J Plant Ecol ›› Advance articles     DOI:10.1093/jpe/rtag198

• Research Article •     Next Articles

Environment-dependent shrub-herb interactions reshape carbon-nutrient coupling in shrub-encroached dryland grasslands

Peng He1, 2, Jiang Chen1, Mengxi Hou1, Heyong Liu3*, Lina Xie1, Liang Man1, Chengcang Ma1, Yong Jiang3, Mai-He Li4, 3, 5   

  1. 1 Tianjin Key Laboratory of Animal and Plant Resistance, College of Life Sciences, Tianjin Normal University, Tianjin, China
    2 Zhejiang Key Laboratory for Restoration of Damaged Coastal Ecosystems, School of Life Sciences, Taizhou University, Zhejiang Taizhou 318000, China
    3 College of Life Sciences, Hebei University, Baoding 071002, China
    4 Forest dynamics, Swiss Federal Institute for Forest, Snow and Landscape Research WSL, CH8903 Birmensdorf, Switzerland
    5 Key Laboratory of Geographical Processes and Ecological Security in Changbai Mountains, Ministry of Education/School of Geographical Sciences, Northeast Normal University, Changchun 130024, Jilin, China.
    *: Correspondence author:Heyong Liu (Correspondence author): liuheyong@hbu.edu.cn
  • Received:2026-02-10 Accepted:2026-08-15 Published:2026-08-31
  • Supported by:
    This research was supported by the Central Government Guided Local Science and Technology Development Fund Projects (Grant No. 2025ZY01033), the National Natural Science Foundation of China (Grant No. 32001147), and the Natural Science Foundation of Tianjin (Grant No. 23JCQNJC01140).

Abstract: Shrub encroachment is a widespread consequence of global change in dryland grasslands, yet how environmental context mediates shrub-herb interactions and their effects on plant carbon-nutrient coupling remains poorly understood. Here, we combined a large-scale field transect spanning a natural aridity gradient with a complementary greenhouse pot experiment to examine how shrub-mediated environmental changes influence foliar nitrogen (N), phosphorus (P), and non-structural carbohydrates (NSCs) in coexisting herbaceous species. Across Inner Mongolian drylands, mature shrub (Caragana stenophylla) patches consistently enhanced foliar N and P concentrations of understory herbs while suppressing NSCs accumulation, indicating a trade-off between nutrient acquisition and carbon storage. However, these inhibitory effects on carbon reserves weakened with increasing aridity, such that shrub-herb interactions shifted toward neutral or facilitative outcomes for NSCs under drier conditions. In contrast, under controlled greenhouse conditions, C. stenophylla seedlings suppressed nutrient uptake and altered NSC dynamics in coexisting herbaceous seedlings, reflecting competitive interactions associated with the limited cumulative environmental modification capacity of early-stage shrub-associated environments. Responses differed markedly between herbaceous functional groups: forbs (e.g. Artemisia frigida) exhibited greater sensitivity to shrub presence and drought than grasses (e.g. Leymus chinensis), driven by coordinated variation in leaf and root functional traits. Together, our results demonstrate that shrub-induced microsite modifications reshape herbaceous carbon-nutrient coupling in an environment-dependent manner, with implications for predicting biogeochemical and vegetation dynamics in dryland grasslands under increasing climatic stress. PTED MANUSCRIPT

Shrub encroachment increased herbaceous N and P but reduced NSCs along the aridity gradient, with this carbon inhibition weakening as aridity increased. Greenhouse experiments, however, revealed consistently competitive shrub seedling effects. Differential responses between grasses and forbs highlight environment dependent shrub-herb interactions that modulate carbon-nutrient coupling in dryland grasslands.