Influence of nitrogen inputs on biomass allocation strategies of dominant plant species in sandy ecosystems

doi:10.1007/s40333-025-0055-1

Journal of Arid Land

2025, Vol. 17

Issue (8): 1118-1146 DOI: 10.1007/s40333-025-0055-1

Research article

Influence of nitrogen inputs on biomass allocation strategies of dominant plant species in sandy ecosystems

CHENG Li¹^,², ZHAN Jin¹^,², NING Zhiying¹^,², LI Yulin¹^,²^,³^,^*(

)

¹Northwest Institute of Eco-Environment and Resources, Chinese Academy of Sciences, Lanzhou 730000, China
²University of Chinese Academy of Sciences, Beijing 100049, China
³Naiman Desertification Research Station, Northwest Institute of Eco-Environment and Resources, Chinese Academy of Sciences, Tongliao 028300, China

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Abstract

Understanding how dominant plants respond to nitrogen (N) addition is critical for accurately predicting the potential effects of N deposition on ecosystem structure and functionality. Biomass partitioning serves as a valuable indicator for assessing plant responses to environmental changes. However, considerable uncertainty remains regarding how biomass partitioning shifts with increasing N inputs in sandy ecosystems. To address this gap, we conducted a greenhouse N fertilization experiment in April 2024, using seeds from 20 dominant plant species in the Horqin Sandy Land of China representing 5 life forms: annual grasses, annual forbs, perennial grasses, perennial forbs, and shrubs. Six levels of N addition (0.0, 3.5, 7.0, 14.0, 21.0, and 49.0 g N/(m²•a), referred to as N0, N1, N2, N3, N4, and N5, respectively) were applied to investigate the effects of N inputs on biomass partitioning. Results showed that for all 20 dominant plant species, the root biomass:shoot biomass (R:S) consistently declined across all N addition treatments (P<0.050). Concurrently, N addition led to a 23.60% reduction in root biomass fraction, coupled with a 12.38% increase in shoot biomass fraction (P<0.050). Allometric partitioning analysis further indicated that N addition had no significant effect on the slopes of the allometric relationships (leaf biomass versus root biomass, stem biomass versus root biomass, and shoot biomass versus root biomass). This suggests that plants can adjust resource investment—such as allocating more resources to shoots—to optimize growth under favorable conditions without disrupting functional trade-offs between organs. Among different life forms, annual grasses, perennial grasses, and annual forbs exhibited increased allocation to aboveground biomass, enhancing productivity and potentially altering community composition and competitive hierarchies. In contrast, perennial forbs and shrubs maintained stable biomass partitioning across all N addition levels, reflecting conservative resource allocation strategies that support long-term ecosystem resilience in nutrient-poor environments. Taken together, these findings deepen our understanding of how nutrient enrichment influences biomass allocation and ecosystem dynamics across different plant life forms, offering practical implications for the management and restoration of degraded sandy ecosystems.

Key words： N addition biomass accumulation biomass partitioning fraction allometric relationship plant life form sandy ecosystems Horqin Sandy Land

Received: 29 December 2024 Published: 31 August 2025

Corresponding Authors: ^*LI Yulin (E-mail: liyl@lzb.ac.cn)

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CHENG Li, ZHAN Jin, NING Zhiying, LI Yulin. Influence of nitrogen inputs on biomass allocation strategies of dominant plant species in sandy ecosystems. Journal of Arid Land, 2025, 17(8): 1118-1146.

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http://jal.xjegi.com/10.1007/s40333-025-0055-1 OR http://jal.xjegi.com/Y2025/V17/I8/1118

Table 1 List of 20 dominant plant species in the Horqin Sandy Land

Fig. 1 Response of biomass accumulation to different nitrogen (N) addition treatments across all 20 dominant plant species. (a), root biomass (RB); (b), stem biomass (SB); (c), leaf biomass (LB); (d), shoot biomass (ShB); (e), total biomass (TB). N0-N5 represent N deposition rates of 0.0, 3.5, 7.0, 14.0, 21.0, and 49.0 g N/(m²•a), respectively. Values represent estimated marginal means and the bar is standard error (SE) derived from the generalized linear mixed models (GLMMs). Different uppercase letters indicate significant differences of the biomass accumulation among different N addition treatments at P<0.050 level.

