Salinity stress ranks among the most prevalent stress globally, contributing to soil deterioration. Its negative impacts on crop productivity stem from mechanisms such as osmotic stress, ion toxicity, and oxidative stress, all of which impede plant growth and yield. The effect of cobalt with proline on mitigating salinity impact in radish plants is still unclear. That's why the current study was conducted with aim to explore the impact of different levels of Co and proline on radish cultivated in salt affected soils. There were four levels of cobalt, i.e., (0, 10, 15 and 20 mg/L) applied as CoSO4 and two levels of proline (0 and 0.25 mM), which were applied as foliar. The treatments were applied in a complete randomized design (CRD) with three replications. Results showed that 20 CoSO4 with proline showed improvement in shoot length (∼ 20%), root length (∼ 23%), plant dry weight (∼ 19%), and plant fresh weight (∼ 41%) compared to control. The significant increase in chlorophyll, physiological and biochemical attributes of radish plants compared to the control confirms the efficacy of 20 CoSO4 in conjunction with 10 mg/L proline for mitigating salinity stress. In conclusion, application of cobalt with proline can help to alleviate salinity stress in radish plants. However, multiple location experiments with various levels of cobalt and proline still needs in-depth investigations to validate the current findings.

Department of Agronomy Faculty of Agricultural Sciences and Technology Bahauddin Zakariya University Multan Punjab Pakistan

Department of Botany and Microbiology College of Science King Saud University PO Box 2455 Riyadh 11451 Saudi Arabia

Department of Botany Hindu College Moradabad Moradabad India

Department of Geology and Pedology Faculty of Forestry and Wood Technology Mendel University in Brno Zemedelska 1 Brno 61300 Czech Republic

Department of Soil Science Faculty of Agricultural Sciences and Technology Bahauddin Zakariya University Multan Punjab Pakistan

Zobrazit více v PubMed

Ali M. Assessment of maize genotypes for salt tolerance based on physiological indices. Pakistan J Bot. 2022;54:1613–1618.

Rafay M, Usman M. Soil salinity hinders plant growth and development and its remediation-A review. J Agric Res. 2023;61:189–200.

Khan RWA, Awan FS, Iqbal RK. Evaluation and identification of salt tolerant wheat through in vitro salinity induction in seeds. Pakistan J Bot. 2022;54:1987–93.

Munns R. Comparative physiology of salt and water stress. Plant Cell Environ. 2002;25:239–50. doi: 10.1046/j.0016-8025.2001.00808.x. PubMed DOI

Rasool S, Hameed A, Azooz MM, Siddiqi TO, Ahmad P. Ecophysiology and responses of plants under salt stress. New York, NY: Springer New York; 2013.

Dietz K-J, Turkan I, Krieger-Liszkay A. Redox-and reactive oxygen species-dependent signaling into and out of the photosynthesizing chloroplast. Plant Physiol. 2016;171:1541–50. doi: 10.1104/pp.16.00375. PubMed DOI PMC

Mansoor S, Ali Wani O, Lone JK, Manhas S, Kour N, Alam P, et al. Reactive oxygen species in plants: from source to sink. Antioxidants. 2022;11:225. doi: 10.3390/antiox11020225. PubMed DOI PMC

Foyer CH. Reactive oxygen species, oxidative signaling and the regulation of photosynthesis. Environ Exp Bot. 2018;154:134–42. doi: 10.1016/j.envexpbot.2018.05.003. PubMed DOI PMC

Foroumandi E, Nourani V, Kantoush SA. Investigating the main reasons for the tragedy of large saline lakes: Drought, climate change, or anthropogenic activities? A call to action. J Arid Environ. 2022;196:104652. doi: 10.1016/j.jaridenv.2021.104652. DOI

Vos R, Bellù LG. Global trends and challenges to Food and Agriculture into the 21st Century. Sustainable food and agriculture. Elsevier; 2019. pp. 11–30.

