Community functional traits shift under long-term nitrogen addition: Roles of intraspecific trait variability and species turnover

doi:10.1093/jpe/rtag101

J Plant Ecol ›› Advance articles DOI:10.1093/jpe/rtag101

Community functional traits shift under long-term nitrogen addition: Roles of intraspecific trait variability and species turnover

Chen Chen^1,2, Jinlong Peng^1,2, Jiaqiang Liao^1,2, Fangfang Ma³, Dashuan Tian^1,2, Han Y. H. Chen^4,5,6*, Shuli Niu^1,2*

1 Key Laboratory of Ecosystem Network Observation and Modeling, Institute of Geographic Sciences and Natural Resources Research, Chinese Academy of Sciences, Beijing 100101, China;
2 College of Resources and Environment, University of Chinese Academy of Sciences, Beijing 100049, China;
3 School of Ecology and Nature Conservation, Beijing Forestry University, Beijing 100083, China;
4 Key Laboratory of Grassland Resources of the Ministry of Education, Key Laboratory of Forage Cultivation, Processing and High Efficient Utilization of the Ministry of Agriculture and Rural Affairs, Inner Mongolia Key Laboratory of Grassland Management and Utilization, College of Grassland Science, Inner Mongolia Agricultural University, Hohhot, 010011 China;
5 Faculty of Natural Resources Management, Lakehead University, Thunder Bay, ON, P7B 5E1 Canada;
6 Institute for Global Change Biology, School for Environment and Sustainability, University of Michigan, Ann Arbor, MI, 48109 USA
^*Corresponding author: Han Y. H. Chen (han.yh.chen@gmail.com) and Shuli Niu (sniu@igsnrr.ac.cn)

Received:2025-10-09 Revised:2026-02-09 Accepted:2026-03-18 Published:2026-05-18
Supported by:
This work was supported by the National Key Technology R & D Program of China (2022YFF0802102), and the International Partnership Program of Chinese Academy of Sciences (177GJHZ2022020BS).

长期氮添加驱动群落功能性状变化：种内性状变异与物种周转的作用

Abstract

Abstract: Plant functional traits at the community level determine ecosystem responses to environmental change. Despite globally increasing nitrogen deposition, the roles of intraspecific trait variability (ITV) and species turnover in shaping community composition of functional traits under long-term nitrogen deposition remain unclear. We conducted a 10-yr nitrogen addition experiment in an alpine meadow and measured five key leaf traits: leaf nitrogen content (LNC), specific leaf area (SLA), leaf dry matter content (LDMC), leaf thickness (LT), and leaf area (LA). Nitrogen addition increased community-weighted means (CWMs) of LNC and SLA but decreased LT. Sum of squares decomposition revealed that ITV primarily drove changes in LNC, SLA, and LDMC, whereas species turnover mainly influenced LT. ITV was shaped by soil inorganic nitrogen and light availability, while species turnover was mainly driven by soil inorganic nitrogen. These findings reflect the complex adaptive strategies of plant communities in nitrogen-enriched environments and offer insights into the mechanisms driving community composition changes of functional traits under nitrogen addition.

Key words: plant functional traits, intraspecific trait variability, species turnover, nitrogen deposition, alpine meadow

摘要：
群落水平的植物功能性状决定了生态系统对环境变化的响应。尽管全球氮沉降持续增加，但在长期氮沉降背景下，种内性状变异和物种周转在塑造群落功能性状组成中的作用仍不明确。本研究依托高寒草甸生态系统开展了为期10年的氮添加实验，并测定了5个关键叶片功能性状，包括叶片氮含量、比叶面积、叶片干物质含量、叶片厚度和叶面积。结果表明，氮添加提高了叶片氮含量和比叶面积的群落加权均值，但降低了叶厚度的群落加权均值。平方和分解结果显示，种内性状变异主要驱动叶片氮含量、比叶面积和叶片干物质含量的变化，而叶厚度的变化则主要由物种周转所驱动。进一步分析表明，种内性状变异主要受土壤无机氮含量和光资源可利用性调控，而物种周转主要由土壤无机氮含量影响。本研究结果揭示了植物群落在富氮环境中的复杂适应策略，并为理解氮沉降背景下群落功能性状组成变化的驱动机制提供了重要认识。

关键词: 植物功能性状, 种内性状变异, 物种周转, 氮沉降, 高寒草甸

Chen Chen, Jinlong Peng, Jiaqiang Liao, Fangfang Ma, Dashuan Tian, Han Y. H. Chen, Shuli Niu. Community functional traits shift under long-term nitrogen addition: Roles of intraspecific trait variability and species turnover[J]. J Plant Ecol, DOI: 10.1093/jpe/rtag101.

