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THE ALLELOPATHIC POTENTIAL OF GINSENOSIDES 169<br />

5.2. Bioactivity of Ginsenosides at Ecologically Relevant Concentrations<br />

As emphasized in Section 1.3, before ginsenosides can be considered allelochemicals,<br />

it has to be demonstrated that they are in fact present in the rhizosphere soil at biologically<br />

active concentrations. Therefore, after demonstrating the presence of<br />

ginsenosides in the soil, in vitro bioassays were conducted using ginsenosides at an<br />

ecologically relevant concentration of 0.06%. At this level, ginsenosides were shown<br />

to remain bioactive. That is, the growth of Py. irregulare was significantly greater<br />

than control, while that of T. hamatum remained unchanged (Nicol et al., 2003).<br />

5.3. Plant-Fungal Allelopathy in Ginseng Gardens and Implications for Disease<br />

The discovery of ginsenosides in ginseng root exudates and rhizosphere soil at biologically<br />

active concentrations suggests that plant-fungal allelopathy could occur within<br />

this crop. This in turn could influence disease levels as the ginseng secondary chemicals<br />

potentially, and differentially, influence the growth of different groups of soilborne<br />

organisms (i.e., pathogens and antagonistic saprotrophs).<br />

Naturally-occurring soilborne antagonists can play a role in preventing or reducing<br />

disease levels in crops (Azad et al., 1987; Miller et al., 1989; Larkin et al., 1993).<br />

However, the applied use of Trichoderma spp. as a biocontrol agent often does not<br />

exert the desired level of disease control. One reason is that isolates of these biocontrol<br />

fungi often suffer from poor rhizosphere competence (Papavizas, 1982; Chao et al.,<br />

1986). Consequently, poor (natural) Trichoderma spp. competence in the rhizosphere<br />

of ginseng could lead to increased disease levels in the plant. Interestingly, in an<br />

attempt to address the issue of the lack of Trichoderma spp. rhizosphere competence,<br />

it was found that exudates from healthy plant roots did not support the growth of T.<br />

harzianum, but did in fact support that of Py. ultimum (Green et al., 2001). More<br />

evidence for the involvement of a soil factor in preventing successful biocontrol by<br />

Trichoderma spp. is found in the work of Hong et al. (2000) where several Trichoderma<br />

isolates were shown to inhibit C. destructans in vitro, but this biocontrol effect<br />

failed to materialize when applied to potted ginseng plants. Again, the specific mechanism<br />

involved in both of these reports was not identified, but our results suggest that<br />

plant-fungal allelopathy is one possible explanation.<br />

Ginsenoside-mediated stimulation of several major pathogens is likely to be<br />

involved in the eventual establishment of ginseng diseases. Three important root<br />

pathogens were repeatedly observed to grow better after the addition of ginsenosides<br />

to their growth medium (i.e., C. destructans, P. cactorum and Py. irregulare).<br />

Therefore, it can be hypothesized that the chemical environment of the ginseng<br />

rhizosphere favours fungal root pathogens over potential antagonists of these pathogens<br />

and that this has a direct consequence on the year-to-year disease levels in ginseng<br />

gardens. We are still working on the exact mechanism of growth stimulation in<br />

pathogens, but our preliminary results lead us to believe that in one case (i.e., Py.<br />

irregulare) the activity is extra-nutritional. For Py. irregulare, increased growth may<br />

be due to the ginsenosides or transformed ginsenosides acting as sterol analogs. Also,

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