9.9
CiteScore
7.1
Impact Factor
Turn off MathJax
Article Contents

The crosstalk between nitrogen utilization and abiotic stress tolerance in rice

doi: 10.1016/j.jgg.2026.01.010
Funds:

This work was supported by the National Natural Science Foundation of China (32061143039), the Zhejiang Provincial Natural Science Foundation of China (LQK26C130004), the Biological Breeding-National Science and Technology Major Project (2023ZD04072), the Agricultural Science and Technology Innovation Program of Chinese Academy of Agricultural Sciences (CAAS-CSNCB-202301 and CAAS-ZDRW202401), and the Central Public-interest Scientific Institution Basal Research Fund (CPSIBRF-CNRRI-202404).

  • Received Date: 2025-10-24
  • Accepted Date: 2026-01-26
  • Rev Recd Date: 2026-01-22
  • Available Online: 2026-02-03
  • Improving nitrogen use efficiency (NUE) in rice is crucial for sustainable agriculture, yet remains a significant challenge due to its complex polygenic and environmental regulation. Although multiple NUE-associated genes have been identified, their intricate regulatory networks are poorly understood, especially under abiotic stresses such as drought, salinity, and extreme temperatures. This review systematically summarizes the genetic basis of NUE in rice, covering key genes involved in nitrogen uptake, translocation, assimilation, and remobilization. It further explores the crosstalk between nitrogen utilization and abiotic stress tolerance, highlighting integrative signaling nodes such as the dual nitrate/ABA receptor OsNRT1.1B. Finally, a comprehensive strategy is proposed to develop elite rice varieties with high NUE and multi-stress resilience, supporting the advancement of resource-efficient and climate-smart agriculture.
  • loading
  • Alfatih, A., Wu, J., Zhang, Z.S., Xia, J.Q., Jan, S.U., Yu, L.H., Xiang, C.B., 2020. Rice NIN-LIKE PROTEIN 1 rapidly responds to nitrogen deficiency and improves yield and nitrogen use efficiency. J. Exp. Bot. 71, 6032-6042.
    Alfatih, A., Zhang, J., Song, Y., Jan, S.U., Zhang, Z.S., Xia, J.Q., Zhang, Z.Y., Nazish, T., Wu, J., Zhao, P.X., Xiang, C.B., 2023. Nitrate-responsive OsMADS27 promotes salt tolerance in rice. Plant Commun. 4, 100458.
    Alvarez-Buylla, E.R., Garcia-Ponce, B., Sanchez, M.P., Espinosa-Soto, C., Garcia-Gomez, M.L., Pineyro-Nelson, A., Garay-Arroyo, A., 2019. MADS-box genes underground becoming mainstream: plant root developmental mechanisms. New Phytol. 223, 1143-1158.
    Cai, H., Zhou, Y., Xiao, J., Li, X., Zhang, Q., Lian, X., 2009. Overexpressed glutamine synthetase gene modifies nitrogen metabolism and abiotic stress responses in rice. Plant Cell Rep. 28, 527-537.
    Cairns, J.E., Impa, S.M., O'Toole, J.C., Jagadish, S.V.K., Price, A.H., 2011. Influence of the soil physical environment on rice (Oryza sativa L.) response to drought stress and its implications for drought research. Field Crops Res. 121, 303-310.
    Cao, X., Zhong, C., Zhu, L., Zhang, J., Hussain, S., Wu, L., Jin, Q., 2017. Glycine increases cold tolerance in rice via the regulation of N uptake, physiological characteristics, and photosynthesis. Plant Physiol. Biochem. 112, 251-260.
    Chen, H., Xu, N., Wu, Q., Yu, B., Chu, Y., Li, X., Huang, J., Jin, L., 2018a. OsMADS27 regulates the root development in a NO3−-Dependent manner and modulates the salt tolerance in rice (Oryza sativa L.). Plant Sci. 277, 20-32.
    Chen, H., Zhang, Q., Lu, Z., Xu, F., 2018b. Accumulation of ammonium and reactive oxygen mediated drought-induced rice growth inhibition by disturbed nitrogen metabolism and photosynthesis. Plant Soil 431, 107-117.
    Chen, J., Qi, T., Hu, Z., Fan, X., Zhu, L., Iqbal, M.F., Yin, X., Xu, G., Fan, X., 2019a. OsNAR2.1 positively regulates drought tolerance and grain yield under drought stress conditions in rice. Front. Plant Sci. 10, 197.
    Chen, L., Zhao, Y., Xu, S., Zhang, Z., Xu, Y., Zhang, J., Chong, K., 2018c. OsMADS57 together with OsTB1 coordinates transcription of its target OsWRKY94 and D14 to switch its organogenesis to defense for cold adaptation in rice. New Phytol. 218, 219-231.
    Chen, T., Yang, S., Wei, T., Li, Y., Wang, S., Su, Y., 2023. Overexpression of OsGS1;2 for improved nitrogen use efficiency and grain yield of rice: A field test. Field Crops Res. 303, 109146.
    Chen, X., Jiang, L., Zheng, J., Chen, F., Wang, T., Wang, M., Tao, Y., Wang, H., Hong, Z., Huang, Y., Huang, R., 2019b. A missense mutation in Large Grain Size 1 increases grain size and enhances cold tolerance in rice. J. Exp. Bot. 70, 3851-3866.
    Chen, Y., Liu, Y., Ge, J., Li, R., Zhang, R., Zhang, Y., Huo, Z., Xu, K., Wei, H., Dai, Q., 2022. Improved physiological and morphological traits of root synergistically enhanced salinity tolerance in rice under appropriate nitrogen application rate. Front. Plant Sci. 13, 982637.
    Cui, L.-G., Shan, J.-X., Shi, M., Gao, J.-P., Lin, H.-X., 2015. DCA1 acts as a transcriptional co-activator of DST and contributes to drought and salt tolerance in rice. PLoS Genet. 11, e1005617.