Ahammed GJ, Li X. Dopamine-induced abiotic stress tolerance in horticultural plants. Sci Hortic (Amsterdam) 2023;307:111506. doi: 10.1016/j.scienta.2022.111506. DOI

Hanaka A, Ozimek E, Reszczyńska E, Jaroszuk-Ściseł J, Stolarz M. Plant tolerance to drought stress in the presence of supporting bacteria and fungi: an efficient strategy in horticulture. Horticulturae. 2021;7:390. doi: 10.3390/horticulturae7100390. DOI

Drobek M, Frąc M, Cybulska J. Plant biostimulants: importance of the quality and yield of horticultural crops and the improvement of plant tolerance to abiotic stress-a review. Agronomy. 2019;9:335. doi: 10.3390/agronomy9060335. DOI

Ehrlich PR, Ehrlich AH, Daily GC. Food Security, Population and Environment. Popul Dev Rev. 1993;19:1–32. doi: 10.2307/2938383. DOI

Mukhopadhyay R, Sarkar B, Jat HS, Sharma PC, Bolan NS. Soil salinity under climate change: challenges for sustainable agriculture and food security. J Environ Manage. 2021;280:111736. doi: 10.1016/j.jenvman.2020.111736. PubMed DOI

Gad N. Interactive effect of salinity and cobalt on tomato plants. II. Some physiological parameters as affected by cobalt and salinity. Res J Agric Biol Sci. 2005;1:270–6.

Gad N, Hassan NM, Sayed S. Influence of Cobalt on tolerating climatic change (Salinity) in onion plant with reference to physiological and chemical approach. Plant Arch. 2020;20:1496–500.

Gad N, Abdel-Moez MR, Fekry Ali ME, Abou-Hussein SD. Increasing salt tolerance in cucumber by using cobalt. Middle East J Appl Sci. 2018;8:345–54.

Zahid A, ul din K, Ahmad M, Hayat U, Zulfiqar U, Askri SMH, et al. Exogenous application of sulfur-rich thiourea (STU) to alleviate the adverse effects of cobalt stress in wheat. BMC Plant Biol. 2024;24:126. doi: 10.1186/s12870-024-04795-1. PubMed DOI PMC

Ali Q, Zia MA, Kamran M, Shabaan M, Zulfiqar U, Ahmad M, et al. Nanoremediation for heavy metal contamination: a review. Hybrid Adv. 2023;4:100091. doi: 10.1016/j.hybadv.2023.100091. DOI

Hu X, Wei X, Ling J, Chen J. Cobalt: an essential micronutrient for Plant Growth? Front Plant Sci. 2021;12:768523. doi: 10.3389/fpls.2021.768523. PubMed DOI PMC

Hayat K, Bundschuh J, Jan F, Menhas S, Hayat S, Haq F, et al. Combating soil salinity with combining saline agriculture and phytomanagement with salt-accumulating plants. Crit Rev Environ Sci Technol. 2020;50:1085–115. doi: 10.1080/10643389.2019.1646087. DOI

Husen A. The harsh environment and resilient plants: an overview. In: Husen A, editor. Harsh environment and plant resilience. Cham: Springer International Publishing; 2021. pp. 1–23.

Thakur P, Nayyar H. Facing the cold stress by plants in the changing environment: sensing, signaling, and defending mechanisms. Plant acclimation to environmental stress. Springer; 2012. pp. 29–69.

Devi EL, Kumar S, Basanta Singh T, Sharma SK, Beemrote A, Devi CP, et al. Medicinal plants and Environmental challenges. Cham: Springer International Publishing; 2017. Adaptation strategies and defence mechanisms of plants during environmental stress; pp. 359–413.

Giordano M, Petropoulos SA, Rouphael Y. Response and defence mechanisms of vegetable crops against drought, heat and salinity stress. Agriculture. 2021;11:463. doi: 10.3390/agriculture11050463. DOI

Nahar K, Hasanuzzaman M, Fujita M. Roles of Osmolytes in Plant Adaptation to Drought and Salinity. In: Iqbal N, Nazar RA, Khan N, editors. Osmolytes and plants acclimation to changing Environment: emerging Omics technologies. New Delhi: Springer India; 2016. pp. 37–68.

