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The Geography of Phytochemical Races

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46 2 Examples Within Continents<br />

southern Finnish ports in that manner. [Other examples <strong>of</strong> seed transport via ballast<br />

will be met later in this review in a discussion <strong>of</strong> beach rocket (Cakile).] <strong>The</strong> more<br />

fundamental question—how to explain the chemical variation in tansy— however,<br />

remains, to use the authors’ word, “elusive.” It seems reasonable to suggest that<br />

tansy, over a considerable period <strong>of</strong> time, has been subjected to human intervention<br />

with an eye to improving one or another <strong>of</strong> its properties, and it is the seeds <strong>of</strong> those<br />

improved lines that have found their way to northern Europe and subsequently into<br />

Scandinavia. Environmental factors may also have played an important role (Sorsa<br />

et al., 1968). As is the case with most <strong>of</strong> these highly variable systems, carefully<br />

controlled studies focusing on likely environmental factors, coupled with a thorough<br />

understanding <strong>of</strong> the biosynthetic processes involved, should lead to a better<br />

appreciation <strong>of</strong> why such complex systems occur in nature.<br />

<strong>The</strong> fi nal Finnish example here deals with the question <strong>of</strong> susceptibility <strong>of</strong> Scots<br />

pine (Pinus sylvestris L.) to insect herbivory. Manninen et al. (1998) studied the terpene<br />

and resin acids and growth characteristics <strong>of</strong> trees from four provenances that<br />

ranged over a distance <strong>of</strong> about 1200 km: Muonio (67°56′N), Suomussalmi (65°10′N),<br />

and Korpilahti (62°0′N) in Finland, and Saaremaa (58°22′N) in Estonia. Experiments<br />

were run either in petri plates or in pots under common garden conditions. Host plant<br />

selection was tested using two species <strong>of</strong> aphids, the gray pine aphid (Schizolachnus<br />

pineti) and the spotted pine aphid (Eulachnus agilis), while measures <strong>of</strong> oviposition<br />

behavior employed the tarnished plant bug (Lygus rugulipennis). <strong>The</strong> terpenes, separated<br />

by HPLC, were identifi ed as bornyl acetate [95] (see Fig. 2.26 for structures<br />

95–103, others above), camphene [96], 3-carene [97], caryophyllene [98], copaene [99],<br />

Fig. 2.26 Compounds 95–105 from Pinus sylvestris in Finland and Estonia

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