Stabilized carbon flux through spatial optimization decouples the trade-off between lodging resistance and population sink capacity in high-density wheat

doi:10.1016/j.jia.2026.05.030

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Fei Gao^1*, Yibo Hu^1*, Zhaolin Wang¹, Yajing Tian¹, Weina Wang¹, Zhikuan Jia¹, Xiaolong Ren¹, Peng Zhang¹, Tiening Liu¹, Enke Liu², Zhenlin Wang³, Weibing Yang^4#, Tie Cai¹^#

¹College of Agronomy, Northwest A&F University, Yangling 712100, China

²Institute of Environment and Sustainable Development in Agriculture, Chinese Academy of Agricultural Sciences, Beijing 100081, China

³College of Agronomy, Shandong Agricultural University, Tai’an 271018, China

⁴Institute of Hybrid Wheat, Beijing Academy of Agriculture and Forestry Sciences, Beijing 100097, China

Highlights

l Spatial heterogeneity stabilizes carbon flux and improves yield and lodging resistance.

l Constraint shifts with density: canopy light variability to carbon-mechanical limit.

l High-density heterogeneous distribution (HD-h) breaks dense-planting trade-off, sustaining high lodging resistance index (LRI) and spike number.

l Stem sugar reserves drive cell-wall building, informing lodging-resistance breeding.

Abstract
References

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摘要

高密度种植是保障粮食安全的重要策略，但通常伴随着倒伏风险的显著升高。本研究旨在阐明空间异质化配置缓解小麦倒伏并稳定产量的作用机制。通过开展田间试验，并结合机器学习方法和路径模型分析，识别不同种植配置下制约小麦抗倒伏能力形成的关键因素。与高密度均匀分布处理（HD）相比，高密度异质分布处理（HD-h）有效缓解了冠层内部遮阴，从而稳定了向茎秆输送的同化碳通量。持续的碳供应进一步提高了参与纤维素生物合成的关键酶活性及相关基因的转录表达，促进了机械组织发育，并增强了茎秆的机械承载能力。机器学习结果进一步表明，抗倒伏能力的主要限制因素存在密度依赖性转变：在资源压力较低的条件下，限制因素主要表现为冠层光环境波动；而在高密度条件下，则转变为以碳代谢和结构建成为主导的机械性瓶颈。通过稳定源端同化碳通量，HD-h在维持较高抗倒伏能力的同时，兼顾了单位面积穗数和单穗性状的形成，从而缓解了密植条件下产量形成与结构稳定性之间的传统权衡关系。由此可见，优化空间配置能够通过促进同化产物向结构性碳的转化，提高高密度条件下小麦的抗倒伏能力和产量稳定性。

Abstract

High-density planting is a pivotal strategy for safeguarding food security, yet it is frequently accompanied by a pronounced escalation in lodging risk. This study aimed to elucidate how a spatially heterogeneous configuration mitigates lodging and stabilizes yield in wheat. Field experiments, together with machine-learning approaches and path modeling, were conducted to identify the key constraints underlying lodging resistance under contrasting planting configurations. Relative to the high-density homogeneous distribution (HD), the high-density heterogeneous distribution (HD-h) effectively alleviated within-canopy shading, thereby stabilizing the assimilate carbon flux delivered to the stem. This sustained carbon supply, in turn, enhanced the activity and transcriptional expression of key enzymes/genes involved in cellulose biosynthesis, promoting the development of mechanical tissues and increasing stem mechanical load-bearing capacity. Machine-learning further revealed a density-dependent shift in the primary limiting factors for lodging resistance: constraints transitioned from fluctuations in canopy light environment under lower resource pressure to a mechanically oriented bottleneck under high density, dominated by carbon metabolism and structural construction. By stabilizing source-derived carbon flux, HD-h coordinated the maintenance of high lodging resistance with concurrent support for spike number per unit area and individual-spike traits, thereby relaxing the conventional trade-off between yield and structural stability under dense planting. Optimizing spatial configuration improves lodging resistance and yield stability under high-density conditions by promoting assimilate conversion into structural carbon.

Keywords: wheat spatially heterogeneous distribution lodging resistance carbon-structural coordination RF and PLS-PM

Online: 22 May 2026

Fund:

This work was funded by the Special Project for Innovation Capacity Building of Beijing Academy of Agriculture and Forestry Sciences, China (KJCX20260904), the National Key R&D Program of China (14th Five-Year Plan; 2023YFD1900503), the National Natural Science Foundation of China (32071955), and the Key R&D Plan of Shaanxi Province, China (2024NC-YBXM-041).

About author: #Correspondence Weibing Yang, E-mail: shennong04@126.com, Tie Cai, E-mail: caitie@nwsuaf.edu.cn * These authors contributed equally to this study.

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Fei Gao, Yibo Hu, Zhaolin Wang, Yajing Tian, Weina Wang, Zhikuan Jia, Xiaolong Ren, Peng Zhang, Tiening Liu, Enke Liu, Zhenlin Wang, Weibing Yang, Tie Cai. 2026. Stabilized carbon flux through spatial optimization decouples the trade-off between lodging resistance and population sink capacity in high-density wheat. Journal of Integrative Agriculture, Doi:10.1016/j.jia.2026.05.030

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