Atta N, Shahbaz M, Farhat F, Maqsood MF, Zulfiqar U, Naz N, et al. Proline-mediated redox regulation in wheat for mitigating nickel-induced stress and soil decontamination. Sci Rep. 2024;14:456. doi: 10.1038/s41598-023-50576-5. PubMed DOI PMC

Mansour MMF, Salama KHA. Proline and Abiotic stresses: responses and adaptation. In: Hasanuzzaman M, editor. Plant Ecophysiology and Adaptation under Climate Change: mechanisms and perspectives II. Singapore: Springer Singapore; 2020. pp. 357–97.

Zulfiqar F, Ashraf M. Proline alleviates abiotic stress Induced oxidative stress in plants. J Plant Growth Regul. 2023;42:4629–51. doi: 10.1007/s00344-022-10839-3. DOI

Spormann S, Nadais P, Sousa F, Pinto M, Martins M, Sousa B, et al. Accumulation of proline in plants under contaminated soils—are we on the same page? Antioxidants. 2023;12:666. doi: 10.3390/antiox12030666. PubMed DOI PMC

Hosseinifard M, Stefaniak S, Ghorbani Javid M, Soltani E, Wojtyla Ł, Garnczarska M. Contribution of exogenous proline to abiotic stresses tolerance in plants: a review. Int J Mol Sci. 2022;23:5186. doi: 10.3390/ijms23095186. PubMed DOI PMC

Gill S, Tuteja N. Reactive oxygen species and antioxidant machinery in abiotic stress tolerance in crop plants. Plant Physiol Biochem. 2010;48:909–30. doi: 10.1016/j.plaphy.2010.08.016. PubMed DOI

Velikova V, Sharkey TD, Loreto F. Stabilization of thylakoid membranes in isoprene-emitting plants reduces formation of reactive oxygen species. Plant Signal Behav. 2012;7:139–41. doi: 10.4161/psb.7.1.18521. PubMed DOI PMC

El Moukhtari A, Cabassa-Hourton C, Farissi M, Savouré A. How does proline treatment promote salt stress tolerance during crop plant development? Front Plant Sci. 2020;11:1127. doi: 10.3389/fpls.2020.01127. PubMed DOI PMC

Mahboob W, Khan MA, Shirazi MU. Induction of salt tolerance in wheat (Triticum aestivum L.) seedlings through exogenous application of proline. Pak J Bot. 2016;48:861–7.

Soroori S, Danaee E, Hemmati K, Ladan Moghadam A. Effect of foliar application of proline on morphological and physiological traits of Calendula officinalis L. under drought stress. J Ornam Plants. 2021;11:13–30.

Hassan A, Fasiha Amjad S, Hamzah Saleem M, Yasmin H, Imran M, Riaz M, et al. Foliar application of ascorbic acid enhances salinity stress tolerance in barley (Hordeum vulgare L.) through modulation of morpho-physio-biochemical attributes, ions uptake, osmo-protectants and stress response genes expression. Saudi J Biol Sci. 2021;28:4276–90. doi: 10.1016/j.sjbs.2021.03.045. PubMed DOI PMC

Sheteiwy MS, Shao H, Qi W, Daly P, Sharma A, Shaghaleh H, et al. Seed priming and foliar application with jasmonic acid enhance salinity stress tolerance of soybean (Glycine max L.) seedlings. J Sci Food Agric. 2021;101:2027–41. doi: 10.1002/jsfa.10822. PubMed DOI

Sultan I, Khan I, Chattha MU, Hassan MU, Barbanti L, Calone R, et al. Improved salinity tolerance in early growth stage of maize through salicylic acid foliar application. Ital J Agron. 2021;16:1–11.

Ali MS, Zahid ZH, Siddike MN, Bappi ZH, Payel NA, Islam T, et al. Effect of different levels of organic fertilizer on growth, yield and economic benefits of radish (Raphanus sativus L) J Biosci Agric Res. 2023;30:2533–40. doi: 10.18801/jbar.300223.306. DOI

Gamba M, Asllanaj E, Raguindin PF, Glisic M, Franco OH, Minder B, et al. Nutritional and phytochemical characterization of radish (Raphanus sativus): a systematic review. Trends Food Sci Technol. 2021;113:205–18. doi: 10.1016/j.tifs.2021.04.045. DOI

Page AL, Miller RH, Keeny DR. Soil pH and lime requirement. In: Page AL, editor. Methods of Soil Analysis: Part 2 Chemical and Microbiological Properties, 9.2.2/Agronomy Monographs. 2nd edition. Madison: American Society of Agronomy, Inc. and Soil Science Society of America, Inc.; 1983. pp. 199–208.

