Modeling trophic pathways, nutrient cycling, and dynamic stability in soils

Soil communities are compartmentalized into pathways of trophic interactions and nutrient flows that originate from plant roots, bacteria and fungi. The pathways differ in terms of the organisms that comprise them, the habitats that the organisms occupy and the rates by which the organisms process and transfer material and energy. The fungi, nematodes and arthropods within the fungal pathway live in air-filled pore spaces and water films, while the bacteria, protozoa, and nematodes within the bacterial pathway occupy water-filled pore spaces and water films. Organisms within the fungal pathway have longer generation times and process matter at slower rates than those within the bacterial pathway. Empirical studies have shown that under natural conditions the pathways co-exist in a stable manner. The relative sizes (indexed by the densities of organisms) and activities (indexed by nutrient-flow rates, excretion rates and respiration rates) of the pathways may change seasonally and in response to minor disturbances, but they persist. However, large anthropogenic and natural disturbances induce shifts in the relative sizes and activities of the pathways. Coincident with these shifts are reports of changes in the aboveground plant community and the availability and retention of plant limiting nutrients. We developed simple models of the bacterial and fungal pathways to explore the consequences of the observed shifts on the dynamic stability of the system. The more stable configurations occurred when there was a balance in the flow of nutrients between the two pathways. Large shifts in nutrient cycling and community structure towards either the fungal pathway or toward the bacterial pathway resulted in less stable or unstable configurations.