*Article not assigned to an issue yet
Sadak Mervat Shamoon, El-Awadi Mohamed El-sayed, Dawood Mona Gergis, Abdel-Baky Yasser Refaai
Keywords: n Glycine max L., Water deficit, Polysaccharides, Chitin, Chitosan
Today, one of the most serious hazards to the world’s population is drought, which is expected to worsen in many areas due to decreased rainfall and increased evaporation triggered by global climate change. So, a pot experiment was conducted at the greenhouse of National Research Centre, Dokki, Giza, Egypt, during summer season 2023 and 2024 to evaluate the physiological role of chitosan (1 gL−1, 2 gL−1, 3 gL−1) in increasing drought tolerance (60% WFC, water field capacity) of soybean. Results show that drought stress at 60% water-holding capacity (WHC) significantly reduced plant growth, photosynthetic pigments, indole acetic acid, seed yield, and seed quality (oil, carbohydrate, and protein). It also increased phenolic compounds, osmolytes (total soluble sugars, proline, total soluble protein), membrane leakage, hydrogen peroxide, lipid peroxidation, and the activities of antioxidant enzymes (SOD, CAT, and POX), compared with well-watered plants (90% WFC). Conversely, foliar-applied chitosan (1 gL−1, 2 gL−1, and 3 gL−1) improves drought tolerance in soybean plants grown at 60% water-holding capacity (WFC). Chitosan application markedly enhanced growth, yield, seed quality, and physiological responses in both well-watered and drought-stressed plants. Chitosan promoted phenolics, osmolytes, membrane stability, and antioxidant enzyme activity while reducing oxidative stress indicators such as membrane leakage, hydrogen peroxide, and lipid peroxidation. It is obvious that 2 gL−1 chitosan is the most effective treatment. It could be concluded that chitosan act as an eco-friendly biostimulant to enhance drought resilience in soybean by improving physiological, biochemical, and growth traits, offering a sustainable strategy for maintaining crop productivity under water-limited conditions.
Afzal S, Chaudhary N, Singh NK (2021) Role of soluble sugars in metabolism and sensing under abiotic stress. Plant growth regulators: signaling under stress conditions. Springer International Publishing, Switzerland, pp 305–334. https://doi.org/10.1007/978-3-030-61153-8_14
Ahluwalia O, Singh PC, Bhatia R (2021) A review on drought stress in plants: implications, mitigation and the role of plant growth promoting rhizobacteria. Resour Environ Sustain 5:100032. https://doi.org/10.1016/j.resenv.2021.100032
Albalasmeh AA, Berhe AA, Ghezzehei TA (2013) A new method for rapid determination of carbohydrate and total carbon concentrations using UV spectrophotometry. Carbohydr Polym 97(2):253–261. https://doi.org/10.1016/j.carbpol.2013.04.072
Alenazi MM, El-Ebidy AM, El-shehaby OA, Seleiman MF, Aldhuwaib KJ, Abdel-Aziz HMM (2024) Chitosan and chitosan nanoparticles differentially alleviate salinity stress in Phaseolus vulgaris L. plants. Plants 13:398. https://doi.org/10.3390/plants13030398
Ali A, Alqurainy F (2006) Activities of antioxidants in plants under environmental stress. In: Motohashi N (ed) The lutein prevention and treatment for diseases. Trans-world Research Network, India, pp 187–256
Ali EF, El-Shehawi AM, Ibrahim OHM, Abdul-Hafeez EY, Moussa MM, Hassan FAS (2021) A vital role of chitosan nanoparticles in improvisation the drought stress tolerance in Catharanthus roseus (L.) through biochemical and gene expression modulation. Plant Physiol Biochem 161:166–175
