journal article·2026-01-23Open access

Mechanistic insights into the role of chitosan nanoparticles in enhancing plant drought tolerance

Nadiyah M. Alabdallah, Salman Latif

Nanotechnology Reviews· Volume 15· Issue 1

Abstract

Abstract Enhancing crop drought resilience is a critical goal for sustainable agriculture under increasing climate variability. Chitosan nanoparticles (ChNPs) have emerged as effective biostimulants; however, their drought-mitigating action remains insufficiently defined at the mechanistic level. Here, we synthesize current evidence to propose a causal mechanistic framework linking ChNP input characteristics (particle size∼20–200 nm; application doses typically 30–400 ppm) to plant physiological and molecular responses under drought stress. Following root or foliar uptake, ChNPs are translocated through apoplastic and symplastic pathways, enabling interaction with photosynthetic and stress-responsive tissues. This uptake initiates signaling cascades involving controlled reactive oxygen species (ROS) modulation and abscisic acid (ABA)-related pathways. Evidence from multiple controlled studies, primarily in leaf and root tissues under moderate drought conditions, indicates that ChNP application is associated with the upregulation of key drought-responsive genes, including LeNCED1 (ABA biosynthesis) and SlAREB1 (an ABA-dependent transcription factor), typically showing modest but consistent induction compared with untreated drought-stressed plants. These molecular responses translate into improved photosynthetic efficiency, enhanced osmolyte accumulation, and stronger antioxidant defenses, collectively reducing oxidative damage and improving drought tolerance. While these findings highlight the mechanistic potential of ChNPs, careful dose optimization and environmental assessment remain essential for sustainable agricultural application.
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Research context

OpenAlex topics: Nanoparticles: synthesis and applications · Plant Growth Enhancement Techniques · Plant Stress Responses and Tolerance

SciVX discovery tags are terminology-based. OpenAlex topics are source-provided classifications.

Affiliations

Department of Biology , College of Science, Imam Abdulrahman Bin Faisal University, Dammam , P.O. Box 1982, 31441 , Saudi Arabia

Basic and Applied Scientific Research Centre, Imam Abdulrahman Bin Faisal University , P.O. Box 1982, 31441 , Dammam , Saudi Arabia

Department of Chemistry , College of Science and Humanities in Al-Kharj, Prince Sattam bin Abdulaziz University , Al-Kharj , 11942 , Saudi Arabia

References (80)

Show source-provided references
1. Yuan, X, Li, S, Chen, J, Yu, H, Yang, T, Wang, C, et al.. Impacts of global climate change on agricultural production: a comprehensive review. Agronomy 2024;14:1360. https://doi.org/10.3390/agronomy14071360.10.3390/agronomy14071360 ↗
2. Khan, AA, Wang, YF, Akbar, R, Alhoqail, WA. Mechanistic insights and future perspectives of drought stress management in staple crops. Front Plant Sci 2025;16:1547452. https://doi.org/10.3389/fpls.2025.1547452.10.3389/fpls.2025.1547452 ↗
