Nitrogen enrichment shifts functional genes related to nitrogen and carbon acquisition in the fungal community

Abstract

To better understand mechanisms of carbon (C) and nitrogen (N) dynamics under anthropogenic N enrichment, we examined frequencies of C- and N-targeting genes in litter fungi. In particular, we tested the hypothesis that N enrichment selects for C-targeting genes but against N-targeting genes, if fungi preferentially invest resources in acquisition of growth-limiting nutrients. We conducted a fully-factorial litter and microbial transplant in a N fertilization experiment in Southern California grassland. The transplant design enabled us to contrast direct effects of N fertilization in the environment, indirect effects of N-induced shifts in the microbial community, and indirect effects of N-induced changes in plant litter chemistry. For each treatment, we assessed frequencies of select well-annotated fungal functional genes: cellulose-targeting AA9 genes (for C acquisition) versus ammonium transporter genes and amino acid permease genes (for N acquisition). We found that our hypothesis was upheld only with regard to shifts in the microbial community. Specifically, when grown in the same environment and litter, fungi from the N-fertilized plots displayed greater frequencies of cellulose-targeting AA9 genes from basidiomycetes, but smaller frequencies of ammonium transporter genes and amino acid permease genes, when compared to fungi from the control plots. In contrast, N fertilization in the plot environment was associated with higher frequencies of amino acid permease genes and ammonium transporter genes. Likewise, plant litter from the N-fertilized plots selected for higher frequencies of ammonium transporter genes. Altogether, we found fairly inconsistent effects of N enrichment on fungal functional genes related to C and N acquisition. Even if the genetic capacity of the fungal community to acquire C versus N changes owing to shifts in the microbial community, direct effects of N fertilization and indirect effects of litter chemistry may offset the response.