Anthropogenic perturbations to the nitrogen cycle, primarily through use of synthetic fertilizers, have caused unprecedented increases in the emission of nitrous oxide (N2O) in recent decades. As a potent greenhouse gas, and an ozone depleting substance, understanding the sources and sinks of N2O is of vital importance. Nitrate (NO3-) reducing microbes are a primary contributor to the biotic production of N2O in anoxic regions of soil, marine systems, and wastewater treatment facilities through the process of denitrification. Thus, developing a better understanding of denitrifying microbial communities, and the environmental factors that influence N2O emissions may provide strategies to mitigate emissions in agriculture and wastewater treatment. Here, through comprehensive genome analysis, we show that pathway partitioning is a common strategy utilized by microbial communities to perform complete denitrification. Through detailed physiological characterization and kinetic modeling of a cooperative synthetic community (SynCom) assembled by pairing bacterial isolates from a field site heavily contaminated with NO3-, we also provide insight into the controls of N2O emissions. We demonstrate that members of this SynCom cooperate to perform complete denitrification through exchange of nitrite (NO2-) and nitric oxide (NO), and that community context drives global physiological changes in each member. We identify links between amino acid metabolism and denitrification activity as well as indicators of competition and amino acid exchange. We also show that NO2- toxicity with unbalanced growth of community members drives N2O production, suggesting that this SynCom provides a simplified, environmentally relevant, model of pathway partitioning in denitrifying communities. This SynCom should provide a framework with which to further explore how environmental context can impact cooperation and lead to the production of N2O