    Delgado, L.D., Nunez-Pascual, V., Riveras, E., Ruffel, S., Gutierrez, R.A., 2024. Recent advances in local and systemic nitrate signaling in Arabidopsis thaliana. Curr. Opin. Plant Biol. 81, 102605.
    Deng, P., Jing, W., Cao, C., Sun, M., Chi, W., Zhao, S., Dai, J., Shi, X., Wu, Q., Zhang, B., et al., 2022. Transcriptional repressor RST1 controls salt tolerance and grain yield in rice by regulating gene expression of asparagine synthetase. Proc. Natl. Acad. Sci. U. S. A. 119, e2210338119.
    Dey, A., Samanta, M.K., Gayen, S., Maiti, M.K., 2016. The sucrose non-fermenting 1-related kinase 2 gene SAPK9 improves drought tolerance and grain yield in rice by modulating cellular osmotic potential, stomatal closure and stress-responsive gene expression. BMC Plant Biol. 16, 158.
    Ding, L., Gao, C., Li, Y., Li, Y., Zhu, Y., Xu, G., Shen, Q., Kaldenhoff, R., Kai, L., Guo, S., 2015. The enhanced drought tolerance of rice plants under ammonium is related to aquaporin (AQP). Plant Sci. 234, 14-21.
    Ding, L., Lu, Z., Gao, L., Guo, S., Shen, Q., 2018. Is nitrogen a key determinant of water transport and photosynthesis in higher plants upon drought stress? Front. Plant Sci. 9, 1143.
    Ding, Y., Shi, Y., Yang, S., 2024. Regulatory networks underlying plant responses and adaptation to cold stress. Annu. Rev. of Genet. 58, 43-65.
    Ding, Y., Yang, H., Wu, S., Fu, D., Li, M., Gong, Z., Yang, S., 2022. CPK28-NLP7 module integrates cold-induced Ca2+ signal and transcriptional reprogramming in Arabidopsis. Sci. Adv. 8, eabn7901.
    Dwivedi, S.L., Ceccarelli, S., Blair, M.W., Upadhyaya, H.D., Are, A.K., Ortiz, R., 2016. Landrace germplasm for improving yield and abiotic stress adaptation. Trends Plant Sci. 21, 31-42.
    El-Kereamy, A., Bi, Y.-M., Ranathunge, K., Beatty, P.H., Good, A.G., Rothstein, S.J., 2012. The rice R2R3-MYB transcription factor OsMYB55 is involved in the tolerance to high temperature and modulates amino acid metabolism. PLoS ONE 7, e52030.
    Fang, Z., Wu, B., Ji, Y., 2021. The amino acid transporter OsAAP4 contributes to rice tillering and grain yield by regulating neutral amino acid allocation through two splicing variants. Rice 14, 2.
    Farooq, M.A., Gao, S., Hassan, M.A., Huang, Z., Rasheed, A., Hearne, S., Prasanna, B., Li, X., Li, H., 2024. Artificial intelligence in plant breeding. Trends Genet. 40, 891-908.
    Fei, H., Li, Y., Liu, Y., Wei, J., Chen, A., Gao, C., 2025. Advancing protein evolution with inverse folding models integrating structural and evolutionary constraints. Cell 188, 4674-4692.
    Fujimura, S., Nishimura, T., Hasegawa, T., 2012. Nitrogen uptake by rice (Oryza sativa L.) exposed to low water temperatures at different growth stages. J. Agron. Crop Sci. 198, 145-151.
    Fukai, S., Cooper, M., 1995. Development of drought-resistant cultivars using physiomorphological traits in rice. Field Crops Res. 40, 67-86.
    Funayama, K., Kojima, S., Tabuchi-Kobayashi, M., Sawa, Y., Nakayama, Y., Hayakawa, T., Yamaya, T., 2013. Cytosolic glutamine synthetase1;2 is responsible for the primary assimilation of ammonium in rice roots. Plant Cell Physiol. 54, 934-943.
    Ganie, S.A., Molla, K.A., Henry, R.J., Bhat, K.V., Mondal, T.K., 2019. Advances in understanding salt tolerance in rice. Theor. Appl. Genet. 132, 851-870.
    Ganie, S.A., Wani, S.H., Henry, R., Hensel, G., 2021. Improving rice salt tolerance by precision breeding in a new era. Curr. Opin. Plant Biol‌. 60, 101996.
    Gao, C., 2021. Genome engineering for crop improvement and future agriculture. Cell 184, 1621-1635.
    Gao, L., Liu, M., Wang, M., Shen, Q., Guo, S., 2016. Enhanced salt tolerance under nitrate nutrition is associated with apoplast Na+ content in canola (Brassica. napus L.) and rice (Oryza sativa L.) plants. Plant Cell Physiol. 57, 2323-2333.
    Gao, Y., Li, Y., Yang, X., Li, H., Shen, Q., Guo, S., 2010. Ammonium nutrition increases water absorption in rice seedlings (Oryza sativa L.) under water stress. Plant Soil 331, 193-201.
    Gao, Y., Xu, Z., Zhang, L., Li, S., Wang, S., Yang, H., Liu, X., Zeng, D., Liu, Q., Qian, Q., et al., 2020. MYB61 is regulated by GRF4 and promotes nitrogen utilization and biomass production in rice. Nat. Commun. 11, 5219.
    Gao, Z., Wang, Y., Chen, G., Zhang, A., Yang, S., Shang, L., Wang, D., Ruan, B., Liu, C., Jiang, H., et al., 2019. The indica nitrate reductase gene OsNR2 allele enhances rice yield potential and nitrogen use efficiency. Nat. Commun. 10, 5207.
    Gautam, H., Sehar, Z., Rehman, M.T., Hussain, A., AlAjmi, M.F., Khan, N.A., 2021. Nitric oxide enhances photosynthetic nitrogen and sulfur-use efficiency and activity of ascorbate-glutathione cycle to reduce high temperature stress-induced oxidative stress in rice (Oryza sativa L.) plants. Biomolecules 11, 305.
