Plant Soil Environ., 2026, 72(3):172-193 | DOI: 10.17221/20/2026-PSE

Nitrogen application rates mediate rice cooking quality by interfering with root anatomical and senescence physiological traitsOriginal Paper

Li Wang1, Jing Cao1, Hao Cheng1, Qinyao Meng1, Haojing Li1, Guowei1
1 College of Agriculture, Henan University of Science and Technology, Luoyang, Henan, P.R. China

Nitrogen fertiliser is a key determinant of rice yield and grain quality; however, the synergistic mechanisms through which nitrogen regulates root anatomical structure, physiological traits, and cooking quality in rice varieties with different eating properties remain unclear. In this study, a pot experiment was conducted using two moderate-eating-quality cultivars (Xudao 3 and Huageng 9) and two superior-eating-quality cultivars (Zhengdao C42 and Nangeng 9308) under four nitrogen levels (0, 0.59, 1.18, and 1.76 g/pot, designated as N0, N1, N2, and N3, respectively). Cooking quality was assessed by amylose content, gel consistency, and alkali spreading value. The results demonstrated that, with increasing nitrogen application, amylose content, alkali spreading value, malondialdehyde (MDA) content, root aerenchyma area, and aerenchyma proportion decreased initially, then increased, reaching their lowest values at the N2 level. In contrast, gel consistency, root antioxidant enzyme activities (SOD, POD, CAT), photosynthetic rate and cortical living cell proportion increased first and then decreased, peaking at N2 treatment. Compared with moderate-eating-quality varieties, superior-eating-quality varieties exhibited significantly lower amylose content, alkali spreading value, MDA content, and aerenchyma proportion, but higher gel consistency, living cell proportion, stele-to-root diameter ratio, antioxidant enzyme activities, and photosynthetic rate. Correlation analysis revealed that root antioxidant enzyme activities, stele diameter and living cell proportion were negatively correlated with amylose content, but positively correlated with gel consistency. Conversely, MDA content, aerenchyma area and aerenchyma proportion showed opposite correlation patterns. These findings indicate that an appropriate nitrogen application rate (1.18 g/pot) enhances root physiological activity, optimises root anatomical structure, and ensures sufficient source supply to the grain sink, thereby synergistically improving cooking quality – an effect particularly pronounced in high-eating-quality rice varieties.

Keywords: Oryza sativa L.; nutrition; root senescence; nitrogen application level; root anatomy; root physiological property

Received: January 15, 2026; Revised: March 3, 2026; Accepted: March 12, 2026; Prepublished online: March 20, 2026; Published: March 31, 2026  Show citation

ACS AIP APA ASA Harvard Chicago Chicago Notes IEEE ISO690 MLA NLM Turabian Vancouver
Wang L, Cao J, Cheng H, Meng Q, Li H, Guowei. Nitrogen application rates mediate rice cooking quality by interfering with root anatomical and senescence physiological traits. Plant, Soil and Environment. 2026;72(3):172-193. doi: 10.17221/20/2026-PSE.
Download citation

References

  1. Averill-Bates D. (2024): Reactive oxygen species and cell signalling. Review. Biochimica et Biophysica Acta (BBA)-Molecular Cell Research, 1871: 119573. Go to original source... Go to PubMed...
  2. Bahmaniar M.A., Ranjbar G.A. (2007): Response of rice (Oryza sativa L.) cooking quality properties to nitrogen and potassium application. Pakistan Journal of Biological Sciences, 10: 1880-1884. Go to original source... Go to PubMed...
  3. Bin Rahman A.R., Zhang J. (2023): Trends in rice research: 2030 and beyond. Food and Energy Security, 12: e390. Go to original source...
  4. Bulas T., Schmidt B.R., Vorburger C., Fabian Y. (2025): Responses of rice paddy dragonflies to fertilisation in a mesocosm experiment. Agriculture, Ecosystems and Environment, 393: 109823. Go to original source...
  5. Chen M., Chen G., Di D., Kronzucker H.J., Shi W. (2020): Higher nitrogen use efficiency (NUE) in hybrid "super rice" links to improved morphological and physiological traits in seedling roots. Journal of Plant Physiology, 251: 153191. Go to original source... Go to PubMed...
