In pot experiment with bread winter wheat varieties Dostatok, Astarta, Malynivka, Natalka, Kuyalnik, Kyivska ostysta there were created different levels of mineral nutrition: 1) high level (N160P160K160 active substances per 1 kg of soil); 2) high level with foliar treatment by 5 % carbamide solution at the end of anthesis (BBCH 69); 3) low mineral nutrition level (N32P32K32); 4) low level with foliar treatment with carbamide. It was shown that functio ning of the photosynthetic apparatus is positively influenced by a high level of nitrogen nutrition and foliar treatment with carbamide. At the milk ripeness stage (BBCH 75), it was found that photosynthetic rate in leaves of individual varieties closely positively correlated with stomatal conductance. At the same time, the varietal specificity of the correlation dependences indicates genotypic differences in the response of the photosynthetic apparatus of plants of different varieties to the conditions of nitrogen nutrition. Water use efficiency (WUE) varied in a fairly wide range — from 4.14 in plants of the Natalka variety to 7.55 mmol CO2/mmol H2O in the Malynivka variety. In general, on a high level of mineral nutrition, WUE was higher than on a low level, and the photosynthetic nitrogen use efficiency (PNUE), on the contrary, was lower. The last index on a high level varied in the range of 76.9—.5, and on a low level — 115.8—262.8 mmol СО2/(mol N · s) depending on the variety and foliar treatment with carbamide. This treatment increased the PNUE of leaves at both high and low nutrition levels, in the latter case the effect was more pronounced due to a relatively greater increase in the photosynthetic rate. Plants of high-yielding varieties on average were characterized by higher PNUE compared to high-protein varieties. A close correlation was found between the photosynthetic rate and PNUE both on a high (r = 0.89) and low (r = 0.91) level of mineral nutrition. The revealed high genotypic variability in winter wheat plants under different conditions of mineral nutrition such important physiological parameters as the water and nitrogen use efficiency during photosynthesis suggests the prospect of improving these traits by genetic means in order to optimize the use of water and mineral resources in the wheat production process.
Keywords: Triticum aestivum L., winter wheat, mineral nutrition, photosynthesis, stomatal conductance, transpiration, nitrogen, photosynthetic nitrogen use efficiency, water use efficiency
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1. Morgun, V.V. & Rybalka, O.I. (2017). A strategy for the genetic improvement of cereals in order to ensure food security, medical and preventive nutrition and the needs of the processing industry. Visn. NAN Ukrainy, No. 3, pp. 54-64 [in Ukrainian].
2. Morgun, V.V. & Kiriziy, D.A. (2012). Prospects and modern strategies for improving the physiological traits of wheat to increase productivity. Fiziol. i biokhimiya kult. rast., 44, No. 6, pp. 463-483 [in Ukrainian].
3. Kiriziy, D.A., Stasik, O.O., Pryadkina, H.O. & Shadchyna, T.M. (2014). Photosynthesis. V. 2. CO2 assimilation and mechanisms of its regulation. Kyiv: Logos.
