The review, based on literature data and the authors’ own research, analyzes the issue of relationship between photosynthetic activity and nitrogen distribution in plant organs in connection to the problems of grain yield and quality formation. Special attention is paid to winter wheat, one of the most widespread and important agricultural crops in the world. The role of nitrogen as a key element that is part of many organic compounds necessary for plant life is considered. The influence of nitrogen on the development of root system and leaf surface, and its role in optimizing water use efficiency are shown. The relationship between nitrogen content and photosynthesis, the role of leaves in the formation of grain yield, and the Rubisco enzyme as a nitrogen depot, as well as the process of redistribution of this element to the grain, are highlighted. It is demonstrated that as the nitrogen level increases, its distribution into the carbon assimilation system first increases and then decreases, while the distribution into the deposition system gradually increases. Foliar fertilization contributes to the formation of grain quality, an increase in its protein content and yield. This measure increases the photosynthetic nitrogen use efficiency (PNUE) and has a positive effect on the water regime of plants. It is emphasized that the issue of the formation of wheat grain productivity and quality has two components: genotypic features of photosynthetic activity and the associated processes of nitrogen remobilization and accumulation in grain, as well as the influence of the main and foliar application of nitrogen-containing fertilizers on these processes. Researchers are focused on finding ways to combine these components to mitigate the contradiction between grain productivity and protein content.
Keywords: wheat, nitrogen, photosynthesis, remobilization, productivity, quality
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1. Zhong, Y., Wang, W., Huang, X., Liu, M., Hebelstrup, K.H., Yang, D., Cai, J., Wang, X., Zhou, Q., Cao, W., Dai, T. & Jiang, D. (2019). Nitrogen topdressing timing modifies the gluten quality and grain hardness related protein levels as revealed by iTRAQ. Food Chem, 277, pp. 135-144. https://doi.org/10.1016/j.foodchem.2018.10.071
2. FAOSTAT (2022). Food and Agricultural Organization of the United Nations. FAOSTAT Food Balance Sheets. Retrieved from: https://www.fao.org/faostat/en/#data/FBS
3. Sandetska, N.V. & Dubrovna, O.V. (2025). Current state of research on wheat grain quality. FTzTol. rosl. genet., 57, No. 3, pp. 187-222 [in Ukrainian]. https://doi.org/10.15407/frg2025.03.187
4. Wang, S., Zhang, X., Liu, K., Fei, P., Chen, J., Li, X., Ning, P., Chen, Y., Shi, J. & Tian, X. (2019). Improving zinc concentration and bioavailability of wheat grain through combined foliar application of zinc and pesticides. Agron. J., 111, pp. 1478-1487. https://doi.org/10.2134/agronj2018.09.0597
5. Stankevych, G., Kats, A. & Vasyliev, S. (2018). Investigation of hygroscopic properties of the spelt grain. Technol. Audit Prod. Reserves, 43, No. 5/3, pp. 37-41. https://doi.org/10.15587/2312-8372.2018.146600
6. Liubych, V. & Zheliezna, V. (2020). Effect of water heat treatment on spelt grain flour quality. Grain Products and Mixed Fodder's, 20, No. 2, pp. 19-25. https://doi.org/10.15673/gpmf.v20i2.1761
7. Hacini, N., Djelloul, R., Hadef, A., Samson, M.F. & Desclaux, D. (2022). Comparative characterization of grain protein content and composition by chromatography-based separation methods (SE-HPLC and RP-HPLC) of ten wheat varieties grown in different agro-ecological zones of Algeria. Separations, 9, No.12, 443. https://doi.org/10.3390/separations9120443
8. Pronin, D., Borner, A., Weber, H. & Scherf, K.A. (2020). Wheat (Triticum aestivum L.) breeding from 1891 to 2010 contributed to increasing yield and glutenin contents but decreasing protein and gliadin contents. J. Agric. Food Chem, 68, No. 26, pp. 13247-13256. https:doi.org/10.1021/acs.jafc.0c02815 https://doi.org/10.1021/acs.jafc.0c02815
9. Bazzo, J.H.B., Garcia, E.B., Cardoso, C.P. & Zucareli C. (2020). Physiological quality of wheat seeds in response to different nitrogen levels applied by soil and leaves. Rev. Terra Cult. Cad. Ensino e Pesqui, 36, pp. 145-156. http://publicacoes.unifil.br/ index.php/Revistateste/article/view/1115/1258