  6. Chen Y., Ding Z., Yu T., Cao L., Duan Y., Zu Y., Li Z. (2024): Accumulation characteristics of heavy metals in three wild rice species and adaptation of root morphology and anatomical structure to native soil heavy metals in Yunnan. Ecological Indicators, 167: 112601. Go to original source...
  7. Chen Z., Hou Y., Yan J., Cheng S., Wang Y., Feng G., Cai H. (2025): Comprehensive responses of root system architecture and anatomy to nitrogen stress in maize (Zea mays L.) genotypes with contrasting nitrogen efficiency. Agronomy, 15: 2083. Go to original source...
  8. Chu G., Xu R., Chen S., Xu C., Liu Y., Abliz B., Zhang X., Wang D. (2022): Root morphological-physiological traits for japonica/indica hybrid rice with better yield performance under low N conditions. Food and Energy Security, 11: e355. Go to original source...
  9. Cui Y., Ma W., Yang C., Bai Y., Xia T., Wei C., Zhang X., Zhou G. (2025): Subsurface drip irrigation combined with ammonium enhances root growth in rice (Oryza sativa L.), leading to improved N uptake and higher yield formation. Plants, 14: 891. Go to original source... Go to PubMed...
  10. Dai L., Zhou X., Jian Z., Tian J., Li Y., Xu G. (2024): Comparison of leaf anatomical structure and photosynthetic characteristics between weedy rice and cultivated rice at the seedling stage. Scientific Reports, 14: 30829. Go to original source... Go to PubMed...
  11. De Bauw P., Vandamme E., Lupembe A., Mwakasege L., Senthilkumar K., Dramé K.N., Merckx R. (2019): Anatomical root responses of rice to combined phosphorus and water stress-relations to tolerance and breeding opportunities. Functional Plant Biology, 46: 1009-1022. Go to original source... Go to PubMed...
  12. Dou Z., Zhou Y., Zhang Y., Guo W., Xu Q., Gao H. (2024): Influence of nitrogen applications during grain-filling stage on rice (Oryza sativa L.) yield and grain quality under high temperature. Agronomy, 14: 216. Go to original source...
  13. Drobnitch S.T., Donovan T.C., Wenz J.A., Flynn N.E., Schipanski M.E., Comas L.H. (2025): Can nitrogen availability impact plant performance under water stress? A review of traits, mechanisms, and whole plant effects. Plant and Soil, 511: 45-67. Go to original source...
  14. Duan R., Lin Y., Yang L., Zhang Y., Hu W., Du Y., Huang M. (2023): Effects of antimony stress on growth, structure, enzyme activity and metabolism of Nipponbare rice (Oryza sativa L.) roots. Ecotoxicology and Environmental Safety, 249: 114409. Go to original source... Go to PubMed...
  15. Eweda M.A., Jalil S., Rashwan A.K., Tsago Y., Hassan U., Jin X.J. (2025): Molecular and physiological characterizations of roots under drought stress in rice: a comprehensive review. Plant Physiology and Biochemistry, 225: 110012. Go to original source... Go to PubMed...
  16. Firouzi S. (2015): Grain, milling, and head rice yields as affected by nitrogen rate and bio-fertilizer application. Acta Agriculturae Slovenica, 105: 241-248. Go to original source...
  17. Fonta J.E., Giri J., Vejchasarn P., Lynch J.P., Brown K.M. (2022): Spatiotemporal responses of rice root architecture and anatomy to drought. Plant and Soil, 479: 443-444. Go to original source...
  18. Fukagawa N.K., Ziska L.H. (2019): Rice: importance for global nutrition. Journal of Nutritional Science and Vitaminology, 65: S2-S3. Go to original source... Go to PubMed...
  19. Galindo-Castañeda T., Lynch J.P., Six J., Hartmann M. (2022): Improving soil resource uptake by plants through capitalizing on synergies between root architecture and anatomy and root-associated microorganisms. Frontiers in Plant Science, 13: 827369. Go to original source... Go to PubMed...
  20. Gharsallah C., Fakhfakh H., Grubb D., Gorsane F. (2016): Effect of salt stress on ion concentration, proline content, antioxidant enzyme activities and gene expression in tomato cultivars. AoB Plants, 8: plw055. Go to original source...