    Giri, A., Heckathorn, S., Mishra, S., Krause, C., 2017. Heat stress decreases levels of nutrient-uptake and-assimilation proteins in tomato roots. Plants 6, 6.
    Gong, Z., Xiong, L., Shi, H., Yang, S., Herrera-Estrella, L.R., Xu, G., Chao, D.-Y., Li, J., Wang, P.-Y., Qin, F., et al., 2020. Plant abiotic stress response and nutrient use efficiency. Sci. China Life Sci. 63, 635-674.
    Guo, L., Lu, Y., Bao, S., Zhang, Q., Geng, Y., Shao, X., 2021. Carbon and nitrogen metabolism in rice cultivars affected by salt-alkaline stress. Crop Pasture Sci. 72, 372-382.
    Guo, S.-Q., Chen, Y.-X., Ju, Y.-L., Pan, C.-Y., Shan, J.-X., Ye, W.-W., Dong, N.-Q., Kan, Y., Yang, Y.-B., Zhao, H.-Y., et al., 2025. Fine-tuning gibberellin improves rice alkali-thermal tolerance and yield. Nature 639, 162-171.
    Guo, S., Chen, G., Zhou, Y., Shen, Q., 2007. Ammonium nutrition increases photosynthesis rate under water stress at early development stage of rice (Oryza sativa L.). Plant Soil 296, 115-124.
    Gupta, K.J., Fernie, A.R., Kaiser, W.M., van Dongen, J.T., 2011. On the origins of nitric oxide. Trends Plant Sci. 16, 160-168.
    Han, M.L., Lv, Q.Y., Zhang, J., Wang, T., Zhang, C.X., Tan, R.J., Wang, Y.L., Zhong, L.Y., Gao, Y.Q., Chao, Z.F., et al., 2022. Decreasing nitrogen assimilation under drought stress by suppressing DST-mediated activation of Nitrate Reductase 1.2 in rice. Mol. Plant 15, 167-178.
    Hoshida, H., Tanaka, Y., Hibino, T., Hayashi, Y., Tanaka, A., Takabe, T., Takabe, T., 2000. Enhanced tolerance to salt stress in transgenic rice that overexpresses chloroplast glutamine synthetase. Plant Mol. Biol. 43, 103-111.
    Hu, B., Wang, W., Ou, S., Tang, J., Li, H., Che, R., Zhang, Z., Chai, X., Wang, H., Wang, Y., et al., 2015. Variation in NRT1.1B contributes to nitrate-use divergence between rice subspecies. Nat. Genet. 47, 834-838.
    Hu, Q., Yan, N., Cui, K., Li, G., Wang, W., Huang, J., Peng, S., 2024. Increased panicle nitrogen application improves rice yield by alleviating high-temperature damage during panicle initiation to anther development. Physiol. Plantarum 176, e14230.
    Huang, G., Zhang, Q., Wei, X., Peng, S., Li, Y., 2017. Nitrogen can alleviate the inhibition of photosynthesis caused by high temperature stress under both steady-state and flecked irradiance. Front. Plant Sci. 8, 945.
    Huang, S., Liang, Z., Chen, S., Sun, H., Fan, X., Wang, C., Xu, G., Zhang, Y., 2019. A transcription factor, OsMADS57, regulates long-distance nitrate transport and root elongation. Plant Physiol. 180, 882-895.
    Huang, X.Y., Chao, D.Y., Gao, J.P., Zhu, M.Z., Shi, M., Lin, H.X., 2009. A previously unknown zinc finger protein, DST, regulates drought and salt tolerance in rice via stomatal aperture control. Gene. Dev. 23, 1805-1817.
    Jagadish, S.V., Murty, M.V., Quick, W.P., 2015. Rice responses to rising temperatures--challenges, perspectives and future directions. Plant Cell Environ. 38, 1686-1698.
    James, D., Borphukan, B., Fartyal, D., Ram, B., Singh, J., Manna, M., Sheri, V., Panditi, V., Yadav, R., Achary, V.M.M., et al., 2018. Concurrent overexpression of OsGS1;1 and OsGS2 genes in transgenic rice (Oryza sativa L.): impact on tolerance to abiotic stresses. Front. Plant Sci. 9, 786.
    Ji, P., Xu, C., Ling, F., Li, X., Qi, Z., Chen, Y., Liu, X., Zhang, Z., Wang, J., Luo, Z., et al., 2025. Improvement in nitrogen-use efficiency increases salt stress tolerance in rice seedlings and grain yield in salinized soil. Plants (Basel) 14, 556.
    Ji, Y., Huang, W., Wu, B., Fang, Z., Wang, X., 2020. The amino acid transporter AAP1 mediates growth and grain yield by regulating neutral amino acid uptake and reallocation in Oryza sativa. J. Exp. Bot. 71, 4763-4777.
    Jia, Y., Liu, H., Qu, Z., Wang, J., Wang, X., Wang, Z., Yang, L., Zhang, D., Zou, D., Zhao, H., 2020. Transcriptome sequencing and iTRAQ of different rice cultivars provide insight into molecular mechanisms of cold-tolerance response in japonica rice. Rice 13, 43.
    Jia, Y., Qin, D., Zheng, Y., Wang, Y., 2023. Finding balance in adversity: Nitrate signaling as the key to plant growth, resilience, and stress response. Int. J. Mol. Sci. 24, 14406.
    Jia, Y., Wang, J., Qu, Z., Zou, D., Sha, H., Liu, H., Sun, J., Zheng, H., Yang, L., Zhao, H., 2019. Effects of low water temperature during reproductive growth on photosynthetic production and nitrogen accumulation in rice. Field Crops Res. 242, 107587.
    Jin, F., Xie, P., Li, Z., Wu, B., Huang, W., Fang, Z., 2024. Blocking of amino acid transporter OsAAP7 promoted tillering and yield by determining basic and neutral amino acids accumulation in rice. BMC Plant Bio. 24, 447.
