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334<br />

M. Zakaria Solaiman and Lynette K. Abbott<br />

vanetti et al. 1993a). Generally, the amount, type and form of root exudates<br />

have the potential to influence fungal growth before the fungus meets the root<br />

as well as after the hyphae contact the root. In addition, the formation of<br />

appressoria and hyphal penetration of the root <strong>surface</strong> may involve recognition<br />

processes (Koske and Gemma 1992).<br />

3 Functioning of Arbuscular Mycorrhizas in Nutrient<br />

Exchange<br />

Nutrient exchange in arbuscular mycorrhizas can occur between the hyphae<br />

and host root cells. External hyphae penetrate the root <strong>surface</strong> and proceed<br />

into the root cortex forming arbuscules and coils (Saito 2000). The arbuscule<br />

is a complex intracellular hyphal structure formed in Arum-type mycorrhizas<br />

(Smith and Smith 1997). It has been assumed that the arbuscule is a likely site<br />

for symbiotic nutrient transfer in Arum-type mycorrhizas (Bonfante-Fasolo<br />

1987; Smith and Smith 1997), but carbon exchange may also occur across<br />

walls of non-arbuscular hyphae (Smith and Read 1997). Variation in arbuscule<br />

development in Arum-type mycorrhizas could reflect different characteristics<br />

of roots and fungi (Smith and Dickson 1991).<br />

Arbuscular mycorrhizal fungi with a range in demand for phosphorus have<br />

been isolated from south-western Australia. In particular, an isolate of S.<br />

calospora either increased or decreased <strong>plant</strong> growth depending on the phosphate<br />

status of the soil (Thomson et al. 1986). This fungus has a high demand<br />

for carbon from the <strong>plant</strong> relative to P transfer (Pearson et al. 1994), and forms<br />

considerably more external hyphae than some other fungi that are highly<br />

effective at enhancing P uptake across a range of P supply (Abbott and Robson<br />

1985). Indeed, under some circumstances, it can be inefficient in P transfer<br />

(Smith et al. 2000). In addition, S. calospora can interact with Glomus invermaium<br />

during colonisation of roots, restricting growth of G. invermaium in<br />

other parts of the same root system during some stages of its colonisation<br />

(Pearson et al. 1993). However, colonisation and activity of S. calospora can be<br />

stopped once sporulation has taken place (Pearson and Schweiger 1994),<br />

resulting in resumption of colonisation (and presumably P uptake and transfer)<br />

by G. invermaium. This example demonstrates the dynamics of activities<br />

of two AM fungi on root <strong>surface</strong>s, but the extent to which this occurs generally<br />

for all isolates of these or other species is not known. Unfortunately, relatively<br />

few isolates have been investigated for most species of AM fungi in any environment<br />

so the extent to which generalisations can be made, even for species,<br />

is unknown (Morton and Bentivenga 1994).<br />

In field soils, as several species of AM fungi would generally be involved in<br />

co-colonisation of roots, P uptake and transfer into the <strong>plant</strong> might be<br />

affected if one fungus interfered with colonisation by another. The outcome<br />

would depend on the functional diversity of the species present, i.e. their effi-

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