  21. Guo Y., Du Y., Niu X., Ma Y., Song G., Cao C., Li P., Chen Y., Siddique K. (2024): Comparison of drought stress responses in large-and small-rooted rice lines: physiological, anatomical, and hormonal changes. Journal of Plant Growth Regulation, 43: 2922-2936. Go to original source...
  22. Hou D., Liu K., Liu S., Li J., Tan J., Bi Q., Zhang A., Yu X., Bi J., Luo L. (2024): Enhancing root physiology for increased yield in water-saving and drought-resistance rice with optimal irrigation and nitrogen. Frontiers in Plant Science, 15: 1370297. Go to original source... Go to PubMed...
  23. Huang S., Zhen H., Zheng Q. (2020): Characterization of eating quality and starch properties of two Wx alleles japonica rice cultivars under different nitrogen treatments. Journal of Integrative Agriculture, 19: 988-998. Go to original source...
  24. Khan M., Al Azzawi T.N.I., Imran M., Hussain A., Mun B.G., Pande A., Yun B.W. (2021): Effects of lead (Pb)-induced oxidative stress on morphological and physio-biochemical properties of rice. Biocell, 45: 1413. Go to original source...
  25. Koper K., de Oliveira M.V., Huß S., Hataya S., Soleymani F., Hawkins C., Rhee S.Y., Takasuka T.E., Nikoloski Z., Maeda H.A. (2025): Mapping multi-substrate specificity of Arabidopsis aminotransferases. Nature Plants, 11: 1863-1876. Go to original source... Go to PubMed...
  26. Kusano M., Fukushima A., Redestig H., Saito K. (2011): Metabolomic approaches toward understanding nitrogen metabolism in plants. Journal of Experimental Botany, 62: 1439-1453. Go to original source... Go to PubMed...
  27. Lee S., Masclaux-Daubresse C. (2021): Current understanding of leaf senescence in rice. International Journal of Molecular Sciences, 22: 4515. Go to original source... Go to PubMed...
  28. Li X., Wang R., Zhou B., Wang X., Wang J., Zhao M., Li C. (2022): Characterization of root morphology and anatomical structure of spring maize under varying N application rates and their effects on yield. Agronomy, 12: 2671. Go to original source...
  29. Li H.J., Li H.R., Zhang D.K., Jiang M.M., Cao J., Xu G.W. (2025): Irrigation methods and nitrogen-form interactions regulate starch-metabolising enzyme activity to improve rice yield and quality. Plant, Soil and Environment, 71: 185-201. Go to original source...
  30. Li J., Yang S., Zhong Z., Xia T., Zhou W., Tu Z.,Chen Z.,Wang H., Dai Z., Jin G., Du Y. (2025): Effects of degree of milling on bran layer structure, physicochemical properties and cooking quality of brown rice. Food Chemistry, 462: 140847. Go to original source... Go to PubMed...
  31. Liang H., Gu B., Wang T., Rong L., Sun W., Wu Z. (2022): Relationship between protein structure and eating quality of rice under different nitrogen application rate. Cereal Chemistry, 99: 692-703. Go to original source...
  32. Liu H., Wang W., He A., Nie L. (2018): Correlation of leaf and root senescence during ripening in dry seeded and transplanted rice. Rice Science, 25: 279-285. Go to original source...
  33. Liu Q., Chen S., Zhou L., Tao Y., Tian J., Xing Z., Wei H., Zhang H. (2022): Characteristics of population quality and rice quality of semi-waxy japonica rice varieties with different grain yields. Agriculture, 12: 241. Go to original source...
  34. Liu A., Li Y., Yang L., Zhang Y., Liao S., Li X. (2024): Applying nitrogen fertilizer improves the indica rice (Oryza sativa L.) quality by coordinating enzyme activity and grain-filling rate. Cereal Chemistry, 101: 1043-1054. Go to original source...
  35. Liu K., Chen Y., Li S., Wang W., Zhang W., Zhang H., Gu J., Yang J., Liu L. (2023): Differing responses of root morphology and physiology to nitrogen application rates and their relationships with grain yield in rice. The Crop Journal, 11: 618-627. Go to original source...
