, Volume 14, Issue 4, pp 519-532

First online:

Open Access This content is freely available online to anyone, anywhere at any time.

Conditions Under Which Nitrogen Can Limit Steady-State Net Primary Production in a General Class of Ecosystem Models

  • Duncan N. L. MengeAffiliated withNational Center for Ecological Analysis and SynthesisDepartment of Ecology and Evolutionary Biology, Princeton University Email author 


Human activity is drastically altering global nitrogen (N) availability. The extent to which ecosystems absorb additional N—and with it, additional CO2—depends on whether net primary production (NPP) is N-limited, so it is important to understand conditions under which N can limit NPP. Here I use a general dynamical model to show that N limitation at steady-state—such as in old-growth forests—depends on the balance of biotically controllable versus uncontrollable N inputs and losses. Steady-state N limitation is only possible when uncontrollable inputs (for example, atmospheric deposition) exceed controllable losses (for example, leaching of plant-available soil N), which is the same as when uncontrollable losses (for example, leaching of plant-unavailable soil N) exceed controllable inputs (biological N fixation). These basic results are robust to many model details, such as the number of plant-unavailable soil N pools and the number and type of N fixers. Empirical data from old-growth tropical (Hawai’i) and temperate (Oregon, Washington, Chile) forests support the model insights. Practically, this means that any N fixer—symbiotic or not—could overcome ecosystem N limitation, so understanding N limitation requires understanding controls on all N fixers. Further, comparing losses of plant-available N to abiotic inputs could offer a rapid diagnosis of whether ecosystems can be N-limited, although the applicability of this result is constrained to ecosystems with a steady-state N cycle such as old-growth forests largely devoid of disturbance.


nitrogen fixation lichen ecosystem theory dissolved organic nitrogen nitrogen deposition biogeochemical theory nitrogen loss