Cytokinins redistributing drives nitrogen remobilization from source to sink in wheat under moderate water limitation during grain filling

doi:10.1016/j.jia.2025.02.032

Journal of Integrative Agriculture

2026, Vol. 25

Issue (5): 1857-1870 DOI: 10.1016/j.jia.2025.02.032

Crop Science

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Cytokinins redistributing drives nitrogen remobilization from source to sink in wheat under moderate water limitation during grain filling

Ying Liu^{1, 2}, Jiangyao Fu^{1, 2}, Haotian Chen^{1, 2}, Yajun Zhang^{1, 2},Siyu Li^{1, 2}, Kuanyu Zhu^{1, 2}, Yunji Xu³, Weilu Wang³, Junfei Gu^{1, 2}, Hao Zhang^{1, 2}, Zhiqin Wang^{1, 2}, Lijun Liu^{1, 2}, Jianhua Zhang^{4, 5}, Weiyang Zhang^{1, 2#}, Jianchang Yang^{1, 2#}

¹Jiangsu Key Laboratory of Crop Genetics and Physiology/Jiangsu Key Laboratory of Crop Cultivation and Physiology/Agricultural College, Yangzhou University, Yangzhou 225009, China

²Jiangsu Co-Innovation Center for Modern Production Technology of Grain Crops, Yangzhou University, Yangzhou 225009, China

³Joint International Research Laboratory of Agriculture and Agri-Product Safety, Yangzhou University, Yangzhou 225009, China

⁴Department of Biology, Hong Kong Baptist University, Hong Kong 999077, China

⁵State Key Laboratory of Agrobiotechnology, The Chinese University of Hong Kong, Hong Kong 999077, China

Highlights
● Post-anthesis moderate water limitation enhances source-to-sink nitrogen remobilization, significantly boosting grain yield and nitrogen use efficiency in wheat.
● Post-anthesis moderate water limitation promotes the redistribution of cytokinins between source and sink in wheat.
● Redistributing cytokinins from source to sink drives elevates nitrogen distribution in sink organs under moderate water limitation post anthesis.

Abstract
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摘要

本研究探讨了细胞分裂素在适度限制供水（MWL）调控小麦灌浆期氮素从源到库转运过程中的作用。采用低氮（LN）、中氮（MN）和高氮（HN）三种施氮水平进行田间实验，并在每个氮肥施用水平下设置了两种土壤水分处理：常规充充分灌溉（CWW）和花后适度限制供水（MWL）。结果表明，MWL处理优化了氮素、总游离氨基酸、反式玉米烯（Z）和反式玉米烯核苷（ZR）从源器官（茎和叶）向库器官（穗）的再分配。与CWW处理相比，MWL处理下茎和叶中的蛋白水解酶（包括内肽酶、羧肽酶和氨基肽酶）的活性，以及穗中总游离氨基酸转运蛋白基因的表达水平显著提高，并且与穗中Z+ZR水平密切相关。向茎和叶喷施6-呋喃基氨基嘌呤（KT，一种合成细胞分裂素）显著抑制了蛋白水解酶活性，促进了氮素在茎和叶中的滞留，减少了库器官中的氮积累，从而降低了氮收获指数。相比之下，向穗喷施KT显著上调了FAA转运蛋白基因的表达，减少了源器官（茎和叶）中的氮滞留，增加了库器官（穗）中的氮积累，并提高了氮收获指数。在HN条件下，MWL处理对氮源库间转运的促进作用明显强于LN和MN条件。研究结果表明，花后适度限制供水通过重新分配小麦中的细胞分裂素（Z+ZR）来增强氮素从源器官到库器官的再分配，协同增加了小麦的产量和氮素利用效率。

Abstract

This study examined the involvement of cytokinins in the process by which moderate water limitation (MWL) mediates nitrogen (N) remobilization from source to sink during the grain-filling phase in wheat. Field experiments were performed using N application rates of low (LN), medium (MN), and high (HN). Two soil moisture regimes were implemented for each N rate: conventional well-watered (CWW) and MWL post anthesis. The MWL application optimized N, total free amino acids (FAA), and trans-zeatin (Z)+trans-zeatin riboside (ZR) reallocation from the source organs (stems and leaves) to the sink organ (spikes) in wheat. Compared to those in the CWW regime, the activities of proteolytic enzymes, including endopeptidase, carboxypeptidase, and aminopeptidase within stems and leaves, and the expression levels of total FAA transporter genes in spikes were significantly elevated in the MWL regime, showing a close correlation with the Z+ZR levels in the spikes. Application of kinetin to stems and leaves significantly inhibited proteolytic enzyme activities, promoting N retention in stems and leaves, decreasing N accumulation in the sink organ, and reducing the N harvest index. In contrast, applying kinetin to spikes significantly upregulated expression levels of FAA transporter genes, reducing N retention in stems and leaves, increasing N accumulation in the sink organ, and raising the N harvest index. Such facilitation induced by the MWL in the remobilization of N from source to sink was greater at HN than at LN or MN. Results demonstrate that post-anthesis MWL can significantly intensify the remobilization of N from source to sink, while also synergistically enhancing grain yield and N use efficiency through strategically redistributing cytokinins (Z+ZR) between source and sink in wheat.

Keywords: cytokinins moderate water limitation nitrogen remobilization nitrogen use efficiency wheat (Triticum aestivum L.)

Received: 11 October 2024 Accepted: 08 January 2025 Online: 19 February 2025

Fund:

This work was supported by the National Natural Science Foundation of China (32572422 and 32272198), the Top Talent Supporting Program of Yangzhou University, China (YZU-2028-01), the Priority Academic Program Development of Jiangsu Higher Education Institutions, China (PAPD), and the Government Funding to the Chinese University of Hong Kong State Key Laboratory of Agrobiotechnology via Innovation and Technology Commission, China (2022/23–2023/24).

About author: #Correspondence Weiyang Zhang, E-mail: wyz@yzu.edu.cn; Jianchang Yang, E-mail: jcyang@yzu.edu.cn

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Ying Liu, Jiangyao Fu, Haotian Chen, Yajun Zhang, Siyu Li, Kuanyu Zhu, Yunji Xu, Weilu Wang, Junfei Gu, Hao Zhang, Zhiqin Wang, Lijun Liu, Jianhua Zhang, Weiyang Zhang, Jianchang Yang. 2026. Cytokinins redistributing drives nitrogen remobilization from source to sink in wheat under moderate water limitation during grain filling. Journal of Integrative Agriculture, 25(5): 1857-1870.

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