    Kim, Y., Chung, Y.S., Lee, E., Tripathi, P., Heo, S., Kim, K.-H., 2020. Root response to drought stress in rice (Oryza sativa L.). Int. J. Mol. Sci. 21, 1513.
    Kobayashi, Y., Yamamoto, S., Minami, H., Kagaya, Y., Hattori, T., 2004. Differential activation of the rice sucrose nonfermenting1-related protein kinase 2 family by hyperosmotic stress and abscisic acid. Plant Cell 16, 1163-1177.
    Konishi, N., Ma, J.F., 2021. Three polarly localized ammonium transporter 1 members are cooperatively responsible for ammonium uptake in rice under low ammonium condition. New Phytol. 232, 1778-1792.
    Lamig, L., Moreno, S., Alvarez, J.M., Gutierrez, R.A., 2022. Molecular mechanisms underlying nitrate responses in plants. Curr. Biol. 32, R433-R439.
    Li, B., Sun, C., Li, J., Gao, C., 2024a. Targeted genome-modification tools and their advanced applications in crop breeding. Nat. Rev. Genet. 25, 603-622.
    Li, G., An, L., Yang, W., Yang, L., Wei, T., Shi, J., Wang, J., Doonan, J.H., Xie, K., Fernie, A.R., et al., 2025a. Integrated biotechnological and AI innovations for crop improvement. Nature 643, 925-937.
    Li, J., Zhang, Z., Chong, K., Xu, Y., 2022a. Chilling tolerance in rice: Past and present. J. Plant Physiol. 268, 153576.
    Li, J.Y., Yang, C., Xu, J., Lu, H.P., Liu, J.X., 2023. The hot science in rice research: How rice plants cope with heat stress. Plant Cell Environ. 46, 1087-1103.
    Li, L., Huang, Z., Wu, S., Zhang, Y., Mu, Y., Li, Y., He, A., Ai, Z., Guo, X., Nie, L., 2025b. The mechanisms of improved grain yield and nitrogen use efficiency in salt-tolerant rice varieties under salt stress. J. Agron. Crop Sci. 211, e70064.
    Li, Q., Lu, X., Wang, C., Shen, L., Dai, L., He, J., Yang, L., Li, P., Hong, Y., Zhang, Q., et al., 2022b. Genome-wide association study and transcriptome analysis reveal new QTL and candidate genes for nitrogen-deficiency tolerance in rice. Crop J. 10, 942-951.
    Li, S., Tian, Y., Wu, K., Ye, Y., Yu, J., Zhang, J., Liu, Q., Hu, M., Li, H., Tong, Y., et al., 2018. Modulating plant growth-metabolism coordination for sustainable agriculture. Nature 560, 595-600.
    Li, W., Liu, J., Li, Z., Ye, R., Chen, W., Huang, Y., Yuan, Y., Zhang, Y., Hu, H., Zheng, P., et al., 2024b. Mitigating growth-stress tradeoffs via elevated TOR signaling in rice. Mol. Plant 17, 240-257.
    Liang, C.-G., Yan, W., Jia, L., Tian, L., 2011. High temperature at grain-filling stage affects nitrogen metabolism enzyme activities in grains and grain nutritional quality in rice. Rice Sci. 18, 210-216.
    Liang, X., Li, J., Yang, Y., Jiang, C., Guo, Y., 2024. Designing salt stress-resilient crops: Current progress and future challenges. J. Integr. Plant Biol. 66, 303-329.
    Liu, K., Deng, J., Lu, J., Wang, X., Lu, B., Tian, X., Zhang, Y., 2019a. High nitrogen levels alleviate yield loss of super hybrid rice caused by high temperatures during the flowering stage. Front. Plant Sci. 10, 357.
    Liu, X., Hu, B., Chu, C., 2022a. Nitrogen assimilation in plants: current status and future prospects. J. Genet. Genomics 49, 394-404.
    Liu, X., Tian, Y., Chi, W., Zhang, H., Yu, J., Chen, G., Wu, W., Jiang, X., Wang, S., Lin, Z., et al., 2022b. Alternative splicing of OsGS1;1 affects nitrogen-use efficiency, grain development, and amylose content in rice. Plant J. 110, 1751-1762.
    Liu, X.X., Zhu, Y.X., Fang, X.Z., Ye, J.Y., Du, W.X., Zhu, Q.Y., Lin, X.Y., Jin, C.W., 2020. Ammonium aggravates salt stress in plants by entrapping them in a chloride over-accumulation state in an NRT1.1-dependent manner. Sci. Total Environ. 746, 141244.
    Liu, Y., Wang, H., Jiang, Z., Wang, W., Xu, R., Wang, Q., Zhang, Z., Li, A., Liang, Y., Ou, S., et al., 2021. Genomic basis of geographical adaptation to soil nitrogen in rice. Nature 590, 600-605.
    Liu, Z.-L., Li, Y.-J., Hou, H.-Y., Zhu, X.-C., Rai, V., He, X.-Y., Tian, C.-J., 2013. Differences in the arbuscular mycorrhizal fungi-improved rice resistance to low temperature at two N levels: aspects of N and C metabolism on the plant side. Plant Physiol. Bioch. 71, 87-95.
    Liu, Z., Tao, L., Liu, T., Zhang, X., Wang, W., Song, J., Yu, C., Peng, X., 2019b. Nitrogen application after low-temperature exposure alleviates tiller decrease in rice. Environ. Exp. Bot. 158, 205-214.
    Lou, D., Chen, Z., Yu, D., Yang, X., 2020. SAPK2 contributes to rice yield by modulating nitrogen metabolic processes under reproductive stage drought stress. Rice 13, 35.
    Lou, D., Wang, H., Liang, G., Yu, D., 2017. OsSAPK2 confers abscisic acid sensitivity and tolerance to drought stress in rice. Front. Plant Sci. 8, 993.