4. Busch, F.A., Ainsworth, E.A., Amtmann, A., Cavanagh, A.P., Driever, S.M., Ferguson, J.N., Kromdijk, J., Lawson, T., Leakey, A.D.B., Matthews, J.S.A., Meacham-Hensold, K., Vath, R.L., Vialet-Chabrand, S., Walker, B.J. & Papanatsiou, M. (2024). A guide to photosynthetic gas exchange measurements: Fundamental principles, best practice and potential pitfalls. Plant Cell Environ., 47, No 9, pp. 3344-3364. https://doi.org/10.1111/pce.14815
5. Arab, M.M., Marrano, A., Abdollahi-Arpanahi, R., Leslie, C.A., Cheng, H., Neale, D.B. & Vahdati, K. (2020). Combining phenotype, genotype, and environment to uncover genetic components underlying water use efficiency in Persian walnut. J. Exp. Bot., 71, No. 3, pp. 1107-1127. https://doi.org/10.1093/jxb/erz467
6. Condon, A.G. (2020). Drying times: plant traits to improve crop water use efficiency and yield. J. Exp. Bot., 71, No. 7, pp. 2239-2252. https://doi.org/10.1093/jxb/eraa002
7. Kang, J., Hao, X., Zhou, H. & Ding, S. (2021). An integrated strategy for improving water use efficiency by understanding physiological mechanisms of crops responding to water deficit: Present and prospect. Agric. Water Manag., 255, 107008. https://doi.org/10.1016/j.agwat.2021.107008
8. Wientjes, E. & Seijger, C. (2024). Less water in agriculture? Potential and challenges in optimizing water use efficiency. J. Exp. Bot., 75, No 13, pp. 3754-3757. https://doi.org/10.1093/jxb/erae227
9. Vadez, V., Kholova, J., Medina, S., Kakkera, A. & Anderberg, H. (2014). Transpiration efficiency: new insights into an old story. J. Exp. Bot., 65, No. 21, pp. 6141-6153. https://doi.org/10.1093/jxb/eru040
10. Gobu, R., Dash, G.K., Lal, J.P., Swain, P., Mahender, A., Anandan, A. & Ali, J. (2022). Unlocking the nexus between leaf-level water use efficiency and root traits together with gas exchange measurements in rice (Oryza sativa L.). Plants, 11, No. 9, 1270. https://doi.org/10.3390/plants11091270
11. Bertolino, L.T., Caine, R.S. & Gray, J.E. (2019). Impact of stomatal density and morphology on water-use efficiency in a changing world. Front. Plant Sci., 10, 225. https://doi.org/10.3389/fpls.2019.00225
12. Kimura, H., Hashimoto-Sugimoto, M., Iba, K., Terashima, I. & Yamori, W. (2020). Improved stomatal opening enhances photosynthetic rate and biomass production in fluctuating light. J. Exp. Bot., 71, pp. 2339-2350. https://doi.org/10.1093/jxb/eraa090
13. Caine, R., Yin, X., Sloan, J., Harrison, E.L., Mohammed, U., Fulton, T., Biswal, A.K., Dionora, J., Chater, C.C., Coe, R.A., Bandyopadhyay, A., Murchie, E.H., Swarup, R., Quick, W.P. & Gray, J.E. (2018). Rice with reduced stomatal density conserves water and has improved drought tolerance under future climate conditions. New Phytol. 221, pp. 371-384. https://doi.org/10.1111/nph.15344
14. Stacey, J., Betts, R., Hartley, A., Mercado, L. & Gedney, N. (2025). Future global water scarcity partially alleviated by vegetation responses to atmospheric CO2 and climate change [preprint]. EGUsphere. https://doi.org/10.5194/egusphere-2025-51
15. Flexas, J., Niinemets, U., Galle, A., Barbour, M.M., Centritto, M., Dias-Espejo, A., Douthe, C., Galmes, J., Ribas-Carbo, M., Rodrigues, P., Rossello, F., Soolana yakanahally, R., Tomas, M., Wright, I.J., Farquhar, G.D. & Medrano, H. (2013). Diffusional conductances to CO2 as a target for increasing photosynthetis and photosynthetic water-use efficiency. Photosynth. Res., 117, pp. 45-49. https://doi.org/10.1007/s11120-013-9844-z
