A “toy model” analysis of causes of nitrogen limitation in terrestrial ecosystems

Abstract

Nitrogen (N) limitation to net primary production is widespread and influences the responsiveness of ecosystems to many components of global environmental change. Logic and both simple simulation (Vitousek and Fieldin in Biogeochemistry 46: 179–202, 1999) and analytical models (Menge in Ecosystems 14:519–532, 2011) demonstrate that the co-occurrence of losses of N in forms that organisms within an ecosystem cannot control and barriers to biological N fixation (BNF) that keep this process from responding to N deficiency are necessary for the development and persistence of N limitation. Models have focused on the continuous process of leaching losses of dissolved organic N in biologically unavailable forms, but here we use a simple simulation model to show that discontinuous losses of ammonium and nitrate, normally forms of N whose losses organisms can control, can be uncontrollable by organisms and can contribute to N limitation under realistic conditions. These discontinuous losses can be caused by temporal variation in precipitation or by ecosystem-level disturbance like harvest, fire, and windthrow. Temporal variation in precipitation is likely to increase and to become increasingly important in causing N losses as anthropogenic climate change proceeds. We also demonstrate that under the conditions simulated here, differentially intense grazing on N- and P-rich symbiotic N fixers is the most important barrier to the responsiveness of BNF to N deficiency.