Ali S, Shakoor A, Ali Q, Chattha MS, El-Sheikh MA, Ali S (2022) Oxidative stress alleviation through enzymatic and non-enzymatic antioxidants and osmoregulators generation in barley (Hordeum vulgare L.) under salt (Nacl) stress by ascorbic acid (ASA). Pak J Bot 54:7–15. https://doi.org/10.30848/PJB2022-1
AOAC (Association of official agriculture chemists) (2000) Official methods of analysis, 17th eds. Association of official agriculture chemists, Gaithersburg
Ashraf M, Iram A (2005) Drought stress induced changes in some organic substances in nodules and other plant parts of two potential legumes differing in salt tolerance. Flora 200:535546
Ashraf M, Shahbaz M, Ali Q (2013) Drought induced modulation in growth and mineral nutrients in canola (Brassica napus L.). Pak J Bot 45:93–98
Avila RG, Magalhaes PC, Silva EM, Lana UGP, Alvarenga AA, Souza TC (2020) Silicon supplementation improves tolerance to water deficiency in sorghum plants by increasing root system growth and improving photosynthesis. Silicon. https://doi.org/10.1007/s12633-019-00349-5
Avila RG, Magalhaes PC, Silva EM, Souza KRD, Campos CN, Alvarenga AA, Souza TC (2021) Application of silicon to irrigated and water deficit sorghum plants increases yield via the regulation of primary, antioxidant, and osmoregulatory metabolism. Agric Water Manag 1016:1016–1070. https://doi.org/10.1016/j.agwat.2021.107004
Avila RG, MagalhAes PC, Vitorino LC, Bessa LA, Souza KR, Queiroz RP, Jakelaitis A, Teixeira MB (2023) Chitosan induces sorghum tolerance to water deficits by positively regulating photosynthesis and the production of primary metabolites, osmoregulators, and antioxidants. J Soil Sci Plant Nutr 23:1156–1172. https://doi.org/10.1007/s42729-022-01111-4
Banon SJ, Ochoa J, Franco JA, Alarcon JJ, Sanchez-Blanco MJ (2006) Hardening of oleander seedlings by deficit irrigation and low air humidity. Environ Exp Bot 56:36–43. https://doi.org/10.1016/j.envexpbot.2004.12.004
Basal O, Szabó A (2020) Physiomorphology of soybean as affected by drought stress and nitrogen application. Scientifica. https://doi.org/10.1155/2020/6093836
Behboudi F, Tahmasebi Sarvestani Z, Kassaee MZ, Modares Sanavi SAM, Sorooshzadeh A, Ahmadi SB (2018) Evaluation of chitosan nanoparticles effects on yield and yield components of barley (Hordeum vulgare L.) under late season drought stress. J Water Environ Nanotechnol 3(1):22–39
Behboudi F, Tahmasebi-Sarvestani Z, Kassaee MZ, Modarres-Sanavy SAM, Sorooshzadeh A, Mokhtassi-Bidgoli A (2019) Evaluation of chitosan nanoparticles effects with two application methods on wheat under drought stress. J Plant Nutr 42(13):1439–1451
Buezo J, Sanz-Saez Á, Moran JF, Soba D, Aranjuelo I, Esteban R (2019) Drought tolerance response of high-yielding soybean varieties to mild drought: physiological and photochemical adjustments. Physiol Plant 166(1):88–104
Chakraborty M, Hasanuzzaman M, Rahman M, Khan MAR, Bhowmik P, Mahmud NU, Tanveer M, Islam T (2020) Mechanism of plant growth promotion and disease suppression by chitosan biopolymer. Agriculture 10:624
Chapman HD, Pratt PF 1978 Method of Analysis for Soil, Plant and Water, California University, Division Agric Sci, Priced Publication, pp 50–169
Chen JX, Wang XF 2006 Plant physiology experimental guide. High Educ Press Beijing, China 24–25:55–56
Chibu H, Shibayama H (2001) Effects of chitosan applications on the growth of several crops. In: Uragami T, Kurita K, Fukamizo T (eds) Chitin and chitosan in life science. Yamaguchi, Japan, pp 235–239
Chow PS, Landhäusser S (2004) A method for routine measurements of total sugar and starch content in woody plant tissues. Tree Physiol 24(10):1129–1136. https://doi.org/10.1093/treephys/24.10.1129
