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Carbon Metabolism, Lipid Composition and Metabolism 161<br />

isolated to date range from nutritionally to morphologically important<br />

genes (review in Harrier 2001; Golotte et al. 2002). More recently, nontargeted<br />

approaches have been used to isolate AM fungal genes. They<br />

include differential display, differential screening, and large-scale EST<br />

sequencing (Gollotte et al. 2002). The use of mycorrhizal defective mutants<br />

and the application of high-density array technology are promising<br />

tools to isolate genes involved in the different steps of plant–AM interactions.<br />

Concerning genes involved in carbon metabolism and transport, the<br />

nature of the plant transporters or fungal transporters involved in the<br />

movement of glucose from the cortical cells to the interface compartment<br />

and to arbuscules and hyphae are still unknown. A gene encoding<br />

a transmembrane sugar transporter (Mtst1) was cloned from mycorrhizal<br />

roots of Medicago truncatula. This transporter was designed as a hexose<br />

transporter by activity measured in yeast (Harrison 1996). In situ hybridization<br />

showed that the sugar transporter transcripts were induced<br />

in cells containing arbuscules and in adjacent cells, suggesting that the<br />

genemaybeactiveintheprocessofhexosetransporttowardscolonized<br />

plant cells. A phosphoglycerate kinase (PGK) cDNA was isolated from<br />

tomato mycorrhizas by differential display (Harrier et al. 1998). Quantitative<br />

immunoblotting using a polyclonal antibody specific for the G. mossae<br />

PGK protein revealed a significantly higher accumulation of the protein<br />

during symbiosis compared with presymbiotic development (Harrier and<br />

Sawczak 2000). These results suggested that there is a differential regulation<br />

of fungal genes during symbiosis. Analysis of the Gmpgk fungal promoter<br />

showed several regulatory elements homologous to carbon sourcecontrolled<br />

upstream activating elements from Saccharomyces cerevisiae<br />

(Harrier 2001).<br />

AM fungi could cause changes in the regulation of plant genes in roots.<br />

Ravnsknov et al. (2003) demonstrated a higher gene expression of both<br />

Sus1 and Sh1, the two isoforms of sucrose synthase cleaving sucrose into<br />

hexoses, in maize roots colonized by different AM fungal isolates. Higher<br />

sucrose synthase gene expression was not related to the concentrations<br />

of sucrose, reducing sugars or starch in the root tissue whereas increasing<br />

soil phosphorus concentrations decreased this gene expression. The<br />

higher gene expression of gene coding for sucrose synthase in AM roots<br />

was measured during the earliest phase of root colonization by fungal<br />

isolates.

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