  36. Liu J., Yang T., Zou J., Zou J., Wu L., Bao X., Zhang B. (2025): Effects of reduced nitrogen fertilizer application rates on grain yield and rice quality of early-and late-season dual-use rice in South China. The Journal of Agricultural Science, 163: 99-109. Go to original source...
  37. Lopez-Valdivia I., Yang X., Lynch J.P. (2023): Large root cortical cells and reduced cortical cell files improve growth under suboptimal nitrogen in silico. Plant Physiology, 192: 2261-2275. Go to original source... Go to PubMed...
  38. Lv R., Zhang W., Xie X., Wang Q., Gao K., Zeng Y., Pan X., Shang Q. (2022): Foliar application uniconazole enhanced lodging resistance of high-quality indica rice (Oryza sativa L.) by altering anatomical traits, cell structure and endogenous hormones. Field Crops Research, 277: 108425. Go to original source...
  39. Lynch J.P., Galindo-Castañeda T., Schneider H.M., Sidhu J.S., Rangarajan H., York L.M. (2023): Root phenotypes for improved nitrogen capture. Plant and Soil, 10: 06301. Go to original source... Go to PubMed...
  40. Ma Z., Chen X., Cao J., Yu J., Zhu Y., Li G., Xu F., Zhang H., Liu G., Wei H. (2025): Analysis of the effects of light and panicle fertilizer on rice eating quality based on morphological structural changes in starch and protein during cooking. International Journal of Biological Macromolecules, 305: 141121. Go to original source... Go to PubMed...
  41. Meng F., Guo J., Feng N., Zheng D., Chen X., Chen Z., Jiang H., Jiang X. (2024): Beneficial effects of hemin on antioxidative capacity and anatomical characters of NaCl-stressed rice plants. Journal of Plant Growth Regulation, 43: 3743-3760. Go to original source...
  42. Owusu E.A., Luthra K., Regonda B., Atungulu G. (2025): Effects of nitrogen application rates on the physicochemical properties of modern rice cultivars: a comprehensive study. Cereal Chemistry, 102: 999-1012. Go to original source...
  43. Qi D., Chen S., Yue W., Duan Y. (2024): Leaf senescence characteristics and economic benefits of rice under alternate wetting and drying irrigation and blended use of polymer-coated and common urea. Frontiers in Plant Science, 15: 1444819. Go to original source... Go to PubMed...
  44. Sachadyn-Król M., Materska M., Chilczuk B., Kara¶ M., Jakubczyk A., Perucka I., Jackowska I. (2016): Ozone-induced changes in the content of bioactive compounds and enzyme activity during storage of pepper fruits. Food Chemistry, 211: 59-67. Go to original source... Go to PubMed...
  45. Salah A., Nwafor C.C., Han Y., Liu L., Rashid M., Batool M., El-Badri A.M., Cao C., Zhan M. (2022): Spermidine and brassinosteroid regulate root anatomical structure, photosynthetic traits and antioxidant defense systems to alleviate waterlogging stress in maize seedlings. South African Journal of Botany, 144: 389-402. Go to original source...
  46. Salsinha Y.C.F., Rini D.S., Indradewa D., Purwestri Y.A., Fatikhasari Z., Lailaty I.Q., Wutsqa Y.U., Harijayanti D., Rachmawati D. (2023): Anatomical characteristics of the upland local Indonesian rice cultivars as structural protection against drought. Brazilian Journal of Botany, 46: 645-659. Go to original source...
  47. Schneider H.M., Postma J.A., Wojciechowski T., Kuppe C., Lynch J.P. (2017): Root cortical senescence improves growth under suboptimal availability of N, P, and K. Plant Physiology, 174: 2333-2347. Go to original source... Go to PubMed...
  48. Shi S., Ma Y., Zhao D., Li L., Cao C., Jiang Y. (2023): The differences in metabolites, starch structure, and physicochemical properties of rice were related to the decrease in taste quality under high nitrogen fertilizer application. International Journal of Biological Macromolecules, 253: 126546. Go to original source... Go to PubMed...
  49. Shiono K., Ejiri M., Sawazaki Y., Egishi Y., Tsunoda T. (2024): Low nitrate under waterlogging triggers exodermal suberization to form a barrier to radial oxygen loss in rice roots. Plant Physiology, 196: 551-563. Go to original source... Go to PubMed...
