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418<br />

A. Javelle et al.<br />

gen to <strong>plant</strong>s of Pinus contorta was also clearly demonstrated (Abuzinadah<br />

and Read 1986a, b; Abuzinadah et al.1986). The use of nitrogen sources not<br />

available to nonmycorrhizal <strong>plant</strong>s contributes, therefore, to an increased<br />

uptake of nitrogen by infected roots.<br />

7.2 Amino Acids Used as Nitrogen and Carbon Sources<br />

Utilization of amino acids by ectomycorrhizal symbionts and ectomycorrhizas<br />

may have important implications, not only for their nitrogen metabolism,<br />

but also for the overall carbon economy of the <strong>plant</strong>. Axenic mycelia of<br />

the ectomycorrhizal basidiomycete Suillus bovinus have been grown in liquid<br />

media in the presence of glucose as the only carbohydrate source and under<br />

such conditions, they produced similar amounts of dry weight with ammonia,<br />

with nitrate or with alanine, 60–80 % more with glutamate or glutamine, but<br />

about 35 % less with urea as the only exogenous nitrogen source (Grotjohann<br />

et al. 2000).<br />

Recently, the fate of carbon derived from alanine, glutamate and glutamine<br />

was investigated in the ectomycorrhizal fungus Paxillus involutus (Chalot et<br />

al. 1994a, b). The result of the 14 C tracer experiments suggested that the carbon<br />

skeletons derived from newly absorbed glutamate were mainly used for<br />

the synthesis of glutamine. The accumulation of [ 14 C]glutamate and the<br />

marked decrease of [ 14 C]glutamine under MSX treatment were consistent<br />

with a rapid utilization of glutamate by the glutamine synthetase (GS)<br />

enzyme. The newly absorbed, as well as the newly synthesized [ 14 C]glutamine<br />

were degraded into [ 14 C]glutamate, suggesting the operation of the glutamate<br />

synthase (GOGAT) enzyme. This was confirmed by the striking accumulation<br />

of [ 14 C]glutamine when the fungus was cultivated in the presence of azaserine,<br />

an inhibitor of GOGAT. In addition, a strong inhibition of glutamine utilization<br />

by aminooxyacetate indicated that glutamine catabolism in Paxillus<br />

involutus might involve a transamination process as an alternative pathway to<br />

GOGAT for glutamine degradation (Chalot et al. 1994a).<br />

The use of 14 C-labelled amino acids also showed a direct involvement of<br />

glutamate and glutamine in the respiration pathways, these amino acids being<br />

obviously channelled through the tricarboxylic acid (TCA) cycle and oxidized<br />

to CO 2. Feeding the fungus with [ 14 C]alanine resulted in a rapid labelling of<br />

pyruvate, citrate, succinate, fumarate and CO 2. Further labelling was detected<br />

in glutamate, glutamine and aspartate. The presence of aminooxyacetate completely<br />

suppressed 14 CO 2 evolution and decreased the flow of carbon to the<br />

Krebs cycle intermediates and amino acids, suggesting that alanine aminotransferase<br />

plays a key role in metabolizing alanine in Paxillus involutus<br />

(Chalot et al. 1994b).<br />

It has been shown by measuring enzyme capacities and metabolite pools<br />

that mycorrhization causes a re-arrangement of the main metabolic pathways

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