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Trichoderma  is a fungal genus that includes species that are currently being used as biological control agents and/or as biofertilizers. In addition to the direct application of  Trichoderma  spp. as biocontrol agents in plant protection, recent studies have focused on the beneficial responses exerted on plants, stimulating the growth, activating the defenses, and/or improving nutrient uptake. The  amdS  gene, encoding an acetamidase of  Aspergillus , has been used as a selectable marker for the transformation of filamentous fungi, including  Trichoderma  spp., but the physiological effects of the introduction of this gene into the genome of these microorganisms still remains unexplored. No evidence of  amdS  orthologous genes has been detected within the  Trichoderma  spp. genomes and the  amdS  heterologous expression in  Trichoderma harzianum  T34 did not affect the growth of this fungus in media lacking acetamide. However, it did confer the ability for the fungus to use this amide as a nitrogen source. Although a similar antagonistic behavior was observed for T34 and  amdS  transformants in dual cultures against  Rhizoctonia solani, Botrytis cinerea , and  Fusarium oxysporum , a significantly higher antifungal activity was detected in  amdS  transformants against  F. oxysporum , compared to that of T34, in membrane assays on media lacking acetamide. In  Trichoderma -tomato interaction assays,  amdS  transformants were able to promote plant growth to a greater extent than the wild-type T34, although compared with this strain the transformants showed similar capability to colonize tomato roots. Gene expression patterns from aerial parts of 3-week-old tomato plants treated with T34 and the  amdS  transformants have also been investigated using  GeneChip Tomato Genome Arrays . The downregulation of defense genes and the upregulation of carbon and nitrogen metabolism genes observed in the microarrays were accompanied by (i) enhanced growth, (ii) increased carbon and nitrogen levels, and (iii) a higher sensitivity to  B. cinerea  infections in plants treated with  amdS  transformants as detected in greenhouse assays. These observations suggest that the increased plant development promoted by the  amdS  transformants was at expense of defenses.

frontiers in microbiology

Nitrogen Metabolism and Growth Enhancement in Tomato Plants Challenged with Trichoderma harzianum Expressing the Aspergillus nidulans Acetamidase amdS Gene