INVITED REVIEW: PART OF AN INVITED ISSUE ON TREE NUTRITION Phosphorus nutrition of mycorrhizal trees

Globally, phosphorus (P) limits productivity of trees in many forests and plantations especially in highly weathered, acidic or calcareous profiles. Most trees form mycorrhizal associations which are prevalent in the organic and mineral soil horizons. This review critically examines mechanisms that enhance the acquisition of P by tree roots. Mycorrhizal roots have a greater capacity to take up phosphate (Pi) from the soil solution than non-mycorrhizal root tips. Factors that contribute to this include the extent of extraradical hyphal penetration of soil and the physiology and biochemistry of the fungal/soil and fungal/plant interfaces. Ectomycorrhizal (ECM) trees are likely to benefit from association with basidiomycetes that possess several high-affinity Pi transporters that are expressed in extraradical hyphae and whose expression is enhanced by P deficiency. To understand fully the role of these putative transporters in the symbiosis, data regarding their localization, Pi transport capacities and regulation are required. Some ECM fungi are able to effect release of Pi from insoluble mineral P through excretion of lowmolecular-weight organic anions such as oxalate, but the relative contribution of insoluble P dissolution in situ remains to be quantified. How the production of oxalate is regulated by nitrogen remains a key question to be answered. Lastly, phosphatase release from mycorrhizas is likely to play a significant role in the acquisition of Pi from labile organic forms of P (Po). As labile forms of Po can constitute the major fraction of the total P in some tropical and temperate soils, a greater understanding of the forms of Po available to the phosphatases is warranted.