Root traits and tree functional groups determine variability in root exudation and N uptake rates in mature temperate trees

While root exudation has the potential to affect soil biogeochemistry profoundly, the process is rarely quantified in mature, field-grown trees. We measured rates of carbon (C) exudation in 11 trees species that exhibit divergent root traits, including gymnosperms and angiosperms that associate with either arbuscular mycorrhizal (AM) or ectomycorrhizal (EcM) fungi. Our goal was to explore how tree species, plant functional groups and root traits collectively influence exudation patterns. Intraspecific variation in exudation rates was larger than interspecific variation, and neither functional groups nor morphological traits alone could sufficiently explain variation in this flux. EcM-associated gymnosperms exuded 2.4 times more C than EcM angiosperms and 1.5 times more than AM gymnosperms. Exudation rates correlated positively with specific root length (SRL) and specific root area (SRA), and were correlated with root tissue density and root diameter in EcM-associated species. Mixed-effect models revealed that exudation rates were best determined by a combination of phylogenetic group, tree-mycorrhizal type and SRA, though a large portion of unexplained variation suggests that contemporary environmental and local edaphic conditions are likely important. Collectively, our results reveal that exudation is a complex physiological process governed by multiple factors and cannot be fully explained by functional groups or root traits alone. Instead, a combined consideration of these factors and new experimental approaches may be needed before exudation patterns can be linked to plant trait frameworks and incorporated into large-scale models.