16. Yan, S., Wu, Y., Fan, J., Zhang, F., U, K.T.P., Zheng, J., Qiang, S., Guo, J., Zou, H., Xiang, Y. & Wu, L. (2020). A sustainable strategy of managing irrigation based on water productivity and residual soil nitrate in a no-tillage maize system. J. Clean. Prod., 262, 121279. https://doi.org/10.1016/j.jclepro.2020.121279
17. Yan, F., Zhang, F., Fan, X., Fan, J., Wang, Y., Zou, H., Wang, H. & Li, G. (2021). Determining irrigation amount and fertilization rate to simultaneously optimize grain yield, grain nitrogen accumulation and economic benefit of drip-fertigated spring maize in northwest China. Agric. Water Manag., 243, 106440. https://doi.org/10.1016/j.agwat.2020.106440
18. Bardhan, K., York, L.M., Hasanuzzaman, M., Parekh, V., Jena, S. & Pandya, M.N. (2021). Can smart nutrient applications optimize the plant's hidden half to improve drought resistance? Physiol. Plant., 172, No. 2, pp. 1007-1015. https://doi.org/10.1111/ppl.13332
19. Bilotto, F., Harrison, M.T., Migliorati, M.D.A., Christie, K.M., Rowlings, D.W., Grace, P.R., Smith, A.P., Rawnsley, R.P., Thornburn, P.J. & Eckard, R.J. (2021). Can seasonal soil N mineralisation trends be leveraged to enhance pasture growth? Sci. Total Environ., 772, 145031. https://doi.org/10.1016/j.scitotenv.2021.145031
20. Turc, B., Sahay, S., Haupt, J., de Oliveira Santos, T., Bai, G. & Glowacka, K. (2024). Up-regulation of non-photochemical quenching improves water use efficiency and reduces whole-plant water consumption under drought in Nicotiana tabacum. J. Exp. Bot., 75, pp. 3959-3972. https://doi.org/10.1093/jxb/erae113
21. Li, K., Liu, D.-N., Li, L.-Y., Gao, Y., Gao, W.-J., Chen, B.-W., Luo, F. & Yao, Y. (2025). Effects of nitrogen application amount on nitrogen distribution and photosynthesis in tea leaves. Front. Plant Sci., 16:1575317. https://doi.org/10.3389/fpls.2025.1575317
22. Archontoulis, S.V., Yin, X., Vos, J., Danalatos, N.G. & Struik, P.C. (2012). Leaf photosynthesis and respiration of three bioenergy crops in relation to temperature and leaf nitrogen: how conserved are biochemical model parameters among crop species? J. Exp. Bot., 63, No. 2, pp. 895-911. https://doi.org/10.1093/jxb/err321
23. Wang, Q., Li, S., Li, J. & Huang, D. (2024). The utilization and roles of nitrogen in plants. Forests, 15, No 7, 1191. https://doi.org/10.3390/f15071191
24. Wei, X., Han, L., Xu, N., Sun, M. & Yang, X. (2024). Nitrate nitrogen enhances the efficiency of photoprotection in Leymus chinensis under drought stress. Front. Plant Sci., 15:1348925. https://doi.org/10.3389/fpls.2024.1348925
25. Lei, Z.Y., Wang, H., Wright, I.J., Zhu, X.G., Niinemets, Ґ., Li, Z.L., Sun, D.S., Dong, N., Zhang, W.F., Zhou, Z.L., Liu, F. & Zhang, Y.Li. (2021). Enhanced photosynthetic nitrogen use efficiency and increased nitrogen allocation to photosynthetic machinery under cotton domestication. Photosynth. Res., 150, pp. 239-250. https://doi.org/10.1007/s11120-021-00872-w
26. Nasar, J., Wang, G.-Y., Ahmad, S., Muhammad, I., Zeeshan, M., Gitari, H., Adnan, M., Fahad, S., Khalid, M.H.B., Zhou, X.-B., Abdelsalam, N.R., Ahmed, G.A. & Hasan, M.E. (2022). Nitrogen fertilization coupled with iron foliar application improves the photosynthetic characteristics, photosynthetic nitrogen use efficiency, and the related enzymes of maize crops under different planting patterns. Front. Plant Sci., 13, 988055. https://doi.org/10.3389/fpls.2022.988055
27. Kiriziy, D.A. (2013). Nitrogen use efficiency during photosynthetic CO2 assimilation in wheat leaves. Fiziol. rosl. genet., 45, No. 4, pp. 296-305 [in Russian].