10. Giordano, N., Sadras, V.O. & Lollato, R.P. (2023). Late-season nitrogen application increases grain protein concentration and is neutral for yield in wheat. A global meta-analysis. Field Crops Res, 290, No. 2-3, 108740. https://doi.org/10.1016/j.fcr.2022.108740
11. Morgun, V.V. & Kots, S.Ya. (2018). The role of biological nitrogen in nitrogen nutrition of plants. Visn. Nats. Akad. Nauk Ukr., No. 1, pp. 62-74 [in Ukrainian]. https://doi.org/10.15407/visn2018.01.062
12. ZШrb, C., Ludewig, U. & Hawkesford, M.J. (2018). Perspective on wheat yield and quality with reduced nitrogen supply. Trends Plant Sci., 23, No. 11, pp. 1029-1037. https://doi.org/10.1016/j.tplants.2018.08.012
13. Hussain, A., Azeem, N., Riaz, T., Khan, T.Y., Pehlwan, B., Amjad, T., Muzammal, M., Sani, I.A., Umm-E-Habiba & Sarfraz, B. (2025). Impact of various nitrogenous fertilizers on wheat crop yield and growth. Indus J. Biosci. Res., 3, No. 3, pp. 364-369. https://doi.org/10.70749/ijbr.v3i3.886
14. Li, J., Wang, Y., Zhang, M., Liu, Y., Xu, X. & Lin, G. (2019). Optimized micro-sprinkling irrigation scheduling improves grain yield by increasing the uptake and utilization of water and nitrogen during grain filling in winter wheat. Agric. Water Manage., 211, pp. 59-69. https://doi.org/10.1016/j.agwat.2018.09.047
15. Zhang, G.X., Liu, S.J., Dong, Y.J., Liao, Y.C. & Han, J. (2022). A nitrogen fertilizer strategy for simultaneously increasing wheat grain yield and protein content: Mixed application of controlled-release urea and normal urea. Field Crops Res., 277, 108405. https://doi.org/10.1016/j.fcr.2021.108405
16. Rosculete, E., Paunescu, R.A., Rosculete, C.A., Paunescu, G., Bonciu, E., Olaru, A.L. & Murtaza, D.F. (2025). Nitrogen management of variety screening of winter wheat cultivated on chernozem soil for yield optimization. Plants, 14, No. 23, 3580. https://doi.org/10.3390/plants14233580
17. Pan, J., Zhao, J., Liu, Y., Huang, N., Tian, K. & Shah, F. (2019). Optimized nitrogen management enhances lodging resistance of rice and its morpho-anatomical, mechanical, and molecular mechanisms. Sci. Rep., 9, pp. 1-13. https://doi.org/10.1038/s41598-019-56620-7
18. Chen, X., Wang, M., Kroeze, C., Chen, X., Ma, L., Chen, X., Shi, X. & Strokal, M. (2022). Nitrogen in the Yangtze River Basin: Pollution reduction through coupling crop and livestock production. Environ. Sci. Technol., 56, No. 24, pp. 17591-17603. https://doi.org/10.1021/acs.est.1c08808
19. Gong, X., Li, L., Qin, L., Huang, Y., Ye, Y., Wang, M., Wang, Y., Xu, Y., Luo, F. & Mei, H. (2022). Targeted metabolomics reveals impact of N application on accumulation of amino acids, flavonoids and phytohormones in tea shoots under soil nutrition deficiency stress. Forests, 13, No. 10, 1629. https://doi.org/10.3390/f13101629
20. Liu, J., Zhang, J., Zhu, G., Zhu, D. & Yan, Y. (2022). Effects of water deficit and high N fertilization on wheat storage protein synthesis, gluten secondary structure, and breadmaking quality. Crop J., 10, No. 1, pp. 216-223. https://doi.org/10.1016/j.cj.2021.04.006
21. Muhammad, I., Yang, L., Ahmad, S., Farooq, S., Al-Ghamdi, A.A., Khan, A., Zeeshan, M., Elshikh, M.S., Abbasi, A.M. & Zhou, X.-B. (2022). Nitrogen fertilizer modulates plant growth, chlorophyll pigments and enzymatic activities under different irrigation regimes. Agronomy, 12, No. 4, 845. https://doi.org/10.3390/agronomy12040845
22. Drobnitch, S.T., Donovan, T.C., Wenz, J.A., Flynn, N.E., Schipanski, M.E. & Comas, L.H. (2024). Can nitrogen availability impact plant performance under water stress? A review of traits, mechanisms, and whole plant effects. Plant Soil, 511, No. 1, pp. 45-67. https://doi.org/10.1007/s11104-024-07006-w
23. Pasa, E.H., Weinert, C., Ferreira, J.P., Ferreira, H.T., Martinez, F.P., Pedo, T., Pasa, M. da S. & Carlos, F.S. (2024). Ammonium nitrate fertilization increases the crude protein content and wheat grain yield in subtropical conditions. Journal of Soil Science and Plant Nutrition, 24, No. 3, pp. 4952-4964. https://doi.org/10.1007/s42729-024-01884-w
24. Srivastava, R., Panda, R., Chakraborty, A. & Halder, D. (2018). Enhancing grain yield, biomass and nitrogen use efficiency of maize by varying sowing dates and nitrogen rate under rainfed and irrigated conditions. Field Crop. Res., 221, pp. 339-349. https://doi.org/10.1016/j.fcr.2017.06.019
25. Li, G., Niu, W., Sun, J., Zhang, W., Zhang, E. & Wang, J. (2021). Soil moisture and nitrogen content influence wheat yield through their effects on the root system and soil bacterial diversity under drip irrigation. Land Degrad. Dev., 32, No. 10, pp. 3062-3076. https://doi.org/10.1002/ldr.3967