    Lu, K., Wu, B., Wang, J., Zhu, W., Nie, H., Qian, J., Huang, W., Fang, Z., 2018. Blocking amino acid transporter OsAAP3 improves grain yield by promoting outgrowth buds and increasing tiller number in rice. Plant Biotechnol. J. 16, 1710-1722.
    Ma, H., Jia, Y., Wang, W., Wang, J., Zou, D., Wang, J., Gong, W., Han, Y., Dang, Y., Wang, J., et al., 2025a. Effects of low-temperature stress during the grain-filling stage on carbon-nitrogen metabolism and grain yield formation in rice. Agronomy 15, 417.
    Ma, X., Wang, W., Zhang, J., Jiang, Z., Xu, C., Zhu, W., Shi, B., Yang, W., Su, H., Wang, X., et al., 2025b. NRT1.1B acts as an abscisic acid receptor in integrating compound environmental cues for plants. Cell 88, 5231-5248.
    Mackay, T.F., 2014. Epistasis and quantitative traits: using model organisms to study gene-gene interactions. Nat. Rev. Genet. 15, 22-33.
    Mao, D., Chen, C., 2012. Colinearity and similar expression pattern of rice DREB1s reveal their functional conservation in the cold-responsive pathway. PLoS ONE 7, e47275.
    Miao, J., Bu, L., Tan, W., Wang, P., Li, X., Li, X., Chen, C., Zhang, K., Shen, W., Gong, Z., 2024. OsPP2C49, a negative regulatory factor in the abscisic acid signaling pathway, positively regulates grain yield in rice. Rice 17, 65.
    Mo, Y., Li, G., Liu, L., Zhang, Y., Li, J., Yang, M., Chen, S., Lin, Q., Fu, G., Zheng, D., Ling, Y., 2023. OsGRF4AA compromises heat tolerance of developing pollen grains in rice. Front. Plant Sci. 14, 1121852.
    Montesinos-Lopez, O.A., Montesinos-Lopez, A., Perez-Rodriguez, P., Barron-Lopez, J.A., Martini, J.W.R., Fajardo-Flores, S.B., Gaytan-Lugo, L.S., Santana-Mancilla, P.C., Crossa, J., 2021. A review of deep learning applications for genomic selection. BMC Genomics 22, 19.
    Mu, X., Luo, J., 2019. Evolutionary analyses of NIN-like proteins in plants and their roles in nitrate signaling. Cell. Mol. Life Sci. 76, 3753-3764.
    Mukhopadhyay, P., Tyagi, A.K., 2015. OsTCP19 influences developmental and abiotic stress signaling by modulating ABI4-mediated pathways. Sci. Rep. 5, 9998.
    Nguyen, H.T.T., Shim, I.S., Kobayashi, K., Usui, K., 2005. Regulation of ammonium accumulation during salt stress in rice (Oryza sativa L.) seedlings. Plant Prod. Sci. 8, 397-404.
    Nie, S., Huang, W., He, C., Wu, B., Duan, H., Ruan, J., Zhao, Q., Fang, Z., 2025. Transcription factor OsMYB2 triggers amino acid transporter OsANT1 expression to regulate rice growth and salt tolerance. Plant Physiol. 197, kiae559.
    Ohashi, M., Ishiyama, K., Kusano, M., Fukushima, A., Kojima, S., Hayakawa, T., Yamaya, T., 2018. Reduction in sucrose contents by downregulation of fructose-1,6-bisphosphatase 2 causes tiller outgrowth cessation in rice mutants lacking glutamine synthetase1;2. Rice 11, 65.
    Phan, N.T.H., Heymans, A., Bonnave, M., Lutts, S., Pham, C.V., Bertin, P., 2023. Nitrogen use efficiency of rice cultivars (Oryza sativa l.) under salt stress and low nitrogen conditions. J. Plant Growth Regul. 42, 1789-1803.
    Pradhan, S.K., Pandit, E., Nayak, D.K., Behera, L., Mohapatra, T., 2019. Genes, pathways and transcription factors involved in seedling stage chilling stress tolerance in indica rice through RNA-Seq analysis. BMC Plant Biol. 19, 352.
    Rajan, H., Ganesan, K.N., Manonmani, S., Gopalakrishnan, C., Senthilkumar, G., Suresh, R., 2025. Unraveling the genetic potential of wild rice (Oryza rufipogon) for sustainable food security. Gene. Resour. Crop Ev. 72, 7639-7663.
    Ranathunge, K., El-Kereamy, A., Gidda, S., Bi, Y.-M., Rothstein, S.J., 2014. AMT1; 1 transgenic rice plants with enhanced NH4+ permeability show superior growth and higher yield under optimal and suboptimal NH4+ conditions. J. Exp. Bot. 65, 965-979.
    Reguera, M., Peleg, Z., Abdel-Tawab, Y.M., Tumimbang, E.B., Delatorre, C.A., Blumwald, E., 2013. Stress-induced cytokinin synthesis increases drought tolerance through the coordinated regulation of carbon and nitrogen assimilation in rice. Plant Physiol. 163, 1609-1622.
    Sade, N., Umnajkitikorn, K., Rubio Wilhelmi, M.D.M., Wright, M., Wang, S., Blumwald, E., 2018. Delaying chloroplast turnover increases water-deficit stress tolerance through the enhancement of nitrogen assimilation in rice. J. Exp. Bot. 69, 867-878.
    Sangjan, W., Kick, D.R., Washburn, J.D., 2025. Improving plant breeding through AI-supported data integration. Theor. Appl. Genet. 138, 132.
    Sathee, L., Jha, S.K., Rajput, O.S., Singh, D., Kumar, S., Kumar, A., 2021. Expression dynamics of genes encoding nitrate and ammonium assimilation enzymes in rice genotypes exposed to reproductive stage salinity stress. Plant Physiol. Bioch. 165, 161-172.
    Sehrawat, A., Gupta, R., Deswal, R., 2013. Nitric oxide-cold stress signalling cross-talk, evolution of a novel regulatory mechanism. Proteomics 13, 1816-1835.