Creelman RA, Mason HG, Bensen RJ, Boyer JS, Mullet JE (1990) Water deficit and abscisic acid causes inhibition of shoots versus root growth in soybean seedlings: analysis of growth, sugar accumulation and gene expression. Plant Physiol 92:205–214
Czekus Z, Iqbal N, Pollak B, Martics A, Ordog A, Poor P (2021) Role of ethylene and light in chitosan-induced local and systemic defence responses of tomato plants. J Plant Physiol 263:153461. https://doi.org/10.1016/j.jplph.2021.153461
Daryanto S, Wang L, Jacinthe PA (2020) Global synthesis of drought effects on cereal, legume, tuber and root crops production: a review. Agric Water Manag 179:18–33
Das M, Das SK, Suthar SH (2002) Composition of seed and characteristics of oil from Karingda. Int J Food Sci Technol 37:893–896. https://doi.org/10.1046/j.1365-2621.2002.00638.x
Dawood MG, Sadak MSh. (2014) Physiological role of glycinebetaine in alleviating the deleterious effects of drought stress on canola plants (Brassica napus L.). Middle East J Agric Res 3(4):943–954
Dawood MG, Khater MA, El-Awadi ME, Badr NM, Shalaby MAF, Gamal El-Din K, Ahmed MA (2023) Alleviation the deleterious impact of water stress on Sorghum bicolor (L.) plants using biostimulant humic acid. Middle East J Agric Res 12(3):520–535. https://doi.org/10.36632/mejar/2023.12.3.34
Diatta AA, Fike JH, Battaglia ML, Galbraith J, Baig MB (2020) Effects of biochar on soil fertility and crop productivity in arid regions: a review. Arab J Geosci 13:595
Din J, Kans SU, Ali J, Gurmani AR (2011) Physiological and agronomic response of canola varieties to drought stress. J of Anim Plant Sci 21:78–82
Dos Reis CO, Magalhaes PC, Avila RG, Almeida LG, Rabelo VM, Carvalho DT, Cabral DF, Karam D, de Souza TC (2019) Action of N-Succinyl and N, O-Dicarboxymethyl chitosan derivatives on chlorophyll photosynthesis and fluorescence in drought-sensitive maize. J Plant Growth Regul 38:619–630
Du X, Zhang X, Chen X, Jin W, Huang Z, Kong L (2024) Drought stress reduces the photosynthetic source of subtending leaves and the transit sink function of podshells, leading to reduced seed weight in soybean plants. Front Plant Sci 15:1337544. https://doi.org/10.3389/fpls.2024.1337544
Duncan DB (1955) Multiple range and multiple F tests. Biometrics 11:1–42
El F, AmeranyRhazi M, Balcke G, Wahbi S, Meddich A, Taourirte M, Hause B (2022) The effect of chitosan on plant physiology, wound response, and fruit quality of tomato. Polymers 14:5006. https://doi.org/10.3390/polym14225006
El-Awadi ME, Sadak MSh., Dawood MG (2021) Comparative effect of potassium and banana peel in alleviating the deleterious effect of water deficit on soybean plants. J Mater Environ Sci 12(7):929–943
Elewa TA, Sadak MSh., Dawood MG (2017) Improving drought tolerance of quinoa plant by foliar treatment of trehalose. Agric Eng Int CIGR J 19(5):245–254
El-Zamily EAM, EL-Shafey AS, Dawood MG, El-Awadi ME (2024) Investigating the physiological roles of bulk chitosan and nano-chitosan in growth, flowering, yield attributes of Brassica napus L. and in nutrient delivery in sandy soils. Middle East J Appl Sci 14(1):161–179. https://doi.org/10.36632/mejas/2024.14.1.11
Eriksen RL, Padgitt-Cobb LK, Townsend MS, Henning JA (2021) Gene expression for secondary metabolite biosynthesis in hop (Humulus lupulus L.) leaf lupulin glands exposed to heat and low-water stress. Sci Rep 11(1):5138. https://doi.org/10.1038/s41598-021-84691-y
Esmaeeli M, Roozbahani A, Daneshian J (2024) Combined effects chitosan and genotype on agronomic, physiologic, and biochemical characteristics of soybean under drought stress conditions. S Afr J Bot 174:678–685. https://doi.org/10.1016/j.sajb.2024.09.036