26. Tahir, M., Arshad, M.A., Akbar, B.A., Bibi, A., Ain, Q.U., Bilal, A., Arqam, S.M., Asif, M., Ishtiaq, M.H. & Rasheed, H.U. (2024). Integrated nitrogen and irrigation management strategies for sustainable wheat production: Enhancing yield and environmental efficiency. J. Pharmacogn. Phytochem., 13, No. 4, pp. 209-222. https://doi.org/10.22271/phyto.2024.v13.i4c.15012
27. Si, Z.Y., Zain, M., Faisal, M., Wang, G.H., Gao, Y. & Duan, A.W. (2020). Effects of nitrogen application rate and irrigation regime on growth, yield, and water-nitrogen use efficiency of drip-irrigated winter wheat in the North China Plain. Agr. Water Manage., 231, 106002. https://doi.org/10.1016/j.agwat.2020.106002
28. Fradgley, N.S., Bentley, A.R. & Swarbreck, S.M. (2021). Defining the physiological determinants of low nitrogen requirement in wheat. Biochem. Soc. Trans., 49, No. 2, pp. 609-616. https://doi.org/10.1042/BST20200282
29. Yao, C., Li, J., Gao, Y., Zhang, Z., Liu, Y., Sun, Z., Wang, Z. & Zhang, Y. (2024). Delayed application of water and fertilizer increased wheat yield but did not improve quality parameters. Field Crops Res., 319, No. 2, 109649. https://doi.org/10.1016/j.fcr.2024.109649
30. Tong, J., Xiong, Y., Lu, Y., Li, W., Lin, W., Xue, J., Sun, M., Wang, Y. & Gao, Z. (2024). Drip fertigation enhances the responses of grain yield and quality to nitrogen topdressing rate in irrigated winter wheat in North China. Plants, 13, No. 11, 1439. https://doi.org/10.3390/plants13111439
31. Tong, J., Xiong, Y., Lu, Y., Li, W., Lin, W., Xue, J., Sun, M., Wang, Y. & Gao, Z. (2025). Drip fertigation with moderate nitrogen topdressing rate achieves high nitrogen and water use efficiencies for irrigated wheat. Agronomy, 15, No. 2, 259. https://doi.org/10.3390/agronomy15020259
32. Cui, H., Luo, Y., Li, C., Chang, Y., Jin, M., Li, Y. & Wang, Z. (2023). Effects of nitrogen forms on nitrogen utilization, yield, and quality of two wheat varieties with different gluten characteristics. Eur. J. Agron., 149, No. 6, 126919. https://doi.org/10.1016/j.eja.2023.126919
33. Chawla, R. & Balasaheb, K.S. (2023). Optimizing water use efficiency and yield of wheat crops through integrated irrigation and nitrogen management: A comprehensive review. Int. J. Environ. Clim. Change, 13, No. 11, pp. 4059-4067. https://doi.org/10.9734/ijecc/2023/v13i113585
34. Hamani, A.K.M., Abubakar, S.A., Fu, Y., Kpalari, D.F., Wang, G., Duan, A., Gao, Y. & Ju, X. (2025). The coupled effects of various irrigation scheduling and split nitrogen fertilization modes on post-anthesis grain weight variation, yield, and grain quality of drip-irrigated winter wheat (Triticum aestivum L.) in the North China Plain. J. Integr. Agric., 24, No. 6, pp. 2123-2137. https://doi.org/10.1016/j.jia.2023.12.037
35. 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
36. 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
37. 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
38. 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
39. 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
40. 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
41. 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
42. 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
43. 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
44. Stasik, O.O., Pryadkina, G.O., Kiriziy, D.A., Sokolovska-Sergiienko, O.G., Sytnyk S.K., Kapitanska, O.S. & Zborivska, O.V. (2021). Photosynthesis and production process of high-intensity varieties of winter wheat in connection with mineral nutrition conditions. Kyiv: Interservice.
45. 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
46. Hamani, A.K.M., Abubakar, S.A., Si, Z., Kama, R., Gao, Y. & Duan, A. (2023). Suitable split nitrogen application increases grain yield and photosynthetic capacity in drip-irrigated winter wheat (Triticum aestivum L.) under different water regimes in the North China Plain. Front. Plant Sci., 13, 1105006. https://doi.org/10.3389/fpls.2022.1105006
47. Shehab, A.E.S. A.E. & Guo Y. (2021). Effects of nitrogen fertilization and drought on hydrocyanic acid accumulation and morpho-physiological parameters of sorghums. J. Sci. Food Agric., 101, No. 8, pp. 3355-3365. https://doi.org/10.1002/jsfa.10965
48. 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, No. 1, pp. 1-14. https://doi.org/10.1038/s41598-020-75388-9
