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The Geography of Phytochemical Races

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260 6 Oceanic Islands<br />

compounds with the styryl double bond reduced (making it the 2-phenylethyl function)<br />

as seen in 7,8-dihydrokawain [548], compounds with a second double bond in<br />

the lactone moiety, as seen in 5,6-dehydrokawain [549], are also known, and compounds<br />

in which the aromatic ring is substituted, such as in yangonin [550], which<br />

bears a methoxy function at the p-position. Several kavalactones are characterized<br />

by the methylenedioxy function, as seen in methysticin [551] and in dihydromethysticin<br />

[552]. Other members <strong>of</strong> the family combine these functional groups in<br />

various ways. <strong>The</strong> six compounds illustrated, however, are the most commonly<br />

encountered ones, <strong>of</strong>ten constituting 95% <strong>of</strong> the total lactone content <strong>of</strong> a particular<br />

cultivar.<br />

Kava has been the subject <strong>of</strong> detailed study by several workers, whose interests<br />

have included usage <strong>of</strong> the preparations in Hawaii (Titcomb, 1948); pyrone analysis<br />

(Young et al., 1966); physiology <strong>of</strong> action <strong>of</strong> the constituents (Hänsel, 1968); monographic<br />

study (Chew, 1972); cultivation (Lebot and Cabalion, 1986); origin <strong>of</strong> the<br />

Oceanian plant (Lebot and Lévesque, 1989); genetic control <strong>of</strong> kavalactone chemotypes<br />

(Lebot and Lévesque, 1996a); and relationships with the related P. wichmannii<br />

C. DC. (Lebot and Lévesque, 1996b). Comprehensive citation lists can be found in<br />

the papers by Lebot and Cabalion (1986) and Lebot and Lévesque (1989).<br />

Lebot and Lévesque’s 1989 work was based upon an exhaustive collection<br />

<strong>of</strong> plant material. Piper wichmannii was obtained from Papua New Guinea, the<br />

Solomon Islands, and Vanuatu, which comprises its natural range. Piper methysticum<br />

was collected from cultivated plots on three islands representing Micronesia,<br />

eight representing Melanesia, and 24 from Polynesia. In all, more than 240 individual<br />

plant acquisitions were subjected to chemical and morphological analysis.<br />

<strong>The</strong> fi rst problem addressed by Lebot and Lévesque (1989), in their comprehensive<br />

ethnobotanical study, dealt with the taxonomic status <strong>of</strong> P. methysticum and<br />

P. wichmannii C. DC. Problems that had to be dealt with included the nearly total<br />

absence <strong>of</strong> female plants, even in cultivation, and the complete lack <strong>of</strong> sexual reproduction<br />

<strong>of</strong> P. methysticum in cultivation, and consequently, the absence <strong>of</strong> seeds.<br />

Piper wichmannii, on the other hand, sets normal seed that germinate and develop<br />

normally. Furthermore, morphological differences are slight, certainly not suffi cient<br />

for specifi c recognition. Both species appear to be decaploids with 2n = 10x = 130<br />

chromosomes (Jose and Sharma, 1985; Okada, 1986). <strong>The</strong> high ploidy level in<br />

P. methysticum cannot be taken as explanation for the sterility <strong>of</strong> the cultivars, since<br />

P. wichmannii, also decaploid, reproduces normally. Electrophoretic studies <strong>of</strong> both<br />

species (Lebot et al., 1991; Lebot and Lévesque, 1996b) revealed a reduced level <strong>of</strong><br />

genetic variability within P. methysticum; only four <strong>of</strong> eight enzyme systems examined<br />

were polymorphic, whereas all systems tested in P. wichmannii were polymorphic.<br />

<strong>The</strong>se studies provided important information relevant to the area <strong>of</strong> origin <strong>of</strong><br />

P. methysticum. Multivariate analysis <strong>of</strong> presence/absence <strong>of</strong> “electromorphs” indicated<br />

that P. wichmannii from the Western Province <strong>of</strong> Papua New Guinea are very<br />

different from those <strong>of</strong> P. methysticum, suggesting that the cultivated form did not<br />

originate from that area. <strong>The</strong> closest “match” came from cultivated plants collected<br />

in Vanuatu and southern Papua New Guinea, an observation that the authors took to<br />

indicate that cultivated kava might have had its origin in that general area.

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