The influence of macrofaunal burrow spacing and diffusive scaling on sedimentary nitrification and denitrification: An experimental simulation and model approach

The influence of burrow spacing on sedimentary nitrification and denitrification was simulated experimentally using sediment plugs of different thicknesses immersed in aerated seawater reservoirs. Different plug thicknesses mimic different distances between oxygenated burrow centers and produce similar changes in aerobic-anaerobic reaction balances as a function of diffusive transport scaling. The thicknesses used were roughly equivalent to transport scales (interburrow spacing) that could be produced by burrow abundances of similar to400 to 50,000 m(-2), depending on burrow lumen radii (e.g., 0.05-1 cm). Following the exposure of anoxic sediment plugs to aerated water, an efficient aerobic nitrification zone was established within the first similar to2-3 millimeters of sediment. At pseudo-steady state, the thinnest plug (2 mm) simulating highest burrow density, was entirely oxic and the denitrification rate nil. Denitrification was stimulated in anoxic regions of the thicker plugs (5, 10, and 20 mm) compared to the initial value in experimental sediment. Maximum nitrification rates and the highest denitrification/nitrification ratio between oxic nitrification and adjacent denitrification zones occurred for the intermediate plug thickness of 5 mm. Of the oxic/anoxic composites, the thickest plug showed the least efficient coupling between nitrification/denitrification zones (lowest denitrification/nitrification ratio). Both the thickness of the oxic layer and the total net remineralization of dissolved inorganic N varied inversely with plug thickness. A set of diffusion-reaction models was formulated assuming a range of possible nitrification kinetic functions. All model forms predicted optimal nitrification-denitrification and ammonification-denitrification coupling with relative oxic-anoxic zonation scales comparable to intermediate plug thicknesses (5-6 mm). However, none of the commonly assumed kinetic forms for nitrification could produce the observed NO3- profiles in detail, implying that natural sediment populations of nitrifiers may be less sensitive to O-2 than laboratory strains. Our experimental and model results clearly show that rates of N remineralization and the balance between stimulation/inhibition of denitrification are highly dependent on sedimentary biogenic structure and the particular geometries of irrigated burrow distributions.