49. Zhang, H., Zhao, Q., Wang, Z., Wang, L., Li, X. & Fan, Z. (2021). Effects of nitrogen fertilizer on photosynthetic characteristics, biomass, and yield of wheat under different shading conditions. Agronomy, 11, No. 10, 1989. https://doi.org/10.3390/agronomy11101989
50. Zhang, X., Du, S., Xu, Y., Qiao, Y., Cao, C. & Li, W. (2022). Response of canopy photosynthesis, grain quality, and harvest index of wheat to different nitrogen applications methods. Plants, 11, No. 18, 2328. https://doi.org/10.3390/plants11182328
51. Zhou, Q., Wang, Q., Zheng, Z. & Liu, W. (2023). Carbon neutrality and nitrogen remediation in agricultural grain production. J. Agric. Environ. Sci., 42, No. 1, pp. 1-10. https://doi.org/10.11654/jaes.2022-1253
52. Ali, N. & Akmal, M. (2022). Wheat growth, yield, and quality under water deficit and reduced nitrogen supply. A Review. Gesunde Pflanz, 74, No. 2, pp. 371-383. https://doi.org/10.1007/s10343-021-00615-w
53. Noor, H., Yan, Z., Sun, P., Zhang, L., Ding, P., Li, L., Ren, A., Sun, M. & Gao, Z. (2023). Effects of nitrogen on photosynthetic productivity and yield quality of wheat (Triticum aestivum L.). Agronomy, 13, No. 6, 1448. https://doi.org/10.3390/agronomy13061448
54. Li, H.R., Wang, H.G., Fang, Q., Jia, B., Li, D.X. & He, J.N. (2023). Effects of irrigation and nitrogen application on NO3 - N distribution in soil, nitrogen absorption, utilization and translocation by winter wheat. Agr. Water Manage., 276, 108058. https://doi.org/10.1016/j.agwat.2022.108058
55. Fan, Y.H., Qin, B.Y., Yang, J.H., Ma, L.L., Cui, G.J. & He, W. (2024). Night warming increases wheat yield by improving pre-anthesis plant growth and post-anthesis grain starch biosynthesis. J. Integr. Agr., 23, No. 2, pp. 6563-6578. https://doi.org/10.1016/j.jia.2023.06.024
56. Noor, H., Ding, P.C., Ren, A.X., Sun, M. & Gao, Z.Q. (2023). Effects of nitrogen fertilizer on photosynthesis characteristics and yield. Agronomy, 13, No. 6, 1550. https://doi.org/10.3390/agronomy13061550
57. Yang, H., Fang, C., Li, Y., Wu, Y., Fransson, P. & Rillig, M.C. (2022). Temporal complementarity between roots and mycorrhizal fungi drives wheat nitrogen use efficiency. New Phytol., 236, pp. 1168-1181. https://doi.org/10.1111/nph.18419
58. 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.L. (2021). Enhanced photosynthetic nitrogen use efficiency and increased nitrogen allocation to photosynthetic machinery under cotton domestication. Photosynth. Res., 150, No. 1-3, pp. 239-250. https://doi.org/10.1007/s11120-021-00872-w
59. 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
60. Kiriziy, D.A. (2013). Nitrogen use efficiency during photosynthetic CO2 assimilation in wheat leaves. Fiziol. rast. genet., 45, No. 4, pp. 296-305 [in Russian].
61. Alpuerto, J.B., Brasier, K.G., Griffey, C.A., Thomason, W.E. & Fukao, T. (2021). Accelerated senescence and nitrogen remobilization in flag leaves enhances nitrogen use efficiency in soft red winter wheat. Plant Prod. Sci., 24, No. 4, pp. 490-504. https://doi.org/10.1080/1343943X.2020.1869044
62. Sheheda, I.M., Kiriziy, D.A., Ivanytska, A.P. & Senina, L.V. (2018). Nitrogen reutilization into grain in different wheat varieties depending on conditions of mineral nutrition. Visn. Kharkiv. nats.ahrar.univ. Ser. Biol., 44, No. 2, pp. 69-80 [in Ukrainian]. https://doi.org/10.35550/vbio2018.02.069
63. Hajibarat, Z. &, Saidi, A. (2022). Senescence-associated proteins and nitrogen remobilization in grain filling under drought stress condition. J. Genet. Eng. Biotechnol., 20, No. 1, p. 101. https://doi.org/10.1186/s43141-022-00378-5
64. Sheheda, I.M., Sandetska, N.V., Ivanytska, A.P. & Kiriziy, D.A. (2025). Nitrogen content in dry matter and parameters of its remobilization during the grain filling in different winter wheat varieties. FTzTol. rosl. genet., 57, No. 6, pp. 521-531 [in Ukrainian]. https://doi.org/10.15407/frg2025.06.521
65. Kiriziy, D.A., Stasik, O.O., Pryadkina, G.A. & Shadchina, T.M. (2014). CO2 Assimilation and Mechanisms of its Regulation. Photosynthesis (V. 2). Kyiv: Logos.
66. Kiriziy, D.A. (2015). Photosynthesis and source-sink relations between the organs as components of the production process of wheat. Fiziol. rast. genet., 47, No. 5, pp. 393-419.
67. Morgun, V.V., Shwartau, V.V., Konovalov, D.V., Mykhalska, L.V. & Skryplov, V.O. (2022). 100 Centners Club. Modern Varieties and Systems of Winter Wheat Nutrition and Protection. 11th edition. Sci. Edition. Kyiv: Vistka, 106 p. [in Ukrainian].
