Seasonal Belowground Ecosystem and Eco-enzymatic Responses to Soil pH and Phosphorus Availability in Temperate Hardwood Forests

Atmospheric acid deposition can increase phosphorus (P) limitation in temperate hardwood forests by increasing N availability, and therefore P demand, and/or by decreasing pH and occluding inorganic P. However, only recently have studies demonstrated that P limitation can occur in temperate forests and very little is known about the temporal aspects of P dynamics in acidic forest soils and how seasonal shifts in nutrient availability and demand influence microbial investment in extracellular enzymes. The objectives of this study were to investigate how P availability and soil pH influence seasonal patterns of nutrient cycling and soil microbial activity in hardwood forests that experience chronic acid deposition. We experimentally manipulated soil pH, P, or both for three years and examined soil treatment responses in fall, winter, spring, early summer, and late summer. We found that site (glaciated versus unglaciated) and treatment had the most significant influence on nutrient pools and cycling. In general, nutrient pools were higher in glaciated soils than unglaciated for measured nutrients, including total C and N (2-3 times higher), extractable inorganic nitrogen, and readily available P. Treatment had no impact on total C and N pools in either region, but did affect other measured nutrients such as ammonium, which was greatest in the elevated pH treatment for both sites. As expected, readily available P pools were highest in the elevated P treatments (3 fold increase in both sites), but raising pH decreased available P pools in the glaciated site. Raising soil pH increased both net N mineralization rates and net P mineralization rates, regardless of site. Nitrification responses were complex, but we observed an overall significant nitrification increase under elevated pH, particularly in the growing season. Extracellular enzyme activity showed more seasonal patterns than site and treatment effects, exhibiting significant growing season activity reductions for all enzymes measured. Phosphatase enzymes did not respond to our treatments and were generally greatest in the unglaciated soils, particularly in winter and spring. Enzyme stoichiometric relationships revealed that soil microbial populations in the glaciated site were consistently less P and N-limited than unglaciated sites but this difference was less pronounced during the growing season. The trajectory of nutrient limitation in response to soil pH and P availability was highly variable, but we observed that enzyme ratios in the early summer were particularly shifted relative to other seasons suggesting that both sites were increasingly P and N-limited during this period. Overall, our results suggest that ecosystem and microbial responses to soil pH and P availability vary with both season and site history and that more spatially and temporally explicit observations are needed to improve our understanding of ecosystem acidification, nutrient limitation, and the cost-benefit relationships of microbial investments in extracellular enzymes.