Tracer experiment and model evidence for macrofaunal shaping of microbial nitrogen functions along rocky shores

Abstract

Seawater microbes as well as those associated with macrobiota are increasingly recognized as a key feature affecting nutrient cycling. Tidepools are ideal natural mesocosms to test macrofauna and microbe interactions, and we quantified rates of microbial nitrogen processing using tracer enrichment of ammonium (15NNH4) or nitrate (15NNO3) when tidepools were isolated from the ocean during low intertidal periods. Experiments were conducted during both day and night as well as in control tidepools and those from which mussels had been removed, allowing us to determine the role of both mussels and daylight in microbial nitrogen processing. We paired time series observations of 15N enrichment in NH4+, NO2- and NO3- with a differential equation model to quantify multiple, simultaneous nitrogen transformations. Mussel presence and daylight increased remineralization and photosynthetic nitrogen uptake. When we compared ammonium gain or loss that was attributed to any tidepool microbes vs. photosynthetic uptake, microbes accounted for 32 % of this ammonium flux on average. Microbial transformations averaged 61 % of total nitrate use; thus, microbial activity was almost 3 times that of photosynthetic nitrate uptake. Because it accounted for processes that diluted our tracer, our differential equation model assigned higher rates of nitrogen processing compared to prior source-product models. Our in situ experiments showed that animals alone elevate microbial nitrogen transformations by 2 orders of magnitude, suggesting that coastal macrobiota are key players in complex microbial nitrogen transformations.