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22 Nitrogen Transport and Metabolism in<br />

Mycorrhizal Fungi and Mycorrhizas<br />

Arnaud Javelle, Michel Chalot, Annick Brun<br />

and Bernard Botton<br />

1 Introduction<br />

1.1 Ecological Significance of Ectomycorrhizas<br />

Unlike most other organisms, <strong>plant</strong>s and fungi are restricted to their habitats,<br />

creating potential problems when nutritional conditions become limited. To<br />

cope with nutrient deficiencies, they have developed a variety of adaptations<br />

that enable them to respond to their internal nutritional status as well as to<br />

the external availability of nutrients.A strategy for <strong>plant</strong>s is mycorrhizal association,<br />

in which expanding mycorrhizal mycelia that grow outward from the<br />

mantle into the surrounding soil is a very efficient nitrogen scavenger owing<br />

to (1) its capacity to explore a larger soil volume than roots alone (Smith and<br />

Read 1997), (2) its ability to provide access to nitrogenous reserves contained<br />

in organic horizons (Chalot and Brun 1998) and (3) its greater capacity for<br />

uptake of nitrogenous compounds (Javelle et al. 1999; Wallanda and Read<br />

1999).<br />

This interconnected network of hyphae (or specialized aggregates, i.e., rhizomorphs)<br />

forms a supracellular compartment for the transport of nutrients<br />

from sites of nutrient capture to sites of nutrient utilization and transfer. It<br />

has been estimated that the external mycelium makes, by far, the greatest contribution<br />

to the overall potential absorbing <strong>surface</strong> area of pine seedlings<br />

inoculated with Pisolithus tinctorius or Cenococcum geophilum (Rousseau et<br />

al. 1994). Fungal hyphae have a number of advantages compared with roots;<br />

(1) hyphae have a low ratio of biomass to absorptive <strong>surface</strong> area and can easily<br />

be regenerated (Harley 1989; Rousseau et al. 1994), (2) they have been<br />

shown to rapidly colonize nutrient-rich sites (Carleton and Read 1991; Bending<br />

and Read 1995) and (3) because of their small diameter, they can exploit<br />

small pores inaccessible to roots. The symbiotic association of higher <strong>plant</strong>s<br />

with mycorrhizal fungi is considered to have been responsible for the colonization<br />

of land by <strong>plant</strong>s (Taylor and Osborn 1995).<br />

Plant Surface Microbiology<br />

A.Varma, L. Abbott, D. Werner, R. Hampp (Eds.)<br />

© Springer-Verlag Berlin Heidelberg 2004

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