68. Yao, C., Li, J., Zhang, Z., Liu, Y., Wang, Z., Sun, Z. & Zhang Y. (2023). Improving wheat yield, quality and resource utilization efficiency through nitrogen management based on micro-sprinkler irrigation. Agric. Water Manag., 282, 108277. https://doi.org/10.1016/j.agwat.2023.108277
69. Liu, Y., Han, M., Zhou, X., Li, W., Du, C., Zhang, Y., Zhang, Y., Sun, Z. & Wang Z. (2022). Optimizing nitrogen fertilizing application under reduced irrigation strategies for winter wheat of the north China plain. Irrig. Sci., 40, pp. 255-265. https://doi.org/10.1007/s00271-021-00764-w
70. Hasanova, I.I., Yerashova, M.V. & Pedash, O.O. (2023). Influence of nitrogen feeding on winter wheat grain yield and quality. Ahronom, 79, No. 1, pp. 58-60.
71. Schwartau, V.V., Mykhalska, L.M. & Miroshnichenko, I.M. (2016). Physiological role of amino acids in the nutrition of highly productive varieties of winter wheat. Sortovyv№enn ohor. prav sorti roslin, No. 3, pp. 52-57. https://doi.org/10.21498/2518-1017.3(32).2016.75980
72. Liu, X., Yang, Y., Wu, B., Lv, C., Wei, H., Gao, P., Zhang, H., Dai, Q. & Chen, Y. (2025). Effects on nitrogen application on crop production and nitrogen use in rice-wheat rotation. Agronomy, 15, No. 5, 1047. https://doi.org/10.3390/agronomy15051047
73. Ren, H., Jiang, Y., Zhao, M., Qi, H. & Li, C.F. (2021). Nitrogen supply regulates vascular bundle structure and matter transport characteristics of spring maize under high plant density. Front. Plant Sci., 11, 602739. https://doi.org/10.3389/fpls.2020.602739
74. Liu, P.Z., Guo, X.Y., Zhou, D., Zhang, Q., Ren, X.L. & Wang, R. (2023). Quantify the effect of manure fertilizer addition and optimal nitrogen input on rainfed wheat yield and nitrogen requirement using nitrogen nutrition index. Agr Ecosyst. Environ., 345, 108319. https://doi.org/10.1016/j.agee.2022.108319
75. Ravier, C., Meynard, J.M., Cohan, J.P., Gate, P. & Jeuffroy, M.H. (2017). Early nitrogen deficiencies favor high yield, grain protein content and N use efficiency in wheat. Eur. J. Agron., 89, pp. 16-24. https://doi.org/10.1016/j.eja.2017.06.002
76. Liu, Z.X., Gao, F., Yang, J.Q., Zhen, X.Y., Li, Y., Zhao, J.H., Li, J.R., Qian, B.C., Yang, D.Q. & Li, X.D. (2019). Photosynthetic characteristics and uptake and translocation of nitrogen in peanut in a wheat-peanut rotation system under different fertilizer management regimes. Front. Plant Sci., 10, p. 86. https://doi.org/10.3389/fpls.2019.00086
77. Chen, L., Liu, X.F. & Fei, L.J. (2020). Evaluating the effects of regulated deficit irrigation on soil water movement, nitrate transport, photosynthetic characteristics, grain yield, water and N use efficiency under furrow irrigation. Int. J. Environ. Polut., 67, No. 2-4, pp. 143-181. https://doi.org/10.1504/IJEP.2020.117791
78. Schwartau, V.V., Mykhalska, L.M. (2016). Physiological basis of high-yielded cereals nutrition. Fiziol. rast. genet., 48, No. 4, pp. 298-309. https://doi.org/10.15407/frg2016.04.298
79. Ma, Q., Sun, Q., Zhang, X.B., Li, F.J., Ding, Y.G. & Tao, R.R. (2022). Controlled-release nitrogen fertilizer management influences grain in winter wheat by regulating flag leaf senescence post-anthesis and grain filling. Food Energy Secur., 11, e361. https://doi.org/10.1002/fes3.361
80. Wang, L., Liu, S., Ma, G., Wang, C. & Sun, J. (2022). Soil organic carbon and nitrogen storage under a wheat (Tritivum aestivum L.)-maize (Zea mays L.) cropping system in Northern China was modified by nitrogen application rates. Peer J., 10, e13568. https://doi.org/10.7717/peerj.13568
81. Li, C., Shi, Y., Yu, Z., Zhang, Y. & Zhang, Z. (2025). Optimizing nitrogen application strategies can improve grain yield by increasing dry matter translocation, promoting grain filling, and improving harvest indices. Front. Plant Sci., 16, 1565446. https://doi.org/10.3389/fpls.2025.1565446
82. Tian, Z.-W., Liu, X.-X., Gu, S.-L., Yu, J.-H., Zhang, L. & Zhang, W.-W. (2018). Postponed and reduced basal nitrogen application improves nitrogen use efficiency and plant growth of winter wheat. J. Integr. Agric., 17, No. 12, pp. 2648-2661. https://doi.org/10.1016/S2095-3119(18)62086-6
83. Zhao, F., Zou, G., Shan, Y., Ding, Z., Dai, M. & He, Z. (2019). Coconut shell derived biochar to enhance water spinach (Ipomoea aquatica forsk.) growth and decrease nitrogen loss under tropical conditions. Sci. Rep., 9, pp. 1-8. https://doi.org/10.1038/s41598-019-56663-w
