Discriminating between Seasonal and Chemical Variation in Extracellular Enzyme Activities within Two Italian Beech Forests by Means of Multilevel Models

Enzymes play a key-role in organic matter dynamics and strong scientific attention has been given to them lately, especially to their response to climate and substrate chemical composition. Accordingly, in this study, we investigated the effects of chemical composition and seasons on extracellular enzyme activities (laccase, peroxidase, cellulase, chitinase, acid phosphomonoesterase, and dehydrogenase) by means of multilevel models within two Italian mountain beech forests. We used chemical variables as the fixed part in the model, season as random variation and layers (decomposition continuum for leaf litter and 0-5, 5-15, 15-30, and 30-40 cm for soil) as nested factors within the two forests. Our results showed that seasonal changes explained a higher amount of variance in enzyme activities compared to substrate chemistry in leaf litter, whereas chemical variation had a stronger impact on soil. Moreover, the effect of seasonality and chemistry was in general larger than the differences between forest sites, soils, and litter layers.