The ectomycorrhizal symbiosis: life in the real world

Ectomycorrhizas (ECM) are dual organs formed between the terminal feeder roots of many plant species and certain soil fungi. The species richness and taxonomic diversity of ECM symbionts is impressive: ca. 7-10,000 fungal and ca. 8,000 plant species may be capable of forming ECM. The latter are the dominant components of forest and woodland ecosystems over much of the earth's surface. The obligate nature of the symbiosis for ECM fungi has been brought into question by reports that some species produce sporocarps under field conditions in the absence of a host plant. We suggest that there is no unequivocal evidence to support this. The spread of tree roots is often underestimated and small, overlooked hosts such as dwarf Salix spp or sedges may explain the appearance of ECM sporocarps in vegetation apparently devoid of ECM hosts. Compared to plant material, the sporocarps of ECM fungi contain high concentrations of N and P. We show that it would take between 3 and 14 million mycorrhizal tips, or 1800 km of hyphae, to supply the N in one sporocarp of Boletus edulis. The mantle formed by the fungus over the root tip is the likely site of storage for the N and P required for sporocarp production, and we discuss the chemical and structural mechanisms developed on mantles by ECM fungi to defend this resource against fungivory.