    Shahzad, N., Nabi, H.G., Qiao, L., Li, W., 2024. The molecular mechanism of cold-stress tolerance: Cold responsive genes and their mechanisms in rice (Oryza sativa L.). Biology 13, 442.
    Singh, K.K., Ghosh, S., 2013. Regulation of glutamine synthetase isoforms in two differentially drought-tolerant rice (Oryza sativa L.) cultivars under water deficit conditions. Plant Cell Rep. 32, 183-193.
    Siringam, K., Juntawong, N., Cha-Um, S., Kirdmanee, C., 2013. Exogenous application of potassium nitrate to alleviate salt stress in rice seedlings. J. Plant Nutr. 36, 607-616.
    Song, L., Yue, L., Zhao, H., Hou, M., 2013. Protection effect of nitric oxide on photosynthesis in rice under heat stress. Acta Physiol. Plant 35, 3323-3333.
    Sun, C., Li, H., Liu, Y., Li, Y., Gao, R., Shi, X., Fei, H., Liu, J., Liang, R., Gao, C., 2025a. Iterative recombinase technologies for efficient and precise genome engineering across kilobase to megabase scales. Cell 188, 4693-4710.
    Sun, L., Di, D.W., Li, G., Kronzucker, H.J., Wu, X., Shi, W., 2020. Endogenous ABA alleviates rice ammonium toxicity by reducing ROS and free ammonium via regulation of the SAPK9-bZIP20 pathway. J. Exp. Bot. 71, 4562-4577.
    Sun, T., Liu, B., Hasegawa, T., Liao, Z., Tang, L., Liu, L., Cao, W., Zhu, Y., 2023. Sink-source unbalance leads to abnormal partitioning of biomass and nitrogen in rice under extreme heat stress: An experimental and modeling study. Eur. J. Agron. 142, 126678.
    Sun, T., Ruan, J., Cao, T., Yao, L., Zhao, Z., Zhang, J., Li, J., Deng, A., Chen, H., Gao, X., et al., 2025b. Effects of low-temperature stress during rice heading stage on carbon and nitrogen allocation in paddy eco-system of northeastern China. Front. Plant Sci. 16, 1484734.
    Tamura, W., Kojima, S., Toyokawa, A., Watanabe, H., Tabuchi-Kobayashi, M., Hayakawa, T., Yamaya, T., 2011. Disruption of a novel NADH-Glutamate Synthase 2 gene caused marked reduction in spikelet number of rice. Front. Plant Sci. 2, 57.
    Tang, S., Zhang, H., Liu, W., Dou, Z., Zhou, Q., Chen, W., Wang, S., Ding, Y., 2019. Nitrogen fertilizer at heading stage effectively compensates for the deterioration of rice quality by affecting the starch-related properties under elevated temperatures. Food Chem. 277, 455-462.
    Teh, C.-Y., Shaharuddin, N.A., Ho, C.-L., Mahmood, M., 2016. Exogenous proline significantly affects the plant growth and nitrogen assimilation enzymes activities in rice (Oryza sativa) under salt stress. Acta Physiol. Plant 38, 151.
    Teng, Z., Zheng, Q., Peng, Y., Li, Y., Meng, S., Liu, B., Peng, Y., Duan, M., Yuan, D., Zhang, J., Ye, N., 2025. Nitrate reductase-dependent nitric oxide production mediates nitrate-conferred salt tolerance in rice seedlings. Plant Physiol. 197, kiaf080.
    Uchida, A., Jagendorf, A.T., Hibino, T., Takabe, T., Takabe, T., 2002. Effects of hydrogen peroxide and nitric oxide on both salt and heat stress tolerance in rice. Plant Sci. 163, 515-523.
    Wang, D., Liang, Z., Gu, C., Chang, Y., Zhang, J., Wu, Y., Zhang, Y., 2025a. The transcription factor OsMADS61 positively regulates both root elongation and nitrogen use efficiency in rice. Plant Commun. 4, 101630.
    Wang, H., Lu, S., Guan, X., Jiang, Y., Wang, B., Hua, J., Zou, B., 2022. Dehydration-responsive element binding protein 1C, 1E, and 1G promote stress tolerance to chilling, heat, drought, and salt in rice. Front. Plant Sci. 13, 851731.
    Wang, H., Zhang, M., Guo, R., Shi, D., Liu, B., Lin, X., Yang, C., 2012. Effects of salt stress on ion balance and nitrogen metabolism of old and young leaves in rice (Oryza sativa L.). BMC Plant Biol. 12, 194.
    Wang, H., Zhong, L., Fu, X., Huang, S., Zhao, D., He, H., Chen, X., 2023. Physiological analysis reveals the mechanism of accelerated growth recovery for rice seedlings by nitrogen application after low temperature stress. Front. Plant Sci. 14, 1133592.
    Wang, J., Wu, B., Lu, K., Wei, Q., Qian, J., Chen, Y., Fang, Z., 2019. The Amino Acid Permease 5 (OsAAP5) regulates tiller number and grain yield in rice. Plant Physiol. 180, 1031-1045.
    Wang, L., Ju, C., Han, C., Yu, Z., Bai, M.Y., Wang, C., 2025b. The interaction of nutrient uptake with biotic and abiotic stresses in plants. J. Integr. Plant Biol. 67, 455-487.
    Wei, S., Li, X., Lu, Z., Zhang, H., Ye, X., Zhou, Y., Li, J., Yan, Y., Pei, H., Duan, F., et al., 2022. A transcriptional regulator that boosts grain yields and shortens the growth duration of rice. Science 377, eabi8455.
    Wu, J., Yang, S., Chen, N., Jiang, Q., Huang, L., Qi, J., Xu, G., Shen, L., Yu, H., Fan, X., 2023. Nuclear translocation of OsMADS25 facilitated by OsNAR2.1 in reponse to nitrate signals promotes rice root growth by targeting OsMADS27 and OsARF7. Plant Commun. 4, 100642.