Fig. 2 Response of biomass partitioning fractions to varying N addition treatments across all 20 dominant plant species. (a), root biomass fraction (RBF); (b), stem biomass fraction (SBF); (c), leaf biomass fraction (LBF); (d), shoot biomass fraction (ShBF); (e), root biomass:shoot biomass (R:S). Values represent estimated marginal means and the bar is SE derived from the GLMMs. Different uppercase letters indicate significant differences of the biomass partitioning fractions among different N addition treatments at P<0.050 level.

Fig. 3 Effects of N addition treatments on the allometric relationships between biomass of different plant organs across all 20 dominant plant species, as assessed using standardized major axis (SMA) regression with statistical significance at P<0.050 level. The colored straight line indicates the slope of the SMA equation under different N treatments, while the colored elliptical area represents the 95% confidence interval of the slope. The pie chart illustrates the explanatory contributions of N treatment, plant species, and residuals to the biomass allometric relationships, based on the variance derived from a Bayesian hierarchical model. The symbol ''--'' means that if the difference in the intercept of the biomass allometric relationship between different N treatments is statistically significant, then the comparison of differences in the shift between N treatments is discontinued.

Fig. 4 Allometric relationships between biomass of different plant organs across all 20 dominant plant species, assessed using SMA regression with statistical significance at P<0.050 level. The orange straight line indicates the slope of the SMA equation under different N treatments, while the orange elliptical area represents the 95% confidence interval of the slope. The two values within parentheses represent the lower 95% confidence limit and the upper 95% confidence limit of the slope, respectively.

Fig. 5 Response of biomass accumulation to varying N addition treatments across different plant life forms. (a), RB; (b), SB; (c), LB; (d), ShB; (e), TB. Values represent estimated marginal means and the bar is SE derived from the GLMMs. Different uppercase letters indicate significant differences of biomass accumulation for the same life form across different N addition treatments at P<0.050 level.

Fig. 6 Response of biomass partitioning fractions to varying N addition treatments across different plant life forms. (a), RBF; (b), SBF; (c), LBF; (d), ShBF; (e), R:S. Values represent estimated marginal means and the bar is SE derived from the GLMMs Different uppercase letters indicate significant differences of biomass partitioning fractions for the same life form across different N addition treatments at P<0.050 level.

Fig. 7 Effects of N addition treatments on the biomass allometric relationships among various plant organs across different life forms, assessed using RMA regression with statistical significance at P<0.050 level. (a1-a4), annual grasses; (b1-b4), annual forbs; (c1-c4), perennial grasses; (d1-d4), perennial forbs; (e1-e4), shrubs. The colored straight line indicates the slope of the SMA equation under different treatments, while the colored elliptical area represents the 95% confidence interval of the slope. The pie charts illustrated the explanatory contributions of N treatment, plant species, and residuals to the biomass allometric relationships, as determined through variance decomposition derived from a hierarchical Bayesian model. The symbol ''--'' means that if the difference in the intercept of the biomass allometric relationship between different N treatments is statistically significant, then the comparison of differences in the shift between N treatments is discontinued.

Fig. 8 Biomass allometric relationships among various plant organs across different life forms, as assessed using RMA regression with statistical significance at P<0.050 level. (a1-a4), annual grasses; (b1-b4), annual forbs; (c1-c4), perennial grasses; (d1-d4), perennial forbs; (e1-e4), shrubs. The orange straight line indicates the slope of the SMA equation under different N treatments, while the orange elliptical area represents the 95% confidence interval of the slope. The two values within parentheses represent the lower 95% confidence limit and the upper 95% confidence limit of the slope, respectively.

Table S1 Summary of the standardized major axis (SMA) regressions of all 20 dominant plant species about the alometric relaionshins of biomass among diferent plant organs under various nitrogen (N) addition treatments

Table S2 Relative contribuions of interspecific variation and N-induced phenotypic plasticity to the allometic relationships of biomass among difrerent organs across all 20 dominant plant species

Table S3 Summary of the SMA regressions about the allometric relationships of biomass among different organs of annual grasses under various N addition treatments

Table S4 Summary of the SMA regressions about the allometric relationships of biomass among different organs of annual forbs under various N addition treatments

Table S5 Summary of the SMA regressions about the allometric relationships of biomass among different organs of perennial grasses under various N addition treatments

Table S6 Summary of the SMA regressions about the allometric relationships of biomass among different organs of perennial forbs under various N addition treatments

Table S7 Summary of the SMA regressions about the allometric relationships of biomass among different organs of shrubs under various N addition treatments

Table S8 Relative contributions of interspecific variation and N-induced phenotypic plasticity to the allometric relationships of biomass among different organs across all 20 dominant plant species with different life forms


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