  50. Singh N., Pal N., Mahajan G., Singh S., Shevkani K. (2011): Rice grain and starch properties: effects of nitrogen fertilizer application. Carbohydrate Polymers, 86: 219-225. Go to original source...
  51. Song Y., Dong M., Chen F., Hu Y., Zhu Y., Gu J., Chen P., Xie Y., Yuan C., Qiao Z., Yu Y., Cao P., Shi L., Wang Y., Zhang M. (2024): Effects of nitrogen fertilizer reduction combined with foliar fertilizer application on the physiological characteristics and yield of high-quality japonica rice. International Journal of Plant Production, 18: 239-254. Go to original source...
  52. Srivastava A., Gupta A., Bishi S.K., Akhila P., Latha P.C., Subrahmanyam D., Brajenndra P., Anantha M.S., Ch S.R., Sakhare A.S., Bhadana V.P., Giri J., Nddraja C.N., Sundaram R.M., Mangrauthia S.K. (2025): Tolerance of Oryza sativa to low phosphate is associated with adaptive changes in root architecture and metabolic exudates. Plant Science, 353: 112415. Go to original source... Go to PubMed...
  53. Subudhi P.K., Garcia R.S., Coronejo S., Tapia R. (2020): Comparative transcriptomic of rice genotypes with contrasting responses to nitrogen stress reveals genes influencing nitrogen uptake through the regulation of root architecture. International Journal of Molecular Sciences, 21: 5759. Go to original source... Go to PubMed...
  54. Tavu L.E.J., Redillas M.C.F.R. (2025): Oxidative stress in rice (Oryza sativa): mechanisms, impact, and adaptive strategies. Plants, 14: 1463. Go to original source... Go to PubMed...
  55. Turk H., Erdal S., Genisel M., Atici O., Demir Y., Yanmis D. (2014): The regulatory effect of melatonin on physiological, biochemical and molecular parameters in cold-stressed wheat seedlings. Plant Growth Regulation, 74: 139-152. Go to original source...
  56. Uga Y., Ebana K., Abe J., Morita S., Okuno K., Yano M. (2009): Variation in root morphology and anatomy among accessions of cultivated rice (Oryza sativa L.) with different genetic backgrounds. Breeding Science, 59: 87-93. Go to original source...
  57. Velikova V., Yordanov I., Edreva A.J.P.S. (2000): Oxidative stress and some antioxidant systems in acid rain-treated bean plants: protective role of exogenous polyamines. Plant Science, 151: 59-66. Go to original source...
  58. Wang R., Sun C., Cai S., Liu F., Xie H., Xiong Q. (2023): Research progress in crop root biology and nitrogen uptake and use, with emphasis on cereal crops. Agronomy, 13: 1678. Go to original source...
  59. Wang C., Jing W., Fa X., Gu J., Wang W., Zhu K., Zhang W., Gu J., Wang Z., Zhang J., Zhang H. (2025): Effect of nitrogen levels on grain yield and quality in improving mid-season japonica rice varieties. Journal of the Science of Food and Agriculture, 105: 3010-3023. Go to original source... Go to PubMed...
  60. Wei G., Wang X., Wu Y., Liu F., Lan T., Liu Q., Lyu C., Kong F., Yuan J. (2025): Post-silking leaf senescence is delayed in low-N-tolerant maize cultivars under low N fertilization. The Crop Journal, 13: 246-256. Go to original source...
  61. Xin W., Liu H., Zhao H., Wang J., Zheng H., Jia Y., Yang L., Wang X., Li J., Li X., Lei L., Zou D. (2021): The response of grain yield and root morphological and physiological traits to nitrogen levels in paddy rice. Frontiers in Plant Science, 12: 713814. Go to original source... Go to PubMed...
  62. Xu C., Chen L., Chen S., Chu G., Wang D., Zhang X. (2020): Effects of rhizosphere oxygen concentration on root physiological characteristics and anatomical structure at the tillering stage of rice. Annals of Applied Biology, 177: 61-73. Go to original source...
