Mitigating Drought Stress in Physalis alkekengi: Synergistic Effects of Foliar Boron and Zinc on Photosynthesis and Metabolite Accumulation

Document Type : Research Paper

Authors

1 Faculty of Basic Science Branch, Faculty of Dentistry, University of Al-Qadisiyah

2 Department of Horticultural Sciences, Faculty of Agriculture & Natural Resources, Ardakan University, Ardakan, Iran

10.22034/jmpb.2026.371603.2114

Abstract

This study investigated the effects of foliar boron (B) and zinc (Zn) on Physalis alkekengi under four levels of drought stress (0%, 25%, 50%, and 75% of field capacity). Plants were treated with B at 0, 100, 200, and 400 ml/L and Zn at 0, 1, 2, and 3 g/L. Drought stress significantly reduced fruit length (3.5–5.2 cm), weight (13.7–22.8 g), and firmness (1.7–3.5 N). The combined foliar application of B (200 ml/L) and Zn (2 g/L) under moderate drought (50% field capacity) produced the best outcomes. Fruit weight increased by 28%, total soluble solids (TSS) reached 11.6%, chlorophyll content rose to 2.1 mg/g FW, and photosystem II efficiency (Fv/Fm) improved to 0.78. Total phenolics (7.2 mg/g FW), flavonoids (32.3 mg/g FW), and proline (38.5 mg/g FW) also peaked under this treatment, indicating enhanced antioxidant metabolism and osmotic adjustment. Strong correlations were observed between proline and total phenolics (r = 0.90), and between flavonoids and total phenolics (r = 0.86), suggesting coordinated stress-response mechanisms. Principal component analysis (PCA) revealed clear clustering of treatments, separating those with higher fruit quality and metabolite accumulation from treatments with improved photosynthetic efficiency. Overall, foliar B and Zn acted synergistically to alleviate drought stress, stabilize photosynthesis, enhance antioxidant defenses, and improve fruit growth and quality in P. alkekengi. These findings demonstrate that integrated micronutrient management can be an effective strategy to improve plant performance under water-limited conditions, benefiting both productivity and the nutritional value of fruits.

