Mechanism of canopy nitrogen spraying after anthesis increasing grain yield and protein content in wheat

doi:10.1016/j.jia.2025.11.036

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Jie Ren¹, Haoran Li¹, Bingjin Zhou¹, Zhongwei Liu¹, Ying Liu¹, Zhimin Wang¹, Zhigan Zhao¹, Zhencai Sun¹, Yong Li², Xubo Zhang^3#, Yinghua Zhang^1,4#

¹ College of Agronomy and Biotechnology, China Agricultural University, Beijing 100193, China

² College of Agronomy, Shandong Agricultural University, Tai’an 271018, China

³ Key Laboratory of Ecosystem Network Observation and Modeling, Institute of Geographic Sciences and Natural Resources Research, Chinese Academy of Sciences, Beijing 100101, China

⁴ Wuqiao Field Scientific Observation and Research Station of Agricultural Environment, Ministry of Agriculture and Rural Affairs, Cangzhou 061807, China

Highlights

l Canopy nitrogen spraying at 15 days after anthesis primarily targeted spike organs rather than other plant tissues, enhancing post-anthesis nitrogen assimilation—accounting for 84.13% of the increase in grain nitrogen accumulation—and promoting the remobilization of pre-anthesis nitrogen to grains.

l Compared with soil-based nitrogen application during the jointing stage, canopy nitrogen spraying at 15 days after anthesis led to significant nitrogen accumulation in grains starting at 12 h after application, contributing 32.52% to the increase in grain nitrogen accumulation.

l Canopy nitrogen spraying at 15 days after anthesis stimulated C₄ pathway activity in wheat spike organs and enhanced nitrogen metabolism in grains, thereby increasing grain yield and total protein content through improved grain weight and storage protein synthesis.

Abstract
References

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摘要

在小麦灌浆期间，适当的外源氮供应可以通过延缓衰老和增加氮储备促进籽粒产量和蛋白质积累。然而，其潜在机制尚不清楚。本文通过盆栽试验证实了花后15天（AS处理）冠层喷氮的有效性，并在节水栽培条件下通过田间试验进一步研究了相关调控机制。盆栽试验表明，AS处理通过促进花前氮素再动员和花后氮同化，提高了小麦的粒重、产量和氮素积累量；喷施后12小时，冠层吸收的氮素在籽粒中开始显著积累，对增加的籽粒氮素积累量贡献了32.52%。田间试验进一步证实，与喷施等量的去离子水（CK处理）相比，AS处理提高了快速生长阶段和缓慢生长阶段的籽粒灌浆速率和氮积累速率，使得籽粒产量提高了5.21%，蛋白质含量提高了7.50%；AS处理增强了C₄途径关键酶的活性，包括磷酸烯醇式丙酮酸羧化酶（PEPC）、NADP苹果酸脱氢酶（NADP-MDH）、NADP-苹果酸酶（NADP-ME）、丙酮酸磷酸二激酶（PPDK），提高了颖壳、外稃和内稃中的苹果酸含量。以上结果表明，AS处理促进了三羧酸（TCA）循环和卡尔文循环，为蛋白质和淀粉的生物合成提供了反应底物，还促进了籽粒中的氮代谢，有利于蛋白质积累。本研究为种植系统和气候条件相似地区提供了一种可以缓解花后的干旱胁迫并提高小麦产量和蛋白质含量的可行栽培策略。

Abstract

During wheat grain filling, exogenous nitrogen supply can enhance grain yield and protein accumulation by delaying senescence and increasing nitrogen reserves. However, the underlying mechanisms remain unclear. The efficacy of canopy nitrogen spraying at 15 days after anthesis (AS) was first evaluated in a pot experiment, and the associated regulatory mechanisms were further investigated in a field trial under water-saving cultivation conditions. The pot experiment demonstrated that AS treatment increased grain weight, yield, and nitrogen accumulation by improving both pre-anthesis nitrogen remobilization and post-anthesis nitrogen assimilation. Canopy-derived nitrogen began accumulating significantly in grains at 12 h after spraying, accounting for 32.52% of the increase in grain nitrogen accumulation. The field experiment further validated that AS treatment increased grain filling rate and nitrogen accumulation rate during fast and slow growth stages, significantly increasing grain yield by 5.21% and protein content by 7.50% compared to spraying equal amounts of deionized water (CK). AS treatment upregulated key enzymes in the C₄ pathway—including phosphoenolpyruvate carboxylase (PEPC), NADP-malate dehydrogenase (NADP-MDH), NADP-malic enzyme (NADP-ME), pyruvate phosphate dikinase (PPDK)—and increased malate levels in glumes, lemmas, and paleae. These responses suggested that AS treatment facilitated the tricarboxylic acid (TCA) cycleand the Calvin cycle, providing reaction substrates for protein and starch biosynthesis. Additionally, AS treatment promoted grain nitrogen metabolism, facilitating protein accumulation. This study presents a viable strategy to mitigate post-anthesis drought stress and improve wheat productivity and grain quality in regions with similar agroclimatic conditions.

Keywords: canopy nitrogen spraying spike organs C₄ pathway C and N metabolism N accumulation wheat

Online: 24 November 2025

Fund:

This study was supported by the National Key Research and Development Program of China (2022YFD2300802), the Higher Level “Qilu Granary” Construction and Capacity Improvement Technology Integration and Demonstration of Shandong Agricultural University (539703006002), and the earmarked fund for China Agriculture Research System (CARS-3).

About author: #Correspondence Yinghua Zhang, E-mail: yhzhang@cau.edu.cn; Xubo Zhang, E-mail: zhangxb@igsnrr.ac.cn

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Jie Ren, Haoran Li, Bingjin Zhou, Zhongwei Liu, Ying Liu, Zhimin Wang, Zhigan Zhao, Zhencai Sun, Yong Li, Xubo Zhang, Yinghua Zhang. 2025. Mechanism of canopy nitrogen spraying after anthesis increasing grain yield and protein content in wheat. Journal of Integrative Agriculture, Doi:10.1016/j.jia.2025.11.036

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