3. Chiappini, S, Cimino, O, Cardillo, C. Climate crises and agricultural drought: evolutions in water scarcity context at the farm level. Earth 2025;6:56. https://doi.org/10.3390/earth6020056.10.3390/earth6020056 ↗
4. Gong, L, Hazzazi, Y, Alfaifi, T, Alabdallah, NM, Alnusaire, TS, Altihani, FA, et al.. Root-to-shoot hormonal and hydraulic signals in stomatal regulation during drought. J Plant Growth Regul 2025;44:1–14. https://doi.org/10.1007/s00344-025-11733-4.10.1007/s00344-025-11733-4 ↗
5. Rao, MJ, Duan, M, Zhou, C, Jiao, J, Cheng, P, Yang, L, et al.. Antioxidant defense system in plants: reactive oxygen species production, signaling, and scavenging during abiotic stress-induced oxidative damage. Horticulturae 2025;11:477. https://doi.org/10.3390/horticulturae11050477.10.3390/horticulturae11050477 ↗
6. Renzetti, M, Funck, D, Trovato, M. Proline and ROS: a unified mechanism in plant development and stress response? Plants 2024;14:2. https://doi.org/10.3390/plants14010002.10.3390/plants14010002 ↗
7. Karamian, R, Ghasemlou, F, Amiri, H. Physiological evaluation of drought stress tolerance and recovery in Verbascum sinuatum plants treated with methyl jasmonate, salicylic acid and titanium dioxide nanoparticles. Plant Biosyst 2020;154:277–87. https://doi.org/10.1080/11263504.2019.1591535.10.1080/11263504.2019.1591535 ↗
8. Nayan, R, Rawat, M, Negi, B, Pande, A, Arora, S. Zinc sulfide nanoparticle mediated alterations in growth and antioxidant status of Brassica juncea. Biologia 2016;71:896–902. https://doi.org/10.1515/biolog-2016-0107.10.1515/biolog-2016-0107 ↗
9. Faqir, Y, Ma, J, Chai, Y. Chitosan in modern agriculture production. Plant Soil Environ 2021;67:12–699. https://doi.org/10.17221/332/2021-pse.10.17221/332/2021-PSE ↗
10. Kumaraswamy, R, Saharan, V, Kumari, S, Choudhary, RC, Pal, A, Sharma, SS, et al.. Chitosan-silicon nanofertilizer to enhance plant growth and yield in maize (Zea mays L.). Plant Physiol Biochem 2021;159:53–66. https://doi.org/10.1016/j.plaphy.2020.11.054.10.1016/j.plaphy.2020.11.054 ↗
11. Behboudi, F, Tahmasebi-Sarvestani, Z, Kassaee, MZ, Modarres-Sanavy, SAM, Sorooshzadeh, A, Mokhtassi-Bidgoli, A, et al.. Evaluation of chitosan nanoparticles effects with two application methods on wheat under drought stress. J Plant Nutr 2019;42:1439–51. https://doi.org/10.1080/01904167.2019.1617308.10.1080/01904167.2019.1617308 ↗
12. Ali, EF, El-Shehawi, AM, Ibrahim, OHM, Abdul-Hafeez, EY, Moussa, MM, Hassan, FAS, et al.. A vital role of chitosan nanoparticles in improvisation of drought stress tolerance in Catharanthus roseus (L.) through biochemical and gene expression modulation. Plant Physiol Biochem 2021;161:166–75. https://doi.org/10.1016/j.plaphy.2021.02.008.10.1016/j.plaphy.2021.02.008 ↗
13. Dowom, SA, Karimian, Z, Mostafaei Dehnavi, M, Samiei, L. Chitosan nanoparticles improve physiological and biochemical responses of Salvia abrotanoides (Kar.) under drought stress. BMC Plant Biol 2022;22:364. https://doi.org/10.1186/s12870-022-03689-4.10.1186/s12870-022-03689-4 ↗
