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The Geography of Phytochemical Races

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28 2 Examples Within Continents<br />

preference for the acyanic form <strong>of</strong> white clover. It is interesting to note that in the<br />

outdoor experiments lasting for 2–3 weeks, animals adjusted their dietary intake <strong>of</strong><br />

cyanogenic clover relative to the acyanic form to a sustainable level. <strong>The</strong> authors<br />

also reported feeding preferences for the acyanic form by two species <strong>of</strong> slugs,<br />

Arion ater and A. subfuscus.<br />

Additional information relating to cyanogenesis polymorphisms can be found in a<br />

recent paper by Schappert and Shore (1999) who studied the phenomenon in Turnera<br />

ulmifolia L. (Turneraceae), which is used by Euptoieta hegesia (Lepidoptera: Nymphalidae)<br />

as its primary host plant.<br />

A further note pertaining to cyanogenic taxa, although not concerned with geographic<br />

differences per se, could have relevance in discussions <strong>of</strong> relationships <strong>of</strong><br />

species or populations based upon cyanogenesis. It has generally been held that a<br />

positive reaction implies the functioning <strong>of</strong> the two genes, Ac and Li, required for<br />

synthesis (one that directs making the glycoside and one that codes for the hydrolyase),<br />

and a negative reaction implies the lack <strong>of</strong> one or both. Kakes and Chardonnens<br />

(2000), working with mixed populations <strong>of</strong> the related species Trifolium<br />

repens and T. occidentale Coombe in Bretagne, demonstrated that the hydrolyase<br />

linamarase was absent from all individual plants examined (763). <strong>The</strong> apparent lack<br />

<strong>of</strong> linamarase from T. occidentale had been reported some years earlier (Gibson<br />

et al., 1972), but only ten plants were used in that work. <strong>The</strong> more recent results<br />

strongly suggest that cyanogenic frequencies in these two closely related species are<br />

controlled by different mechanisms. In other words, cyanogenesis does not have the<br />

same genetic meaning in these two species.<br />

It is useful to include a recently described example <strong>of</strong> cyanogenic polymorphism<br />

here, although the species involved is neither related to the clovers nor does it occur<br />

in Europe. <strong>The</strong> work in question comes from a study <strong>of</strong> variation in cyanogenesis<br />

in two populations <strong>of</strong> Eucalyptus polyanthemos Schauer subsp. vestita L. Johnson<br />

and K. Hill from the Brisbane Ranges National Park in Victoria, Australia (Goodger<br />

et al., 2002). Two sets <strong>of</strong> trees were sampled, 100 from a population at Stony<br />

Creek, and 201 from a population at Sutherland Creek. All trees (100/100) from<br />

the Stony Creek set were cyanogenic with concentrations in the range 0.17–1.98 mg<br />

cyanide per gram dry weight leaves. Six acyanogenic trees were detected within the<br />

Sutherland Creek set (195/201) with maximum concentration <strong>of</strong> cyanide in the others<br />

reaching 2.07 mg cyanide per gram dry weight leaves. <strong>The</strong> authors, assuming no<br />

sampling error, speculated that the absence <strong>of</strong> acyanogenic trees in the Stony Creek<br />

population might be the result <strong>of</strong> selection against some localized herbivore. Again,<br />

follow-up studies <strong>of</strong> this system would be welcomed.<br />

2.3.3 Lotus corniculatus (Fabaceae)<br />

A few paragraphs above, we saw the part played by Lotus corniculatus in the cyanogenic<br />

glycoside story. We now turn our attention to another aspect <strong>of</strong> this species<br />

that has attracted a considerable amount <strong>of</strong> attention and has, coincidentally,

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