Faqir Y, Ma J, Chai Y (2021) Chitosan in modern agriculture production. Plant Soil Environ 67(12):679–699. https://doi.org/10.17221/332/2021-PSE
Fazeli F, Ghorbanli M, Niknam V (2007) Effect of drought on biomass, protein content, lipid peroxidation and antioxidant enzymes in two sesame cultivars. Biol Plant 51(1):98–103
García-García AL, García-Machado FJ, Borges AA, Morales-Sierra S, Boto A, Jiménez-Arias D (2020) Pure organic active compounds against abiotic stress: a biostimulant overview. Front Plant Sci 11:575829. https://doi.org/10.3389/fpls.2020.575829
Garcı́a-Garcı́a AL, Matos AR, Feijão E, Cruz de Carvalho R, Boto A, Marques da Silva J, Jime´ nez-Arias D (2023) The use of chitosan oligosaccharide to improve artemisinin yield in well watered and drought-stressed plants. Front Plant Sci 14:1200898. https://doi.org/10.3389/fpls.2023.1200898
Ge TD, Sui FG, Bai LP, Yin-yan LU, Guang-sheng Z (2006) Effects of water stress on the protective enzyme activities and lipid per- oxidation in roots and leaves of summer maize. Agric Sci China 5(4):291–298. https://doi.org/10.1016/S1671-2927(06)60052-7
Geng W, Zhou L, Muhammad JH, Yan P (2020) Chitosan regulates metabolic balance, polyamine accumulation, and Na+ transport contributing to salt tolerance in creeping bentgrass Geng. BMC Plant Biol 20:506
Gill SS, Tuteja N (2010) Reactive oxygen species and antioxidant machinery in abiotic stress tolerance in crop plants. Plant Physiol Biochem 48:909–930. https://doi.org/10.1016/j.plaphy.2010.08.016
Gonzalez MB, Guzman R, Rudkyk E, Romano M, Molina AA (2003) Spectrophotometric determination of phenolic compounds in propolis. Lat Am J Pharm 22(3):243–248
Guan Y, Hu J, Wang X, Shao C (2009) Seed priming with chitosan improves maize germination and seedling growth in relation to physiological changes under low temperature stress. J Zhejiang Univ Sci B 10:427–433. https://doi.org/10.1631/jzus.b0820373
Gusmiaty M, Restu A, Payangan RY (2019) Production of IAA (Indole Acetic Acid) of the rhizosphere fungus in the suren community forest stand. IOP Conf Ser Earth Environ Sci 343:012058
Hasanuzzaman M, Bhuyan M, Zulfiqar F, Raza A, Mohsin SM, Mahmud JA, Fujita M, Fotopoulos V (2020) Reactive oxygen species and antioxidant defense in plants under abiotic stress: revisiting the crucial role of a universal defense regulator. Antioxidants 9:681
Hassnain K, Alam M, Ahmad I, Basit A, Ullah I, Alam N, Ullah I, Khalid MA, Muhammad B, Shair M (2020) Efficacy of chitosan on performance of tomato (Lycopersicon esculentum L.) plant under water stress condition. Pak J Agric Res 33(1):27–41. https://doi.org/10.17582/journal.pjar/2020/33.1.27.41
Hidangmayum A, Dwivedi P, Katiyar D, Hemantaranjan A (2019) Application of chitosan on plant responses with special reference to abiotic stress. Physiol Mol Biol Plants 25(2):313–326
Hodges DM, DeLong JM, Forney C, Prange PK (1999) Improving the thiobarbaturic acid reactive substances assay for estimating lipid peroxidation in plant tissues containing anthocyanin and other interfering compounds. Planta 207:604–611
Hong ZL, Lakkineni K, Zhang ZM, Verma DPS (2000) Removal of feedback inhibition of Δ1-pyrroline-5-carboxylate synthetase results in increased proline accumulation and protection of plants from osmotic stress,. Plant Physiol 12:1129–1136
Ibrahim EA, Ebrahim NE, Mohamed GZ (2023) Effect of water stress and foliar application of chitosan and glycine betaine on lettuce. Sci Rep 13(1):17274
Jiao Z, Li Y, Li J, Xu X, Li H, Lu D, Wang D (2012) Effects of exogenous chitosan on physiological characteristics of potato seedlings under drought stress and rehydration. Potato Res 55(3):293–301
