Available nitrogen and enzyme activity in rhizosphere soil dominate the changes in fine-root nutrient foraging strategies during plantation development

The variation in fine-root traits in response to soil resources (i.e., fine-root nutrient foraging strategy) is critical for plants to adapt to environmental changes and even win in intra- or interspecific resource competition. However, the patterns and driving mechanisms that change in fine-root traits and nutrient foraging strategies during the development of plantations remain unclear. We analyzed the relationships among fine-root traits, rhizosphere soil nutrient variables, and enzyme activities at four stages of Pinus massoniana plantations: 18 years (young), 30 years (middle-aged), 43 years (mature), and 63 years (overmature). We found that specific root area (SRA) and specific root length (SRL), which represent the speed of nutrient acquisition, decreased with stand development. Root tissue density (RTD) and specific root tip number (SRT), which represent the cost of nutrient foraging, decreased with stand development after peaking in the mature and middle-aged forests, respectively. Compared with young stands, fine-root organic carbon and total phosphorus contents decreased by 23 % in the overmature forest, but total nitrogen increased by nearly 50 %. Principal component analysis (PCA) showed that those fine-root traits significantly varied from young to overmature stands on PC1, and such a shift was consistent with changes in the growth period from fast (in the young forest) to slow (in the overmature forest) for P. massoniana. We suggest that those changes in fine-root traits along the PC1 represents a change in the fine-root foraging strategy of P. massoniana. This strategy (i.e., PC1) was negatively correlated with rhizosphere soil total nutrients (i.e., soil organic carbon and total nitrogen) and carbon (C)-nitrogen (N)-phosphorus (P) stoichiometry but positively correlated with available carbon, nitrogen, and C and N cycling-related enzyme activities. Specifically, the variation in rhizosphere soil NH4+ had the highest amount of explication (72 %) for the variation in the root nutrient foraging strategy. These results demonstrate that the intensity of the fine-root response to soil resources is related to the structure of rhizosphere soil available nutrients (especially soil available nitrogen) as well as carbon and nitrogen conversion enzyme activities during plantation development. Such relationships can reshape fine-root traits and change the fine-root nutrient foraging strategies during the development of plantations.