14. Mohamed, NG, Abdel-Hakeem, MA. Chitosan nanoparticles enhance drought tolerance in tomatoes (Solanum lycopersicum L.) via gene expression modulation. Plant Gene 2023;34:100406. https://doi.org/10.1016/j.plgene.2023.100406.10.1016/j.plgene.2023.100406 ↗
15. Giglou, MT, Giglou, RH, Esmaeilpour, B, Azarmi, R, Padash, A, Falakian, M, et al.. A new method in mitigation of drought stress by chitosan-coated iron oxide nanoparticles and growth stimulant in peppermint. Ind Crops Prod 2022;187:115286. https://doi.org/10.1016/j.indcrop.2022.115286.10.1016/j.indcrop.2022.115286 ↗
16. Sivanesan, I, Gopal, J, Muthu, M, Shin, J, Mari, S, Oh, J, et al.. Green synthesized chitosan/chitosan nanoforms/nanocomposites for drug delivery applications. Polymers 2021;13:2256. https://doi.org/10.3390/polym13142256.10.3390/polym13142256 ↗
17. Al-Nemrawi, NK, Okour, AR, Dave, RH. Surface modification of PLGA nanoparticles using chitosan: effect of molecular weight, concentration, and degree of deacetylation. Adv Polym Tech 2018;37:3066–75. https://doi.org/10.1002/adv.22077.10.1002/adv.22077 ↗
18. Luangtana-anan, M, Opanasopit, P, Ngawhirunpat, T, Nunthanid, J, Sriamornsak, P, Limmatvapirat, S, et al.. Effect of chitosan salts and molecular weight on a nanoparticulate carrier for therapeutic protein. Pharm Dev Technol 2005;10:189–96. https://doi.org/10.1081/pdt-54388.10.1081/PDT-54388 ↗
19. Calvo, P, Remunan-Lopez, C, Vila-Jato, JL, Alonso, MJ. Novel hydrophilic chitosan-polyethylene oxide nanoparticles as protein carriers. J Appl Polym Sci 1997;63:125–32. https://doi.org/10.1002/(sici)1097-4628(19970103)63:1<125::aid-app13>3.0.co;2-4.10.1002/(SICI)1097-4628(19970103)63:1<125::AID-APP13>3.0.CO;2-4 ↗
20. Algharib, SA, Dawood, A, Zhou, K, Chen, D, Li, C, Meng, K, et al.. Preparation of chitosan nanoparticles by ionotropic gelation technique: effects of formulation parameters and in vitro characterization. J Mol Struct 2022;1252:132129. https://doi.org/10.1016/j.molstruc.2021.132129.10.1016/j.molstruc.2021.132129 ↗
21. Alehosseini, E, Shahiri Tabarestani, H, Kharazmi, MS, Jafari, SM. Physicochemical, thermal, and morphological properties of chitosan nanoparticles produced by ionic gelation. Foods 2022;11:3841. https://doi.org/10.3390/foods11233841.10.3390/foods11233841 ↗
22. Kafshgari, MH, Khorram, M, Mansouri, M, Samimi, A, Osfouri, S. Preparation of alginate and chitosan nanoparticles using a new reverse micellar system. Iran Polym J 2012;21:99–107. https://doi.org/10.1007/s13726-011-0010-1.10.1007/s13726-011-0010-1 ↗
23. Taokaew, S, Ofuchi, M, Kobayashi, T. Size distribution and characteristics of chitin microgels prepared via emulsified reverse-micelles. Materials 2019;12:1160. https://doi.org/10.3390/ma12071160.10.3390/ma12071160 ↗
24. Liu, G, Shao, L, Ge, F, Chen, J. Preparation of ultrafine chitosan particles by reverse microemulsion. China Particuology 2007;5:384–90. https://doi.org/10.1016/j.cpart.2007.08.002.10.1016/j.cpart.2007.08.002 ↗
25. Kotsaeng, N, Karnchanajindanun, J, Baimark, Y. Chitosan microparticles prepared by the simple emulsification-diffusion method. Part Sci Technol 2010;28:369–78. https://doi.org/10.1080/02726351003702574.10.1080/02726351003702574 ↗
