Biogeochemical niche: optimality, acclimation and adaptation principles in nutrient cycling under global change.

There are several processes underlying the shifts in organism's functions, species composition and ecosystem adaptation to the fast rates of environmental changes resulting from global change drivers. These environmental changes imply a shift in the use and cycling of resources, and in particular of nutrients, by organisms, communities and ecosystems. We will review the different use of bio-elements related to global change drivers such as climate change (warming and drought), increased concentrations of atmospheric CO2, or expansion of invasive species among others. Thereafter, we will discuss the resulting progressive change in nutrient cycling and its coupling with organism's, species, communities and ecosystem function in the frame of the biogeochemical niche hypothesis (Peñuelas et al., 2008; 2010). This hypothesis, based on the fact that each bio-element participates in different proportion in distinct organism's structures and functions, claims that each species has an optimal elemental composition and stoichiometry that allows reaching an optimal functioning. Species are nonetheless expected to exhibit a certain degree of stoichiometric flexibility (adaptive capacity) necessary to respond to environmental changes and competition, probably with a trade-off with stability. We will present data for the dominant tree species in Europe showing that the elemental foliar composition differences among species can be explained by their phylogenetic distances, current climate differences in their distribution areas and niche speciation in sympatric species, but also by some more recent human-driven impacts such as N deposition, thus showing the suitability and sensitivity of the "biogeochemical niche" concept to understand recent organism's, species, and ecosystem responses to novel environmental conditions imposed by human activity. We will finally discuss possible clues to improve the projections of ecosystem shifts in global change scenarios based on this concept.