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The Geography of Phytochemical Races

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2.3 Europe 31<br />

conditions revealed that pigment deposition (presumably synthesis) does not occur<br />

if the temperature is maintained above 20°C.<br />

2.3.4 Silene latifolia (= alba, = pratensis) (Caryophyllaceae)<br />

This discussion features an analysis <strong>of</strong> the fl avonoid genetics <strong>of</strong> Silene latifolia and<br />

how those data, among others, provide insights into the development <strong>of</strong> agriculture<br />

in Europe. Before looking at the fl avonoid chemistry <strong>of</strong> the group, it is necessary<br />

to deal with some nomenclatural matters. In the earlier literature, this taxon was<br />

referred to as Melandrium album. Referral to Silene required the taxon to be named<br />

S. pratensis (Rafn.) Godron and Gren., since S. alba (Miller) E. H. L. Krause had<br />

been shown to be illegitimate by McNeill and Prentice (1981). A further change was<br />

occasioned by the discovery <strong>of</strong> a specimen under the name S. latifolia Poiret that<br />

carried an earlier publication date (see comments in Mastenbroek and van Brederode,<br />

1986, p. 166). Silene latifolia is closely related to S. dioica (L.) Clairv., and<br />

is also widely distributed in Europe. <strong>The</strong>y differ in fl ower color; S. dioica has red<br />

fl owers, whereas S. latifolia, as its earliest name recognizes, has white fl owers.<br />

Before embarking on a discussion <strong>of</strong> Silene, a few comments on the signifi -<br />

cance <strong>of</strong> the work are in order. Biochemical systematics or chemotaxonomy as the<br />

subject was more <strong>of</strong>ten called in the 1960s and early 1970s, consisted in the early<br />

days essentially <strong>of</strong> comparing chromatographic patterns and developing a table <strong>of</strong><br />

presence/absence data for the taxa under consideration. This was the “spot counting”<br />

phase <strong>of</strong> the subject. Occasionally, statistical treatments <strong>of</strong> presence/absence<br />

data were employed to establish levels <strong>of</strong> signifi cance <strong>of</strong> the arrays <strong>of</strong> chemical<br />

data, which, incidentally, was an important contribution made by another fl edgling<br />

discipline <strong>of</strong> the time, namely, numerical taxonomy. As the subject matured, more<br />

and more attention was given to determination <strong>of</strong> exact structures and, logically,<br />

how these structures were constructed in the cell. <strong>The</strong> establishment <strong>of</strong> biosynthetic<br />

pathways led to the study <strong>of</strong> the enzymes responsible for individual steps<br />

and, ultimately, to the question <strong>of</strong> the genetic control <strong>of</strong> these reactions. Flavonoids<br />

provided particularly good test subjects for genetic studies owing to the ease <strong>of</strong><br />

scoring the products <strong>of</strong> controlled crosses, that is, the accumulation <strong>of</strong> intermediates<br />

when an enzyme was absent, or, more readily, by correlating fl ower color with<br />

genotype.<br />

<strong>The</strong> subject more recently has progressed to the stage <strong>of</strong> probing <strong>of</strong> the genome<br />

using allozyme/isozyme analyses, endonuclease restriction fragment analysis<br />

(RFLP), or sequence analysis <strong>of</strong> a variety <strong>of</strong> genes themselves, although very few<br />

<strong>of</strong> these focuses on enzymes or genes coding for fl avonoid biosynthetic reactions.<br />

Ready availability <strong>of</strong> these techniques has tended to overshadow use <strong>of</strong> secondary<br />

metabolites (not just fl avonoids) in systematic studies. <strong>The</strong> current situation can be<br />

appreciated by quoting from a work by Crawford et al. (1992b) that addressed the<br />

comparative value <strong>of</strong> the two types <strong>of</strong> data: “Flavonoid chemistry <strong>of</strong>fers few advantages<br />

over morphology because it is diffi cult, if not impossible, to infer genetic

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