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384<br />

Uwe Nehls<br />

the utilization of alternative nitrogen sources under nitrogen limitation (e.g.,<br />

Tazebay 1997). Nitrogen-dependent gene repression is presumably regulated<br />

by the internal nitrogen status of cells, and not the external nitrogen availability.<br />

Either, the intracellular ammonium concentration (Ter Schure et. al.<br />

2000) and/or the activity of the glutamine synthetase (Sophianopoulou and<br />

Diallinas 1995) are supposed to sense the endogenous nitrogen status.<br />

In ectomycorrhizal fungi, nitrogen importer gene expression is presumably<br />

also regulated by the internal nitrogen status of the hyphae (Nehls et. al. 1999;<br />

Javelle et al. 2001; Wipf et al. 2002). This could indicate how nitrogen uptake by<br />

soil-growing hyphae and nitrogen export by hyphae of the Hartig net might<br />

be managed (Fig. 3). Since the nitrogen content of forest soil is quite low and<br />

part of the nitrogen is transported to other parts of the growing fungal colony<br />

(e.g., mycorrhizas), soil-growing hyphae are presumably nitrogen-limited,<br />

resulting in a low endogenous nitrogen status and a strong expression of<br />

nitrogen importer genes. On the other hand, mycorrhizas are well supplied<br />

with nitrogen by soil-growing hyphae, thus revealing a high nitrogen status<br />

and a strongly reduced nitrogen importer gene expression. This nitrogen-<br />

Fig. 3. Regulation of fungal nitrogen uptake from soil and nitrogen excretion at the<br />

<strong>plant</strong>/fungus interface: a model. Nitrogen export to other parts of the fungal colony<br />

together with a low nitrogen content in soil results in a low endogenous nitrogen state in<br />

soil growing hyphae. In consequence, nitrogen importer genes are highly expressed and<br />

nitrogen uptake capacity is high. Nitrogen import from soil growing hyphae causes a<br />

high endogenous nitrogen status in hyphae of the Hartig net. This results in a repression<br />

of nitrogen importer gene expression and, together with posttranslational inactivation<br />

processes, in a low nitrogen uptake capacity. Together with export mechanisms, this<br />

leads to a net export of nitrogen at the <strong>plant</strong>/fungus interface

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