The canopy layer, a biogeochemical actor in the forest N-cycle

Atmospheric nitrogen (N) depositions have increased sharply since the industrial revolution. Numerous authors forecasted severe impacts on forest ecosystem services and functioning when new N input exceeds both biotic and abiotic holding capacity, in what is known as the N saturation process. However, most of the studies performed so far to quantify these impacts have largely neglected the potential role of the canopy and associated biota in biogeochemical processes. The present review collects and summarizes the existing literature about all the interactions of atmospheric N deposition and the canopy, evidencing different processes that may interfere with them. By summarizing the research of the last three decades, we found that a share of both wet and dry N deposition can be retained within the canopy, partially absorbed by the leaves or other aerial parts and utilized for plant metabolism, substantially changing the fate of N in the ecosystem. Focusing on canopy-associated biota, we analyzed how this compartment can represent a highly reactive layer and have an active role in different processes involved in the N cycle such as nitrification, fixation and volatilization. It emerges clearly that the canopy hosts a complex network of processes; however, despite this, most of the manipulative experiments failed to include the canopy into the studies. The documented manipulative experiments which included the canopy are mainly short-term and represent very few forest types. Moreover, we highlighted a lack of research concerning the interaction between the complex network of canopy processes, atmospheric N deposition and other global change factors, such as increasing atmospheric CO2 concentration, increasing temperature and drought events.