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78<br />

Ramesh Chander Kuhad et al.<br />

roots (Torrisi et al. 1999). Extra radical AM hyphae augment the uptake of<br />

nutrients from up to 12 cm away from the root <strong>surface</strong> (Cui and Caldwell<br />

1996).<br />

The network of hyphae may increase the availability of nutrients like N or<br />

P from locked sources by decomposing large organic molecules (George et al.<br />

1995). Mycorrhizal fungi are also known to develop bridges connecting the<br />

root with the surrounding soil particles to improve both nutrient acquisitions<br />

by the <strong>plant</strong> and soil structure (Varma 1995; Hodge 2000). Unlike N 2-fixing<br />

bacteria that function as biological fertilizers, AM fungi do not add P to the<br />

soil. They only improve its availability to the <strong>plant</strong>. There is evidence that<br />

phosphatase activity is higher in the rhizosphere around AM than in nonmycorrhizal<br />

roots. P uptake is enhanced with the increase in root colonization by<br />

mycorrhizae.A system of barter operates, the colonized <strong>plant</strong> provides photosynthate<br />

to the fungus, in return, its extraradical hypha makes more P available<br />

to the host (Merryweather and Fitter 1995). Plants rely more on AM when<br />

growing in soils deficient in P (Bationo et al. 2000). Depriving a <strong>plant</strong> in its<br />

natural environment of mycorrhizae on a long-term basis can also reduce P<br />

acquisition. Plants that are nonmycorrhizal invest more in their vegetative tissues<br />

like shoots and roots. In contrast, in mycorrhizal <strong>plant</strong>s, the functions of<br />

the roots are taken over by the AM hyphae, thereby permitting the host <strong>plant</strong><br />

to invest its resources in reproductive organs.<br />

Nitrogen occurs in the soil predominantly in the form of nitrate and<br />

ammonia, which is water-soluble and readily available for absorption. Studies<br />

with labelled N have revealed that the AM increases N uptake by <strong>plant</strong>s (Bijbijen<br />

et al. 1996; Faure et al. 1998; Mädder et al. 2000). AM fungal hyphae have<br />

been credited with the uptake and transfer of large amounts of N from the soil<br />

to the host (Johansen et al. 1996; Hodge et al. 2000). However, there is little reciprocal<br />

transfer of N from the <strong>plant</strong> to the fungi, which makes uptake and<br />

assimilation of N by the symbiont essential for its growth (Bijbijen et al. 1996).<br />

Since AM form underground hyphal links between <strong>plant</strong>s, N transfer between<br />

<strong>plant</strong>s by means of such links is possible. Using labelled 15 N, Frey and Schüepp<br />

(1993) demonstrated that N flows from Trifolium alexandrium to Zea mays<br />

via AM fungal network. AM are believed to enhance N 2-fixation by symbiotic<br />

legumes by increasing root and nodule biomass, N 2-fixation rates, root N<br />

absorption rates, and <strong>plant</strong> N and P content (Olesniewicz and Thomas 1999).<br />

Mycorrhizae have also been reported to be involved in the uptake of other<br />

micro- and macro-nutrients like K, S, Mg, Zn, Cu, Ca and Na (Díaz et al. 1996;<br />

Hodge 2000).<br />

Soil microorganisms, particularly saprophytic fungi affect the development<br />

and function of AM symbiosis. Fracchia et al. (2000) investigated the<br />

effect of the saprophytic fungus Fusarium oxysporum on AM colonization<br />

and <strong>plant</strong> dry matter was studied in greenhouse and field experiments using<br />

host <strong>plant</strong>s, maize, sorghum, lettuce, tomato, wheat, lentil and pea and AM<br />

fungi, Glomus mosseae, G. fasciculatum, G. intraradices, G. clarum and G.

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