Estefan G, Sommer R, Ryan J. Methods of Soil, Plant, and Water Analysis: A manual for the West Asia and North Africa region. 3rd edition. Beirut, Lebanon: International Center for Agricultural Research in the Dry Areas (ICARDA); 2013.

Rhoades JD, et al. Salinity: electrical conductivity and total dissolved solids. In: Sparks DL, Page AL, Helmke PA, Loeppert RH, Soltanpour PN, Tabatabai MA, et al., editors. Methods of Soil Analysis, Part 3, Chemical methods. Madison, WI, USA: Soil Science Society of America; 1996. pp. 417–35.

Nelson DW, Sommers LE, Total, Carbon . Organic Carbon, and Organic Matter. In: Page AL, editor. Methods of Soil Analysis: part 2 Chemical and Microbiological properties. Madison, WI, USA: American Society of Agronomy, Crop Science Society of America, and Soil Science Society of America; 1982. pp. 539–79.

Bremner M, et al. Nitrogen-total. In: Sumner DL, Sparks AL, Page PA, Helmke RH, Loeppert NP, Soltanpour AM, et al., editors. Methods of Soil Analysis Part 3. Madison, WI, USA: John Wiley & Sons, Inc; 1996. pp. 1085–121.

Kuo S, et al. Phosphorus. In: Sparks DL, Page AL, Helmke PA, Loeppert RH, Soltanpour PN, Tabatabai MA, et al., editors. Methods of Soil Analysis Part 3: Chemical methods. Madison, Wisconsin: John Wiley & Sons, Ltd: SSSA; 2018. pp. 869–919.

Pratt PF. Potassium. In: Norman AG, editor. Methods of Soil Analysis, Part 2: Chemical and Microbiological properties. Madison, WI, USA: John Wiley & Sons, Ltd; 2016. pp. 1022–30.

Donald AH, Hanson D. Determination of potassium and sodium by flame emmision spectrophotometery. In: Kalra Y, editor. Handbook of Reference Methods for Plant Analysis. 1st edition. Washington, D.C.: CRC Press; 1998. pp. 153–5.

Gee GW, Bauder JW. Particle-size Analysis. In: Klute A, editor. Methods of soil analysis. Part 1. Physical and mineralogical methods. 2nd edition. Madison, WI, USA: John Wiley & Sons, Inc.; 2018. pp. 383–411.

Boutraa T, Akhkha A, Al-Shoaibi AA, Alhejeli AM. Effect of water stress on growth and water use efficiency (WUE) of some wheat cultivars (Triticum durum) grown in Saudi Arabia. J Taibah Univ Sci. 2010;3:39–48. doi: 10.1016/S1658-3655(12)60019-3. DOI

Arnon DI. Copper enzymes in isolated chloroplasts. Polyphenoloxidse in beta vulgaris. Plant Physiol. 1949;24:1–15. doi: 10.1104/pp.24.1.1. PubMed DOI PMC

Durak I, Yurtarslanl Z, Canbolat O, Akyol Ö. A methodological approach to superoxide dismutase (SOD) activity assay based on inhibition of nitroblue tetrazolium (NBT) reduction. Clin Chim Acta. 1993;214:103–4. doi: 10.1016/0009-8981(93)90307-P. PubMed DOI

Cakmak I, Strbac D, Marschner H. Activities of hydrogen peroxide-scavenging enzymes in germinating wheat seeds. J Exp Bot. 1993;44:127–32. doi: 10.1093/jxb/44.1.127. DOI

Aebi H. Catalase in vitro. Methods Enzymol. 1984;105:121–6. doi: 10.1016/S0076-6879(84)05016-3. PubMed DOI

Nakano Y, Asada K. Hydrogen peroxide is scavenged by ascorbate-specific peroxidase in spinach chloroplasts. Plant Cell Physiol. 1981;22:867–80.

