Diffusion-Linked Microbial Metabolism in the Vadose Zone

Figure 7.1 is a schematic of nutrient and contaminant transformations and cycling in the vadose zone. As detailed in Harris and Arnold (1995), higher plants take up C, N, P, and S in their most oxidized forms and use, via photosynthesis, the Sun’s energy and low-energy electrons from the oxygen in water to convert the oxidized forms of these essential elements into the relatively high energy reduced forms comprising plant biomass. Following plant death, the biomass residues enter the soil and are attacked by soil organisms as a source of food. The plant residues are depolymerized and the reduced, high-energy monomers are assimilated in part into soil organism biomass, and in part are used as electron donors to combine with the most thermodynamically efficient electron acceptors for dissimilatory energy generation to drive growth and maintenance reactions. In aerobic zones, oxygen is the preferred electron acceptor as long as it is nonlimiting. Death of soil organisms produces dead biomass which re-enters the biological reactor. Ultimately, via respiration in aerobic soils, all the reduced C, N, P, and S materials are released as their oxidized forms, and oxygen is reduced to water to complete the cycle. Ideally, the cycle is conservative, particularly from the standpoint of nonleakage of nutrients, such as nitrate, into the groundwater. Similarly, contaminants entering the vadose zone, either as a function of agronomic use or by accident, should ideally be integrated into the natural nutrient cycles and converted to harmless by-products for assimilation and dissimilation by soil organisms and higher plants (Liu, 1994). Management of nutrient and contaminant transformations by the soil organisms requires a thorough understanding of the ecophysiological and solute transport ground rules that control the nature and rates of transformation options available to the soil organisms. In models of chemical transport and transformation through the vadose zone, colonies of microorganisms are frequently treated as a homogeneous biofilm reactor (Grant and Rochette, 1994). Often, modeling efforts are focused on environmental conditions external to the microbial colony. This consideration of the colony as a biofilm with relatively constant nutrient uptake rates ignores the growth differentiation that occurs as the colony develops