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19 Functional Diversity of Arbuscular Mycorrhizal Fungi on Root Surfaces 335<br />

ciency in P uptake and transfer. Circumstantial evidence of seasonal variation<br />

in the function of different species of AM fungi present in a woodland environment<br />

has been clearly demonstrated (Merryweather and Fitter 1998b).<br />

There have been few direct measurements of P uptake by communities of AM<br />

fungi (Jakobsen et al. 2001), but it is difficult to attribute P transferred to the<br />

activity of a particular fungus within a community.<br />

The relative abundance of AM fungi in field-grown <strong>plant</strong>s can be manipulated<br />

by environmental and management processes that occur in the field,<br />

where dual AM fungal occupancy of roots is the norm (Merryweather and Fitter<br />

1998a). With manipulation of P supply, the important observations of<br />

Thomson et al. (1986) can be used to identify the P status of the <strong>plant</strong> at which<br />

there is a point of transition from growth enhancement to growth depression<br />

in <strong>plant</strong>s colonised by S. calospora. In contrast, G. invermaium did not display<br />

such a physiological transition in this study.<br />

3.1 Metabolic Activity During Mycorrhiza Formation<br />

Changes in metabolic activities during mycorrhiza formation provide evidence<br />

for hypotheses related to biochemical mechanisms of C and P exchange<br />

between symbionts (Saito 2000). These have mostly been examined by comparing<br />

mycorrhizal and non-mycorrhizal roots, but without identifying the<br />

mechanisms of nutrient exchange. Changes in quantity and concentration of<br />

soluble carbohydrates in roots have not shown consistent trends with mycorrhizal<br />

colonisation (Pakovsky 1989; McArthur and Knowels 1993; Pearson et<br />

al. 1994; Solaiman and Saito 1997).<br />

The localisation of alkaline phosphatase (ALPase) in arbuscular hyphae<br />

was observed by histochemical study (Saito 1995). ALPase has also been<br />

located in vacuoles in intraradical hyphae of AM fungi (Gianinazzi et al.<br />

1979), and its activity varied with environmental conditions (Jabaji-Hare et al.<br />

1990). A correlation between the number of ALPase-active arbuscules and P<br />

uptake by mycorrhizal <strong>plant</strong>s indicates that arbuscular ALPase plays a significant<br />

role in phosphorus transformation from the AM fungus to the host <strong>plant</strong><br />

(Tijssen et al. 1983). Histochemical observation of intraradical hyphae has<br />

located lipid and polyphosphate (polyP) granules in hyphae. Although polyP<br />

molecules are likely to play an important role in P translocation, their existence<br />

has been contradicted. Recently, a successive extraction method showed<br />

that a granular fraction of polyP was contained in hyphae of Gigaspora margarita<br />

(Solaiman et al. 1999). The contribution of polyP was calculated from<br />

these data and it has been concluded that the contribution is not significant<br />

(Smith et al. 2001).

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