Hernández JA, Almansa MS. Short-term effects of salt stress on antioxidant systems and leaf water relations of pea leaves. Physiol Plant. 2002;115:251–7. doi: 10.1034/j.1399-3054.2002.1150211.x. PubMed DOI

Bates LS, Waldren RP, Teare ID. Rapid determination of free proline for water-stress studies. Plant Soil. 1973;39:205–7. doi: 10.1007/BF00018060. DOI

Griffith OW. Determination of glutathione and glutathione disulfide using glutathione reductase and 2-vinylpyridine. Anal Biochem. 1980;106:207–12. doi: 10.1016/0003-2697(80)90139-6. PubMed DOI

Carlberg I, Mannervik B. Glutathione reductase. Methods in Enzymology. Elsevier Inc.; 1985. pp. 484–90. PubMed

Kampfenkel K, Vanmontagu M, Inzé D. Extraction and determination of ascorbate and dehydroascorbate from plant tissue. Anal Biochem. 1995;225:165–7. doi: 10.1006/abio.1995.1127. PubMed DOI

Lutts S, Kinet JM, Bouharmont J. NaCl-induced Senescence in leaves of Rice (Oryza sativa L.) cultivars Differing in Salinity Resistance. Ann Bot. 1996;78:389–98. doi: 10.1006/anbo.1996.0134. DOI

Loka DA, Oosterhuis MD, Ritchie GL. Water stress and reproductive development in cotton. Stress Physiol Cott. 2011;7:37–72.

Barrs HD, Weatherley PE. A re-examination of the relative turgidity technique for estimating water deficits in leaves. Aust J Biol Sci. 1962;15:413–28. doi: 10.1071/BI9620413. DOI

Steel RG, Torrie JH, Dickey DA. Principles and Procedures of Statistics: A Biometrical Approach. 3rd edition. Singapore: McGraw Hill Book International Co.; 1997.

OriginLab Corporation . OriginPro. Northampton. MA, USA: OriginLab; 2021.

Atta K, Mondal S, Gorai S, Singh AP, Kumari A, Ghosh T, et al. Impacts of salinity stress on crop plants: improving salt tolerance through genetic and molecular dissection. Front Plant Sci. 2023;14:1241736. doi: 10.3389/fpls.2023.1241736. PubMed DOI PMC

Godfray HCJ, Garnett T. Food security and sustainable intensification. Philos Trans R Soc B Biol Sci. 2014;369:20120273. doi: 10.1098/rstb.2012.0273. PubMed DOI PMC

Munns R, James RA, Läuchli A. Approaches to increasing the salt tolerance of wheat and other cereals. In: J Exp Bot. 2006. p. 1025–43. PubMed

Abdelaal K, Alsubeie MS, Hafez Y, Emeran A, Moghanm F, Okasha S, et al. Physiological and biochemical changes in vegetable and field crops under drought, salinity and weeds stresses: control strategies and management. Agriculture. 2022;12:2084. doi: 10.3390/agriculture12122084. DOI

Khan I, Muhammad A, Chattha MU, Skalicky M, Bilal Chattha M, Ahsin Ayub M, et al. Mitigation of salinity-induced oxidative damage, growth, and yield reduction in fine rice by sugarcane press mud application. Front Plant Sci. 2022;13:840900. doi: 10.3389/fpls.2022.840900. PubMed DOI PMC

Fahad S, Hussain S, Matloob A, Khan FA, Khaliq A, Saud S, et al. Phytohormones and plant responses to salinity stress: a review. Plant Growth Regul. 2015;75:391–404. doi: 10.1007/s10725-014-0013-y. DOI

Yildrim E, Donmez MF, Turan M. Use of bioinoculants in ameliorative effects on radish plants under salinity stress. J Plant Nutr. 2008;31:2059–74. doi: 10.1080/01904160802446150. DOI

Arif Y, Singh P, Siddiqui H, Bajguz A, Hayat S. Salinity induced physiological and biochemical changes in plants: an omic approach towards salt stress tolerance. Plant Physiol Biochem. 2020;156:64–77. doi: 10.1016/j.plaphy.2020.08.042. PubMed DOI