26. Zhang, K, Xu, Y, Lu, L, Shi, C, Huang, Y, Mao, Z, et al.. Hydrodynamic cavitation: a feasible approach to intensify the emulsion cross-linking process for chitosan nanoparticle synthesis. Ultrason Sonochem 2021;74:105551. https://doi.org/10.1016/j.ultsonch.2021.105551.10.1016/j.ultsonch.2021.105551 ↗
27. Lee, M, Cho, YW, Park, JH, Chung, H, Jeong, SY, Choi, K, et al.. Size control of self-assembled nanoparticles by an emulsion/solvent evaporation method. Colloid Polym Sci 2006;284:506–12. https://doi.org/10.1007/s00396-005-1413-3.10.1007/s00396-005-1413-3 ↗
28. Estevinho, BN, Rocha, F, Santos, L, Alves, A. Microencapsulation with chitosan by spray drying for industry applications: a review. Trends Food Sci Technol 2013;31:138–55. https://doi.org/10.1016/j.tifs.2013.04.001.10.1016/j.tifs.2013.04.001 ↗
29. He, P, Davis, SS, Illum, L. Chitosan microspheres prepared by spray drying. Int J Pharm 1999;187:53–65. https://doi.org/10.1016/s0378-5173(99)00125-8.10.1016/S0378-5173(99)00125-8 ↗
30. Kasapoğlu, KN, Gültekin-Özgüven, M, Kruger, J, Frank, J, Bayramoğlu, P, Barla-Demirkoz, A, et al.. Effect of spray drying on physicochemical stability and antioxidant capacity of Rosa pimpinellifolia fruit extract-loaded liposomes conjugated with chitosan or whey protein during in vitro digestion. Food Bioprocess Technol 2024;17:3162–76. https://doi.org/10.1007/s11947-024-03317-z.10.1007/s11947-024-03317-z ↗
31. Atma, Y, Sadeghpour, A, Murray, BS, Goycoolea, FM. Chitosan-alginate polyelectrolyte complexes for encapsulation of low molecular weight fish bioactive peptides. Food Hydrocoll 2025;160:110789. https://doi.org/10.1016/j.foodhyd.2024.110789.10.1016/j.foodhyd.2024.110789 ↗
32. Raja, MA, Arif, M, Feng, C, Zeenat, S, Liu, CG. Synthesis and evaluation of pH-sensitive, self-assembled chitosan-based nanoparticles as efficient doxorubicin carriers. J Biomater Appl 2017;31:1182–95. https://doi.org/10.1177/0885328216681184.10.1177/0885328216681184 ↗
33. Wang, XY, Wang, J, Rousseau, D, Tang, CH. Chitosan-stabilized emulsion gels via pH-induced droplet flocculation. Food Hydrocoll 2020;105:105811. https://doi.org/10.1016/j.foodhyd.2020.105811.10.1016/j.foodhyd.2020.105811 ↗
34. Gadkari, RR, Suwalka, S, Yogi, MR, Ali, W, Das, A, Alagirusamy, R, et al.. Green synthesis of chitosan–cinnamaldehyde cross-linked nanoparticles: characterization and antibacterial activity. Carbohydr Polym 2019;226:115298. https://doi.org/10.1016/j.carbpol.2019.115298.10.1016/j.carbpol.2019.115298 ↗
35. Abdallah, Y, Liu, MJ, Ogunyemi, SO, Ahmed, T, Fouad, H, Abdelazez, A, et al.. Bioinspired green synthesis of chitosan and zinc oxide nanoparticles with strong antibacterial activity against rice pathogen Xanthomonas oryzae pv. oryzae. Molecules 2020;25:4795. https://doi.org/10.3390/molecules25204795.10.3390/molecules25204795 ↗
36. Qian, L, Zhang, HF. Green synthesis of chitosan-based nanofibers and their applications. Green Chem 2010;12:1207–14. https://doi.org/10.1039/b927125b.10.1039/b927125b ↗
