Nutrient-mediated tree growth responses are governed by internal resource allocation: intra-organ water-carbon...

Nutrient deposition is reshaping forest productivity, yet the physiological mechanisms linking nitrogen (N) and phosphorus (P) enhancement to tree growth remain incompletely resolved. We propose that internal resource allocation strategies-specifically intra-organ trade-offs between water and carbohydrate resources and inter-organ carbohydrate concentration gradient-mediate nutrient-driven tree growth responses. Using a long-term fertilization experiment in subtropical Cunninghamia lanceolata (Lamb.) Hook. plantations, we measured relative water content (RWC) and traditional resource concentrations [soluble sugar (SS) concentration; starch (ST) concentration] during drought season. We rank-transformed RWC, SS and ST within each organ type across all samples. Intra-organ water-carbon trade-offs were quantified as two normalized ratios: ln(RWCrank/SSrank) for RWC:SS and ln(RWCrank/STrank) for RWC:ST, where higher values indicate greater solute dependence for turgor maintenance or greater ST storage volume fraction, respectively. The inter-organ carbon concentration gradient was quantified as differences in SS or ST between leaves and twigs (SSleaf-twig, STleaf-twig); and between absorptive roots and transport roots. We demonstrated three key nutrient-specific responses: (i) N addition promoted transport root ln(RWCrank/SSrank) while enhancing leaf ln(RWCrank/STrank), amplifying STleaf-twig; (ii) P addition reduced leaf and twig ln(RWCrank/SSrank) and (iii) combined N+P addition reduced twig ln(RWCrank/SSrank), while increasing leaf and twig ln(RWCrank/STrank). These reorganization patterns had direct growth consequences: while traditional resource concentrations explained 26.0% of growth variation, incorporating derived attributes increased explanatory power by 43.5% (to 37.3% total variance explained). Notably, the twig ln(RWCrank/STrank) emerged as the single strongest growth predictor, where N+P induced ST dominance correlated with enhanced growth rate. Nitrogen enrichment enhanced SS dominance in transport roots, a pattern that may occur at the expense of stem growth. Our results establish an internal resource allocation framework that mechanistically links nutrient-mediated carbon management patterns to forest productivity under global change, revealing how N+P co-enrichment synergistically optimizes carbon resource storage and utilization beyond single-nutrient effects.