[1]	Yimin Zhao, Zhen Zhang, Li Zhang, Miaojun Ma, Guorui Hu, Shurong Zhou. Nitrogen addition regulates community biomass resistance and recovery to drought by altering species asynchrony rather than the diversity response [J]. J Plant Ecol, 2026, 19(2): 1-43.
[2]	Jiatao Zhang, Lan Du, Yonghong Luo, Yann Hautier, Ru Tian, Yan Shen, Mohsin Mahmood, Zhuwen Xu. Distinct effects of drought and nitrogen enrichment on the compositional and productivity stability of temperate grasslands [J]. J Plant Ecol, 2026, 19(2): 1-.
[3]	Na Li, Suke Hou, Mai-He Li, Xi Qi, Liangjun Zhu, Jitang Li, Paolo Cherubini, Zhaoyong Hu, Zhengfang Wu, Renkai Dong, Lei Wang, Youwen Wang, Danyang Yuan, Haibo Du. Elevation gradient modulates the effects of herbaceous encroachment on the long-term growth trends and climate sensitivity of Rhododendron aureum in alpine tundra [J]. J Plant Ecol, 2026, 19(2): 1-.
[4]	Yaping Lin, Qiufang Zhang, Hao Sun, Jiaohong Lu, Zehong Yao, Quanxin Zeng, Chenxing Lian, Huizhi Pi, Xiaochun Yuan, Yuehmin Chen. Parallel patterns of microbial carbon and nitrogen use efficiency in response to nitrogen addition across soil depths in a Castanopsis faberi forest [J]. J Plant Ecol, 2026, 19(2): 1-.
[5]	Xiao-Hua Yang, Xiao-Sa Liang, Yuan Shi, Yuan-Xiu Wu, Xiao-Ru Zhang, Cong Ding, Xiao-Jing Zhang, Alemayehu Adugna, Xiao-Tao Lü. Grassland plant species with slow-growing strategy are more resistant to extreme drought [J]. J Plant Ecol, 2026, 19(2): 1-.
[6]	Jiahui Chen, Nan Jia, Youcun Suo, Nan Hu, Guofu Zhao, Haiyang Gong, Zhiqiang Wang. Long-term yak-grazing alters plant fine-root C:N:P stoichiometry in a Tibetan alpine meadow [J]. J Plant Ecol, 2026, 19(1): 1-.
[7]	Peipei Liu, Cai-E Wang, Mei Huang, A. Allan Degen, Wenyin Wang, Lingyan Qi, Binyu Luo, Tianyun Qi, Jianxin Jiao, Zhanhuan Shang. The shift from dominant Kobresia species to toxic plants is associated with bare patch encroachment on alpine meadow hillslopes [J]. J Plant Ecol, 2026, 19(1): 1-.
[8]	Jiaoneng Yu, Fei Ren, Li Ma, Huakun Zhou, Xiaotao Huang, Yongsheng Yang, Songbo Qu, Yuxuan Cui, Ribu Shama, Yuxia Zhang, Xu-Ri, Almaz Borjigidai. Community dynamics of alpine meadows under fencing durations: based on diversity, niches, and interspecific associations [J]. J Plant Ecol, 2026, 19(1): 1-.
[9]	Yuhan Zou, Xiying Tang, Lijuan Cui, Xinsheng Zhao, Xiajie Zhai, Yinru Lei, Jinzhi Wang, Huazhe Li, Wei Li. Disturbance-driven variation in soil characteristics, plant functional traits, and biodiversity in wetlands [J]. J Plant Ecol, 2026, 19(1): 1-.
[10]	Gianalberto Losapio, Lucía Mottet, Nora Khelidj, Bao Ngan Tu, Bruno E.L. Cerabolini, Stephanie Grand, Natasha de Vere, Antoine Guisan. Ecological indicators predict functional diversity dynamics following glacier retreat [J]. J Plant Ecol, 2025, 18(6): 1-37.
[11]	Yuguang Ke, Hongqiang Wang, Yushu Zhang, Jiale Chen, Yalong Shi , Yadong Yang, Qiang Yu, Xiwen Li, Ang Li, Jianguo Wu, Honghui Wu. Ecosystem stability declines consistently with nitrogen addition across a gradient of organic to inorganic nitrogen ratios [J]. J Plant Ecol, 2025, 18(5): 1-28.
[12]	Fandong Meng, Jianping Sun, Wangwang Lü, Zhenhua Zhang, Bowen Li, Yang Zhou, Jingya Lü, Lanying Chen, Tsechoe Dorji, Shiping Wang. Trade-off between growing season length and herbaceous growth on the Third Pole [J]. J Plant Ecol, 2025, 18(4): 1-11.
[13]	Ya-Jing Zhang, Ya-Li Si, Shi-Bo Ju, Wan-Ying Jiang, Ming-Zhen Zhao, Jing Liu, Mulualem Tigabu, Xiang-Qing Ma, Ming Li. Synergistic Effects of Nitrogen Deposition and Phosphorus Distribution Patterns on Root morphological and chemical traits, and Nutrient Efficiency in Chinese Fir [J]. J Plant Ecol, 2025, 18(4): 1-50.
[14]	Chao Wang, Jiayin Feng, Jian Song, Hongpeng Wang, Jinhua Zhang, Jingyi Ru, Yaru Lyu, Wenjing Ma, Zixuan Yang, Mengyu Yan, Xueli Qiu, Shiqang Wan. Nighttime warming reshapes thermal regimes in freshwater wetlands: cascading effects on carbon dynamics [J]. J Plant Ecol, 2025, 18(4): 1-41.
[15]	Yang Wang, Yu-Ping Dong, Zhao-Jie Ren, Ling Hu, Tong-Yue Deng, Zun-Tian Zhao. Mosses record historical variations of atmospheric nitrogen deposition in a mountain area of northern China [J]. J Plant Ecol, 2025, 18(3): 1-10.