Stable isotopes as tracers in aquatic ecosystems

The addition of stable isotopes (SI) of 13C and 15N has been used to study several aquatic processes, thus avoiding environmental disturbance by the observer. This approach, employed for the last three decades, has contributed to expanding our knowledge of food-web ecology and nutrient dynamics in aquatic systems. Currently, SI addition is considered a powerful complementary tool for studying several ecological and biogeochemical processes at the whole-aquatic-ecosystem scale, which could not be addressed otherwise. However, their contributions have not been considered jointly nor have they been evaluated with a view to assessing the reliability and scope of their results from an ecosystem perspective. We intend to bridge this gap by providing a comprehensive review (78 scientific publications reporting in situ 13C/15N additions at the whole-aquatic-ecosystem scale) addressing the main results arising from their use as tracers. Specifically, we focus on: (i) reasons for SI additions at the whole-ecosystem scale to study ecological processes, (ii) the paradigms resulting from its use and the insights achieved, (iii) uncertainties and drawbacks arising from these SI addition experiments, and (iv) the potential of this approach for tackling new paradigms. SI tracer addition at the ecosystem scale has provided new functional insights into numerous ecological processes in aquatic sciences (importance of subsidies in lakes; heterotrophy dominance in benthic food webs in lakes, wetlands and estuaries; the decrease in N removal efficiency in most aquatic ecosystems due to anthropogenic alteration; the recognition of hyporheic zones and floodplains as hot spots for stream denitrification; and high rates of internal N recycling in tidal freshwater marshes). However, certain constraints such as the high cost of isotopes, the maintenance of the new isotopic steady state, and avoidance of biomass changes in any compartment or pool during tracer addition bear witness to the difficulties of applying this approach to all fields of aquatic ecology and ecosystems. The future development of this approach, rather than expanding to larger and complex aquatic ecosystems, should include other stable isotopes such as phosphorus (P18O4).