Modeling interplant competition in maize: Integrating density-dependent specific leaf area plasticity enhances canopy light and nitrogen use efficiency

doi:10.1016/j.jia.2026.03.049

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Pengpeng Zhang^{1, 2, 3}, Cheng Li¹, Hongbo Wang¹, Xiujuan Wang⁴, Mengzhen Kang⁴, Jun Xue², Youhong Song^1,^5,⁶^#

¹School of Agronomy, Anhui Agricultural University, Hefei 230036, China

²Key Laboratory of Crop Physiology and Ecology, Institute of Crop Sciences, Chinese Academy of Agricultural Sciences, Beijing 10081, China

³Institute of Western Agriculture, Chinese Academy of Agricultural Sciences, Changji 831100, China

⁴State Key Laboratory of Multimodal Artificial Intelligence Systems, Institute of Automation, Chinese Academy of Sciences, Beijing 100190, China

⁵Ministry of Education Key Laboratory of Crop Physiology, Ecology and Genetic Breeding, Jiangxi Agricultural University, Nanchang 330045, China

⁶Centre for Crop Science, Queensland Alliance for Agriculture and Food Innovation, The University of Queensland, Brisbane, QLD 4350, Australia

Highlights

l The adjustment of SLA under dense planting optimizes canopy photosynthesis and resource use efficiency

l GreenLab-Maize with parameterized SLA can predict the effects of intensified interplant competition

Abstract
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摘要

通过模型策略优化种植密度对于提高现代玉米生产系统的资源利用效率和生产力至关重要。功能-结构植物模型诸如GreenLab等为模拟作物生长和产量形成提供了强大的框架。然而，它们在模拟密植条件下株间竞争方面的能力仍然有限，这主要是由于对比叶面积（SLA）参数过于简化。为弥补这一不足，本研究开展了一项为期两年（2022-2023年）的田间试验，试验设置四个玉米种植密度（3、6、9和12 plants m^-2，分别为PD3、PD6、PD9和PD12），旨在探究密度驱动的叶片结构和功能性状变化，并评估SLA可塑性在调控冠层光能及氮素利用效率中的作用。增加种植密度显著改变了冠层结构和资源分布。与PD3相比，高密度处理（PD6、PD9和PD12）提高了叶向值、SLA、光合氮利用效率（PNUE）、辐射利用效率（RUE）和作物生长速率，但同时降低了单株叶面积、比叶氮含量和光饱和光合速率。进一步分析表明，密度诱导的SLA可塑性提高了PNUE，并调整了氮素与光的消光系数之比，从而部分补偿了因SLN降低所导致的叶片光合能力下降。在冠层尺度上，SLA与RUE呈显著正相关，突显了其在株间竞争下促进冠层光合作用的作用。随后，将观察到的SLA对种植密度的响应进行参数化，并整合到GreenLab-Maize模型中。与标准模型相比，修订后的模型显著改善了对密植条件下叶面积、叶面积指数、RUE和累积生物量的预测。总体而言，本研究确立了SLA可塑性作为密植条件下资源优化的关键适应性性状，并提供了一种经过验证的方法，以增强功能-结构植物模型中株间竞争模拟的真实性。

Abstract

Optimizing plant density through modeling strategies is essential for improving resource-use efficiency and productivity in modern maize production systems. Functional–structural plant models (FSPMs) such as GreenLab provide a powerful framework for simulating crop growth and yield formation, yet their ability to represent interplant competition under high-density conditions remains limited, largely due to the oversimplified treatment of specific leaf area (SLA). To address this gap, a two-year field experiment (2022–2023) composed of four maize plant densities (3, 6, 9 and 12 plants m⁻², denoted as PD3, PD6, PD9, and PD12, respectively), was conducted to examine density-driven changes in leaf structural and functional traits and to evaluate the role of SLA plasticity in regulating canopy light and nitrogen use efficiency. Increased plant density substantially reshaped canopy architecture and resource distribution. Compared to PD3, higher densities (PD6, PD9, and PD12) increased leaf orientation value, SLA, photosynthetic nitrogen-use efficiency (PNUE), radiation-use efficiency (RUE) and crop growth rate (CGR), while reducing individual leaf area (LA), specific leaf nitrogen content (SLN) and light-saturated photosynthetic rate (A_max). Further analyses showed that density-induced SLA plasticity enhanced PNUE and adjusted the ratio of nitrogen to light extinction coefficients (K_N/K_L), thereby partially compensating for the decline in leaf photosynthetic capacity caused by reduced SLN. At the canopy scale, SLA was strongly and positively associated with RUE, highlighting its role in enhancing canopy photosynthesis under interplant competition. Subsequently, the observed SLA responses to plant density were parameterized and incorporated into the GreenLab-Maize model. Compared to the standard model, the revised model markedly improved predictions of LA, leaf area index, RUE and accumulated biomass under high-density conditions. Overall, this study establishes SLA plasticity as a key adaptive trait for resource optimization in dense stands and provide a validated method to enhance the realism of competition simulations in FSPMs.

Keywords: Zea mays L. interplant competition light and nitrogen distribution canopy structure specific leaf area

Online: 20 March 2026

Fund:

This work was financially supported by the Natural Key R&D Program of China (2023YFD2301500).

About author: #Correspondence Youhong Song, E-mail: uqysong@163.com

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Pengpeng Zhang, Cheng Li, Hongbo Wang, Xiujuan Wang, Mengzhen Kang, Jun Xue, Youhong Song. 2026. Modeling interplant competition in maize: Integrating density-dependent specific leaf area plasticity enhances canopy light and nitrogen use efficiency. Journal of Integrative Agriculture, Doi:10.1016/j.jia.2026.03.049

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