Jie Z, Yuncong Y, Streeter JG, Ferree DC (2010) Influence of soil drought stress on photosynthesis, carbohydrates and the nitrogen and phophorus absorb in different section of leaves and stem of Fugi/M.9EML, a young apple seedlings. Afr J Biotechnol 9:5320–5325
Jogawat A, Yadav B, Chhaya LN, Singh AK, Narayan OP (2021) Crosstalk between phytohormones and secondary metabolites in the drought stress tolerance of crop plants: a review. Physiol Plant 172(2):1106–1132. https://doi.org/10.1111/ppl.13328
Jomo M, Netondo W, Musyimi M (2016) Drought inhibition of chlorophyll content among seven Amaranthuss species. Int J Adv Res Sci Eng Technol 3:1362–1371
Kalsoom U, Bennett IJ, Boyce MC (2016) A review of extraction and analysis: methods for studying osmoregulants in plants. Chromatogr J Sep Tech 7:1. https://doi.org/10.4172/2157-7064.1000315
Karimi S, Abbaspour H, Sinaki JM, Makarian H (2012) Effects of water deficit and chitosan spraying on osmotic adjustment and soluble protein of cultivars castor bean (Ricinus communis L.). J Stress Physiol Biochem 28:160–169
Katiyar D, Hemantaranjan A, Singh B (2015) Chitosan as a promising natural compound to enhance potential physiological responses in plant: a review. Indian J Plant Physiol 20(1):1–9. https://doi.org/10.1007/s40502-015-0139-6
Kayes A, Araf T, Mustaki S, Islam N, Choudhury S (2024) Effects of biochar and chitosan on morpho-physiological, biochemical and yield traits of water-stressed tomato plants. Int J Hortic Agric Food Sci 8(2):1–9. https://doi.org/10.22161/ijhaf.8.2
Kazimi R, Saxena D (2023) Significance of chitosan foliar spraying on the growth and yield of vegetable crop under protected cultivation: a review. Plant Sci Today 10(3):140–148. https://doi.org/10.14719/pst.2204
Khalil HA, BadrEldin RM (2021) Chitosan improves morphological and physiological attributes of grapevines under deficit irrigation conditions. J Hortic Res 29(1):9–22. https://doi.org/10.2478/johr-2021-0003
Khan WM, Prithiviraj B, Smiyh D (2002) Effect of foliar application of chitin oligosaccharides on photosynthesis of maize and soybean. Photosynthetica 40(4):621–626. https://doi.org/10.1023/A:1024320606812
Kijowska-Oberc J, Dylewski T, Ratajczak E (2023) Proline concentrations in seedlings of woody plants change with drought stress duration and are mediated by seed characteristics: a meta-analysis. Sci Rep 13(1):15157
Kirova E, Pecheva D, Simova-Stoilova L (2021) Drought response in winter wheat: protection from oxidative stress and mutagenesis effect. Acta Physiol Plant 43:1–11. https://doi.org/10.1007/S11738-020-03182-1
Kocaman B (2025) Effects of foliar application of chitosan on some vegetative growth and biochemical parameters of strawberry under salt stress. Indian J Pharm Educ Res 59(1):1–8
Ksouri R, Megdiche W, Debez A, Falleh H, Grignon C, Abdelly C (2007) Salinity effects on polyphenol content and antioxidant activities in leaves of the halophyte Cakile maritima. Plant Physiol Biochem 45(3–4):244–249
Kumar KB, Khan PA (1982) Peroxidase and polyphenol oxidase in excised ragi (Eleusine coracana cv. PR 202) leaves during senescence. Indian J Exp Biol 20:412–416
Kumar A, Memo M, Mastinu A (2020) Plant behaviour: an evolutionary response to the environment? Plant Biol 22(6):961–970. https://doi.org/10.1111/plb.13149
Landi L, De Miccolis Angelini RM, Pollastro E, Feliziani S, Faretra F, Romanazzi G (2017) Global transcriptome analysis and identification of differentially expressed genes in strawberry after preharvest application of Benzothiadiazole and chitosan. Front Plant Sci 8:23. https://doi.org/10.3389/fpls.2017.002