    Wu, J., Yang, S., Rehman, M., Chen, N., Liu, B., Xu, G., Fan, X., Gan, Y., 2025. Nuclear accumulation of OsMADS27 was promoted by OsNAR2.1 in response to nitrate to control root growth by directly targeting OsMADS57 in rice. Physiol. Plantarum 177, e70553.
    Wu, J., Yu, C., Huang, L., Gan, Y., 2021a. A rice transcription factor, OsMADS57, positively regulates high salinity tolerance in transgenic Arabidopsis thaliana and Oryza sativa plants. Physiol. Plantarum 173, 1120-1135.
    Wu, J., Zhang, Z.S., Xia, J.Q., Alfatih, A., Song, Y., Huang, Y.J., Wan, G.Y., Sun, L.Q., Tang, H., Liu, Y., et al., 2021b. Rice NIN-LIKE PROTEIN 4 plays a pivotal role in nitrogen use efficiency. Plant Biotechnol. J. 19, 448-461.
    Wu, K., Wang, S., Song, W., Zhang, J., Wang, Y., Liu, Q., Yu, J., Ye, Y., Li, S., Chen, J., et al., 2020. Enhanced sustainable green revolution yield via nitrogen-responsive chromatin modulation in rice. Science 367, eaaz2046.
    Wu, Y., Yang, W., Wei, J., Yoon, H., An, G., 2017. Transcription factor OsDOF18 controls ammonium uptake by inducing ammonium transporters in rice roots. Mol. Cells 40, 178-185.
    Xia, K., Pan, X., Chen, H., Xu, X., Zhang, M., 2023. Rice miR168a-5p regulates seed length, nitrogen allocation and salt tolerance by targeting OsOFP3, OsNPF2.4 and OsAGO1a, respectively. J. Plant Physiol. 280, 153905.
    Xin, W., Zheng, H., Yang, L., Xie, S., Xia, S., Wang, J., Wang, M., Liu, Y., Zou, D., Liu, H., et al., 2025. Genome-wide association studies identify OsNLP6 as a key regulator of nitrogen use efficiency in rice. Plant Biotechnol. J. 23, 5110-5125.
    Xu, J., Huang, X., Lan, H., Zhang, H., Huang, J., 2016. Rearrangement of nitrogen metabolism in rice (Oryza sativa L.) under salt stress. Plant signal Behav. 11, e1138194.
    Xu, N., Chu, Y., Chen, H., Li, X., Wu, Q., Jin, L., Wang, G., Huang, J., 2018. Rice transcription factor OsMADS25 modulates root growth and confers salinity tolerance via the ABA-mediated regulatory pathway and ROS scavenging. PLoS Genet. 14, e1007662.
    Xu, Y., Guan, X., Han, Z., Zhou, L., Zhang, Y., Asad, M.A., Wang, Z., Jin, R., Pan, G., Cheng, F., 2022. Combined effect of nitrogen fertilizer application and high temperature on grain quality properties of cooked rice. Front. Plant Sci. 13, 874033.
    Yamamoto, A., Shim, I.-S., Fujihara, S., 2012. Chilling-stress responses by rice seedlings grown with different ammonium concentrations and its relationship to leaf spermidine content. J. Plant Biol. 55, 191-197.
    Yan, M., Fan, X., Feng, H., Miller, A.J., Shen, Q., Xu, G., 2011. Rice OsNAR2.1 interacts with OsNRT2.1, OsNRT2.2 and OsNRT2.3a nitrate transporters to provide uptake over high and low concentration ranges. Plant Cell Environ. 34, 1360-1372.
    Yang, A., Dai, X., Zhang, W.H., 2012. A R2R3-type MYB gene, OsMYB2, is involved in salt, cold, and dehydration tolerance in rice. J. Exp. Bot. 63, 2541-2556.
    Yang, J., Shi, W., Xiao, G., Zhang, X., Wang, D., Xu, H., Wu, J., Yang, Z., Lai, Y., Duan, M., Zhang, J., 2022a. Optimum total nitrogen application is required to reduce the yield loss of hybrid rice to high temperature. Field Crops Res. 288, 108696.
    Yang, X., Nian, J., Xie, Q., Feng, J., Zhang, F., Jing, H., Zhang, J., Dong, G., Liang, Y., Peng, J., et al., 2016. Rice Ferredoxin-dependent Glutamate Synthase regulates nitrogen-carbon metabolomes and is genetically differentiated between japonica and indica subspecies. Mol. Plant 9, 1520-1534.
    Yang, Y., Guo, Y., 2018. Unraveling salt stress signaling in plants. J. Integr. Plant Biol. 60, 796-804.
    Yang, Y., Yu, J., Qian, Q., Shang, L., 2022b. Enhancement of heat and drought stress tolerance in rice by genetic manipulation: A systematic review. Rice 15, 67.
    Ye, J.Y., Tian, W.H., Jin, C.W., 2022. Nitrogen in plants: From nutrition to the modulation of abiotic stress adaptation. Stress Biol. 2, 4.
    Yi, Y., Peng, Y., Song, T., Lu, S., Teng, Z., Zheng, Q., Zhao, F., Meng, S., Liu, B., Peng, Y., et al., 2022. NLP2-NR module associated NO is involved in regulating seed germination in rice under salt stress. Plants (Basel) 11, 11060795.
    Yoshida, T., Mogami, J., Yamaguchi-Shinozaki, K., 2014. ABA-dependent and ABA-independent signaling in response to osmotic stress in plants. Curr. Opin. Plant Biol. 21, 133-139.
    Yu, C., Liu, Y., Zhang, A., Su, S., Yan, A., Huang, L., Ali, I., Liu, Y., Forde, B.G., Gan, Y., 2015. MADS-box transcription factor OsMADS25 regulates root development through affection of nitrate accumulation in rice. PLoS ONE 10, e0135196.
