Soil extracellular enzyme activities, soil carbon and nitrogen storage under nitrogen fertilization: A meta-analysis

Nitrogen (N) fertilization affects the rate of soil organic carbon (SOC) decomposition by regulating extracellular enzyme activities (EEA). Extracellular enzymes have not been represented in global biogeochemical models. Understanding the relationships among EEA and SOC, soil N (TN), and soil microbial biomass carbon (MBC) under N fertilization would enable modeling of the influence of EEA on SOC decomposition. Based on 65 published studies, we synthesized the activities of α-1,4-glucosidase (AG), β-1,4-glucosidase (BG), β-D-cellobiosidase (CBH), β-1,4-xylosidase (BX), β-1,4-N-acetyl-glucosaminidase (NAG), leucine amino peptidase (LAP), urease (UREA), acid phosphatase (AP), phenol oxidase (PHO), and peroxidase (PEO) in response to N fertilization. The proxy variables for hydrolytic C acquisition enzymes (C-acq), N acquisition (N-acq), and oxidative decomposition (OX) were calculated as the sum of AG, BG, CBH and BX; AG and LAP; PHO and PEO, respectively. The relationships between response ratios (RRs) of EEA and SOC, TN, or MBC were explored when they were reported simultaneously. Results showed that N fertilization significantly increased CBH, C-acq, AP, BX, BG, AG, and UREA activities by 6.4, 9.1, 10.6, 11.0, 11.2, 12.0, and 18.6%, but decreased PEO, OX and PHO by 6.1, 7.9 and 11.1%, respectively. N fertilization enhanced SOC and TN by 7.6% and 15.3%, respectively, but inhibited MBC by 9.5%. Significant positive correlations were found only between the RRs of C-acq and MBC, suggesting that changes in combined hydrolase activities might act as a proxy for MBC under N fertilization. In contrast with other variables, the RRs of AP, MBC, and TN showed unidirectional trends under different edaphic, environmental, and physiological conditions. Our results provide the first comprehensive set of evidence of how hydrolase and oxidase activities respond to N fertilization in various ecosystems. Future large-scale model projections could incorporate the observed relationship between hydrolases and microbial biomass as a proxy for C acquisition under global N enrichment scenarios in different ecosystems.