84. Hafiza, B.S., Ishaque, W., Akhtar, M., Shani, M.Y., Azmat, M., Bauerle, W.L., Rahimi, M. & Ashraf, M.Y. (2025). Optimizing nitrogen management for sustainable wheat production in semi-arid subtropical environments: impact on growth, physio-biochemical, and yield attributes. Nitrogen, 6, No. 2, p. 36. https://doi.org/10.3390/nitrogen6020036
85. Liang, W.X., Zhang, Z.C., Wen, X.X., Liao, Y.C. & Liu, Y. (2017). Effect of non-structural carbohydrate accumulation in the stem pre-anthesis on grain filling of wheat inferior grain. Field Crop Res., 211, pp. 66-76. https://doi.org/10.1016/j.fcr.2017.06.016
86. Luo, Y.L., Li, W.Q., Huang, C., Yang, J.H., Jin, M. & Chen, J. (2021). Exogenous abscisic acid coordinating leaf senescence and transport of assimilates into wheat grains under drought stress by regulating hormones homeostasis. Crop J., 9, No. 4, pp. 901-914. https:doi.org/10.1016/j.cj.2020.08.012 https://doi.org/10.1016/j.cj.2020.08.012
87. Liu, Y., Liao, Y.C. & Liu, W.Z. (2021). High nitrogen application rate and planting density reduce wheat grain yield by reducing filling rate of inferior grain in middle spikelets. Crop J., 9, No. 2, pp. 412-426. https://doi.org/10.1016/j.cj.2020.06.013
88. Yue, K., Li, L.L., Xie, J.H., Liu, Y.Q., Xie, J.H. & Anwar, S. (2022). Nitrogen supply affects yield and grain filling of maize by regulating starch metabolizing enzyme activities and endogenous hormone contents. Front. Plant Sci., 12, 798119. https://doi.org/10.3389/fpls.2021.798119
89. Fang, L., Struik, P.C., Girousse, C., Yin, X.Y. & Martre, P. (2024). Source-sink relationships during grain filling in wheat in response to various temperature, water deficit, and nitrogen deficit regimes. J. Exp. Bot., 75, No. 20, pp. 6563-6578. https://doi.org/10.1093/jxb/erae310
90. Du, K., Zhao, W.Q., Lv, Z.W., Xu, B., Hu, W. & Zhou, Z.G. (2024). Optimal rate of nitrogen fertilizer improves maize grain yield by delaying the senescence of ear lives and thereby altering their nitrogen remobilization. Field Crop Res., 310, 109359. https://doi.org/10.1016/j.fcr.2024.109359
91. Meng, F.G., Zhao, L.D., Li, W.L. & Zhao, C.X. (2024). Optimizing nitrogen application for enhanced yield and quality of strong-gluten wheat: A case study of Zhongmai 578 in the north China plain. Agronomy, 14, No. 6, 1301. https://doi.org/10.3390/agronomy14061301
92. Ye, J.Y., Zhong, X.F., Harrison, M.T., Kang, K., Sheng, T., Shang, C., Wang, C.H., Deng, J., Huang, L.Y. & Tian, X.H. (2023). Towards improved grain yield and soil microbian communities of super hybrid rice through sustanaible management. Agronomy, 13, No. 9, 2259. https://doi.org/10.3390/agronomy13092259
93. Zain, M., Si, Z., Li, S., Gao, Y., Mehmood, F. & Rahman S.-U. (2021). The coupled effects of irrigation scheduling and nitrogen fertilization mode on growth, yield and water use efficiency in drip-irrigated winter wheat. Sustainability, 13, No. 5, 2742. https://doi.org/10.3390/su13052742
94. Liu, Z., Gao, F., Liu, Y., Yang, J., Zhen, X. & Li, X. (2019). Timing and splitting of nitrogen fertilizer supply to increase crop yield and efficiency of nitrogen utilization in a wheat-peanut relay intercropping system in China. Crop J., 7, No. 1, pp. 101-112. https://doi.org/10.1016/j.cj.2018.08.006
95. Zhang, Y., Wang, H., Wang, R., He, F., Jiang, G. & Xu, J. (2025). Improvement of the photosynthetic characteristics and yield of wheat by regulating the proportion of nitrogen fertilizer base and topdressing. Agronomy, 15, No 4, 899. https://doi.org/10.3390/agronomy15040899
96. Chen, X.G. (2018). Optimized nitrogen fertilizer application mode increased culms lignin accumulation and lodging resistance in culms of winter wheat. Field Crops Res., 228, pp. 31-38. https://doi.org/10.1016/j.fcr.2018.08.019
97. Yao, C.S., Li, J.P., Gao, Y.M., Zhang, Z., Liu, Y., Sun, Z.C., Wang, Z.M. & Zhang, Y.H. (2024). Delayed application of water and fertilizer increased wheat yield but did not improve quality parameters. Field Crops Res., 319, 109649. https://doi.org/10.1016/j.fcr.2024.109649
98. Wang, L., Palta, J.A., Chen, W., Chen, Y. & Deng X. (2018). Nitrogen fertilization improved water-use efficiency of winter wheat through increasing water use during vegetative rather than grain filling. Agric. Water Manag., 197, pp. 41-53. https://doi.org/10.1016/j.agwat.2017.11.010
