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THE ALLELOPATHIC POTENTIAL OF GINSENOSIDES 161<br />

shown to be biologically active at their ecologically relevant concentration. Lastly,<br />

the allelopathic role of the compounds has to be demonstrated at the field level.<br />

3.1. The Host Plant<br />

3. GINSENG AND GINSENG SAPONINS<br />

American ginseng (Panax quinquefolius L.) is a native North American member of<br />

the Araliaceae, a family whose more than 800 species are found mostly in the tropics<br />

(Lawrence, 1951). Panax quinquefolius is a perennial understory herb that is associated<br />

with deciduous forests (Fountain, 1986; Anderson et al., 1993) and ranges from<br />

Ontario and Quebec, south to northern Florida and west to Minnesota (Small and<br />

Catling, 1999). The aboveground tissues senesce at the end of each growing season<br />

and estimates of the maximum age of this plant are 23-30 yr (Anderson et al., 1993)<br />

to >50 yr (Lewis and Zenger, 1982). Ginseng typically has one aerial stem, with<br />

three to five palmately compound leaves and an umbelliferous inflorescence. The<br />

main pollinators of the small greenish-white to greenish-yellow flowers are bees<br />

(Catling and Spicer, 1995) and the mature red fruits usually contain two seeds, but<br />

range from one to three seeds (Anderson et al., 1993; Schlessman, 1985). Before<br />

germinating, the seeds of American ginseng require an after-ripening period of one<br />

to two winters (Lewis and Zenger, 1982; Anderson et al., 1993). Panax quinquefolius<br />

is threatened in Canada (Small and Catling, 1999), probably due to over-collection of<br />

the root for economic gain.<br />

The main commercial product from ginseng is the taproot, which is sought for its<br />

purported medicinal properties. Commercially, the roots are harvested after 3-5 yr of<br />

intensive cultivation. American ginseng has been commercially cultivated in Canada<br />

since the late nineteenth century (Proctor and Bailey, 1987), under artificial shade or<br />

in interplanted woodlands. In 2002 Ontario, which is one of Canada’s largest provincial<br />

producers, exported over one thousand metric tons of ginseng root representing a<br />

value of almost $ 40 million (OMAF, 2003). However, commercial productivity is<br />

hindered by susceptibility to several fungal diseases of the leaves, stem, fruit and<br />

roots.<br />

3.2. Ginsenoside Structure and Biosynthetic Origins<br />

Ginsenosides are triterpenoid saponins primarily based on two tetracyclic dammarane<br />

ring structures: (20S)-protopanaxadiol and (20S)-protopanaxatriol (Figure 1), with<br />

one representative (Ro) derived from oleanoic acid. While more than 25 ginsenoside<br />

structures have been identified in Panax spp. (Fuzzati et al., 1999), six are considered<br />

major: the (20S)-protopanaxadiol-derived Rb 1 , Rb 2 , Rc and Rd and the (20S)protopanaxatriol-derived<br />

Re and Rg 1 . With the exception of a few monodesmosidic<br />

compounds (e.g., Rh 2 , Rg 3 , Rh 1 , Rf and Rg 2 ), ginsenosides are generally bisdesmosidic.<br />

(20S)-Protopanaxadiols are 3-O and 20-O diglycosides, while (20S)-protopanaxatriols

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