28. Stasik, O.O., Kiriziy, D.A. & Pryadkina, H.O. (2016). Photosynthesis and productivity of agricultural plants. Fiziol. rosl. genet., 48, No. 3, pp. 232-251 [in Russian]. https://doi.org/10.15407/frg2016.03.232
29. Zhang, Y., Wang, J., Gong, S., Xu, D. & Sui, J. (2017). Nitrogen fertigation effect on photosynthesis, grain yield and water use efficiency of winter wheat. Agric. Water Manag., 179, pp. 277-287. https://doi.org/10.1016/j.agwat.2016.08.007
30. Ye, M., Peng, S.B. & Li, Y. (2019). Intraspecific variation in photosynthetic nitrogen-use efficiency is positively related to photosynthetic rate in rice (Oryza sativa L.) plants. Photosynth., 57, No. 1, pp. 311-319. https://doi.org/10.32615/ps.2019.011
31. Hong, M.J. & Kim, D.Y. (2025). Recent advances in nitrogen use efficiency (NUE) research in wheat. Korean J. Breed. Sci., 57, No 3, pp. 251-270. https://doi.org/10.9787/KJBS.2025.57.3.251
32. Lal, S.K., Gaggar, P., Kumar, S., Mallikarjuna, M.G., Vishwakarma, C., Rakshit, S., Pandey, A., Achary, V.M.M. & Mehta, S. (2024). Recent Advancements in Nitrogen Use Efficiency in Crop Plants Achieved by Genomics and Targeted Genetic Engineering Approaches. Plant Mol. Biol. Rep., 42, pp. 435-449. https://doi.org/10.1007/s11105-024-01439-4
33. Ali, A., Jabeen, N., Farruhbek, R., Chachar, Z., Laghari, A.A., Chachar, S., Ahmed, N., Ahmed, S. & Yang, Z. (2025). Enhancing nitrogen use efficiency in agriculture by integrating agronomic practices and genetic advances. Front. Plant Sci., 16: 1543714. https://doi.org/10.3389/fpls.2025.1543714
34. Ren, W., Li, X., Liu, T., Chen, N., Xin, M., Qi, Q. & Liu, B. (2025). Controlled-release fertilizer affects leaf nitrogen allocation and photosynthesis to improve nitrogen use efficiency and yield in the sunflower field. Front. Plant Sci., 16: 1622766. https://doi.org/10.3389/fpls.2025.1622766
35. Zhang, Z., Zhang, Y., Shi, Y. & Yu, Z. (2020). Optimized split nitrogen fertilizer increase photosynthesis, grain yield, nitrogen use efficiency and water use efficiency under water-saving irrigation. Sci. Rep., 10: 20310. https://doi.org/10.1038/s41598-020-75388-9
36. Laisk, A. & Oja, V. (1998). Dynamics of leaf photosynthesis: rapid response measurements and their interpretations. Collingwood: CSIRO Publishing. https://doi.org/10.1071/9780643105294
37. Sheheda, I.M., Pochynok, V.M., Kiriziy, D.A. & Mamenko, T.P. (2018). Effect of nitrogen fertilization conditions on photosynthesis, productivity and protein content of winter wheat grain. Fiziol. rosl. genet., 50, No. 2, pp. 105-114 [in Ukrainian]. https://doi.org/10.15407/frg2018.02.105
38. Endres, L., Silva, J.V., Ferreira, V.M. & De Souza Barbosa, G.V. (2010). Photosynthesis and water relations in brazilian sugarcane. Open Agric. J., No. 4, pp. 31-37. https://doi.org/10.2174/1874331501004010031
39. Li, S., Xie, Y., Liu, G., Wang, J., Lin, H., Xin, Y. & Zhai, J. (2020). Water Use effciency of soybean under water stress in different eroded soils. Water, 12, 373. https://doi.org/10.3390/w12020373
40. Lopez, M.A., Xavier, A. & Rainey, K.M. (2019). Phenotypic variation and genetic architecture for photosynthesis and water use efficiency in soybean (Glycine max L. Merr). Front. Plant Sci., 10, 680. https://doi.org/10.3389/fpls.2019.00680
41. Kiriziy, D.A. & Sheheda, I.M. (2019). Distribution of nitrogen in the donor-acceptor system of plants and its role in the production process. Fiziol. rosl. genet., 51, No. 2, pp. 114-132 [in Ukrainian]. https://doi.org/10.15407/frg2019.02.114
42. Tian, Z., Chai, H., Guo, H., Lu, Y., Yang, S., Liu, X., Jiang, D., Cao, W. & Dai, T. (2024). Genetic improvement of photosynthetic nitrogen use efficiency of winter wheat in the Yangtze River Basin of China. Field Crops Res., 305, 109199. https://doi.org/10.1016/j.fcr.2023.109199
43. Feller, U., Anders, I. & Mae, T. (2008). Rubiscolytics: fate of Rubisco after its enzymatic function in a cell is terminated. J. Exp. Bot., 59, No 7, pp. 1615-1624. https://doi.org/10.1093/jxb/erm242