bBecause microorganisms have different abilities to utilize nitrogen (N) through various assimilatory and dissimilatory pathways, microbial composition and diversity likely influence N cycling in an ecosystem. Terrestrial plant litter decomposition is often limited by N availability; however, little is known about the microorganisms involved in litter N cycling. In this study, we used metagenomics to characterize the potential N utilization of microbial communities in grassland plant litter. The frequencies of sequences associated with eight N cycling pathways differed by several orders of magnitude. Within a pathway, the distributions of these sequences among bacterial orders differed greatly. Many orders within the Actinobacteria and Proteobacteria appeared to be N cycling generalists, carrying genes from most (five or six) of the pathways. In contrast, orders from the Bacteroidetes were more specialized and carried genes for fewer (two or three) pathways. We also investigated how the abundance and composition of microbial N cycling genes differed over time and in response to two global change manipulations (drought and N addition). For many pathways, the abundance and composition of N cycling taxa differed over time, apparently reflecting precipitation patterns. In contrast to temporal variability, simulated global change had minor effects on N cycling potential. Overall, this study provides a blueprint for the genetic potential of N cycle processes in plant litter and a baseline for comparisons to other ecosystems.
Microorganisms play a key role in the decomposition of terrestrial plant litter (1-3), a process that controls the balance of plant carbon (C) released into the atmosphere as CO 2 with that stored in the soil. Less often considered is the role that litter microorganisms play in nitrogen (N) cycling. The N available to microorganisms degrading plant litter comes from several sources. One source is organic N bound in plant tissues and microorganisms. Because the average C/N ratio is much higher in plant litter than in microbial decomposers, N availability is thought to limit litter decomposition (4-6). Fungal hyphae can further translocate N from the soil into plant litter (7). And in some ecosystems, atmospheric deposition of inorganic N from human-driven nitrogen oxide (NO x ) emissions can also be an important source (8-11).Microbes can rapidly alter the forms of N in plant litter through a variety of different N cycle pathways, and these changes in N availability can feed back to influence overall ecosystem functioning (12, 13). During decomposition, bacteria utilize N in both assimilatory and dissimilatory pathways. Assimilatory pathways require energy and lead to the conversion of inorganic N to organic N in microbes (e.g., utilizing N for protein, nucleic acid, and cellular component assembly). Dissimilatory pathways use N compounds to provide energy to microbes. Thus, the pathways by which microbes use N affect the fate of N in the ecosystem, specifically whether it is converted into microbi...