Effects of soil type, fertilization and drought on carbon allocation to root growth and partitioning between secondary metabolism and ectomycorrhizae of Betula papyrifera

Paper birch (Betula papyrifera Marsh) seedlings were grown in a greenhouse in either subsoil or topsoil in factorial combination with two fertilization and drought regimes to investigate how different soil environments and nutrient availability drive belowground partitioning between growth, secondary metabolism and ectomycorrhizal (EM) associations, and impact drought tolerance of paper birch. Root and total seedling dry biomass, starch, soluble sugars, soluble phenolics, lignin and EM abundance were quantified. In unfertilized topsoil, total plant biomass and root biomass were approximately nine times higher than in unfertilized subsoil, but the root weight ratios did not differ between soils. Root soluble phenolics and lignin were higher in unfertilized subsoil than in unfertilized topsoil, whereas EM abundance was significantly higher in unfertilized topsoil than in unfertilized subsoil. In topsoil, fertilization decreased root biomass and EM abundance and increased root phenolics and lignin. In contrast, fertilization of subsoil increased root biomass but decreased root phenolics and lignin, while EM abundance was unaffected. In both soil types, fertilization reduced root weight ratios. Across soil types, EM abundance was negatively correlated with root soluble sugars, root phenolics and lignin, but this was driven mainly by the responses in the topsoil treatment. Our results show that soil fertility mediates carbon tradeoffs among defense, growth and EM associations.