis and stress responses in Arabidopsis plants","volume":"10","author":"Marco","year":"2019","journal-title":"Front. Plant Sci."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"15159","DOI":"10.1007\/s11356-018-1742-7","article-title":"Interaction of 24-epibrassinolide and salicylic acid regulates pigment contents, antioxidative defense responses, and gene expression in Brassica juncea L. seedlings under Pb stress","volume":"25","author":"Kohli","year":"2018","journal-title":"Environ. Sci. Pollut. Res."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1007\/s11738-021-03276-4","article-title":"Silicon attenuates the negative effects of chromium stress in tomato plants by modifying antioxidant enzyme activities, ascorbate\u2013glutathione cycle and glyoxalase system","volume":"43","author":"Alam","year":"2021","journal-title":"Acta Physiol. Plant."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"245","DOI":"10.1016\/j.ecoenv.2015.12.026","article-title":"Polyamine and nitric oxide crosstalk: Antagonistic effects on cadmium toxicity in mung bean plants through upregulating the metal detoxification, antioxidant defense and methylglyoxal detoxification systems","volume":"126","author":"Nahar","year":"2016","journal-title":"Ecotoxicol. Environ. Saf."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"905","DOI":"10.1007\/s12298-019-00672-6","article-title":"24-epibrassinolide and spermidine alleviate Mn stress via the modulation of root morphology, stomatal behavior, photosynthetic attributes and antioxidant defense in Brassica juncea","volume":"25","author":"Hussain","year":"2019","journal-title":"Physiol. Mol. Biol. Plants"},{"key":"ref_62","doi-asserted-by":"crossref","unstructured":"Choudhary, S.P., Kanwar, M., Bhardwaj, R., Yu, J.-Q., and Tran, L.-S.P. (2012). Chromium stress mitigation by polyamine-brassinosteroid application involves phytohormonal and physiological strategies in Raphanus sativus L. PLoS ONE, 7.","DOI":"10.1371\/journal.pone.0033210"},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"988","DOI":"10.2135\/cropsci2003.0678","article-title":"Effects of salicylic acid on heat tolerance associated with antioxidant metabolism in Kentucky bluegrass","volume":"45","author":"He","year":"2005","journal-title":"Crop Sci."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"317","DOI":"10.1016\/j.envexpbot.2007.11.003","article-title":"Effects of exogenous salicylic acid on manganese toxicity, element contents and antioxidative system in cucumber","volume":"63","author":"Shi","year":"2008","journal-title":"Environ. Exp. Bot."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"189","DOI":"10.1007\/s10725-020-00571-x","article-title":"Crosstalk amongst phytohormones from planta and PGPR under biotic and abiotic stresses","volume":"90","author":"Khan","year":"2020","journal-title":"Plant Growth Regul."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"53","DOI":"10.1016\/j.nbt.2018.07.003","article-title":"Phytohormones and polyamines regulate plant stress responses by altering GABA pathway","volume":"48","author":"Ugena","year":"2019","journal-title":"New Biotechnol."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"101","DOI":"10.1007\/s10311-016-0584-0","article-title":"Salicylic acid to decrease plant stress","volume":"15","author":"Wani","year":"2017","journal-title":"Environ. Chem. Lett."},{"key":"ref_68","doi-asserted-by":"crossref","unstructured":"Sharma, A., Sidhu, G.P.S., Araniti, F., Bali, A.S., Shahzad, B., Tripathi, D.K., Brestic, M., Skalicky, M., and Landi, M. (2020). The role of salicylic acid in plants exposed to heavy metals. Molecules, 25.","DOI":"10.3390\/molecules25030540"},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"967","DOI":"10.1007\/s10529-008-9639-z","article-title":"Fatty acid unsaturation, mobilization, and regulation in the response of plants to stress","volume":"30","author":"Upchurch","year":"2008","journal-title":"Biotechnol. Lett."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"12839","DOI":"10.1038\/s41598-018-31297-6","article-title":"Interaction of polyamines, abscisic acid and proline under osmotic stress in the leaves of wheat plants","volume":"8","author":"Tajti","year":"2018","journal-title":"Sci. Rep."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"1077","DOI":"10.4236\/ajps.2012.38129","article-title":"Effects of cadmium on growth, photosynthetic pigments, photosynthetic performance, biochemical parameters and structure of chloroplasts in the agarophyte Gracilaria domingensis (Rhodophyta, Gracilariales)","volume":"3","author":"Eder","year":"2012","journal-title":"Am. J. Plant Sci."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"209","DOI":"10.1111\/aab.12660","article-title":"Polyamines as key regulatory players in plants under metal stress\u2014A way for an enhanced tolerance","volume":"178","author":"Spormann","year":"2021","journal-title":"Ann. Appl. Biol."}],"updated-by":[{"DOI":"10.3390\/antiox14030312","type":"correction","label":"Correction","source":"publisher","updated":{"date-parts":[[2022,8,30]],"date-time":"2022-08-30T00:00:00Z","timestamp":1661817600000}}],"container-title":["Antioxidants"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2076-3921\/11\/9\/1704\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,8,3]],"date-time":"2025-08-03T14:18:09Z","timestamp":1754230689000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2076-3921\/11\/9\/1704"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2022,8,30]]},"references-count":72,"journal-issue":{"issue":"9","published-online":{"date-parts":[[2022,9]]}},"alternative-id":["antiox11091704"],"URL":"https:\/\/doi.org\/10.3390\/antiox11091704","relation":{"correction":[{"id-type":"doi","id":"10.3390\/antiox14030312","asserted-by":"object"}]},"ISSN":["2076-3921"],"issn-type":[{"value":"2076-3921","type":"electronic"}],"subject":[],"published":{"date-parts":[[2022,8,30]]}}}