Lawlor DW, Cornic G (2002) Photosynthetic carbon assimilation and associated metabolism in relation to water deficits in higher plants. Plant Cell Environ 25:275–294. https://doi.org/10.1046/j.0016-8025.2001.00814.x
Laxa M, Liebthal M, Telman W, Chibani K, Dietz KJ (2019) The role of the plant antioxidant system in drought tolerance. Antioxidants Basel 8:94
Li Y, Chen M (2015) Novel chlorophylls and new directions in photosynthesis research. Funct Plant Biol 42:493–501
Li Z, Zhang Y, Zhang X, Merewitz E, Peng Y, Ma X, Huang L, Yan Y (2017) Metabolic pathways regulated by chitosan contributing to drought resistance in white clover. J Proteome Res 16:3039–3052
Li R, He J, Xie H, Wang W, Bose SK, Sun Y, Hu J, Yin H (2019) Effects of chitosan nanoparticles on seed germination and seedling growth of wheat (Triticum aestivum L.),. Int J Biol Macromol 126:91–100
Liu F, Andersen MN, Jensen CR (2003) Loss of pod set caused by drought stress is associated with water status and ABA content of reproductive structures in soybean. Funct Plant Biol 30:271–280. https://doi.org/10.1071/FP02185
Makhdum MI, Shababuddin M (2006) Effects of different doses of glycine betaine and time of spray application on yield of cotton (Gossypium hirsutum L). J Res Sci 17:241–245
Malerba M, Cerana R (2020) Chitinand chitosan-based derivatives in plant protection against biotic and abiotic stresses and in recovery of contaminated soil and water. Polysaccharides Basel 1(1):21–30
Marcinska I, Czyczylo-Mysza I, Skrzypek M, Filek E, Grzesiak S (2013) Impact of osmotic stress on physiological and biochemical characteristics in drought-susceptible and drought-resistant wheat genotypes. Acta Physiol Plant 35:451–461
Mecozzi M (2005) Estimation of total carbohydrate amount in environmental samples by the phenol-sulphuric acid method assisted by multivariate calibration. Chemom Intell Lab Syst 79:84–90
Methela NJ, Pande A, Islam MS, Rahim W, Hussain A, Lee D-S, Mun B-G, Raj MNP, Kim S-J, Kim Y, Yun B-W (2023) Chitosan-GSNO nanoparticles: a positive modulator of drought stress tolerance in soybean. BMC Plant Biol 23(1):639. https://doi.org/10.1186/s12870-023-04640-x
Mittal D, Kaur G, Singh P, Yadav K, Ali SA (2020) Nanoparticle-based sustainable agriculture and food science: recent advances and future outlook. Front Nanotechnol 2:579954
Moolphuerk N, Lawson T, Pattanagul W (2022) Chitosan mitigates the adverse effects and improves photosynthetic activity in rice (Oryza sativa L.) seedlings under drought condition. J Crop Improv 36(5):638–655
Muley AB, Shingote PR, Patil AP, Dalvi SG, Suprasanna P (2019) Gamma radiation degradation of chitosan for application in growth promotion and induction of stress tolerance in potato (Solanum tuberosum L.). Carbohydr Polym 210:289–301
Muthukrishnan S, Murugan I, Selvaraj M (2019) Chitosan nanoparticles loadedwith thiamine. stimulate growth and enhances protection against wilt disease in chickpea. Carbohyd Polym 15(212):169–177. https://doi.org/10.1016/j.carbpol.2019.02.037
Nazarli H, Faraji F, Zardashti MR (2011) Effect of drought stress and polymer on osmotic adjustment and photosynthetic pigments of sunflower. Cercet Agron Moldova 44:35–41. https://doi.org/10.2478/v10298-012-0022-9
Noctor G, Veljovic-Jovanovic S, Foyer CH (2000) Peroxide processing in photosynthesis: antioxidant coupling and redox signalling. Proc R Soc Lond B Biol Sci 355:1465–1475
Nurliana S, Fachriza S, Hemelda NM, Yuniati R 2022 Chitosan application for maintaining the growth of lettuce (Lactuca sativa) under drought condition. In: The 4th Int Conf Food Agric IOP Conf Series: Earth Environ Sci, vol 980. IOP, p 012013. https://doi.org/10.1088/1755-1315/980/1/012013
