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The orchid Dactylorhiza sambucina shows a stable and dramatic flower-color polymorphism, with both yellow-and purple-flowered individuals present in natural populations throughout the range of the species in Europe. The evolutionary significance of flower-color polymorphisms found in many rewardless orchid species has been discussed at length, but the mechanisms responsible for their maintenance remain unclear. Laboratory experiments have suggested that behavioral responses by pollinators to lack of reward availability might result in a reproductive advantage for rare-color morphs. Consequently, we performed an experiment varying the relative frequency of the two color morphs of D. sambucina to test whether rare morph advantage acted in the natural habitat of the species. We show here clear evidence from this manipulative experiment that rare-color morphs have reproductive advantage through male and female components. This is the first demonstration, to our knowledge, that negative frequency-dependent selection through pollinator preference for rare morphs can cause the maintenance of a flower-color polymorphism.

SUMMARY In many parts of the world soils are detrimental to plant growth owing to elevated levels of metal ions, caused either by natural processes or by the result of man's activities. Many plants have evolved ecotypes or varieties that are able to grow more‐or‐less normally on these soils. This paper reviews our knowledge of the genetics of this phenomenon. The nature of tolerance and the problems of its measurement are discussed. Tolerance is frequently measured by an index produced by comparing growth in a contaminated environment with growth in a control environment. It is argued that this measurement is inappropriate for many genetical studies, and that it is frequently more useful to use growth at a single critical level of metal as a measure of tolerance. Polygenic inheritance provides a null hypothesis that has to be tested in a genetical analysis. Examples of major genes for tolerance to aluminium, arsenic, boron, cadmium, copper and manganese are discussed. Even where major genes have been demonstrated, it is probable that other minor genes, ‘modifiers’, are present as well. Because of the nature of tolerance as a character, dominance and epistasis are likely to vary with the level of metal at which an analysis is performed. Tolerance is generally found to be dominant at some levels of the metal. Studies which have mapped tolerance genes, particularly to aluminium and salt, are discussed. The specificity of tolerance is a matter of some confusion. Some studies indicate that tolerances evolve independently to different metals, but others have suggested that tolerance to one metal may often confer a degree of tolerance to some other metals. Very little is known about the molecular genetics of tolerance, and the mechanisms of tolerance to most metals. The possible role of phytochelatins and metallothionein‐like proteins in metal tolerance is discussed. The distribution of tolerance in natural populations suggests that tolerance is a disadvantage in uncontaminated environments, but how this ‘cost’ arises is not known. There is some evidence that the disadvantage to tolerance may be associated more with the modifiers of tolerance than with the primary tolerance gene. The study of the genetics of tolerance is of importance in planning breeding programmes to produce tolerant crops for use in areas where metal contamination is a limiting factor in productivity. It can also assist in understanding the mechanisms of tolerance, as exemplified by the study of the mechanism of arsenic tolerance in Holcus lanatus. Important areas for further research are discussed.

Zinc tolerance was investigated in five populations of Arabidopsis halleri (syn. : Cardaminopsis halleri) raised from seeds collected from contaminated and uncontaminated sites. Tolerance was measured by determining the concentration which inhibited root growth (EC "!! ). A. halleri populations from contaminated and uncontaminated sites were found to be Zn-tolerant compared with the Zn-nontolerant species Arabidopsis thaliana and A. lyrata subsp. petraea. At very high Zn concentrations, populations of A. halleri from uncontaminated sites were slightly less Zn-tolerant than those from contaminated sites. These observations support the hypothesis that in A. halleri, Zn tolerance is largely a constitutive property. One population from an uncontaminated site and one population from a contaminated site were studied for Zn uptake. Zinc content was measured in shoots and roots using a colorimetric test under laboratory conditions. The results showed that whatever their origin, individuals from both populations are Zn accumulators compared with the nonaccumulator species A. thaliana. Moreover, the population from the uncontaminated area accumulated Zn in its shoots and roots more quickly than the population from the contaminated site. These results suggest that, in A. halleri, Zn accumulation to very high concentration is a constitutive property.