37. Gopal, J, Muthu, M, Dhakshanamurthy, T, Kim, KJ, Hasan, N, Kwon, SJ, et al.. Sustainable ecofriendly phytoextract mediated one-pot green recovery of chitosan. Sci Rep 2019;9:13832. https://doi.org/10.1038/s41598-019-50133-z.10.1038/s41598-019-50133-z ↗
38. Santiago, TR, Bonatto, CC, Rossato, M, Lopes, CAP, Lopes, CA, Mizubuti, ESG, et al.. Green synthesis of silver nanoparticles using tomato leaf extract and their entrapment in chitosan nanoparticles to control bacterial wilt. J Sci Food Agric 2019;99:4248–59. https://doi.org/10.1002/jsfa.9656.10.1002/jsfa.9656 ↗
39. Paulkumar, K, Gnanajobitha, G, Vanaja, M, Pavunraj, M, Annadurai, G. Green synthesis of silver nanoparticle and silver-based chitosan bionanocomposite using stem extract of Saccharum officinarum and assessment of its antibacterial activity. Adv Nat Sci Nanosci Nanotechnol 2017;8:035019. https://doi.org/10.1088/2043-6254/aa7232.10.1088/2043-6254/aa7232 ↗
40. Kim, JH, Yu, D, Eom, SH, Kim, SH, Oh, J, Jung, WK, et al.. Synergistic antibacterial effects of chitosan–caffeic acid conjugate against antibiotic-resistant acne-related bacteria. Mar Drugs 2017;15:167. https://doi.org/10.3390/md15060167.10.3390/md15060167 ↗
41. Shameli, K, Ahmad, MB, Yunus, WMZW, Rustaiyan, A, Ibrahim, NA, Zargar, M, et al.. Green synthesis of silver/montmorillonite/chitosan bionanocomposites using the UV irradiation method and evaluation of antibacterial activity. Int J Nanomed 2010;5:875–87. https://doi.org/10.2147/IJN.S13632.10.2147/IJN.S13632 ↗
42. Huang, SQ, Yu, ZM, Zhang, Y, Qi, CS, Zhang, SF. In situ green synthesis of antimicrobial carboxymethyl chitosan–nanosilver hybrids with controlled silver release. Int J Nanomed 2017;12:3181–91. https://doi.org/10.2147/ijn.s130397.10.2147/IJN.S130397 ↗
43. Jia, J, Gai, Y, Wang, W, Zhao, Y. Green synthesis of biocompatible chitosan–graphene oxide hybrid nanosheet by ultrasonication method. Ultrason Sonochem 2016;32:300–6. https://doi.org/10.1016/j.ultsonch.2016.03.027.10.1016/j.ultsonch.2016.03.027 ↗
44. Shehabeldine, A, Hasanin, M. Green synthesis of hydrolyzed starch–chitosan nanocomposite as drug delivery system to gram-negative bacteria. Environ Nanotechnol Monit Manag 2019;12:100252. https://doi.org/10.1016/j.enmm.2019.100252.10.1016/j.enmm.2019.100252 ↗
45. Saeed, RM, Dmour, I, Taha, MO. Stable chitosan-based nanoparticles using polyphosphoric acid or hexametaphosphate for tandem ionotropic/covalent crosslinking and subsequent investigation as novel vehicles for drug delivery. Front Bioeng Biotechnol 2020;8:4. https://doi.org/10.3389/fbioe.2020.00004.10.3389/fbioe.2020.00004 ↗
46. Al-Naamani, L, Dobretsov, S, Dutta, J. Chitosan–zinc oxide nanoparticle composite coating for active food packaging applications. Innov Food Sci Emerg Technol 2016;38:231–7. https://doi.org/10.1016/j.ifset.2016.10.010.10.1016/j.ifset.2016.10.010 ↗
47. Syame, SM, Mohamed, WS, Mahmoud, RK, Omara, ST. Synthesis of copper–chitosan nanocomposites and its application in treatment of local pathogenic isolates bacteria. Orient J Chem 2017;33:2959–69. https://doi.org/10.13005/ojc/330632.10.13005/ojc/330632 ↗