Oh MM, Trick HN, Rajashekar C (2009) Secondary metabolism and antioxidants are involved in environmental adaptation and stress tolerance in lettuce. J Plant Physiol 166(2):180–191
Okorie VO, Mphambukeli TN, Amusan SO (2019) Exploring the political economy of water and food security nexus in BRICS. Afr Insight 48:21–38
Phothi R, Theerakarunwong CD (2017) Effect of chitosan on physiology, photosynthesis and biomass of rice (Oryza sativa L.) under elevated ozone. Aust J Crop Sci 11(5):624–630
Pirbalouti AG, Malekpoor F, Salimi A, Golparvard A (2017) 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. Sci Hortic 217:114–122
Pongprayoon W, Siringam T, Panya A, Roytrakul S (2022) Application of chitosan in plant defense responses to biotic and abiotic stresses. Appl Sci Eng Progress 15(1):1–10
Qi J, Song CP, Wang B, Zhou J, Kangasjarvi J, Zhu JK, Gong Z (2018) Reactive oxygen species signaling and stomatal movement in plant responses to drought stress and pathogen attack. J Integr Plant Biol 60:805–826
Rabêlo VM, Magalhães PC, Bressanin LA, Teixeira CD, dos Oliveira RC, Karam DK, Antônio Carlos DA, dos Henrique SM, dos Rodrigues Santos SF, de Thiago Corrêa ST (2019) 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 9:8164. https://doi.org/10.1038/s41598-019-44649-7
Rady MM, Sadak MSh., El-Bassiouny HMS, Abd El-Monem AA (2011) Alleviation the adverse effects of salinity stress in sunflower cultivars using nicotinamide and 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 α-tocopherol. Aust J Basic Appl Sci 5(10):342–355
Rahman M, Mukta JA, Sabir AA, Gupta DR, Mohi-Ud-Din M, Hasanuzzaman M, Islam MT (2018) Chitosan biopolymer promotes yield and stimulates accumulation of antioxidants in strawberry fruit. PLoS ONE 13(9):e0203769
Ray SR, Bhuiyan MH, Hossain MA, Hasan AK, Sharmin S (2016) Chitosan ameliorates growth and biochemical attributes in mungbean varieties under saline condition. Res Agric Livest Fish 3:45–51
Reddy TY, Reddy VR, Anbumozhi N (2003) Physiological response of groundnut to drought stress and its amelioration. Plant Growth Regul 41(3):75–78. https://doi.org/10.1023/A:1027353430164
Sachdev S, Ansari SA, Ansari MI, Fujita M, Hasanuzzaman M (2021) Abiotic stress and reactive oxygen species: generation, signaling, and defense mechanisms. Antioxidants 10(2):277
Saharan V, Pal A (2016) Chitosan based nanomaterials in plant growth and protection. Springer, New Delhi, India, pp 33–41
Salehi-Lisar SY, Bakhshayeshan-Agdam H (2020) Agronomic crop responses and tolerance to drought stress. Agronomic crops. Springer, Berlin/Heidelberg, Germany, pp 63–91
Sanchez-Rodriguez E, Moreno DA, Ferreres F, del Mar RW, Ruiz JM (2011) Differential responses of five cherry tomato varieties to water stress: changes on phenolic metabolites and related enzymes. Phytochemistry 72(8):723–729
Seleiman MF, Al-Suhaibani N, Ali N, Akmal M, Alotaibi M, Refay Y, Dindaroglu T, Abdul Wajid HH, Battaglia ML (2021) Drought stress impact on plants and different approaches to alleviate its adverse effects. Plants 10:259
Shehzad MA, Nawaz F, Ahmad F, Ahmad N, Masood S (2020) Protective effect of potassium and chitosan supply on growth, physiological processes and antioxidative machinery in sunflower (Helianthus annuus L.) under drought stress. Ecotoxicol Environ Saf 187(109841):1–11
Shinde NA, Kawar PG, Dalvi SG (2024) Chitosan-based nanoconjugates: a promising solution for enhancing crops drought-stress resilience and sustainable yield in the face of climate change. Plant Nano Biol 7:100059. https://doi.org/10.1016/j.plana.2024.100059