  63. Xu Y., Li K., Zhu K., Tian Y., Yu Q., Zhang W., Wang Z. (2021): Effect of nitrogen rates on the growth and grain yield of a leaf early-senescent rice mutant. Agronomy Journal, 113: 1503-1522. Go to original source...
  64. Xu Y., Huang Y., Weng X., Traye I., Tang S., Jiang X., Zhu K., Zhang W., Zhang H., Wang Z., Yang J. (2025): Cooking and eating quality of rice as affected by nitrogen fertilizers and alternate wetting and moderate drying irrigation at panicle stage. Agricultural Water Management, 318: 109715. Go to original source...
  65. Yang X., Li Y., Ren B., Ding L., Gao C., Shen Q., Guo S. (2012): Drought-induced root aerenchyma formation restricts water uptake in rice seedings supplied with nitrate. Plant Cell Physiology, 53: 495-504. Go to original source... Go to PubMed...
  66. Ye M., Wang Z., Wu M., Li H., Gu J., Yang J., Zhang H., Zhang Z. (2024): Optimized leaf anatomy improves photosynthetic producing capacity of mid-season indica rice in the Yangtze River Basin during the genetic improvement. European Journal of Agronomy, 158: 127196. Go to original source...
  67. York L.M., Galindo-Castañeda T., Schussler J.R., Lynch J.P. (2015): Evolution of US maize (Zea mays L.) root architectural and anatomical phenes over the past 100 years corresponds to increased tolerance of nitrogen stress. Journal of Experimental Botany, 66: 2347-2358. Go to original source... Go to PubMed...
  68. Zhang D.K., Li Y.Y., Li H.R., Li H.J., Zhao X.H., Cao J., Xu G.W. (2025): Phosphorus application rates affect the grain yields of different phosphorus-tolerant rice cultivars by regulating grain filling and leaf senescence characteristics. Plant, Soil and Environment, 71: 363-380. Go to original source...
  69. Zhang X., Xing Y., Wang Q., Yan G., Wang M., Liu G., Wang H., Huang B., Zhang J. (2020): Effects of long-term nitrogen addition and decreased precipitation on the fine root morphology and anatomy of the main tree species in a temperate forest. Forest Ecology and Management, 455: 117664. Go to original source...
  70. Zhang J., Zhang Y.Y., Song N.Y., Chen Q.L., Sun H.Z., Peng T., Huang S., Zhao Q.Z. (2021): Response of grain-filling rate and grain quality of mid-season indica rice to nitrogen application. Journal of Integrative Agriculture, 20: 1465-1473. Go to original source...
  71. Zhao T., Wang J., Li R., Zhang P., Guo X., Qi Y., Li Y., Cheng S., Ji J., He A., Ai Z. (2025): Effects of nitrogen fertilizer types and planting density on the yield and nitrogen use efficiency of salt-tolerant rice under salt stress conditions. Plants, 14: 501. Go to original source... Go to PubMed...
  72. Zhen B., Li H., Niu Q., Qiu H., Tian G., Lu H., Zhou X. (2020): Effects of combined high temperature and waterlogging stress at booting stage on root anatomy of rice (Oryza sativa L.). Water, 12: 2524. Go to original source...
  73. Zhou M., Yang J. (2023): Delaying or promoting? Manipulation of leaf senescence to improve crop yield and quality. Planta, 258: 48. Go to original source... Go to PubMed...
  74. Zhu D.W., Zhang H.C., Guo B.W., Xu K., Dai Q.G., Wei H.Y., Gao H., Hu Y.J., Cui P.Y., Huo Z.Y. (2016): Effects of nitrogen level on yield and quality of japonica soft super rice. Journal of Integrative Agriculture, 16: 1018-1027. Go to original source...
  75. Zhu S., Sun S., Zhao W., Yang X., Chen Z., Mao H., Sheng L. (2024): Comprehensive physiology and proteomics analysis revealed the resistance mechanism of rice (Oryza sativa L.) to cadmium stress. Ecotoxicology and Environmental Safety, 278: 116413. Go to original source... Go to PubMed...

This is an open access article distributed under the terms of the Creative Commons Attribution 4.0 International License (CC BY 4.0), which permits use, distribution, and reproduction in any medium, provided the original publication is properly cited. No use, distribution or reproduction is permitted which does not comply with these terms.