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The Geography of Phytochemical Races

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2.7 North and Central America 105<br />

confi rmed the visual comparisons. <strong>The</strong> mass <strong>of</strong> information generated by the HPLC<br />

analyses was subjected to multivariate statistical analysis. Principal- components<br />

analysis revealed a large degree <strong>of</strong> dispersion in the data, with orientin dominating<br />

axis I (axis I accounted for 72% <strong>of</strong> total variation), isoörientin [213, R 1 = glucose,<br />

R 2 = hydrogen] dominating axis II (12%), and orientin 3′-O-glucoside [214] dominating<br />

axis III (10%). Despite the large contribution from orientin, no clear-cut<br />

grouping <strong>of</strong> populations appeared to exist in most <strong>of</strong> the northern and central<br />

parts <strong>of</strong> the range. <strong>The</strong> southernmost populations are more discernible than those<br />

throughout the rest <strong>of</strong> the range, but the major outcome <strong>of</strong> these analyses was<br />

recognition <strong>of</strong> clinal variation in a north–south axis, with the high-orientin plants<br />

originating from the southernmost populations. This agrees well with an observation<br />

made by Fleak (1971) on plants from the southern populations, which consistently<br />

display heavy anthocyanin pigmentation in the lower part <strong>of</strong> the stems. Stem<br />

pigmentation was suffi ciently variable in plants from the rest <strong>of</strong> the range <strong>of</strong> the<br />

species that Fleak and Dunn (1971) recognized the southern plants as L. sericeus<br />

subsp. huffmanii. Accumulation <strong>of</strong> high concentrations <strong>of</strong> orientin in these individuals<br />

provided support for their view. Flavonoid data provide no support, however,<br />

for three other subspecies defi ned by Fleak (1971).<br />

A resampling <strong>of</strong> populations, again representing the entire range <strong>of</strong> the species,<br />

provided seed so that plants could be grown under controlled conditions <strong>of</strong> day<br />

length and temperature. <strong>The</strong> fl avonoid fraction from each individual was then subjected<br />

to HPLC analysis as before. Under uniform conditions, all quantitative differences<br />

in the fl avonoid pr<strong>of</strong>i les disappeared! Further studies to determine what<br />

environmental factor or factors is/are responsible for the quantitative differences in<br />

orientin concentration in nature have not been undertaken. It is interesting to note<br />

in passing that plants from the southern populations retained their red pigmentation<br />

under the controlled conditions <strong>of</strong> the growth experiment, thus providing support<br />

for the use <strong>of</strong> anthocyanic pigmentation as a character in suggesting subspecifi c<br />

status for those individuals.<br />

2.7.15 Satureja douglasii (Lamiaceae)<br />

Terpene pattern variation has provided a useful means <strong>of</strong> investigating evolutionary<br />

and ecological situations in a variety <strong>of</strong> plant groups. <strong>The</strong> next example, a study <strong>of</strong><br />

the monoterpenes <strong>of</strong> Satureja douglasii (Benth.) Briq. by Lincoln and Langenheim<br />

(1976), is an excellent example <strong>of</strong> an investigation aimed at establishing possible<br />

factors responsible for chemical variation exhibited by a species throughout its<br />

range. Satureja douglasii occurs in a narrow band paralleling the coast from southern<br />

British Columbia to southern California and disjunctly in a narrow band in the<br />

western half <strong>of</strong> the Idaho panhandle and adjacent British Columbia. Sampling for<br />

this study involved the coastal and near-coastal sites and constituted a north–south<br />

transect between approximately 50°N and 34°N latitude (Fig. 2.67). (A specimen<br />

from Idaho was analyzed but does not fi gure in the study.)

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