Silva V, Singh RK, Gomes N, Soares BG, Silva A, Falco V (2020) Comparative insight upon chitosan solution and chitosan nanoparticles application on the phenolic content, antioxidant and antimicrobial activities of individual grape components of Sousao variety. Antioxidants 9(2):178
Singh S, Sinha S (2005) Accumulation of metals and its effects in Brassica juncea (L.) Czern. (cv. Rohini) grown on various amendments of tannery waste. Ecotoxicol Environ Saf 62:118–127. https://doi.org/10.1016/j.ecoenv.2004.12.026
Snedecor GW, Cochran WG (1990) Statistical methods, 8th edn. Iowa State University Press, Ames, p 609
Sorrequieta A, Ferraro G, Boggio SB, Valle EM (2010) Free amino acid production during tomato fruit ripening: a focus on L‐glutamate. Amino Acids 38:1523–1532
Su Y, Xu L, Wang S, Wang Z, Yang Y, Chen Y, Que Y (2015) Identification, phylogeny and transcript of chitinase family genes in sugarcane. Sci Rep 5(1):10708
Sweeney DW, Long JH, Kirkham MB (2003) A single irrigation to improve early maturing soybean yield and quality. Soil Sci Soc Am J 67:235–240
Ullah N, Basit A, Ahmad I, Ullah I, Shah ST, Mohamed HI, Javed S (2020) Mitigation the adverse effect of salinity stress on the performance of the tomato crop by exogenous application of chitosan. Bull Natl Res Cent 44:181
Ullah A, Bano A, Khan N (2021) Climate change and salinity effects on crops and chemical communication between plants and plant growth-promoting microorganisms under stress. Front Sustain Food Syst 5:618092
Vahala J, Ruonala R, Keinänen M, Tuominen H, Kangasjärvi J (2003) Ethylene insensitivity modulates ozone-induced cell death in birch. Plant Physiol 132:185–195
Vaseva-Gemisheva I, Lee D, Karanov E (2005) Response of Pisum sativum cytokinin oxidase /dehydrogenase expression and specific activity to drought stress and herbicide treatments. Plant Growth Regul 46:199–208
Walker-Simmons M, Hadwiger LA, Ryan CA (1983) Chitosan and peptic polysaccharides both induce the accumulation of the antifungal phytoalexin pisatin in pea pods and antinutrient proteinase inhibitors in tomato leaves. Biochem Biophys Res Commun 110:194–199
Wang X, Wu Z, Zhou Q, Wang X, Song S, Dong S (2022) Physiological response of soybean plants to water deficit. Front Plant Sci 12:809692. https://doi.org/10.3389/fpls.2021.809692
Wardlaw I, Willenbrink J (2000) Mobilization of fructan reserves and changes in enzyme activities in wheat stems correlate with water stress during kernel filling. New Phytol 148(3):413–422
Wu HH, Zou YN, Rahman MM, Ni QD, Wu QS (2017) Mycorrhizas alter sucrose and proline metabolism in trifoliate orange exposed to drought stress. Sci Rep 7:42389
Xie WM, Xu PX, Liu Q (2001) Antioxidant activity of water-soluble chitosan derivatives. Bioorg Med Chem Lett 11:1699–1701
Yang X, Lu M, Wang Y, Wang Y, Liu Z, Chen S (2021) Response mechanism of plants to drought stress. Horticulturae 7:50
Yin H, Fretté XC, Chrestensen LP, Grevsen K (2012) Chitosan oligosaccharides promote the content of polyphenols in Greek oregano (Origanum vulgare ssp. hirtum). J Agric Food Chem 60(1):136–143. https://doi.org/10.1021/jf204376j
Yu CW, Murphym TM, Lin CH (2003) Hydrogen peroxide-induced chilling tolerance in mung beans mediated through ABA-independent glutathione accumulation. Funct Plant Biol 30(9):955–963. https://doi.org/10.1071/FP03091
Zhang XQ, Li KC, Xing R, Liu S, Li P (2017) Metabolite profiling of wheat seedlings induced by chitosan: revelation of the enhanced carbon and nitrogen metabolism. Front Plant Sci 8:2017–7029
Botany Department, Agriculture and Biological Institute, National Research Centre, Giza, Egypt