48. Rico, CM, Majumdar, S, Duarte-Gardea, M, Peralta-Videa, JR, Gardea-Torresdey, JL. Interaction of nanoparticles with edible plants and their possible implications in the food chain. J Agric Food Chem 2011;59:3485–98. https://doi.org/10.1021/jf104517j.10.1021/jf104517j ↗
49. Raliya, R, Biswas, P, Tarafdar, JC. TiO2 nanoparticle biosynthesis and its physiological effect on mung bean (Vigna radiata L.). Biotechnol Rep 2015;5:22–6. https://doi.org/10.1016/j.btre.2014.10.009.10.1016/j.btre.2014.10.009 ↗
50. Dashtmian, AD, Hosseini Mazinani, SM, Pazoki, A. Exogenous chitosan nanoparticles modulated drought stress through changing yield, biochemical attributes, and fatty acid profile of common bean (Phaseolus vulgaris L.) cultivars. Gesunde Pflanz 2023;75:2463–76. https://doi.org/10.1007/s10343-023-00912-6.10.1007/s10343-023-00912-6 ↗
51. Oh, MM, Carey, EE, Rajashekar, CB. Regulated water deficits improve phytochemical concentration in lettuce. J Am Soc Hortic Sci 2019;135:223–9. https://doi.org/10.21273/jashs.135.3.223.10.21273/JASHS.135.3.223 ↗
52. Shin, YK, Bhandari, SR, Jo, JS, Song, JW, Lee, JG. Effect of drought stress on chlorophyll fluorescence parameters, phytochemical contents, and antioxidant activities in lettuce seedlings. Horticulturae 2021;7:238. https://doi.org/10.3390/horticulturae7080238.10.3390/horticulturae7080238 ↗
53. Javanmard, A, Ashrafi, M, Morshedloo, MR, Machiani, MA, Rasouli, F, Maggi, F, et al.. Optimizing phytochemical and physiological characteristics of balangu (Lallemantia iberica) by foliar application of chitosan nanoparticles and myco-root inoculation under water supply restrictions. Horticulturae 2022;8:695. https://doi.org/10.3390/horticulturae8080695.10.3390/horticulturae8080695 ↗
54. Das, D, Bisht, K, Chauhan, A, Gautam, S, Jaiswal, JP, Salvi, P, et al.. Morpho-physiological and biochemical responses in wheat foliar sprayed with zinc–chitosan–salicylic acid nanoparticles during drought stress. Plant Nano Biol 2023;4:100034. https://doi.org/10.1016/j.plana.2023.100034.10.1016/j.plana.2023.100034 ↗
55. Brown, A, Al-Azawi, TNI, Methela, NJ, Rolly, NK, Khan, M, Faluku, M, et al.. Chitosan–fulvic acid nanoparticles enhance drought tolerance in maize via antioxidant defense and transcriptional reprogramming. Physiol Plant 2024;176:e14455. https://doi.org/10.1111/ppl.14455.10.1111/ppl.14455 ↗
56. Dawood, MG, El-Awadi, MES, Sadak, MS. Chitosan and its nanoform regulates physiological processes and antioxidant mechanisms to improve drought stress tolerance of Vicia faba plant. J Soil Sci Plant Nutr 2024;24:5696–709. https://doi.org/10.1007/s42729-024-01934-3.10.1007/s42729-024-01934-3 ↗
57. Faluku, M, Al-Azawi, TNI, Methela, NJ, Khan, M, Huy, VN, Brown, A, et al.. Fulvic acid-releasing chitosan nanoparticles promote the growth and drought stress tolerance of rice plants. J Crop Health 2024;76:739–51. https://doi.org/10.1007/s10343-024-00979-9.10.1007/s10343-024-00979-9 ↗
58. Jabeen, Z, Mahmood, B, Rehman, S, Butt, AM, Hussain, N. Biodegradable glycine betaine encapsulated chitosan nanoparticles induce the expression of antioxidant enzyme genes to improve drought tolerance in maize. S Afr J Bot 2024;171:571–82. https://doi.org/10.1016/j.sajb.2024.06.012.10.1016/j.sajb.2024.06.012 ↗