    Yu, C., Su, S., Xu, Y., Zhao, Y., Yan, A., Huang, L., Ali, I., Gan, Y., 2014a. The effects of fluctuations in the nutrient supply on the expression of five members of the AGL17 clade of MADS-box genes in rice. PLoS ONE 9, e105597.
    Yu, J., Xuan, W., Tian, Y., Fan, L., Sun, J., Tang, W., Chen, G., Wang, B., Liu, Y., Wu, W., et al., 2021. Enhanced OsNLP4-OsNiR cascade confers nitrogen use efficiency by promoting tiller number in rice. Plant Biotechnol. J. 19, 167-176.
    Yu, L.H., Miao, Z.Q., Qi, G.F., Wu, J., Cai, X.T., Mao, J.L., Xiang, C.B., 2014b. MADS-box transcription factor AGL21 regulates lateral root development and responds to multiple external and physiological signals. Mol. Plant 7, 1653-1669.
    Zayed, O., Hewedy, O.A., Abdelmoteleb, A., Ali, M., Youssef, M.S., Roumia, A.F., Seymour, D., Yuan, Z.C., 2023. Nitrogen journey in plants: From uptake to metabolism, stress response, and microbe interaction. Biomolecules 13, 1443.
    Zeng, D.D., Qin, R., Li, M., Alamin, M., Jin, X.L., Liu, Y., Shi, C.H., 2017. The ferredoxin-dependent glutamate synthase (OsFd-GOGAT) participates in leaf senescence and the nitrogen remobilization in rice. Mol. Genet. Genomics 292, 385-395.
    Zhang, B., Ma, L., Wu, B., Xing, Y., Qiu, X., 2022a. Introgression lines: Valuable resources for functional genomics research and breeding in rice (Oryza sativa L.). Front. Plant Sci. 13, 863789.
    Zhang, F., Zeng, D., Huang, L., Shi, Y., Chen, T., Zhang, F., Zhou, Y., 2019a. Stress-activated protein kinase OsSAPK9 regulates tolerance to salt stress and resistance to bacterial blight in rice. Rice 12, 80.
    Zhang, H., Forde, B.G., 1998. An Arabidopsis MADS box gene that controls nutrient-induced changes in root architecture. Science 279, 407-409.
    Zhang, H., Zhu, J., Gong, Z., Zhu, J.-K., 2022b. Abiotic stress responses in plants. Nat. Rev. Genet. 23, 104-119.
    Zhang, J., Liu, Y.-X., Zhang, N., Hu, B., Jin, T., Xu, H., Qin, Y., Yan, P., Zhang, X., Guo, X., et al., 2019b. NRT1.1B is associated with root microbiota composition and nitrogen use in field-grown rice. Nat. Biotechnol. 37, 676-684.
    Zhang, Y., Tateishi-Karimata, H., Endoh, T., Jin, Q., Li, K., Fan, X., Ma, Y., Gao, L., Lu, H., Wang, Z., et al., 2022c. High-temperature adaptation of an OsNRT2.3 allele is thermoregulated by small RNAs. Sci. Adv‌. 8, eadc9785.
    Zhang, Z., Chu, C., 2020. Nitrogen-use divergence between indica and japonica rice: variation at nitrate assimilation. Mol. Plant 13, 6-7.
    Zhang, Z.S., Xia, J.Q., Alfatih, A., Song, Y., Huang, Y.J., Sun, L.Q., Wan, G.Y., Wang, S.M., Wang, Y.P., Hu, B.H., et al., 2022d. Rice NIN-LIKE PROTEIN 3 modulates nitrogen use efficiency and grain yield under nitrate-sufficient conditions. Plant Cell Environ. 45, 1520-1536.
    Zhao, H., Ma, H., Yu, L., Wang, X., Zhao, J., 2012. Genome-wide survey and expression analysis of amino acid transporter gene family in rice (Oryza sativa L.). PloS one 7, e49210.
    Zhao, S., Xu, Z., Zhang, Z., Zhu, Z., Li, H., 2025a. Optimal nitrate supply improves plant thermotolerance through NLP7-mediated transcriptomic changes in Arabidopsis. Plant Commun. 6, 101485.
    Zhao, Y., Zhao, Y., Peng, Y., Sun, Y., Zhang, D., Zhang, C., Ran, X., Shen, Y., Liu, W., Ding, Y., Tang, S., 2025b. Nitrogen regulated reactive oxygen species metabolism of leaf and grain under elevated temperature during the grain-filling stage to stabilize rice substance accumulation. Environ. Exp. Bot. 229, 106037.
    Zhen, F., Liu, Y., Ali, I., Liu, B., Liu, L., Cao, W., Tang, L., Zhu, Y., 2020. Short-term heat stress at booting stage inhibited nitrogen remobilization to grain in rice. J. Agr. Food Res. 2, 100066.
    Zheng, X., Peng, Y., Qiao, J., Henry, R., Qian, Q., 2024. Wild rice: unlocking the future of rice breeding. Plant Biotechnol. J. 22, 3218-3226.
    Zhu, D.-J., Zhang, Z.-S., Qing, T., Gao, J., Xiang, C.-B., Liu, J.-X., 2025. The NLP-HSF regulatory module contributes to nitrogen-mediated thermotolerance in rice. Plant Commun. 6, 101522.
    Zhu, J.K., 2002. Salt and drought stress signal transduction in plants. Annu. Rev. Plant Bio. 53, 247-273.
    Zhu, J.K., 2016. Abiotic stress signaling and responses in plants. Cell 167, 313-324.
  • 加载中

Catalog

    通讯作者: 陈斌, bchen63@163.com
    • 1. 

      沈阳化工大学材料科学与工程学院 沈阳 110142

    1. 本站搜索
    2. 百度学术搜索
    3. 万方数据库搜索
    4. CNKI搜索

    Article Metrics

    Article views (31) PDF downloads (0) Cited by ()
    Proportional views
    Related

    /

    DownLoad:  Full-Size Img  PowerPoint
    Return
    Return