Root-soil facilitation in mixed Eucalyptus grandis plantations including nitrogen-fixing species

Abstract

The success of mixed plantation systems is ultimately the net result of positive and negative interactions, including belowground interactions, among the respective species. Despite increasing knowledge on positive interactions on biomass and productivity in mixed-species plantations, relatively little is known about the mechanisms underlying belowground interactions. Based on biodiversity data from four mixed Eucalyptus grandis plantations, we determined the optimal ecological facilitative interactions of mixed plantation, i.e., E. grandis and Alnus formosana. We then investigated in situ root exudation of E. grandis using a cuvette-based method and analysed root patterns, defence chemicals, bacterial communities, and biochemical properties of rhizosphere soils in mixed E. grandis with alder. Compared with pure Eucalyptus plantations, the fine root production by E. grandis was higher in subsoil layers (40–80 cm) in mixed plantations with alder. Autotrophic root respiration and associated enzymes were also increased in mixed plantations. Root exudates and defence allelochemicals of E. grandis changed in the presence of alder. Compared with the pure Eucalyptus plantations, E. grandis roots in mixed plantations reduced the release of potential allelochemicals, such as phenolic acids, flavonoids, unsaturated lactone, and glycosides. Methyl jasmonate, a common signal chemical, was significantly decreased in mixed-species plantations. The bacterial community of E. grandis rhizosphere soil was improved in mixed stands and recruited more nitrifying, N-fixing, and cellulose-decomposing bacteria, such as family Nitrosomonadaceae, genera MND1, Marmoricola, and Rikenellaceae RC9 gut groups. Thus, belowground ecological facilitative interactions occurred in mixed plantations with alder, which were due to E. grandis altering its rooting pattern, reducing the levels of released allelochemicals, recruiting more beneficial bacteria, and improving the biochemical properties of rhizosphere soil. This mechanism may be useful in reforestation programs for Eucalyptus monocultures that are suffering from problems associated with low biodiversity and reduced soil fertility.