99. Ju, X.T. & Zhang, C. (2021). The principles and indicators of rational N fertilization. Acta Pedol. Sin., 58, pp. 1-13. https://doi.org/10.11766/trxb202006220322
100. Trevisan, S., Khorshidi, A.S. & Scanlon, M.G. (2022). Relationship between nitrogen functionality and wheat flour dough rheology: Extensional and shear approaches. Food Res. Int., 162, No. 8, 112049. https://doi.org/10.1016/j.foodres.2022.112049
101. Wu, W., Wang, Y., Wang, L., Xu, H.S., Zorb, C., Geilfus, G.M., Xue, C., Sun, Z.M. & Ma, W.Q. (2022). Booting stage is the key timing for split nitrogen application in improving grain yield and quality of wheat - A global meta-analysis. Field Crops Res., 287, 108665. https://doi.org/10.1016/j.fcr.2022.108665
102. An, H.-Y., Han, J.-J., He, Q.-N., Zhu, Y.-L., Wu, P., Wang, Y.-C., Gao, Z.-Q., Du, T.-Q. & Xue, J.-F. (2024). Influence of nitrogen application rate on wheat grain protein content and composition in China: A meta-analysis. Agronomy, 14, No. 6, 1164. https://doi.org/10.3390/agronomy14061164
103. Abdelrhman, A.A., Abdel-Fattah, I.M., Mostafa, M.O., Fadl, M.E., Drosos, M. & Scopa, A. (2025). Optimizing drip irrigation and nitrogen fertilization for sustainable wheat production in arid soils: Water-nitrogen use efficiency. Water, 17, No. 18, 2708. https://doi.org/10.3390/w17182708
104. Horvat, D., Simic, G., Dvojkovic, K., Ivik, M., Plavsin, I. & Novoselovic, D. (2021). Gluten protein compositional changes in response to nitrogen application rate. Agronomy, 11, No. 2, p. 325. https://doi.org/10.3390/agronomy11020325
105. Sun, M., Feng, H., Zhang, X., Liu, S., Han, Z., Hao, X., Ma, G., Wang, L. & Wang C. (2022). Effects of nitrogen rates on wheat yield and quality under different soil fertility regimes. J. Triticaceae Crops, 42, pp. 826-834.
106. Wang, W.Z., Li, Y.J., Zhang, M., Bi, J.X., Yang, M., Guo, Z.Q. & Cai, R.Q. (2024). Effects of nitrogen application rate on photosynthesis characteristics, yield, protein content, and processing quality of strong gluten wheat. J. Triticaceae Crops, 44, pp. 462-471.
107. Wang, J.J., Sun, X., Hussain, S., Yang, L.H., Gao, S.S., Zhang, P., Chen, X.L. & Ren, X.L. (2024). Reduced nitrogen rate improves post-anthesis assimilates to grain and ammeolirates grain-filling characteristics of winter wheat in dry land. Plant Soil, 499, No. 1-2, pp. 91-112. https://doi.org/10.1007/s11104-023-06276-0
108. Kozlowski, O., Balik, J., Cerny, J., Kulhanek, M., Kos, M. & Prasilova, M. (2022). Regulation of nitrogen application on grain yield and quality of strong-gluten wheat in Huaibei region. J. Triticaceae Crops, 42, pp. 99-112.
109. Gu, X., Yang, X., Jiang, L. & Huang, S. (2023). Nitrogen foliage application at anthesis improves grain yield and quality of wheat in genotype-dependent manner. Agronomy, 13, No. 12, 3077. https://doi.org/10.3390/agronomy13123077
110. Maignan, V., Coquerel, R., Geliot, P. & Avice, J.C. (2021). VNT4, a derived formulation of glutacetine biostimulant, improved yield and N-related traits of bread wheat when mixed with urea-ammonium-nitrate solution. Agronomy, 11, No. 5, 1013. https://doi.org/10.3390/agronomy11051013
111. Lyu, X., Liu, Y., Li, N., Ku, L., Hou, Y. & Wen, X. (2022). Foliar applications of various nitrogen (N) forms to winter wheat affect grain protein accumulation and quality via N metabolism and remobilization. Crop J., 10, No. 4, pp. 1165-1177. https://doi.org/10.1016/j.cj.2021.10.009
112. Ochieng', I.O., Gitari, H.I., Mochoge, B., Rezaei-Chiyaneh, E. & Gweyi-Onyango, J.P. (2021). Optimizing maize yield, nitrogen efficacy and grain protein content under different N forms and rates. J. Soil Sci. Plant Nutr., 21, pp. 1867-1880. https://doi.org/10.1007/s42729-021-00486-0
113. Yao, Y., Ma, W., Jin, X., Liu, G., Li, Y., Liu, B. & Cao, D. (2025). Foliar nitrogen application enhances nitrogen assimilation and modulates nitrogen expression in spring wheat leaves. Agronomy, 15, No. 7. 1688. https://doi.org/10.3390/agronomy15071688
114. Mi, X.T., Bai, N., Liu, Y.R., He, G. & Wang, Z.H. (2023). Exploring nitrogen management methods for depressing the decline of wheat grain protein in plastic film mulch via 15N-labeling technique. Soil Tillage Res., 228, 105632. https://doi.org/10.1016/j.still.2022.105632