59. Machiani, MA, Javanmard, A, Ostadi, A, Alizadeh, K. Improvement in essential oil quantity and quality of thyme (Thymus vulgaris L.) by integrative application of chitosan nanoparticles and arbuscular mycorrhizal fungi under water stress conditions. Plants 2023;12:1422. https://doi.org/10.3390/plants12071422.10.3390/plants12071422 ↗
60. Silveira, NM, Seabra, AB, Marcos, FC, Pelegrino, MT, Machado, EC, Ribeiro, RV. Encapsulation of S-nitrosoglutathione into chitosan nanoparticles improves drought tolerance of sugarcane plants. Nitric Oxide 2019;84:38–44. https://doi.org/10.1016/j.niox.2019.01.004.10.1016/j.niox.2019.01.004 ↗
61. Kavi Kishor, PB, Sreenivasulu, N. Is proline accumulation per se correlated with stress tolerance or is proline homeostasis a more critical issue? Plant Cell Environ 2014;37:300–11. https://doi.org/10.1111/pce.12157.10.1111/pce.12157 ↗
62. Rolland, F, Baena-Gonzalez, E, Sheen, J. Sugar sensing and signaling in plants: conserved and novel mechanisms. Annu Rev Plant Biol 2006;57:675–709. https://doi.org/10.1146/annurev.arplant.57.032905.105441.10.1146/annurev.arplant.57.032905.105441 ↗
63. Alizadeh, A, Moghaddam, M, Asgharzade, A, Sourestani, MM. Phytochemical and physiological response of Satureja hortensis L. to different irrigation regimes and chitosan application. Ind Crops Prod 2020;158:112990. https://doi.org/10.1016/j.indcrop.2020.112990.10.1016/j.indcrop.2020.112990 ↗
64. Jiao, Z, Li, Y, Li, J, Xu, X, Li, H, Lu, D, et al.. Effects of exogenous chitosan on physiological characteristics of potato seedlings under drought stress and rehydration. Potato Res 2012;55:293–301. https://doi.org/10.1007/s11540-012-9223-8.10.1007/s11540-012-9223-8 ↗
65. Rabêlo, VM, Magalhães, PC, Bressanin, LA, Carvalho, DT, Reis, COD, Karam, D, et al.. The foliar application of a mixture of semisynthetic chitosan derivatives induces tolerance to water deficit in maize, improving the antioxidant system and increasing photosynthesis and grain yield. Sci Rep 2019;9:1–13. https://doi.org/10.1038/s41598-019-44649-7.10.1038/s41598-019-44649-7 ↗
66. Divya, K, Thampi, M, Vijayan, S, Shabanamol, S, Jisha, MS. Chitosan nanoparticles as a rice growth promoter: evaluation of biological activity. Arch Microbiol 2021;204:95. https://doi.org/10.1007/s00203-021-02669-w.10.1007/s00203-021-02669-w ↗
67. Rizwan, M, Akhtar, G, Shehzad, MA, Aloufi, S, Suliman Alghanem, SM, Zulfiqar, F, et al.. Chitosan induces water deficit stress tolerance in Gladiolus grandiflorus via regulating water relations, gaseous exchange traits, and antioxidative defense. Plant Biosyst 2025;159:1552–68. https://doi.org/10.1080/11263504.2025.2543867.10.1080/11263504.2025.2543867 ↗
68. Dumanović, J, Nepovimova, E, Natić, M, Kuča, K, Jaćević, V. The significance of reactive oxygen species and antioxidant defense system in plants: a concise overview. Front Plant Sci 2021;11:552926. https://doi.org/10.3389/fpls.2020.552969.10.3389/fpls.2020.552969 ↗
69. Ji, H, Wang, J, Chen, F, Fan, N, Wang, X, Xiao, Z, et al.. Meta-analysis of chitosan-mediated effects on plant defense against oxidative stress. Sci Total Environ 2022;851:158212. https://doi.org/10.1016/j.scitotenv.2022.158212.10.1016/j.scitotenv.2022.158212 ↗