Amin I, Rasool S, Mir MA, Wani W, Masoodi KZ, Ahmad P. Ion homeostasis for salinity tolerance in plants: a molecular approach. Physiol Plant. 2021;171:578–94. doi: 10.1111/ppl.13185. PubMed DOI

Singh M, Kumar J, Singh S, Singh VP, Prasad SM. Roles of osmoprotectants in improving salinity and drought tolerance in plants: a review. Rev Environ Sci Bio/Technology. 2015;14:407–26. doi: 10.1007/s11157-015-9372-8. DOI

Akeel A, Jahan A. Contaminants in Agriculture. Cham: Springer International Publishing; 2020. Role of cobalt in plants: its stress and alleviation; pp. 339–57.

Ali S, Gill RA, Mwamba TM, Zhang N, Lv MT, Ul Hassan Z, et al. Differential cobalt-induced effects on plant growth, ultrastructural modifications, and antioxidative response among four Brassica napus (L.) cultivars. Int J Environ Sci Technol. 2018;15:2685–700. doi: 10.1007/s13762-017-1629-z. DOI

Roychoudhury A, Chakraborty S. Cobalt and molybdenum: deficiency, toxicity, and nutritional role in plant growth and development. Plant nutrition and food security in the era of climate change. Elsevier; 2022. pp. 255–70.

Jahani M, Khavari-Nejad RA, Mahmoodzadeh H, Saadatmand S. Effects of foliar application of cobalt oxide nanoparticles on growth, photosynthetic pigments, oxidative indicators, non-enzymatic antioxidants and compatible osmolytes in canola (Brassica napus L). Acta Biol Cracov Ser Bot. 2019;61.

Brengi SH, Khedr AAEM, Abouelsaad IA. Effect of melatonin or cobalt on growth, yield and physiological responses of cucumber (Cucumis sativus L.) plants under salt stress. J Saudi Soc Agric Sci. 2022;21:51–60.

Tourky SMN, Shukry WM, Hossain MA, Siddiqui MH, Pessarakli M, Elghareeb EM. Cobalt enhanced the drought-stress tolerance of rice (Oryza sativa L.) by mitigating the oxidative damage and enhancing yield attributes. South Afr J Bot. 2023;159:191–207. doi: 10.1016/j.sajb.2023.05.035. DOI

Hasanuzzaman M, Raihan MRH, Masud AAC, Rahman K, Nowroz F, Rahman M, et al. Regulation of reactive oxygen species and antioxidant defense in plants under salinity. Int J Mol Sci. 2021;22:9326. doi: 10.3390/ijms22179326. PubMed DOI PMC

Sofo A, Scopa A, Nuzzaci M, Vitti A. Ascorbate peroxidase and catalase activities and their genetic regulation in plants subjected to drought and salinity stresses. Int J Mol Sci. 2015;16:13561–78. doi: 10.3390/ijms160613561. PubMed DOI PMC

Raj Rai S, Bhattacharyya C, Sarkar A, Chakraborty S, Sircar E, Dutta S, et al. Glutathione: role in oxidative/nitrosative stress, antioxidant defense, and treatments. ChemistrySelect. 2021;6:4566–90. doi: 10.1002/slct.202100773. DOI

Salam A, Afridi MS, Khan AR, Azhar W, Shuaiqi Y, Ulhassan Z et al. Cobalt induced toxicity and tolerance in plants: insights from omics approaches. Heavy Met Toxic Toler Plants Biol Omi Genet Eng Approach. 2023;:207–29.

Gómez-Merino FC, Trejo-Téllez LI. The role of beneficial elements in triggering adaptive responses to environmental stressors and improving plant performance. Biot Abiotic Stress Toler Plants. 2018;:137–72.

Isah T. Stress and defense responses in plant secondary metabolites production. Biol Res. 2019;52:39. doi: 10.1186/s40659-019-0246-3. PubMed DOI PMC

Siddique A, Kandpal G, Kumar P. Proline accumulation and its defensive role under diverse stress condition in plants: an overview. J Pure Appl Microbiol. 2018;12:1655–9. doi: 10.22207/JPAM.12.3.73. DOI

Mansour MMF, Ali EF. Evaluation of proline functions in saline conditions. Phytochemistry. 2017;140:52–68. doi: 10.1016/j.phytochem.2017.04.016. PubMed DOI

Hayat S, Hayat Q, Alyemeni MN, Wani AS, Pichtel J, Ahmad A. Role of proline under changing environments: a review. Plant Signal Behav. 2012;7:1456–66. doi: 10.4161/psb.21949. PubMed DOI PMC

Khanna-Chopra R, Semwal VK, Lakra N, Pareek A. Proline–A key regulator conferring plant tolerance to salinity and drought. Plant tolerance to environmental stress. CRC; 2019. pp. 59–80.