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  1. Farooq M., Wahid A., Kobayashi N., Fjita D., Basra S.M.A. Plant drought stress: Effects, mechanisms and management. Agronomy for Sustainable Development. 2009; 29(1): 185-212. https://doi.org/10.1007/978-90-481-2666-8_12
  2. Qiao M., Hong C., Jiao Y., Hou S., Gao H. Impacts of drought on photosynthesis in major food crops and the related mechanisms of plant responses to drought. Plants. 2024; 13(13): 1808. https://doi.org/10.3390/plants13131808
  3. Szabados L., Savouré A. Proline: a multifunctional amino acid. Trends in Plant Science. 2010; 15(2): 89-97. https://doi.org/10.1016/j.tplants.2009.11.009
  4. Arnon A.N. Method of extraction of chlorophyll in the plants. Agronomy Journal. 1967; 23(1): 112-21.
  5. Bates L.S., Waldren R.P., Teare I.D. Rapid determination of free proline for water-stress studies. Plant and Soil. 1973; 39(1): 205-7. https://doi.org/10.1007/BF00018060
  6. Chirinos R., Rogez H., Campos D., Pedreschi R., Larondelle Y. Optimization of extraction conditions of antioxidant phenolic compounds from mashua (Tropaeolum tuberosum Ruíz & Pavón) tubers. Separation and Purification Technology. 2007; 55(2): 217-25. https://doi.org/10.1016/j.seppur.2006.12.005
  7. Adak N., Gubbuk H., Tetik N. Yield, quality and biochemical properties of various strawberry cultivars under water stress. Journal of the Science of Food and Agriculture. 2018; 98(1): 304-11. https://doi.org/10.1002/jsfa.8471
  8. Semida W.M., Abdelkhalik A., Mohamed G.F., Abd El-Mageed T.A., Abd El-Mageed S.A., Rady M.M., Ali E.F. Foliar application of zinc oxide nanoparticles promotes drought stress tolerance in eggplant (Solanum melongena L.). Plants. 2021; 10(2): 421. https://doi.org/10.3390/plants10020421
  9. Singh R., Saffeullah P., Umar S., Abass S., Ahmad S., Iqbal N. Comparing the individual and combined effects of nano zinc and conventional zinc fertilization on growth, yield, phytochemical properties, antioxidant activity, and secoisolariciresinol diglucoside content in linseed. Plant Nano Biology. 2024; 10: 100098. https://doi.org/10.1016/j.plana.2024.100098
  10. Hanif S., Javed R., Cheema M., Kiani M.Z., Farooq S., Zia M. Harnessing the potential of zinc oxide nanoparticles and their derivatives as nanofertilizers: trends and perspectives. Plant Nano Biology. 2024; 10: 100110. https://doi.org/10.1016/j.plana.2024.100110
  11. Khatoon F., Kundu M., Mir H., Nahakpam S. Efficacy of foliar feeding of brassinosteroid to improve growth, yield and fruit quality of strawberry (Fragaria× ananassa Duch.) grown under subtropical plain. Communications in Soil Science and Plant Analysis. 2021; 52(8): 803-14. https://doi.org/10.1080/00103624.2020.1869765
  12. Li X., Ahammed G.J., Li Z.X., Zhang L., Wei J.P., Shen C., Yan P., Zhang L.P., Han W.Y. Brassinosteroids improve quality of summer tea (Camellia sinensis L.) by balancing biosynthesis of polyphenols and amino acids. Frontiers in Plant Science. 2016; 7: 1304. https://doi.org/10.3389/fpls.2016.01304
  13. Martinez-Perez M.E., Ruiz-Anchondo T.D., Jacobo-Cuellar J.L., Calderón-Zavala G. Responses to foliar sprays of strawberry variety ‘Portola’to biostimulants on growth, yield, quality, and bioactive compounds. Notulae Botanicae Horti Agrobotanici Cluj-Napoca. 2024; 52(3): 13513. https://doi.org/10.15835/nbha52313513
  14. Gulen H., Kesici M., Cetinkaya C., Ergin S. Proline and antioxidant enzyme activities in some strawberry cultivars under drought and recovery. Notulae Botanicae Horti Agrobotanici Cluj-Napoca. 2018; 46(2): 570-8. https://doi.org/10.15835/nbha46211077
  15. Luo Y., Wang X.R., Zhang Y., Liu Z.J., Ren Y.N., Yang T., Tang H.R. Effect of extraction conditions on antioxidant components and antioxidant activity of strawberry fruit extracts. Food Science; 2011. 10(6): 124-136.
  16. Liu A., Xiao W., Lai W., Wang J., Li X., Yu H., Zha Y. Potential application of selenium and copper nanoparticles in improving growth, quality, and physiological characteristics of strawberry under drought stress. Agriculture. 2024; 14(7): 1172. https://doi.org/10.3390/agriculture14071172
  17. Renzetti M., Funck D., Trovato M. Proline and ROS: a unified mechanism in plant development and stress response? Plants. 2024; 14(1): 2. https://doi.org/10.3390/plants14010002
  18. Gall H.L., Philippe F., Domon J.M., Gillet F., Pelloux J., Rayon C. Cell wall metabolism in response to abiotic stress. Plants. 2015; 4(1): 112. https://doi.org/10.3390/plants4010112
  19. Marschner H. Marschner's mineral nutrition of higher plants (3rd ed.). Academic Press; 2011.
  20. Pejam F., Ardebili Z.O., Ladan-Moghadam A., Danaee E. Zinc oxide nanoparticles mediated substantial physiological and molecular changes in tomato. PloS One. 2021; 16(3): e0248778. https://doi.org/10.1371/journal.pone.0248778
  21. Silva F.L., Vieira L.G., Ribas A.F., Moro A.L., Neris D.M., Pacheco A.C. Proline accumulation induces the production of total phenolics in transgenic tobacco plants under water deficit without increasing the G6PDH activity. Theoretical and Experimental Plant Physiology. 2018; 30(3): 251-60. https://doi.org/10.1007/s40626-018-0119-0
  22. Wang X.P., Li Q.Q., Pei Z.M., Wang S.C. Effects of zinc oxide nanoparticles on the growth, photosynthetic traits, and antioxidative enzymes in tomato plants. Biologia Plantarum. 2018; 62(4): 801-8. https://doi.org/10.1007/s10535-018-0813-4
  23. Furio R.N., Salazar S.M., Mariotti-Martínez J.A., Martínez-Zamora G.M., Coll Y., Díaz-Ricci J.C. Brassinosteroid applications enhance the tolerance to abiotic stresses, production and quality of strawberry fruits. Horticulturae. 2022; 8(7): 572. https://doi.org/10.3390/horticulturae8070572.
  24. Nakhaei F., Foliar Nutrient Applications to Barberry (Berberis vulgaris). II: Effects on Leaf Nutrient Content and Physico-Chemical Characteristics of Fruit and Yield. Journal of Medicinal plants and By-products, 2023; 12(4): 357-363. doi: 10.22092/jmpb.2022.354742.1364.
  25. Semsarzadeh M., Sayfzadeh S., Darvishi Zeidabadi D., Zakerin H., Hadidi Masuleh, the Role of Zeolite in Mitigating Physiological and Biochemical Stress Responses and Enhancing Seed Yield of Coriander (Coriandrum sativum L.) under Drought Conditions. Journal of Medicinal plants and By-products, 2026; 15(2): 188-195. doi: 10.22034/jmpb.2025.369791.1996.
  26. Salehpour S., Mojaradi M., Heydari Sadegh B., Poormasoumi H., Ahmadi T., Dahmardeh N., Aghayari M., Sanjari S., Saravani K. Study on Green Zinc Nanoparticles from Plant Extract and Antimicrobial Effects on Pathogen Urease-producing Bacteria. Journal of Medicinal plants and By-products, 2026; 15(2): 232-235. doi: 10.22034/jmpb.2025.369677.1982.
  27. Olad M., Filizadeh Y., Roodpeyma M. Influence of Foliar Applications of Micronutrients and Biofertilizers on Plant Growth and Fruit Yield of the Sponge Gourd [Luffa cylindrica (L.) M.Roem.]. Journal of Medicinal plants and By-products, 2025; 14(5): 435-440. doi: 10.22034/jmpb.2025.366827.1749