The arbuscular mycorrhizal symbiosis is mutualistic, based on reciprocal transfer of P from the fungus to the plant and carbon from the plant to the fungus. Thus P is a most important `currency' in the symbiosis. After absorbing P from the soil solution, the fungi first incorporate it into the cytosolic pool, and the excess P is transferred to the vacuoles. The vacuolar P pool probably plays a central role in P supply to the plant. The main forms of inorganic P in fungal vacuoles are orthophosphate and polyphosphate, but organic P molecules may also be present. Long distance translocation of P from the site of uptake in the external mycelium to the site of transfer to the plant is probably achieved via transfer of vacuolar components. This transport would be mediated either by protoplasmic streaming or the motile tubular vacuole-like system. The site of release of P into the interfacial apoplast and thence to the plant is most probably the fungal arbuscules. The biochemical and biophysical processes involved in P metabolism and transfer between cellular compartments in the symbiosis are currently not well understood. Some recent investigations of substrate specificities of phosphatase-type enzymes in AM fungi and other eukaryotic microorganisms, however, have shed new light on earlier results and permit the construction of a hypothetical scheme of P-flow, including possible regulatory factors. Steps in this scheme are experimentally testable and should stimulate future research.
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