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340<br />

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

4.2 Relationship Between Hyphae in the Root and in the Soil<br />

The quantity of hyphae in the vicinity of roots associated with mycorrhizal<br />

roots can vary greatly (Sylvia 1986) and change with time (Bethlenfalvay et al.<br />

1982). Hyphae of AM fungi play key roles in the formation and functioning of<br />

mycorrhizas (Abbott et al. 1992). Hyphae in soil, originating from either an<br />

established hyphal network or from other propagules (spores, vesicles and<br />

root fragments), lead to the recognition and subsequent colonisation of roots.<br />

The distribution of hyphae and associated sporulation will determine where<br />

propagules are located in relation to newly formed roots. The roles of the<br />

hyphae in both P uptake and soil stabilisation are dependent on their distribution<br />

within the soil matrix and their interaction with the root <strong>surface</strong>.<br />

Abbott and Robson (1984) hypothesised for G. invermaium that low initial<br />

levels of infective hyphae in the soil would lead to small amounts of hyphae in<br />

soil relative to the amount formed inside the root. For high initial densities of<br />

infective hyphae of this fungus in soil, the exponential phase of colonisation<br />

of roots was expected to occur in parallel with extensive development of<br />

hyphae in the soil. In contrast, there was no similar relationship between the<br />

formation of hyphae in soil and within the root expected for S. calospora<br />

which consistently produced large amounts of hyphae in soil, irrespective of<br />

the density of hyphae within the root.<br />

There is relatively little information about the longevity of hyphae in soil<br />

(Sylvia 1988; Hamel et al. 1990). This would be important for predicting the<br />

activities of hyphae for both colonisation and P uptake. The majority of studies<br />

of mycorrhizas measure the extent of colonisation of roots at one point in<br />

time. This measure is of little value for understanding functional diversity of<br />

AM fungi because the AM fungi within the root may either be highly active or<br />

have ceased activity for some time. Most routine techniques for assessing<br />

mycorrhizas do not assess any functional attribute of the fungus. Therefore,<br />

care is required in extrapolating from levels of mycorrhizal root colonised to<br />

functional diversity of communities of AM fungi present in soil.<br />

5 Role of Arbuscular Mycorrhizal Fungi Associated with<br />

Roots in Soil Aggregation<br />

The AM fungal hyphae can enhance soil aggregation by using more than one<br />

mechanism. In clayey soils, entanglement of soil particles by hyphae can<br />

occur (Tisdall and Oades 1982; Oades 1984). There is insufficient hyphal<br />

length to extend around particles of sand and a more likely mechanism is<br />

cross-linking of particles by hyphae in sandy soils (Degens 1997). AM fungal<br />

hyphae may bind soil aggregates by exuding a glycoprotein (Wright et al.<br />

1996; Wright and Upadhyaya 1998). Some information on species differences<br />

in soil aggregation is available which indicates that AM fungi commonly pro-

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