70. Yu, J, Wang, D, Geetha, N, Khawar, KM, Jogaiah, S, Mujtaba, M, et al.. Current trends and challenges in the synthesis and applications of chitosan-based nanocomposites for plants: a review. Carbohydr Polym 2021;261:117904. https://doi.org/10.1016/j.carbpol.2021.117904.10.1016/j.carbpol.2021.117904 ↗
71. Liu, J, Gai, L, Zong, H. Foliage application of chitosan alleviates the adverse effects of cadmium stress in wheat seedlings (Triticum aestivum L.). Plant Physiol Biochem 2021;164:115–21. https://doi.org/10.1016/j.plaphy.2021.04.038.10.1016/j.plaphy.2021.04.038 ↗
72. Sheikhalipour, M, Esmaielpour, B, Behnamian, M, Gohari, G, Giglou, MT, Vachova, P, et al.. Chitosan–selenium nanoparticle (Cs–Se NP) foliar spray alleviates salt stress in bitter melon. Nanomaterials 2021;11:684. https://doi.org/10.3390/nano11030684.10.3390/nano11030684 ↗
73. Rasheed, R, Ashraf, MA, Arshad, A, Iqbal, M, Hussain, I. Interactive effects of chitosan and cadmium on growth, secondary metabolism, oxidative defense, and element uptake in pea (Pisum sativum L.). Arab J Geosci 2020;13:847. https://doi.org/10.1007/s12517-020-05871-0.10.1007/s12517-020-05871-0 ↗
74. Choudhary, RC, Kumaraswamy, RV, Kumari, S, Sharma, SS, Pal, A, Raliya, R, et al.. Cu–chitosan nanoparticle boosts defense responses and plant growth in maize (Zea mays L.). Sci Rep 2017;7:9754. https://doi.org/10.1038/s41598-017-08571-0.10.1038/s41598-017-08571-0 ↗
75. Oliveira, HC, Gomes, BC, Pelegrino, MT, Seabra, AB. Nitric oxide-releasing chitosan nanoparticles alleviate the effects of salt stress in maize plants. Nitric Oxide 2016;61:10–19. https://doi.org/10.1016/j.niox.2016.09.010.10.1016/j.niox.2016.09.010 ↗
76. Singh, BN, Dwivedi, P, Sarma, BK, Singh, GS, Singh, HB. Trichoderma asperellum T42 reprograms tobacco for enhanced nitrogen utilization efficiency and plant growth when fed with N nutrients. Front Plant Sci 2018;9:163. https://doi.org/10.3389/fpls.2018.00163.10.3389/fpls.2018.00163 ↗
77. Kashyap, PL, Xiang, X, Heiden, P. Chitosan nanoparticle-based delivery systems for sustainable agriculture. Int J Biol Macromol 2015;77:36–51. https://doi.org/10.1016/j.ijbiomac.2015.02.039.10.1016/j.ijbiomac.2015.02.039 ↗
78. Chandra, S, Chakraborty, N, Dasgupta, A, Sarkar, J, Panda, K, Acharya, K, et al.. Chitosan nanoparticles: a positive modulator of innate immune responses in plants. Sci Rep 2015;5:15195. https://doi.org/10.1038/srep15195.10.1038/srep15195 ↗
79. Asgari-Targhi, G, Iranbakhsh, A, Ardebili, ZO. Potential benefits and phytotoxicity of bulk and nano-chitosan on the growth, morphogenesis, physiology, and micropropagation of Capsicum annuum. Plant Physiol Biochem 2018;127:393–402. https://doi.org/10.1016/j.plaphy.2018.04.013.10.1016/j.plaphy.2018.04.013 ↗
80. Abdel-Aziz, H. Effect of priming with chitosan nanoparticles on germination, seedling growth and antioxidant enzymes of broad beans. Catrina Int J Environ Sci 2019;18:81–6. https://doi.org/10.21608/cat.2019.28609.10.21608/cat.2019.28609 ↗