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21 Carbohydrates and Nitrogen: Nutrients and Signals in Ectomycorrhiza 385<br />

dependent repression of amino acid transporter gene expression (indicated<br />

by AmAAP1), together with posttranslational events (e.g., increased degradation<br />

of plasma membrane transport proteins) that are described for yeasts<br />

(Springael and Andre 1998), could thus result in a highly reduced fungal<br />

capacity for re-uptake of amino acids at the <strong>plant</strong>/fungus interface. In combination<br />

with efflux mechanisms (e.g., nitrogen leakage), this would thus result<br />

in a net export of nitrogen.<br />

13 Carbohydrate and Nitrogen-Dependent Regulation of<br />

Fungal Gene Expression<br />

Carbohydrates as well as nitrogen are essential components of biological molecules<br />

(e.g., amino acids or nucleotides), and obviously have a great impact on<br />

fungal gene expression (e.g., Gonzales et al. 1997).<br />

With regard to carbon and nitrogen nutrition, four different patterns of<br />

regulation have been observed in A. muscaria. The amino acid importer gene<br />

AmAAP1 is only regulated by nitrogen nutrition, while the hexose transporters<br />

AmMst1 and AmMst2 (Nehls et al. 1998) are only regulated by carbohydrate<br />

nutrition. On the other hand, AmProt1 (protease; Nehls et al. 2001b)<br />

and AmTPS1 (trehalose-6-phosphate synthase) are regulated by both nitrogen<br />

as well as carbon nutrition. Nevertheless, the impact of carbon and nitrogen<br />

nutrition differs significantly for both genes. While AmProt1 is mainly<br />

regulated by nitrogen, AmTPS1 is mainly regulated by carbon availability.<br />

Comparable gene expression patterns have been described for fungi (Gonzales<br />

et al. 1997) as well as <strong>plant</strong>s (Coruzzi and Zhou 2001), revealing a universal<br />

and phylogenetically old regulation strategy.<br />

14 Conclusions<br />

Since large EST projects of ectomycorrhizal model systems are currently<br />

under progress (Tagu and Martin 1995; Johansson et al. 2000; Voiblet et al.<br />

2001; Wipf et al. 2003), macro- and micro-array hybridization will enable an<br />

overview of the general impact of carbon and nitrogen nutrition on gene<br />

expression for different ectomycorrhizal fungi.<br />

Present data suggest that carbon- and nitrogen-dependent gene repression<br />

in ectomycorrhizal fungi is presumably similar to that of saprophytic<br />

ascomycetes (yeast, Neurospora). Ascomycotic model organisms could thus<br />

help to develop working models for ectomycorrhizal function (e.g., nitrogen<br />

uptake from soil and release at the <strong>plant</strong>/fungus interface; see Fig. 3) that<br />

could be investigated in turn in an ectomycorrhizal model system. In addition,<br />

differences, e.g., in carbon-dependent gene regulation for an ectomycorrhizal<br />

fungus (A. muscaria) and saprophytic ascomycetes (yeast, Neurospora)

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