115. Chen, Y.F., Chen, H.L., Chen, R.H., Yang, H.K., Zheng, T., Huang, X.L. & Fan, G.Q. (2023). The impacts of nitrogen accumulation, translocation, and photosynthesis on simultaneous improvements in the grain yield and gluten quality of dryland wheat. Agronomy, 13, No. 5, 1283. https://doi.org/10.3390/agronomy13051283
116. Xia, H.Y., Li, X.J., Qiao, Y.T., Xue, Y.H., Yan, W., Ma, L., Zhao, Q.Y., Kong, L.A., Xue, Y.F. & Cui, Z.L. (2023). Dissecting the relationship between yield and mineral nutriome of wheat grains in double cropping as affected by preceding crops and nitrogen application. Field Crops Res., 293, 108845. https://doi.org/10.1016/j.fcr.2023.108845
117. Ben Marien, S., Gonzales-Torralba, J., Collar, C., Aranjuelo, I. & Morales, F. (2020). Durum wheat grain yield and quality under low and high nitrogen conditions: Insights into natural variation in low- and high-yielding genotypes. Plants, 9, No. 12, 517313. https://doi.org/10.3390/plants9121636
118. Shiwakoti, S., Jeliazkov, V., Gollany, H., Kleber, M. & Xing, B. (2019). Micronutrients decline under long-term tillage and nitrogen fertilization. Sci. Rep., 9, No. 1, 12020. https://doi.org/10.1038/s41598-019-48408-6
119. Rekowski, A., Wimmer, M.A., Henkelmann, G. & ZШrb, C. (2019). Is a change of protein composition after lane application of nitrogen sufficient to improve the baking quality of winter wheat? Agriculture, 9, No. 5, p. 101. https://doi.org/10.3390/agriculture9050101
120. Shang, Y., Wang, S., Lin, X., Gu, S. & Wang, D. (2023). Supplemental irrigation at jointing improves spike formation of wheat tillers by regulating sugar distribution in ear and stem. Agr. Water Manage., 279, 108160. https://doi.org/10.1016/j.agwat.2023.108160
121. Roy, S. & Verma, O. (2018). Seed quality and storage of wheat (Triticum aestivum L.) as influenced by basal and foliar application of nitrogen. Natl. Acad. Sci. Lett., 41, No. 6, pp. 337-340. https://doi.org/10.1007/s40009-018-0688-0
122. Wu, W., Wang, Y., Xu, H., Liu, M. & Xue, C. (2025). Enhancing wheat yield and quality through late-season foliar nitrogen application: A global meta-analysis. Agronomy, 15, No. 5, 1058. https://doi.org/10.3390/agronomy15051058
123. Roychowdhury, R., Zilberman, O., Chandrasekhar, K., Kurzon, A.Y., Nashef, K., Abbo, S., Slafer, G.A., Bonfil, D.J. & Ben-David, R. (2023). Pre-anthesis spike growth dynamics and its association to yield components among elite bread wheat cultivars (Triticum aestivum L. spp.) under Mediterranean climate. Field Crops Res., 298, 108948. https://doi.org/10.1016/j.fcr.2023.108948
124. Morgun, V.V., Schwartau, V.V. & Kiriziy, D.A. (2009). Physiological bases of formation of high productivity of grain cereals. In Fiziologiia roslyn: Problemy i perspektyvy rozvytku (Vol. 1) (pp. 11-42). Kyiv: Lohos [in Ukrainian].
125. Rossman, A., Buchner, P., Savill, G.P., Hawkesford, M.J., Scherf, K.A. & Mтhling, K.H. (2019). Foliar N application at anthesis alters grain protein composition and enhances baking quality in winter wheat only under a low N fertilizer regimen. Eur. J. Agron., 109, p. 4. https://doi.org/10.1016/j.eja.2019.04.004
126. Tedone, L., Alhajj, Ali, S., Verdini, L. & De Mastro, G. (2018). Nitrogen management strategy for optimizing agronomic and environmental performance of rainfed durum wheat under Mediterranean climate. J. Clean. Prod., 172, p. 2058-2074. https://doi.org/10.1016/j.jclepro.2017.11.215
127. Pan, Y., Han, X., Xu, H., Wu, W., Liu, X., Li, Y. & Xue, C. (2023). Elevated atmospheric CO2 delays the key timing for split N applications to improve wheat (Triticum aestivum L.) protein composition. Front. Plant Sci., 14, 1186890. https://doi.org/10.3389/fpls.2023.1186890
128. Hou, P., Hu, C., Yu, J., Gao, Q., Zhou, M., Gao, L., Jiang, D., Dai, T. & Tian, Z. (2024). Increasing topdressing ratio of nitrogen fertilizer improves grain yield and grain use efficiency of winter wheat under winter and spring night-warming. J. Soil Sci. Plant Nutr., 24, No. 2, pp. 3459-3473. https://doi.org/10.1007/s42729-024-01767-0
129. Dong, L., Li, Y., Li, P., Liu, Y., Ma, F., Hao, X. & Guo, L. (2023). Growth response of wheat and maize to different nitrogen supply forms under the enrichment of amospheric CO2 concentrations. Agronomy, 13, No. 2, 485. https://doi.org/10.3390/agronomy13020485