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170<br />

MARK A. BERNARDS, LINA F. YOUSEF, ROBERT W. NICOL<br />

rhizosphere secondary chemicals in general may be serving as signal molecules for<br />

plant pathogens in a manner analogous to Rhizobium-legume and tree-mycorrhizae<br />

symbioses (Peters and Long, 1988; Larange et al., 2001). The ability of rhizosphere<br />

phytochemicals to fulfill a molecular need for the pathogen or to induce cellular<br />

pathways important for pathogenesis or virulence is a relatively unexplored avenue of<br />

plant-microbe allelopathy, even though this has obvious implications for diseases in<br />

economically important plants.<br />

Anecdotal evidence suggests that ginseng crops suffer from a replant syndrome<br />

and a common farming practice is to avoid planting consecutive ginseng gardens in<br />

the same area. Although there is no data on the cause of the replant problem in<br />

ginseng, it has been suggested that pathogens could be involved (Reeleder et al.,<br />

1999). In fact the ginseng replant problem bears a similarity to the situation in apple<br />

orchards. Using our results, a replant syndrome in ginseng gardens could be postulated<br />

to have the same basic ingredients (i.e., phytochemicals, pathogens and antagonists)<br />

present in the model of the apple replant syndrome. For example, it is well<br />

established that the “dominant causal agents” (Mazzola, 1998) of the apple replant<br />

syndrome are a complex of soilborne fungal pathogens, especially species in the genera<br />

Cylindrocarpon, Fusarium, Phytophthora and Pythium (Mazzola 1999; Isutsa<br />

and Merwin, 2000) including C. destructans, P. cactorum and Py. irregulare (Braun,<br />

1991; 1995; Mazzola, 1998). Although pathogens play a primary role, antagonists<br />

have also been implicated through results obtained from their application as biocontrol<br />

microbes (Utkhede et al., 2001) or surveys of orchard soil (Mazzola, 1999). Finally,<br />

it has been suggested that an important component of apple replant syndrome is the<br />

induction of virulence in microbes by root exudates (Szabo and Wittenmayer, 2000).<br />

Similarly, unidentified allelochemicals may also be involved in the asparagus replant<br />

syndrome (Peirce and Colby, 1987; Hartung and Stephens, 1983; Pedersen et al.,<br />

1991). Therefore, a ginseng replant syndrome, like that of apple and asparagus,<br />

could be due to a phytochemical-mediated alteration in the soil microbial population<br />

resulting in the stimulation of a complex of pathogens coupled with an inhibition of<br />

beneficial microbes. In other words, allelopathy may play a role in year-to-year soilborne<br />

crop disease cycles as well as replant syndromes.<br />

6. CONCLUDING REMARKS<br />

Plant disease researchers often focus on causal agents in relative isolation. Our research<br />

further demonstrates that other host-derived factors may also play an important role<br />

in the etiology and severity of plant diseases and replant syndromes. That is, allelopathy<br />

may contribute to plant diseases through the ability of certain microbes to grow and/<br />

or survive preferentially in the phytochemical profile of the rhizosphere surrounding<br />

specific crop roots. Because pathogenicity is a common nutritional mode in microbes,<br />

it is not surprising that pathogens are adapted to the chemical environment of their<br />

host plants. Manipulation of rhizosphere chemistry, therefore, either through genetic<br />

modification of plants, identification and use of microbe-influencing genotypes or the

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