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Contents XVII<br />

6 Carbohydrates as Signal, Regulating Fungal<br />

Gene Expression in Ectomycorrhizas . . . . . . . . . . . . . 377<br />

7 Nitrogen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 380<br />

8 Utilization of Inorganic Nitrogen . . . . . . . . . . . . . . . 381<br />

9 Utilization of Organic Nitrogen . . . . . . . . . . . . . . . . 382<br />

10 Proteolytic Activities of Ectomycorrhizal Fungi . . . . . . . 383<br />

11 Uptake of Amino Acids . . . . . . . . . . . . . . . . . . . . . 383<br />

12 Regulation of Fungal Nitrogen Export in Mycorrhizas<br />

by the Nitrogen-Status of Hyphae . . . . . . . . . . . . . . . 385<br />

13 Carbohydrate and Nitrogen-Dependent Regulation<br />

of Fungal Gene Expression . . . . . . . . . . . . . . . . . . . 385<br />

14 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . 385<br />

References and Selected Reading . . . . . . . . . . . . . . . . . . . . . 386<br />

22 Nitrogen Transport and Metabolism<br />

in Mycorrhizal Fungi and Mycorrhizas . . . . . . . . . . . . 393<br />

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

and Bernard Botton<br />

1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . 393<br />

1.1 Ecological Significance of Ectomycorrhizas . . . . . . . . . 393<br />

1.2 Nitrogen Uptake and Translocation by Ectomycorrhizas . . 394<br />

2 Nitrate and Nitrite Transport . . . . . . . . . . . . . . . . . . 395<br />

2.1 Uptake Kinetics . . . . . . . . . . . . . . . . . . . . . . . . . 395<br />

2.2 Characterization of Nitrate and Nitrite Transporters . . . . 395<br />

3 Ammonium Transport . . . . . . . . . . . . . . . . . . . . . 398<br />

3.1 Physico-Chemical Properties of Ammonium:<br />

Active Uptake Versus Diffusion . . . . . . . . . . . . . . . . 398<br />

3.2 Physiology of Ammonium Transport in Ectomycorrhizas . . 399<br />

3.3 Isolation of Ammonium Transporter Genes . . . . . . . . . 400<br />

3.4 Regulation of the Ammonium Transporters . . . . . . . . . 400<br />

3.5 Other Putative Functions of Ammonium Transporters . . . 402<br />

4 Amino Acid Transport . . . . . . . . . . . . . . . . . . . . . 403<br />

4.1 Utilization of Amino Acids by Ectomycorrhizal Partners . . 403<br />

4.2 Molecular Regulation of Amino Acid Transport . . . . . . . 404<br />

5 Reduction of Nitrate to Nitrite and Ammonium . . . . . . . 405<br />

5.1 Reduction of Nitrate to Nitrite . . . . . . . . . . . . . . . . . 405<br />

5.2 Reduction of Nitrite to Ammonium . . . . . . . . . . . . . . 406<br />

5.3 Molecular Characterization of Nitrate Reductase<br />

and Nitrite Reductase . . . . . . . . . . . . . . . . . . . . . . 406<br />

6 Assimilation of Ammonium . . . . . . . . . . . . . . . . . . 409<br />

6.1 Role and Properties of Glutamate Dehydrogenase . . . . . . 410

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