Hossain MA, Hoque MA, Burritt DJ, Fujita M. Proline protects plants against abiotic oxidative stress: biochemical and molecular mechanisms. Oxidative damage to plants. Elsevier; 2014. pp. 477–522.

Carillo P, Annunziata MG, Pontecorvo G, Fuggi A, Woodrow P. Others. Salinity stress and salt tolerance. Abiotic Stress plants-mechanisms Adapt. 2011;1:21–38.

Yaqoob H, Akram NA, Iftikhar S, Ashraf M, Khalid N, Sadiq M, et al. Seed pretreatment and foliar application of proline regulate morphological, physio-biochemical processes and activity of antioxidant enzymes in plants of two cultivars of Quinoa (Chenopodium quinoa Willd) Plants. 2019;8:588. doi: 10.3390/plants8120588. PubMed DOI PMC

Leiva-Ampuero A, Agurto M, Matus JT, Hoppe G, Huidobro C, Inostroza-Blancheteau C, et al. Salinity impairs photosynthetic capacity and enhances carotenoid-related gene expression and biosynthesis in tomato (Solanum lycopersicum L. Cv. Micro-tom) PeerJ. 2020;8:e9742. doi: 10.7717/peerj.9742. PubMed DOI PMC

Zahedi SM, Abolhassani M, Hadian-Deljou M, Feyzi H, Akbari A, Rasouli F, et al. Proline-functionalized graphene oxide nanoparticles (GO-pro NPs): a new engineered nanoparticle to ameliorate salinity stress on grape (Vitis vinifera l. Cv sultana) Plant Stress. 2023;7:100128. doi: 10.1016/j.stress.2022.100128. DOI

Irshad I, Anwar-Ul-Haq M, Akhtar J, Maqsood M. Enhancing maize growth and mitigating salinity stress through foliar application of proline and glycine betaine. Pakistan J Bot. 2024;56:9–17.

Semida WM, Abdelkhalik A, Rady MOA, Marey RA, Abd El-Mageed TA. Exogenously applied proline enhances growth and productivity of drought stressed onion by improving photosynthetic efficiency, water use efficiency and up-regulating osmoprotectants. Sci Hortic (Amsterdam) 2020;272:109580. doi: 10.1016/j.scienta.2020.109580. DOI

Shafi A, Zahoor I, Mushtaq U. Proline accumulation and oxidative stress: diverse roles and mechanism of tolerance and adaptation under salinity stress. Salt stress, microbes, and plant interactions: mechanisms and molecular approaches. Springer; 2019. pp. 269–300.

Rohman MM, Begum S, Akhi AH, Ahsan A, Uddin MS, Amiruzzaman M, et al. Protective role of antioxidants in maize seedlings under saline stress: exogenous proline provided better tolerance than betaine. Bothalia J. 2015;45:17–35.

Hussain RA, Anjum S, Khalid MA, Saqib MF, Zakir M. Plant abiotic stress tolerance. Cham: Springer International Publishing; 2019. Oxidative stress and antioxidant defense mechanisms in plants under salt stress; pp. 191–205.

Liang X, Zhang L, Natarajan SK, Becker DF. Proline mechanisms of stress survival. Antioxid Redox Signal. 2013;19:998–1011. doi: 10.1089/ars.2012.5074. PubMed DOI PMC

Khalid M, Rehman HM, Ahmed N, Nawaz S, Saleem F, Ahmad S, et al. Using exogenous melatonin, glutathione, proline, and glycine betaine treatments to combat abiotic stresses in crops. Int J Mol Sci. 2022;23:12913. doi: 10.3390/ijms232112913. PubMed DOI PMC