Nutrient-limited conditions determine the responses of foliar nitrogen and phosphorus stoichiometry to nitrogen addition: A global meta-analysis

To test the hypothesis that nutrient-limited conditions can determine the responses of nitrogen (N) and phosphorus (P) stoichiometry to N addition, a meta-analysis was conducted to identify the different responses of foliar N and P concentrations and N-to-P ratios to N addition under N limitation, N and P co-limitation and P limitation. N addition increased the foliar N-to-P ratios and N concentrations by 46.2% and 30.2%, respectively, under N limitation, by 18.7% and 19.7% under N and P co-limitation, and by 4.7% and 12.9% under P limitation. However, different responses of foliar P concentrations to N addition were observed under different nutrient limitations, and negative, positive, and neutral effects on P concentrations were observed under N limitation, P limitation and N and P co-limitation, respectively. Generally, the effects of N addition on N-to-P ratios and N concentrations in herbaceous plants were dramatically larger than those in woody plants (with the exception of the N-to-P ratio under N limitation), but the opposite situation was true for P concentrations. The changes in N-to-P ratios were closely correlated with the changes in N and P concentrations, indicating that the changes in both N and P concentrations due to N addition can drive N and P stoichiometry, but the relative sizes of the contributions of N and P varied greatly with different nutrient limitations. Specifically, the changes in N-to-P ratios may indicate a minimum threshold, which is consistent with the homeostatic mechanism. In brief, increasing N deposition may aggravate P limitation under N-limited conditions but improve P limitation under P-limited conditions. The findings highlight the importance of nutrient-limited conditions in the stoichiometric response to N addition, thereby advancing our ability to predict global plant growth with increasing N deposition in the future.